ISSN 0198-7356

BARTONIA

JOURNAL OF THE
PHILADELPHIA BOTANICAL CLUB
No. 57, Supplement
Symposium on Rare Plants of Pennsylvania and Adjacent States
Held March 28, 1991. Jointly sponsored by the

Philadelphia Botanical Club and the
Morris Arboretum of the University of Pennsylvania.

CONTENTS
fg Dg Ps eae eS re ee ee re a TED GorDon 1
Rare Plants and Plant C bties GF Presiue 1ale ioc rss ye. JAMES K. BISSELL 2
The Collecti f Albert C Delmarva, pe tes with Attention to August 4-5, 1874
and September 9-10; 1875... 3 5 ee RTHUR O. TUCKER AND NORMAN H. Dut <9
Rare Vascular Pl 4 A 3 re a eed 4 : c 4k YQ. ape Qn nia é <A Z 2
Larry H. KLorz AND esac + Wes 6

Hill Creek as a Case Study -biyn aes eee RODNEY L. BARTGIS AND DAVID MADDOX 4.

42
Extinct, Extant, Extirpated or Historical? Or in Defense of Historical Species . . . DAvip B. SNYDER ae
-

Commie Matiral Attest laveliede: S50 = ee ae ANTHONY F. pas: ee
The Serpentine Barrens of T: te Eastern North America: Critical I in tk a 3G
of Rare Species and Commaniiies oo = he ee ee ROGER EARL . LATHAM A a

Selected R a) eer oMon LV: scular DI + £ Tol.

Bartonia No. 57, Supplement: 1, 1993
Introduction

TED GORDON
President, Philadelphia Botanical Club

This is the first of three symposia presented by the Philadelphia Botanical Club to
celebrate its centennial. Indeed, it is fitting that the co-sponsor of a symposium on rare
plants of Pennsylvania and adjacent states be our host, the Morris Arboretum of the
University of Pennsylvania. In 1979, the Arboretum published the Atlas of the Flora of
Pennsylvania, and since that time, annual field surveys to update our information on critical
species have been conducted under the leadership of Ann Rhoads, Chair of Botany here at
the Arboretum. The development of a computerized database and the investigation of the
applications for this database, in the use of development planning, for example, are vital
components of this undertaking.

Although collecting-trips were initiated by the Morris Arboretum in the mid 1930s,
documentation of the local flora by the Philadelphia Botanical Club began even earlier. It
was 1891 when the Club organized at the Academy of Natural Sciences “to further the
interest of botany and to make a checklist and herbarium of the plants within a radius of 50
miles of Philadelphia,” a jurisdiction that, of course, extends beyond the borders of
Pennsylvania into New Jersey, Delaware, and Maryland. In the past 100 years, club
members have contributed thousands of specimens to the Academy herbarium. The local
collection is one of the finest in the world! Thanks to the steadfast encouragement of
Alfred E. Schuyler, editor for more than twenty years of Bartonia, our Club’s journal,
interest in monitoring the local flora continues to mount. Judging from the ambitious field
trip schedules of the past several years, the Club has not lost any of the enthusiasm
exhibited by its founders.

MISSOURI BOTANICAL
JAN 25 1995

GARDEN LIBRARY

Bartonia No. 57, Supplement: 2-8, 1993
Rare Plants and Rare Plant Communities of Presque Isle

JAMES K. BISSELL
The Cleveland Museum of Natural History, Wade Oval, University Circle, Cleveland, OH 44106

The flora and vegetation of Presque Isle Peninsula, the finest example of a recurved,
compound sand spit along the southern shore of Lake Erie, were surveyed by the Botany
Department of The Cleveland Museum of Natural History from May 1985 through 1990. A
floristic survey for all vascular plants was conducted and rare plant communities and
specific occurrences of rare vascular plants were mapped for the peninsula and adjacent
waters of Lake Erie. Responses of rare natural communities to fluctuating lake levels were
documented. Observations were made on apparent threats to rare species including exotic
species encroachment, deer overpopulation, and i tly poor t practices

The Museum initiated this study because 79 years had elapsed since the last thorough
study of plants on the peninsula. That study, conducted in 1905 and 1906 and published in
1909 by Otto E. Jennings, documented the occurrence of several dozen plants currently
listed as Plants of Special Concern in Pennsylvania (POSCIP) by the Pennsylvania Natural
Diversity Inventory (PNDI), Department of Environmental Resources. John Miller’s
Flora of Erie County (1923) provided some important historical rare plant records not
included in Jennings (1909).

DESCRIPTION OF AREA
Presque Isle Peninsula extends northeast into Lake Erie from Mill Creek Township,

Erie County, Pennsylvania. The City of Erie is located on the mainland south of the

peninsula, and the body of water between the city and the peninsula is known as Presque
Isle Bay. Most of the peninsula (approximately 6 miles of its length) is located on the Erie
North, PA U.S.G.S. 7 1/2 minute quadrangle map. One mile of the peninsula, including
the attachment to the mainland is located on the Swanville, PA quadrangle map. All
features referred to in this paper can be found on the topographic maps.

Most of the 3,000 acres of the peninsula are included within Presque Isle State Park,
established in 1921. PA Bureau of Parks manages the park for both recreation and
conservation. At this time most of the park is undeveloped except for 11 bathing beach
areas and a perimeter state highway (PA Rt. 832) that provides access to beach areas.
There are relatively few buildings within the park but each beach area has restrooms and
there are several park buildings.

For thousands of years, waves generated by prevailing westerly winds have eroded lake
bluffs to the west of Presque Isle. The sand component from the bluffs moves eastward
along the shoreline by littoral drift to the base of the peninsula and along the lake side
(northwest shore) of the peninsula. Relatively light, wind-blown silica sand tends to
accumulate on the upper beach shelf, and is crucial to maintaining dunes along the lake
side of the peninsula.

Shoreline jetties, sheet piling, and offshore breakwaters placed along the lake side of the
peninsula during the last century have accumulated sand from the littoral drift system to
the windward (westerly) side of structures and contributed to shoreline erosion downshore
(east) of the structures. In an attempt to counter severe beach erosion that has taken place
along the lake side of the peninsula between the mainland and the signal tower, PA

4

RR RIT ermg sears etrret conn co i

RARE PLANTS OF PRESQUE ISLE 5

Bureau of Parks and the Corps of Engineers have carried out a sand replenishment
program. The program began in the 1950s but annual nourishment did not commence until
1974 (Gorechi 1985). Unfortunately, much of the replenishment sand is bank gravel
instead of angular silica sand that is so important to dune building.

The major ridges within the park trend from northwest to southeast. The development
of these ridges has been linked by Jennings (1909) to major northeasterly storm surge
events. New ridges and ponds form at the northeast edge of the peninsula and successively
older ridges and ponds are located to the southwest.

A great diversity of natural communities has developed on the complex system om sume
ridges, ponds, and bay shore waters. The following
the six year study: mixed hardwood forests (dominated by Quercus borealis—Quercus velutina—
Prunus serotina—Acer rubrum), Populus deltoides, sand dunes, emergent marshes, shrub
swamps, mixed swamp forests, palustrine sand plains, dry sand plains, and oak savannahs.

METHODS

More than 100 full field days were spent searching for POSCIP species historically
reported for the peninsula. (Plants recommended for addition to the POSCIP list at the
January 1989, 1990, and 1991 meetings of the PA Rare Plant Committee are included as
rare species for this paper.) Many Carnegie Museum specimens from the area were
examined along with the John Miller Collection at Edinboro University. (The Miller
Collection is now at Carnegie Museum.)

All POSCIP species were mapped on the U.S.G.S. quad maps. Prints made from high
resolution aerial negatives on loan to the Museum from the U.S. Army Corps of Engineers
were used to map rare natural communities. The scale of the aerials is 1 inch:400 feet.

Voucher specimens of all rare species, except Opuntia humifusa that was too rare to
collect, have been placed in the Cleveland Museum of Natural History Herbarium, and all
rare plant occurrences have been reported to PNDI.

RESULTS AND DISCUSSION

Sixty-four species listed as Plants of Special Concern, or suggested as additions to the list
at the January 1991 PA Rare Plant Committee Meeting, were documented at Presque Isle
from 1985 through 1990. More than 400 occurrences of the 64 species were mapped.

The following 15 species, historically reported from the peninsula, were not found
(those marked with an “‘*” have not been examined by the author): Arethusa bulbosa, Carex
sartwelli*, Carex tetanica, Eriophorum gracile, Populus balsamifera*, Potamogeton gramineus,
Potamogeton praelongus, Potamogeton robbinsii, Potamogeton strictifolius*, Salix candida*,
Salix serissima*, Scirpus torreyi, Scleria verticillata, Sparganium minimum, Triglochum
palustre, and Utricularia resupinata.

The 64 rare species are found in the following six rare communities. No rare species
were found within full canopy forests but rare dry sand plain species were found
establishing within recent sand blowouts on the forested ridges. The number of rare
species in each community is shown in parentheses.

Palustrine Sand Plain (27)

Dry Sand Plain and Oak Savannah (8)
Dunes and Dry Beaches (10)

Aquatic Bed (11)

Mixed Emergent Marshes (7)

Shrub Swamp (1)

4 BARTONIA

+1 P|

A seventh rare community, a sphagnum wetland communi ity, described as the “Sphagnum-
Oxycoccus Formation” by Shank (1909), was not found during the study. Two of the 15
rare plants known only from historical records, Arethusa bulbosa and Eriophorum gracile,
were probably restricted to this rare habitat.

There is a historical record for an eighth community. A specimen of the rare tree,
Populus balsamifera, was presumably taken from the swamp forest community at Presque
Isle by Jennings in 1906.

Four rare plants were found that had never before been reported for Presque Isle:
Eleocharis caribaea, Eleocharis intermedia, Geranium bicknellii, and Myriophyllum verticilla-
tum. Eleocharis caribaea and Myriophyllum verticillatum are additions to the flora of
Pennsylvania.

Based upon Wherry et al. (1979), examination of Carnegie specimens, and field work by
the author, the following 12 rare taxa have been reported only from Presque Isle within
Pennsylvania: Ammophila breviligulata, Artemisia campestris ssp. caudata, Carex, aquatilis,
Carex garberi, Carex viridula, Eleocharis caribaea, Eleocharis pauciflora, Prunus pumila, vat.
pumila, Schizachyrium scoparium var. littoralis, Sparganium minimum, Triglochin palustris,

nd Utricularia resupinata.

Successional changes within all rare communities at Presque Isle, except the highest
dunes and dry sand plains, are not unidirectional or generally predictable because Lake
Erie levels drive those changes. When the lake level rises: (1) Swamp forest and shrub
swamps die back and are replaced by emergent marshes. (2) Emergent marshes along
interior pond shores and Presque Isle Bay are replaced by the aquatic bed community. (3)
Dry sand plains on interdunal flats, trail flats, and mowed flats are replaced by the
palustrine sand plain community. When the lake level falls: (1) Swamp forests and shrub
swamps establish again within emergent marshes. (2) Palustrine sand plain replaces the
aquatic bed along the shores of interior ponds and Presque Isle Bay. (3) Emergent marsh
species, especially Calamagrostis canadensis, advance into the palustrine sand plain
community as the palustrine sand plain moves downslope onto newly exposed beach flats.
(4) Dry sand plains replace palustrine sand plain communities on interdunal flats, trail
flats, and mowed flats.

PALUSTRINE SAND PLAIN. The palustrine sand plain is an ephemeral community that
occupies open, moist, sparsely vegetated, sandy flats. Standing water is often present in the
spring and the water table often drops below the surface during the summer. This
community contains 25 rare plants, more than any other in the park. The distribution and
abundance of the community is tied to the level of Lake Erie. Typical palustrine sand plain
species include: Cyperus flavescens, Cyperus bipartitus, Hypericum majus, Juncus alpinus,
Juncus articulatus, and Juncus brachycephalus.

Many of the rarest species of this community are not present in the park every year.
During the high water levels of 1985 and 1986, the community flourished upon moist flats
within depressions along inland trails and on wet flats between dunes. When the lake level
dropped in 1987 and 1988, visits to the interior trail flats and interdunal flats revealed that
the palustrine sand plain community had apparently disappeared from those sites and
been replaced largely by dry sand plain species.

Seeds of many of the palustrine sand plain species are apparently stored within dry
sands along trail flats during periods of low lake levels and in the moist sandy or peaty
substrates beneath the water surface of the interior ponds and the Presque Isle Bay
shoreline during periods of high water levels.

RARE PLANTS OF PRESQUE ISLE 5

During 1985 and 1986, when the lake was very high, no palustrine sand plain community
was found along the shore of Presque Isle Bay from Big Bend to the mainland. When the
level of Lake Erie dropped in 1987 and 1988, thousands of individuals of the rare plants
that compose the community appeared on newly exposed low-relief beach shelves along
the Presque Isle Bay shore. This phenomenon was also observed along interior pond
shores

Park management has a profound impact on this community. Many of the rarest species
within the community tolerate heavy foot traffic and flourish on trails and frequently
mowed areas of the park. Through the six year study, Calamagrostis canadensis and other
species including Typha x glauca, Phragmites australis, Epilobium hirsutum, Lythrum
salicaria and Phalaris arundinacea were observed to invade and quickly replace the
community except on areas receiving heavy foot traffic or frequent mowing during the
growing season. During the high lake levels of 1985 and 1986, large populations of several
rare palustrine sand plain species were found to be commonplace on wet flats that were
mowed for public use or on heavily used wet flats along trails. Adjacent unmowed and
untrampled areas, typically dominated by Calamagrostis canadensis, supported no palus-
trine sand plain species. The following rare species were found only on trail flats: Carex
garberi, Eleocharis caribaea, and Eleocharis pauciflora.

There is great potential for increasing the acreage of this rare community at Presque
Isle. Hundreds of acres of Myrica pensylvanica—Calamagrostis canadensis flats could be
regularly mowed during the growing season. Park maintenance activities have proved the
effectiveness of this succession management technique in several areas.

Three POSCIP species published by Jennings in 1909 that would have inhabited this
community were discovered to be other species. Carex schweinitzii was determined to be
Carex hystericina, Juncus brachycarpus was determined to be Juncus torreyi, and Eleocharis
compressa var. atrata was determined to be Eleocharis elliptica. Four historical species,
presumably members of this community, were not found (the last year reported is shown in
parentheses): Carex tetanica (1869), Scleria verticillata (1911), Triglochin palustre (1905),
and Utricularia resupinata (1879). The following rare species were found within the
community (an “*” denotes that the species was found associated with wet trail flats):
Agalinis paupercula*, Aster dumosus*, Carex bebbii*, Carex garberi*, Carex viridula*, Cyperus
diandrus*, Cyperus engelmannii*, Cyperus odoratus*, Eleocharis caribaea, Eleocharis ellipti-
ca*, Eleocharis intermedia*, Eleocharis olivacea*, Eleocharis pauciflora*, Equisetum variega-
tion, Hemicarpha micrantha*, Hypericum majus*, Juncus alpinus*, Juncus balticus*, Juncus
biflorus*, Juncus brachycephalus*, Juncus torreyi*, Lathyrus palustris*, Lobelia kalmii,
Parnasia glauca, Potentilla anserina*, Potentilla paradoxa, and Scirpus smithii.

DRY SAND PLAIN AND OAK SAVANNAH. The dry sand plain is an open, dry grassland
usually dominated by Sorghastrum nutans and Schizachyrium scoparium vat. littoralis. This
community often occurs as a savannah with widely scattered Quercus velutina, Quercus
rubra, and Sassafras albidum. The community also establishes on blowouts within
hardwood forests. Other species common in the dry sand plain include: Panicum virgatum,
Carex muhlenbergii, Carex tonsa, Rumex acetosella, and Dichanthelium sabulorum. The alien
species, Lonicera morrowii, has invaded much of this community within the park. ’

One species in this community, Lithosperum caroliniense, is on the brink of extirpation
within Pennsylvania due to severe grazing by deer. Less than 100 plants occur on the dry
sand plains on the peninsula. Arctostaphylos uva-ursi has apparently been extirpated from
both the peninsula and the state. Shepherdia canadensis is another shrub that has been

6 BARTONIA

extirpated from the peninsula. Geranium bicknellii was found only during 1986 at a site
disturbed during construction of a restroom. The species is known to seed bank between
episodes of fire within this community at the Indiana Dunes National Lakeshore (Wilhelm
1980). Fire suppression at the Indiana Dunes area has been blamed for deterioration of
the community (Wilhelm 1980) and fire suppression has likely had the same negative
impact at Presque Isle.

Eight rare species are found on the dry sand plain: Dichanthelium sabulorum,
Dichanthelium oligosanthes, Digitaria cognatum, Geranium bicknellii, Lithospermum carolin-
iense, Lupinus perennis, Opuntia humifusa, and Schizachyrium scoparium vat. littoralis.

DUNES. Sand Dunes and drift beaches at Presque Isle support ten rare species:
Ammophila breviligulata, Artemisia campestris ssp. caudata, Cakile edentula, Chamaesyce
polygonifolia, Cyperus schweinitzii, Lathyrus japonicus, Ptelea trifoliata, Schizachyrium
scoparium var. littoralis, Sporobolus cryptandrus, and Triplasis purpurea.

Ammophila breviligulata and Populus deltoides are the most frequent dune builders, but
in some areas Panicum virgatum, Schizachyrium scoparium var. littoralis, and Elymus
canadensis are also important. Active dunes at Presque Isle are best developed from the
Light House eastward to the tip of the peninsula and fine Ammophila dunes are located
along the shore of Thompson Bay, northeast of Horseshoe Pond.

The dunes at Presque Isle are the highest impact area from the 5 million visitors that
annually use the park. Ammophila breviligulata stands on some of the dunes between the
Lighthouse and the Signal Tower have breaks in the most heavily trampled sections.

Some POSCIP species characteristic of dunes, including Cyperus schweinitzii and
Sporobolus cryptandrus, grow in abundance on trampled grounds at the highest use areas
on the dunes and beaches. Only one rare species, Prunus pumila var. pumila, reported as a
member of the dune vegetation by Jennings in 1909, was not found during the present
study. Deer browsing may account for the absence of this plant. The beach areas between
the Light House and Signal Tower are browsed so heavily that Salix exigua and Salix
eriocephala seldom exceed a foot in height.

The on-going construction of 58 off-shore breakwaters along the lake side of the park
from just north of the mainland to east of the Light House may have a severe impact on the
dune community at Presque Isle. Twenty of the breakwaters have been installed.
According to information presented at a public hearing held by the U.S. Army Corps of
Engineers and PA Bureau of Parks on July 22, 1988, the 58 off-shore breakwaters may
reduce the littoral drift of sand by as much as 65%. This reduction may have a negative
impact on the formation of new dunes.

AQUATIC BED. This plant community is found along the entire Presque Isle Bay
shoreline and within open ponds in the interior section of the park. The interior ponds
which have well developed aquatic beds are sometimes sparsely covered with Nuphar
advena, Nymphaea odorata, Brasenia schreberi, and Potamogeton natans.

One alien submersed aquatic plant, Myriophyllum spicatum, apparently poses a threat to
submersed aquatic plants listed as POSCIP species. Based upon examination of herbarium
specimens, Myriophyllum spicatum was first collected at Presque Isle in 1950. Prior to that
time several specimens of the rare Myriophyllum sibiricum had been collected from Presque
Isle, but it has not been reported since. According to Cooperrider (1982), this replacement
has also taken place in Ohio, and Reznicek and Catling (1989) reported Myriophyllum
sibiricum to be confined to a single small pond at Long Point, across the lake from Presque
Isle. Presque Isle Bay and all interior ponds at Presque Isle that have open water channel

NOP RRResinsen di cchome

RARE PLANTS OF PRESQUE ISLE 7

connections to Lake Erie contain large populations of Myriophyllum spicatum. Three rare
submersed species, Myriophyllum verticillatum, Myriophyllum heterophyllum, and Najas
gracillima occur within interior ponds that do not have populations of Myriophyllum
spicatum. Continued isolation of interior ponds may be necessary for long term survival of
rare submersed aquatics now inhabiting them.

The only historical record for Myriophyllum heterophyllum was collected by John Miller
from Big Pond in 1899. Big Pond currently supports large beds of Myriophyllum spicatum
and no Myriophyllum heterophyllum, but Myriophyllum heterophyllum is abundant within
Ridge Pond and one other pond without a channel connection to Lake Erie. Myriophyllum
verticillatum is confined to the unnamed pond between the east shore of Horseshoe Pond
and Thompson Bay. Jennings (1909) labeled this pond Aa and estimated the age of this
pond to be less than 100 years.

Typical members of the aquatic bed community within Presque Isle Bay include:
Vallisneria americana, Elodea canadensis, Ceratophyllum dermersum, Najas flexilis, Najas
guadalupensis, Potamogeton pusillus var. pusillus, and Potamogeton richardsonii. The single
collection of Potamogeton perfoliatus from Misery Bay in 1989 may be the only known
extant population of this species within Lake Erie. According to Cooperrider (1982), the
species was last collected from Ohio in 1896 and Reznicek and Catling (1989) did not find
the species at Long Point during their recent survey there.

Eleven submersed POSCIP species were documented within aquatic beds at Presque
Isle: Megalodonta beckii, Najas gracillima, Myriophyllum heterophyllum, Myriophyllum
verticillatum, Potamogeton perfoliatus, Potamogeton illinoensis, Potamogeton richardsonii,
Potamogeton zosteriformis, Ranunculus longirostris, Utricularia intermedia, and Utricularia
minor. Four species, historically reported from the aquatic bed community of Presque Isle,
were not located (the most recent collection date for each is shown in parentheses):
Potamogeton gramineus (1948), Potamogeton praelongus (1930), Potamogeton robbinsii
(1928), and Potamogeton strictifolius (1868).

MIXED EMERGENT MARSHES. The emergent wetland marshes at Presque Isle cover
hundreds of acres and include: Typha swamp, Carex stricta—Carex aquatilis tussock
meadows, non-persistent Nuphar-Nymphaea—Pontederia marshes, and Acer rubrum—
Quercus palustris-Nyssa—Cephalanthus savannahs with extensive mixed herbaceous marsh
openings. Typha x glauca and Phragmites australis have aggressively invaded much of this
community type within the park. Phalaris arundinacea also poses a threat to the emergent
marshes. There are no historical records for Phalaris, small populations have become
established between Big Bend and the mainland.

Seven rare emergent wetland species were documented within this community type:
Carex aquatilis, Carex pseudocyperus, Cladium mariscoides, Eleocharis quadrangulata,
Scirpus acutus, Scirpus fluviatilis, and Zizania aquatica. Three species historically reported
from the community were not found (the last date of collection is shown in parentheses):
Carex sartwellii (1867), Scirpus torreyi (1879), and Sparganium minimum (1880).

SHRUB SWAMP. Shrub swamps within the peninsula are often dominated by pure stands
of Cephalanthus occidentalis. Mixed shrub swamps, dominated by mixtures of Cephalan-
thus, Vaccinium corymbosum, Viburnum recognitum, Ilex verticillata, Alnus rugosa, Cornus
amomum spp. obliqua, and Rosa palustris also occur frequently within the peninsula. A
single rare shrub, Salix gracilis, was found in this community. Two rare shrubs, historically
reported from the peninsula were not found (the last date for each is shown in
parentheses): Salix candida (1916), and Salix serissima (1916).

8 BARTONIA

CONCLUSION

Presque Isle contains more Plants of Special Concern in Pennsylvania than any other
area of similar size within the state. Long term survival of the present diversity of rare
species and rare communities can be accomplished by informed park management.

ACKNOWLEDGEMENTS

The author thanks Frederick Utech, Associate Curator of Carnegie Museum Herbar-
ium for providing assistance in examining Presque Isle specimens and Don Snyder, of
Edinboro University for providing access to John Miller’s Presque Isle specimens while
that collection was housed at Edinboro. Financial support was received from the PA Wild
Resources Conservation Fund, PA Coastal Zone Management, Western Pennsylvania
Conservancy and Zurn Industries, Inc. Special acknowledgement goes to Museum
Associate, Beverly W. Danielson, who gave more than 100 days to the Presque Isle
inventory.

LITERATURE CITED

BISSELL, J. K. AND C. K. Bier. 1987. aa oid vein Natural pian d sonar oye for Presque Isle
State Park, Erie County, PA. Pennsylvan
COMMONWEALTH OF PA DEPARTMENT OF See ee RESOURCES. me 28 "PA Code Chapter 82.
Conservation of PA Native Wild Plants
COOPERRIDER, T. S., ed. 1982. Endangered and Threatened Plants of Ohio. Ohio Biol. Surv. Biol. Note No. 16,
Columbus
GorECKI, R. 1985. Coastal Engineering At Presque Isle and the Erie Harbor—Past, Present and Future.
omorrow’s Oasis Today: Proc. Conf. Presque Isle and Erie Bayfront. Erie
JENNINGS, O. E. 1909. A Botanical Survey of Presque Isle, Erie Co., Phsemiicain Ann. Carnegie Museum
:289-42

MILLER, J. 1923. Flora of Erie County, PA. Publ. by the author.

PA RARE PLANT COMMITTEE pommel PA Natural Diversity Inventory. January 1989, bist 1990, January
1991. Lists of Suggested A the Plants of Special Concern in PA. Harrisbur

REZNICEK, A. A DP. M. CaTLING. 1989. Flora of Long Point, Regional Municipality of Haldimand-Norfolk,
Ontario. Michigan Bot 28(3): 99-175.

US Army Corps OF ENGINEERS AND PA STATE BUREAU OF Parks Public Hearing on proposed 58 off-shore
breakwaters. July 22, 1991. Villa Maria College, Er =

Wuerry, E. T., J. M. Foca, JR. AND H. A. WAHL. 1979. Atlas of the Flora of Pennsylvania. Philadelphia: The
Morris Arboretum of the University of Pennsylvania, Philadelphia.

WILHELM, G. 1980. Report on the special vegetation of the Indiana Dunes National Lakeshore. Indiana Dunes
National Lakeshore Research Program Report 80-01. National Park Service, Midwest Region, Omaha.

MER esc e sh

RSet

Bartonia No. 57, Supplement: 9-15, 1993

The Collections of Albert Commons on Delmarva, 1861-1901,
with Attention to August 4-5, 1874 and September 9-10, 1875

ARTHUR O. TUCKER AND NORMAN H. DILL
Department of Agriculture and Natural Resources and Department of Biological Sciences,
Delaware State College, Dover, DE 19901-2275

When Albert Commons died in 1919, his nephews, Frank W. and Howard W. Commons,
donated his herbarium to the Academy of Natural Sciences of Philadelphia. Francis
Pennell (1943) called these collections the “most ample botanical collections ever
assembled from Delaware.” Robert Tatnall (1946) stated: “It gives evidence of care and
discrimination on the part of the collector in the identification of his specimens, and of his
keenness in detecting new or rare material in the field.” At his death, Albert Commons
was also known as a mycologist, having collected 1300 species of fungi (now at the
Carnegie Museum of Natural History, and the Academy) and 160 species of lichens
(Harshberger 1899). Many duplicates of Commons’ collections were sent out, notably to
the Plant Pathology Herbarium of Cornell University, Herbarium Universitatis Floren-
tinae, Gray Herbarium of Harvard University, Missouri Botanical Garden, New York
Botanical Garden (639 sheets), and the Smithsonian Institution (Lanjouw and Stafleu
1954). His importance was such that Ashe (1898) named Panicum commonsianum after
him.

Albert Commons was born the son of John and Ann (Phipps) Commons in the village of
Doe Run, Chester Co., PA on January 23, 1829. Albert was sickly and greatly influenced by
his older brother Franklin. In 1839, Albert, while a student at the Academy at Unionville,
purchased a copy of Darlington’s ‘Flora Cestrica,’ and had a tin collecting box made
(Harshberger 1899). At his brother’s demise in 1842, they had collected about 500
specimens. Albert’s first botanical trip in Delaware appears to have been in 1842, soon
after he moved to the Commons farm near Centreville, New Castle Co., DE.

The 4262 specimens of Albert Commons that we have located are of great interest for
botanists to understand the historical flora of the Delmarva Peninsula. His collections of
Rhynchospora knieskernii and Eupatorium resinosum in 1874 and 1875 are of concern due to
the potential status of these taxa as federal endangered or threatened species. This paper
reconstructs Commons’ field activities for this period and concludes that both species were
collected by Commons in Delaware.

THE CASE OF RHYNCHOSPORA KNIESKERNII

Albert Commons never knew that he had collected Rhynchospora knieskernii. He died
without making such a claim, leaving us his notebook (now at the Claude E. Phillips
Herbarium) with this species marked as something to be sought in Delaware. The facts in
the case are these: (1) On August 5, 1874 Albert Commons collected a specimen that he
labelled “Rhynchospora (?) capillacea. Torr. swamps near Gumboro, Delaware”
(PH540107). (2) On September 10, 1875, he collected a specimen that he labelled
“Rhynchospora axillaris microcephala. Britt. swamps, (Baltimore Hundred) Sussex County,
Delaware” (PH540117). (3) Both of these specimens languished under these names for
67-68 years until Shirley Gale annotated them in 1941 as R. knieskernii Carey (see Gale,

44).

9

10 BARTONIA

What floras did a collector use in 1874 and 1875? Wood (1861) and Gray (1867) were
widely available, but the latter would have been more commonly used. Gray (1867) does
not have keys, and illustrations are given only for genera. Rhynchospora knieskernii is
lumped together with R. capillacea under “spikes chestnut-colored or darker ...,
few-several flowered; stamens 3; bristles usually 6.” The other name that Commons used,
R. axillaris var. microcephala, was not published by Britton until 1892 and did not appear in
Gray’s Manual until the seventh edition (Gray 1908). The handwriting for this specimen is
on a different slant, indicating that it was added later, probably between 1908 and
Common’s death in 1919. A key to Rhynchospora and an illustration of R. knieskernii were
also published by Britton and Brown (1913), but Commons apparently did not check his
sheets of R. knieskernii.

Rawinski and Cassin (1986) stated of R. knieskernii, “‘the 1875 record may have actually
been collected from NJ.” This speculation later was noted in the Endangered Species
Technical Bulletin, a publication of the US Department of the Interior, Fish and Wildlife
Service: “It is believed to be endemic to the Pinelands area of southern New Jersey...”
and “‘Historically, it was found at 38 confirmed sites in New Jersey.” (Anonymous 1990,
1991), without mention of Delaware. The official report from Delaware to the federal
government (Trew and Clancy 1991) furthermore stated: “There is doubt as to the validity
of these records.”

How did these two specimens of Albert Commons assume doubtful status? Conversa-
tions with some individuals hired by a private conservation group revealed their belief that
Delaware lacked any real importance in rare plants and that Commons actually collected
many of his rare plants in New Jersey. Tracing this claim further revealed the rumor that
Bayard Long (1885-1969) had remarked that Albert Commons did not always label his
specimens until later and that he often confused his collections. No one has offered proof,
however, that Albert Commons actually did this. Bayard Long was only 16 at the time of
Commons’ last collection from Delmarva in 1901. We must also remember that R.
knieskernii is found in early successional wetlands within oak/pitch pine forest stands.
Even in New Jersey, only 27 out of 38 sites for this species still exist (Anonymous, 1991).

THE CASE OF EUPATORIUM RESINOSUM

On August 5, 1874, Albert Commons collected Eupatorium resinosum from “wet places,
near Gumboro” (PH542918). On September 10, 1875, Commons collected E. resinosum
from “low pine barrens (Baltimore Hundred), Sussex Co.” (PH542917). Note also that
Commons misidentified a collection of E. hyssopifolium as E. resinosum collected in sandy
woods of Rehoboth on August 1, 1895 (PH542901). The 1874 and the 1875 specimens of
this composite were collected on the same dates as Rhynchospora knieskernii. In his report
on Eupatorium resinosum Torrey ex D.C. for the US Fish and Wildlife Service, Mowbray
(1986) reported this species as once found in Delaware but now extirpated. A similar
report was made by Rawinski and Cassin (1986).

In our study of the floristic geography of Delmarva, we have come to realize that rarity is
often a natural and successful “strategy” for survival. We have also come to appreciate the
Delmarva Peninsula as a significant link in the Long Island-New Jersey-Delmarva-Carolinas-
Georgia chain of pinelands, wetlands, and coastal habitats.

We rediscovered Oxypolis canbyi in 1980 in South Carolina, and established its
taxonomic status as a distinct species (Tucker et al. 1983). We later rejoiced at the
discovery of O. canbyi in Maryland (Boone, Fenwick, and Hirst, 1984). We have explored
R. knieskernii sites in New Jersey and E. resinosum sites in New Jersey, South Carolina, and

COLLECTIONS OF ALBERT COMMONS i

Kentucky (Tucker and Dill, 1989b). We have reconstructed, at least in part, Rafinesque’s
lost Florula Delawarica (Tucker and Dill 1989a), and we have no problem accepting his
collection of Drosera filiformis in Sussex Co., Delaware, especially since Clyde Reed has
re-collected this species (personal communication, 1985). Aeschynomene species, sedges,
and many other rare plants also share this distribution on Delmarva (Broome et al, 1979;
Carulli, Tucker, and Dill 1988; Naczi et al. 1986; Tucker et al. 1979).

We therefore have no problem with the collections of Albert Commons. In fact, we
noted (Dill and Tucker 1986) that “... contrary to the classic philosophical problem of
science being able to establish the negative but not the positive, the biogeographer
establishes concretely the presence of a taxon through voucher collections but is never able
(over large areas) to state that a taxon is not present. ‘Extirpated’ taxa have an uncanny
(and embarassing) way of reappearing.”

RETRACING THE STEPS OF ALBERT COMMONS

In order to assimilate the flora of Delmarva for his book, Tatnall (1946) compiled a card
file of all Delmarva herbarium records of the Academy of Natural Sciences, his herbarium,
and the herbarium of the Society of Natural History of Delaware (the latter two now at the
Claude E. Phillips Herbarium). Manual examination of this card file revealed many other
species that had been collected on the same dates as the Rhynchospora knieskernii and
Eupatorium resinosum. Robert Tatnall did not have access to the herbarium of Edward
Tatnall, which had been sent out to Colorado College in 1902 (Anonymous, 1902), but
these specimens were returned on permanent loan to the Claude E. Phillips Herbarium in
1980. We also acquired the Delmarva specimens of the Langlois Herbarium at Catholic
University upon its dissolution in 1986 (Tucker, Poston and IItis, 1989). In addition, we
have examined specimens of Commons at the New York Botanical Garden and the
Smithsonian Institution.

The vascular plant collections of Commons, totalling at least 4262 specimens, were
collated for computer analysis. These records were then sorted by date, species, and
location. In addition, we are fortunate to have Commons’ notebook, which is actually an
annotated copy of Edward Tatnall’s 1860 Catalogue.

Our first record of a Commons herbarium specimen from Delmarva is Tephrosia
virginiana, collected near Centreville. Commons made this collection in 1861 at the age of
32. Many of his collections were near the family farm in Centreville, but many excursions
were made as far south as Salisbury, MD. The last recorded collections were made on
September 9, 1901, when Commons was 72, from the Stanton-Ogletown area of Delaware.
Our computerized records comprise a nearly complete survey of Albert Commons’
activities in Delmarva over a 40-year period. This resource makes it possible for us to
compare and contrast Commons’ botanical travels with his collections of R. knieskernii and
E. resinosum.

On August 3, 1874, Albert Commons collected Potamogeton nodosus from the Red Clay
Creek at Greenbank (probably while waiting for the train).

On Tuesday, August 4, he collected the following specimens (names have been
modernized to Kartesz and Kartesz (1980) but not all sheets have been annotated
recently): Polypodium polypodioides, near Gumboro, Cypress Swamp; Chasmanthium
laxum (Uniola laxa), Gumboro, sand; Eleocharis robbinsii, Laurel, bank of Broad Creek;
Paronychia fastigiata, Laurel, Peppers Mill; Centrosema virginianum, near Laurel; Clitoria
mariana, near Gumboro; Phyllanthus caroliniensis, near Laurel, sandy soil, road to Peppers
Mill; Toxicodendron toxicarium (Rhus quercifolia), Laurel, pine woods south; Vitis aestivalis,

12 BARTONIA

near Gumboro, Cypress Swamp; Vitis rotundifolia, Gumboro; Oenothera fruticosa, near
Gumboro; Nyssa sylvatica, near Gumboro; Monotropa uniflora, Gumboro; Symplocos
tinctoria, near Gumboro; Cypress Swamp; Polypremum procumbens, Gumboro; Verbena
officinalis, Gumboro, sandy soil; Pycnanthemum setosum, Laurel, dry pine woods; Urtricu-
laria juncea, Laurel; Lobelia nuttallii, near Gumboro, sand; and Senecio tomentosus,
Gumboro.

On Wednesday, August 5, 1874, he collected the following specimens: Potamogeton
epihydrus, Laurel, Broad Creek at wharf; Najas gracillima, Laurel, Broad Creek; Cyperus
compressus, Laurel, sand; Cyperus dentatus, sandy swamps, Sussex Co.; Eleocharis equisetoi-
des, ponds between Gumboro & Laurel; Eleocharis melanocarpa, between Gumboro &
Laurel, wet sand; Rhynchospora gracilenta, near Laurel; Rhynchospora knieskernii, near
Gumboro, swamp; Rhynchospora torreyana, near Laurel; along road to Gumboro; Scirpus
etuberculatus, ponds, Sussex Co.; Smilax pseudo-china, Laurel; Xerophyllum asphodeloides,
near Laurel, dry pine woods, rare; Lophiola aurea, pine barren bogs between Gumboro &
Laurel; Tipularia unifolia near Laurel; Myrica cerifera Laurel; Myrica pensylvanica Laurel,
Quercus ilicifolia near Laurel; Quercus marilandica near Laurel; Paronychia fastigiata
Gumboro, dry woods; Minuartia caroliniana (Arenaria caroliniana) near Little Hill, dry
sand; Desmodium marilandicum Laurel; Rhynchosia tomentosa Pepperbox, 74 mi NE, sandy
pine barrens, Little Hill Church; Hypericum denticulatum vat. denticulatum (H. denticula-
tum var. ovalifolium), near Laurel, pine barrens; Viola pedata, Laurel, pine barrens;
Chimaphila maculata, Laurel; Vaccinium stamineum, Gumboro, woods; Eupatorium
resinosum, near Gumboro, wet places; and Pityopsis graminifolia (Chrysopsis graminifolia),
Laurel, pine barrens SE. By August 7, he returned to Centreville, where he collected Vitis
vulpina, Physalis alkekengii, and Lobelia cardinalis.

On September 3, 1875, Albert Commons collected Impatiens capensis, Solanum nigrum,
and Scrophularia marilandica at Mt. Cuba and near Centreville. On the morning of
Thursday, September 9, he collected Leersia virginica from Centreville, and, on continuing
his travels south that day, he collected the following specimens: Lycopodium appressum,
Cedar Neck, sandy bogs; Scleria reticularis (S. muhlenbergii), Felton; Ilex glabra, Frankford;
Eryngium aquaticum, Felton, sandy ditches; Bartonia virginica, near Frankford, Cypress
swamp; and Pluchea camphorata, Gumboro, cypress swamp E. He probably arrived in
Sussex Co. late in the day on September 9, so his list for that day is quite naturally short.

On Friday, September 10, he collected the following specimens: Ruppia maritima, Cedat
Neck; Triglochin striata, salt marshes below mouth Indian River; Eragrostis refracta, Cedar
Neck, moist sand; Aristida lanosa, Cedar Neck; Paspalum dissectum, Cedar Neck, pond 2 ft
deep; Paspalum floridanum var. glabratum, Rehoboth, $; Panicum amarum, Cedar Neck;
Dichanthelium aciculare (Panicum angustifolium), Frankford; Cenchrus tribuloides, Cedar
Neck, seashore; Schizachyrium scoparium var. scoparium (Andropogon scoparius), Cedar
Neck; Schizachyrium scoparium var. littorale (Andropogon littoralis), Cedar Neck (N of
Ocean View); Cyperus tenuifolius, moist soil, Baltimore Hundred; Eleocharis tortilis (E.
simplex), Baltimore Hundred; Psilocarya scirpoides, Baltimore Hundred; Rhynchospora
capitellata, Frankford, moist soil; Rhynchospora knieskernii, Baltimore Hundred; Sc/eria
reticularis (S. muhlenbergii), Cedar Neck; Scleria pauciflora, Baltimore Hundred; Eriocau-
lon decangulare, Cedar Neck, ocean shore; Xyris difformis, Baltimore Hundred; Juncus
acuminatus, Frankford; Juncus roemerianus, salt marsh of Indian River; Lachnanthes
caroliniana (L. tinctoria), Cedar Neck; Spiranthes tuberosa, Frankford; Quercus falcata,
Frankford; Polygonum glaucum, sea shore S of Indian River; Polygonum ramosissimum, dry
sand, S of Indian River, Baltimore Hundred; Amaranthus pumilus, Baltimore Hundred,

sedis aie ee

COLLECTIONS OF ALBERT COMMONS 13

sea beach; Atriplex arenaria, sandy seashore, Baltimore Hundred; Salicornia virginica (S.
bigelovii), Cedar Neck, salt marsh; Salicornia europaea, Cedar Neck, salt marsh; Suaeda
linearis, Cedar Neck, salt marsh; Desmodium laevigatum, near Frankford, woods; Kosteletz-
kya virginica, Cedar Neck, salt marsh; Hudsonia tomentosa, Cedar Neck, S of Indian River;
Viola lanceolata, Cedar Neck; Lythrum lineare, Cedar Neck; Oenothera humifusa, dry sand,
coast S of Indian River Inlet; Myriophyllum pinnatum, Cedar Neck, ponds; Eryngium
aquaticum, Cedar Neck, wet places; Centella erecta, Cedar Neck, S of Indian River;
Chimaphila umbellata, Frankford; Polypremum procumbens, near Frankford; Sabatia
stellaris, Cedar Neck near Ocean View; Centaurium pulchellum, Cedar Neck N of Ocean
View; Gentiana autumnalis (G. porphyrio), Sussex Co. pine barrens; Asclepias lanceolata,
Cedar Neck N of Ocean View; Gratiola pilosa, Frankford, sandy soil; Agalinis maritima
(Gerardia maritima), below Indian River Inlet, salt marsh; Utricularis inflata, Cedar Neck,
Baltimore Hundred; Galium pilosum, Frankford, sandy soil; Lobelia puberula, Cedar Neck,
N of Ocean View; Eupatorium resinosum, Baltimore Hundred, low pine barrens; Eupato-
rium serotinum, Baltimore Hundred; Pityopsis graminifolia (Chrysopsis graminifolia), sandy
pine barrens, Sussex Co.; Solidago erecta, Baltimore Hundred; Euthamia galetorum
(Solidago tenuifolia), Cedar Neck, salt marsh; Aster concolor Baltimore Hundred, dry sand;
Aster nemoralis Frankford, swamp E; Aster novi-belgii Frankford; Aster tenuifolius Baltimore
Hundred; Pluchea foetida Cedar Neck, swamps; and Prenanthes autumnalis Ocean View,
N, Baltimore Hundred. By September 15, he had returned to New Castle Co., where he
collected Heliopsis helianthoides from Mt. Cuba.

Students of Delmarva plant geography may wish to note that there is a Salisbury in Kent
Co., Delaware and a Centerville in Sussex Co., Delaware (Heck et al., 1966). According to
Robert Tatnall’s card file, there are also two Cedar Necks; one between Cedar Creek and
Mispillion R., the other south of Indian River Bay, between White Creek and the ocean.
All but one of Commons’ ‘Cedar Neck’ records (18 in number) are under date of 10 Sept
1875 as are also his ‘Baltimore Hundred’ or ‘seashore S. of Indian R.’ (10 in number). The
one exception is Spergularia marina (A.C. ‘Cedar Neck’ 16 Jy 1896), but Heliotropium
(Slaughter Beach 16 Jy 96) shows that Commons was in upper Sussex Co. at that time. It
seems clear that all Cedar Neck records of 10 Sept ’75 refer to the Indian River locality.

We have assembled the location information from these specimens and correlated them
with Beers (1868) and Delaware place names (Heck et al. 1966). Two putative routes are
revealed on today’s highway maps. The trip August 4-5, 1874 was apparently east from
Laurel on Rt. 24, south on Rd. 422A, east on Rd. 422, east on Rd. 427, south on Rt. 26 to
Gumboro, and then east on Rt. 54 to the Cypress Swamp. The trip on September 9-10,
1875 was apparently east from Gumboro on Rt. 54, north through the Cypress Swamp on
the “South West Bridge” (no longer extant), north on Rd. 406, south on Rd. 405 to
Frankford, south on Rd. 376, north on Rd. 375 to Omar, east on Rd. 54 to Clarksville, east
on Rt. 26, north on Rd. 357 through Cedar Neck, east on Rd. 360, and then north on Rt. 1
to Rehoboth. .

We can also postulate potential habitats along these routes. Many of the species,
particularly R. knieskernii and E. resinosum, could have been collected from wet meadows
on the edges of forests and in forest clearings, such as those which persisted until recently
east of Ellendale, DE. Atlantic white cedar swamps in this area are particularly important
for rare species on the Delmarva Peninsula (Dill et al. 1987). We note that the current site
of Ryan’s blueberry farm, just north of the Cypress Swamp, is dominated by Panicum
hemitomon, a species which we have only otherwise found correlated with Delmarva Bay
or bay-like habitats.

14 BARTONIA

Much environmental change has occurred in the Cypress Swamp since Commons’ trips
in 1874 and 1875. The standing cypress trees were extensively harvested for shingles. After
the Civil War, the buried logs were mined. Then, in 1930, a devastating fire hit the Cypress
Swamp; it burned for eight months. After the fire, attempts were made to drain the swamp
and farm it. In 1961, Edmund H. Harvey formed Delaware Wild Lands, Inc. and
subsequently purchased 11,000 acres of the Cypress Swamp.

SUMMARY AND CONCLUSIONS

Our research supports the hypothesis that Albert Commons collected Rhynchospora
knieskernii and Eupatorium resinosum in Delaware in 1874 and 1875:

(1) Ninteenth century field botanists travelled by foot, horse and buggy, boat, and/or
train, all of which facilitated close botanical observation and collection. The roads of this
era were dirt with few highway markers. Consequently, labels of this era were also vague.

(2) Information from specimen labels and notes conclusively place Albert Commons in
Delaware on August 5, 1874 and September 10, 1875. At this time we find no evidence that
he was in New Jersey on either of these dates. Polypodium polypodioides, Symplocos
tinctoria, Rhynchosia tomentosa, Pityopsis graminifolia, Triglochin striata, Eragrostis refracta,
Scirpus etuberculatus, and Juncus roemeriianus were all collected by Commons on August
4-5, 1874 and September 9-10, 1875. These species reach their northern limit in Delaware
and have never been collected from New Jersey.

(3) Many of the sites of Albert Commons’ botanizing have succumbed to residential,
industrial, recreational, and agricultural development, especially in the vicinity of the
Great Cypress Swamp in Sussex Co. Habitats, communities, and ecosystems may be
extirpated as well as local populations of species. The failure to find R. knieskernii and E.
resinosum today in Delaware does not mean that Albert Commons mislabelled his
collections, either intentionally or accidentally.

(4) The 4262 Delmarva specimens which Albert Commons collected from 1861 to 1901
represent a significant portion (estimated 78%) of the 2259 vascular taxa listed by Tatnall
(1946). Albert Commons’ collections, often of overlooked taxa such as grasses and sedges,
represent an historical treasure painstakingly assembled over a lifetime. His work firmly
establishes him, along with Edward Tatnall and William M. Canby, as a member of the
trinity of founding botanists of the Delmarva Peninsula.

ACKNOWLEDGMENTS

We thank Mr. Robert Naczi and Mrs. Nancy Seyfried for staunchly defending the honor
of Albert Commons on numerous occasions. We also thank Mrs. Nancy Seyfried and Mr.
Joseph M. McLaughlin for checking and summarizing these records and Ms. Dana
Degano for her expertise on FileMaker® Plus. This is contribution No. 14 from the Claude
E. Phillips Herbarium.

LITERATURE CITED

ANONYMOUS. 1902. Gift of Tatnall eon sca ——— sense: 7 692.

—.. Knieskern’s beaked-r d Species Tech. Bull. 15(9): 4.
——. 1991. Knieskern’s beaked-rush (Rhynchospora knieskernii). Endangered Species Tech. Bull. 16(7-8): 11.
ASHE, * on 1898. The dichotomous group of Panicum in the eastern United States. J. Elisha Mitchell Sci. Soc.

BEERS, D. G. 1868. Atlas of the state of Delaware. Pomeroy and Beers, Philadelphia.
Boone, D. D., G. H. FENWICK, AND F. Hirst. 1984. The rediscovery of Oxypolis canbyi on the Delmarva
Peninsula. Bartonia 50: 21-22.

COLLECTIONS OF ALBERT COMMONS 15

BRITTON, N. L., AND A. BROWN. 1913. An pea flora of the northern United States, Canada and the British
Possessions. Charles Scribner’s Sons, New

BROOME, C. R., J. L. REVEAL, A. O. TUCKER, pe ‘HL sabi 1979, Rare and endangered vascular plant species
in Maryland. US Fish and Wildlife Serv., Newton Corner, M

CARULLI, J. P., A. O. TUCKER, AND N. H. DILL. 1988. Mee rudis Benth. (Fabaceae) in the United States.

DENNIS, J. V. spe The t Louisiana State Univ. Press, Baton Rouge.
DILL, N. H., AND A. O. TUCKER. 1986. Characteristics and conservation of rare plants in the Delaware and
Maryland portion of the Del-Mar-Va Peninsula. Pages 113-155 in S. V. S. Rana, ed. Recent trends in
biotechnology and biosciences. Soc. of biosciences, Muzaffarnagar, India.
. SEYFRIED, AND R. F. C, Naczi. 1987. Atlantic white cedar on the Delmarva Peninsula. Pages

41-55 i in 4 D. Laderman, ed. Atlantic white cedar wetlands. Westview Press, Boulder

GALE, S. 1944. Rhynchospora, section Spy sini in Canada, the United States eae the West Indies.
Rhodora 46: 89-134, 159-197, 207-249, 2 78.

Gray, A. 1867. Manual of the botany of the sa United States. Fifth ed. Ivison, Blakeman, Taylor, & Co.,
New York.

———. 1908. Gray’s new manual of botany: A handbook of the fl ing plants and f Seventh ed. American
Book Co., New York.

HARSHBERGER, J. W. 1899. The botanists of Philadelphia and their work. T. C. Davis and Sons, Philadelphia.

CK, L. W., A. J. WRAIGHT, D. J. ORTH, J. R. CARTER, L. G. VAN WINKLE, AND J. HAZEN. 1966. Delaware place

names. US Geol. Survey Bull. 1245.

Hirst, F. 1990. habe new taxa for the Delmarva Peninsula. Bartonia 56: 70-71.

KARTESZ, J. T., AND R. KARTESZ. 1980. A synonymized checklist of the vascular flora of the United States,

ill.

LANJouw, J., AND F. A. STAFLEU. 1954, Index herbariorum. Part II. Collectors. International Bureau of Plant
Taxonomic aah sieaeargh Utrecht.

Mowsray, T. B. 1986. Final status report: Eupatorium resinosum Torrey ex D.C. US Fish and Wildlife Service,
Newton Corner.

Naczl, R. ° = Rod: — E. B PENNELL, Fg E. SEYFRIED, A. O. TUCKER, AND N. H. DILL. 1986. New
record Bartonia 52: 49-57.

PENNELL, F. W. 1943. Botani f the Philadelphia area (concluded). Bartonia 22: 10-31.

RAWINSKI, T., AND J. CASSIN. 1986. Final status survey reports for 32 plants. Eastern Heritage Task Force, The
Nature Comsrne Boston

TATNALL, E. 1860. Catalogue of the phaenogamous and filicoid plants of Newcastle County, Delaware. J. T.
Heald, Wilmington

TATNALL, R. R. 1946. Blora of Delaware and the Eastern Shore. Society of Natural History of Delaware,

Trew, L. D., AND K. CLANCY. 1991. Survey and protection of federally listed and candidate plant species in
Delaware: 1989-1990 project report, Federal Aid Project Agreement E-2-5, Grant No. 5131. Delaware
Natural Heritage Inventory, Department of Natural Resources and Environmental Control, Dover.

TuckER, A. O., AND N. H. DILL. 1989a. Rafinesque’s Florula Delawarica. Bartonia 55: 4-14

, AN 1989b. Nomenclature and distribution of Eupatorium x truncatum, with comments on the
status of E. resinosum var. kentuckiense (Asteraceae). Castanea 54: 4

2 E, C. E. PHILLIPS, AND M. J. MACIARELLO. 1979. Rare and endangered vascular plant
species in arate ae Fish and Wildlife Service, Newton Comer
ees silacepcp AND R. D. KRAL. 1983. }
byi and O. filiformis (Apiaceae). Syst. Bot. 8: 209-304.
——,.M. E. Poston, AND H. ILtis. 1989. History of the LCU eg ca 1895-1986. Taxon 38: 196-203.
Woon, A. 1861. Class-book of botany. A. S. Barnes & Burr, New Yor

distribution, ct bers, and

Bartonia No. 57, Supplement: 16-41, 1993

Rare Vascular Plants Associated with Limestone
in Southwestern Franklin County, Pennsylvania

LARRY H. KLotTz
Department of Biology, Shippensburg University, Shippensburg, PA 17257

JEFFREY L. WALCK'
Morris Arboretum of the University of Pennsylvania, 9414 Meadowbrook Avenue, Philadelphia, PA 19118
and Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103

We have observed 16 species of “Plants of Special Concern in Pennsylvania” (PNDI
1991) in the limestone lands of southwestern Franklin County, a region which adjoins
Maryland along Pennsylvania’s southern border. Several of these species have robust
populations, and four of them are not known to occur elsewhere in Pennsylvania. Some of
the natural communities are of such high floristic quality that they would be noteworthy
even if they lacked rare species.

e summarize the 16 rare species and their populations as a group; describe these
species and their communities individually; and list additional rare species that have been
collected historically in southwestern Franklin County, probably on limestone. We report
our field observations for the 16 species and review the literature on their range, habitat,
rarity, morphological variation, and breeding system. For the communities, we list the
most constant species that we have observed in Franklin County and correlate the most
recent community classifications available to us for Pennsylvania and surrounding states.
Also, we note the species of Franklin County that are included in two recent reviews on
rare vascular plants associated with limestone in Maryland (Boone 1984; Riefner and Hill

84).

We performed our field work between June 1985 and June 1991 and supplemented our
observations with the field reports available from the Pennsylvania Natural Diversity
Inventory (PNDI). The voucher specimens cited are deposited in the Shippensburg
University Herbarium unless otherwise indicated. Nomenclature follows Kartesz and
Kartesz (1980). The range and habitat of each species were summarized from all of the
pertinent literature listed in the bibliography, but only representative references are cited
in each section. The community classifications cited for Pennsylvania and surrounding
states include the systems currently employed by the respective natural heritage programs.

OVERVIEW OF RARE SPECIES AND THEIR POPULATIONS

CHARACTERISTICS OF THE RARE SPECIES. Taxonomic groups (Table 1). The 16 species are
diverse taxonomically: they are distributed among fifteen genera in as many families and
orders. The list contains one eusporangiate fern, one leptosporangiate fern, 12 dicotyle-
dons, and two monocotyledons. Five of the six subclasses of dicotyledons (Cronquist 1988)
are represented; only the Hamamelidae are lacking. The monocotyledons belong to two of
the five recognized subclasses (Cronquist 1988): the Commelinidae and the Liliidae,
respectively.

' Present address: School of Biological Sciences, University of Kentucky, Lexington, KY 40506.
16

LIMESTONE PLANTS 17

Habit (Table 1). Only three of the species are woody. The 13 herbaceous species include
one annual and 12 perennials, one of which is a vine.

Global rarity (Table 1). Eleven of the species are ranked G5, i.e. “demonstrably secure
globally,” according to the classification employed by The Nature Conservancy. Two have
the rank of G4 (‘apparently secure globally”) while two others have an intermediate
designation, G4GS. Only one of them, Dodecatheon amethystinum, has the rank of
“G3” species have only 21 to 100 occurrences and are either very rare throughout their
range, local in a restricted range, or vulnerable to extinction throughout their range (The
Nature Conservancy 1982; PNDI 1991).

Rarity in Pennsylvania and surrounding states (Table 1). Four of the species are
“endangered” in Pennsylvania, i.e. in danger of extinction within the state if they are
exploited or their habitats are not maintained. One of them is “threatened,” which means
that the species could become endangered in the state if exploited or deprived of critical
habitat. Five of them are “rare,” which means uncommon within the state. At present, six
of them are classified as “undetermined” because more data are needed to determine
their degree of rarity (Pa. Dept. Environmental Resources 1987; PNDI 1991). Of the seven
surrounding states and one Canadian province, Maryland is most similar to Pennsylvania
in the rarity of these 16 species: 13 of a are found in Maryland, and ten are of “special
concern” (MNHP 1991; Bartgis pers. comm.; also Argus et al. 1982-1987; Delaware
Natural Heritage Inventory 1991; Mitchel 1 1986; New Jersey Natural Heritage Program

TABLE 1. Characteristics of the Rare Species.

North-
State rarity‘ eastern Calcar-
Global Limit of — eous
Species Family Habit* rarity’ NY NJ DE PA MD VA WV OH ON Range® Affinity‘
Asplenium resiliens Aspleniaceae ph GS ami alee Sar: detainee < F* °
Bouteloua i
curtipendula Poaceae ph GS A See Bis. Se US tS ne* i
Chrysogonum
virginianum Asteraceae ph GS ¢+) Bk Be SRS? Re F g
Corallorhiza
isteri Orchidaceae ph GS xX ie 38, ocr Ee g
Dodecatheon
eth Primulaceae ph G3? ly r i
Hydrophyllum
macrophyllum Hydrophyllaceae ph GS Se oe hia Fr i
Matelea obliqua Asclepiadaceae phy G4? aoe a od x °
Ophioglossum ;
engelmannii Ophioglossaceae ph GS ER | ea. S F o
Opuntia humifusa __ Cactaceae ses GS ope ee Re ae . ne : g
Ptelea trifoliata Rutaceae ds/t GS §$5=285004) Rss + + + S ne i
Ranunculus
micranthus Ranunculaceae ph Gs?” Ss ee Seale eR ne i
amnus lanceolata pnt naceae ds G4G5 U ar ee i 0
Ruellia humilis canthaceae ph Baas cet eee eo F i
Ruellia strepens Acantacee ph G4G5 > 4 Uae. fom een ee P i
Salvia reflexa ae ah GS U (+) at Some May 8 UME g
Sedum telephioides dain ph Gt <(+) x i ea as ee Gs Mnf g

“ah: annual herb, ph: perennial herb (non-vine), b ine, d b, ds/t: decid hrut
— tree, see: succulent evergreen swede "see ‘text for —— ‘in PA: capone? FB: thre tened, R: rare, U:
ee aca (i.e. rare, saieenat or endangered—

jorth America, bet. piaenree

including | Diack List” i. MD), Me extirpated, +: native sad pet rare, (+): native to eastern

locally, ¢F wl di
“we a2 : Penne, fies t
south and west beyond eastern North America. © ig phile, g: edaphic g i: intermediate

18 BARTONIA

1990; Ohio Division of Natural Areas and Preserves 1990; Virginia Natural Heritage
Program 1990; West Virginia Natural Heritage Program 1990; Zika 1990; plus sources
cited in the next section on community types).

Northeastern limit of range (Table 1). According to the literature reviewed below in the
section on community types, most of the 16 species are native to the states neighboring
Pennsylvania to the south and west (Maryland, Virginia, West Virginia, and Ohio). Only a
few of them (i.e. one to four) are native and currently extant in the states to the north and
east (New York, New Jersey, Delaware) and in Ontario. Five of the species reach their
northeastern limit in southwestern Franklin County, and three are restricted to that region
within Pennsylvania (Wherry et al. 1979): Ophioglossum engelmannii, Asplenium resiliens,
Ruellia humilis, Hydrophyllum macrophyllum, and Chrysogonum virginianum. Seven other
species reach their northern or northeastern limit in Pennsylvania beyond southwestern
Franklin County (Wherry et al. 1979): Dodecatheon amethystinum, Sedum telephioides,
Rhamnus lanceolata, Matelea obliqua, Salvia reflexa, Ruellia strepens, and Corallorhiza
wisteriana. The remaining four species range north of Pennsylvania into southern New
England, New York, and/or southern Canada.

None of the 16 species have distributions centered on the Coastal Plain, and none are
restricted to the Appalachians. All range widely to the south and west in eastern North
America; and six extend beyond this region into western North America and Mexico:
Ophioglossum engelmannii, Asplenium resiliens, Ptelea trifoliata, Salvia reflexa, Bouteloua
curtipendula, and Corallorhiza wisteriana. At least two species also extend to Argentina:
Asplenium resiliens (Mickel and Beitel 1988) and Bouteloua curtipendula (Hitchcock 1971).

Wherry et al. (1979) recognized four groups of taxa that “reach a geographic limit in
Pennsylvania or extend only sparingly beyond its borders.’”’ Five of the 16 species are
among Wherry’s “Midland taxa,” which are “widespread in the Mississippi and tributary
valleys, ranging eastward in habitats where the soil-reaction tends to be circum-neutral
owing to a base-rich underlying rock . . .”: Asplenium resiliens, Dodecatheon amethystinum,
Rhamnus lanceolata, Matelea obliqua, and Bouteloua curtipendula. The “Appalachian
taxa,”’ which are “widespread in the southern Appalachians from northern Alabama .. .
[to] Maryland,” include “O. calcicola” (Opuntia humifusa in part), Sedum telephioides, and
Chrysogonum virginianum. Only “O. compressa” (Opuntia humifusa in part) and Corallo-
rhiza wisteriana are among the “Lowland taxa,’ which are “widespread ... on the
Atlantic—-Gulf Coastal Plain . . . [and] in the Great Lakes lowland.” None of the 16 species
are “Northern taxa.”

Keener and Park (1986) elucidated the distributional patterns of plant species within
Pennsylvania. They listed Asplenium resiliens, Sedum telephioides, Ruellia humilis, and
Chrysogonum virginianum among the species that are restricted to the southern part of the
Valley and Ridge Province in Pennsylvania and reach their northern limits there. These
authors included Ranunculus micranthus among the species that range across the southern
part of the state.

Calcareous affinity (Table I). Species of plants that occur over limestone differ in their
degree of fidelity to it. The literature reviewed below implies that the 16 species compose a
spectrum in this feature. At one extreme are the obligate calciphiles such as Ophioglossum
engelmannii, Asplenium resiliens, Rhamnus lanceolata, and Matelea obliqua. They may be
edaphically specialized, for they occur exclusively on substrates that are high in calcium,
namely calcitic or dolomitic limestone, or soils derived from these parent materials
(Riefner and Hill 1984). At the opposite extreme are edaphic generalists, which occur ona
broad variety of substrates, calcareous and non-calcareous, throughout their ranges.

LIMESTONE PLANTS 19

Opuntia humifusa and Sedum telephioides are obvious examples; but Salvia reflexa,
Chrysogonum virginianum, and Corallorhiza wisteriana are also in or near this group. The
seven remaining species are intermediate, occurring on limestone but also other sub-
strates, especially those of basic or ci tral pH such as diabase, serpentine, calcareous
shale or sandstone, and alluvial soils derived from non-acidic parent materials (Riefner
and Hill 1984). Listed in an approximate, descending order of fidelity to limestone, the
“intermediate” species are: Dodecatheon amethystinum, Ruellia humilis, R. strepens, Ptelea
trifoliata, Hydrophyllum macrophyllum, Ranunculus micranthus, and Bouteloua curtipendula.

CHARACTERISTICS OF THE POPULATIONS (TABLE 2). The entries after each species in
Table 2 represent the known populations on limestone in southwestern Franklin County.
Ten of the species have only one or two populations, five species have three or four
populations, and one species has eight populations.

Physiographic sections (Table 2). In southwestern Franklin County, as in Pennsylvania as
a whole, limestone bedrock underlies extensive portions of the Great Valley Section and
the Appalachian Mountain Section, which are the two subdivisions of the Valley and
Ridge Province (Anon. 1990; Berg et al. 1989; Long 1975; O’Neill 1964). The 16 species are
about equally divided between the two physiographic sections. Six of them are only known
from the Great Valley Section, five others have been found only in the Appalachian
Mountain Section, and the remaining five occur in both regions.

Of the 40 populations of the rare species, 19 are in the Great Valley Section and 21 are
in the Appalachian Mountain Section. Most of the Great Valley populations are in two
clusters. Eight populations occur just west of the Conococheague Creek while seven others
are centered around its main tributary, the West Branch. The Appalachian Mountain
populations are concentrated around Licking Creek and one of its tributaries, Little Cove
Creek. Three of the populations along Licking Creek are actually in Maryland, within a
few meters of Pennsylvania. Both Licking Creek and the Conococheague Creek are
tributaries of the Potomac River.

Geologic formations (Table 2). The rare species occur on nine geologic formations
representing three successive geologic periods of the Paleozoic era (Berg et al. 1980; Berg
and Dodge 1981; Clark 1970; Geyer and Wilshusen 1982; Vokes 1957). The standardized
abbreviations of the formations are given in parentheses:

Devonian: Onondaga and Old Port Formations, undivided (Doo);

Devonian and Silurian: Keyser and Tonoloway Formations, undivided (Dskt);

Silurian: Bloomsburg and Mifflintown Formations, undivided (Sbm); Clinton Group
(Sc); Wills Creek Formation (Swe);

Ordovician: Chambersburg Formation (Oc), Rockdale Run Formation (Orr), St. Paul
Group (Osp), Stonehenge Formation (Os).

All of the species with more than one population occur on more than one of the above
formations. Richest in rare plants are the Onondaga and Old Port Formations, undivided,
with eight populations representing eight species; and the St. Paul Group (plus the
adjoining Chambersburg Formation), with 14 populations among eight species. The
remaining six formations each support only one to six populations. ; i
In some locations, the limestone bedrock of southwestern Franklin County exhibits
characteristics of karst topography, including outcrops, caves, and sinkholes (Davies and
Legrand 1972). Southwestern Franklin County also contains shale and sandstone (Berg
and Dodge 1981; Long 1975). On Ordovician shale of this region, the junior author has
discovered four state-rare species: a single plant of Trifolium virginicum (Walck 474,

20 BARTONIA

TABLE 2. Characteristics of the Populations.

Physio- Slope Population
icra Geol. Elev. Soil Map Comm. Shad- Mois-s —————
Species Section? Form? (m) Unit® Position? Aspect Degrees type’ _—ing® ture" Area’ Size!
Asplenium GV Orr 146 HkD i N s Cc s m B b
resiliens AM Doo 152. HkD?-Md. 1 NNW S-v c s m B b
AM DSkt 146 HkD u NE s € f m B b
Swe 140 Ph: HkD? I NNE s Cc s m B a
loua GV Oc/Osp 171 HkD m Ee g Ww ° x Cc d
curtipendula
Chrysogonum GV Os 152. HkD u NW g U s-f m D c
virginianum AM DSkt 143 HhD3 u SW m U m A a
Corallorhiza AM Doo 150 HkD 1 SE m U s-f m Cc b
wisteriana
Dodecatheon GV Os 152. HkD u NW g-m R f-o m D d
amethystinum AM Doo 247 HeB u- NW g-m U f m € e
AM DSkt 152 3 m Ww m U s-f m B c
AM Swe 143. HhD3 u-c NW-SW g U s-f m Cc c
lum AM 150°- ~ HED 1 SE m U s-f m D d
macrophyllum AM Dskt 143. RyC I Ww s U s m B a
Matelea obliqua GV Oc/Osp 174 HkD m E g Ww f m/x B a
AM Doo 140 1 SE m U s-f m B a
Swe 143 HhD3 I S m U f m B a
GV Osp 165 m-u NE g Ww te) m/x B c
ssn ene
Opuntia humifusa GV Oc/Osp 158 HkB I-c E g-v RC f-o x Cc a
Ptelea trifoliata AM Sbm 137° PR b — F f m € a
AM DSkt 137. Du-Md b _— F s m A a
Ranunculus 50 HkD i SE m U s-f m 2 b
micranthus AM Sc 168 DIF/Vd m S m U f m Cc c
AM Swe 146 HhD3 l-m S m U f m c c
Rhamnus lanceolata GV Oc/Osp 168 HkB m E g Ww f-o m/x Cc b
GV Osp 168 HgB3 f = — U s-f m/x B a
GV Oc/Osp 174 HkD m E g W f m/x A a
Ruellia humilis GV Oc/Osp 168 HkB m E g WwW f-o m/x Cc d
GV Oc/Osp 174 HkD m var. g WwW ° x D d
Ruellia strepens GV Orr 146 HkD l-m NE m U s-f m B a
GV Os 146 HkB i SW g F f-o m A b
GV Osp 149 Ph b — — F f m B a
GV 152. HkB m m U s-f m B a
GV Osp 1 HgB3 f _ U f-o m/x Cc b
GV Oc/Osp 17 HkD c — U s m B a
AM 137. Ph, HkD b-1 SE g-m F,U s-f m B b
AM Dskt 3 | S g Ss ° m B b
Salvia reflexa GV Oc/Osp 162 HkB m ENE g Ww f m/x B b
ui AM Doo 168 HkD?-Md. l-u N S-v Cc f-o m B c
telephioides AM Swe 143. Ph: HkD? I-u N-NW s ec f-o m/x B b
*GV: Great ag AM: ICAO wnmne ®Slash (/): locati in relati °DIF: Dekalb and Lehew extremely
stony soils, 25-75% (in MD- ng 1975); HeB: Hagerstown silt loam, +-8% 4 lopes; ae
pepe rocky phe er “is 3-8% sake pit ds) HhD3: Hagerstown silty clay, oe slope:
—— > HKD: Hage agerstown- “Rock E onberep ccpaples, eo slopes (HkD?-Md: in Mix Icareous rock o1 ~ p); Ph: Philo silt loam (Ph:
% slopes; Vd: Very stony lan ekalb soil material. 4c: crest, u: upper slope, m: mid
soe 1: lower slope é : hota, f: level upland; °g: gradual ies degrees), m: : moder = (1045), Ss: £ sto P (45-90), v: vertical, —: level. fCalcareous
community types—U: upland forest, R: summit, C: cliff, F: floodplain fo: is mociiiciadd field. £s: shade, f: filtered, 0:
mesic, x: xeric, m/x: feliisiaaaie 'A: <1 m’, B: >1 m? - 100m ey C: > 100 m? — I ha, D: "> 1 ba; Jaz 1-10, b: 1 11-100, c: 101-1000, d:

open. ra
1001-10,000, e: > 10,000.

MOAR), plus populations of Senecio antennariifolius (Walck 1987), Ranunculus micran-
thus, Opuntia humifusa, and Ruellia strepens. The first two are considered to be endemics of
the onan pase shale barrens (Keener 1983).

Elevation (Table 2). The elevation in Franklin County ranges from 130 to 750 meters
(U.S. Geological Survey 1979). The 16 rare species occur below 250 meters. Table 2 gives
the estimated midpoint of the elevational ranges of the respective populations.

Soil mapping units (Table 2). The soils supporting the rare species lie chiefly within five

LIMESTONE PLANTS 21

mapping units of the Hagerstown Series, which includes well-drained, upland soils
“formed in material weathered from relatively pure limestone” (Long 1975). Fifteen of the
16 species occur wholly or partly within Hagerstown soil mapping units. Thirty of the 40
populations are wholly within Hagerstown mapping units, as are two populations on
calcareous cliffs in Maryland. One other population occurs partly on Hagerstown soils and
partly on Philo silt loam, a floodplain soil formed in alluvium “from uplands underlain by
sandstone and shale” (Long 1975); and two populations are wholly on Philo silt loam.
(Two other populations mapped as “Philo” are actually on cliffs that would correspond to
““Hagerstown-Rock outcrop complex.”) The three remaining populations are within
additional mapping units: Dunning silty clay loam, a floodplain soil formed in alluvium
“from uplands underlain by calcareous material”; Ryder silt loam, which is formed from
thin bedded limestone; and the Dekalb Series, which is derived from “acid sandstone and
some shale” (Long 1975). The bedrock of this last population is the Clinton Group (Berg
and Dodge 1981), which includes some limestone (Geyer and Wilshusen 1982).

Slope (Table 2). All of the species occur wholly or largely on slopes except Ptelea
trifoliata, which was seen only on bottomlands. Thirty-four of the 40 populations occur on
slopes, most commonly mid or lower slope. Only six of them occur entirely on crests,
bottomlands, or nearly level uplands. The aspect of the slope differs among the
populations. The most frequent aspects are east and southeast, with seven and five
populations, respectively. The remaining aspects are represented by one to four popula-
tions each.

Community types (Table 2). The limestone regions of southwestern Franklin County
support a spectrum of community types, which differ in topographic location, vegetational
physiognomy, and taxonomic composition. The communities are not always distinct and
extensive: they intergrade or interdigitate finely at some sites. This study deals with five
community types on limestone (adapted from PNDI 1983): upland calcareous forest,
calcareous cliff community, calcareous rocky summit community, floodplain forest, and
successional red cedar woodland.’ In Maryland, four “habitat types” on limestone have
been recognized that retain the original character of the vegetation: (1) cliffs and ledges
that are partly shaded to fully exposed; (2) well-shaded outcrops along streams and rivers;
(3) wooded slopes, often on talus, with accumulation of humus; and (4) calcareous alluvial
soils (Riefner and Hill 1984). These habitat types also occur in southwestern Franklin
County and partly correspond to the categories employed in the present study. In Franklin
County, as in Maryland, most of the upland terrain on limestone has been cleared for
agriculture, quarrying, residences, and highways. Nevertheless, some interesting examples
of level, upland forests and successional communities are still extant on this type of land.

The populations and species of rare plants are not distributed equally among the five
community types. Eighteen of the 40 populations are entirely in upland forest, and eight of
the 16 species occur partly or wholly in this habitat type. Next richest is successional red
cedar woodland, with eight populations in five species. The remaining community types
are each represented by one to six populations and only one or two species.

Shading (Table 2). Degree of shading depends on the aspect and degree of slope as well
as the physiognomy of the vegetation. The three categories (open, filtered, and shaded)
represent qualitative visual estimates when the vegetation is in leaf. In only eight of the 16

'In addition, one population of Ruellia strepens occurs in a small, mesic “successional field” community (PNDI
1983) along a roadside. The flora is a mixture of native and naturalized grasses, forbs, and shrubs. It resembles
the forest edges that R. strepens commonly inhabits in the other communities listed.

22 BARTONIA

species are all of the populations in shaded to filtered lighting. In the other eight species, at
least some of the populations are more brightly illuminated. Twenty-eight of the 40
populations occur in shaded to filtered lighting. The other populations occur in open or
nearly open lighting.

Moisture (Table 2). The general moisture regime of the populations was inferred from
the combined factors of position, aspect, and shading. More of the populations occurred in
mesic than in xeric sites: 28 of the 40 populations (and seven of the 16 species) occurred
solely in mesic habitats (mesic upland forest, floodplain forest, shaded cliff). The
remaining 12 populations occurred in xeric or intermediate habitats (xeric upland forest,
exposed cliff, calcareous rocky summit, red cedar woodland

Population area and size (Table 2). The populations differ greatly in area and in number
of aerial shoots. Areas range from less than one square meter to four hectares. The
numbers range from a solitary shoot to more than ten thousand. Whether the shoots
represent ramets or genets was not determined, but inferences are made for some of the
species listed below. A majority of the populations are small, occupying areas of up to 100
square meters and comprising less than 100 shoots.

CALCAREOUS COMMUNITY TYPES AND THEIR RARE SPECIES

XERIC TO MESIC UPLAND FOREST. These communities commonly have a diverse and
luxuriant canopy and understory. Many of the species are local: they were observed in only
one example of this community type. Such floristic variation among the communities may
be related partly to edaphic or topographic differences. However, these communities also
exhibit a strong floristic similarity. The following species of canopy trees occur at six or
more of the nine examples that were tabulated: Juniperus virginiana, Celtis occidentalis,
Ulmus americana, Carya cordiformis, C. ovata, Juglans nigra, Quercus muhlenbergii, Q. rubra,
Ostrya virginiana, Tilia americana, Prunus avium, Cercis canadensis, Acer saccharum S.l.,
Fraxinus americana and/or F. pennsylvanica. By the same criterion, the characteristic
shrubs and vines are: Lindera benzoin, Menispermum canadense, Rosa multiflora, Toxicoden-
dron radicans, Lonicera japonica, Viburnum prunifolium, Rubus occidentalis, Parthenocissus
quinquefolia, and Smilax hispida. Ericaceae are absent or scarce on the calcareous
substrates but are frequent on the adjoining formations of shale or sandstone.

In some of these communities, the native herbaceous flora appears threatened by the
vigorous growth of several naturalized Eurasian species, especially Rosa multiflora,
Lonicera Japonica, Alliaria petiolata, and Hesperis matronalis. These alien species also
abound in floodplain forests described below. In addition to being pressured by
nica many of the native herbaceous species are easily damaged by trampling, some
of them are vulnerable to browsing by deer or other herbivores, and a few are so scarce that
they could be extirpated by collecting. In some of these communities, for example the
stand described by Klotz and Walck (1990), many of the rare or unusual native species
occur close to the road. Since the application of herbicides for roadside maintenance could
damage or eliminate these populations, infrequent mowing is preferable in such areas.

Similar communities or forest types, not necessarily on limestone, have been described
in New York (Reschke 1990): “Appalachian oak-hickory forest” and “rich mesophytic
forest”; New Jersey (Breden 1989): “dry-mesic calcareous forest” and “dry-mesic inland
mixed oak forest—mixed oak-hardwood subtype”; Pennsylvania (PNDI 1983): “‘dry-mesic
calcareous central forest” and “mesic central forest”; Maryland (Brush et al. 1980): “sugar
maple—basswood association”; Virginia (Harvill et al. 1977): “beech—maple-tuliptree
forest’; (Rawinski 1990): “‘submesotrophic to eutrophic forest’; and West Virginia (Core

LIMESTONE PLANTS 23

TABLE 3. Floristic Spectrum (%) of a Mesic Calcareous Forest Community in Southcentral Pennsylvania.

Geographic
Taxonomic Group* Habit” Origin®
F G D M WwW FP A Ne Nd
Southcentral PA 4 1 80 15 23 63 14 81 19
Northeastern US* 2 1 70 27 16.5 68 155 80 20

aE: eipragias hey G; daria ra D: Dicotyledons, M: Monocotyledons. "W: woody, P: perennial or
biennial, A: ual. ‘Ne: native, Nd: naturalized. “Source: Klotz and Walck 1990. Sources: Gleason 1952,
Gleason and Choinit 1964, ie 1950.

1966): “mixed hardwood forest: mesic.” All of the above (except Rawinski 1990) list Fagus
grandifolia as an indicator, but we did not see it in our study areas.

The sites each contain one to five of the eight rare species that we have observed in this
community type: Ranunculus micranthus, Dodecatheon amethystinum, Rhamnus lanceolata,
Matelea obliqua, Hydrophyllum macrophyllum, Ruellia strepens, Chrysogonum virginianum,
and Corallorhiza wisteriana. Thirty of the species that we have observed in this community
type are generally restricted to limestone or other basic to circumneutral soils in Maryland:
Aristolochia serpentaria, Anemone canadensis (extirpated in Maryland), Delphinium tr-
corne, Jeffersonia diphylla, Quercus muhlenbergii, Ostrya virginiana, Tilia americana, Hyban-
thus concolor, Arabis laevigata, A. patens (perhaps better included in the calcareous cliff or
rocky summit community types), Desmodium glutinosum, Zanthoxylum americanum,
Floerkea proserpinacoides, Chaerophyllum procumbens, Matelea obliqua, Hydrophyllum
macrophyllum, Phacelia purshii, Lithospermum canescens, Mertensia virginica, Ruellia
strepens, Campanula americana, Triosteum perfoliatum, Aster shortii, Chrysogonum virginia-
num, Carex hitchcockiana, C. jamesii, C. oligocarpa, C. platyphylla, Oryzopsis racemosa, and
Trillium sessile (Boone 1984; Riefner and Hill 1984).

Floristic relationships of a mesic calcareous forest. The flora of one of the mesic calcareous
forest communities has been studied in detail (Klotz and Walck 1990). The floristic list
consisted of the species occurring within the extent of the population of Hydrophyllum
macrophyllum. A total of 253 species in 69 families was found within an area of about four
hectares. The factors contributing to the richness of this flora possibly include calcareous
substrate, topographic heterogeneity, edge effect, periodic mowing along the roadside,
past removal of trees and bedrock, and invasion by introduced species. Because the flora of
this community was surveyed thoroughly, features of its floristic spectrum can be
calculated (Table 3). These data are a potential baseline for future floristic studies in this
region. This community is proportionally richer than the northeastern United States as a
whole in the percentages of dicotyledons and woody species, poorer in the percentage of
monocotyledons, and similar in the percentages of annuals and naturalized species. The 12
largest families in the northeastern United States (Fernald 1950) include the 11 largest
families in this community.” Carex is the largest genus in both floras.

This mesic calcareous forest community is floristically related to the Potomac and Ohio
drainages of Maryland, West Virginia, southwestern Pennsylvania, and southeastern Ohio

? The families with seven or more species at this site are: Asteraceae (36 species); Poaceae (16); Rosaceae and
Ranunculaceae (11); Fabaceae, Brassicaceae, and leo (10); Crpermrese as all i ‘arex spp.); Liliaceae and
Caryophyllaceae (7). Th uld large family eastern United States.

24 BARTONIA

(Klotz and Walck 1990). The affinity with the Ohio drainage can be discerned from the
analysis of the geographic relations of the flora of Pennsylvania by Wherry et al. (1979).
These authors listed 22 of the species that occur in this community. Eighteen of them are
“Midland taxa.” In contrast, only one is an “Appalachian taxon,” three are “Lowland
taxa,” and none are “Northern taxa.”* (The explanation of these categories is given above
in the section on Overview of the Rare Species and Their Populations, after the heading:
Northeastern limit of range.)

The geographic analysis of the flora of Pennsylvania by Keener and Park (1986)
elucidates the southern affinity of this community. Their lists include 76 of its species.
Within Pennsylvania, three of them are otherwise confined to the Pittsburgh Plateaus
Section (Delphinium tricorne, Aster shortii, and Trillium sessile); and one of them (Phacelia
purshii) is confined to “southwestern/southeastern” regions of the state. Fifteen of the 7
listed species range across southern Pennsylvania, 21 are generally south of the glacial
boundaries and the Allegheny High Plateaus Section, and six are confined to the provinces
southeast of the Allegheny Front (i.e. Valley and Ridge, Piedmont, Coastal Plain). In
contrast, only one of the species (Magnolia acuminata) ranges across the western part of
the state, and none of them have a northern or eastern distribution within Pennsylvania.
The remaining 29 species are among the “common native species,” which are frequent
throughout Pennsylvania.

Rare species of xeric to mesic calcareous upland forest. a. Ranunculus micranthus (Gray)
Nutt. ex Torr. & Gray, small-flowered crowfoot, ranges from Massachusetts to North
Carolina and Oklahoma and also occurs in the Black Hills of South Dakota (Benson 1948;
Gleason and Cronquist 1991; Keener 1976). In Pennsylvania, it is found in the southern
third of the state (Wherry et al. 1979).

Besides noting R. micranthus on limestone in Franklin County, we have also seen it on
soils derived from diabase in Adams County (Berg and Dodge 1981; Speir 1967) and shale
in Franklin and Fulton counties (Berg and Dodge 1981; Berg et al. 1980). In degree of
moisture, these sites range from the relatively mesic communities on limestone to the more
xeric communities on shale. Most of the sites are on slopes of various aspects and degrees;
only one is on level upland terrain. Our observations agree with published accounts on the
range of parent materials and slopes that this species inhabits (Deam 1970; Keener 1976;
Mohlenbrock 1981; Steyermark 1963). This species is known to be calciphilic in the Blue
Ridge (Wofford 1989), and it is reported from diabase and floodplain soils in northern
Virginia (Allard and Leonard 1962).

Fernald (1939) recognized two eastern varieties in this species on the basis of leaf
morphology. Pennsylvania lies within the range of var. delitescens (Fernald 1950). Some
subsequent authors have accepted this treatment (Fassett 1942; Steyermark 1963) while
others have considered R. micranthus to be a single polymorphic taxon (Benson 1948;
Keener 1976; Mohlenbrock 1981)

We have not satisfactorily distinguished R. micranthus from R. abortivus at some sites in
southwestern Franklin County. These species are best identified in late April or early May;
by early June, the aerial shoots shed their achenes, become lax, and disintegrate. Also, the

3 “Midland”: Carex amphibola, Trillium sessile, Dioscorea quaternata, Quercus muhlenbergii, Delphinium tricorne,
Jeffersonia diphylla, Magnolia acuminata, Agrimonia rostellata, Vicia caroliniana, Oxalis grandis, Rhus aromatica,
Ei rpureus, Matelea obliqua, Phlox divaricata, Phacelia purshii, abies: concinnum, Hedyotis
nuttalliana (as H. purpurea var. tenuifolia), Aster shortii; ““Appalachian”: Penstemon canescens; “Lowland”:
Corallorhiza wisteriana, Krigia virginica, Lactuca floridana var. floridana

LIMESTONE PLANTS 25

two species are polymorphic and partly overlap in their diagnostic characters (Fernald
1950, Mohlenbrock 1981). Two of our herbarium specimens (Klotz 1236, Walck 7)
exemplify the latter difficulty. Putative hybrids have been noted in southwest Ohio. These
morphological intermediates occurred near the two parental species and were probably
infertile (Blackwell 1977).

These two species possibly exhibit a partial ecological separation in some of the regions
where they are sympatric. In southcentral Pennsylvania, we have observed both species on
upland sites but only R. abortivus on floodplains. Similarly, along the Potomac River in
Washington, DC and Maryland, R. micranthus tends to grow in more well-drained places
than R. abortivus (Terrell 1970). In southwest Ohio, R. micranthus is confined to wooded
slopes while R. abortivus is often weedy or adventive (Blackwell 1977).

We have not yet observed R. allegheniensis, a close relative of these two species, even
though it has been reported from Franklin County (Wherry et al. 1979). According to
Sobel (1974), R. micranthus and R. allegheniensis are perhaps merely ecological varieties of
the polymorphic R. abortivus, at least in the Carolinas.

b. Dodecatheon amethystinum (Fassett) Fassett, jeweled shooting-star, is native to
eastern and southcentral Pennsylvania. It also occurs along the upper Mississippi River in
the “Driftless Area” of Wisconsin, Minnesota, and Iowa (Fassett 1944); and at scattered
localities in Kentucky (Fassett 1944), western Illinois (Schwegman 1984), and central
Missouri (Steyermark 1963). The closely similar D. meadia is widespread from Washing-
ton, DC to western Wisconsin, south to Georgia and Texas (Fassett 1944).

Both D. amethystinum and D. meadia have been reported from Pennsylvania, sometimes
from the same location, as in Franklin and Columbia Counties (Wherry et al. 1979).
However, the populations in eastern Pennsylvania are now considered to be D. amethysti-
num (R. Sutter pers. comm.; Fassett 1944). Although Fassett (1944) believed that both
species might be ing in the eastern panhandle of West Virginia, D. meadia is the only
Dodecatheon currently reported from West Virginia (Strausbaugh and Core 1977),
Maryland (Brown and Brown 1984), and Virginia (Harvill et al. 1986; Wofford 1989).

In Franklin County we have observed D. amethystinum in a calcareous rocky summit
community on bluffs along West Branch Conococheague Creek, and at several sites in
mesic calcareous upland forests. These habitats in Franklin County more nearly resemble
those reported for D. meadia than those reported as most typical for D. amethystinum.
Throughout its range, D. meadia inhabits meadows, railroad rights-of-way (Fassett 1931),
mesic or dry prairies, open deciduous woods, oak openings, and wooded to open bluffs or
cliffs. In contrast, D. amethystinum inhabits damp rock outcrops or earth slopes, deep
ravines (IItis and Shaughnessy 1960), moist crevices, and “steep slopes of north-facing
wooded limestone bluffs” (Steyermark 1963). These situations agree with the habitat of D.
amethystinum at sites in Lancaster and Columbia counties in Pennsylvania. However, in
the Midwest this species also sometimes occurs on wooded slopes (Mohlenbrock 1978) or
in small upland prairies (Iltis and SI y 1960). These situations resemble locations of
D. amethystinum in Franklin County.

Both species are generally calciphilic throughout their respective ranges. Dodecatheon
amethystinum occurs on limestone in Missouri (Steyermark 1963) and Wisconsin (Iltis and
Shaughnessy 1960). In Illinois, it nearly always occurs on outcrops of limestone or
dolomite; but a few populations are on outcrops of sandstone (Schwegman 1984). D.
meadia is usually in basic or circumneutral soil in the Carolinas (Radford et al. 1968). It
occurs on calcareous substrates in the Blue Ridge (Wofford 1989) and in Maryland (Boone
1984; Riefner and Hill 1984). In Ohio, it grows on dolomitic cliffs (Brockett and

26 BARTONIA

Cooperrider 1983); and in central Tennessee, it occurs in calcareous woodlands and even
in open cedar glades (Turner and Quarterman 1968). However, this species also occurs on
serpentine in Maryland (Boone 1984) and on sandstone as well as “high lime” prairies in
Wisconsin (Iltis and Shaughnessy 1960).

Dodecatheon amethystinum was initially described as a variety of D. meadia (Fassett
1929) and was later elevated to the rank of species (Fassett 1931). Subsequently, some
authors have included it within D. radicatum Greene subsp. radicatum of western North
America (Thompson 1953; Gleason and Cronquist 1991) despite morphological and
ecological differences between it and this western taxon (IItis and Shaughnessy 1960;
Steyermark 1963).

Fassett (1944) considered the shape, thickness, and color of the capsule walls to be the
most reliable characters for distinguishing D. amethystinum from D. meadia in both the field
and herbarium. However, Schwegman (1984) showed that only the capsule wall thickness
(measured in microns) is fully reliable; the shape and color exhibit overlap and variation.

Some of our recent field observations have exemplified the difficulty of distinguishing
these two species. In April, 1991, the senior author observed D. meadia (with white
corollas) in the Great Smoky Mountains National Park, Tennessee; the dried capsules
from 1990 resembled D. amethystinum. In June, 1991, R. Bartgis showed us a large
population of Dodecatheon (considered to be D. meadia) on the forested bluffs above the
Potomac River near Sharpsburg, Maryland. The plants are known to have dark pink
corollas (Bartgis pers. comm.) but had nearly mature fruit on the date of our visit. The
shape of the capsules (Klotz, Bartgis, and Walck 1869) varies among the plants and
apparently spans the alleged difference between the two species. We subsequently
documented a similar array of variation in capsule shape in one of the populations in
Franklin County (Klotz and Walck 1872). Our measurements of capsule wall thickness in
these populations agree with the values reported for D. amethystinum (Fassett 1944).

The observed populations of Dodecatheon in Franklin County have pale to deep pink
corollas, and some of the plants exhibit a degree of red pigmentation in the leaf bases. In
these characters, the Franklin County populations resemble D. amethystinum more than D.
meadia (Fassett 1944; Schwegman 1984). Both of these characters are unreliable in
herbarium specimens. On drying, the corollas of the two species fade to the same range of
colors; and the red pigmentation in the base of the leaves fades more in D. amethystinum
than in D. meadia (Fassett 1944).

Possible hybrids have been collected in Wisconsin at a location where the two species
occur in close proximity even though D. amethystinum blooms earlier than D. meadia by at
least one week (Fassett 1931). Both species are pollinated by bees, including some of the
same species of bumblebees (Bombus spp.). ““The existence of genetic barriers to crossing
remains to be demonstrated” (Macior 1970). Perhaps this possibility of hybridization in
the Midwest is related to the indistinctness of these two species in the eastern part of their
range. We believe that the populations in the Potomac River drainage of Pennsylvania,
Maryland, West Virginia, and Virginia should be re-examined in fruit to determine
whether they represent the same species.

c. Rhamnus lanceolata Pursh, lanceolate buckthorn, ranges from southcentral Pennsylva-
nia to southern Wisconsin and Nebraska south to Alabama and eastern Texas (Brizicky
1964; Fernald 1950). This species is known from West Virginia (Strausbaugh and Core
1977) and northern Virginia (Harvill et al. 1986) but has not been reported from Maryland
(Boone 1984; Brown and Brown 1972). Most of the records in Pennsylvania are from the
southcentral counties west of the Susquehanna River (Wherry et al. 1979).

LIMESTONE PLANTS | 27

Although it is a large shrub, this species is not visually distinctive and is easily
overlooked. It occurs in upland habitats such as bluffs, cliffs, rocky or gravelly wooded
slopes, glades, and thickets; but it can also be found in springy places, fens, or floodplains
(Deam 1970; Mohlenbrock 1982a; Steyermark 1963). We have not yet observed this
species in a floodplain or other wetland but only in upland habitats, including a
successional red cedar woodland described below, and a xeric upland forest. The species
tends to be calciphilic (Braun 1961; Brizicky 1964; Deam 1970; Gleason and Cronquist
1991; Great Plains Flora Association 1986; Steyermark 1961).

Two varieties, perhaps merely “trivial” forms distinguished by pubescence (Johnston
1975), have been described within R. lanceolata. Variety glabrata predominates in the
eastern and southeastern parts of the range of the species, but our specimen (Klotz 1254)
fits var. lanceolata, which predominates in the western part of the range (Brizicky 1964).

Some descriptions suggest that the species is heterostylous (Fernald 1950; Mohlenbrock
1982a) or possibly functionally dioecious (Great Plains Flora Association 1986). We have
not determined which floral morphs occur in the study area. The senior author has
observed some plants in fruit in one of the populations.

d. Matelea obliqua (Jacq.) Woods., oblique milkvine, ranges from Pennsylvania to
Missouri south to Georgia (Fernald 1950; Gleason and Cronquist 1991). In Pennsylvania it
occurs sparsely throughout the southeastern and southcentral counties (Wherry et al.
1979).

We have observed three small populations of this species: two near the roadside edges of
mesic calcareous forest, and another in an open, grassy area in successional red cedar
woodland. At both sites M. obliqua grows in partial shade, which is the optimal condition
for the vegetative growth of this species in cultivation (Rosatti 1989). Throughout its
range, it occurs in woods, thickets, and riverbanks (Small 1933). However, the species of
Matelea are reported to grow best in well-drained soils and do not occur in areas subject to
flooding (Rosatti 1989). Matelea obliqua is known to be calciphilic in Maryland (Riefner
and Hill 1984), North Carolina (Radford et al. 1968), the Blue Ridge (Wofford 1989), and
the southeastern states in general (Rosatti 1989).

e. Hydrophyllum macrophyllum Nutt., large-leaved waterleaf, has two known populations
in Franklin County, both in mesic calcareous forest. The larger population has been
described previously (Klotz and Walck 1990). It occurs with H. virginianum in that
community, and H. canadense is located on the nearby floodplain of Licking Creek.

f. Ruellia strepens L., limestone petunia, ranges from northcentral New Jersey to lowa
and Kansas south to South Carolina and eastern Texas (Fernald 1950; Gleason and
Cronquist 1991). This species is frequent in southwestern Franklin County and along the
Conodoquinet Creek in neighboring Cumberland County. There are four other records in
Pennsylvania, all in the southern counties (Wherry et al. 1979). However, the species is
easily overlooked. Our observation agrees with other authors that R. strepens occurs in
upland forests, floodplain forests, and along mesic roadsides (Boone 1984; Deam 1970;
Fernald 1950; Steyermark 1963). It is noted to be somewhat calciphilic (Allard and
Leonard 1962; Boone 1984; Fernald 1950; Long 1970; Small 1933). The junior author has
observed this species from shale in addition to limestone and alluvium in southwestern
Franklin County.

We have seen chasmogamous flowers infrequently, chiefly in early June. The observed
populations mostly appeared vegetative or belonged to “forma cleistantha” (Fernald
1950), which produces cleistogamous flowers that yield “numerous fruits with good seeds”
(Long 1970). This so-called “form” is probably only a phase that develops during the latter

28 BARTONIA

part of the growing season. However, these two phases reportedly differ in leaf width and
length. Differences between the cleistogamous and chasmogamous flowers include
peduncle length, frequency of glands on the calyx-lobes, and size and form of the
corolla—i.e. small and tubular in the cleistogamous flowers (Long 1970; McCoy 1937). The
species is reported to be predominantly autogamous in the northern portion of its range
(Long 1970). This situation possibly explains the infrequency of chasmogamous flowers in
our observations.

g. Chrysogonum virginianum L., green-and-gold, ranges from southern Pennsylvania and
eastern Ohio to western Florida and southern Mississippi (Gleason and Cronquist 1991;
Stuessy 1977). The populations in Pennsylvania are reported from southern Franklin and
Fulton Counties (Wherry et al. 1979) and from Lancaster County (Porter 1903). They
compose part of the northeastern limit of the range of the species (Stuessy 1979).
Specimens from the northern part of the range are mostly var. virginicum, which is
cespitose and has well developed internodes. In the Carolinas, it gives way to var. australe,
which is stoloniferous and has mostly basal leaves on the erect stems (Cronquist 1980;
Steussy 1977). Morphological intermediates between these varieties have been collected
throughout much of the total range, including southcentral Pennsylvania (Stuessy 1977).

The habitats of Chrysogonum throughout its range include rich woods, dry woods, and
shaded rocks (e.g. Fernald 1950; Small 1933). In Maryland, the species is known to occur
on limestone but is not restricted to it (Boone 1984). Chrysogonum occurs with
Dodecatheon amethystinum at one location that we have observed. However, the two
species occupy different microhabitats at this site: Dodecatheon chiefly occurs in grassy
openings, which resemble the calcareous rocky summit community type described below,
whereas Chrysogonum occurs in the shaded understory of the mesic upland forest.

h. Corallorhiza wisteriana Conrad, spring coral-root, ranges from Pennsylvania to
Wisconsin and Kansas south to central Florida and eastern Texas. It also occurs in the
Western Cordillera from southern Montana and eastern Idaho to southern Mexico, and in
the Black Hills of South Dakota (Case 1987; Correll 1950). Most of the records in
Pennsylvania are from southeastern counties, but one is southwestern (Wherry et al.
1979). In eastern North America this species is found in deciduous, mesic to xeric rich
forests and swamp forests (Case 1987; Correll 1950; Sheviak 1974). In Indiana, this species
grows on “beech slopes, sometimes in black or black and white oak woods, and rarely in
white oak woods” (Deam 1970). It may grow mostly in circumneutral, nutrient-rich humus
(Correll 1950 fide Wherry), but it is not restricted to limestone. In Illinois, the soils in
which it grows are generally “moderately acidic” (Sheviak 1974).

Our two sightings of C. wisteriana were in the same community: one shoot in 1987 and a
cluster of 15 shoots in 1991. This scarcity is not surprising. The species rarely occurs in
colonies, and individuals are usually five meters or more apart (Deam 1970). The plants do
not bloom every year, long periods may pass between blooming (Cusick 1984a), and
populations vary annually in the number of individuals that bloom. Furthermore, the
inflorescence is inconspicuous and is the only part of the plant above ground. The plants
are essentially rootless and do not produce significant amounts of chlorophyll (Case 1987).
Instead, they are dependent on a mycorrhizal fungus that envelops and invades the
rhizome (Campbell 1970; Case 1987; Cusick 1984a). The prevention of trampling is
especially important in the habitat of Corallorhiza because the mycorrhizal relationship
makes it sensitive to soil disturbance (Cusick 1984a).

CALCAREOUS ROCKY SUMMIT COMMUNITY. This community type is characterized by
conspicuous rock outcrops and a discontinuous canopy with herbaceous openings (PNDI

LIMESTONE PLANTS 29

1983). We have observed two communities that somewhat approach this category. Both of
them contain only one rare species. The community with Opuntia humifusa is artificial and
not diverse: it occurs along the edge of a limestone quarry. In contrast, the community with
Dodecatheon amethystinum appears natural and is species-rich: it consists of a suite of
grassy openings in the calcareous upland forest on the crest of north-facing bluffs along the
West Branch of the Conococheague Creek. In both of these rocky summit communities,
the trees include Juniperus virginiana and Celtis occidentalis; but several other canopy
species characteristic of calcareous upland forest (e.g. Quercus muhlenbergii) are present in
the community with D. amethystinum. The latter community has considerable floristic and
esthetic value. Similar rocky summit communities are reported in New York (Reschke
1990): “red cedar rocky summit community” (specified as Saneareons), and “rocky summit
grassland”; New Jersey (Breden 1989): a similar category, “‘traprock glade/rock outcrop
community”; and Pennsylvania (PNDI 1983): “calcareous rocky summit community.”

Opuntia humifusa (Raf.) Raf., prickly-pear cactus, ranges from Massachusetts to Iowa,
south to Florida and Texas (Benson 1982). In Pennsylvania it occurs sparsely in the
southern half of the state and along the Delaware River (Wherry et al. 1979). Throughout
its range, it occurs on well-drained, exposed sandy soil and on rock outcrops, which include
granite, sandstone, limestone, or shale (Benson 1982; Brown and Brown 1984). Distinctive
community types that contain this species include shale barrens (Strausbaugh and Core
1977), cedar glades (Braun 1964; Baskin and Baskin 1978), and coastal dunes (Brown and
Brown 1984). In New Jersey, it occurs on level terrain and on south (to southeast) slopes
(Hanks and Fairbrothers 1969a). In Franklin County we have observed a large population
of this species on shale and a small population on the forested brink and cliff face of a
limestone quarry.

Two species were formerly recognized in Pennsylvania (Wherry 1926; Wherry 1964;
Wherry et al. 1979): O. calcicola Wherry and O. compressa (Salis.) Macbr. Both are now
considered conspecific with O. humifusa (Kartesz and Kartesz 1980), a polymorphic
species (Hanks and Fairbrothers 1969b) exhibiting a high degree of morphological
variability within populations (Romeo et al. 1987).

CALCAREOUS CLIFF COMMUNITY. This community type occurs on steep exposures and
ledges of resistant, calcareous bedrock. Soil is nearly non-existent and the herbaceous
vegetation is sparse and open (PNDI 1983). The degree of moisture ranges from mesic to
xeric depending on aspect and cover. Similar communities have been described in New
York (Reschke 1990) and Pennsylvania (PNDI 1983): “calcareous cliff community”;
Maryland (Riefner and Hill 1984): limestone habitats one and two; and West Virginia
(Core 1966): “limestone cliffs.”

Mesic calcareous cliffs were observed in two types of locations. The first consists of
small, vertical limestone outcrops in upland forest. We did not find any rare species on
them. The second comprises wooded, north-facing bluffs along clearwater creeks, namely
Licking Creek (in Pennsylvania and Maryland), Little Cove Creek, and the West Branch of
the Conococheague Creek. Two rare species were observed on this type of mesic
calcareous cliff: Asplenium resiliens and Sedum telephioides. On the rocky bluffs along
Licking Creek, the two species occur in both Pennsylvania and Maryland. These two
species plus seven others that we have observed in this community type in Franklin County
are largely restricted to limestone or other basic and circumneutral substrates in
Maryland: Pellaea atropurpurea, Asplenium rhizophyllum, A. ruta-muraria, Cystopteris
bulbifera, Arabis lyrata, Galium boreale, and Solidago flexicaulis (Boone 1984; Riefner and
Hill 1984).

30 BARTONIA

The mesic calcareous cliff communities contain twelve species of epipetric ferns,
including six species of Asplenium. The most constant are A. platyneuron, A. rhizophyllum,
and. A. trichomanes, which occur in five or all six examples of this community type that were
tabulated. Their most constant associates are two herbaceous dicotyledons, Aquilegia
canadensis and Thalictrum dioicum. The senior author discovered the county record of
Asplenium pinnatifidum as a small population on a vertical limestone rockface in this region
(Klotz 1767). This species usually inhabits non-calcareous rocks (Cobb 1956; Mickel 1979;
Reed 1953) including sandstone (Lellinger 1985), and igneous and metamorphic outcrops
(Wofford 1989).

Xeric calcareous cliffs also occur in southwestern Franklin County. We have observed
two rare species on them: Opuntia humifusa on the edge and side of the limestone quarry
described above in the calcareous rocky summit community, and one population of Sedum
telephioides described below.

a. Asplenium resiliens Kunze, black-stemmed spleenwort, ranges from southcentral
Pennsylvania to southern Nevada, south to the West Indies and northern Argentina
(Lellinger 1985; Mickel and Beitel 1988; Small 1964). In Pennsylvania the species is known
only from southwestern Franklin County (Wherry et al. 1979), which constitutes the
northeastern limit of the range. This fern is an epipetric calciphile occurring on limestone,
dolomite, and marl of cliffs, outcrops, and sinkholes (Gleason and Cronquist 1991;
Lellinger 1985; Radford et al. 1968; Steyermark 1963). In the southeastern states it is
noted as “preferring limestone, but also [occurring] on igneous rocks” (Small 1964).

Although A. resiliens is an apogamous triploid, it hybridizes with A. heterochroum, a
species of Florida, Bermuda, and the West Indies (Lakela and Long 1976; Lellinger 1985).
Asplenium resiliens is not known to hybridize with the Appalachian species of Asplenium
even though the latter have produced a “complex” of hybrids and backcrosses (Lellinger
1985).

The populations of A. resiliens along Licking Creek occur in close proximity to Sedum
telephioides, but the two species do not share the same microhabitats. In one instance, S.
telephioides occurs fairly high on the bluffs, mostly on mesic but well-drained, sloping to
vertical, forested rock faces. The adjoining stand of A. resiliens at this site occurs lower on
the bluffs on moist, vertical rock faces. In the other instance, the two species are
horizontally separated by about 200 meters. The Sedum occurs on an exposed grayish-red
outcrop without an overarching canopy whereas the Asplenium occurs in shaded, moist
crevices on forested bluffs.

b. Sedum telephioides Michx., Allegheny stonecrop, ranges from southern Pennsylvania
to southern Illinois south to western South Carolina (Clausen 1975). It is also reported
from New York, but these populations are other, misidentified species of Sedum except
one which may represent an escape from cultivation (Clausen 1975; Spongberg 1978). In
Pennsylvania the plant is restricted to six known sites in Bedford, Fulton, and southwest-
ern Franklin Counties. All but one are near the Maryland border (Wherry et al. 1979) and
represent the northeastern limit of the species.

Sedum telephioides occurs on dry, rocky ledges; exposed or shaded cliffs; rocky woods;
rock crevices; and steep bluffs (Mohlenbrock and Voigt 1959; Gleason and Cronquist
1991; Radford et al. 1968; Spongberg 1978). It is not a calciphile, but is known from
greenstone, gneiss, granite, sandstone, shale, and slate in addition to limestone (Clausen
1975). It occurs on sandstone outcrops and shale barrens in West Virginia (Strausbaugh
and Core 1977), on limestone and shale outcrops in Maryland (Riefner and Hill 1984), and

LIMESTONE PLANTS 31

on sandstone and limestone cliffs in Indiana (Deam 1940; Spongberg 1978). At one of the
two sites in Fulton County, PA, the species occurs on a south-facing bluff of reddish rock of
the Catskill Formation, “a complex unit consisting of shale, siltstone, sandstone, and
conglomerate” (Geyer and Wilshusen 1982). The community at this site combines species
of the acidic and calcareous cliff community types (PNDI 1983). The other population in
Fulton County occurs in a shale barren community on calcareous Devonian shale (Cusick
1990). It also grows on shale barrens in Bedford County (Henry 1978). One of the two
populations in Franklin County occurs sympatrically with S. ternatum.

FLOODPLAIN FOREST. Floodplain forests are hardwood forests on mineral soils of low
terraces along rivers and creeks that flood periodically (PNDI 1983). The following species
of canopy trees were observed in three or more of the five examples that were tabulated:
Platanus occidentalis, Ulmus americana, Carya cordiformis, Juglans nigra, Acer negundo, A.
saccharinum, and Fraxinus americana and/or F. pennsylvanica. The shrubs and vines with
this degree of constancy include: Lindera benzoin, Rosa multiflora, Rubus spp., Cornus
amomum, Vitis vulpina, and Toxicodendron radicans. The herb layer is commonly luxuriant
and diverse. The examples of this community type containing rare species occur along
Licking Creek and the West Branch of the Conococheague Creek. The floodplain forests
in our study area are floristically similar to those described in New York (Reschke 1990),
New Jersey (Breden 1989), Pennsylvania (PNDI 1983), West Virginia (Core 1966):
“floodplain forest”; Maryland (Brush et al. 1980): “‘sycamore—green ash—box elder-silver
maple association”; and Virginia (Harvill et al. 1977): “floodplain forest,” (Rawinski
1990): “eutrophic seasonally flooded palustrine system.”

The rare species that we have observed are Ruellia strepens (discussed above) and Ptelea
trifoliata. They and four other species that we have seen in this community type are
generally restricted to limestone or other basic to circumneutral soils in Maryland: Carya
laciniosa, Hydrophyllum canadense, Arisaema dracontium, and Carex shortiana (Boone
1984; Riefner and Hill 1984).

Ptelea trifoliata L., common hop-tree, occurs from Connecticut and southern Quebec to
Nebraska, south to Florida and northern Mexico, and west to Arizona (Fernald 1950;
Gleason and Cronquist 1991). Our specimens belong to subsp. trifoliata var. trifoliata. In
Pennsylvania, the species is known from a few localities, mostly in the southeastern
counties and along Lake Erie (Wherry et al. 1979). It occurs in a wide range of habitats
including alluvial and upland rocky woods, swamps, glades, prairie openings, dry blufts,
thickets, fencerows, sandy fields, and wooded to open dunes along the Great Lakes
(Billington 1949; Radford et al. 1968; Steyermark 1963; Voss 1985). Along the north shore
of Lake Erie, it especially occurs on the windward side of beaches and sandspits (Ambrose
et al. 1985). It is known to be calciphilic in Maryland (Boone 1984; Riefner and Hill 1984),
Missouri (Steyermark 1963), and the Great Plains (Great Plains Flora Association 1986);
and it is often in limestone areas in southern Illinois (Mohlenbrock 1982b) and southern
Ontario (Ambrose et al. 1985).

In 1985, we observed ten trees of P. trifoliata on the floodplain along the north bank of
Licking Creek. Some of them were two to three meters tall, but we observed no saplings or
seedlings. In 1990, we observed only two of these trees; and they appeared unhealthy. We
also found one tree of P. trifoliata along the south bank.

The species is predominantly dioecious, rarely polygamous (Ambrose et al. 1985;
Brizicky 1962). In Ontario, the sex ratio of native populations is skewed toward the males
(Ambrose et al. 1985). We have observed both sexes in the local population, but we have

a2 BARTONIA

not determined the sex ratio. Despite the potential for root sprouts (Brizicky 1962), the
species is not clonal and depends on seeds for establishment. However, flowering and
fruiting are reduced or suppressed by shading (Ambrose et al. 1985).

SUCCESSIONAL RED CEDAR WOODLAND. Our three examples of this community type
occur in different situations: an abandoned field and pasture, a powerline cut, and the
cleared areas around a limestone quarry. The first of these is the most extensive and
possibly the oldest. It occurs on karst terrain with a mosaic of vegetation: an upland
savanna dominated by Juniperus virginiana surrounds numerous sinkholes that support
closed deciduous forest. The ground layer of the savanna contains the rare elements
Ruellia humilis, Rhamnus lanceolata, Matelea obliqua, Bouteloua curtipendula, plus many
other herbaceous and woody species. The sinkholes support “islands” of mixed deciduous
forest with a canopy that includes Celtis occidentalis, Ulmus rubra, Carya ovalis, Quercus
muhlenbergii, Q. rubra, Prunus serotina, Acer negundo, and Fraxinus americana. Communi-
ties that are similar to the red cedar woodlands of Franklin County have been described in
New York (Reschke 1990): “successional red cedar woodland”; New Jersey (Breden
1989): “limestone glade”; Pennsylvania (Duppstadt 1972): “cedar barrens,” (PNDI 1983):
intermediate between “successional field” and “young miscellaneous forest’; Virginia
(Rawinski 1990): “oligotrophic scrub” or, as the trees grow taller, “oligotrophic woodland”;
and West Virginia (Bartgis 1985): “limestone glade”; (Core 1966): intermediate between
“artificial prairies” and “red cedar forest type.”

The cedar glades of the unglaciated eastern United States can be defined as
communities of low herbaceous vegetation with “limestone or dolomite bedrock at or near
the surface” (Baskin and Baskin 1986). The soils are very shallow or restricted to crevices,
saturated from late autumn through early spring, and droughty during the remainder of
the year (Baskin and Baskin 1974). Slightly deeper soils support cedar barrens, which are
forest openings dominated by mid-to-tall, native perennial grasses (DeSelm 1989;
Quarterman 1989). Franklin County lacks the extensive bedrock pavement that character-
izes the cedar glades of the Ridge and Valley Province from West Virginia to Alabama; the
Interior Low Plateaus of Kentucky and Tennessee to southern Ohio, Indiana, and Illinois;
and the Ozark region (Baskin and Baskin 1986). However, some of the successional
woodlands on limestone in Franklin County resemble the cedar barrens of these regions in
having shallow soils and several of the characteristic species, including Juniperus virginiana,
Ophioglossum engelmannii, Ruellia humilis, and Bouteloua curtipendula (Baskin and Baskin
1974; DeSelm 1989).

Before European settlement in the mid eighteenth century, the limestone regions of the
Great Valley in Franklin County contained luxuriant grasslands, which were called
“barrens” because of the scarcity of trees (Finafrock 1942; M’Cauley 1878; Orr 1904). The
extensiveness of these grasslands has been debated (Cooper 1903). They were maintained
annually by the Indians in November, when “the woods and grass of the mountains and
prairies were burned and their game was driven from concealment” (Richard 1887). The
practice of burning ceased after European settlement; and within less than a century, the
grassland developed into forest, most of which was cleared during the nineteenth century
(Orr 1904).

The prairies that formerly occupied the Big Barrens region of Kentucky may have had a
similar origin. The karstic topography helped to dry the soil and was therefore conducive
to fire. After the Indians ceased to burn the vegetation, “the swift return of the forests . . .
caused a large part of this prairie country to be rewooded before it could be subjected to

LIMESTONE PLANTS 33

the plough” (Baskin and Baskin 1981). The prairies of the Middle West have had a similar
history except that the period of reforestation began much later, e.g. after 1860 in Illinois
(Gleason 1922).

In southwestern Franklin County, the presence of unusual native flora on karst
topography suggests that some examples of red cedar woodland may be relics of the former
grassland vegetation. Of course, many stands of red cedar woodland in this region are
simply the result of recent secondary succession on agricultural land. For example, in
Bedford County, Pennsylvania, “cedar barrens” have developed as Juniperus virginiana has
spread in heavily pastured areas on limestone soils, sometimes in association with Cercis
canadensis. The herbaceous plants are suppressed by grazing (Duppstadt 1972). The
presence of Juniperus virginiana indicates the absence of burning since the thin bark and
the roots near the soil surface are injured by light surface fires (Fowells 1965). In this
respect, red cedar woodlands differ from the midwestern oak savannas and “barrens,” in
which the characteristic woody species are more tolerant of fire. However, in all of these
community types, burning would enhance the native prairie species such as Ruellia humilis
and Bouteloua curtipendula (Anderson 1981; Anderson 1982; Anderson and Schwegman
1991; Curtis 1959).

The four rare species that we have observed only in this community type are
Ophioglossum engelmannii, Salvia reflexa, Ruellia humilis, and Bouteloua curtipendula. In
addition, we have seen Rhamnus lanceolata and Matelea obliqua, which also occur in
calcareous upland forest. Five of the species that we have seen in this community type are
generally restricted to limestone or other basic to circumneutral soils in Maryland: Matelea
obliqua, Trichostema brachiatum, Ruellia humilis, Eupatorium altissimum, and Bouteloua
curtipendula. We have also seen Panicum flexile and Sporobolus neglectus in this community
type. These two grasses are “endangered” in Maryland but are not yet known from
limestone in that state (Boone 1984; MNHP 1991; Riefner and Hill 1984; Bartgis pers.
comm.).

a. Ophioglossum engelmannii Prantl, Engelmann’s adder’s-tongue, was discovered in
June 1990 for the first time in Pennsylvania and represents the known northern limit of the
range of the species (Walck and Klotz 446 PH). Otherwise, it ranges from northern
Virginia to Kansas and Arizona south to Florida and southern Mexico (Fernald 1950;
Mickel and Beitel 1988). It is not known to occur in Maryland (Boone 1984) but was
collected in 1989 in a glade woodland on dry limestone in Hardy County, West Virginia
(Bartgis 1252 PH). This species is calciphilic: it inhabits soils derived from limestone or
calcareous shale in pastures, barrens, cedar glades, and open woods (Baskin and Baskin
1974; Cusick 1984b; Lellinger 1985; Mickel and Beitel 1988; Wherry 1961; Wofford 1989).
In Illinois, it is restricted to limestone ledges whereas O. vulgaris occurs in moist woods and
shaded sandstone ledges (Mohlenbrock 1967). In the Great Plains, it occurs on shallow
soil over limestone and rarely on sandstone in prairies, pastures, or open woods (Great
Plains Flora Association 1986).

The population in Pennsylvania occurs in an upland portion of a powerline cut through a
deciduous forest. The population is limited but dense and occurs within an area of four
Square meters. The visible part consisted of 69 fertile leaves plus 149 vegetative leaves. The
species is known to be clonal: the roots grow horizontally and produce adventitious buds,
which produce new shoots. The short, vertical rhizomes produce one or two, rarely three
leaves during a growing period (Baskin and Baskin 1974; Shaver 1954). After summer
droughts the leaves die, but a reduced number of new ones appear in late summer or fall

34 BARTONIA

following rainy periods (Steyermark 1963; Baskin and Baskin 1974). Because the fronds
wither quickly after the spores are released, the populations can be located only within a
limited period (Cusick 1984b).

Population studies of this species differ markedly in the reported percentages of plants
with fertile leaves (Baskin and Baskin 1974; Shaver 1954; Steyermark 1963). These
discrepancies suggest that populations may differ in vigor or may vary in vigor from year to
year because of differences in soil moisture (Baskin and Baskin 1974).

b. Salvia refleca Hornem., lance-leaved sage, ranges from Ohio and Michigan to
Montana and Utah, south to Arkansas and Mexico. It is sometimes adventive in the
northeastern United States (Fernald 1950; Gleason and Cronquist 1991) and southern
Canada (Scoggan 1978-1979). In Pennsylvania, it is known from a few scattered locations
in the southern half of the state (Wherry et al. 1979)—for example, in ballast and on
curbing in Lehigh County (R.L. Schaeffer 99984, 100033, 101149). It is recorded from one
location in Maryland (Hill 1988) and is sparingly established in West Virginia (Straus-
baugh and Core 1977). In Indiana, it is established in disturbed sandy soil in a few locations
(Deam 1970). Throughout its range it occurs in disturbed habitats including dry pastures,
roadsides, railroads, fields, wash areas in prairies, dry rocky open ground, open woods, and
alluvial ground (Great Plains Flora Association 1986; Steyermark 1963). Salvia reflexa
shows no indication of being calciphilic. Nevertheless, our single site for this species is
close to a limestone quarry. The immediate vegetation is highly disturbed and consists
largely of two aggressive, non-native species: Ailanthus altissima and Lonicera japonica.

c. Ruellia humilis Nutt., fringed-leaved petunia, ranges from Pennsylvania to Nebraska
south to western Florida and Texas (Gleason and Cronquist 1991; Long 1970). In
Pennsylvania, it is only known from Franklin County (Wherry et al. 1979). Throughout its
range, it occurs in open fields, dry prairies, glades, dry or rocky woods and the escarpment
of bluffs (Great Plains Flora Association 1986; Long 1970; Steyermark 1963). It is present
in cedar glades of the Big Barren region of Kentucky (Baskin and Baskin 1981). In
southern Illinois, it occurs in the upland grass communities called hillside prairies, which
are on the crest of bluffs, especially limestone, that border the Mississippi River (Evers
1955; Voigt and Mohlenbrock 1964). It also occurs in similar communities in Ohio and
Indiana (Gordon 1969). This species is known to be calciphilic in Maryland where it
“grows in dry woods and basic habitats that simulate prairies” (Riefner and Hill 1984).

Ruellia humilis is morphologically variable. Two to five varieties have been recognized
(Fernald 1950; Gleason and Cronquist 1991), but their importance is doubtful (Long
1970). Our specimen (Klotz 1253) combines the characters that are supposed to separate
var. humilis and var. frondosa Fern., the two varieties that have been reported from
Pennsylvania (Fernald 1950). In some parts of its range, R. humilis intergrades morpholog-
ically with R. caroliniensis; and these two species can hybridize (Long 1961).

Like R. strepens, R. humilis can produce cleistogamous flowers (Fernald 1950); but we
have observed that it also produces abundant chasmogamous flowers. We have not found
R. strepens and R. humilis growing sympatrically; R. humilis occurs in more open habitat.
We have observed R. humilis in two of the red cedar woodlands described above, namely
the disturbed vegetation surrounding a quarry and the more mature vegetation on karst
terrain.

d. Bouteloua curtipendula (Michx.) Torr., tall grama, ranges from Maine to southern
Saskatchewan south to Georgia, southern California, and Argentina (Fernald 1950;
Hitchcock 1971; Scoggan 1978-1979). In Pennsylvania, it occurs in a few localities in the
southcentral and eastern counties (Wherry et al. 1979). It is one of the chief grasses of the

LIMESTONE PLANTS 35

prairies of the midwestern states (Curtis 1959; Weaver 1968) and can dominate the xeric
extreme of the topographic moisture gradient (Bazzaz and Parrish 1974). It is frequent on
limestone glades, barrens, and prairie openings in rocky woodlands in Missouri (Steyer-
mark 1963) and southwestern Virginia (Carr 1965). It is absent from the cedar glades of
Kentucky but is present in cedar glades in Tennessee, Alabama, and Georgia (Baskin and
Baskin 1981 and pers. comm.). In Ohio, Indiana, and Illinois, the species is important in
the upland grass communities or hillside prairies that were mentioned above for Ruellia
humilis (Deam 1970; Evers 1955; Gordon 1969; Voigt and Mohlenbrock 1964). It occurs on
serpentine barrens in southeastern Pennsylvania (Braun 1964; Pohl 1947; Rhoads 1986;
Wherry et al. 1979); shale barrens in Maryland (Morse 1984) and West Virginia
(Strausbaugh and Core 1977); and dry, open calcareous slopes in New Jersey (Snyder
1986), Pennsylvania (T. Smith pers. comm.), Maryland (Boone 1984; Riefner and Hill
1984), and West Virginia (Bartgis 1985; Strausbaugh and Core 1977). In our study area it
occurs with R. humilis on the karst terrain described above.

EARLIER RECORDS OF RARE PLANTS ON LIMESTONE
IN SOUTHWESTERN FRANKLIN COUNTY

Thomas C. Porter, author of the Flora of Pennsylvania (1903), collected plants in
Franklin County in the 1850s and recognized the distinctiveness of its flora within the state.
We have observed four of the species that he considered “rare” or “peculiar” at that time:
Rhamunus lanceolata, Matelea obliqua (as Gonolobus obliquus), Ruellia strepens (as Diptera-
canthus strepens), and Chrysogonum virginianum (Porter 1872).

Besides the 16 species described above, 11 additional “Species of Special Concern”
(PNDI 1991) have been collected previously in southwestern Franklin County (Wherry et
al. 1979—except Clematis viorna, which was noted by Porter 1872) and are likely to be
associated with calcareous substrates there. They can be classified into three categories of
habitat:

(1) RICH WOODS AND BOTTOMS. Clematis viorna (Boone 1984; Radford et al. 1968;
Riefner and Hill 1984; Steyermark 1963). Quercus shumardii (Boone 1984; Fernald 1950;
Small 1933; Waldron et al. 1987; also on rocky slopes and in swamps—Grimshaw and
Bradley 1973). Spiranthes ovalis (Boone 1984; Fernald 1950; Riefner and Hill 1984; also in
disturbed sites: abandoned fields and pastures, edges of woods and thickets, wooded soil
bank of a strip mine—Sheviak 1974).

(2) PRAIRIES AND DRY, OPEN woops. Ranunculus fascicularis (Boone 1984; Fernald
1950; Keener 1976; Riefner and Hill 1984; Wofford 1989).* Onosmodium hispidissimum
(Baskin and Baskin 1986; Boone 1984; Fernald 1950; Riefner and Hill 1984; Strausbaugh
and Core 1977). Pycnanthemum torrei (Riefner and Hill 1984). Solidago rigida (Radford et
al. 1968; Snyder 1986).

(3) WETLANDS (POSSIBLY MARL WETLANDS). Juncus brachycephalus (Boone 1984; Deam
1970; Fernald 1950). Carex buxbaumii (Bartgis and Lang 1984; Deam 1970; Small 1933;
Steyermark 1963; Voss 1972). Carex lasiocarpa (Bartgis and Lang 1984; Voss 1972). Carex
tetanica (Bartgis and Lang 1984; Boone 1984; Deam 1970; Fernald 1950; Keener and Park
1986; Rhoads 1989; Riefner and Hill 1984; Westerfeld 1961).

Additional examples of the first two community types remain to be explored in

4 After this paper was completed, John Kunsman informed us that he had recently rediscovered a historical
population of this species on limestone bluffs along the Conococheague Creek in Franklin County.

36 BARTONIA

southwestern Franklin County, and most of the sites that we have examined still lack a
complete floristic survey conducted over the entire growing season. Calcareous wetlands
remain to be discovered and described in this region, but marl deposits are present: they
are associated with the Warner soil series (Long 1975). Marl wetlands like those of the
Great Valley Province in eastern West Virginia (Bargis and Lang 1984) could contain
some of the wetland species listed above. Besides the eleven historic species that remain to
be rediscovered on limestone in southwestern Franklin County, additional rare species
associated with limestone are known from the neighboring portions of Maryland (Boone
1984; Riefner and Hill 1984), eastern West Virginia (Bartgis 1985; Bartgis and Lang 1984),
and other parts of southern Pennsylvania (e.g. Pachistima canbyi—Duppstadt 1972;
Wherry et al. 1979). Also, the hybrids and backcrosses of the “Appalachian Asplenium
complex” (Lellinger 1985) could be present in this region. All of these taxa should be
sought in southwestern Franklin County.

SUMMARY

Sixteen species of vascular plants that are rare (i.e. “Plants of Special Concern’) in
Pennsylvania are currently known from calcareous soils or outcrops in southwestern
Franklin County: Ophioglossum engelmannii, Asplenium resiliens, Ranunculus micranthus,
Opuntia humifusa, Dodecatheon amethystinum, Sedum telephioides, Rhamnus lanceolata,
Ptelea trifoliata, Matelea obliqua, Hydrophyllum macrophyllum, Salvia reflexa, Ruellia
humilis, R. strepens, Chrysogonum virginianum, Bouteloua curtipendula, and Corallorhiza
wisteriana. For each of these species, the habit, habitat, range, and rarity are described;
and the literature on intraspecific variation, breeding system, and hybridization is
summarized. Each population is characterized according to physiographic section,
geologic formation, elevation, soil mapping unit, slope, community type, shading, area, and
number of aerial shoots. Additional species are listed that have been collected historically
in this region, probably from calcareous substrates. Floristic information is provided on the
communities and community types on calcareous substrates in southwestern Franklin
County: upland forest, rocky summit, cliff, floodplain forest, and red cedar woodland. The
flora of one mesic upland forest community is analyzed in detail.

ACKNOWLEDGMENTS

We are grateful to the following residents of Franklin County for their congenial
cooperation in this project. Mr. Robert Hunter discovered the three known populations of
Dodecatheon amethystinum in the vicinity of Licking Creek and the second population of
Hydrophyllum macrophyllum in that area. Mr. Charles Brightbill provided several leads:
the rich, calcareous slope along Licking Creek (described by Klotz and Walck 1990);
Asplenium resiliens along Little Cove Creek; and Dodecatheon amethystinum along the
West Branch of the Conococheague Creek. Mr. and Mrs. John J. McCulloh and family,
Mrs. Margaret Weller, Mr. and Mrs. Harry Heinbaugh, and the owners of Valley Quarries,
Inc. personally granted us access to their respective properties.

We also thank all of the following persons and agencies for their assistance and support
in this project. Thomas Smith, Phillip de Maynadier (both formerly of PNDI), Dr. Roger
Latham (The Nature Conservancy, Philadelphia Office), and Dr. Ann Rhoads (Morris
Arboretum, University of Pennsylvania) accompanied us in some of our field work. The
Pennsylvania Natural Diversity Inventory provided copies of their pertinent field reports.
Botanists and ecologists of the natural heritage programs in the surrounding states
contributed current editions of their rare plant listings and community classifications:

LIMESTONE PLANTS 37

Steve Young (New York), David Snyder (New Jersey), Keith Clancy (Delaware), Gene
Cooley (Maryland), Chris Ludwig and Tom Rawinski (Virginia), P.J. Harmon (West
Virginia), and Allison Cusick (Ohio). Robert Sutter (Southern Heritage Task Force)
annotated our initial collections of Dodecatheon. Charles Bier (Western Pennsylvania
Conservancy) provided data on Sedum telephioides in western Pennsylvania. Drs. Jerry an
Carol Baskin (University of Kentucky) provided information on Bouteloua curtipendula in
the cedar glades. Dr. Alfred E. Schuyler (Academy of Natural Sciences), Dr. Carl Keener
(Pennsylvania State University), Rodney Bartgis (Maryland Natural Heritage), and Dr.
Ann Rhoads (Morris Arboretum of the University of Pennsylvania) provided helpful
suggestions on the manuscript. Shippensburg University provided the use of a microcom-
puter and the support for numerous interlibrary loans and computerized bibliographic
searches (DIALOG).

LITERATURE CITED

ALLARD, H. A. AND E. C. LEONARD. 1962. List of vascular plants of the Northern Triassic area of Virginia.
Castanea 27: 1-56.

AMBROSE, J. D., P. G. KEVAN, AND R. M. GADAWSKI. 1985. Hop tree (Ptelea trifoliata) in Canada: populati d
reproductive biology of a rare species. ana. J. Bot. 63: 1928-1935.

ANDERSON, D. 1981. Ohio plant community ¢ l tion and survey. Ohio Dept. Natural Resources, Columbus.

ANDERSON, R. C. 1982. An | i ke f fire, climate, and grazing animals in the
origin and maintenance oy grasslands: an igus pape Pages 297-308 in J. R. Estes, R. L. Tyrl, and J. N
Brunken, eds. G logy. Univ. of Oklahoma Press, Norman.

————-AND J. E. pera se 1991. Twenty years of vegetational ‘change on a southern Illinois barren. Natural
Areas J. 11:

Anon. 1990. aioe and dolomite distribution in Pennsylvania. Map 15. Topographic and Geologic Survey,
Harrisburg.

ArGus, G. W., K. M. PRYER, D. J. WHITE, AND C. J. KENNEDY, eds. 1982-1987. Atlas of the rare vascular plants of
Ontario. Four parts. Natl. Mus. Nat. Sci., Ottawa. (looselea

BartToIs, R. L. 1985. ‘ eee glade in West Virginia. Bartonia 51: 34-36.

DG E. 1984. Marl wetlands in eastern West Virg distributi plant species, and recent

history. Caines 49: 17-25.

BASKIN, J. M. AND C. C. BASKIN. 1974. Some aspects of the ecology of Ophioglossum engelmannii in the cedar
glades of Kentucky and Tennessee. Amer. Fern J. 64: 65-73.

———anp———. 1978. Plant ecology of cedar glades in the Big Barren region of Kentucky. Rhodora 80:
545-557.

——anp———. 1981. The Big Barrens of Kentucky not a part of Transeau’s Prairie Peninsula. Pages 43-48 in
R. L. Stuckey and K. J. Reese, eds. The Prairie Peninsula—in the “shadow” of Transeau. Biol. Notes No. 15.
Ohio Biol. Surv., Columbus

———_AND———.. 1986. Distribution and geographical/evolutionary relationships of cedar glade endemics in
southeastern United States. ASB Bull. 33: 138-1

BAZZAZ, F. A. AND J. A. D. PARRISH. 1982. Organization of grassland communities. Pages 233-254 in J. R. Estes,
R. L. Tyrl, and J. N. Brunken, eds. Grasses and grasslands: systematics and ecology. Univ. of Oklahoma
Press, Norman.

BENson, L. 1948. A treatise on the North American Ranunculi. Amer. Midl. Naturalist 40: 1-261.

——. 1982. The cacti of the United States and Canada. Stanford Univ. Press, Stanford, Cal.

BERG, T. M., et al. 1980. Geologic map of Pennsylvania. Pa. Dept. Environ. Res., Harrisbur

oP H. BARNES, W. D. SEVON, V. W. SKEMA, J. P. WILSHUSEN, AND D. S. YANNACCI. 1989. Physiographic

provinces of Perma! Map 13, Pa. Topogr. Geol. Surv., Harrisburg.

AND C DGE. 1981. Atlas of preliminary geologic quadrangle maps of Pennsylvania. Map 61, Pa.

Geol. Surv., 4th Ser., Harrisburg.
BILLINGTON, C. 1949. Shrubs of Michigan. 2nd 265 Bull. Cranbrook Inst. Sci. No. ri Bloomfield Hills.

CKWELL, W. H., JR. 1977. A hybrid R Castanea
Boone, D. D. 1984. Rare and Maryland plants associated with limestone formations. Pages e214
in A. W. Norden, D. C. Forester, and G. H. Fenwick, e

Maryland. Maryland Natural Heritage Program, Special Publication 84-1, Md. Dept. Nat. Res., Annapolis.

38 BARTONIA

BRAUN, E. L. 1961. The woody plants of Ohio. Ohio State Univ. Press, Columbu
ae eas forests of eastern North America. Facsimile of the sition of 1950. Hafner Publishing
ork.

Co..

BREDEN, = i 1989. A preliminary natural community classification for New Jersey. Pages 157-191 in E. F.
Karlin, ed. New Jersey’s rare and endangered plants and animals. Institute for Environmental Studies,

mapo College, Mahwah.
Brizicky, G. K. 1962. The genera of Rutaceae in the southeastern United States. J. Arnold Arbor. 43: 1-22.
964. The genera of Rhamnaceae in the southeastern United States. J. Arnold Arbor. 45: 439-463.

BROCKETT, B. L. AND T. S. COOPERRIDER. 1983. The Primulaceae of Ohio. Castanea 48: 3740.

Brown, M. L. AND R. G. BRown. 1984. Herbaceous plants of Maryland. Port City Press, Baltimore.

Brown, R. G. AND M. L. BRown. 1972. Woody plants of Maryland. Port City Press, Baltimore.

Brush, G. S., C. LENK, AND J. SMITH. 1980. The natural forests of Maryland: an explanation of the vegetation
map of Maryland. Ecol. Monogr. 50: 77-92.

CAMPBELL, E. O. 1970. Morphology of the fungal association in three species of Corallorhiza in Michigan.
Michigan Bot. 9: 108-113.

Carr, L. G. ey 1965. Floristic elements in southwestern Virginia I hical ideration. Castanea
30: 1

tae 4 o Ss sa

CASE, F. W., i 1987. Orchids of the Western Great Lakes Region. 2nd ed. Bull. Cranbrook Inst. Sci. No. 48,
Bloom field Hills.

CLARK, J. H. 1970. Geology of the ea rocks in western Franklin County, Pennsylvania. Progress Report
180. Topogr. Geol. Surv., Harris’

CLAUSEN, . - 1975. Sedum of North iter north of the Mexican Plateau. Cornell Univ. Press, Ithaca.

Coss, B. 1 A field guide to the ferns. Houghton Mifflin Co., Boston

COOPER, i ; ps 1903. The tradition concerning our limestone lands. Pages 74-93 in Papers read before the
Kittochtinny Historical Soc. from February 1899 to February 1901. Public Opinion Print, Chambersburg.

Core, E. L. 1966. Vegetation of West Virginia. McClain Printing Company, Parsons

CorRELL, D. S. 1950. Native orchids of North America north of Mexico. Chronica Botanica Company, Waltham.

CronauisT, A. 1980. Vascular flora of the soil United States. Vol. I. Asteraceae. Univ. of North
Carolina Press, Chapel H

Curtis, J. T. 1959. The ctieteon ee of Wisconsin: an ordination of plant communities. Univ. of Wisconsin Press,

Madison.

Cusick, A. W. 1984a. Corallorhiza wisteriana Conrad, spring coral-root. Page A-150 in R. M. McCance, Jr. and
J. F. Burns, eds. Ohio endangered and threatened vascular plants: abstracts of state-listed taxa. Dept. Nat
Res., Columbus

a, 1984b. Cnbdenlosune engelmannii Prantl, limestone adder’s-tongue. Page A-7 in R. M. McCance, Jr. and
J. F. Burns, eds. Ohio endangered and threatened vascular plants: abstracts of state-listed taxa. Dept. Nat.

Res., Columbus

. 1990. Danibuton and current status of three state-listed goldenrods (Solidago: Asteraceae) in
Petinsstvaeins Bartonia 56: 9-12.

Davies, W. E. AND H. E. LEGRAND. 1972. Karst of the United States. Pages 467~505 in M. Herak and V. T.
Stringfield, eds. Important karst regions of the Northern Hemisphere. Elsevier Publishing Co., New York.

Dea, C. C. 1970. Flora of Indiana. Reprint of 1940 edition. J. Cramer, New York.

aoe NATURAL HERITAGE INVENTORY. 1991. Plants of special concern. Dept. Nat. Res. Environ. Contr.,
Dov

DESELM, . R. 1989. The barrens of Tennessee. J. Tenn. Acad. Sci. 64: 89-95.

DuppstapT, W. H. 1972. Flora of Bedford County, Pennsylvania. I. Plant communities. Castanea 37: 86-94.

Evers, R. A. ao me pees of rar Ill. Nat. Hist. Survey Bull. 26: 366-446.

FASSETT, N. C. 19: the herbarium of the University of Wisconsin—IV. fp 31: 49-53.

. 19351. Sia from the herbarium of the University of Wisconsin—VII. Rhodora 33: 224-228.

. 1944. Dodecatheon in eastern North America. Amer. Mid]. Naturalist 31: 455-486.

FERNALD, M. L. 1939, Last survivors in the flora of Tidewater Virginia. Rhodora 41: 541-544 (on Ranunculus).

. 1950. Gray’s manual of botany. 8th ed. American Book Co., New York.

FINAFROCK, J. L. 1942. Notes on Franklin County History. Kittochtinny Historical Society, Chambersburg.

FowELLs, H. A. 1965. Silvics of forest trees of the United States. Agric. Handbook 271, U.S. Dept. Agric.,
Washington.

GEYER, A. R. AND J. P. WiLSHUSEN. 1982. Engineering characteristics of the rocks of Pennsylvania. 2nd ed.
Environmental Geology Report 1. Pa. Geol. Surv., 4th Ser., Harrisburg.

LIMESTONE PLANTS 39

GLEASON, H. A. 1922. The vegetational history of the Middle West. Ann. Assoc. Amer. Geogr. 12: 39-85.
1 The new Britton and Brown No flora of the northeastern United States and adjacent

Canada. Three vols. Hafner Press, New

AND A, ee 1964. The natural ae) of plants. Columbia Univ. Press, New York.

AND . 1991. Manual of nia plants of the northeastern United States and adjacent Canada. 2nd
ed. New ms epee Garden

GORDON, R. B. 1969. The natural er of Ohio in pioneer days. Ohio Biol. Survey Bull., New Ser. III(2),
Ohio State at Columbus

GREAT PLAINS FLORA ASSOCIATION. apis Flora of the Great Plains. Univ. Press of Kansas, Lawrence.

GRIMSHAW, S. AND T. R. oer sale 973. The vascular flora of Great Falls National Park, Fairfax County,
Virginia. Castanes 38: 229-26
HANKS, S. AND D. E. FAIRBROTHERS. 1969a. una and associations of Opuntia compressa (Salisb.) Macbr. in

New very Bull. ng Bot. Club 96: 592
69b. Diversity of de fed of Opuntia compressa (Salisb.) Macbr. in New Jersey.

Rhodors 96: oat-2

HARVILL, A. M., C. E. STEVENS, AND D. M. E. Ware. 1977. Atlas of the Virginia flora, part 1: pteridophytes
through sere Virginia Botanical Associates, Farmville.

——, T. R. BRADLEY, C. E. STEVENS, T. F. WIEBOLDT, D. M. E. WARE, AND D. W. OGLE. 1986. Atlas of the
Virginia flora. 2nd ed. Virginia Botanical Associates, Farmville.

Henry, L. K. 1978. Vascular flora of Bedford County, Pennsylvania: an annotated checklist. Carnegie Museum of
Natural History, Pittsburgh

HILL, S. R. 1988. New plant records for Maryland with an additional note on Nymphaea tetragona (Nymphaea-
ceae) pollination. Castanea 53: 164-166.

Hitcucock, A. S. 1971. Manual of the grasses of the United States. 2nd ed. Revised by Agnes mcs U.S. Dept.
Agric. Pigs Ph No. 200. 1950 Dover unabridged republication. Dover Publications, New Y

ILTis, H. M. SHAUGHNESSY. 1960. Preliminary reports on Bi flora of wis No. 43.
Prim mules primrose family. Wisc. Acad. Sci. Arts Letters 49: 113-13

JOHNSTON, L. A. 1975. Revision of the Rhamnus serrata complex. Sida 6: ne

KARTESz, J. T. AN D R. KartTesz. 1980. A synonymized checklist of oe vascular flora of the United States,

Canada, and Greenland. Univ. of North Carolina Press, Chapel
KEENER, C. S. 1976. Studies in the Ranunculaceae of the Oe aes Pale: States. V. Ranunculus L. Sida 6:
Z 83.

66-2
———. 1983. Distribution and biohistory of the endemic flora of the mid-Appalachian shale barrens. Bot. Rev.
49: 65-115.
— AND M. M. — 1986. An overview of the vascular plant geography of Pennsylvania. Pages 111-144 in
K r, F. J. Brenner, and A. F. Rhoads, eds. Endangered = Narencoaths species a ne in
Pennsylvania se other states: part issues and management. Penns
OTZ,. sb, D J 1990. isa records of icin eateeat \Eiydroplothim
macrophyllum, Shotts tan Bartonia 56:
LA, O. AND R. W. Lona. 1976. Ferns of Florida: an illustrated manual and identification guide. Banyan

Books, Mia
LELLINGER, D. B. i9as. A field oe of the ferns and fern-allies of the United States and Canada. Smithsonian
Institution Press, Washingto
Lona, R. S. 1975. Soil survey of Franklin County, Pennsylvania. U.S. Dept. Agric., Washin
Lona, R. W. 1961. Convergent patterns of variation in Ruellia caroliniensis and R. cae ier Bull.
agent Bot. Club 88: 387-396
0. The = igs of Acanthaceae i in Sigel ite einai beeps States. a: ae Arbor. 51: 257-309
eee ‘i W. 197 ( | ). Bull. Torrey Bot. Club 97:
150-153.

M’cAULEY, I. H. 1878. Historical sketch of Franklin panies Rena NONA. D. F. Pursel, Chambersburg.

McCoy, S. 1937. A cleistogamous Ruellia. Amer. Botan 24.

ae J. T. 1979. How to know the ferns and fern ra ae C. Brown Co., Dubuque

D J. M. BEITEL. 1988. Pteridophyte flora of Oaxaca, Mexico. Memoirs New York Botanical Garden,
Vo e 46. New York Botanical Garden, Bro

MITCHELL, R. S. 1986. A checklist of New York State plants. Bull. 458, New York State Museum, Alban

MARYLAND NATURAL HERITAGE PROGRAM. 1991. Rare, threatened, and endangered plants of Maryland. Dept.
Nat. Res., Annapolis

40 BARTONIA

yanargorinies: R. H. 1967. The illustrated flora of Illinois: ferns. Southern Illinois Univ. Press, Carbondale.
8. The illustrated flora of Ilinois: flowering plants, hollies to loasas. Southern Illinois Univ. Press,
Carbondale
——.. 1981.

a5 + ig £ Tn = gq : 1 + | + ae

gp g p plants. Southern Illinois Univ.
Press, Carbondal
. 1982a. The stated flora of Illinois: flowering plants, basswoods to spurges. Southern Illinois Univ.
Press, Carbonda
. 1982b. Woody oe of the Shawnee National Forest (Illinois). Castanea 47: 347-359.

AND J. W. VoicrT. 1959. A flora of southern Illinois. Southern Illinois Univ. pie Carbondale.

Morse, L. E. 1984. Some rare plants of Maryland’s shale barrens. Pages 215-218 in A. W. Norden, D. C.
Forester, and G. H. Fenwick, eds. Threatened and endangered plants and erat of Maryland. Maryland
Natural Heritage Program, Special Publication 84-1, Md. Dept. Nat. Res., Annapolis.

THE NATURE CONSERVANCY. 1982. Natural Heritage Program operations manual. The Nature Conservancy,

New JERSEY NATURAL HERITAGE PROGRAM. 1990. Special plants of New Jersey. N.J. Dept. Environ. Protect.,
Trenton.

OuI0 DivisiON OF NATURAL AREAS AND PRESERVES. 1990. Rare native Ohio plants: 1990-91 status list. Ohio

ept. Nat. Res., Columbus.

O’NEILLL, B. J., JR. 1964. Atlas of Pennsylvania’s mineral resources. Part 1. Limestones and dolomites of
Pennsylvania. Bull. M50. Pa. Geol. Surv., 4th Ser., Harrisbur.

Orr, J. G. 1904. The traditions relating to the barrens of the limestone lands of the Cumberland Valley, with

special reference to Franklin County. Pages 18-31 in Papers read before the Kittochtinny Historical Soc.
from March 1901 to February 1903. Repository Printing House, Chambersburg.

PENNSYLVANIA DEPT. OF ENVIRONMENTAL RESOURCES. 1987. Title 25, Chapter 82: Conservation of Pennsylvania
native wild plants. Pennsylvania Bull. 17-5027, Harrisbur,

PENNSYLVANIA NATURAL DIVERSITY INVENTORY. 1983. Chigattiontines of natural communities in Pennsylvania.
Unpublished draft. The Nature Conservancy, Western Pennsylvania Conservancy, and Pa. Dept. Environ.
Res., Middletown

—-. 1991. Plants of special concern. The Nature Conservancy, Western Pennsylvania Conservancy, and Pa.

Dept. Environ. Res., Middletow:
POHL, R. W. 1947. A testniie secs on the grasses of Pennsylvania. Amer. Midl. Naturalist 38: 513-604.
PorTER, T. C. 1872. Botany. Pages 25~26 in H. F. Walling and O. W. Gray, eds. New topographical atlas of the
state of Pennsylvania with descriptions historical, scientific, and statistical. Stedman, Brown, and Lyon,
Philadelphia.

1903. Flora of Pennsylvania. Edited by J. K. Small. span & ss Peete.

QUARTERMAN, E. 1989. Structure and dynamics of the | ities in Tennessee. J. Tenn.
Acad. Sci. 64: 155-1

RADFORD, A. E., H. E. AHLES, AND C. R. BELL. 1968. Manual of the vascular flora of the Carolinas. Univ. North
Carolin Press, Chapel Hill.

RAWINSKI, T. J. 1990. A classification of Virginia’s indigenous biotic communities: phase 1. Upper levels of the
hierarchy. Review draft. Dept. Conserv. Recr., Richmond.

REED, C. F. 1953. The ferns and fern-allies of Maryland and Delaware including District of Columbia. Science
Press, Lancaster.

RESCHKE, C. 1990. Ecological communities of New York State. New York Natural Heritage Program, Latham.

Ruoabs, A. F, 1986. Rare plants of eastern Pennsylvania. Pages 103-110 in S. K. Majumdar, F. J. Brenner, and
A. F. Rhoads, eds. Endangered and threatened species programs in Pennsylvania and other states: causes,
issues and management. Pennsylvania Academy of Science, Easton

. 1989. Endangered and seme bse of Pennsylvania wetlarids. Pages 139-146 in S. K. nan
R. P. Brooks, F. J. Brenner, an er, Jr., eds. Wetlands ecology and conservation: emphasis i
Pennsylvania. Pennsylvania as Me cig Easton

RICHARD, J. F. 1887. History of Franklin County. Pages 137-614 in S. P. Bates, J. F. Richard, et al. History of
Franilin County, Pennsylvania. Warner, Beers and Co., Chicago

RIEFNER, JR., R. E. AND S. R. HILL. 1984. Ecology and floristics of limestone areas: an important habitat for
Maryland plants. Pages 161-193 in A. W. Norden, D. C. Forester, and G. H. Fenwick, eds. Threatened and
endangered plants and animals of Maryland. Maryland Natural Heritage Program, Special Publication 84-1,
M Res., Annapolis.

Romeo, P. A., S. L. STEPHENSON, AND R. B. CLARKSON. 1987. Systematics of Opuntia (Cactaceae) in the central
Appalachians. Bull. Torrey Bot. Club 114: 8-12.

ee

LIMESTONE PLANTS 41

RosatTi, T. J. 1989, The a. of eo RAN peel (Apocynaceae and Asclepiadaceae) in the
southeastern United Sta d Arbor. 70: 443-514.

SCHWEGMAN, J. E. 1984. The pai shooting star or anes amethystinum) in eae Castanea 49: 74-82.

ScoGGAN, H. J. 1978-1979. The flora of Canada. Four parts. Nat. Mus. Canada, Otta

SHAVER, J. M. 1954. Ferns of Tennessee. George Peabody College for Teachers, Nashville

SHEVIAK, C. J. 1974. An introduction to the ecology of the Illinois Orchidaceae. Scientific Paper XIV, Illinois
State Mus., Springfield.

SMALL, J. K. 1933. Manual of the southeastern flora. Univ. North Carolina Press, Chapel Hill.

———.. 1964. Ferns of the southeastern states. Facsimile of the edition of 1938. Hafner Publishing Co., New

York.

SNYDER, D. B. 1986. Rare New Jersey plant species rediscovered. Bartonia 52: 44-48,

SOBEL, K. C. 1974. Systematics of three species of Ranunculus (Ranunculaceae). M. A. Thesis, Univ. North
Carolina, Chapel Hill.

SPEIR, R. 1967. Soil survey of Adams County, Pennsylvania. U.S. Dept. Agric., Washington

SPONGBERG, S. A. 1978. The genera of Crassulaceae in the southeastern United States. : Arnold Arbor, 59:
197-

248.

STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames

STRAUSBAUGH, P. D. AND E. L. Core. 1977. Flora of West Virginia. 2nd ad. Seneca Books, Grantsville.

StuEssy, T. F. 1977. Revision of Chrysogonum (Compositae, Heliantheae). Rhodora 79: 190-202.

TERRELL, E. E. 1970. ea flora of the Chesapeake and Ohio Canal Area, Washington, D.C. to Seneca,
Maryland. Castanea 35: 1

THOMPSON, H. J. wigs The pa ieciadslics of Dodecatheon. Contr. Dudley Herb. 4: 73-154.

TURNER, B. H. AND E. QUARTERMAN. 1968. Ecology of Dodecatheon meadia L. (Primulaceae) in Tennessee
glades and inet Ecology 49: 909-915.

SG SuRVEY. 1979. Franklin Co., Pa. (topographic map) U. ni Geol. Surv., Reston.

VIRGINIA Saher Sgirchaps in PROGRAM. 1990. Rare plants of Virginia. Dept. Conserv. Recr., Richmond.

VoicT, J. W. AND R. H. MOHLENBROCK. 1964. Plant communities of southern Illinois. Southern Illinois Univ.
Press, Cishanditi e:

Vokes, H. E. 1957. Geography and geology of Maryland. Bull. 19. Dept. Geol. Mines Water Res., Baltimore.

Voss, E. G. 1972. Michigan flora, part I: gymnosperms and monocots. Bull. Cranbrook Inst. Sci. No. 55,
Bloomfield Hills and Univ. Michigan Herbarium, Ann Arbor.

———. 1985. Michigan flora, part II: dicots ee Bull. Cranbrook Inst. Sci. No. 59,
Bloomfield Hills and Univ. Michigan Herbarium, Ann Ar

WALCK, J. L. 1987. Shale barren ragwort (Senecio ssenbyancoa Asteraceae) in the Great Valley, Franklin
County, Pennsylvania. Bartonia 53: 13-14

WALDRON, G. E., S. W. ABouD, J. D. AMBROSE, AND G. A. MEYERS. 1987. Shumard oak, Quercus shumardii, in
Canada. Canad. Field-Nat. ate 532-538.

WEAVER, J. E. 1968. Prairie plants and their environment: a fifty-year study in the Midwest. Univ. Nebraska
Press, Lincoln.

WEST VIRGINIA NATURAL HERITAGE PROGRAM. 1990. State element list: plants. Div. Nat. Res., Elki

WESTERFELD, W. F. 1961. An annotated list of vascular plants of Centre and Huntingdon Counties, secre te
Castanea 26: 1-80.

Wuerry, E. T. 1926. A new circumneutral soil prickly-pear from the Middle Atlantic States. J. Wash. Acad. Sci.

42
———. 1961. The fern guide. Doubleday & Co., Garden Ci
———. 1964. Our Northeastern cacti: one species or three? Morris Arbor. Bull. ek ve
, J. M. Foca, JR., AND H. A. WAHL. 1979. Atl y rris Arb

of Pennsylvania, Philadelphia.

Worrorp, B. E. 1989. Guide to the vascular plants of the Blue Ridge. Univ. of Georgia Press, Athens.

ZIkA, P. F., ed. 1990. New York rare plant status list. New York State Dept. Environ. Conserv., Albany.

+ f eho T Ins
Willy.

Bartonia No. 57, Supplement: 42-49, 1993

Designing and Implementing a Riverine Preserve
for the Endangered Ptilimnium nodosum:
Sideling Hill Creek as a Case Study

RODNEY L. BARTGIS

Maryland National Heritage Program, MD Dept. Natural Resources, Tawes State Office Bldg.,
Annapolis, MD 21401.

DAviID MADDOX
The Nature Conservancy, 1815 North Lynn Street, Arlington, VA 22209

A frequent criticism of research and recovery efforts for endangered species is that they
tend to focus on a single species and consequently fail to address concerns about other
species or functions of the landscape in which that species occurs (Grumbine 1990, Hutto
et al. 1987). Landscape and ecosystem features may often be critical to long-term
persistence of the critical microsite. Consequently, even if a protection strategy focused on
a single species is pursued, a broader view of habitat may be warranted.

Endangered riparian plant species exemplify these issues and therefore are particularly
difficult to protect. Events critical to the survival of the species may occur at considerable
distances from individual plants. Also, and perhaps consequently, riparian populations
and habitats are often dramatically dynamic (Bowles and Apfelbaum 1989, Menges 1990).
We suspect that such remote, ecosystem-level effects on populations probably occur in
many habitats; in riparian systems they may simply be more dramatic and apparent.

Recent studies we have completed on the demographics, life history and habitat
requirements of the endangered riparian plant Ptilimnium nodosum demonstrate the
importance of landscape scale factors to the conservation of endangered species. In this
paper we will discuss Sideling Hill Creek, MD, and its population of Prilimnium, as a case
study to encourage similar approaches elsewhere.

Sideling Hill Creek is a 40 km long stream which originates in Bedford and Fulton
counties, PA, and flows south forming the boundary between Allegany and Washington
counties, MD. It meets the Potomac River about 130 km northwest of Washington DC.
Approximately 80% of the watershed in Maryland is forested, agricultural or residential
use is limited. The stream is within the Ridge and Valley physiographic province and, in
Maryland, crosses Devonian shales and sandstones of the Chemung and Hampshire
formations as well as Quaternary alluvium and terrace gravels (Edwards 1978). The
topography is rolling to steep over much of the watershed. In Maryland, the stream
elevation drops from 204 m at the Pennsylvania line to 125 m at the Potomac; the highest
Maryland elevation in the watershed is 536 m.

Ptilimnium nodosum is a short-lived member of the Apiaceae. It occurs in riparian and
wetland habitats in Maryland, West Virginia, North Carolina, Georgia, Alabama and
Arkansas. It was listed as an endangered species by the federal government in 1988
because of habitat loss from destruction of wetlands, impoundments, shoreline develop-
ment, and declines in water quality (US Fish and Wildlife Service 1988). In the Potomac
River watershed the species is currently known only from Sleepy Creek and the Cacapon
River in West Virginia and Sideling Hill Creek and Fifteen Mile Creek in Maryland.

42

PTILIMNIUM NODOSUM 43

Historically it also occurred along the Potomac River in Maryland and probably along the
Shenandoah River in West Virginia (US Fish and Wildlife Service 1991).

PTILIMNIUM LIFE HISTORY AND DEMOGRAPHICS

In 1988 we initiated a series of studies on the life history, demographics, and habitat of
Ptilimnium nodosum in order to better define threats to the species and conservation
needs. Here, we briefly review those findings which are guiding habitat protection efforts
on Sideling Hill Creek.

Ptilimnium occurs along the stream in relatively discrete stands of one plant to
thousands. Distribution is not uniform, with some areas supporting many stands and other
areas having hundreds of unoccupied meters between stands. Ptilimnium nodosum
occupies cobblestone shoals, gravelbars, and bedrock outcrops which are seasonally
flooded but above water for part of the summer. Shoals co-dominated by Justicia americana
support the largest stands of Ptilimnium, but Ptilimnium is restricted to a much narrower
range of water depth than Justicia (Maddox and Bartgis 1992), indicating the species is
poorly suited to changes in stream flow. Ptilimnium is usually absent from areas with
significant silt accumulation. Both field transplant studies and greenhouse experiments
indicate that sedimentation can reduce the vigor and success of Ptilimnium (Maddox and
Bartgis unpublished data).

In July, about when water levels are lowest, Ptilimnium plants develop new basal leaves
and flower from leafy stems 10 to 50 cm tall. By September the seeds germinate, either on
the inflorescence or after falling. During high water levels in late summer flowering plants
are knocked over by the flowing water. Rootlets and leafy shoots form at nodes of repent
flowering stems, providing for asexual recruitment. Although asexual recruitment appears
to occur frequently, sexual recruitment rates vary from year to year in response to water
regimes. During the winter young and adult Ptilimnium nodosum persist under water as
basal rosettes, although stands may be exposed briefly during periods of low water
(Maddox and Bartgis 1989, 1992).

Stand size distribution is considerably skewed. Over a third of all stands are small (less
than a hundred individuals), although big stands (over 1000 individuals) are not
uncommon. We monitored 56 locations along Sideling Hill Creek which supported a stand
of Ptilimnium for at least one year during the period 1988-1990. During this period the
number of stands increased, but the mean stand size stayed about the same. However, only
43% of the locations with stands in 1988 had stands in 1990. Nearly all stands which were
lost during this period were small, with fewer than 100 individuals (Maddox and Bartgis
1992). Overall, the population of Ptilimnium nodosum on Sideling Hill Creek is relatively
large and stable, with an estimated total population size of 25,000 to 35,000 individuals
during the study period.

OTHER BIODIVERSITY CONCERNS

In addition to the large population of Ptilimnium, the Sideling Hill Creek valley is also
noteworthy for other biological diversity resources. The creek and its surroundings support
a number of rare species populations and high quality natural communities in both
Maryland and Pennsylvania (Table 1). In both states the slopes adjacent to the stream
include mid-Appalachian shale barrens, which support a number of endemic plants
(Keener 1983). In Maryland the Sideling Hill Creek barrens support a variety of state en-
dangered and threatened plants, including Prunus allegheniensis, Woodsia ilvensis, Melica

— BARTONIA

nitens, Adlumia fungosa, Minuartia michauxii, and the endemics Trifolium virginicum and
Taenidia montana. The adjacent diverse forest communities also support a variety of
threatened and endangered species including Polygala polygama, and the only known
Maryland population for the Appalachian endemic Calamagrostis porteri (Bartgis 1991).
Floodplain forests support Cacalia suaveolans and the largest population known in the
state of Iris cristata. The upland and many of the floodplain forests along the creek are
remarkably free of alien weeds.

Sideling Hill Creek itself is relatively unpolluted (Duigon and Dine 1991) and supports a
diverse native freshwater mussel fauna including the largest population known in the state
of the globally uncommon Lasmigona subviridus, the globally uncommon Elliptio producta,
and several other state-rare mussels (Ta

Farther east in Maryland upland forests have been heavily fragmented by human
land-use patterns, resulting in a decline in bird species which require large blocks of forest
land to maintain viable breeding populations (Whitcomb et al. 1981). The Sideling Hill
Creek watershed includes some of the largest blocks of unbroken forest in this part of
Maryland. Most forest-interior dwelling bird species of the Ridge and Valley breed in the
area.

CONSERVATION ISSUES

Realistically, certain potential goals of landscape-level conservation are not attainable.
For example, corridors connecting areas of suitable habitat have been suggested for
several reasons, including maintaining movement pathways in anticipation of long-term
climate change (Hunter et al. 1988). Unfortunately, it is unlikely that a corridor which
effectively connects Sideling Hill Creek with other streams can be created since
experimental transplants suggest that the Potomac River no longer provides suitable
habitat for the species (Maddox and Bartgis 1989).

The high turn-over rates among Ptilimnium nodosum stands indicate that protection
efforts aimed at protecting specific stands, especially smaller ones, will probably be
unsuccessful. Although turn-over rate may vary among streams, we suspect that they will
be fairly high in most streams within the Potomac watershed. On Sideling Hill Creek, less

TABLE 1. Rare species of the Sideling Hill Creek watershed in Maryland. Status ranks are defined by Maryland
Natural Heritage Program (1991a, 1991b), except for mussels which are partially based on more recent
information.

Butterflies: Satyrium edwardsii (State Endangered); Euchloe olympia (State Rare); and Fixsenia ontario (State
Endangered

Mussels: Alasmidonta undulata (State Rare); Alasmidonta varicosa (State Endangered, Federal Candidate);
Lasmigona subviridus (State Endangered, Federal Candidate); Elliptio producta (State Rare); and Strophitus

undulatus (State Rare).

Vertebrates: anne floridana (In Need of Conservation (State), Federal Candidate); and Dendroica fusca
(State Threatene

Plants: Adlumia fungoss (State Threatened); Amelanchier spicata (State Threatened); Astragalus canadensis
(State Endangered); Cacalia suaveolans (State Endangered); Calamagrostis porteri (State Endangered); Calyste-
gia cara. (State Rare); Carya laciniosa (State Endangered); Cyperus houghtoni (Status Uncertain, State);
Euphorb tusata (State Endangered); Iris cristata (State Endangered); Liatris turgida (State Endangered);
Melica nitens (State Threatened); Minuartia michauxii (State Threatened); Polygala ponies (State Threatened);
Polygala senega (State Threatened); Prunus alleghaniensis (State Endangered); Ptilimnium nodosum (State
Endangered, Federal Endangered); Solidago hispida (State Endangered); Stellaria pil peo Endangered);

aenidia montana (State Threatened); Trifolium virginicum (State Threatened); Woodsia ilvensis (State

Threatened); and Zanthoxylum americanum (State Endangered).

PTILIMNIUM NODOSUM 45

than half of the locations which supported Ptilimnim nodosum contained plants all three
years of the study. While large stands are more stable, they are occasionally lost, or may
decline through time. A stand on Sleepy Creek, WV, declined from over 100,000
individuals in 1983 to about 10,000 individuals in 1986. Although Ptilimnium populations
have not been tracked through major storms (see below), it is likely that even large stands
are at risk of being lost during such events.

Small stands, however, probably play significant roles in overall population dynamics.
Small stands are not just less “successful” stands, they also include pioneering stands
which may become larger through time. For example, on Sideling Hill Creek, stands which
had only about 40, 60, and 100 individuals in 1988 had grown to support about 150, 750,
and 1100 individuals, respectively in 1990.

Habitat characteristics at a given site change through time as deposition and erosion
alter shoal and gravelbar areas. These events result in changes in the distribution of
Ptilimnium habitat. On Sideling Hill Creek, some sections of the creek are entrenched,
with little or no floodplain, and the channel has limited options for movement. Other
sections are on a well-developed floodplain and have undergone substantial changes in
channel location through time. Ptilimnium occurs in both landforms: stands are slightly
more frequent in entrenched sections but typically largest where there are well-developed
floodplains (Maddox and Bartgis 1992).

In 1985 Hurricane Juan had devastating effects on streams further upstream in the
Potomac watershed. Although the storm was not as severe in the range of Ptilimnium
nodosum, observations at severely impacted streams suggest habitat loss would vary
depending on the nature of the stream channel (Bartgis, pers. obs.). During severe, but not
huge, storms, entrenched segments get scoured more severely than segments with
well-developed floodplains. This may be why the largest stands of Ptilimnium are often
associated with the floodplain areas; these stands have had longer periods to grow. In huge
storms such as Hurricane Juan, scouring is severe everywhere, although sheltered bedrock
outcrops in the entrenched areas may be less severely affected.

Because of the normal dynamics of stand loss and creation, and the shifting mosaic of
habitat along the creek, we believe protected stream segments should include a mosaic of
small to large stands over a variety of stream morphologies. Furthermore, protected
segments must contain areas of appropriate habitat that do not currently contain
Ptilimnium. This will increase the probability that new stands can be created as they are
naturally lost.

Identifying potential sources of sedimentation and other pollutants has been a major
concern. If disturbed, steep slopes near the stream and along tributaries could introduce
substantial sediment to the stream system. Transplant studies of Ptilimnium have shown
that mortality was substantially greater near the National Freeway than at control sites on
the creek (Maddox and Bartgis 1992). The cause of the higher mortality is not known,
although chloride levels are substantially higher near the freeway than elsewhere.

Based upon what had been learned about Ptilimnium nodosum and Sideling Hill Creek,
we established certain criteria to help guide habitat conservation activities on the stream.
These are: (1) Protection efforts for Ptilimnium nodosum should emphasize stream
segments containing a variety of stand sizes, a mosaic of potential habitat areas, varied
stream morphology, and uplands which can be protected to prevent sedimentation.
Protecting such habitat mosaics increases the chance that Ptilimnium stands will be able to

“move” in response to shifting stream morphology. (2) Stream segments upstream from
concentrations of Ptilimnium should be considered for preservation in order to maintain

46 BARTONIA

water quality and conserve other biodiversity features. (3) Conservation programs should
not overlook activities in the watershed away from the stream itself that may impact water
quality and flow characteristics. For example, reduced stream flow would subject
Ptilimnium to extended periods of dryness, of which the species is intolerant (Maddox and
Bartgis 1992).

The effective protection of Ptilimnium requires a landscape-level approach. At Sideling
Hill Creek this landscape also includes many other biodiversity features of interest.
Consequently, conservation efforts for Ptilimnium on Sideling Hill Creek should incorpo-
rate other rare species, special community examples, and other resources such as habitat
for forest-interior dwelling birds. Where possible, examples of common communities
should also be included in conservation programs so that a full suite of natural
communities can be maintained along the stream.

IMPLEMENTING LANDSCAPE-LEVEL HABITAT PROTECTION

Existing land-use patterns along Sideling Hill Creek are currently suited to protecting
large stream segments. In Maryland, fewer than fifty privately owned tracts, averaging 25
ha, border the stream. Two large public land areas also occur along the creek, Green Ridge
State Forest and Sideling Hill Wildlife Management Area. The Maryland Department of
Natural Resources and The Nature Conservancy have begun to protect landscapes along
the stream. It was recognized from the beginning that such a program would require a
wide-range of tools, including designation and management of public lands, acquisition,
cooperation with private landowners and education.

During this study period most stands of Ptilimnium nodosum occurred on or near lands
owned by the State of Maryland (Figure 1). This area coincides with concentrations of
shale barrens, mussels, and other rare species and has a diversity of land-forms.
Consequently, this section of stream was selected as a focus for habitat protection efforts.

e state has taken several actions directed at the conservation of Ptilimnium nodosum
and other biodiversity resources on state lands. These include erosion prevention directed
at slopes which contribute run-off (and potentially sediment) to the stream, preservation
of large blocks of forest, and protection of lands which contain other rare species.

Two segments of Sideling Hill Wildlife Management Area have been designated as
Natural Heritage Areas (Figure 1). This is a regulatory designation which recognizes the
significance of the areas for biological conservation. The northern Natural Heritage Area
(Figure 1) has been nominated as a state Wildland. If approved, the designation would
offer protection similar to designation of federal lands as wilderness.

A Habitat Protection Area has been designated for the portion of Green Ridge State
Forest which borders and drains into Sideling Hill Creek. Although this administrative
designation carries no legal protection, Habitat Protection Areas define state lands on
which management actions are to be reviewed for potential impacts on biodiversity
features.

During this study a significant percentage of stands of Ptilimnium nodosum not on state
lands were located on a 320+ ha tract near the creek’s mouth. This tract also supports rare
mussels, high-quality forest communities, and shale barrens. In 1991 the Maryland
Department of Natural Resources, with the help of The Nature Conservancy, acquired
this tract using monies from the Heritage Conservation Fund (Figure 1). A youth group
and the National Guard had existing use rights to the property, so the acquisition effort
included a series of leases and management agreements to define management and use of
the tract. The highlight of this effort is the establishment of a primary protection area in

PTILIMNIUM NODOSUM

PENNSYLVANIA g
Meee Pe eee oe ee eek
| MARYLAND Se maa)
‘
f
y
(<
s
oO i
\ be o
% -
‘a ) a
/ (
——
\W
= a
4
Lo 3S od
NS % Sf
15 = Ss Pe
4
$,
\a

@
7
ee
<

~~
il

=

ES aa
Netanatereneeetou *.

end

.
“ip
Gy
Ny A

ea |
2 Km

Fic. 1. Sideling Hill Creek watershed in Maryland and lands associated with the landscape conservation

program. The hatched areas are designated Natural Heritage Areas within Sideling Hill Wildlife Management

Ar

ea. The cross-hatched area is also the proposed Sideling Hill Creek Wildland. The stippled area within the
1991 acquisition tract is the primary protectio
Administrati atura ) private |

n area. The numbered areas represent: (1) the State Highway
rea; (2) Pp f the Maryland C p tive Nat 1 AreasR gi t y; d (3) The
Nature Conservancy/ Western Pennsylvania Conservancy acquisition in Pennsylvania.

which protection of the stream, adjacent slopes, special communities, and endangered
species is the primary management goal (Figure 1)

Most private landowners along this critical section of creek have been notified of the
significance of the stream and the impacts of shoreline erosion and other activities. The
Maryland Cooperative Natural Areas Registry, a voluntary land-protection program (see
Hoose 1981) of The Nature Conservancy and Maryland Natural Heritage Program, have

48 BARTONIA

also been involved on Sideling Hill Creek. Most private stream frontage between
Pennsylvania and the National Freeway has been added to the Registry, as has a Maryland
State Highways Administration tract, which supports the northernmost known stand of
Ptilimnium nodosum (Figure 1).

Pennsylvania conservationists have also shown an interest in protecting segments of
Sideling Hill Creek in their state. Not only will this effort help protect species and
ecosystems of concern in Pennsylvania, it will hopefully help maintain stream quality
downstream. The cornerstone of the effort was the acquisition of land along the stream at
the PA-MD border by The Nature Conservancy and Western Pennsylvania Conservancy
(Figure 1).

CONCLUSIONS

Although existing land-use patterns along Sideling Hill Creek have been conducive to a
comprehensive program for landscape-level conservation, many issues remain. Progress to
date has been on establishing a conservation corridor along the stream itself. The opening
of the National Freeway across Sideling Hill Creek in 1988 may lead to land-use changes
throughout the watershed, including residential and commercial development. It will be
necessary to work with local landowners and planners to minimize impacts of changes in
land-use on water quality and hydrologic characteristics of the stream.

Currently, no activity in the Sideling Hill Creek watershed is known to be significantly
impacting water quality or flow characteristics, but potential impacts need to be identified
and anticipated. For example, an analysis of riparian habitat dynamics along the St. John’s
River in Maine indicates that timbering practices upstream in the watershed have
increased peak river stage levels concommitant with a decline in the endangered plant
Pedicularis furbishiiae (Menges 1990).

Site-specific stewardship problems also need to be identified and addressed. For
example, a significant amount of the sediment which enters the stream is from dirt roads
which border or ford Sideling Hill Creek (Maddox and Bartgis unpublished data).

To create an effective program, though, the most difficult problems will be dealing with
the complexities of protecting a riparian system at the landscape level. There is a
possibility that landowners, managers, and planners will not recognize the potential
implications of their actions on the creek beyond some distance within which the
relationship to the creek is obvious. The viability and security of Ptilimnium nodosum,
other endangered species, and the natural communities must be continuously moni-
tored. The interest of all participants involved in maintaining this landscape, including
private landowners, public land managers, and public land users, must be continuously
cultivated.

ACKNOWLEDGEMENTS

Our research was funded by the Chesapeake Bay and Endangered Species Fund, the
Maryland State Highways Administration, and the US Fish and Wildlife Service. Habitat
protection efforts have been conducted by The Nature Conservancy, the Western
Pennsylvania Conservancy, Maryland Program Open Space, and the Maryland Natural
Heritage Program. Our research activities were assisted in the field by Ashton Berdine,
Laurie Maclvor, and Johanna Thomas. Josephine Thoms prepared the figure. D. Daniel
Boone, Edward Thompson, and Charles Bier were instrumental in developing a broad
view of the biodiversity resources of Sideling Hill Creek.

PTILIMNIUM NODOSUM 49

LITERATURE CITED

BARTGIS, R. L. 1991. Cypripedium candidum and ther additions to the native flora of Maryland. Castanea
56: 220-222.

Bow Les, M. L. AND S. I. APFELBAUM. 1989. omuaicdaenanite d stochasti ts on the heart-shaped plantai
(Plantago cordata Lam.) in an Illi Nat. Areas J. 9: 9

Duicon, M. T. AND J. R. DINE. hing Water resources of Washington aig MD Geol. Survey Bull. 36.
D Dept. Nat. Res. Baltim

EDWARDS, J., JR. 1978. Geologic ae oat Washington County. William and Heintz Map Corp., Washington.

GRUMBINE, E. 1990. Protecting biological diversity through the greater ecosystem concept. Nat. Areas J. 10:

120.
Hoose, P. M. 1981. Building an ark. Island Press. Covelo.
HutTo, R. L., S. REEL, AND P. B. LANDRES. 1987. A critical evaluation of the species approach to biological
conservation. Endangered Species Update 6: 1—
KEENER, C. S. 1983. Distribution and biohistory of the endemic flora of the mid-Appalachian shale barrens. The
Bot. Rev. 49: 65-115.
Mappox, G. D. AND R. L. BARTGIS. 1989. Ptilimnium eae rei in Maryland: a progress report on
conservation and ronan — MD Dept. Nat. Res., Anna
AND 19972. s and causes of peor anaes variation ne Ptilimnium nodosum

(Apiaceae), an ee pt Unpublished report to the MD Dept. Nat. Res napolis

MARYLAND NATURAL HERITAGE PROGRAM. 1991a. Rare, threatened and endangered ae of Maryland. MD
Dept. Nat. ‘ie Vispiicicm

. 1991b. Rar d end d plants of Maryland. MD Dept. Nat. Res., Annapolis.

MENGES, E. S. 1990. Population pegged analysis for an endangered plant. Conserv. Biol. 4: 5

US FIsH AND WILDLIFE SERVIC . Endangered and threatened wildlife and plants: determination of
Ptilimnium mae bed sa an pe Speers. Federal Register 50: 37978-37982.
1991. Harperella y plan. US Fish and Wildlife Service. Newton Corner.

Wuitcoms, RAF S: ROBBINS, JS; LyNcH, B. L. WuITcomB, M. K. KLIMKIEWICZ, AND D. BysTRAK. 1981.
Effects of forest fase mn leon of the eames) deciduous pias Pages 125-205 in R. L. Burgess
and D. M. Sharpe, eds. Forest island dynamics in man-domina scapes. Springer-Verlag, New York.

Bartonia No. 57, Supplement: 50-57, 1993

Extinct, Extant, Extirpated, or Historical?
or In Defense of Historical Species

DAVID B. SNYDER

Natural Heritage Program, New Jersey Dep t of Envi | Protection Division of Parks and Forestry,
Office of Natural es pit con CN-404, Trenton, NJ, 08625

“Down the bluff I where the water had cut its banks into the limestone and here and there in the crevices of
dry limestone was Siete curtipendula growing just as Porter said it was a hundred years ago. Porter is dead
and long gone and aaas forgotten but the small grass is thrifty and annually casts its seeds despite the passing
of its discoverer . the constancy of the limestone and the Delaware River and the curious grass continues de-
spite th ye of the inconstant passer-by.” Vincent Abraitys.

Relocating a rare species at the exact location discovered many years before by a long
dead predecessor is an satan that is particularly satisfying to the field botanist.
Foremost, it creates a conservation opportunity that did not previously exist. And, as
Vincent Abraitys (1975) cena related, it is also a leveling experience for us humans,
because it puts our brief existence in its proper perspective. An individual lifespan is often
the measure and the extent of our experience of time. To us then, one hundred years
becomes a significant chunk of time, which is why we often are amazed when we hear that
someone has relocated a rare species after such a lapse of time. But considering the
thousands of years that many of New Jersey’s plant species have been in the state, is it
really all that amazing? Given this anthropocentric view of time, is it any wonder that we
tend to underestimate the tenacity that many rare and endangered plant species exhibit in
persisting in a human-altered landscape?

CLASSIFICATION OF RARE SPECIES

Unfortunately, this human-skewed bias of time has become manifested in our approach
to the classification of rare and endangered plant species. Nowhere is this more evident
then in the process of how species (or populations) are determined to be extant. Often
time is used as the single criterion in separating the extant species from the extinct, the
extirpated, or the historical. When time is used as the sole measure of the “‘extantness”’ of a
plant species, the distinctions and value of these different ranks of classification become
obscured, and even redundant.

A ranking system based entirely on time is certainly attractive in its convenience. For
example, if an extant plant is defined as having been documented within the last five years,
it is very easy to sort out the extant from the non-extant. But is this a realistic
representation? Do plant species cease to exist after the fifth year passes? Or even the
fiftieth?

Another approach is to simplify the classification system by eliminating ranks. After all,
common sense tells us that a plant species is either extant or it isn’t. Set in this context, the
rank of historical becomes a logical impossibility—obviously a plant species can’t both exist
and not exist at the same time. But the problem with this either/or approach is that it does
not take into consideration that our ability to document the simple reality of a plant’s
existence is limited and sometimes inconclusive. There is no room for uncertainties in a
classification system based entirely on yes or no. A system that forces all species to be

50

EXTINCT, EXTANT, EXTIRPATED OR HISTORICAL bs 3

pigeonholed as either extant or not aa in arbitrary decisions being made when
supporting data is lacking or ambiguo

To those insisting on absolutes, ae idea of historical species may be viewed as a
conservation contrivance—a fudge factor used in the absence of quantitative data. It
would be nice to be able to devise a ranking system based strictly on quantifiable data, but
the numbers, diversity, and biological complexities of the species under consideration,
preclude reducing data to a few, easily measured common denominators.

The dilemma then is in achieving congruency between our limited ability to document a
plant species’ existence with its absolute reality. The most unfortunate and uncomfortable
aspect of this dilemma is that what we are trying to prove is ultimately knowable, but our
resources to do so are finite. The reality of conservation biology is that time, staff, and
financial resources are all in short supply dictating that concessions be made. The
conservation rank of historical is one of these necessary concessions.

The concept of historical species allows us to tackle head-on the limitations of our
methods and abilities to document a plant’s existence. It acknowledges the uncertainties
and gaps in our knowledge without sidestepping common sense by employing convenient
short cuts that only work on paper or are valid only in theory. The need for an historical
rank becomes apparent when we try to sort out the extant species from the less than
extant—not as lists on paper, but as living organisms, each presenting a unique set of
problems.

EXTANT, EXTINCT OR EXTIRPATED? I consider a plant to be extant when its presence on
the landscape is, or can be, verified by direct experience. That is, when someone can go
out, see it, touch it, collect it, photograph it, or by any other method document its
existence. The fundamental problem in documenting a plant species extant is that since it
is always subject to destruction by natural or human caused events, its existence must be
continuously re-verified. As a result, all practical methods used to document species extant
are valid only for the specific points in time in which they were made. Mackenzie’s 1917
collection of Lechea tenuifolia! documents only that it was extant at the time he collected it;
to determine whether it is extant today requires a field survey.

Because of the uncertainty of a plant’s continued existence and the limitation of our
documentation, it is sometimes much easier to determine that a plant no longer exists,
then to prove that it does. Once destroyed, there is obviously no need to re-verify a plant’s
nonexistence. When a plant species is determined to no longer exist, it is classified as either
extinct or extirpated. In conservation biology these terms are used to distinguish species
that no longer exist in a living form anywhere on the planet (extinct) from those that are
only absent from specific portions of their geographical ranges (extirpated).

Currently, the only New Jersey species conclusively meeting the definition of extinct are
an indeterminable number of species known only from fossil or pollen records. Micranthe-
mum micranthemoides may be extinct, since no extant populations are currently docu-
mented throughout its geographical range. However, persuasive evidence of its extinction
has yet to be presented (US Fish & Wildlife Service, 1990). Narthecium americanum and
Rhynchospora knieskernii are potential candidates for extinction since both are currently
documented extant only from a small area of New Jersey’s Pine Barrens. Several
populations of these species occur on protected state-owned land so their continued
Survival is at present reasonably secure.

IN lat foll Kartesz & Kartesz (1980)

52 BARTONIA

Applied to levels or ranks of New Jersey plant endangerment, extirpated is used to
identify species which are believed to no longer occur anywhere in the state as a native wild
plant; either because known populations have been confirmed destroyed or because their
existence on the landscape is beyond practical verification.

DOCUMENTING EXTIRPATION. Documenting a species as extirpated can be a straightfor-
ward procedure as it was for Ledum groenlandicum. Ledum is a conspicuous shrub with
showy flowers that is restricted to quaking bogs or peatlands throughout glaciated portions
of North America. In New Jersey, it was documented from a single bog near Mount Hope,
Morris County, which was destroyed in the late 1970’s by the construction of a shopping
mall (Snyder 1987). True quaking bogs are extremely rare natural communities in New
Jersey and all known occurrences have been surveyed without success for Ledum. Like the
bogs themselves, Ledum was probably of relictual occurrence in the state and the
probability of new colonizations from out of state populations seems remote. For species
like Ledum, which occur in well-defined or geographically restricted habitats, field work
can be a directed, methodical elimination of esae sland and a determination of
extirpation can be made with a reasonable degree of confiden

But if a species occurs in a widespread or common habitat, this level of confidence is not
always possible. Cacalia muhlenbergii is a good example. It was collected a few times
between 1862 and 1869 from a single location along the Delaware River near Camden at a
site which has been destroyed by urban and industrial sprawl. The only habitat information
contained on specimen labels is “banks of the Delaware River.” By extrapolating general
habitat information from regional manuals and from what is known of the New Jersey
location, potential habitat can be interpreted as rich woods and thickets along rivers and
streams in the Coastal Plain. This eliminates highly urbanized areas, most of the Pine
Barrens, and probably all of the coastal strip, but it still leaves thousands of acres of
potential habitat to search. If the objective is to be able to state with absolute certainty
whether or not Cacalia muhlenbergii still exists in New Jersey, it would be necessary to
search every last square yard of potential habitat. The amount of effort necessary to
accomplish this is both prohibitive and unrealistic as a conservation goal. As far as the
preservation of Cacalia muhlenbergii in New Jersey is concerned, there is not much more
that can be done—since recent searches of appropriate areas along the Delaware River
near Camden have not located suitable habitat and more than 100 years of field work in the
general area have not been successful in rediscovering the species.

For some species, even less information is available. Arnica acaulis is known from a
single 1852 collection labeled, “Atlantic County.”’ Chasmanthium latifolium is known from
a single undated, but certainly pre-1863 (the year of its collector’s death) specimen,
labeled, “NJ” General rangewide habitat information for these species is not very helpful
in directing field work. Using Gleason’s (1952) habitat characterization for Arnica as
“sandy pine woodlands, often in damp soil’ nets you all of the Coastal Plain, while
Fernald’s (1950) “shaded slopes and wooded thickets” is virtually useless in narrowing the
search for Chasmanthium. In situations like these, biogeographical information may be of
some use, such as checking their distribution in adjacent states, but for these particular
species it is of little value. The remaining option is to conduct random searches in seemly
appropriate habitat. A formidable, and again, unrealistic task, when the amount of ground
to cover is considered. If, as in these examples, the only means of relocating a species is by
random or serendipitous field searches, then its practical conservation value has been
extinguished and a rank of extirpated is appropriate.

irpated species have no tangible significance in land conservation or regulatory

EXTINCT, EXTANT, EXTIRPATED OR HISTORICAL 53

programs; you can’t preserve what doesn’t exist. The ranking of extirpated should
therefore be reserved only for those species for which there is no realistic expectation of
their being found. Our flora is of course dynamic, and species presumed extirpated are
sometimes rediscovered. Plants continue to migrate and species are reestablished.
Overlooked populations are occasionally discovered. But these events are rare and
unpredictable, and remain beyond the bounds of practical conservation efforts.

VALUE OF ‘HISTORICAL’ CATEGORY

Because determining whether a plant species is extant or extirpated can be both a
lengthy and difficult process, many species are caught somewhere midstream—their
presence on the landscape is neither currently verifiable, nor have they been determined
extirpated. It is this circumstance that defines historical species.

The primary reason that species are classified as historical is because not all their
documented or potential localities have been adequately surveyed. An assumption is made
that with additional field work, historically ranked species can be relocated. This
assumption makes historical species an active conservation priority, and the subject of
intensive field work to identify which populations still exist.

In the past 30 years of field work in the state, a number of species that were once
presumed beyond relocation have been found at the same locations documented many
years before. For example, in 1959 Vincent Abraitys rediscovered Dicentra canadensis at a
site discovered by Mackenzie in 1922 (Snyder 1984). In 1956, Fables (1960-61) listed it as
one of his “lost plants,” stating in his preface, “possibly someone knows a station for one or
more of [these species], but this information was unknown to the compiler.” By this
statement it is apparent to me that Fables was unaware of the explicit directions to the site
contained on Mackenzie’s specimens, or if he was aware of the directions, he never used
them to relocate Dicentra canadensis.

Asimina triloba is another example of plant that was never truely lost. Britton (1889) lists
three sites in Atlantic, Hunterdon, and Mercer counties. Stone (1911) quotes the same
three localities adding, “I have been unable to find any specimens from the State or to see
the tree growing.” In 1915, Mackenzie relocated Asimina on the same Hunterdon County
island cited by Britton. By 1956, Asimina was once again a “lost plant” (Fables 1960-61). In
1965 Abraitys found the Hunterdon site, and in 1976, he relocated what is probably the
Crosswicks Creek locality cited in Britton’s catalog (Snyder 1984).

Under today’s ranking criteria, neither Dicentra canadensis nor Asimina triloba would
have been classified as extirpated since obviously neither were beyond relocation. For each
example, exact locations had been documented, both in the literature and on herbarium
specimens, but only Abraitys seems to have made use of the information in recent years.

RELOCATING HISTORICAL COLLECTION SITES. No matter how old the literature reference
or the herbarium specimen, it is often the best source of information for relocating
historical species. For example, in 1990, I found Svenson’s 1931 Passaic County location
for Pycnanthemum torrei. | relocated in 1989 Chrysler’s Watchung Mountain site for Cercis
canadensis within 30 minutes of examining his 1935 specimen at CHRB. In 1988, I
relocated New Jersey’s only locality for Schizachne purpurascens at Branchville, Sussex
County where it was last collected in 1935 (Snyder 1989). Prior to my 1986 collection, the
last collection of Geum vernum was made in 1936 at a site along the Rancocas River
(Snyder 1989). In 1910, Mackenzie noted that Vicia caroliniana was locally frequent on
Sparta Mountain; it was still frequent there in 1985 (Snyder 1986). There is nothing
extraordinary or exceptional about these relocations. All were found by using the location

54 BARTONIA

data contained on specimen labels. Since in each case I found the species on my first field
visit, I suspect that I was probably the first botanist in recent time to seriously search them
out. These species are clearly examples of historical, not extirpated, species. To have
ranked them as extirpated, would have been grossly premature. Certainly extreme lapses
of time justify reasonable doubt as to whether a species is still extant, but it is not
persuasive evidence of extirpation.

Rediscovering historical species often is not as easy as were preceding examples. I spent
many years searching documented locations for historically ranked species like Onosmo-
dium virginianum, Polygonum glaucum, Pyrola chlorantha, and Solidago rigida before
rediscovering them. Each of these species had been documented from numerous locations.
For example, Pyrola chlorantha has been collected from more than 30 different locations,
essentially statewide. Since there is no easy way of telling which historical population may
be extant, all sites may have to be searched before the species is relocated. I have been
searching for Cacalia atriplicifolia? for more iar a decade and, at this point, have only
adequately searched 12 of the 32 documented locations.

Not all historical plants are conspicuous like ne atriplicifolia, a three to six foot high
composite. If you don’t find Cacalia at a particular locality, you can be reasonably sure it is
no longer present. But with a small, inconspicuous species like Micranthemum micranthe-
moides, you never can be quite certain that you haven’t overlooked it. The search for
Micranthemum is further complicated because not only does it look like other species
occupying the same habitat, but its habitat of muddy tidal flats is both difficult and
unpleasant to search. Not only do you have to worry about sinking into tidal ooze, but you
need to time your field work to correspond to a low or outgoing tide. I suspect this is in
large part why no one has yet given up on relocating an extant population of Micranthe-
mum.

For some historical species not only is the habitat difficult and unpleasant to search, but
it can be nearly inaccessible, even outright dangerous. Dicentra eximia and Asplenium
pinnatifidum come to mind. Dicentra was discovered in 1933 on ledges and cliffs high up on
the face of Mount Tammany at the Delaware Water Gap (Edwards 1933). With the aid of
a spotting scope (and while standing firmly on solid ground at the foot of the mountain)
Vincent Abraitys confirmed a few plants extant in 1963 (Snyder 1984). It is not known to
have been observed there since and the speculation is that the construction of Interstate 80
may have impacted its habitat. My few feeble attempts at its relocation have been both
inconclusive and harrowing. The species is still extant on the Pennsylvania side of the
Water Gap, and I’m confident that with additional searching, it will be relocated at its New
Jersey location. As for Asplenium pinnatifidum, Rick Radis braved the heights and
venomous snakes when he recently relocated Edward’s 1939 station for it on the vertical
cliff face of the Kittatinny Mountain in Sussex Coun

Occasionally, a species may remain classified as historical because a documented
occurrence is on private property to which the landowner refuses access. The only
currently known population for Scleria verticillata was not re-verified extant until 1977
when Vincent Abraitys secured permission from the landowners to search for it. The

? In September, 1992, Charles and Mary Leck rediscovered this species i Burlington County along Crosswicks
Creek, near Yardville. It was growing in an moist, open thicket on the flood plain, and not in the drier woods on
the se a slopes = ine ravine as be rnao - the majority of the herbarium Specamnems, In my rhc St
reloca is
the ace of historical species obviously confound our efforts to relocate them.

ia

EXTINCT, EXTANT, EXTIRPATED OR HISTORICAL 55

current land owners are trying to sell the property and have flatly stated that they don’t
want any botanists snooping around. Scleria verticillata has been documented at this
locality for more than 100 years, and as long as its habitat remains undisturbed, I expect it
to be there for many more, whether or not any botanist gets to see it. Another example is
Linum intercursum. This species had eluded me for more than a decade. The most recent
collection for the state was a 1939 collection by Long from Burlington County. This site
had always been my first choice to search, since I had a pretty good hunch exactly where
the plant had been collected. Unfortunately, it is also the most widely known location for
the highly prized Platanthera integra, and the resulting abuses by orchid hunters as well as
nonbotanical vandals, have forced the landowner to restrict access. In August, 1990, Ted
Gordon obtained permission for us to search the site, and, within a few minutes we
relocated Linum intercursum.

The rediscovery of some historical species has been impeded by their own biological
peculiarities. New Jersey’s only confirmed extant population of Triphora trianthophora had
not been seen for many years until Tom Halliwell and I relocated it in 1979. We got the
location from Vincent Abraitys, who had been taken to its Morris County site by J. L.
Edwards, one of its discoverers. Although Abraitys was taken to the exact spot where
Edwards had last seen the plant, no Triphora could be found. Vince spent another twenty
years or so unsuccessfully trying to find Triphora there; Tom and I found it on our first
search. Triphora, like many other orchid species, is notorious for its wide population
fluctuations and, at times, produces no above ground vegetation.

Species like Rhynchospora knieskernii and Scirpus longii, both listed as “lost species” by
Fables (1960-61), are suspected seed bankers, and apparently lie dormant in the soil for
many years, until conditions again become favorable. Rhynchospora knieskernii had not
been collected at Atsion for more than 100 years, but after the great fire of 1983, Jim Stasz
located a sizable colony. After the fire, Scirpus longii was found there by the thousands
(Schyuler and Stasz 1985), where previously it was known from only a few isolated patches.

Weather patterns also affect searching for historical species. Some, restricted to Coastal
Plain pond habitats, appear only during seasons when the water levels are very low. Others
can be inventoried only in years of normal or high water levels. The Bennett Bogs in Cape
May County provides a remarkable number of examples. One historical species I’m
currently trying to relocate there is Rhynchospora rariflora, which grows in sedgy turf on the
exposed pond bottom. It has been collected or observed at Bennett Bogs at least once
every decade since it was discovered there in 1907 (Stone 1911). The last known collection
was made in 1973 by Abraitys and the last known observation was made by Gil Cavileer in
the early 1980’s (Gil’s population which was one of three documented for Bennett Bogs
may have been destroyed by some recent adverse impacts to the habitat). The 1990’s may
very well be the decade of Rhynchospora rariflora’s rediscovery, but so far it’s been a
washout; 1990 being the second consecutive year of exceptionally high water levels.

There is something extremely unsettling in watching the population of Coastal Plain
pond species decline from hundreds of plants in one year, to few or none in succeeding
years. But we must always be on guard not to allow our anthropocentric view of time to
undermine the conservation of species that display cyclical adaptations to their environ-
ment by manufacturing a conservation crisis when in actuality all may be well.

_ SEEKING NEW SITES. Searching potential habitat can be just as challenging and reward-
i d historical locations. Carex jamesii, Carex oligocarpa, Desmodium
Bie eian Linum sulcatum, and Muhlenbergia capillaris are all examples of historical
species recently relocated as a result of searching potential habitat (Snyder 1986, 1989).

56 BARTONIA

One of the greatest benefits obtained from the systematic field searching of historical
locations, is the assimilation of habitat data into a search image which allows you to more
narrowly focus future searching or to predict new locations. I rediscovered Muhlenbergia
capillaris this way. Once I had developed a feel for its traprock glade habitat at historical
collection sites on the Watchung Mountains, I began systematically searching similar sites
in the general area. In late September of 1985 I found a small population about three miles
east of where Miller had last collected it in 1918. Within two weeks, Tom Breden and I
found a second population in Passaic County, approximately 20 miles northeast of the first.
Its discovery there was purely predictive based on previous knowledge of the presence of
suitable habitat. In September of 1987, the same search image was used to relocate
Mackenzie’s 1904 Sussex County location. I had searched this location once before, but as
I later realized, I had focused on the wrong habitat.

My rediscovery of Lechea tenuifolia in the fall of 1990 succinctly underscores the
necessity of confirming through first hand field work that an historically documented
location has truly been destroyed and that no unsearched potential habitat remains in the
immediate vicinity. The only vouchered collections for this species in New Jersey had been
Mackenzie’s 1916 and 1917 specimens collected at Ringwood Junction, Passaic County.

Ringwood Junction, and much surrounding acreage, was destroyed in the late 1920’s by
the creation of Wanaque Reservoir. From the detailed directions contained in Macken-
zie’s collecting journal, it is evident that the original collection locality is now submerged
under millions of gallons of water. But neither Mackenzie’s specimens labeled “dry fields”
and “dry sunny banks,” nor Fernald’s (1950) general rangewide habitat of, “dry sandy or
rocky open woods and slopes,” describes a plant of exacting habitat requirements. Aerial
photoquads indicated that plenty of this habitat still existed near the original locality. I
further reasoned that because Mackenzie described the species as being “local” and
because so much seemingly suitable habitat still existed, the probability of finding it was
good. On my first field visit to the area, at my first stop at what seemed like a good place for
Lechea tenuifolia, I found it. Fifty-eight plants were growing on a dry rock ledge in open
woods, approximately 1.5 miles south of what once had been Ringwood Junction. Like the
Bouteloua curtipendula that Abraitys found at Porter’s last century locality, Lechea
tenuifolia is still alive and well in New Jersey.

CONCLUSIONS

It is experiences like these that have shaped and continuously reinforce my thinking on
the necessity of having a conservation rank of historical. The search for historical species
doesn’t always end successfully; species are, and will continue to be, extirpated from New
Jersey. But let’s not be premature in ranking a species extirpated before anyone has
bothered to look for it, or because some arbitrary statute of limitations has expired. If the
habitat for a plant still exists there is a more than even chance that the historically
documented plant species is still present, regardless of how much time has passed.

I am certainly not advocating that we spend conservation dollars on maybes, but only
that we recognize the conservation potential of species that have not been proved beyond a
reasonable doubt to be extirpated. Historically ranked species are among New Jersey’s
rarest plants and we should make it a priority to resolve their status as promptly as
possible. As the destruction of our natural landscape escalates, the resulting loss and
degradation of habitats and ecosystems remind us that we no longer have all the time in the
world in which to act. Several of our rare species may not survive the next ten years, let
alone the next 100 years.

EXTINCT, EXTANT, EXTIRPATED OR HISTORICAL ay

ACKNOWLEDGMENTS

I thank the following for their help in the preparation of this paper: Maude Backes, Tom
Breden, Gil Cavileer, Ted Gordon, John Konkle, Tom Halliwell, Larry Morse, Rick Radis,
Joanne Ruscio, and the curators of CHRB, NY, and PH. I want to especially thank all
those who have contributed to, and continuously work to refine The Nature Conservancy’s
Global Ranking System. The final measure of a system of classification is not whether it is
absolutely perfect, but whether it meets the objective of practical biological conservation.

LITERATURE CITED

ABRAITYS, V. 1975. Alive and well. The backyard wilderness. Columbia Publishing Co., Inc. Frenchtown.

BRITTON, N. L. 1889. Catalog of plants found in New Jersey. Geol. Surv. N. J., Final Rep. State ei 2: 27-642.

EDWARDS, J. L. 1933. Dicentra eximia at the Delaware Water Gap, New Jersey. Torreya 33: 136-13

FABLES, D. 1960-61. 20 “lost” plants. Bartonia 31: 7-10.

FERNALD, M. L. 1950. Gray’s manual of botany. 8th ed. American Book Co., New York

GLeEason, H. A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and
adjacent Canada. Hafner eye New York.

KarTEsz, J. T. AND R. KArRTESZ. 1980. A synonymized checklist of the vascular flora of the United States,
Canada, and Greenland. neces of North Carolina Press, Chapel Hill.

SNYDER, D. B. 1984. Botanical discoveries of Vineeat Abraitys, Bartonia 50: 54-56.

——. 1986. Rare New Jersey d. Bartonia 52: 44-48.

aie On baie edge of extirpation: — pereey s — hagenn’ imperiled flora. In E. F. Karlin ed., New
Jers Ins v. Studies, Ramapo College, Mahwah.

—. 1989. Notes on some recently rediscovered New aren plant species. Bartonia 55: 40-46

SCHUYLER, A. E. AND J. L. STASZ. 1985. Influence of fire on the reproduction of Scirpus longii. Bartonia 51:
105-107

~ * 4 £ Pier eer £thio Di R Ann

STONE, W. 1911. The plants of southern New Jersey, wi Pp
Rep. New Jersey State Museum, 1910, II.
US FisH AND WILDLIFE SERVICE, Dept. of the Interior. Federal Register, 1990. 55 (35): 6184-6229.

Bartonia No. 57, Supplement: 58-60, 1993
County Natural Areas Inventory

ANTHONY F. DAVIS
Pennsylvania Science Office, The Nature Conservancy, 34 Airport Drive, Middletown, PA 17057

The County Natural Areas Inventory (CNAI) program is an effort to locate, evaluate
and map the locations of rare, threatened, and endangered species, exemplary natural
communities, and other areas of significant local interest. This information can be useful
for guiding development away from the most environmentally sensitive areas through
land-use planning and zoning, and for proactive conservation efforts such as acquiring land
for nature preserves.

Much of the information on species of special concern in PA is available in the museums
and libraries of the Commonwealth and in the files of the Pennsylvania Natural Diversity
Inventory (PNDI). Since its inception in 1982, the PNDI data base has become
increasingly utilized by the public and private sectors for information about locations of
species of special concern when development or other land use change is proposed. This
increased use has pointed to the need for “upfront” information to make the process of
evaluating development proposals more efficient and less costly for all parties concerned.
The CNAIT has grown out of this need for information on the locations of rare, threatened,
and endangered species and unique natural habitats early in the decision-making process
and in a format that is useful to planners and developers. The inventory provides
government officials and public and private development agencies with one more tool with
which to make decisions for siting open space and development, and targeting land to
preserve in its natural state. Through field survey, the inventory provides biologists with
more information on biotic diversity in the county and the Commonwealth.

To date, county inventories in eastern Pennsylvania have been funded through a variety
of sources. Most prominent among these has been the Department of Community Affairs
via the Recreational Improvement and Rehabilitation Act grant program. Additional
funds have been obtained through Community Development Block Grants, the Eastern
Pennsylvania Chapter of The Nature Conservancy, county government, legislative appro-
priation, and local nonprofit conservation groups.

METHODS

Methodologies for county natural areas inventories began with surveys in Illinois (White
1978) and Indiana (Anonymous 1985). The primary focus of the CNAI is to locate and
assess exemplary natural communities, areas that are relatively undisturbed by humans or
have recovered sufficiently from prior disturbances to closely resemble their natural state,
and are of a size to maintain their integrity into the foreseeable future. Examples of
undisturbed natural communities are old-growth forest, glacial bogs dominated by
Sphagnum spp., leatherleaf (Chamaedaphne calyculata) and sedges (Carex spp.) on a
floating peat mat, or eastern serpentine barrens with their mosaic of woodland, savannah,
and prairie.

Plants and animals of special concern in Pennsylvania are included in the inventory
survey using the PNDI database as a guide for relocating old sites and for locating new
ones. When new sites are located, population status is evaluated to permit ranking the site
against other known locations for the species.

58

NATURAL AREAS INVENTORY 59

Areas of local significance are also included in the inventory. These are areas that have
been disturbed and contain no known species of special concern but do have other
attributes that make them potentially valuable for preservation. For example, relatively
large tracts of successional woodland are valuable for wildlife and low-impact recreation.
These “sites of local significance” become increasingly important in highly urbanized
landscapes.

The first step in the county inventory process is to review the PNDI data base file for the
county. The file is a compilation of historical and extant locations for the elements (natural
communities and species of special concern). Given this information, we then review aerial
photography of the county stereoscopically. Usually the county planning office is the
source of the photos, and it typically consists of 9 in. x 9 in., black-and-white prints ranging
from 2 to 8 years old. We have also been able to use large-scale, color-infrared photos in
Pike and southern Wayne counties courtesy of the US Army Corps of Engineers. Natural
communities often have distinctive “signatures” (pattern, tone, texture, reflectance, etc.)
on air photos that make them readily identifiable. Species of special concern are often
associated with particular habitat types that can be identified from the photos. Known
locations of natural communities and species of special concern are identified and these
same photo patterns are then looked for in other areas of the county. Soil and geologic
survey maps are also utilized to help identify potential locations for elements. When a
possible area is located, it is marked on a USGS topographic map as a Potential Natural
Area (PNA) and rated for the likelihood of finding elements during a field survey.

An aerial survey may be done at any stage during the inventory but the dormant season
seems to provide the most information on previous land use and forest species
composition. If the aerial photography is relatively old and the county is experiencing rapid
growth, the landscape may have changed dramatically since the time of photography.
Areas that appear to have potential on the photos may no longer exist and this can be most
easily and quickly verified from the air. A few hours in the air early in the inventory can
save a great deal of time on the ground. When the photography is recent, an aerial survey
toward the end of the field season can aid in assessing which of the remaining sites to be
surveyed actually hold much potential.

RESULTS AND DISCUSSION

New information on species and natural communities has differed with each county
studied. In those counties that have been studied for a long time or where much of the
natural habitat has been altered, little new information has been forthcoming. In counties
that may have been little studied, or where vegetation is still intact, much new information
has been uncovered. For example, in Pike and Wayne counties the number of elements
was doubled (Table 1). This may also be due, in part, to the quality of the photography
available. Recent, large-scale, color-infrared photography (used in Pike and southern
Wayne counties) provided much more information about the landscape, species composi-
tion, and subtle changes in soil wetness, etc., than did the black-and-white photography
available for most counties.

The addition of information on animal species of concern has been far more difficult to
generate in inventories designed for one field season. However, the CNAI process does
produce information for potential new occurrences for those species that have specific
habitat or food requirements. These sites can be checked in the future. Although any new
information may be too late for inclusion in the original CNAI report, addendums are
planned for each county as new information is acquired.

60 BARTONIA

ABLE 1. Pike and Wayne County element occurrences added to the on Natural Diversity Inventory
data base as the result of county natural areas inventories in those counti

Extant Element

Occurrences Pike Wayne
Total pre-1989 a2 98
Additions

Natural Communities 44 24
Plants 38 49
Animals J 0
Total Additions 87 73

To date, in eastern PA,' six inventories have been completed and five more are in
various stages of preparation. The reports we produce consist of USGS topographic maps,
each accompanied by a table and text. The maps show extant natural communities, species
of special concern, and locally significant site locations. The table lists the rarity of the
element, its protection status, quality of the site, and the date last observed. The text
describes each site, its importance, and possible threats or recommendations for
protection. Species of special concern are identified by map code only because of the
potential for over-collection or malicious destruction.

Because many of the sites are wetlands, we usually map the watershed boundary as well
as the site itself. The watershed often provides an ideal buffer zone around the element; if
the entire watershed can be protected the elements have a reasonable chance for
long-term survival. Even if the entire watershed cannot be protected, planners can take
steps to ensure that only compatible activities are allowed to take place to avoid serious
impacts to the elements. Given the inventory information, developers can avoid areas
where they are likely to encounter delays in approval for their plans, if not denial.
Developers can also improve their chances for approval of projects by incorporating
protection of rare elements located in the watershed.

In some cases, county and township governments will be able to use information derived
from the CNAI proactively to site parks and nature preserves. The information will also be
useful for local and national conservancies who need information on the priority
protection sites within their purview. Since the inventory is public, there is also potential
for increasing the awareness of local citizens to the importance and uniqueness of the land
around them.

County inventories generate more information on the elements of special concern that
groups like The Nature Conservancy are attempting to protect. New information on
locations will aid researchers in determining the true rarity and site requirements of some
species. Location information will also help researchers who may wish to study the habitat
requirements of certain species.

LITERATURE CITED
ANONYMOUS. 1985. A preliminary inventory of natural areas on the Hoosier National Forest. Indiana
Department of Natural Resources, Indianapolis. Unplub. rept.

WHITE, J. 1978. Illinois Natural Areas soir! Technical Report, Volume 1: Survey methods and results.
Illinois Natural Areas Inventory, Urban

'The Western Pennsylvania Conservancy i ducting i ies i counti

Bartonia No. 57, Supplement: 61-74, 1993

The Serpentine Barrens of Temperate Eastern North America:
Critical Issues in the Management of Rare Species
and Communities

ROGER EARL LATHAM
The Nature Conservancy, 1211 Chestnut Street, 12th Floor, Philadelphia, PA 19107

Startlingly out of place in a forested and farmed landscape, a few patches of native
prairie and savanna in southeastern P y and | Maryland attracted the
curiosity of early naturalists. They named them serpentine barrens. Further exploration of
the continent revealed that the native eastern grasslands resemble misplaced bits of the
American West. The largest and best remaining examples now lie only a few miles from
major metropolitan centers but few outside a handful of botanists, ecologists and
neighboring residents know of their existence.

Serpentine barrens are best known to botanists as hotspots for disjunct and endemic
plant species. These exceptional ecosystems are ranked high in priority for biodiversity
conservation because they are rich in rare species of both plants and animals, including
several that are globally rare and in danger of extinction. The barrens rate high in
importance also because they comprise one of the rarest and most unusual sets of natural
communities in the eastern North American temperate forest region.

Effective biodiversity conservation requires a thorough understanding of ecosystems,
communities, and key species such as those in danger of extinction. Despite sporadic
attention to serpentine vegetation by scientists, our understanding of its unusual
properties remains far from complete. Nonetheless, we have a set of working hypotheses
about how serpentine barrens communities work, how they are affected by human activity,
and what needs to be done to insure their continued well being. This presentation is a
summary of those hypotheses, including a brief review of the natural history of the
serpentine barrens of temperate eastern North America.

SERPENTINE BARRENS CHARACTERISTICS

GEOLOGY. Farmers attempting to cultivate areas of serpentine vegetation called the
land barren on learning that the thin soils were unproductive of crops and pasturage.
Serpentine is the name of the rock type underlying the barrens.

The origin of the name serpentine is commonly attributed to the long, sometimes wiggly
veins of minerals seen in some serpentine rocks. An alternative etymology points to
ancient Italy. In northern and northwestern Italy lie many outcrops, similar to those of
Pennsylvania and Maryland, of the mottled greenish bedrock. Around outcrops of the
rock, the story goes, lives a mottled greenish snake. The match in coloration is not
surprising, given the propensity for sharp-eyed birds of prey to eat snakes that do not blend
in well with the colors of their surroundings. A cryptically colored snake basking on a rock
is nearly invisible until it suddenly slithers away at the approach of a predator or a human
observer. People assumed the snake and the rock were two different phases of the same
mottled greenish substance. Since the animate phase was a serpent, they named the
inanimate phase of this apparently mystical rock serpentine.

Serpentine was highly valued as a building stone in the nineteenth and early twentieth

61

62 BARTONIA

centuries in and around Philadelphia and Baltimore because of its striking light green
color. Old serpentine buildings—typically churches, banks, or farm houses—still stand
here and there throughout the region. Noteworthy examples of buildings made mostly of
serpentine include College Hall and Logan Hall at the University of Pennsylvania,
Philadelphia, and the coincidentally named Green Library at West Chester University,
West Chester, P

Geologists call the rock serpentinite and the most abundant constituent minerals,
serpentines. Serpentine minerals are hydrated magnesium silicates, mainly chrysotile (the
most common form of asbestos) and lizardite. Serpentinites include variable quantities of
other minerals, commonly including olivine, magnetite, chromite, talc and iron oxides
(Pearre and Heyl 1960).

Serpentine minerals are formed beneath the ocean floor at fracture zones—areas
riddled with giant cracks from stresses created by the movement of oceanic crustal plates
(Saleeby 1984). Seawater leaking into deep cracks in the ocean’s foundations reacts with
the minerals olivine and pyroxene, constituents of the common oceanic crustal rock
peridotite. The hydration of olivine-rich igneous rock is a type of seieceead Seth called
serpentinization. Serpentinized peridotites are termed ultramafic—the “ma” for magne-
sium and the “f” for ferrum, Latin for iron—because magnesium is the most abundant
metallic constituent followed by iron.

Magnesium’s abundance in serpentine minerals and this metal’s strong reactivity with
certain acids explains why buildings clad in serpentine stone are particularly susceptible to
damage in urban centers. There, smoke and car exhaust enrich the air with carbon dioxide
which combines with fog and rain to form carbonic acid. The acid reacts with the stone to
form highly soluble magnesium carbonate. The problem is severe in Philadelphia:
crumbling serpentine walls at the University of Pennsylvania are routinely patched with
green-dyed concrete.

If serpentinization occurs in oceanic crust, it follows that most serpentine rock must
remain beneath the bottom of the ocean or disappear into the earth’s interior in deep
ocean trenches where the heavier ocean floor dives (subducts) beneath lighter continental
crust. maith in a process that is still debated among geologists, slivers of serpentinite
" up,” most likely between continents moving toward each other just before they
collide cee 1984). Serpentinite is far less dense than other oceanic rock and less dense
than many continental rock types because serpentinization adds oxygen and hydrogen to
the mineral structure. Serpentinite formations are buoyant relative to other rocks and float
upward during the ponderously slow but violent process of continental collision. They are
further boosted by the uplift of mountains which occurs along the sutures between
colliding continents. The upward thrust of serpentinite coupled with the rapid surface
erosion associated with mountain ranges eventually bring some serpentine rock to the
surface.

Pieces of oceanic crust perched on the continents are called ophiolites—a word coined
from the Greek words for snake and rock. Serpentinite’s low density and tendency to float
upward through other kinds of rock may be what drives the movement of ophiolites to the
continental surface (Saleeby 1984). Ophiolite emplacement has been a relatively rare
event in the earth’s history. A true-to-scale world map of continental serpentine rock
outcrops looks blank except for coastlines, at first glance. Only on close inspection does
one notice widely scattered wispy streaks and specks. Every continent has a few outcrops,
but nowhere does serpentinite occupy more than a fraction of one percent of any
continent’s surface. For example, in North America there are four extremely sparse

SERPENTINE BARRENS 63

clusters of serpentinite outcrops: Georgia to Newfoundland, Quebec to Manitoba,
Guatemala to Cuba, and central California to Alaska (Irwin and Coleman 1972).

VEGETATION. Only a tiny fraction of the area of even these small outcrops bears a
distinctive vegetation. Whether the vegetation overlying serpentine rock stands out as
different from the surrounding plant cover depends on the mineral content of the
serpentinite and on regional peculiarities of soil formation, present climate and climate

istory. The most extensive serpentine barrens in North America are those of California
and Oregon. In temperate eastern North America, serpentine barrens range from Georgia
to New York with more than 90% of the acreage lying in Pennsylvania and Maryland.

One of the first written accounts of serpentine vegetation in Pennsylvania was by John
Bartram, in a letter dated 6 December 1745 to the Dutch naturalist John Frederic
Gronovius: “The Loadstone [magnetite] lieth in a vein of a particular kind of stone that
runs near east and west for sixty or seventy miles or more, appearing even with, or a little
higher than its surface, at three, five, eight, or ten miles distance, and from ten to twenty
yards broad, generally mixed with some veins of cotton [asbestos]. The earth of each side is
very black, and produceth a very odd, pretty kind of Lychnis, with leaves as narrow and
short as our Red Cedar, of humble growth, perennial, and so early as to flower, sometimes,
while the snow is on the ground [Minuartia michauxii (Fern.) Farw., synonymous with
Arenaria stricta Michx.]; also a very pretty Alsine [Cerastium velutinum Raf.?]. Hardly
anything else grows here. Our people call them Barrens. . .” (Darlington 1849).

From the air, serpentine barrens look like islands. Most eastern serpentine barrens are
surrounded by forests or farmland. The boundaries between serpentine and non-
serpentine vegetation often appear as distinct as a true island’s shore. Few plant species
occur on both sides of these boundaries and those that do constitute an insignificant
fraction of the local biomass. The vegetation boundaries in many cases mark a bedrock
transition as abrupt as the boundary in the masonry of Penn’s College Hall between the
schist of the first story and the serpentine of the upper stories. The analogy is apt:
serpentine formations in Pennsylvania and Maryland most often adjoin schist or gneiss.

The typical natural community on schist or gneiss is a mesic forest with a nearly closed
canopy of deciduous forest trees. In contrast the quintessential serpentine barrens
landscape is a prairie of mainly native grasses with scattered trees—usually Pinus, Quercus,
or Juniperus—and exposed rock. Serpentine barrens are, quite literally, a textbook
example of the influence of soil variation on vegetation structure and composition. Plant
associations growing atop serpentine outcrops all over the world are cited by ecology,
botany, and soil science textbooks as models of azonal vegetation, that is, vegetation
resembling the norm of some other climatic zone.

Large, well developed serpentine communities are actually mosaics of different
serpentine plant associations, reflecting variation in soil development, bedrock mineral
content, moisture availability, successional age since the last episode of natural distur-
bance, and the type of disturbance (K. S. Dougherty, unpubl. ms.). The open-canopy rock
exposure called glades) and the grasslands are richest in disjunct,

| Ul Vai

endemic and rare species. Wetlands overlying serpentine are highly minerotrophic and
have a distinct flora. Several forest types also occur on serpentine including closed-canopy
Pinus rigida, Quercus stellata-Quercus marilandica oak woods and, at one location only (in
Lancaster County, PA), what appears to be a giant clone of Populus tremuloides, possibly a
relict of periglacial climate. Serpentine forests usually have a nearly impenetrable
understory of Smilax rotundifolia and Smilax glauca.

DISTRIBUTION. At the core of the eastern serpentine archipelago lie the state line

64 BARTONIA

barrens—a string of seven sites along 20 km of the Mason-Dixon Line in Chester and
Lancaster Counties, PA, and Cecil County, MD. Six are in Pennsylvania, three extend a
short distance into Maryland, and one is entirely in Maryland. Totalling approximately 850
ha, these sites comprise the largest area of serpentine vegetation in the eastern temperate
zone across which serpentine-restricted plant and insect species are likely to form locally
interbreeding populations. The two largest of the seven sites, Goat Hill and Nottingham
Barrens, lie near the geographic center of the state line barrens. Separated by less than 2
km they encompass approximately 400 ha of serpentine vegetation, mainly savanna
dominated by Schizachyrium scoparium, Sporobolus heterolepis, and Pinus rigida. Scattered
glades include the only two known eastern temperate serpentine endemics: Aster
depauperatus (Porter) Fern. (serpentine aster) and Cerastium velutinum Raf. (lumped with
C. arvense L. in most floral manuals) var. villosissimum Pennell (long-haired barrens
chickweed).

Soldiers Delight in Baltimore County, MD, a 540-ha area of serpentine vegetation 60 km
southwest of the state line barrens, is the largest single eastern serpentine barrens site. Its
flora is among the richest in species diversity of all the barrens but it lacks the endemics
and the typical state line barrens dominants Pinus rigida and Sporobolus heterolepis.

Discounting a few very small sites (<2 ha), the remaining eastern serpentine barrens
cover up to 40 ha per site and are scattered from Georgia (1 site) through North Carolina
(1) and Maryland (3) to northern Chester and Delaware Counties, Pennsylvania (6), and
New York (4 small sites on Staten Island). Altogether there are 23 serpentine barrens sites
>2 ha in temperate eastern North America. A number of small areas of serpentine
vegetation recorded historically has been destroyed by incompatible land use including
nearly a dozen sites in the western suburbs of Philadelphia and one in northern Delaware.

All temperate eastern North American serpentine barrens occur on serpentine rock in
the Piedmont physiographic province except for the one in North Carolina, which occurs
on olivine and serpentinized dunite in the Blue Ridge province (Mansberg and Wentworth

984).

DISJUNCT AND ENDEMIC SPECIES. Serpentine barrens form archipelagoes of specialized
habitats for species with disjunct and endemic distributions. The characteristic plant
species of the barrens can be classified in several ways: regional fidelity to serpentine, the
region or habitat where they are most abundant off serpentine, and regional or global
rarity. A few examples will illustrate.

One of the showiest characteristic serpentine species, Phlox subulata, is an example of a
species restricted to serpentine locally within the counties with serpentine outcrops.
Elsewhere it occurs on dry rock exposure communities including the Appalachian shale
barrens. Pink Hill, a serpentine barren at Tyler Arboretum, Delaware County, PA, takes
its name from the masses of P. subulata flowers that appear there in early spring during
favorable years. Another showy species, Lilium philadelphicum, inhabits a variety of dry,
relatively open sites, but is still considered a characteristic member of the serpentine flora.

A group with special biogeographical interest includes those species whose main range
lies in the western plains. Their populations on serpentine are generally considered as
relicts of a warmer and drier interval called the hypsithermal, which occurred from 7,000 to
2,500 years ago following the most recent glacial maximum (Deevey and Flint 1957).
During the hypsithermal, these species may have been far more abundant across central
and eastern North America. Carex bicknellii, Bouteloua curtipendula, and Sporobolus
heterolepis fall into this category. Asclepias verticillata—an unusual milkweed with almost
needle-like leaves—is also a western prairie disjunct.

SERPENTINE BARRENS 65

Fimbristylis annua, with a mainly tropical and subtropical distribution, behaves as if it
were a desert annual. Its soil seed bank suddenly germinates to form lush carpets of turf
only in years when rainfall occurs abundantly and at the right time during the season.
Pennsylvania’s serpentine populations lie at the northernmost limit of the species’ global
range. Senecio anonymus Wood (S. smallii Britt. in most floral manuals) is another species
reaching its northern limit in Pennsylvania’s serpentine barrens. Scleria pauciflora, with
fruits resembling miniature golf balls, inhabits soils and sands that are low in mineral
nutrients. All three species occur almost exclusively on serpentine in southeastern
Pennsylvania and northcentral Maryland and are classified as rare in both states.

Talinum teretifolium (fameflower) inhabits several types of rock outcrops throughout its
highly fragmented range. As is typical of the Portulacaceae, its stems and leaves are
succulent. The serpentine populations in Pennsylvania and Maryland define the northern
limit of its range. They are separated by hundreds of kilometers from the next-nearest
population on a granite outcrop in southern Virginia (W. H. Murdy, pers. comm.). How
the species’ tiny seeds found their way to nearly a dozen widely scattered serpentine
outcrops in the northern Piedmont is a mystery that may keep biogeographers guessing for
a long time. Dr. William Murdy of Emory University conjectures that they may have been
lofted by windstorms or possibly by a single windstorm. Such a scenario requires a
fortuitous combination of a tornado or other strong updraft raking across one or more
southern rock outcrops, storm movement along a track intersecting the northern
serpentines, and deposition on the serpentines. The absence of differences detectable by
gel electrophoresis at 23 gene loci between serpentine and other populations of T.
teretifolium suggests that the species’ colonization of serpentine was relatively recent
(Murdy and Carter 1985).

Among the rarest species living on the barrens are the two eastern temperate serpentine
endemics. Aster depauperatus, with its distinctive summer-deciduous cauline leaves and
winter-persistent basal rosette, is known from about a dozen sites in Pennsylvania and
adjacent areas in Maryland. Recently, its endemic status was challenged by the discovery
that it occurs at three diabase glades in north-central North Carolina (Levy and Wilbur
1990)..A. depauperatus currently is regarded as a suspect taxonomic entity. Dr. Robin Hart
has presented morphological evidence obtained from common-garden experiments that A.
depauperatus may actually be an eastern disjunct of A. parviceps, a prairie species with a
restricted range in Illinois, lowa, and Missouri (Hart 1990).

The taxonomy of the other eastern temperate serpentine endemic is also in question.
The species name Cerastium velutinum Raf. has recently been resurrected to refer to a
hairy broad-leaved form of C. arvense L. (vars. villosum (Muhl.) Hollick & Britt, and
villosissimum Pennell in floral manuals) restricted to serpentine and limestone glades
(Morton 1987). The discovery that the distinctions of anatomy and habitat distribution
parallel a different chromosome number makes a convincing case for the revival of the
species split. C. velutinum may be a rare species in its own right. Its revival as a separate
species is so recent that no one has yet tried assessing its global abundance.

Var. villosissimum, with distinctively long white hair covering leaves and stem, is often
Said to occur in only one place in the world—bluffs overlooking Octoraro Creek at Goat
Hill, Chester County, PA, from which the type specimen was collected and described by
Francis W. Pennell (1930). Herbarium specimens from at least three other Pennsylvania
Serpentine sites were labelled as var. villosissimum and a recent discovery, still unpub-
lished, places var. villosissimum at a serpentine barren in Maryland. However, Cerastium
monographer Dr. John Morton of the University of Waterloo considers var. villosissimum

66 BARTONIA

to be a taxonomically trivial form, based on several lines of experimental and observational
evidence (J. K. Morton pers. comm.).

Adiantum pedatum subsp. calderi (serpentine maidenhair) is the only endemic shared
between serpentine outcrops of both eastern and western North America. It is the most
shade-tolerant of all the plants that have high regional fidelity to eastern serpentine
outcrops, commonly occurring in the understory of serpentine woods. Recent biosystem-
atic research (Paris and Windham 1988) suggests it may be better considered as a full
species distinct from A. pedatum. The study also showed evidence that caldert is likely to be
more closely related to maidenhair ferns of the western mountains than to the common
maidenhair of eastern forests.’

Other globally rare and endangered species are also found on the barrens. The rarest
may be Agalinus acuta (sandplain gerardia). It lives at one Maryland serpentine barren and
fewer than a half dozen non-serpentine sites northward along the Atlantic coast. Another
is Elliottia racemosa (Georgia plume), an ericaceous shrub or small tree whose entire range
is confined to scattered sites on Georgia’s coastal plain and to the Burke Mountain, GA,
serpentine barren.

Euphorbia purpurea (glade spurge) is known to occur at less than 20 places globally
including the Goat Hill, PA serpentine barren and a site in Virginia underlain by a gabbro
formation considered to be mafic (ferro-magnesian but to a lesser degree than rock classed
as ultramafic) and sharing some mineral characteristics with serpentine. E. purpurea, a
wetland species, directs attention to the fact that not all serpentine habitats are dry
uplands. Some groundwater-fed wetlands at serpentine outcrops have a distinctive flora,
most often typified by Deschampsia cespitosa or Sanguisorba canadensis.

Not all rare species characteristic of serpentine barrens are plants. The unusual plant
communities host numerous specialist-feeder insect species. As with plants, many of the
serpentine populations of insects are disjunct from the species’ main ranges. The
buckmoth (Hemileuca maia) is one of the most conspicuous of the rare insect species with
high regional fidelity to serpentine. Unlike most moths it flies during the day. It can be
spotted in late October and early November by its distinctive black and white wing
markings, large size, and rapid bouncing flight. Buckmoth larvae, spectacularly armed with
stinging hairs, feed on the leaves of Quercus ilicifolia.

PHYSIOGNOMY. The salient fact of serpentine biogeography is the convergence of
community physiognomy. Most serpentine communities look like dry prairies and
savannas even where they are well watered and surrounded by lush forests. The
southernmost temperate eastern North American serpentine barren—Burke Mountain,
GA—is a pine-grass savanna in which Pinus palustris (an isolated population) and Pinus
echinata substitute for the Pinus rigida of the state line barrens. More remarkably, the
aspect of much more distant serpentine barrens often closely resembles the appearance of
the sites in Pennsylvania and Maryland. For example, Gasquet Mountain, CA, has a
pine-grass savanna dominated by Pinus jeffreyi, Pinus monticola, Festuca idahoensis and Poa
piperi. The presence of savanna on serpentine may not seem remarkable in California
where oak savanna is one of the most widespread natural community types. However,
Gasquet Mountain stands adjacent to coastal redwood forests and the serpentine barrens
there, on average, receive more than four times the annual rainfall (Goforth 1984) as those
in Pennsylvania and Maryland!

A paper published after this symposium places serpentine maidenhair in A. aleuticum (Rhodora 93:105-121).

SERPENTINE BARRENS 67

An amazingly large proportion of the plant life at Gasquet Mountain looks quite
familiar to an eastern serpentine devotee. One is struck by the high degree of convergence
in phylogeny—some higher taxa appear to be “preadapted” for life on serpentine—and in
morphology—one sees similar, apparently adaptive traits across a wide phylogenetic
spectrum. Phlox adsurgens is strongly reminiscent of Phlox subulata, Senecio macounii looks
like Senecio anonymus, and the serpentine endemic Lilium bolanderi closely resembles
Lilium philadelphicum. The eastern naturalist may spot a plant that looks very much like
Asclepias verticillata and, on finding a flower, discover it to be a lavender-rayed composite
instead, Erigeron foliosus var. confinis.

Despite the similarities, many species appear totally unfamiliar to an easterner. For
example, the flowers of Calochortus tolmiei (Liliaceae) have purple anthers surrounded by
large blunt white petals covered with hundreds of projections resembling tentacles. The
rare Sedum laxum subsp. heckneri is a sprawling succulent with dramatic upright
candelabras of large pink and white flowers. Darlingtonia californica, a pitcher plant, has
insectivorous leaves up to 60 cm tall that look like a cross between a rearing cobra and a
giant banana slug. These and many other stunningly unfamiliar plants remind the eastern
naturalist that the California serpentines are far larger in area and have been less buffeted
by severe climate changes during the Pliocene and Quaternary compared with those in
Pennsylvania and Maryland, allowing the processes of colonization and evolution greater
scope in generating diversity.

Even within the flora of the temperate eastern North American serpentine barrens,
convergence in morphology across taxa is striking. For example, the basal rosette habit is
well developed in species from a wide range of families. Arabis lyrata (Brassicaceae), Aster
depauperatus (Asteraceae), and Panicum sphaerocarpon (Poaceae) all have basal rosettes
that are nearly identical in diameter, height, and general aspect. Asclepias verticillata, Phlox
subulata, and Polygonum tenue have linear leaves atypical of their respective families and
genera. These and other morphological traits such as leaf pubescence (Cerastium
velutinum) and succulence (Talinum teretifolium) are most often associated with hot or dry
habitats. Although the serpentine barrens are not dry, the rocky glades inhabited by all the
plants listed in this paragraph can be quite hot in summer. Infrared radiation reflected and
re-radiated from bare shales, granites, limestones, serpentine, and other rock types
commonly occurring with little or no soil cover may well have been the selective force
favoring characteristics such as basal rosettes, linear leaves, pubescence, and succulence.

SOIL FACTORS. But why is the rock bare in the first place? In regions of high rainfall such
as Pennsylvania and Maryland, what prevents weathering and biological processes from
covering serpentine with a thick layer of soil and vegetation as is the case with virtually all
neighboring bedrock formations? Before considering these important questions, I will
address one that is more fundamental. Soil does cover the bedrock beneath the other types
of serpentine vegetation besides that growing on exposed rock or glades—the serpentine
prairies, savannas, woodlands, and minerotrophic wetlands. Why are these types of
vegetation still quite different from nearby vegetation on soils weathered from non-
serpentine bedrock?

Most explanations focus on mineral nutrient conditions in soils weathered from
serpentine bedrock. Serpentine soils have exceptionally low levels of nitrogen, phospho-
rus, potassium, and calcium—the minerals needed by plants in greatest quantity. The soils
are also exceptionally high in nickel, chromium, and cobalt—metals known to be toxic to
plants in high concentrations. However, experiments conducted by manipulating these
factors in ordinary soils have failed to show a consistent link between any one of them and

68 BARTONIA

the species-specific effects of actual serpentine soils on plant growth and survival (Proctor
and Woodell 1975; Brooks 1987).

Arthur Kruckeberg’s experiment (1954) with plants from serpentine and non-serpentine
populations of Phacelia californica is one of several studies that point to the magnesium:
calcium ratio as the main factor limiting plant growth on serpentine sites. Ordinary soils
typically have approximately one or two exchangeable calcium ions for every exchangeable
magnesium ion. In serpentine soils the ratio is dramatically reversed. A survey of 156
serpentine soil samples from four continents showed a mean of 7.5 exchangeable
magnesium ions for every exchangeable calcium ion (Proctor and Woodell 1975); Mg:Ca
ratios of 20-100 are common and at least one sample has exceeded 300 (Brooks 1987).

Apparently many plant species, especially plants that are capable of fast growth, are
physiologically “confused” by an excess of magnesium relative to calcium. Their calcium-
handling enzymes may begin to bind with magnesium under conditions of magnesium
abundance combined with calcium deficiency (Proctor and Woodell 1975). Even though
magnesium is an essential plant nutrient (it forms the core of the chlorophyll molecule), at
high concentrations or in combination with calcium deficiency it interferes with the uptake
of other minerals or acts as a toxin, or both, leading to retarded growth, disease, or death.

Little is known of the physiological mechanisms that enable certain plants to handle
high magnesium:calcium ratios. Presumably some have means of preventing magnesium
from substituting for calcium. For example, some plants growing on serpentine may have
highly discriminating versions of the enzymes responsible for calcium uptake, calcium
transport, and the assembly of calcium-bearing biochemicals.

There is a common misconception that plants apparently restricted to serpentine
habitats actually require serpentine soils. To the contrary, virtually all serpentine plants
grow better on ordinary soils than they do on serpentine soils in pots or gardens where
potential competitors are absent. The key to this paradox is a trade-off, apparently
universal in the plant kingdom, between tolerance of nutrient deficiency and maximum
growth rate (Chapin 1980). Constrained by the incompatibility of key physiological and
morphological traits, members of a species can be good at tolerating nutrient scarcity or
other nutrient stresses such as high magnesium:calcium ratio or they can be good at
growing rapidly where nutrients are abundant or normally balanced, but not both. An
inherently low upper limit on growth rate apparently is a side effect of the adaptations
enabling plants of some species to grow under difficult nutrient conditions (Chapin 1980).
Plants that can take advantage of nutrient abundance by growing fast and quickly
depriving their neighbors of sunlight usually die in situations of nutrient stress. Thus,
certain plants that are tolerant of nutrient stress can achieve competitive dominance on
serpentine soils despite their low growth rates. They grow somewhat faster on nonserpen-
tine soils but not fast enough to prevent succumbing to shading and crowding by much
faster-growing plants that are intolerant of nutrient stress.

ther factors may be important in limiting the local distributions of some plants to
serpentine sites. For example, even though one might expect plants with nitrogen-fixing
root symbionts to have a competitive edge in generally high-light, low-nitrogen environ-
ments such as the serpentine barrens, very few such plants are present (exceptions on the
Pennsylvania and Maryland barrens are Cassia fasciculata and the introduced Robinia
pseudoacacia). Some scientists have attributed this paradox to exceptionally low concentra-
tions of molybdenum in serpentine soils (Walker 1948; White 1967). Molybdenum is the
central ion in the enzyme employed by nitrogen-fixing bacteria and actinomycetes to
reduce N, (atmospheric nitrogen) to NH,+ (ammonium) usable by plants.

SERPENTINE BARRENS 69

At least one investigator has suggested that there may be exceptions to the rule that
serpentine plants grow better on ordinary soils than on serpentine soils. T. M. Tadros
(1957) assayed the growth responses of a serpentine-restricted species in California,
Emmenanthe rosea (Hydrophyllaceae), to serpentine and non-serpentine soils with half of
each soil type exposed to enough heat to kill all microorganisms. The plants thrived on
sterilized non-serpentine soils but failed to germinate on those that were not sterilized.
The scientist hypothesized from this result that the serpentine-restricted species lacks
innate resistance to some fungal pathogen of seedlings but that it is protected in its native
habitat by the fungus’s intolerance of serpentine conditions.

The hypothesis to my knowledge remains untested but it is an appealing idea. One of the
most common causes of plant mortality is damping-off—a generic term for lethal attacks
upon seedlings by various fungi. If such fungi avoid serpentine soil then they are not
capable of selectively favoring resistance in potential host populations growing on
serpentine. It follows that plant populations living entirely on serpentine and not
interbreeding with populations living off serpentine could lose resistance to fungal attack
by genetic drift. In this way, hypothetically, entire plant populations could become obligate
serpentine dwellers.

CONSERVATION THREATS

So far I have reviewed a number of explanations for why the vegetation of serpentine
barrens is so distinct from the surrounding plant life. But none of these explanations has
addressed why the soil is thin and why there is often much bare rock on the barrens. It is
true that the rate of plant growth on the barrens and the resulting rate of organic matter
production are slow on the barrens relative to adjacent communities. However, soil
formation depends on net organic matter accumulation, a function of both production and
decomposition (Olson 1963). Areas free of soil should result only if serpentine conditions
impede rates of plant growth more than they impede rates of decomposition. Little
evidence exists to support or refute this hypothesis. In order to establish a foundation for
what I believe is a more compelling explanation, I will take a short detour from biology and
focus attention on conservation issues. What are the threats to this unique ecosystem in
the eastern USA?

MINING AND QUARRYING. Pennsylvania and Maryland’s serpentine outcrops were the
world center of chromium mining in the late nineteenth century, when the main use for
chromium was as a pigment in paint. Other minerals including talc, asbestos, and
corundum as well as building stone were also extracted commercially. The serpentine
barrens at Gasquet Mountain, California, have been seriously threatened by a proposed
large-scale nickel mining operation. Currently there is a huge active serpentine quarry in
southern Lancaster County producing crushed stone for concrete and asphalt paving.

Serpentine barrens conservation in Pennsylvania began after a plea from neighbors of
Goat Hill worried about a proposal to strip-mine the rock there. To date, land totalling 278
ha has been acquired for the Goat Hill serpentine barrens nature preserve by The Nature
Conservancy (a private organization dedicated to conserving biological diversity by
identifying the best remaining occurrences of natural communities and rare species,
protecting the land, and redressing any adverse effects of human land use).

DEVELOPMENT. Northern Chester County and central Delaware Counties, PA, are
littered with more than a dozen small dead and dying serpentine barrens, eaten away by
housing developments, two golf courses, and other types of intensive land use. Until
recently, serpentine barrens have enjoyed a modest level of natural protection: the soils

70 BARTONIA

are poor for growing crops and they usually fail to meet percolation requirements for
septic tank construction. But newer high density developments and the gradual coales-
cence of the patchwork of suburban sprawl make sewer systems economically feasible.
Equipped with sewers and treatment plants, serpentine barrens suddenly become
developable. Riddle Hospital currently plans to build a nursing home atop one of two
remaining barrens in Delaware County, which once harbored at least nine such sites. In
response to protest by citizens and the Pennsylvania Department of Environmental

esources, construction plans were revised to show a set-aside—a tiny patch labelled
“serpentine aster’”—with recontoured ground, landscape planting, and a wing of the
proposed building still occupying most of the actual serpentine barrens community. This
plan exemplifies what might be termed the flower-garden approach to rare species
conservation, doomed to failure. No law exists in Pennsylvania that can be invoked to
protect this rare natural community or the globally rare species that lives there.

INVASION BY NON-SERPENTINE VEGETATION. The other remaining serpentine barren in
Delaware County, Pink Hill, is protected from development as part of the Tyler
Arboretum. But is protection from bulldozers enough? A closer look suggests otherwise.
Robinia pseudoacacia has invaded the serpentine prairie at Pink Hill and appears to be
spreading. It was introduced to the region from its pre-European settlement range in the
Appalachian and Ozark mountains and the Ohio valley (Fowells 1965). Without
intervention this aggressively invasive tree would doubtless cover tiny Pink Hill’s entire
prairie area in only a few years

Even at Goat Hill, with a barrens area almost 100 times larger than Pink Hill, there are
signs of invasion by species not considered as characteristic members of the serpentine
flora. Around the fringes of the barrens, large Pinus rigida, many dead or dying, stand
among young woods dominated by Acer rubrum, Prunus serotina, Robinia pseudoacacia, and
Ailanthus altissima. An aerial view of the heart of the Goat Hill barrens reveals that the
Pinus rigida canopy is nearly closed across most of the savanna. Prairie openings—common
on other barrens of the state line group—are nearly absent at Goat Hill.

A comparison of historic and modern aerial photos reveals a dramatic pattern. An aerial
photo taken at Pilot Barrens, Cecil County, MD, in 1938 shows vast areas of open prairie
and sparse savanna. A photo of the same site taken in 1988 presents quite a different
picture. Only one comparatively small area of prairie remains. Most of the area formerly in
barrens now has closed-canopy forests of Pinus virginiana

Why do the barrens appear to be declining or even disappearing in just a few decades
when we can infer with confidence from the distributions of so many indicator plant
species that the communities have existed for thousands of years? Searching has turned up
numerous clues.

IMPORTANCE OF FIRE IN PERPETUATING BARRENS COMMUNITIES

Charred bark is a common sight on Pinus rigida of serpentine savannas and glades.
Trunks and limbs of many Pinus rigida also sport a green “fur” of adventitious shoots. It is
not uncommon to see Quercus marilandica and Quercus stellata with leaves bunched tightly
around a stout trunk on disproportionately small twigs, with larger branches either dead or
absent. Adventitious shoots spring from epicormic buds present in these particular pine
and oak species but not in other tree species living in the region. In the pine barrens of the
New Jersey coastal plain and at certain other sites, these buds are most often released from
dormancy by fire.

The serpentine dominant Sporobolus heterolepis is a bunch grass common in the
shortgrass prairies of the Dakotas, Nebraska, and other parts of the West. Bunch grasses

SERPENTINE BARRENS 71

build up masses of living and dead plant tissue around their crowns. The material acts as
effective insulation shielding crown meristems from the heat of prairie ground fires, which
occur frequently in shortgrass country. Furthermore, seeds in the soil from this and some
other prairie species reportedly break dormancy and germinate most profusely immedi-
ately following a fire.

A study conducted in the 1960’s and 1970’s confirmed the implications of numerous
fire-adapted traits in the serpentine flora. Botanist Gary Miller undertook a vegetation
distribution analysis on a serpentine barren in southern Lancaster County, PA, for his
master’s thesis. Fortuitously, a major wildfire swept through many of his sampling plots five
years after he had collected species abundance data. In resampling six and nine years
post-fire, he found increases or little change in cover by many of the characteristic
serpentine species, including the addition of previously unrecorded Aster depauperatus. He
also found substantial decreases in cover by species more commonly found on non-
serpentine soils, including the virtual disapperance of formerly abundant Pinus virginiana
(Miller 1981

Many plant species inhabiting regions or natural communities where wildfires are
common have traits enabling them to survive a fire if it is not too severe. Some of these
species are even suspected to have traits that facilitate relatively cool, fast-burning fires
(Mutch 1970; Platt, Evans and Rathbun 1988). These may include air pockets, volatile
resins, resistance to absorption and retention of moisture, and other flammability-
enhancing traits in cast-off leaves and other tissue. Vegetation dominated by species with
such characteristics tends to ignite, especially in the dormant season, at any provocation—
lightning, a discarded cigarette, a spark from burning trash or a campfire—while ordinary
vegetation is likely at most merely to smolder except during severe drought. Species acting
as “ecological arsonists” may have hit upon, evolutionarily, a means of eliminating
competitors of fire-intolerant species.

The barren-dwelling plants have apparently undergone an evolutionary trade-off. In
acquiring the abilities to tolerate infertile soil and frequent wildfire, they have sacrificed
the ability to grow fast and aggressively compete for sunlight (Chapin 1980; Grime 1977).
Thus, the vegetation of the serpentine barrens is fire-dependent. In the prolonged absence
of fire, mesic vegetation may invade serpentine barrens producing litter that readily
absorbs and retains moisture, is resistant to igniting and carrying flame, and decays to form
soil rich in organic matter. The enriched soil may shift the entire system over to a different
self-perpetuating community dominated by non-barren plant species (Streng and Harb-
combe 1982).

Frequent wildfire would explain the presence of bare rock and thin soil, two of the key
characteristics of the barrens that may be crucial to maintaining the distinctiveness of the
vegetation. Fire turns carbohydrates, the largest constituent of organic matter by volume,
into carbon dioxide, water vapor, and smoke. The organic matter that is so important to
building soil and holding it in place against erosion is periodically lost from the system in
massive quantities. Exceptionally hot fires, for example, where the wood of large trees
provides abundant fuel, may burn organic matter even after it has been incorporated into
soil. Thin soils and bare rock most likely result from the interaction of fire and water
erosion.

FIRE SUPPRESSION. Accounts from the Daily Local News, a Chester County, PA,
newspaper, give clues about why the serpentine barrens are shrinking. From 20 April 1908:
“A forest fire is raging in the pine barrens just to the south of Oxford. . . . No effort is being
made to stop the progress of the fire, it being allowed to burn itself out.” > From 7 May 1962:
“More than 300 volunteers from 14 fire companies were called into the battle to save the

72 BARTONIA

area, known as the Barrens ...” (italics are mine). Sometime between 1908 and 1962,
active fire suppression came to the rural areas where eastern serpentine barrens occur.

Not all fire suppression involves pumpers and hoses. An aerial photograph of Chrome
Barrens, Chester County, PA, shows a plethora of inadvertent firebreaks including the
road bisecting the barrens, driveways, farm fields, pastures, and lawns. Long-time
neighbors recall only one relatively small wildfire at Chrome Barrens in the past 50 years.

PRESCRIBED BURNING AS A MANAGEMENT TOOL. What can be done about this most
subtle of threats to the integrity and persistence of the eastern serpentine barrens? At the
state line barrens, staff of The Nature Conservancy in Pennsylvania and Maryland have
embarked upon a stewardship program designed to counter the effects of decades of fire
suppression and to restore periodic fire to its rightful position as an integral force in
serpentine community dynamics. A similar program is underway at Soldiers Delight by the
Maryland Natural Heritage Program, an agency of the state Department of Natural
Resources.

The program starts with research, beginning with an assay of herbaceous and woody
plant species abundances on permanently marked sampling plots. The next phase is
prescribed burning, the carefully planned and controlled use of fire by trained experts
under optimum conditions of wind, humidity, fuel moisture, and other critical factors. The
ignition, spread and movement of the fire is tightly controlled to burn a precisely specified
area. Burn units include only half of the vegetation sampling plots, allowing researchers to
tease apart the effects of fire from other influences on vegetation such as insect or disease
outbreaks, weather, soil nutrients and soil moisture.

Prescribed burning accomplishes several ends. It provides information critical to the
long-term protection of fire-dependent systems by allowing experimental evaluation of the
effects of fire on species abundances and distributions. By imitating the effects of wildfire it
restores a vital component of the natural disturbance regime to communities that show
evidence of decline due to fire suppression. It also reduces the natural fuel load and with it
the threat of catastrophic, uncontrollable wildfire.

In some places, many tons per acre of highly flammable natural fuels have accumulated
due to the artificial suppression of fire. Ironically, in a community of plants with traits
conferring high flammability, fire prevention in the short term increases fire hazard in the
long term. This is a lesson resource managers have learned over and over the hard way.
The principle made the news after the 1988 fires at Yellowstone National Park, where
decades of fire suppression had resulted in massive fuel build-up which in turn led to
unusually hot and widespread fires during an ordinary episode of drought. Yellowstone
covers a large enough area and is so sparsely settled that land managers have the option of
allowing wildfires to burn unimpeded. In an area as densely populated as the northeastern
USA, a “let it burn” wildfire policy is virtually unthinkable. Here, prescribed burning
offers a practical and environmentally sound way of preventing massive fuel build-up in
fire-dependent ecosystems and staving off catastrophic wildfires.

The first of many planned ecological burns at the state line barrens took place in fall,
1990, at Pilot Barrens under the direction of Conservancy biologist Mary Droege. The first
burn at Chrome Barrens, to be led by Conservancy biologist Michael Batcher, and the
second burn at Pilot Barrens are planned for late 1991-early 1992.

OTHER FIRE-DEPENDENT NATURAL COMMUNITIES. Serpentine barrens are not the only
natural communities considered to be fire-dependent in the mid-Atlantic region of eastern
North America’s deciduous forest biome. New Jersey’s pine barrens, especially the dwarf
forests known as the Plains, are probably the most extensive and best known example. The
Albany Pine Bush and Long Island’s pine barrens in New York exhibit the syndrome of

SERPENTINE BARRENS 73

fire-dependency. All are sites of active prescribed burning programs conducted by state
agencies, The Nature Conservancy, and other groups charged with the stewardship of
natural areas.

In Pennsylvania a few small fragments of limestone glades and prairies have escaped
cultivation, including communities characterized by Dodecatheon meadia and Poa com-
pressa at Conococheague Bluffs, Franklin County, and by Bouteloua curtipendula at
Westfall Ridge Prairie, Juniata County. The Appalachian sand barrens near Gatesburg in
Centre and Huntingdon Counties and the Appalachian shale barrens of Bedford, Fulton,
and Huntingdon Counties also probably have varying degrees of fire-dependence.

he Pocono till barrens in northeastern Pennsylvania are the state’s most extensive
fire-dependent community. They cover an area several times greater than the serpentine
barrens and have similarly rich complements of rare species. The Pocono barrens lie
overtop the state’s largest intact deposit of Illinoisan glacial till, a remnant of the
second-most recent glacial advance 140,000 years ago, in Monroe and Carbon Counties.
Dominant and characteristic species of these barrens include Pinus rigida, Quercus
ilicifolia, Rhododendron canadense, Kalmia angustifolia, Vaccinium angustifolium, Oryzopsis
pungens, Carex polymorpha, Carex vestita, and Amianthium muscaetoxicum.

Like serpentine soils, the soils weathered from Illinoisan till on the Pocono Plateau have
unusually low mineral nutrient concentrations. Like serpentine vegetation, the plant life of
the Pocono till barrens is well watered but has the aspect of vegetation native to a far drier
region. Like the serpentine barrens, the Pocono till barrens are prone to wildfire and many
of the plants exhibit traits associated with fire dependence. The community is so distinctive
that its one large and several small occurrences on the Pocono Plateau are considered to
be the planet’s entire inventory of the type. It is host to many rare species including the
globally rare and endangered Carex polymorpha. At least seven globally rare and
endangered insect species inhabit the Pocono barrens including the only insect species
whose entire known range is confined to Pennsylvania—the flypoison bulb-borer moth
(Papaipema sp. 1), which feeds on the highly toxic root of Amianthium mascaetoxicum.

CONCLUSIONS

Fire-dependent natural communities make up less than a tenth of one percent of
Pennsylvania’s total area, yet they include about 10% of the state’s known occurrences of
globally rare plants and animals (species or subspecies represented by 20 or fewer total
populations) and natural communities ranked as having global significance. Unlike many
southern, midwestern and western states, Pennsylvania has almost no recent tradition of
using fire in farming, forestry or wildlife management. Much of what people in
Pennsylvania and other northeastern states typically know about wildfire they learned
from Smokey the Bear. Prescribed burning is the best way to achieve certain important
research, conservation, and wildfire prevention goals but it cannot succeed without
increased awareness and acceptance by government officials, neighbors of fire-dependent
natural areas, and the general public. Unfortunately, the bias that all fires are tragedies
permeates the public perception and is often exacerbated by the style and content of
journalists’ coverage (Smith 1989).

The long-term persistence of Pennsylvania’s rarest and most unusual terrestrial
communities depends on scientists and resource managers adopting fire as a tool Oo
biodiversity conservation. It also depends on a convincing, well targeted campaign by
knowledgeable and concerned people to spread the word about the ecological importance
of fire.

74 BARTONIA

ACKNOWLEDGMENTS

I thank Ann Rhoads and Steve Demos for helpful comments on an earlier version of the
manuscript. Thanks are also due Kevin Dougherty, Mary Droege, George Harlow, John
Morton, Bill Murdy, Gail Newton, Carol Palmer, Dale Schweitzer and Conrad Smith for
information on particular topics.

LITERATURE CITED

Brooks, R. R. 1987. Serpentine and its vegetation. va Press, Portlan

CHAPIN, F. S., II. 1980. The mineral nutrition of wild plants. Ann. Rev. Ecol. ca 11: 233-260.

DARLINGTON, W. 1849. Memorials of John Bartram wre Humphry Marshall. Classica Botanica Americana,
Suppl. 1 (repr. 1967, J. A. Ewen, ed., Hafner, New York).

DEEVEY, E. S. AND R. F. FLINT. 1957. Postglacial hypsithermal interval. Science 125: 182-184.

FowELLs, H. A. 1965. Silvics of forest trees of the United States. Agr. Handbk. 271. U.S. Dept. of Agr., Forest
Service, Washington.

GoForRTH, D. 1984. Gasquet Mountain. Fremontia 11: 11-12

GRIME, J. P. 1977. Evidence for the existence of three primary strategies in plants and its relevance to ecological
and evolutionary theory. Am. Nat. 111: 1169-1194.

Hart, R. 1990. Aster depauperatus: a midwestern migrant on eastern serpentine barrens? Bartonia 56: 23-28.

IRWIN, W. P. AND R. G. COLEMAN. 1972. Preliminary map showing pie distribution of alpine-type ultramafic
rocks and blueschists. U.S. Geol. Surv. Misc. Field Studies Map MF-340.

KRUCKEBERG, A. R. 1954. The ecology of serpentine soils. III. Plant species in relation to serpentine soils.
ppd - 267-274.

EVY, F. AND R. L. WILBUR. 1990. Disjunct populations of the alleged serpentine endemic, Aster depauperatus

éPovten\| Fern., on diabase ial a in North Carolina. Rhodora 92: 17-21

MANSBERG, L. AND T. R. WENTWORTH. 1984. Vegetation and soils of a serpentine barren in western North
Carolina. met Torrey | Bot. Club iL 1-256,

MILLER, G. L. 1 tine b . Proc. Pennsylvania Acad. Sci. 55:

o pg

62-64.

Morton, J. K. 1987. Caryophyllaceae. Looseleaf unnumbered pages in G. W. Argus, K. M. Pryer, D. J. White
and es i Keddy, eds. 1982-1987. Atlas of the Rare Vascular Plants of Ontario. Nat. Mus. of ‘Natal Sci.,
Otta

Murpy, W. H. AND M. E. CarTErR. 1985. pearepnerntc study of the Seas Pe ome of Talinum
teretifolium in ee specific status of 1590-1597
Mutcu, R. W. 1970. Wildland fires and echeystene =n hypothesis. Ecology He aie

OLson, J. S. 1963. Ener storage and the balance of producers and decomposers in ecological systems. Ecology
44: 322-331

PARIS, C. A. AND M. D. WINDHAM. 1988. A biosystematic investigation of the Adiantum pedatum complex in
eastern North America. Syst. Bot. 13: 240-255.

PEARRE, N. C. AND A. V. HEYL, JR. 1960. Chromite and other mineral deposits in ag rocks of the

Piedmont upland, Maryland, shapes isacepes a shiasdhasi Us Geol. Surv. Bull. 1082-

PENNELL, F. W. 1930. O i =<

PLaTT, W. J., G. W. EVANS, AND S. J. RATHBUN. 1988. The population dynamics of a long-lived conifer (Pinus
palustris). Am. Nat. 131: 491-525.

ROCTOR, J. AND S. R. J. WOODELL. 1975. The ecology oils. Adv. Ecol. Res. 9: 255-366.

SALEEBY, J. B. 1984. Tectonic significance of ser tentinite ed pice ophiolitic melange. Pp. at in L. A.
Raymond, ed. Melanges: their nature, origin, and significance. Geol. Soc. of Am. Spec. Paper 198.

SMITH, C. 1989. Reporters, news sources, accuracy, and the Yellowstone forest fires. Paper pontine at Int.
Communications Assn. ann. mtg., May 1989, San Francisco.

STRENG, D. R. AND P. A. HARCOMBE. 1982. Why don’t East Texas savannas grow up to forest? Amer. Midl. Nat.

78-2
Tapros, T. M. 1957. Evidence of the presence of an edapho-biotic factor in the problem of tine tolerance.
Ecology 38: 14-23.
WALKER, R. B. 1948, Molybdenum defici i tine b il Science 108: 473-475.
WHITE, C. D. 1967. Absence of nodule f ti C th tl ils. Nature 215: 875.

=

Bartonia No. 57, Supplement: 75-92, 1993

Selected Rare and Historical Vascular Plants of Delaware

KEITH CLANCY
Delaware Natural Heritage Inventory, Department of Natural Resources and Environmental Control,
89 Kings Highway, Dover, DE 19903

Botanists and zoologists with the Delaware Natural Heritage Inventory (DNHI) are
responsible for surveying the entire state for rare species and significant natural
communities. A major challenge of this work is to assess the rare flora and fauna, a task
made even more difficult by the dynamic nature of the biota. A formerly common species
may become rare due to habitat destruction, and conversely, as more field work is
undertaken, a species previously believed rare may be found to be more common.

DNHI has developed a list of rare native plants (McAvoy 1993). The list, which is
updated annually, depends on input from many sources: experts in the field, literature,
herbarium collections, and field surveys. Tatnall (1946) provided substantial, and
important, historical information on the Delaware flora including species he considered
rare or no longer extant. Phillips (1978) was unable to relocate 338 species reported by
Tatnall. Tucker et al. (1979) designated 450 taxa as rare, endangered, possibly extirpated,
or status uncertain, and recognized the need for more thorough field work.

Over the past five years, intensive botanical field work has resulted in new information
on the distribution and abundance of many species. These data are used to assess status.
Although information from literature sources and herbarium collections form the frame-
work, the development of the list is driven by intensive field surveys.

When a rare species is discovered, detailed information is gathered on population
ecology, areal extent, co-occurring species, habitat description, etc. Locations are mapped
on USGS topographic maps, and the record of the species is entered into the DNHI
database (DNHI 1993). Herbarium voucher collections are made with prudence (i.e.
collecting is avoided if population is small, or species is known from only a few
populations).

Of the 2259 taxa listed by Tatnall (1946) as native, naturalized, or otherwise occurring
on the Delmarva Peninsula; nearly 93% were present in Delaware (Tucker et al. 1979).
Inventory work undertaken since Tatnall’s publication, and particularly during the past
five to ten years, has shown that many of the remaining 7% are also present in the state
(e.g. Alopecurus aequalis, Hypericum drummondii, Juncus torreyi, Mecardonia acuminata,
Passiflora incarnata). In addition, many other species which have been found in Delaware
since 1946, are also new to the flora of the Delmarva Peninsula (e.g. Carex projecta, Isotria
medeoloides, Listera australis, Panicum hirstii, Pellaea atropurpurea, Sanicula trifoliata,
Schizaea pusilla).

The information presented in this paper has been updated since the 1991 symposium to
reflect new data from 1991 and 1992 field surveys. Nomenclature generally follows
Gleason and Cronquist (1991).

RARE SPECIES

On-going botanical field surveys in potentially suitable habitats and at historical sites
throughout the state suggest that the following species are rare in Delaware. All those
discussed below are ranked $1 (extremely rare; 5 or fewer known occurrences in the state)

75

76 BARTONIA

by DNHI. A total of 296 species are currently included in this category; an effort was made
to select for inclusion here, species for which rarity could be well substantiated.

Federal status designations are included (US Fish and Wildlife Service 1991a, 1991b,
1992a, 1992b, 1992c): LE = Listed Endangered; LT = Listed Threatened; PT = Proposed
Threatened; C1 = Candidate, Category 1; C1* = Taxa which are believed to be extinct but
persuasive documentation has not been made; C2 = Candidate, Category 2.

Actaea alba (L.) Miller (= A. pachypoda Ell.), white baneberry. Tatnall (1946)
considered this species to be “rare, in rocky woods along Brandywine Creek,” and cited
several collections: (1) Wills Rock, Wilmington, 1866 and 1893 (A. Commons s.n., PH), 13
May 1899 (W. M. Canby s.n., DOV), and along the railroad at Wills Rock, 13 May 1944
(R. R. Tatnall 5085, DOV); and (2) Rockford Woods, 14 May 1897 (E. Tatnall, DOV). It is
currently known from two sites in the Brandywine drainage of the Piedmont Province: (1)
Rockland Park, Wilmington, several dozen plants were observed on steep slopes above the
Brandywine Creek in 1988 (Ebert et al. 1990), this may be a relocation of E. Tatnall’s 1897
Rockford Woods site; and (2) near Hoopes Reservoir, one plant was observed in a mature
hardwood forest in 1991 by J. Ebert & J. Holt.

Amianthium muscaetoxicum (Walt.) Gray, fly-poison. Tatnall (1946) considered this
plant to be “infrequent.” Historical collections include the following: (1) along Penrose
Branch, 5.5 miles west of Dover, 24 Jun 1930 (H. H. Hanson s.n., DOV); (2) near
Greenbank, low ground, Work horse farm, 8 Jun 1922 (J. P. Otis s.n., PH); (3) between
woods and Greenbank farm, low meadow, 2 Jul 1923 (J. P. Otis s.n., PH); (4) Mt. Cuba, dry
woods, 8 Jul 1875, (A. Commons s.n., PH), 19 Jul 1891, (J. B. Brinton s.n., PH) and 24 Jun
1906, (S. S. VanPelt s.n., PH). It is currently known from five New Castle Co. sites, all in
the Red Clay Creek drainage: (1) near Mt. Cuba, over 80 individuals were observed in a
dry oak-beech woods, discovered in 1989 (Ebert et al. 1990); (2) near Wooddale, a small
population of several plants was observed in a wooded ravine of mixed hardwoods in 1991
by J. Ebert & J. Holt; (3) near Wooddale three plants were observed in a mature mesic
forest dominated by Fagus grandifolia in 1991 by J. Ebert & J. Holt; (4) near Ashland, along
small tributary, four plants were observed in a mature woods of Fagus grandifolia, Quercus
rubra, Hamaemelis virginiana, and Viburnum acerifolium in 1991 by J. Ebert & J. Holt; and
(5) within 0.25 mile of previous site, more than 30 individuals were noted growing within 10
meters of a small tributary of the Red Clay Creek in 1991 by J. Ebert & J. Holt.

Arenaria caroliniana Walter, pine barren sandwort. According to Tatnall (1946) this
plant was known from “a single collection: near Little Hill Church, 0.5 mile ne of
Pepperbox, in dry pine barrens,” (Commons, 5 Aug 1874, PH). The pine barren sandwort
had been ranked SH by DNHI until several hundred plants were found in the dunes at
Cape Henlopen State Park by F. Hirst & R. Wilson in 1992.

Asclepias lanceolata Walt., few-flowered milkweed. Tatnall (1946) considered this
milkweed to be “frequent on beaches and in salt marshes” of Kent and Sussex counties.
Recent field surveys conducted by DNHI botanists have resulted in its discovery at four
Sussex Co. sites: (1) near Bethany Beach, a small population was observed in 1989 in an
interdunal swale with several other state-wide rare species, Eupatorium leucolepis, Fuirena
squarrosa, and Vaccinium macrocarpon by F. Hirst. In addition three distinct populations
were located in the Assawoman Bay Wildlife Area; one in a coastal plain pond, one at the
edge of a pond and along a woods road, both discovered by F. Hirst in 1989 and 1991; and
the third at the edge of a brackish high-marsh within the general area of other rare species
(Agalinis maritima, Centella erecta, and Juncus roemerianus), discovered in 1992 by W.
McAvoy. All populations consisted of less than 10 individuals.

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 77

Asclepias rubra L., red milkweed. Tatnall (1946) reported this species to be “infrequent,
in all three counties of Delaware.” There are two known extant populations on the Coastal
Plain: (1) near Dagsboro, less than 10 individuals were observed, with other state rare
species (Paspalum dissectum, Platanthera blephariglottis, P. lacera, Rhynchospora gracilenta,
R. microcephala) along an approximately 0.5 mile long swale in a powerline right-of-way in
1984 by F. Hirst: and (2) south of Woodland Ferry, a population of about 50 plants was
observed along the roadside adjacent to a tidal red maple swamp in 1989 by R. Radis.

Asplenium trichomanes L., maidenhair spleenwort. Tatnall (1946) stated that this fern
was “rare, in moist rock-crevices along the Brandywine and Red Clay Creeks. ... Not
collected since 1893.” Tucker et al. (1979) considered it extirpated, and at a December
1990 meeting of DNHI botanists it was decided to rerank this species from SH to SX.
However, that decision was premature, as J. Holt & J. Ebert located three small
populations of this fern growing on rocks above the Brandywine Creek near the Hagley
Museum in April 1991. This site is a relocation of the Soda House Woods site discussed by
Fleming (1978).

Bidens bidentoides (Nutt.) Britton var. bidentoides, bur-marigold, C2. This plant has
rather narrow habitat requirements, i.e. fresh to slightly brackish tidal mud flats (Ferren
and Schuyler 1980). Tatnall (1946) stated that it was “infrequent” on the Delmarva. It is
known from four extant sites, although population viability is tenuous, and several
historical sites. It is found growing in tidal mud amid rocks and concrete blocks from near
Delaware City to just below the PA state line, with the majority of plants having been
observed in and near New Castle. Historically, it has been collected on several occasions
along the Delaware River, the Brandywine Creek, and possibly along the Appoquinimink
River (W. M. Canby, undated PENN; W. M. Canby Oct. 1865, PENN; ex Herb. E. Tatnall
1865, GH; Ferren 1159 and 1107, PH; R. Tatnall s.n., PENN). Population numbers in 1989
and 1990 ranged from a few individuals to less than twenty per site. Surveys in the summer
and fall of 1991 located only one individual, a seedling (K. Clancy, B. McAvoy, A. E.
Schuyler pers. observs.). This species is threatened throughout its range by dredging,
filling, bulkheading, salinity increases, and pollution. A recent oil spill on the Delaware
River (Presidente Rivera oil spill of 1989) resulted in many individuals of Bidens being
covered in oil (R. Radis pers. comm.) and during surveys in September, 1991 a thick layer
of oil-soaked mud was discovered at a site near Claymont (K. Clancy and B. McAvoy pers.
obs.).

Botrychium matricariaefolium A. Br., chamomile grape-fern. This species was not
reported to occur on the Delmarva by Tatnall (1946). It is now known from four Delaware
sites: (1) Pike Creek Valley near Rte 2, in 1989 eight individuals were counted in a young
tulip poplar woods (Ebert et al. 1990), this population may already have been destroyed by
a housing development (B. McAvoy pers. comm.); (2) White Clay Creek Preserve, several
dozen plants were observed in a young red maple woods in 1988 (Ebert et al. 1990); (3)
Brandywine Creek Valley near Rockland, only one sporophyte was observed at the
interface between mature hardwood forest and young maple woods in 1989 (Ebert et al.
1990); and (4) adjacent to the Christiana River near Newark, three sporophytes were
observed, growing with B. virginianum, in a mature beech-tulip woods in 1991 by J. Ebert &
J. Holt.

Cacalia atriplicifolia L., pale Indian plantain. Tatnall (1946) reported this species to be
“infrequent in pastures, thickets and open woods of the Piedmont Province; rare on the
Coastal Plain. . . .” It is known from several historical collections from New Castle Co.: (1)
field near Thompson Station, 6 Aug 1932, (R. R. Tatnall s.n., PH); (2) Kiamensi, 9 Sep

78 BARTONIA

1901, (A. Commons s.n., PH); and (3) Greenbank, in woods, 13 Sep 1884, (A. Commons
s.n., PH). The latter two sites occur near each other in the Red Clay Creek drainage. Since
1988 this rare composite has been discovered by J. Ebert & J. Holt at three localities in the
state: (1) along the Red Clay Creek north of Wooddale, eight plants were observed
growing in an old field; (2) Brandywine Creek, several dozen plants were observed along
the floodplain near Ramsey Run; and (3) Sandy Branch east of MD line, nearly 100 plants
were observed on slope above the creek.

Chamaelirium luteum (L.) A. Gray, devil’s-bit, blazing star. This species was considered
“infrequent, in rich woods and thickets of the Piedmont province” by Tatnall (1946), and
was listed by Tucker et al. (1979) as “possibly extirpated. ...” Historical collections
include: (1) Wilmington, thickets, 25 May and Sep 1896, (A. Commons s.n., PH); (2)
Brandywine, rocky woods, May 1843 (Rev. J. H. B. s.n., PH); (3) Mt. Cuba, 3 Jun 1894 (A.
John s.n., PH). According to Tucker et al. (1979) the species was last collected in Delaware
in 1938, however, during the 1992 field season, three plants were observed in disturbed
habitat along a road bank in northern New Castle Co. by J. Ebert & J. Holt.

Coreopsis rosea Nutt., pink tickseed. This species was reported by Tatnall (1946) to be
“frequent ... from New Castle to Sussex . . . counties.” Phillips (1978) stated that it was
not found on the Delmarva, and Tucker et al. (1979) concluded that the species needed
further field work to determine its true status in Delaware. According to Tucker et al.
(1979) there were seven historical populations, the most recent from 1938, distributed in
New Castle, Kent, and Sussex counties. At one time, Ellendale wet meadow was home to
the pink tickseed (Dill and Tucker 1982), but searches at the site since 1984 have proved
unsuccessful. Recent field surveys have resulted in its discovery at three Sussex Co.
locations: (1) near Lewes, two populations were found in separate coastal plain ponds
(Delmarva Bays), by D. Boone & F. Hirst in 1982, and F. Hirst in 1988; and (2) near
Milton, an exceptionally vigorous population, estimated at >10,000 in 1990, was
discovered in 1982 by D. Boone and F. Hirst in a pristine coastal plain pond.

Corydalis flavula (Raf.) DC., yellow corydalis. Tatnall (1946) considered the species to
be “locally abundant in rocky woods . . . rare in northeastern New Castle Co.” and cited
one historical collection: Naamans Creek, below Harvey (Arden) Station, (Commons and
Tatnall, 27 April 1894, PH). Although Tucker et al. (1979) mentioned two populations
seen in 1971, they believed the plant to be extirpated in the state. There is one known
population: near White Clay Creek, north of Newark, thousands of individuals were
observed along a nearly 1.0 mile length of terrain, growing with a number of alien species
(Cerastium vulgatum, Stellaria media, Valerianella locusta) in 1992 by J. Ebert & J. Holt.

Desmodium strictum (Pursh) DC., pine-barren tick-trefoil. Tatnall (1946) considered
this species to be “infrequent, ... from Caroline Co. to Worcester Co.,” and did not
include Delaware in the species’ range. Nevertheless, since 1988, four populations have
been discovered: (1) Nanticoke Wildlife Area, three populations of one, two, and over 200
individuals, respectively were discovered in xeric, sandy pine woods by C. Ludwig in 1988;
and (2) near Nanticoke Wildlife Area north of Broad Creek, several dozen fruiting
individuals were observed in dry sandy pine woods by F. Hirst in 1989.

Diervilla lonicera Miller, bush-honeysuckle. This species was reported by Tatnall (1946)
to be “rare, in rocky woods of New Castle Co.” Historical collections include: (1
Rattlesnake Run, Wilmington, 1 Jun 1844 (“G. W. T.” s.n., DOV); (2) Mt. Cuba, 1873 and
1896 (A. Commons s.n., PH) and 29 Jun 1893 (E. Tatnall s.n., DOV); (3) Point Lookout,
0.5 mile nw of Granogue, 20 Jun 1937 and earlier collections (R. R. Tatnall 3425, DOV, G,
PENN, PH); and (4) above Rockland, rocky woods along Brandywine, 8 May 1897 (A.

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 79

Commons s.n., PH). Since 1988 it has been discovered at two sites on the Piedmont: (1)
along the Brandywine Creek west of Talleyville, a small population was noted in a rocky,
steep-sloped Linodendron tulipifera woods, discovered in 1988; and (2) northwest of Mt.
Cuba, two plants were observed in a rocky mature oak-beech woods, discovered in 1989
(Ebert et al. 1990).

Dirca palustris L., leatherwood. Tatnall (1946) considered this plant to be “rare, in rich,
rocky woods along Brandywine and Red Clay Creeks, New Castle Co.” Historical
collections include the following: (1) Mt. Cuba, 3 Jun 1864 (A. Commons s.n., PH) and at
least seven other collections; and (2) west bank of Brandywine Creek, 0.25 mile below
Henry Clay bridge, 10 Apr 1888 (R. R. Tatnall 1369, PH), and several additional
collections made from, or near, this site. Intensive surveys during the past five years
resulted in the 1989 discovery of a site near Mt. Cuba, one fruiting shrub was observed in
1992 in a mature mixed hardwood forest; this may be a relocation of Commons’ 1864 site
(Ebert et al. 1990).

Eleocharis rostellata (Torr.) Torr., small-beaked spikerush. This plant was considered to
be “rare” by Tatnall (1946), who cited several historical collections: (1) Collins Beach, 20
Jun 1866 (A. Commons s.n., PH); and (2) Love Creek, near Rehoboth, 20 Jun 1926 (True
s.n., PENN). This rare sedge is known from four extant sites: (1) Love Creek near Rte 24, a
small population, surrounded on three sides by Phragmites australis, was observed in tidal
brackish marsh in 1992 by C. Ludwig & B. McAvoy; (2) Angola Neck near Bookhammer
Landing, an abundant and dominant species in more than 4 acres of a slightly brackish to
freshwater bog-like habitat discovered in 1992 by C. Ludwig; (3) Assawoman Bay Wildlife
Area, several thousand stems were observed covering approximately 6 acres of high tidal
marsh by B. McAvoy in 1992; and (4) Angola Neck near Bookhammer Landing, more than
1000 stems was observed at edge of salt marsh by F. Hirst & R. Wilson in 1992. The Love
Creek site may represent a rediscovery of True’s historical site.

Equisetum fluviatile L., water horsetail. Tatnall (1946) considered this species to be
“infrequent to rare’? on the Delmarva and cited the following collections: (1) Bellevue,
near Station, Jun 1868 (Commons s.n., PH); (2) Port Penn, 28 Jul 1884 (J. Carter s.n., PH);
(3) south bank of Drawyers Creek, above DuPont Highway 1 mile north of Odessa, 4 Jul
1929 (R. R. Tatnall s.n., DOV); (4) streamlet west side of DuPont Highway, 1 mile north of
Fieldsboro, 3 Jun 1939 (R. R. Tatnall 4188, DOV, PH). The last site is still extant. I
followed Tatnall’s directions precisely, and discovered this horsetail at the exact location
specified. More than 100 stems were observed in a narrow mixed hardwood swamp forest
adjacent to an emergent freshtidal marsh (K. Clancy 2617, 2625, DNHI Reference
Herbarium).

Eriophorum virginicum L., tawny cotton-grass. Tatnall (1946) considered this species to
be “infrequent,” and intensive searches by DNHI botanists over the past few years would
support that characterization. Since 1986, two sites have been discovered in Sussex Co.: (1)
Angola Neck near Bookhammer Landing, a small population was observed in a narrow
Alnus maritima dominated scrub-shrub wetland at the interface between a mixed forest
and an emergent freshwater bog-like community by C. Ludwig in 1992; and (2) west of
Frankford, in moist sandy soil, where it was observed associated with Bartonia virginica,
Eupatorium leucolepis, Rhynchospora microcephala, R. capitellata, Polygala lutea, and Juncus
canadensis (Naczi et al. 1986).

Eryngium aquaticum L., button snakeroot. This species was reported by Tatnall (1946) to
be “frequent; ditches, rivers, beaches, dune hollows and swamps of the coastal plain, in
fresh or brackish habitats.” Recent intensive floristic surveys have resulted in its discovery

80 BARTONIA

at three (possibly four) Sussex Co. sites: (1) along Love Creek, near Rt 24, a large
population covering approximately five acres occurs in fresh tidal marsh, discovered in
1992 by C. Ludwig & B. McAvoy; (2) along Burton Prong, near Burton Millpond, ca. 50
plants in bud and flower were observed at the edge of a brackish marsh, co-occurring with
such species as Baccharis halamifolia, Hydrocotyle verticillata var. verticillata, Phragmites
australis, Rumex verticillatus, and Sabatia dodecandra in 1992 by F. Hirst; and (3) Miller
Neck, adjacent to Dirickson Creek, several dozen plants were observed in an open,
freshwater bog-like habitat above salt marsh in 1992 by B. McAvoy. Associated species
include: Centella erecta, Cladium mariscoides, Eleocharis palustris, Carex crinita, and
Sphagnum sp. A possible fourth site is in the Assawoman Bay Wildlife Area north of Miller
Creek, several plants were observed in relatively dry habitat (K. Clancy 2046, DNHI
Reference Herbarium). Identification is tentative as these plants appear to be intermedi-
ate between E. aquaticum and E. yuccifolium.

Fimbristylis perpusilla Harper, Harper’s fimbristylis, C2. This species was not reported
for the Delmarva by Tatnall (1946). Until very recently this sedge was only known from
Georgia (Kral 1983), however, field work in the last twelve years has resulted in discoveries
in several southeastern states (Leonard 1981, Hirst 1983, Leonard 1987, Wofford and
Jones 1988, Harvill et al. 1992). This species is sporadic and appears irregularly over the
years; at one locale in Georgia it went nine years between appearances (Kral 1983).
Populations in Delaware exhibit similar erratic fluctuations (F. Hirst pers. comm.). It is
known from three Coastal Plain ponds: (1) west of Hartly, a small population was observed
in a woodland pond in 1984 by F. Hirst, this population has not been seen here since its
discovery; (2) in a pond east of Delaneys Corner, a small population restricted to the
center of the pond was first discovered by F. Hirst in 1983 and subsequently observed in
1991 and 1992 (K. Clancy 2373, DNHI Reference Herbarium); and (3) west of Delaneys
Corner, a small population was observed in a woodland pond in 1991 by B. McAvoy, no
plants were observed in 1992, although the pond was flooded into October.

Hudsonia ericoides L., golden heather. Tatnall (1946) described this species as “rare, in
dry sandy soil of the seacoast,” and cited several collections: (1) Cape Henlopen, in 1895
and 1898, (A. Commons s.n., PH); and (2) along the railroad, 3 mile nw of Rehoboth, 30
May 1935, (R. R. Tatnall 2578, DOV, PH), and 16 May 1938, (R. R. Tatnall 3695, DOV).
There are currently three known extant sites, all within the confines of Cape Henlopen
State Park: (1) in dunes, several hundred plants were observed by F. Hirst in 1988; (2) in
xeric sand dunes near the US Naval Reservation, only a few clumps were seen growing
alongside H. tomentosa, Lechea sp., and Panicum commonsianum in 1992 by F. Hirst & R.
Wilson; and (3) in maritime pine forest, a few plants were seen in 1992 by F. Hirst & R.
Wilson.

Isoetes riparia Engelm., riverbank quillwort. This species was reported by Tatnall (1946)
to have been “collected in tidal mud of Delaware River, at and north of Wilmington, by
Commons, Canby and E. Tatnall, between 1862 and 1896; not seen since.” Today, there
are three known localities: (1) Delaware River near Delaware City, New Castle Co.,
between 50-100 plants were observed in mud of a brackish tidal marsh in 1988 by R. Radis,
searches at this site in 1992 were unsuccessful; (2) along the Nanticoke River, near
Middleford, Sussex Co., two distinct populations of ca. 20 plants, and between 50-100
individuals, respectively, were observed in the freshwater intertidal zone by F. Hirst in
1988. There appears to be ample habitat for this species, especially in light of its discovery
along the Nanticoke River.

Isotria medeoloides (Pursh) Raf, small-whorled pogonia, LE. In Delaware this rare

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 81

orchid is known from a single population near Blackbird State Forest, New Castle Co.
where a few plants were observed in a young mesic oak-beech-hickory forest by M.
McLaughlin in 1985. Since 1985, population numbers at this site have varied from 0-11
stems, no plants were observed in 1992.

Lachnanthes caroliniana (Lam.) Dandy, Carolina redroot. Tatnall (1946) stated that this
species was “frequent in bogs and sandy swamps, Kent and Sussex counties.” However,
intensive surveys over the past five years indicate that it has suffered a decline since
Tatnall’s time. The species is now known from five Sussex Co. sites: (1) near Lewes, several
hundred plants were observed in a coastal plain pond (Delmarva Bay) by F. Hirst in 1988;
(2) near Milton, a large population of several thousand plants was observed in a pristine
coastal plain pond by F. Hirst & D. Boone in 1982 and still extant in 1992; (3) near
Robbins, 50-100 plants were observed in a wet clearing of a re-planted pine plantation by
F. Hirst in 1989; (4) north of Georgetown, two distinct populations, located more than 1.0
mile apart in openings in wet pine woods, one with less than 10 plants, the other with
several hundred plants, were discovered by F. Hirst & R. Wilson in 1990 and 1992.

Listera australis Lindl., southern twayblade. Tatnall (1946) did not list this species for the
Delmarva. There are currently three known sites in Delaware, all in Sussex Co.: (1)
Nanticoke Wildlife Area, hundreds of plants were observed in 1992 growing on hummocks
in a white cedar-red maple-black gum swamp, discovered in 1990 by F. Hirst; (2) Great
Cypress Swamp, hundreds of plants were observed growing in a poorly drained loblolly
pine woods by B. McAvoy in 1992; and (3) Nanticoke River near Middleford, a small
population of approximately 30 plants was observed growing on hummocks in an Atlantic
white cedar-red maple swamp by K. Clancy & B. McAvoy in 1992.

Lobelia elongata Small, elongated lobelia. Tatnall (1946), who cited collections by
Commons (1877, 1880) from “ponds, Millsboro,” and a 1907 record from Brown collected
along “Indian River, Millsboro,” considered this species to be “rare, in wet places of the
Coastal Plain.” Tucker et al. (1979) reported that it was last seen in Delaware in 1907.
Today this Lobelia is known from two extant Sussex Co. sites: (1) along Guinea Creek, less
than 10 plants were observed above brackish marsh in a freshwater seepage habitat by C.
Ludwig & B. McAvoy in 1992; and (2) Upper Love Creek near Rt 24, an abundant
flowering and fruiting population was observed in fresh-tidal marsh on both sides of the
creek by C. Ludwig & B. McAvoy in 1992. The latter population co-occurs with other
rarities including: Bidens coronata, Eryngium aquaticum, Sabatia dodecandra, and Sacciole-
pis striata.

Lupinus perennis L., wild lupine. This species was reported by Tatnall (1946) to be
“infrequent in sandy soil . . . southern New Castle Co. southward. . . .” However, extensive
inventories indicate that this species is much rarer than Tatnall reported. It is currently
known from three locations: (1) near Killens Pond State Park, Kent Co., a vigorous
flowering population of several hundred stems was observed in sandy soil along a roadside
in 1991 (K. Clancy 2210, DNHI Reference Herbarium); and (2) southwestern Sussex Co.,
in the Nanticoke Wildlife Area, two populations of less than 50 stems and ca. 200 stems,
respectively, were observed along roadsides (several miles apart) in dry, sandy pine-oak
woods in 1988 by C. Ludwig, and 1989 by F. Hirst.

Malaxis unifolia Michx., green adder’s-mouth orchid. Tatnall (1946) listed this species as
“infrequent but widespread in both Piedmont and Coastal Plain provinces,” while Phillips
(1978) stated that it was “infrequent throughout.” Currently it is known from three
locations: (1) near Sandhill, Sussex Co., less than ten plants were observed in roadside
woods by F. Hirst in 1984; (2) south of Ellendale, Sussex Co., one fruiting individual was

82 BARTONIA

observed in a rich, mesic woods by F. Hirst in 1989; and (3) near the Cedar Swamp, New
Castle Co., about 30 individuals were observed in a wet sweetgum woods in 1992 by J.
Ebert & J. Holt.

Matelea carolinensis (Jacq.) Woodson, Carolina angelpod. Tatnall (1946) reported that
this species was “infrequent, in thickets, or climbing on fences, in southern New Castle, . . .
Kent ... counties.’”’ Brown and Brown (1972) stated that this species was “infrequent in
Cecil, Kent, and Queen Anne counties” on Maryland’s Eastern Shore. Harvill et al. (1992)
did not list it for Virginia’s portion of the Delmarva. This plant is known from one location:
along the MD-DE state line, about 40 plants were observed at the edge of road and
scattered in woods by J. Ebert & J. Holt in 1992. The habitat suggests that it may be more
common than currently believed.

Nelumbo lutea (Willd.) Pers., American or yellow lotus. This plant was reported to be
“rare” by Tatnall (1946) who cited one locality: St Jones Creek, 2.5 miles below Dover,
1863 (A. Commons s.n., PH), 3 August 1930 (R. R. Tatnall 965, DOV, GH, PH). In
addition, several undocumented records were also mentioned by Tatnall; Drawyers Creek
below Shallcross Lake; Moore’s (Wiggins) Pond, 1 mile nw of Townsend; and the sw end of
McGinnis Pond, 2 mile e of Canterbury. Tucker et al. (1979) suggested that it was
‘probably extirpated.” There is currently one known extant site: in Dover south of the Hwy
13 bridge, in tidal marsh along the St. Jones River. This population was brought to the
attention of DNHI in December 1990 and a small population was observed in July 1991 (K.
Clancy pers. obs.); it may represent what remains of the historical St. Jones’ population.

Nymphoides cordata (Ell.) Fern., little floating heart. This species was reported by
Tatnall (1946) to be “infrequent, on ponds, from central Kent.” It is known from one site:
west of Lewes, where less than 100 individuals occur with 13 other state rare species, in a
coastal plain pond discovered by F. Hirst in 1988. More field work is needed to accurately
assess this species’ status in Delaware, as it could easily be confused with the often
sympatric, and more common N. aquatica.

Panicum hirstii Swallen, Hirst’s panic grass, C2. One of the rarest grasses known, this
plant occurs in wet meadows or temporary ponds in NJ, DE, NC, and GA (A. Weakley
pers. comm., Kral 1983). P. hirstii is known from one Sussex Co. location: Assawoman Bay
Wildlife Area, a small population, varying from 10 to several hundred culms, occurs in a
coastal plain pond (Delmarva Bay) within a pine-hardwood forest, discovered by F. Hirst
in

Passiflora lutea L., yellow passion-flower. Tatnall (1946) stated that P. Jutea was “local
... and cited collections from only the Maryland and Virginia portions of the Delmarva.
However, a check of herbaria uncovered two historical collections: (1) “Delaware” 1824?
(Nuttall?, GH); and (2) “Delaware” 1833 (H. C. Beyrich, MO). There are currently two
known populations (Clancy 1993): (1) near Dover and the St. Jones River, Kent Co.,
several flowering vines at the edge and several dozen seedlings just inside a narrow
second-growth mixed deciduous forest, discovered in 1991 (K. Clancy 2315, DNHI
Reference Herbarium, PH); and (2) near Lewes, Sussex Co., an extensive flowering and
fruiting population was observed in a clearing of a mixed oak-pine forest in 1991 (K-
Clancy 2381, DNHI Reference Herbarium, PH).

Pellaea atropurpurea (L.) Link, purple cliff-brake. This fern was not reported by Tatnall
(1946) for the Delmarva. It is currently known from two locations on the Piedmont: (1)
near Rockland, about 100 plants were observed growing on a stone wall above the
Brandywine Creek, (Ebert et al. 1990); and, (2) along Rockland Road, approximately 100
plants were observed, also growing on a stone wall, discovered in 1990 by J. Ebert & J

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 83

Holt. These are the first known sites for this species in Delaware, and perhaps in the
Delmarva

Platanthera peramoena (Gray) Gray, purple fringeless orchid. Tatnall (1946) reported
this species to be “rare and local ...” and cited several historical collections from New
Castle Co.: (1) moist meadows above Rockland along Brandywine Creek, 8 Aug 1864 (A.
Commons s.n., PH); (2) Granogue, (Canby, no date); (3) meadow along White Clay Creek,
“rather abundant at this station,” ca. 1 mile north of Newark, 4 Aug 1944 (G. R. Proctor
1113 pers. herb.) and 28 Jul 1945 (R. R. Tatnall 5222, DOV, PH). There is currently one
known extant population, which may be a relocation of the 1944 Proctor and/or the 1945
Tatnall sites: White Clay Creek drainage near Newark, between 1988-91, a small
population, ranging from 1-6 individuals, was observed in an open wet meadow at the edge
of floodplain woods, no plants were observed in 1992, this population was discovered by C.
Pattison (see Ebert et al. 1990).

Polypodium polypodioides (L.) Watt, resurrection a siete (1946) reported this
species to be “infrequent ... from Sussex Co. to... Accomac Co.” It is currently known
from one site: near Laurel, a small population of a me vigorous, spore-producing
individuals, was observed on a tree in 1979 (R. Radis pers. comm.). Field surveys are
needed to determine the current status of this species.

Rhexia aristosa Britt., awned-meadowbeauty, C2. Tatnall (1946) considered this plant to
be “rare and local . . .,” and cited several historical collections: (1) Ellendale, 24 Jul 1893
(Canby s.n., DOV), and several later collections; and (2) at Wilmington, “south side of
Christina Creek,” 21 Sep 1896 (Commons, s.n., PH). R. aristosa is currently known to be
extant from six Sussex Co. sites: (1) Ellendale wet meadow, less than 50 plants were
observed growing in a ditch, it has been monitored at this locality since 1984; (2)
Assawoman Bay Wildlife Area, in a woodland pond (Delmarva Bay) with other rare
species (Coelorachis rugosa, Panicum hirstii, and Sclerolepis uniflora), population numbers
have fluctuated dramatically over the years (1984-1992), discovered by F. Hirst in 1984; (3)
south of Ellendale, between 1988 and 1992 four separate populations have been located, in
ditches in sandy, open pine plantation, ca. 5000 plants were observed in 1991, but fewer
than 100 in 1992; in a clear-cut, ca. 3000 plants in 1991 and 1992; at the edge of an
excavated pond, three plants in flower; and in a ditch, five flowering individuals, discovered
by F. Hirst, R. Wilson, and B. McAvoy. The latter four sites, contain an assemblage of state
rare species (Amphicarpum purshii, Boltonia asteroides, Eleocharis robbinsii, Fuirena pumila,
F. squarrosa, Hypericum adpressum, H. denticulatum, Lobelia canbyi, Rhynchopsora fusca, R.
gracilenta, R. torreyena, Scleria reticularis.) Since all but one of the sites occur in degraded
habitats and because the awned-meadow beauty is a candidate for federal listing, the
species has been ranked S1. ;

Sagittaria calycina Engelm., Mississippi arrow-head. Tatnall (1946) reported that this
species “infrequent in ditches, and tidal mud of Delaware River, in New Castle Co., where
not recently collected. ...” There are currently four known extant sites: (1) Along the
Delaware River near the Delaware Memorial Bridge, more than 100 plants were observed
at the upper-edge of the inter-tidal zone in 1990 by A. E. Schuyler; (2) near New Castle,
along the Delaware River, more than one hundred seedlings were observed in tidal mud
growing with typical marsh species such as Amaranthus cannabinus, Bidens winds
Echinochloa walter, Impatiens capensis, Scirpus pungens, S. tabernaemontanii, and Spartin
alterniflora in 1992 (K. Clancy 2551, with A. E. Schuyler, DNHI Reference Herbarium); 3)
Blackbird Creek, near Blackbird Landing, numerous plants were observed in mud at edge
of freshtidal marsh in 1990 by F. Hirst & R. Wilson; and (4) Drawyers Creek, near DuPont

84 BARTONIA

Hwy, abundant in exposed mud of brackish tidal marsh (K. Clancy 2661, DNHI Reference
Herbarium).

Sagittaria teres Wats., slender arrowhead. This species was reported by Tatnall (1946) as
“rare.” One Delaware collection was cited by Tatnall and later by Tucker et al. (1979):
“Millpond at Milton, (Commons, 17 Aug 1899, PH).” The site of Common’s collection was
likely Wagamons Millpond, where S. teres has been unsuccessfully searched for during the
past 10 years. Presently there is one known site: Reynolds Millpond, north of Milton,
where one plant was noted in 1988 by C. Ludwig. Subsequent searches in 1990 and 1992
failed to relocate this plant (B. McAvoy pers. comm.).

Sanicula marilandica L., black snakeroot. This species was described by Tatnall (1946)
as “rare” on the Delmarva and was known from several historical collections: (1) near
Wilmington, 1896, (W. M. Canby s.n., DOV); (2) near Centreville, 1878, 1898, (A.
Commons s.n., PH); (3) Choptank Mills, 1904, (Stone s.n., PH); and (4) Arden, New
Castle County, in open woods, 1945, (Morris & D. Berd 17514, NY). The latter collection
was cited by Moldenke (1945). There is currently one known extant site: five plants were
observed along Snuff Mill Road near the PA state line by J. Ebert & J. Holt in 1992.

Sanicula trifoliata Bickn., long-fruited snakeroot. This species was not reported by
Tatnall (1946). Brown and Brown (1984) considered this species to be “scarce” and listed
Harford Co. and Baltimore City for its distribution in MD. Harvill et al. (1992) do not list
this species in the Virginia portion of the Delmarva. The one known locality occurs along a
roadside, se of Hoopes Reservoir, where less than 50 mature fruiting plants were observed
in 1989 (Ebert et al. 1990).

Schizaea pusilla Pursh, curly grass fern. This diminutive fern was recently discovered
near Milton where a vigorous population was observed on hummocks and downed logs in
an Atlantic white cedar swamp by F. Hirst & R. Wilson (Hirst 1990). Subsequent surveys
have shown it to be more abundant at this site than previously reported (Hirst pers.
comm.).

Scirpus etuberculatus (Steudel) Kuntze, Canby’s bulrush. According to Tatnall (1946),
Canby’s bulrush was “rare . . . reaches the northern limit of its range in Sussex Co.” It is
known from two historical sites: (1) “. . . ponds, rare,” 5 Aug 1874, (A. Commons s.n., PH);
and (2) Ellis pond, 4.75 mile s of Laurel, 9 Aug 1961 (F. Hirst 59, PH). Surveys between
1989 and 1992 failed to relocate this population, Ellis Pond is in the latter stages of
succession to a shrub swamp and very little open water remains. Canby’s bulrush may have
last been seen at the site in 1972 (A. E. Schuyler pers. comm.). There is currently one
known extant population: Raccoon Pond, several thousand plants were observed growing
in shallow water by C. Ludwig in 1988; surveys in 1990 revealed the presence of only a few,
scattered clumps.

Sclerolepis uniflora (Walter) BSP, pink bog-button. Tatnall (1946) cited one New Castle
Co. location for this species: “Sassafras Crossroads” (=Green Springs Station), 1866, (A.
Commons s.n., DOV, PH).” In Delaware, it usually co-occurs with Boltonia asteroides and
Coreopsis rosea. Recent surveys by Natural Heritage botanists have resulted in its discovery
at five Sussex Co. sites: (1) Assawoman Bay Wildlife Area, two populations occur in
woodland ponds (Delmarva Bays), one has remained relatively stable between 1984 and
1992 forming dense patches, while the other has declined; (2) between Ellendale and
Robbins, less than 100 mature plants were observed in wet spots of a recently re-planted
pine plantation in 1988 by F. Hirst; (3) Ellendale Wet Meadow, a small population of ca.
50 plants was observed in 1991 after being re-discovered by F. Hirst in 1984; and (4) near
Georgetown, several hundred plants were observed in wet deciduous woods by F. Hirst &

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 85

R. Wilson in 1992. In addition to the presence of Sclerolepis, site 4 is also home to the rare
Coelorachis rugosa, Eupatorium leucolepis, Hypericum dentatum, Lobelia canbyi, Polygala
cruciata, Rhynchospora fusca, and Scleria reticularis, to name a few.

Senecio anonymus A. Wood, Small’s ragwort. This species was considered by Tatnall
(1946) to be “infrequent” on the Delmarva. Currently it is known from one site: northeast
of Mt Cuba, near Owls Nest Road, a relatively large population (more than 100
individuals) growing in a mowed lawn underlain by serpentinite was discovered in 1992 by
J. Ebert, J. Holt, & B. McAvoy.

HISTORICAL SPECIES

Selected SH or SX ranked species are discussed below. These species have not been
seen or collected in Delaware for at least 15 years. Taxa classified SX have been searched
for repeatedly at all known historical sites and in all potential habitats. In contrast,
although taxa ranked SH have not been seen or collected for many years, there appears to
be potential habitat yet to survey. In many cases, these species were also believed to be
historical, or extirpated, by Tatnall (1946) and Tucker et al. (1979).

Adlumia fungosa (Ait.) Green ex BSP., climbing fumitory, SX. Tatnall (1946) stated that
this species was “infrequent” in Delaware, while Tucker et al. (1979) thought that it was
“probably extirpated....” Apparently it was never abundant in the state and was
restricted to the rocky banks of the Brandywine Creek: (1) opposite Bancroft’s Factory, 15
Jun 1869 (A. Commons, s.n., PH); and (2) rocky woods at Wills Rocks, below the High
Bridge, 8 Jul 1865, 11 Aug 1897 (A. Commons, s.n., PH).

Aeschynomene virginica (L.) BSP., sensitive joint-vetch, LT, SH. According to Tatnall
(1946) this species was “formerly frequent on tidal shores of the Delaware River, from
Holly Oak to Delaware City.”’ Tucker et al. (1979) considered the species to be “definitely
extirpated.” Historical localities all occur in New Castle Co.: (1) river shores, near
Wilmington, Aug 1878 (W. M. Canby s.n., NY); (2) Holly Oak, 23 Aug 1888 (J. B. Brinton
s.n., NY); and (3) tidewater along Brandywine-Christina Rivers, 1846 (B. Hoopes 486,
NY). Although much of Delaware’s tidal habitat has been degraded, areas requiring more
thorough surveys include the Christina, Appoquinimink, and Nanticoke Rivers and
Drawyers Creek.

Amaranthus pumilus Raf., seabeach pigweed, PT, SH. The sea-beach amaranth is
extirpated from most of the northern half of its range south to Virginia, although it has
reappeared in New York, possibly as a result of severe storms in 1990 that may have
transported propagules from the Carolinas (S. Young pers. comm.), or unearthed a seed
bank (A. Weakley pers. comm.). It was reported by Tatnall (1946) to be “rare, on sea
beaches; southeastern Sussex Co.,” with a single, documented collection: Baltimore
Hundred (area from the Indian River Inlet south to state line), 10 Sep 1875 (A. Commons
s.n., PH). A winter storm that battered Delaware’s coastal beaches in January 1992
brought some hope of its reappearance in the state, but surveys of the beaches from Cape
Henlopen to Fenwick Island in the summer of 1992 failed to relocate this plant.

Arabis drummondii A. Gray, Drummond rock-cress, SH. Tatnall (1946) cited one
Delmarva collection: Concord Station, Wilmington, Jun 1897 (A. Commons, s.n., PH).

Arethusa bulbosa L., dragon’s-mouth, SH. Tatnall (1946) stated that this distinctive
orchid was “rare and local” and Tucker et al. (1979) believed it “definitely extirpated.”
Historical locations include: (1) Farnhurst, McCrones Swamp (now State Hospital), New
Castle Co., 30 May, 2 Jun 1866, 30 May 1882 (A. Commons s.n., PH), and 5 May 1866, 30
May 1882 (W. M. Canby s.n., PH); (2) Millsboro, Sussex Co., sandy swamp, 23 May 1876

86 BARTONIA

(A. Commons s.n., PH); (3) Milford, 5 miles south at margin of Clendaniel Pond (Hudson
Pond?), Sussex Co., 30 May 1928, 11 May 1929, 30 May 1930 (R. R. Tatnall, DOV, PENN);
and (4) near Dagsboro, Sussex Co., Jun 1809 (T. Nuttall s.n., PH).

Arnica acaulis (Walter) BSP, leopard’s-bane, SH. Tatnall (1946) listed this species as
“infrequent; sandy woods and roadsides on the coastal plain ...,” while Tucker et al.
(1979) thought that it was extirpated from the state. Historical collections, all from New
Castle Co., include: (1) ‘““New Castle Co.” Jul 1862 (W. M. Canby s.n., PH); (2) Townsend,
30 May 1890 (J. B. Brinton s.n.), and sandy woods, 11 Jun 1890 (A. Commons s.n., PH); (3)
4.0 mile south of Odessa, DuPont Hwy, meadow, 19 Jun 1933 (H. E. Stone s.n., PH); (4)
5.0 mile south of Odessa, open grass roadside, DuPont Hwy, 0.25 mile south of Union
Church, 3 Jun 1934 (R. R. Tatnall s.n., PH), and border of low woods, near Union Church,
ca. 1.0 mile northwest of Blackbird, 15 Jun 1937 (B. Long 50096, 50097, PH).

Asclepias longifolia Michx., long-leaf milkweed, SH. This species was last collected in
1934 and according to Tatnall (1946) was “very rare ... now apparently extinct in our
area.” Tucker et al. (1979) considered it to be “definitely extirpated.” It was known from
one location: Ellendale, in ditches along the railroad, south of the village (DOV, GH, PH).

Asclepias quadrifolia Jacq., whorled milkweed, SH. Tatnall (1946) considered this
distinctive milkweed “infrequent, in dry hilly woods of the Piedmont; rare on the Coastal
Plain ...,” while Tucker et al. (1979) believed that it had been extirpated. Historical
collections include: (1) near Centreville, dry woods, Jun 1865 (A. Commons, s.n., PH); (2)
Mt. Cuba, 3 Jun 1894 (J. B. Brinton, s.n., PH) and 30 May 1895 (I. A. Keller, s.n., PH); and
(3) below Yorklyn along the Red Clay Creek, loamy wooded slope, 24 May 1924 (B. Long
30251, PH).

Asplenium rhizophyllum L., walking fern, SH. According to Tatnall (1946) this plant was
“rare, on wooded rocky slopes of the Piedmont.” Tucker et al. (1979) considered it to be
“definitely extirpated.” Historical collections include: (1) Mt. Cuba, Jul 1865 (A.
Commons, s.n., PH); near foot of rocky slope, just above deep railway cut, 0.5 mile north of
Mt. Cuba 27 Jun 1931 and 17 April 1942 (R. R. Tatnall 1195, 5004 DOV); (2) near
Centreville, 13 Jun 1877 (A. Commons, s.n., PH); (3) White Clay Creek Valley, reported
by Linehan et al. 1970 (cited in Fleming 1978); and (4) high up the rocky slope, along east
side of Brandywine Creek above Rockland, 24 Mar 1929 (R. R. Tatnall 232, DOV).

Carex bicknellii Britton, Bicknell sedge, SX. This sedge was known from a single
Delaware location: near Centreville, on serpentine barren, dry soil, 26 May 1863 (A.
Commons, PH).

Carex polymorpha Muhl., variable sedge, C2, SH. Tatnall (1946) stated there was one
known collection of this species on the Delmarva (Cecil Co., MD). Apparently he
overlooked the only known Delaware collection: “Sussex Co.,” 1874 (W. M. Canby s.n.,
DOV). Because this sedge grows in xeric to mesic woods, it is hoped that it will yet be
relocated in the state.

Carex striatula Michx., lined sedge, SH. Tatnall (1946) considered the species to be
“rare; wooded banks, Piedmont and Coastal Plain. . . .” Historical locations include the
following: (1) Mt. Cuba, rocky woods, 26 May 1884, 19 Jun 1895 (A. Commons s.n., PH),
and 4 Jul 1903 (W. Stone s.n., PH); (2) Greenbank, woods, 27 May 1879 (A. Commons s.1.,
PH); (3) 1.5 mile north of Choate, road bank, edge of woods along Pike Creek, 18 May
1940 (R. R. Tatnall 4495, DOV, PH); and (4) Wilmington, May 1893 (W. M. Canby s.n.,

DOV).
Chamaedaphne calyculata (L.) Moench., leatherleaf, SH. Tatnall (1946) stated that
leatherleaf was “‘...not recently found within our limits, in spite of careful search.”

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 87

Tucker et al. (1979) considered it “definitely extirpated.” Historical locations: (1)
Thompson’s Swamp, 1.5 mile northwest of New Castle, 1858 (W. M. Canby s.n., DOV); (2)
Frankford, 1875 (A. Commons s.n., PH); and (3) Townsend, sandy swamps, 11 Jun 1890
(A. Commons s.n., PH).

Cheilanthes lanosa (Michx.) D. C. Eat., hairy lip-fern, SX. Tatnall (1946) reported that
this species was “rare, on rocky hills ... in and near Wilmington, where now extinct.”
Tucker et al. (1979) considered the hairy lip-fern to be “definitely extirpated.” Historical
collections include: (1) near Wilmington, 1861 (I. Hoopes, s.n., PH); (2) Wilmington, east
side of Brandywine Creek below Jessup’s papermill (site now destroyed), rocky ridge, 13
Sep 1864 (A. Commons, s.n., PH); and (3) Wilmington, banks of Brandywine opposite
Bancroft’s Factory, on rocks, 3 Dec 1889 (A. Commons, s.n., PH). All three of the above
collections may have been made at the same location. As shale outcrops, the habitat that
typically supports this species, are lacking in Delaware, it is doubtful that this fern will be
relocated.

Cicuta bulbifera L., bulb-bearing water hemlock, SX. Although Tatnall (1946) did not
specifically comment on the abundance or rarity of this species, Tucker et al. (1979)
thought that it was “definitely extirpated.” There is one historical site: New Castle Co., wet
ground at top of north bank of C & D Canal, west of Summit Bridge, 24 Aug 1869 (A.
Commons, s.n., PH).

Cleistes divaricata (L.) Ames, spreading pogonia, SH. Tatnall (1946) considered this
species “rare” and Tucker et al. (1979) thought it “probably extirpated.” Historical
collections, all in Sussex Co. include: (1) meadow and thicket-pasture east of Ellendale, 22
Jul 1908 (B. Long s.n., PH), and 7 Sep 1938, 12 Jun 1939 (A. V. Smith & R. R. Tatnall s.n.,
DOV); (2) 3 miles northwest of Rehoboth, in ditch along railroad, 4 Jul 1930, 5 Sep 1936
(R. R. Tatnall s.n., DOV); and (3) near Dagsboro, Jun 1809 (T. Nuttall s.n., PH).

Crassula aquatica (L.) Schonl., pygmy-weed, SX. Tatnall (1946) stated that this species
was “rare, on muddy shores .. .” and listed one historical site: Brandywine Creek above
railroad bridge, collected by Canby in 1867, but no specimen was seen. Ferren and
Schuyler (1980) stated that it “has not been collected in the Delaware system for over 50
years.”

Cyperus plukenetii Fern., a flatsedge, SX. Tatnall (1946) considered this species to be
“rare, in dry ground. ...” Historical localities include: (1) Rehoboth, Sussex County, 13
Aug 1896 (A. Commons s.n., PH); (2) Millsboro, 21 Sep 1907 (S. Brown, PH); and (3)
Wilmington, Giant’s Cave, on Brandywine, 28 Jul 1866 (A. Commons s.n., PH).

Cyperus tenuifolius (Stuedel) Dandy, thinleaf flatsedge, SH. Tatnall (1946) cited one
known collection from the Delmarva: “moist soil,” Baltimore Hundred (Sussex Co., south
of Indian River Inlet), 10 Sep 1875 (A. Commons s.n., PH).

Desmodium humifusum (Muhl.) Beck, spreading tick-clover, C2, SH. Tatnall (1946)
reported the species to be “rare, in sandy soil.” Historical collections, all in New Castle
Co., include: (1) sandy woods near Collins Beach, 27 Aug 1867 (A. Commons s.n., NY); (2)
wooded hillside of Red Clay Creek, near Delaware Western RR tank, Sep 1879 and 1 Oct
1879 (W. M. Canby s.n., NY); (3) cedar swamp, 9 Aug 1866 (A. Commons s.n., NY); and
(4) New Castle City, rare, 27 Aug 1867 (A. Commons s.n., NY). Considering that it is
known to occur in disturbed habitats (i.e. along powerline clearings) it may yet be
relocated in Delaware.

Eriophorum gracile Koch ex Roth, slender cotton-grass, SX. This species was thought to
be “rare ...” in Delaware (Tatnall 1946). Historical locations include: (1) Thompson’s
swamp, 1.5 miles northeast of New Castle (E. Tatnall, 1858, DOV); (2) McCrone’s swamp,

88 BARTONIA

near Farnhurst, 20 Jun 1876 (A. Commons s.n., PH); and (3) in sphagnum, east of
VanDyke, 7 Jun 1881 (A. Commons, s.n., PH). The State Hospital now occupies the
former McCrone’s swamp site, and Thompson’s swamp has been destroyed (or severely
degraded) due to expansion of Wilmington and New Castle. The best area for relocating
this species is the wetlands near Vandyke.

Eupatorium resinosum Torr., pine barrens boneset, C2, SH. Tatnall (1946) considered
the pine-barrens boneset to be “rare, in low pine barrens . . .,” while Tucker et al. (1979)
stated that this species was “‘definitely extirpated.” There are two historical sites: (1) in low
pine barrens near Gumboro, 5 Aug 1874 (A. Commons, s.n., PH); and (2) Baltimore
Hundred (south of Indian River Inlet), 10 Sep 1875 (A. Commons, Sti, PE).

Euphorbia sg sealei (Raf. ) Fern., Darlington’s or glade spurge, C2, SH. Tatnall (1946)
stated that it was “rare, in the Piedmont Province,” while Tucker et al. (1979) considered it
to be ‘definitely extirpated.” There is one known historical location: swamp west of
Hockessin, 8 Jun 1881 (A. Commons s.n., PH). This site is in the northwest part of the state
within 10 km of an extant PA population (Pennsylvania Natural Diversity Inventory pers.
comm.).

Fimbristylis annua (All.) Roemer & Schultes, annual fimbry, SX. Tatnall (1946) stated
that the annual fimbry was “rare, known from serpentine soil of New Castle Co., ... no
recent collections.” Tucker et al. (1979) thought that it was “possibly extirpated.”
Historical location: near Centreville, New Castle Co., moist soil (serpentine barrens), Sep
1972 (A. Commons s.n., PH

Gentiana autumnalis L., pine barren gentian, SH. Tatnall (1946) cited one historical
location: Sussex Co., pine barrens, 10 Sep 1875 (A. Commons, s.n., PH). Botanists should
continue to search for this species in the moist pinewoods of southern Delaware.

Gentianopsis crinita (Froelich) Ma., fringed gentian, SH. Tatnall (1946) reported this
species “rare, New Castle Co. . . . now extinct in most or all of the above localities.” Tucker
et al. (1979) reported that it was “definitely extirpated.’ Collections include: (1) near
Centreville, 1865, 1878 (A. Commons s.n., PH); (2) Faulkland, 24 Sep 1886 (R. R. Tatnall
s.n., DOV); (3) Southwood, Oct 1894 (W. M. Canby, DOV); (4) Brandywine Springs, 8
Oct 1896 (W. M. Canby, DOV); and ©) roadside bank one mile west of Centreville, 12 Oct
1928 (R. R. Tatnall 221, DOV, PEN

Hydrangea arborescens L., heehee hydrangea, SX. Tatnall (1946) reported that this
species was “... rare in New Castle Co.” It was known from one site: Mt. Cuba, 13 May
1892 (J. Crawford s.n., PH).

Lobelia boykinii T. & G., Boykin’s lobelia, C2, SH. Tatnall (1946) considered the species
“rare; meadows and ditches ...,”’ and Tucker et al. (1979) reported it “probably
extirpated.” It was known from one location: Ellendale, 24 Jul 1893 (W. M. Canby, s.n.
DOV); ditches along RR, 9 Jul 1908, 20 Jun 1909 (C. S. Williamson, s.n., PH), and 6 Jul
1913 (Long & Bartram PH).

Lophiola aurea Ker Gawler (=L. americana (Pursh) Wood), gold-crest, SH. This species
was cited by Tatnall (1946) as “very rare... .” There is one historical site: Sussex Co.,
pine-barren bogs between Gumboro and Laurel, 5 Aug 1874 (A. Commons, s.n., PH).

Lygodium palmatum (Bernh.) Sw., climbing fern, SH. This distinctive fern was known
from two historical collections in New Castle Co., and more recently, from southwestern
Sussex Co.: (1) border of wooded swamp ca. 1 mile north of streamlet tributary to
Christina Creek, 22 Apr 1933 (B. Long s.n., CHRB); (2) woods along bank above Christina
Creek 1.25 miles wnw of Bear Station (A. W. Leeds s.n., CHRB); and (3) low woods near
Woodland Ferry (discovered by R. Radis in 1976, pers. comm.). The two New Castle

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 89

collections may represent the same locality reported by Tatnall (1946). The Sussex Co. site
needs to be revisited to determine the current status of this plant.

Menyanthes trifoliata L., buckbean, SX. Tatnall (1946) stated that the species was
“formerly rare in swamps of New Castle Co. . . . now probably extinct in our area.”’ Tucker
et al. (1979) thought it “probably extirpated.” It was known from two locations: (1)
Thompson’s Swamp, nw of New Castle, 1840 (E. Tatnall s.n., DOV); and (2) Cedar
Swamp, 1866 (Tatnall 1946). Thompson’s Swamp has been virtually destroyed, but the
Cedar Swamp area still has extensive “intact’’ habitat.

Micranthemum micranthemoides (Nutt.) Wettst., Nuttall’s micranthemum, C1*, SX.
This species is presumed to be extinct and has not been seen anywhere since 1941 (The
Nature Conservancy 1987). According to Tatnall (1946) it was “infrequent ... in New
Castle Co.,” and was known to occur “near Claymont, Delaware River between tides,” 3
Oct 1866 (A. Commons s.n., DOV).

Monotropsis odorata Elliott, sweet pinesap, SH. Tatnall (1946) did not mention the
species as occurring on the Delmarva. In Maryland it is known from Anne Arundel and
Baltimore counties (G. Cooley pers. comm, Brown and Brown 1984). Apparently, there
were two Delaware collections: (1) “Delaware,” no date (E. Durand s.n. US 968638); and
(2) “Delaware,” no date (E. Durand s.n., PH). These may have been made in the 1860s,
since Durand is known to have collected M. odorata in Maryland in 1867.

Muhlenbergia torreyana (Schultes) A. Hitche., Torrey’s dropseed, SH. This grass was
considered “rare” by Tatnall (1946), while Tucker et al. (1979) thought it “definitely
extirpated. ...” Historical sites: (1) Felton, Kent Co. 25 Sep 1873, 25 Sep 1875 (A.
Commons, s.n., NY, PH, US); and (2) bogs 0.5 mile east of Ellendale at RD 231, Sussex
Co., Sep 1875 (W. M. Canby, s.n., NY); 27 Sep 1895 (A. Commons, s.n., PH); 12 Oct 1940
(A. Chase 12619, US); and also 12 Oct 1940 (R. R. Tatnall 4473, GH).

Narthecium americanum Hudson, bog-asphodel, C1, SX. Tatnall (1946) believed the
species “rare ...” while Tucker et al. (1979) stated that it was “definitely extirpated.”
Historical locations include: (1) swamp near Lewes, Sussex Co., 1 Aug 1895 (A. Commons
s.n., PH); and (2) damp railroad bank at Vandyke, New Castle Co., 26 Sep 1894 (Tatnall
1946).

Ophioglossum vulgatum L. var. pseudopodum (Blake) Fawr., northern adder’s tongue,
SH. Tatnall (1946) considered this fern to be “rare,” while Tucker et al. (1979) state that
this species is “definitely extirpated.” Historically, it is known from one New Castle Co.
locale: near Centreville, moist place in woods, 14 Jul 1873 (A. Commons s.n., PH).

Oxypolis canbyi (Coult & Rose) Fern., Canby’s dropwort, LE, SH. This plant was last
collected in Delaware in 1894. The historical site, the type locality, was the meadows and
bogs at Ellendale. According to Dill and Tucker (1986) a total of seven field trips to the site
from 1867 to 1894 produced 23 herbarium sheets. Its recent discovery in Maryland (Boone
et al. 1984) has renewed hope that O. canbyi will be rediscovered in Delaware.

Polygala paucifolia Willd., gay-wing mikwort, SX. Tatnall (1946) considered this plant
“very rare, in the Piedmont of New Castle Co. . . .,” while Tucker et al. (1979) thought it
“possibly extirpated.” It was known from a single locality: 0.25 mile north of Mt. Cuba
Station, on railroad bank, May 1890 (W. Tatnall, Jr. s.n., DOV) and many later collections
deposited at DOV, PENN, PH, and US.

Polygala senega L., seneca snakeroot, SX. Tatnall (1946) stated of this species “formerly
rather frequent in rocky woods of the Piedmont, now rare or perhaps extinct in our range.
Tucker et al. (1979) also thought P. senega extirpated. Historical collections include: (1)
Wilmington, May 1890, (W. M. Canby, s.n., PH); (2) Mt. Cuba, rocky woods, 15 & 21 Jun

90 BARTONIA

1865, 27 May 1873 (A. Commons, s.n., PH); and (3) Guyencourt, south of P. & R. Station,
tributary of Brandywine Creek, loamy wooded slopes, 23 Jun 1923 (B. Long 27524) and 1
Jun 1924 (H. Williams, s.n. PH).

Rhynchospora knieskernii Carey, Knieskern’s beakrush, LT, SH. This species is only
known from Delaware and New Jersey (US Fish and Wildlife Service 1991b). Both
historical localities were in Sussex Co.: (1) swamp near Gumboro, 5 Aug 1874 (A.
Commons s.n., PH); and (2) Baltimore Hundred (south of Indian River Inlet), 10 Sep 1875
(A. Commons s.n., PH). Considering that R. knieskernii is found along powerline
right-of-ways and other disturbed habitats in New Jersey (D. Snyder pers. comm.), there is
still hope that it may be relocated in Delaware.

Rhynchospora oligantha A. Gray, few-flowered beakrush, SH. Tatnall (1946) considered
this species to be “rare,” while Tucker et al. (1979) thought it “possibly extirpated.” There
are two Sussex Co. collections, perhaps from the same site: (1) bogs, 1.5 mile south of
Lewes, Jul 1878 (W. M. Canby s.n., DOV, PH); and (2) near Lewes, 26 Aug 1895 (A.
Commons s.n., PH).

Schwalbea americana L., chaffseed, LE, SH. Tatnall (1946) considered the species
“rare” and Tucker et al. (1979) stated that it was “definitely extirpated.” Collections, all
from New Castle Co. include: (1) St. Georges, Jun 1866 (W. M. Canby s.n., NY), along the
canal (or south side of canal, below St. Georges), St. Georges, 22 Jul 1875 (A. Commons
s.n., NY, PH); and (2) on a hill south side of C & D Canal, 1.0 mile below the crossing of
the Delaware Railroad, Jun—Aug 1862 (W. M. Canby s.n., NY). Most of the habitat in this
area has been destroyed or degraded during the past 100 years and there is little hope that
the chaffseed will be relocated.

Scutellaria saxatalis Riddell, rock-skullcap, C2, SH. Tatnall (1946) reported the species
to be “rare, in rocky woods” and Tucker et al. (1979) thought it “probably extirpated.”
Historical sites were in New Castle Co.: (1) stony places above Rockford, 10 Jun 1836 (E.
Tatnall s.n., PH); (2) Brandywine, 1888 (W. Tatnall s.n., PH); and (3) Wilmington, rocky
woods, 19 Jun 1897 (A. Commons s.n., PH).

Solidago uliginosa Nutt., bog-goldenrod, SH. Tatnall (1946) stated that this species was
“infrequent, in bogs, Coastal Plain, New Castle, Sussex, . . . counties,” and Tucker et al.
(1979) thought it was extirpated. All of Delaware’s collections were from New Castle Co.:
(1) swamps, New Castle, no date (W. M. Canby s.n.); (2) McCrone’s swamps, near
Wilmington, 10 Oct 1876 (A. Commons s.n.); (3) Wilmington, 10 Sep 1890 (A. Commons
s.n.); (4) Pencader Hundred, Porter’s Station, in dry soil, 9 Oct 1890 (A. Commons s.n.);
(5) Kiamensi, swamps, 8 Aug 1894 (A. Commons s.n.); (6) swamps, near Townsend, 21 Sep
1894 and 9 Oct 1896 (A. Commons s.n.); and (7) swampy margins of woods, southwest of
Vandyke, 9 Oct 1908 (B. Long s.n.), all collections at PH.

Tofieldia racemosa (Walter) BSP, coastal false-asphodel, SH. This species was reported
by Tatnall (1946) to be “rare, in wet ground or shallow water ...,” while Tucker et al.
(1979) considered it “probably extirpated.” Historical collections may all refer to the same
locality: (1) “Glades south of Leweston,” and swamp 1.0 mile south of Lewes, Sussex Co.,
13 Jul, 22 Jul 1878 (W. M. Canby s.n., PH, NY); (2) near Lewes, 15 Aug 1895 (A. Commons
s.n., PH); (3) three miles northwest of Rehoboth, 14 Aug 1923 (Otis s.n. GH, PH), 5 Sep
1936, and 5 Aug 1937 (R. R. Tatnall 3173, 3461, DOV, GH); and (4) wet ground on east
side of railroad, 0.75 mile northwest of Howlands Glade bridge (Otis s.n., GH, PH).

Triphora trianthophora (Swartz) Rydb., three-birds orchid, SH. This species has not been
collected in Delaware since before 1900 (Tucker et al. 1979). Tatnall (1946) considered it

SELECTED RARE AND HISTORICAL VASCULAR PLANTS OF DELAWARE 91

“rare and local; meadows and low woods of the Piedmont.” The one historical collection
was from Hockessin, rich woods, no date (Tatnall 1946).

Xerophyllum asphodeloides (L.) Nutt., turkeybeard, SH. This species was known from
one historical site: dry pine woods near Laurel, Sussex Co., 5 Aug 1874 (A. Commons s.n.,
PH)

ACKNOWLEDGEMENTS

I would like to extend my sincere appreciation to all who have contributed to this paper
and to furthering our knowledge of Delaware’s rare plants, particularly those individuals
who have carried out rare plant surveys: J. Ebert, J. Holt, F. Hirst, C. Ludwig, B. McAvoy,
R. Radis, T. Rawinski, W. Rittenhouse, and A. E. Schuyler. A special thanks are also due
to B. McAvoy and D. Rothstein for their encouragement and support during the writing of
this paper. Thanks go to B. McAvoy and A. E. Schuyler for reviewing previous drafts of this
manuscript. Finally, I would like to thank the curators of CHRB, GH, NY, PENN, PH, and
US for providing access to their collections.

LITERATURE CITED

Boong, D., G. FENWICK, AND F. Hirst. 1984. The rediscovery of Oxypolis canbyi on the Delmarva Peninsula.
Bartonia 50: 21-22.

Brown, M. L., AND R. G. BRown. 1972. Woody plants of Maryland. Port City Press, Baltimore.

Brown, M. L., AND R. G. BRown. 1984. Herbaceous plants of Maryland. Port City Press, Baltimore.

CLANCY, K. 1993. The yellow passion-flower, Passiflora lutea L., rediscovered in Delaware. Castanea, in press.

DNHI 1993. Unpublished database. Delaware Department of Natural Resources and Environmental Control,

over.

DILL, N. H., AND A. O. TUCKER. 1982. Areas of floristic diversity tion of plants in Delaware. Claude
E. Phillips Herbarium. Dover.

DILL, N. H., AND A. O. TUCKER. 1986. Delaware rare plant survey; 1985. Prepared by Claude E. Phillips
Herbarium, Dover.

EBERT, J., J. Hott, A. NEWBOLD, AND A. E. sesapes 1990. Delaware rare plants survey—finding and
relocating rare plants of the Piedmont of Delaware. Bartonia 56: 65-67.

FERREN, W. R., AND A. E. SCHUYLER. 1980. snip vascular plants of river systems near Philadelphia.
Proceedings of the Academy of Natural Sciences of Philadelphia 132: 86-120.

FLEMING, L. 1978. Delaware’s outstanding natural areas and their preservation. Delaware Nature Education
Society, Hockessin.

GLEASON, H. A., AND A. CRONQUIST. 1991. an e vascular plants of northeastern United States and adjacent
Canada. The New York Botanical Garden, B

—— A. M., JR., T. R. BRADLEY, C. E. emi "7. F, WiEBOLDT, D. E. WARE, D. W. OGLE, G. W. RAMSEY,

DG. P. FLEMING. 1992. Atlas of the Virginia flora: III. Privately published. ae

aching F. 1983. Field report on the Delmarva flora. Bartonia 49: 59-68.

Hirst, F. 1990. Three new taxa for the Delmarva Peninsula. Bartonia 56: 70-71.

KRAL, R. 1983. A report on some rare, threatened, or endangered forest-related vascular plants of the south. 2
Vols. Technical Publication R8-TP 2. United States Department of Agriculture, U. S. Forest Service
publication. Atlanta.

LEONARD, S. W. 1981. Fimbristylis perpusilla Harper in South Carolina. Castanea 46: 235-236.

LEONARD, S. W. 1987. Fimbristylis perpusilla in North Carolina. Castanea 52: 150

McAvoy, W. 1993. Rare Native Plants of Delaware. Delaware Natural Heritage Inventory, Department of
Natural Resources and Environmental Control, Dover.

sina areni H. N. 1945. A contribution to our knowledge of the wild and cultivated flora of Delaware. Torreya

106-109.
NAczI, R. F.C., R. J. DRIscout, E. L. PENNELL, N. E. SEYFRIED, A. O. TUCKER, AND N. DILL. 1986. New records
of some rare Delmarva sedges. Bartonia 52: 49-57.
Puitups, C. E. 1978. Wildflowers of Delaware and the Eastern Shore. Delaware Nature Education Society.
Hockessin

92 BARTONIA

TATNALL, R. R. 1946. Flora of Delaware and the Eastern Shore. The Society of Natural History of Delaware,
Wilmington

THE NATURE CONSERVANCY. 1987. Micranthemum micranthemoides; element global ranking form. Prepared by
L. E. Morse and E. Roth. Arlington

Tucker, A. O., N. H. DILL, C. R. BROOME, C. E. PHILLIPS, AND M. J. MACIERELLO. 1979. Rare and endangered
vascular plant species in Delaware. Prepared by the Society of Natural History of Delaware, in cooperation
with the U. S. Fish and Wildlife Service.

US FisH AND WILDLIFE SERVICE. 1991a. Endangered and Threatened Wildlife and Plants. 50 CFR 17.11 and
17.12.

US FisH AND WILDLIFE SERVICE. 1991b. Endangered and threatened wildlife and plants; determination of the
plant Rhynchospora knieskernii (Knieskern’s beakrush), to be a threatened species. Federal Register 56:
32978-32983.

US FisH AND WILDLIFE SERVICE. 1992a. Endangered and threatened wildlife and plants; determination of
threatened status for the joint-vetch (Aeschynomene virginica). Federal Register 57: 21569-21574.

US FIsH AND WILDLIFE SERVICE: 1992b. Endangered and threatened wildlife and plants; proposed threatened
status for the plant A (sea-beach amaranth). Federal Register 57: 21921-21925.

US FisH AND WILDLIFE SERVICE. 1992c. Fadsuaeve and Threatened wildlife and plants; determination of
endangered status for the plant Schwalbea americana (American chaffseed). Federal Register 57: 44708.
WoororpD, B. E., AND R. L. JONEs. 1988. ai perpusilla Harper (Cyperaceae) from the Cumberland

Plateau of Tennessee. Castanea 53: 299-302

es
ees

a

ate

‘Z
eth ate

bat

ISSN 0198-7356

BARTONIA

JOURNAL OF THE
PHILADELPHIA BOTANICAL CLUB

No. 58
CONTENTS
Satellite Remote Sensing of Pinelands Ecosystems .............. RICHARD G, LATHROP, JR. 1
Mosses of the New Jersey Pine Barrens and Adjacent Coastal Plain ......... Eric F. KARLIN 11

Interactions Between Research and Management: Examples from the New Jersey Pine Barrens
EMILY W. B. RUSSELL 23

Taxonomy of Scirpus, Trichophorum, and Schoenoplectus (Cyperaceae) in Virginia
MARK T. STRONG 29

Noteworthy Plant Collections from Pennsylvania .. ALLISON W. CUSICK AND SUE A. THOMPSON 69
Magnolia twipetala tn Pennsylvatia’ . - 5 30 i ee

Additions, Range Extensions, Reinstatements, and Relocations in the New Jersey Flora
DAVID B. SNYDER 79

The Vegetation of Little Beach Island, New Jersey ............-...---+ RICHARD STALTER 97
The Vegetation of the Glades Region, Cumberland County, New Jersey .. STEVENS HECKSCHER 101

Ecological Observations of Bear Swamp, Cumberland County, New Jersey
RICHARD STALTER AND DANIEL O'CONNOR [15

Juncus caesariensis Coville (New Jersey Rush) in Nova Scotia, Canada
RUTH E. NEWELL AND REG. B. NEWELL 121

CRGmaV ies a i ee ee ee 125
POV NCWS a as ek a a ee 130
Pies snd Notes: = os a a ee ee 138
EO5t Wield Tring 2 ie a a a ee 141
ROOD Wield Tite os a a ae a ee 148

155

Mbrmaivaralley List. = oo yn ee as eee
Program of Meetings 1991 and 1992 ........ 2-2. se eee ee reece etn eens
PUBLISHED BY THE CLUB
ACADEMY OF NATURAL SCIENCES, 1900 BENJAMIN FRANKLIN PARKWAY
PHILADELPHIA, PENNSYLVANIA 19103-1195
Issued 28 June 1994

The Philadelphia Botanical Club

Editor: Alfred E. Schuyler
Editorial Assistant: Elizabeth B. Farley

Editorial Board
DAVID E. FAIRBROTHERS CARL S. KEENER
WILLIAM M. KLEIN, JR. JAMES D. MONTGOMERY
RONALD L. STUCKY
Officers of the Philadelphia Botanical Club for 1991-1993
President: TED GORDON Vice President: ROBERT J. HOLT

Treasurer: ROY L. HILL, JR. Recording Secretary: WILLIAM F. OLSON
Curator: HANS WILKENS Corresponding Secretary: ELIZABETH B. FARLEY

MISSOURI BOTANICAL

AUG 1 4 1998

Bartonia No. 58: 1-9, 1994
P F GARDEN LIBRARY
Satellite Remote Sensing of Pinelands Ecosystems

RICHARD G. LATHROP, JR.
Department of Environmental Resources, Cook College—Rutgers University
New Brunswick, NJ 08903-0231

Satellite remote sensing has found increasing application to the investigation of ecological
systems at landscape and regional global scales. The synoptic and repetitive nature of
satellite remote sensing provides a consistent data set to monitor the earth’s surface at
different spatial scales and at different temporal frequencies. Passive remote sensing systems
measure reflected or emitted energy in the visible, near, middle and thermal infrared (IR)
wavelength regions of the electromagnetic spectrum. Each of these different wavelength
regions provides different information as to the state of the Earth’s surface.

In forested regions, as in the case of the Pinelands, the observed remotely sensed signal
is largely determined by the state of the forest overstory vegetation: the community type; the
density/biomass; and the leaf area (Peterson and Running 1989). Other factors also have an
influence: the understory vegetation; the background soil/litter reflectance; and the
topographic slope and aspect. Remote sensing data provides information on the spatial
patterns of the vegetation or ecosystem state; the challenge is to translate the information on
pattern into information on process at either the ecosystem or landscape level.

This paper reports on an investigation of the utility of Landsat Thematic Mapper (TM)
remote sensing imagery for classifying and mapping the major Pinelands vegetation
communities. The objectives of this study were: 1) to gain a better understanding of the
factors determining the remotely sensed signal observed for Pinelands vegetation; 2) to
classify and map the major forest community types in the Pinelands National Reserve
Preservation area as of 1988; and 3) to document spatial patterns in forest vegetation type.

METHODS

This study focussed on the central core of the Pinelands National Reserve Preservation
Area (Fig. 1). The image used for this study was acquired by the Landsat-5 TM on 8
August 1988 (Path 14/Row 32; Scene ID Y5162815103X0). The TM sensor has seven
bands spanning the visible, near IR, middle IR and thermal IR spectral regions. Bands 1-5
and 7 are acquired at a 30 m ground cell resolution, Band 6 (Thermal IR) is acquired at a
coarser 120 m grid cell resolution. It should be noted that the image was acquired during
the height of a severe drought period. A subset of cloud-free data of high radiometric
quality was extracted for analysis. The study area contained a diversity of forest
communities typical of NJ Pinelands conditions (McCormick 1979).

A Geophysical Environmental Research Corporation Mark V spectroradiometer was used
to measure the spectral radiance (from 0.4 to 2.5 pm) of clipped pitch pine (Pinus rigida)
and white cedar (Chamaecyparis thyoides) foliage in a laboratory setting. This data was
compared to the spectral radiance curves extracted from the Landsat TM imagery subsets of
relatively homogeneous pitch pine and white cedar forest stands, as well as additional forest
community types.

BARTONIA

NEW YORK CITY

OCEAN

PHILADELPHIA,

N.J. Pinelands
National Reserve

Study Area

CJ

—

Delaware
Bay @) 25
_e_
km
Map of NJ Pinelands National Reserve and study area

Fic. 1.

PINELANDS REMOTE SENSING 3

TABLE 1. Land Cover/Vegetation Types.

1. Pine/oak forest — >50% Pinus rigida 5. Pitch pine lowland forest — Pinus rigida/heath
2. Oak/pine forest — >50% Quercus spp. understo

3. Cedar swamp — Chamaecyparis thyoides 6. Non-Pine Barrens vegetation

4. Hardwood swamp — Acer rubrum, Nyssa sylvatica 7. Water

All digital image analysis for this study was undertaken at the Cook College Remote
Sensing Center using ERDAS image processing and GRASS geographic information system
software. Supervised classification techniques were used to classify the image into
appropriate land cover/community types (Lillesand and Kiefer 1986). A classification scheme
was used that was similar to that employed by the NJ Pinelands Commission to map
vegetation in 1979 (Pinelands Commission 1980) (Table 1). Aerial photo-interpretation and
ground reconnaissance were used to select training sites (representative samples) of each land
cover/vegetation type.

The results of the supervised image classification were then compared to visual
interpretation of the TM image (in conjunction with aerial photographs and the Pinelands
Commission vegetation maps) at approximately 500 random points. Due to particular
concern over the status of the cedar swamps in the Pinelands National Reserve (Roman et
al. 1987), the utility of TM imagery to delineate these critical ecosystems was further
analyzed. Those areas classified as cedar swamps were extracted as a separate digital map
layer and compared with a digital map of cedar swamps produced by the Division of Parks
and Forestry of the NJ Department of Environmental Protection. The DEP classified cedar
swamps into 3 classes: <50% cedar; >50% cedar; and >95% cedar. The DEP’s map was
based on a visual interpretation of 1986 aerial photographs. A subarea of the Pinelands
Reserve Preservation area and a smaller subarea centered on the Great Swamp in the Mullica
River Drainage were selected for analysis.

RESULTS

Spectroradiometric analysis of the clipped pitch pine and white cedar foliage shows that
the two species have very similar spectral response curves in the 0.4-2.5 um wavelength
range (Fig. 2). In contrast, the spectral radiance curves of selected forest stands from the
Landsat TM imagery shows a greater difference between the two vegetation types (Figure
3). In the near IR wavelength range, pine shows a 33% higher response over cedar in the
TM data (Fig. 3) as compared to a 6% higher response in the clipped foliage (Fig. 2). This
difference between handheld and satellite sensor observations illustrates the fact that in
addition to leaf spectral properties, the satellite observed signal is also a function of the
canopy structure and background reflectance. White cedar swamps appear extremely dark
on the image; "black holes" that effectively trap solar radiation. The high va alues in TM
Band 1 — at 0.49 um) (Fig. 3) are due to atmospheric scattering adding to the earth
surface reflectan

Further as of Landsat spectral radiance curves shows that deciduous forest stands
(e.g. oak and hardwood swamps), as expected, have much higher reflectance in the near and
middle IR as compared to the coniferous forest stands (Fig. 3). Pitch pine lowlands have
an intermediate response between pine uplands and white cedar swamps. The pres-
ence/absence of vegetation greatly influences the satellite observed signal. Fig. 4 illustrates
the transition from bare soil to a sparsely vegetated section of the Pine Plains to a closed
canopy pitch pine stand. As the vegetation canopy cover increases, the red (TM Band 3,

4 BARTONIA

rs actions. Bl is
/ Vat
= j \ moe, Sat
1 47 i \
‘C |
<b)
oe
w
SS
" a be
E {
- \
= \
E 27 \
Py \
e \
= \
x = |
oO 14 \ ee
[a - a
L a.
UD T tf T T 1
0.40 0.82 1.24 1.66 2.08 2.50

Wavelength (micrometers)
Fic. 2. Plot of spectral radiance vs. wavelength for clipped foliage of Pinus rigida and Chamaecyparis thyoides.

centered at 0.66 um) reflectance decreases due to chlorophyll absorption, the near IR (TM
ae 4, centered at 0.83 um) reflectance increases due to intra-leaf scattering and the middle

R (TM Band 5, Si at 1.65 um) reflectance decreases due to cellular water absorption
rae and Runn 1989).

Ellipse plots of ie training set data show similar relationships as above (Fig. 5). In
closed canopy forest stands, there is little difference in red reflectance (TM Band 3) but a
gradient in the near IR (TM Band 4) from cedar swamps at the lower extreme to oak forests
and hardwood swamps at the upper extreme (Fig. 5a). The response from mixed pine and
oak stands is a function of the relative amounts of the darker pine versus the brighter oak
in the overstory. In the middle IR (TM Band 5), the pitch pine lowlands generally show a
lower response than upland pine/oak forests (Fig. 5b). This lower response may be due to
a variation in the understory (e.g. Kalmia angustifolia versus Quercus ilicifolia) rather than
in the overstory.

One unexpected result was the discrimination provided by the thermal IR (TM Band 6,
si gee on 11.5 um) (Fig. Sc). The lowland forest types, the cedar and hardwood swamps
and to a lesser extent the pitch pine lowlands, showed significantly cooler thermal IR
cater than the upland forest types. This difference in thermal response, on the order
of 3°C, is presumably a function of evapotranspirative cooling due to either standing water
or elevated soil moisture levels. The drought conditions at the time of image acquisition
may have heightened this difference by further restricting soil moisture levels and
evapotranspirative cooling in the upland forests.

Results show that some of the major Pinelands land cover/vegetation types can be
discriminated and mapped with Landsat TM data with a reasonable degree of accuracy. The
comparison of the automated classification vs. a visual interpretation of the imagery at 489
randomly selected points, shows an overall agreement of 68.3% (Table 2). The percent
agreement varies with land cover/vegetation type. For example, the automated classification

PINELANDS REMOTE SENSING

10.00
0 OAK
© Sw HDWD
7 * PINE
a : 4 PINE LWLD
sp 8.00 O WCEDAR
a a
oO al
ere) a
” 7
i 6.00 4
. :
O 7
z 4.00 4
a
aD)
eS)
(x
i 2.00
AB
[@)
~
0.00

0.00 0,50 1.00 ry

50 2.00 2.50
Wavelength (micrometers)

FIG. 3. Plot of spectral radiance vs. TM wavelength bands for pinelands forest vegetation types.

8.00 5
: O B SOIL
7 0 PINE PLNS
oe : oO PINE
i
- 6.00 4
~ Hl
8) a
N 7
| 1
- a
O 4.00 5
= :
2 ,
ae el
"i
© ‘
ae 4
O 5
Oo Ai
Ce :
0.00 Ce eee EEE EE eeBEEnn Beeee
0.00 0.50 1.00 1.50 2.00 2.50

Wavelength (micrometers)

FIG. 4. Plot of spectral radiance vs. TM wavelength bands for transition from bare soil to pine plains to mature pitch
pine overstory.

6 BARTONIA

TABLE 2. Annotated classification vs. visual interpretation agreement matrix.

Reference Data

Row Number Producer’s User’s
Classified Data PO OP HS PL CS Totals Correct Accuracy Accuracy
pine-oak (PO) 95 18 1 22 1 146 95 68.8% 65.1%
oak-pine (OP) 29 78 14 9 0 133 78 69.6% 58.6%
hardwood swamp (HS) 0 5 7 0 0 12 w) 22.6% 58.3%
P pine lowland (PL) 12 9 9 118 4 157 118 78.1% 75.2%
cedar swamp (CS) 0 0 0 2 14 16 14 73.7% 88.0%
non-PB vegetation 2 1 0 0 0 26 22 57.9% 84.6%
Column Totals 138 112 31 151 19 489 334

Producer’s Accuracy: # correct/column total; User’s Accuracy: # correct/row total; Overall Classification Agreement
8.3%.

gave poor results for hardwood swamps which were readily confused with oak-pine forests.
This result is not unexpected as the automated classification is based solely on the spectral
information and there is a large spectral overlap between hardwood swamps and oak forests.
Visual interpretation utilizes spatial as well as spectral information, permitting more subtle
discrimination based on spatial context. As in the case of the hardwood swamps,
ae on patch shape and distance from streams allows more accurate discrimination.

Comparison between the automated classification of Landsat TM and visual interpretation
of aerial photographs shows a reasonable agreement (Table 3). The TM classification most
closely corresponds to the second two categories: >50% and >95% cedar. The aggregated
number of hectares for these two categories varied by +20-30% from the hectare results of
the TM classification. Some of the discrepancies are due to differences in the two methods.
The photo-interpreter drew smooth polygons around the cedar swamps often including small
inliers of different vegetation that the automated image classification would exclude. The
automated methods may misclassify a feature to be a cedar swamp based on spectral
information that a photo-interpreter can discriminate as a different land cover/vegetation type
based on spatial and textural information.

TABLE 3. Comparison of TM classification and photo-interpretation results of cedar swamps.

Percentage Area of TM Classified Cedar Swamps

Preservation Area Great Swamp
Photo-Interpretation
other 40.7% 17.3%
<50% 5.4 5.9
<95% sre cedar 20.5 6.8
>95% cedar 33.4 70.0

Area of Cedar Swamp in Hectares

Preservation Area Great Swamp
Photo-Interpretation
>95% cedar 1278 241
>50% cedar 2245 272

TM Classification
cedar swamp 2670 210

PINELANDS REMOTE SENSING 7

SUMMARY AND CONCLUSIONS

As demonstrated in other studies of the application of remotely sensing for forest
vegetation classification and mapping, one channel from the visible, near IR, and middle IR
is needed to adequately characterize the vegetation (Benson and DeGloria 1985, Horler and
Ahern 1986, Jensen et al. 1986, Lathrop et al. 1987, Spanner et al. 1984). To further
discriminate upland from lowland forest types, the thermal IR band should also be included.
The visible wavelength region (e.g. the red band) provides information on the pres-
ence/absence of vegetation. The near IR and middle IR provide information to discriminate
community types (e.g. oak vs. pine in the near IR, or pine uplands vs. lowlands in the middle
IR). The thermal infrared provides critical information on surface temperature and the
hydrological regime helping to differentiate upland from lowland forest types which appear
cooler. Due to the often sparse pine overstory, the understory vegetation may greatly
influence the remote sensing observed signal; the extent of this influence has yet to be
determined.

Analysis of the original and classified Landsat TM imagery further confirms ground-based
studies (Forman 1979) that the Pinelands landscape is characterized by a complex mosaic
of ecological land types: discrete patches or more often corridors of white cedar and
hardwood swamps amidst a background matrix of upland pine and oak forests. The upland
forests are better characterized by a continuous gradient with pure pine stands on one end
grading into pure oak at the opposite. These upland forests in turn grade into pitch pine
lowland forests depending on the hydrological regime. The effects of fire disturbance
provide an additional dimension overlaid on the above patterns.

Traditional supervised image classification techniques provide reasonably accurate results
but can be improved by the incorporation of additional data sets (e.g. soil type, elevation,
hydrologic regime) in the context of a geographic information system ). The
incorporation of spatial information (e.g. distance from streams) into classification procedures
should provide further discrimination among land cover/vegetation types. As part of a longer
term investigation of the landscape ecology of the Pinelands system, the Cook College
Remote Sensing Center is developing a GIS data base for the Pinelands National Reserve.
This GIS data base includes an approximately twenty year archive of Landsat Multispectral
Scanner (starting in 1972) and Thematic Mapper (starting in 1982) images. Remote sensing,
both satellite and airborne, in concert with GIS technology should provide a more effective
means of monitoring the spatial and temporal dynamics of the Pinelands ecosystem.

ACKNOWLEDGMENTS

I wish to acknowledge John Bognar, Sandra Luque, Jim Gasprich and Merlyn Liberty for
their efforts in all phases of this study. I thank the NJ Division of Parks and Forestry and
the Department of Environmental Protection, specifically Craig Coutros, for making the
state’s white cedar coverage available. This research was supported by the NJ Agricultural
Experiment Station with funding through the MclIntire-Stennis program.

REFERENCES

BENSON, A. S. AND S. D. DEGLORIA. 1985. nig oso of Landsat-4 Thematic Mapper and Multispectral Scanner

Data for Forest Surveys. Photogram. Eng. Remote Sensing 51:1281-1289.
FORMAN, R. T. T. 1979. The Pine Barrens of < Jersey: An Ecological Mosaic. Pages 569-586 in R. T. T.
rk.

Fort, ed. Pine Barrens: Ecosystem and Landscape. Academic Press, New Yo

BARTONIA

TM Band 4 - NIR DN

1
0 TM Band 3 - RED DN 65

Fic. 5A. Plot of TM classification training set ellipses for TM Band 4 (near IR) vs. Band 3 (red).

TM Band 4 - NIR DN

L
0 125
TM Band 5 - MIR DN

FIG. 5B. Plot of TM classification training set ellipses for TM Band 4 (near IR) vs. Band 5 (mid IR).

PINELANDS REMOTE SENSING 9

1225

TM Band 4 - NIR DN

fal

qT
134 159
TM Band 6 - TIR DN

Fic. 5c. Plot of TM classification training set ellipses for TM Band 4 (near IR) vs. Band 6 (thermal IR).

HORLER, D.N.H. AND AHERN. 1986. Forestry Information Content of Thematic Mapper Data. Int. J. Remote Sensing
7:405-428

JENSEN, J. R., M. E. HODGSON, E. CHRISTENSEN, H. E. MACKEY, JR., L. R. TINNEY, AND R. SHARITZ. 1986. Remote
Sensing Inland Wetlands: A Multispectral Approach. Photogram Eng Remote Sensing 52:87-100.

LATHROP, R. G., T. M. LILLESAND AND B. S. YANDELL. 1987. An Evaluation of Thematic Mapper Data for Forest

Cover Mapping in Northern Wisconsin. Pages 386-393 in Proc. 11th Pecora Sym., Sioux Falls, S

LILLESAND, T. M. AND R. W. KIEFER. 1987. Remote Sensing and Image PARES Wiley, New York.

McCormick, J. 1979. The Vegetation of the New Jersey Pine Barrens. Pages 229-244 in R. T. T. Forman, ed.
Pine Barrens: Ecosystem and Landscape. Academic Press, New York.

PINELANDS COMMISSION. 1980. New Jersey Pinelands Draft Comprehensive Management Plan, Appendix 3.

PETERSON, D. L. AND S. W. RUNNING. 1989. Applications in Forest Science and Management. Pages 429-473 in
G. Asrar, ed. Preys and Applications of Optical Remote Sensing. Wiley, New York.

ROMAN, C. T., R. E. GOOD, AND S. LITTLE. 1987. Atlantic White Cedar Swamps of the New Jersey Pinelands.
Pages 35-39 i % i Laderman, 7 gee Mi Cedar Wetlands. Westview, Boulde

SPANNER, M. A. BRASS, AND 1984. Feature Selection and the <n Content of
Thematic ie Simulator Data eo ee Assessment. IEEE Trans. Geosci. Remote Sensing, Vol.
22:482-489,

_? :
”
: heals
= By ai f |
: iam CA he aan ea ceay-| p62 actin
; | |
| ae

aes = aa) , :

at

7

ee

oe

Bartonia No. 58: 11-21, 1994

Mosses of the New Jersey Pine Barrens and
Adjacent Coastal Plain

ERIC F. KARLIN
Institute for Environmental Studies, Ramapo College of New Jersey, Mahwah, NJ 07430-1680

This paper presents the first comprehensive published list of the mosses occurring in the
New Jersey Pine Barrens, a region which has long been recognized for its unique and highly
significant ecosystems (Good and Good 1984). Mosses are a notable component of the Pine
Barrens vegetation, especially in the wetlands which occupy 25-35% of the region (Ehrenfeld
1986; Roman and Good 1990). Reviews of the ecological and taxonomical literature on the
mosses of the New Jersey Pine Barrens have been made by Forman (1979) and Karlin and
Andrus (1988). Although not focused on the Pine Barrens or New Jersey, Crum and
Anderson’s (1981) treatment of eastern North American mosses is an invaluable source of
information. Recent studies have focused on the development of a checklist of New Jersey
mosses and the determination of the rare and endangered species of mosses in the state
(Andrus and Karlin 1989; Karlin 1990; Karlin and Schaffroth 1992). Listings of endangered
moss species already exist for New York (Clemants and Ketchledge 1990), Ohio (Snider
1990) and North Carolina, and one is currently being developed for Pennsylvania (Webster
and Manville 1990).

The most prolific collector in southern New Jersey in the 1800’s was Coe Finch Austin,
who collected both in the Pine Barrens and also in the Palisades of Bergen County. His
collections served as the primary source upon which the first listing of New Jersey mosses
was formulated (Britton 1881 & 1889). Other collectors between 1850 and 1900 include E.
G. and N. L. Britton, G. N. Best, D. C. Eaton, T. P. James and E. A. Rau. The next period
of notable activity was during the 1930’s, when Bayard Long collected extensively
throughout the Pine Barrens region. A number of collections were also made during this
time by A. T. Beals, H. Koster and G. M. Tees. Edwin T. Moul collected in the late 1940s
and 1950s, and also identified specimens collected by others. Marian Robertson collected
extensively in the 1960s; unfortunately many of her specimens were not identified and her
collections have yet to be curated. Other active collectors from 1960 until the present include
R. E. Andrus, T. Gordon, E. F. Karlin, A. List, W. S. G. Maass, C. F. Reed, W. Olson and
G. L. Smith.

METHODS

All available moss specimens collected from the coastal plain region of New Jersey
(southern New Jersey) at the Brooklyn Botanic Garden (BKL), Chrysler Herbarium (CHRB),
New York Botanical Garden (NY), Academy of Natural Sciences (PH) and State University
of New York at Binghamton (BING) were examined. Collections from the Piedmont
portions of Mercer and Middlesex Counties were excluded. Using the Pine Barrens
boundaries given by McCormick and Forman (1979), all taxa having been collected from the
Pine Barrens region were noted. Nomenclature follows Anderson (1990) for Sphagnum and
Anderson et al. (1990) for all other mosses.

12 BARTONIA

RESULTS AND DISCUSSION

A total of 168 species and 3 varieties of mosses (35 families and 71 genera) have been
collected in the New Jersey Pine Barrens; another 36 species occur on the adjacent coastal
plain of New Jersey (Table 1). Six of these taxa have only recently been reported for New
Jersey (Bryum uliginosum, Fontinalis hypnoides, Fontinalis flaccida, Leskella nervosa,
Rhizomnium magnifolium and Tetraplodon angustatus) (Karlin and Schaffroth 1992). This
makes a total of 204 species and 3 varieties of mosses (in 37 families and 84 genera) known
from southern New Jersey, which is significantly lower than the number of vascular plants:
about 850 species reported for the Pine Barrens (Fairbrothers 1979) and approximately 1400
species for all of southern New Jersey (Stone 1911).

The most recent checklist of mosses for North America (Anderson et al. 1990) includes
a number of significant taxonomic changes affecting New Jersey taxa. Following Koponen
(1968), the genus Mnium has been divided up into several genera, with Mnium, Rhizomnium
and Plagiomnium being present in southern New Jersey (Table 1). Drepanocladus was also
split up into several genera, following Tuomikoski (1973), with Warnstorfia being the only
representative of the complex found in southern New Jersey to date (Table 1). At the
species level, all southern New Jersey material of Plagiomnium affine is now referable to
either P. ciliare or P. ellipticum, and all southern New Jersey material of Atrichum
undulatum is now referable to A. altecristatum and perhaps A. oerstedianum. In addition,
Sphagnum viridum was split from S. cuspidatum (Flatberg 1988) and S. isoviitae was split
from S. fallax (Flatberg 1992). All four of these Sphagnum species occur in the Pine
Barrens.

Unlike vascular plants, there are no moss species which are endemic to the Pine Barrens.
Most of the species are widespread in eastern North America; only 22 taxa (about 11% of
the southern New Jersey moss flora) have strong coastal plain affinities (Crum and Anderson
1981; Karlin and Andrus 1988). Several taxa reach the northern (5 taxa) or southern (24
taxa) limits of their geographical coastal plain limits in the Pine Barrens and adjacent coastal
plain of New Jersey (Table 1). Interestingly, the ratio of taxa at their northern geographical
limits to taxa at their southern geographical limits in this region is quite different for vascular
plants (about 7:1) (Fairbrothers 1979) and mosses (about 1:5). This indicates that the
biology and ecology of these two groups of plants may be quite different.

Sphagnum are the most ecologically and economically significant mosses in the Pine
Barrens, and they also represent the most distinctive aspect of the Pine Barrens moss
vegetation. Representing 20% of the total number of moss taxa found in the Pine Barrens
(16% of the total in southern New Jersey), Sphagnum is the second most species rich plant
genus in this region. Only Carex has a greater diversity. The Sphagnum floras of the
coastal plain of New Jersey (33 species and 1 variety) and Long Island (33 species) are the
most diverse found on the North American coastal plain (Andrus 1980; Lane and Dubois
1981; Karlin and Andrus 1988; Karlin, Andrus and Reed 1991). Such a high diversity is
partly explained by the many species which reach their geographical limits in the Pine
Barrens. Several taxa are at their northern (3) or southern (8) geographical coastal plain
limits in the Pine Barrens and adjacent coastal plain of New Jersey (Table 1), with three
other species being very near their southern limits (Karlin and Andrus 1988; Karlin, Andrus
and Reed 1991). Over half of the Sphagnum taxa present in this region (16 taxa) have
strong coastal plain affinities (Karlin and Andrus 1988).

MOSSES OF NEW JERSEY PINE BARRENS 13

TABLE 1. The moss taxa presently known to occur in the New Jersey Pine Barrens and the adjacent coastal plain.
Taxa occurring on the adjacent coastal plain but not yet collected in the Pinelands are indicated with an asterisk.
The date of the most recent collection in each county (or region) is indicated as follows: A = 1975-1990; B =
1950-1974; C = 1900-1949; D = prior to 1900; Z = collection — Upper case letters indicate two
or more collections from that county (or region) and lower case letters stand for only one collection. Taxa at their
geographical coastal plain limits in southern New Jersey are nde NL (northern limit) or SL (southern limit).
The counties and regions are: CM = Cape May; At = Atlantic; Cu = Cumberland; Sa = Salem; Gl = Gloucester; Ca
= Camden; Bu = Burlington; Oc = Ocean; Mo = Mon une Mi = Middlesex; Me = Mercer; PB = no legis
site given other than the Pine Barrens region (Pinelands); SNJ = no collection site given other than
Jersey. For Plagiomnium ciliare and P. ellipticum, a question mark indicates that Mnium affine had ns ed
from that county, but it was not determined whether the specimen was P. ciliare or P. ellipticum.

Counties
Species CM At Cu Sa Gl Ca Bu Oc Mo Mi Me PB SNJ
aere distant
m (Hedw.) C. Mull.* ere ee Ve dS ay noe hse eal Se re
Ambystegtim pa ease
serpens ) Schimp. v
“alan eae on & Herv.* Sore. Fone i es Se Bo trot Gh ae ean gee
m (Hedw.) Lindb.* Oe ee ao ee ee ee ee We ee
Prepon e into
Pen accom het ex Brid.) Brid. oe ee ee ee a oe eee ee ee ne
modon (An ontaceae)
attenuatus pa Hub. hoes “OR Ge Cn De a? eras
rostratus (Hedw.) Schimp. Bh CE ee Se Oe See ey
Atrichum (Polytrichaceae)
altecristatum (Ren. & Card.) Smyth &
Smyth = oe ee ® ee ee
angustatum sera "hal & Schimp. CEO eB ae ee Cea ee
crispum (James) S bi Gis ee eRe A a DB oe eas
Aulacomnium nn eee

pietegien 2 (Hedw.) Bruch & Schimp. OS ae a Be oe ee ee

paulstre (Hedw.) Schwaegr. BBB C&C eA eS ee Sh UG
Barbula (Pottiaceae)

co. ta Hedw. Se ee ee ee ae

unguiculata Hedw. 2 =e) ha Oe ee OB CBee aang
a (Bartramiaceae)

omiformis Hedw.* ee es ee ee ee

iteeclianbcuen (Brachytheciaceae)

campestre (C. Mull.) Schimp. SL Seg ed Lee re ae ere i | eee ee eee eee

oxycladon (Brid.) Jae Se Gy oS Os Sule OB ae ee ee

plumosum (Hedw.) Schimp. oe Se ee Ee eee

rutabulum (Hedw.) S 3 ere NRG ae ey SN iy es rg eee ee etre

salebrosum (Web. & Mohr.) Schimp ot Cait 1, A ee en Ae ee ee ee
Sele ouctsy (Sematophylace ae)

recurvans (M spat Fleisch. . eo ee OR hee oe a ce
Bruchic ( Chi ae)

flexuo . ex Schwaegr.) C. Mull. eet Se as VE ee ee eee ee ee
Bryhnia space tie ciaceae)

novae-angliae (Sull. & Lesq.) Grout* Soe ee ee ee ee eM oe See

ryoandersonia (Brachytheciaceae)
illecebra (Hedw.) Robins Ce HE OE Se eR ee SS eee

recurvirostre (Hedw.) Chen gt igh Nigh an oe cae oe 2

14

TABLE 1. Continued

BARTONIA

Species

Counties

CM At Cu

Sa

Ca Bu Oc

Mo

Mi

Me PB SNJ

sateen Wenning
crophyllum w.) Wat. & Iwats.
virginianum a Wat. & Iwats.
Bryum (Bry )

IW
lisae De Not. var. capitan (Bruch &
wean. oa
eudotrique ae .) Gaertn. et al.*
pride ais Bruch & Schimp. SL
Buxbaumia (Buxbaumiaceae)
hylla Hedw
Callicladium aero
haldanianum (Grev.) Crum.

chrysophyllum A sea J. Lange
hispidulum (Brid.) M
radicale (P. Beauv.) fed
stellatum (Hedw.) C. Jens. SL
Ceratodon oe
re )

dene (Hedw.) Web. & Mohr SL
dbergii (Ren. & Card.) Grout
Cratoneuron Sereg conings

glomerata Bruch & Schimp. NL
Desmatodon (Pottiaceae)
obtusifolius i Schimp.
Dicranella (Dicranaceae)
cerviculata (Hi ‘die Schimp. SL
heteromalla (Hedw.) Schimp.
rufescens (With.) Schimp.*
varia (Hedw.) Schimp.
Dicranum (Dicranaceae)
condensatum Hedw.
flagellare Hedw.
Sus m

fallax (Hedw.) Zand.
Diphyscium ae reigy
foliosum (Hedw.) M

im 2 SoS

MOSSES OF NEW JERSEY PINE BARRENS

TABLE 1. Continued

Species

Counties

CM

At

Cu

Ca Bu Oc

Mo Mi

Me

PB SNJ

Ditrichum (Ditrichaceae)
Lindb.

pallidum (Hedw.) Hampe
pusillum (Hedw.) Hampe*
Drummondia (Orthotrichaceae)
prorepens (Hed = E.G. Britt.
Entodon — ae)
cladorrhizans poke C. Mull.
compressus (Hedw.) C. Mull. NL

Eurhynchium ewes gs
pulchellum (Hedw.) Jen
Fissidens (Fissidentaceae)
bryoides Hedw.*

dubius P
fontanus B. ein Steud.*
osmundioides Hedw.
Fontinalis (Fontinalaceae)
laccida Ren. & Card
hypnoides Hartm. SL
oe Sull. var. novae-angliae
ar. cymbifolia (Aust.) Welch
Pca Lindb.

Forsstroemia (Le ptodonaceae)
b

striatella (Brid.) Iwats.

turfacea (Lindb.) Iwats.
Homalotheciella eee neal

subcapillata (Hedw.) B

num (Hypnaceae

cupressiforme Hedw.

curvifolium Hedw.*

imponens Hedw.

lindbergii Mitt.

sana (Hedw.) P. Beauv.

pratense (Rabenh.) W. Koch

Leptobryum (Bryaceae)
pyriforme (Hedw.) Wils.

d BY @
B

ee

FF. = +t
'wonwwn

16

TABLE 1. Continued

BARTONIA

Species

Counties

At

Cu

Sa

Gl

Ca

Bu Oc

Mo

Mi

Me PB SNJ

Leptodictyum (Amblystegiaceae)
riparium (Hedw.) Warnst.*
humile (P. Beauv.) Ochyra
Leskea (Leskeaceae)
gracilescens Hedw.*

9
a
8
8
en)
o
*

julaceus (Hedw.) Sull.

marginatum (With.) Brid.
stellare Hedw.*
Orthotrichium (Orthotrichaceae)
hioense Sull. & Lesq.

sordidum Sull. & Lesq. SL
speciosum Nees var. shen (Schwaegr.)
L

stellatum Brid.
Phascum (Pottiaceae)
cuspidatum es
Physcomitrium (Funariaceae)
immersum Sull.
pyriforme (Hedw.) Hampe*
Plagiomnium (Mniaceae)
ciliare (C. gone T. Kop.
cuspidatum (Hedw.) T. Kop.
peda "(Brid) T. Kop, SL
Plagiothecium (Plagiotheciaceae)
cavifolium (Brid ts.

ae)
palustre (Bruch & ‘Sching’) Bruch &
Schimp.
ravenelii A
subulatum Phi Rabh.
Pleurozium (Hylocomiaceae)
schreberi (Brid.) Mitt.

aw
1

wm >
= 2

MOSSES OF NEW JERSEY PINE BARRENS

TABLE 1. Continued

Species

Counties

At

Cu

Ca Bu. Oc

Mo

Mi

Me PB SNJ

Pogonatum Ny ciario oe ae)
brachyphyllum (Michx.) P. Beauv.
nan (Hedw.) P. Beauv.
ts (Bryaceae
annotina *(Hedw) Lindb.
L

wahlenbergii (Web. & Mohr) Andrews*
Polytrichum ischemia
commune Hedw. v omm
var. ar. perigoniale Oleic) Hampe

Pseudo area
legans (Brid.) Iw
Pychomirium choix)
m (Schwaegr.) Spruce*
Pratap (Hynaceae
tenuirostris (Bruch & Schimp.) Buck*
ae)

selwynii (Kind) Crum et al.
Rhizomnium (M ae)
magnifolium Gick) T. Kop.* SL
punctatum (Hedw.) T. Kop
Rhytidiadelphus (Hiylnowmniacens)
triquetrus (He ain Bh Warnst.
he ta (Grimmiaceae)
apocarpum ffiede:) Bruch. & Schimp.
Sphagnum (Sphagnaceae)
affine Ren. & Card
angermanicum Melin SL
austinii Sull. ex Aust. SL
bartlettianum Warnst.
carolinianum Andrus NL
compactum
spidatum Hoffm.'
cyclophyllum Sull. & ia
fallax =A Klinggr.
fimbriatu
panne pea arnst.
fuscum fing King SL
henrye
pry a
macrophyllum Brid. var. macrophyllum
Aust. NL

aw

Cy Ou u

Pr-Wwon:

-» s

>a

>a > ocd >d' >>>dD:

oS Cy

a ee ae

>' app:

i oe |
B

BDmwwo p>

>>

(> PrPrrrrrr:
>rPrirrrirrrrrr: Fore

>
rr Pr: ap»

Cc
Cc

18

TABLE 1. Continued

BARTONIA

Species

Counties

=

Cu

Ca Bu Oc

Mo Mi

Me PB SNJ

m Lindb.
perichaetiale Hampe NL
latyp aoa —— Warnst. SL
portoric ampe
puchram "ih Warnst. SL
pylaesi
recurvum ae Beauv.
a aoe Pony SL
stric 5
can sol Brid. SL

Sphagnum (continued)
pillosu

ms

e C. Mu
Pie (Splachnaceae)
edw

m (Bri ; ae
)

Ss e
Taxiphyllum (Hypnaceae)
deplanatum (Bruch - eras Fleisch.
taxirameum (Mitt.) Fleisch.
isn (etapa)
p Hed

oe ee (Siiltickanaceac’
angustatus (Hewd.) Bruch & Schimp. SL

Thelia sae ae)

asprella §

hirtella sw) Sull.

lescurii
Thuidium ps eee

delicatulum (Hedw.) Schimp.

crispa (Hedw.) Brid.
acti ii Fe Samp
tus (S ue Loeske SL
sa “* edw.) Loe SL
Weissia (Pottiaceae)
controversa Hedw.

lool

rPrpreaoaors prepa

ory |

a6

rrrrrrrrrr: Py

eu
a &

>rPrmrr: >

>rPrPr:

‘Includes Sphagnum viridum Flatb.
"Includes Sphagnum isoviitae Flatb.

MOSSES OF NEW JERSEY PINE BARRENS 19

Three (S. austinii, S. macrophyllum var. floridanum, S. strictum) of the nine Sphagnum
species that are critically imperiled (having an S1 status) or nee (SH) in the state are
limited, in New Jersey, to the coastal plain region (Andrus and Karlin 1989, Office of
Natural Lands Management 1992; Karlin and Shaffroth 1992), Sphagnum austinii, for which
the type collection was made by C.F. Austin in the Pine Barrens, has not been collected in
New Jersey in over 100 years (Table 1), and attempts to relocate it have not been successful.
As habitat suitable for its existence still occurs in the Pine Barrens, there is a possibility that
S. austinii still persists there. Sphagnum platyphyllum, ae Pipeaas imperiled in New
Jersey, was recently found in the Pine Barrens for the firs

In contrast to Sphagnum, the dung mosses pb ene are among the rarest mosses in
the Pine Barrens. These unique mosses are usually found growing on dung or animal
remains, with some species having their spores dispersed by insects (Crum and Anderson
1981). Three species of dung mosses have been collected in the Pine Barrens: Splachnum
pensylvanicum, S. ampullaceum, and Tetraplodon angustatus. All three have apparently not
been found in the Pine Barrens in over 100 years (Table 1). The two genera have different
habitat requirements. Splachnum is associated with the dung of herbivores and Tetraplodon
is associated with the dung of carnivores.

A large percentage of the mosses of southern New Jersey are currently only known from
historical collections. For instance, 79 taxa (38% of the total) were last collected prior to
1950, with 34 of these having not been collected since the late 1800s (Table 1). And,
although the more common and easily noted species have been collected from a number of
locations, 48 taxa (23% of the total) are only known from one site each (Table 1). Aside
from Sphagnum (Karlin and Andrus 1988; Andrus and Karlin 1989), there is a significant
lack of current data on the occurrence, distribution and ecology of most mosses in the Pine
Barrens.

ACKNOWLEDGEMENTS

I am grateful for the assistance provided by Dick Andrus (BING), William Buck (NY),
Janice Meyer (CHRB), Steve Clemants (BKL), Kerry Barringer (BKL), Christine Manville
(PH) and Erie Schuyler (PH). I also greatly appreciate Kim Schaffroth’s help with the
updating of the taxonomy and the recording of specimens at NY. Bruce Allen (Missouri
Botanical Garden) verified some of the Fontanis collections.

LITERATURE CITED

ALLEN, B. 1990. Delaware mosses. Evansia 7: 1-8.
ANDERSON, L. E. 1990. A checklist of Sphagnum in North America north of Mexico. Bryologist 93(4): 550-501.
, H. A. CRUM, AND W. R. Buck. 1990. List of the mosses of North America north of Mexico. The

Bryologist 93: 448-499.

ANDRUS, pe E. 1980. Sphagnaceae of New York State. New York State Mus. Bull. 442. 89 pp.

E. F. KARLIN. 1989. A preliminary report on New Jersey’s rare and endangered species of Sphagnum:
A a ae at codengered and threatened bryophytes in New Jersey. Pages 23-57 in E.F. a ed., New
Jersey’s rare gr endangered plants and animals. Inst. Env. Stud., Ramapo College, Mahwah

BRITTON, N. L. 1881. A preliminary catalogue of the flora of New Jersey. Annual Rep. a Geol., New
Brunswick.

. 1889. Catalogue of plants found in New Jersey. N.J. Geol. Surv., Final Rep. State Geol., No. 2

CLEMANTS, S. AND E. KETCHLEDGE. 1990. Flora paerg the question of endangered mosses in New York State.
Pages 211-216 in R. S. Mitchell, C.J. Sheviak, an J. Leopold eds., Ecosystem arg ae rare species and
neongrr habitats. Proc. 15th Ann. Natural Mee pe coat New York State Mus. Bull.
RUM, H. A. AND L. E. ANDERSON. 1981. Mosses of eastern North America. 2 Vols. Columbia tae Press, New

20 BARTONIA

York.
EHRENFELD, J. G. 1986. Wetlands of the New Jersey Pine Barrens [U.S.A.]: The role of species composition in
oe function. Amer. Midl. Nat. 115: 301-313.
AIRBR Endangered, threatened and rare vascular plants of the Pine Barrens and their
Roecaeete Pages 395-405 in R. T. T.Forman ed., Pine Barrens: ecosystem and landscape. Academic Press,

FLATBERG, K. I. 1988. Sphagnum viridum sp. nov., and its relation to S. cuspidatum. K. Norske Vidensk. Selks.
Skr. 1: 5-63.

— . 1992. The European taxa in the Sphagnum recurvum complex. 1. Sphagnum isoviitae sp. nov. Journal of
Bivens Lil

FORMAN, R. T. T. 1979. Common bryophytes and lichens of the New Jersey Pine Barrens. Pages 407-424. in
R York.

m
Goop, R. E. AND N. F. Goon. . The Pinelands National Reserve: an ecosystem approach to management.
Bioscience 34(3): 169- 173.
. 1990. Endangered bryophytes in New Jersey: aferagenan- protection and management. Pages
208-210 in R. S.Mitchell, C.J. Sheviak, and D.J. Leopold eds., Ecosystem management: eo species and
significant habitats. Proc. 15th Ann. Natural Areas Conference. Sate ork State Mus. Bull. 4
R. E. ANDRUS. 1988. The Sphagnum species of New Jersey. Bull. Torrey Bot. ia: ine 168-195.
—— , ——— and C. F. REED. 1991. The Sphagnum flora of Delaware. Bull. oh Bot. Club 118: 43-51.
— A. SCHAFFROTH. 1992. The mosses of New Jersey. Evansia 9: 11-32.
een = H. 1980. Revised checklist of the mosses of New York State. hie York State Mus. Bull. No.
440

KOPONEN, T. 1968. Generic revision of Mniaceae Mitt. (Bryophyta). Ann. Bot. Fenn. 5: 117-151.
D. M.

LANE, AND S. DuBois. 1981. Water analysis of habitats of Sphagnum in the Coastal Plain of North and
South Lm ss Bryologist 84: 408-413.
McCoRM . AND R. T. T. FORMAN. 1979. Introduction: Location and boundaries of the New Jersey Pine

ei gee XXXV-xiii in R. T. T. Forman ed., Pine Barrens: ecosystem and landscape. Academic Press, New
York.

OFFICE OF NATURAL LANDS MANAGEMENT. 1992. Special Plants of New Jersey. New Jersey Dept. of Env.
Protection Energy, sag

ROMAN, C. T. AND R. E. Goop. 1990. A regional study for wetlands protection. Pages 1-5 in P. F. Whigham,
R.E. Good, me J. Kvet an Wetland ecology and management: case studies. Kluwer Academic Publishers,

London.

SNIDER, J. A. 1990. An endangered species list for ig pesa endangered and threatened species of Ohio. Pages
205-207 in R. S. Mitchell, C.J. Sheviak and D.J. Leopold eds., Ecosystem management: me species and
significant habitats. Proc. 15th Ann. Natural Areas oo New York State Mus. Bull. 4

STONE, W. 1911. The plants of southern New Jersey, with special reference to ee a of the oa Barrens and
the geographic distribution of the species. N.J. State Museum Annual Report

TUOMIKOSKI, R., T. KOPONEN AND T. AHTI. 1973. The mosses of the island of uae Ann. Bot. Fenn.
10: 217-264.

WEBSTER, H. J. AND C. MANVILLE. 1990. Rare species of Pennsylvania mosses: an assessment. Pages 202-204 in
R. S. Mitchell, C. J. Sheviak, and D. J. Leopold eds., Ecosystem management: rare species and significant
habitats. Proc. 15th Ann. Natural Areas Conference. New York State Mus. Bull. 471

APPENDIX
SELECTED SPECIMENS EXAMINED

nly those species which are represented by one collection or which have recently been reported as new state
records (Karlin and Schaffroth 1992) are oe
B IN

um: BURLINGT!
Zanoni 4874 (NY). Campylium stellatum: SALEM: N of Brotmanville, Long 999a (PH). Climacium dendiliek

MOSSES OF NEW JERSEY PINE BARRENS 21

OCEAN: in swamps about Closter eek a and Tom’s River, ae Musci Appal. 286 (NY). Cratoneuron
filicinum: MONMOUTH: banks of Manas , N.L. Britton 26 VII 1882 (CHRB). Cryphaea glomerata
naereonen ed Batsto, Austin VI poe a. gama poeta Closter and southern New Jersey, pres
Musci Appal. 122 (NY). Dicranella rufescens;: MONMOUTH: Swimming River above Red Bank, N.L. Britton 2
X 1886 (CHRB). Dicranum polysetum: CUMBERLAND: woods 0.25 mile NNW Menantico Creek, Robertson 3143
(PH). Didymodon fallax: BURLINGTON: Harrisville, 6.5 miles NW of New Gretna, Robertson 4812 (PH).
Entodon na sua s: OCEAN: Lakewood, near head of lake, Beals 15 VI 1942 (NY). Ephemerum crassinervum:
CAMDEN a Austin X 1868 (NY). Fissidens sane BURLINGTON: Pompeston Creek, N Broad Street,
Riverton si , Robertson 4807 (PH). Fissidens Seika : BURLINGTON: Riverton Park, between old lily pond
and Pompeston Creek, Robertson 4790 (PH). Fissidens cals CAMDEN: Atco, 1882 (NY). Fontinalis

flaccida: ATLANTIC: May’s rect eee 20 VI 1914 (NY); OCEAN: Tom’s River, Austin 1868 (NY).
Fontinalis hypnoides: as RLAND: Manumuskin River at Manumuskin, Forren 26 VI 1976 (PH). Funaria
flavicans. CUMBERLAND: S of Delmont, Long 1925 ye Homalotheciella subcapillata. ATLANTIC: Dorothy,

Long 2165 (PH). pours nervosa: CUMBERLAND: Bridgeton, along Racops Run, Long 1124 (PH);
GLOUCESTER: 0.5 mile S of Downer, Long 2184 (PH). eran ep austinii: MONMOUTH: Swan’s Pond,
Highlands, Haynes 7 X 1911 (NY). Micromitrium megalosprum: CAMDEN: broken damp ground about Closter
(Bergen Co.) and Camden, sas Musci Novae-Caesarenses 52 (NY). ace ambiguum: MIDDLESEX: Ireland
Brook, 2.5 miles NNW of Helmetta, Moul 6993 (CHRB). Mnium marginatum: OCEAN: Tom’s River, between
Lakewood and Lakehurst, ae 27 IX 1941 (NY). Mnium stellare:. BURLINGTON: W of Vincentown, Long
1572 (PH). Orthotricum speciosum var. elegans: BURLINGTON: Batsto, James VIII 1858 (NY). Phascum
cuspidatum: MERCER: near North Crosswicks, Camberlin ef a/. 16 II] 1912 (NY). Pleuridium palustre:
BURLINGTON: James 25 IV 1854 (NY). Pleuridium ravenelii: OCEAN: Farrago, Austin 14 VI 1869 (NY).
Pohlia annotina: BURLINGTON: Sim Place, Robertson 5736 (PH). Pohlia me ie Pines of New Jersey, C.F.
Austin VI 1863 (BKL). vilecngaie commune vat. igi ogee ON: 2 mi. ENE of Berlin, Mout 2452
(PH). Polytrichum oe OCEAN: Manchester, Eaton 26 VI 1862 (NY). Ptychomitrium incurvum:
MIDDLESEX: 1.5 mi NW of Mostanouth Junction, tec 2966 (PH). Pylaisiadelpha tenuirostris: MONMOUTH:
Prospertown, near beri Beals 14 V 1933 (NY). Rardin intricata. BURLINGTON: Brown’s Mills, Best 28
V 1892 Rhizomnium magnifolium. CUMBERLAND: Sarah Run, N of Marlboro, eyes 1040 (PH).
— triquetrus: MONMOUTH: Prins Small 14 IV 1894 (NY). Schistidium veal SALEM:

Mannington Hill, Portertown, Long 2080 (PH). Bde te ge MIDDL yess! ie Imetta pond, Karlin
me 0125 tee Splachnum pales INGTON: Quaker Bridge, T 1 PED. ——
taxirameum: CUMBERLAND: S of Leaning i, Long 607 (PH). Tetraplodon piconet INGTON
Quaker Bridge, i (NY). Thuidium recognitum: GLOUCESTER: Jennings, Swedesboro, Lippinicott 13 XI
1898 (PH). Ulota coarctata. GLOUCESTER: Newhdld, Ellis 29 X 1893 (NY).

7 - 7 : re fe Rg > ,

eee a pons ete ee ery ae lie

‘Oe ' ria nye Pe : . ;
i”

Bartonia No. 58: 23-28, 1994

Interactions Between Research and Management:
Examples from the New Jersey Pine Barrens

EMILY W. B. RUSSELL
Department of Geological Sciences, Rutgers University, Newark, NJ 07102

There is a special mystique about the odd and unique. Taxonomists who tire of
cataloguing the common flora are often drawn to what is rare or unusual. Many ecologists
also are attracted to unusual or unique ecosystems. An ecosystem is deemed of special value
because it has species at the edges of their ranges, disjuncts, endangered species, or other
unusual features, not usually because it contains the main part of the ranges of many fairly
common species. Farmers, on the other hand, are drawn to more mesic, ordinary conditions,
which will support the best plant growth. This serendipitous juxtaposition, of scientific
curiosity about rare species or ecosystems and agricultural propensity for mesic sites, has in
New Jersey left us the Pine Barrens. While heavy use for timber products has seriously
disturbed the forest, it has not had the sustained impact on the plant communities of plowing
and continued agriculture.

Research has flourished there, since taxonomists travelled out from Philadelphia to
botanize in the 18th century (Fairbrothers 1979). As the flora became well known and
published in 1911 (Stone 1911), ecological studies were well under way, and first published
in a comprehensive text in 1916 (Harshberger 1970). Research has continued on the many
aspects of the flora, ecosystems, geology, soils, and other features of the Pine Barrens since,
much of it presented and summarized in Forman’s compendium on the Pine Barrens in 1979
(Forman 1979). Just scanning that volume and the references in it will convince anyone that
the Pine Barrens have been a focus of exciting research for most of the 20th century.

But what of management? Farmers did not like the poor sandy soils, but others found
abundant resources to exploit in the Pine Barrens. After all, until recently, and even today
for many, land that has no clear economic use to people is waste land (Glacken 1967,
preface and passim). The Pine Barrens were not waste land; they supplied iron ore and the
trees with which to make charcoal to process the ore, glass sands, pitch and tar, and native
plant products such as cranberries, white cedar, Lycopodium, peat, and herbs. Management
is probably too strong a word to apply to the use of most of these materials; plunder would
be more appropriate. But pitch pines, oaks, ericaceous shrubs, and other typical species
survived this onslaught, producing the landscape we have today.

Threats to the integrity of this landscape, which would provide much more permanent
damage than the preceding centuries of use, inspired preservation efforts in the 1960s and
1970s, culminating in designation of the region as the nation’s first National Reserve in 1979
(Collins and Russell 1989, chapters 2 and 3). The activities leading to this designation
paralleled the growth of conservation movements in the nation at large. Our current
understanding of systematics and ecology indicated that the Pinelands had an intrinsic value
to the biosphere in terms of its unique nature, and led to designation of the Pinelands as an
International Biosphere Reserve (UNESCO 1984).'

"The name was changed from Pine Barrens to Pinelands during the political battles leading to creation of the
Pinelands National Reserve. 3

24 BARTONIA

The managers of the Pinelands National Reserve were then left with the problem of how
to maintain this value. It did not come to us unsullied by human activities, rather the
opposite. To manage it wisely, one needs to know which activities will sustain and which
damage the system as we think it should be: in some cases what it is today and in others
what we think that it was sometime in the past.

Decisions about research and management do not occur within a cultural vacuum.
Ecologists and land managers have, for example, changed their attitudes toward the role of
fire many times in the past. Fire management will be used here as an example to illustrate
the complex interactions between cultural values, research, and management.

Until the late 19th century, fire was viewed as both a tool in farm management and a
threat to forest vegetation. Laws from the early 18th century had limited the timing and
extent of fires used to clear land, in order to protect timber. Such fires still frequently
escaped into the surrounding forested land. By the end of the 19th century and into the 20th,
sparks from locomotives were added as a leading cause of forest fires, and because trains
penetrated very widely, fires were common everywhere. Fires that escaped from land-
clearing or were maliciously set in slash after logging were also major threats to forest
regeneration (Hough 1882, pp. 128-207; Meier 1903, 1904, 1905). Harshberger’s
observation in 1916 that "forest fires have done great damage to the wild plants [of the Pine
Barrens]" was typical of the attitudes toward fire at that time (Harshberger 1970, p. vi). By
the late 1930s in New Jersey such changes as improved fire-suppression and decreased
railroad usage contributed to a decrease in total area burned and number of fires (Forman and
Boerner 1981).

In the southeastern United States long leaf pine (Pinus palustris) grows in open groves
with a grassy understory. By 1924 in this region bobwhite quail, a very popular and income-
producing game bird, were becoming scarce (Landers and Mueller 1986). Research into the
decline in an important economic resource for the region suggested that a shift to larger
farms, cleaner cultivation with fewer hedgerows, less fire used to clear brush, and
overhunting had contributed to the decline (Stoddard 1936, Landers and Mueller 1986).
Prescribed burning was initiated to improve habitat for the quail, and led to extensive
research on the relationship between controlled fires and game management (Stoddard 1936).
Such research on the use of fire for game management continued into the 1970s, much of
it reported in the Proceedings of the Tall Timbers Fire Ecology Conferences, e.g., papers in
1970 on blue grouse and ruffed grouse and in 1972 on prairie chickens and sage grouse. In
England, as well, fire was used to manage game lands (Gimingham 1970). In other words,
an economic incentive led in the early days to the setting of controlled fires to manage
habitats of valued wildlife.

In 1936 a controlled burning regime was established in the Pinelands to study the effect
of prescribed burning (Little 1979). The goals of burning there were to reduce fuels, and
thus crown fires, and to improve the pine timber, that is, a concern with improving safety
and productivity. Again, the goals were related to economic return, and in this case also
human well-being and protecting investments in structures that might be destroyed by
wildfires.

However, by the 1960s, ecologists began to appreciate the value of fire for more general
ecosystem preservation, based on observations of ecosystems where fire had been suppressed.
For example, in 1969 Oberle published in Science that "forest fire suppression policy has its
ecological drawbacks" (Oberle 1969) and in 1973 Tiixen published in the Proceedings of
the 13th Tall Timbers Fire Ecology Conference on "the use of fire in nature conservation”

PINE BARRENS RESEARCH AND MANAGEMENT 25

(Titixen 1973). For the Pinelands, specifically, ecologists observed that "the importance of
fire to the maintenance of the Pinelands ecosystem is undeniable. If burning practices are
eliminated, the character of the Pinelands will change," understood as a change for the worse
ecologically (Bartlett and others 1978). This statement in a report done for the National Park

ervice signalled a new emphasis on fire as a tool for reconstructing the Pinelands as they
"should be." While the Comprehensive Management Plan for the Pinelands National Reserve
stated more conservatively that "... many ecologists believe that the existing Pinelands
landscape is in part the product of wildfires of varying intensities, including crown fires
which kill the overstory" (Pinelands Commission 1980, p 69), a report on research objectives
in the Pinelands two years later left no question about the beneficial effect of fire: "The
New Jersey Pine Barrens is clearly a fire dependent system in which periodic fires are not
only inevitable but are a natural feature of the landscape" (Good 1982, p. 11).

This new attitude, of using fire to mimic nature, must then be related to the overall goals
of management in the Pinelands National Reserve, and thus management and research have
needed to establish new relationships. The overall goal for natural resources in the
Pinelands, as stated in the Pinelands Comprehensive Management Plan, is to "Preserve,
protect and enhance the overall ecological value of the Pinelands, including its large forested
areas, its essential character, and its potential to recover from disturbance" (Pinelands
Commission 1980, p.193). Just to arrive at such a goal required research, based in part on
the present commitment of systematists and ecologists to the rare and unique ("essential
character") and conviction that disturbances such as fire were necessary for their preserva-
tion. But this goal is quite different from managing w acon —— en or Pages
timber and it requires techniques and research which different from the mo
economically-motivated purposes. It has been the noe of the ioeatione to cctira
convincingly to the managers the rationale and consequences of the new interpretations.
Further research may indicate changes in interpretations, that is, that certain kinds and
distributions of fires may have very different effects, for example. The importance of such
changes in interpretations must be incorporated by managers into their plans. In addition,
they need to indicate to the researchers what questions they have with relation to how to use
fire to reach their goals. While researchers cannot set the goals, they can provide
information that is relevant to setting as well as implementing them

Fire is one of many issues about which the interpretations have henintd over the years,
and thus research and management related to fire have changed as well. Both the research
and the applications of the research have been closely related to contemporary societal value
systems. This example illustrates the complexity of relating research and management, and
the continuing readjustments that must be made.

The papers presented in this symposium have clearly demonstrated exciting frontiers of
research in the Pinelands, and by extrapolation, in general, and all bear relations to managing
the resources of the Pinelands. Forman and Lathrop both present landscape-scale studies.
While decisions must be made at the local scale, these decisions must be set in the context
of the regional, or landscape, scale. The models presented by Forman provide a theoretical
way of testing the consequences of patterns of landscape change, which are generalized
across many kinds of landscapes (Franklin and Forman 1987). To relate these models to the
management decisions requires making them responsive to specific kinds of disturbances in
the Pinelands, and local patterns of disturbance, such as "nibbling at the edges." They
provide a framework in which to ask questions about many otherwise unexpected
consequences of management decisions before these decisions are implemented. The concept

26 BARTONIA

of sustainability can provide a framework in which to set the goals for management of the
resource.

Lathrop’s use of remote sensing provides a way of analyzing the elements of natural
landscapes at different scales and monitoring changes over time. The im
consistency in collecting data, so that they will be comparable over time, is critical. "For
example, the pixel size of the image determines the spatial scale of activities that can be
detected. Remote images can provide more details of spatial diversity than can be detected
visually in a photograph, so that, for example, cedar swamps which appear on photographs
to be continuous corridors, may appear as discrete patches in the remote images. Such detail
provides information that can be used in management decisions regarding preservation of
cedar swamps, depending on the importance of the continuity of the corridors. It is critical
that those interpreting, and planning, the remote images interact with those making
management decisions, so that the scales of the information provided and needed coincide.

Research on major components of the ecosystem, oaks (Quercus spp.) and mosses, as
reported by Fairbrothers and Karlin provides information on the scale of individual
organisms and species that leads to the broader patterns recognized by Forman and Lathrop.
These two taxa, which have suffered from unmerited neglect relative to pines (Pinus spp.,
especially P. rigida), are integral parts of the unique vegetation of the Pinelands. Because
the diversity of the oaks is due as much to intra- as to inter-population variation their
contribution to the diversity of the plant communities and the ability to recover from
disturbance is much greater than might appear from overall vegetation patterns. This
diversity is reflected also in a major pattern of recovery from disturbance by sprouting from
lignotubers. Both of these characteristics of oaks need to be integrated into management of
the Pinelands. It is interesting to note that in another pitch pine (P. rigida) community, the
Albany (New York) "Pine Bush," it is the scrub oak (Quercus ilicifolia) that is most
characteristic of the vegetation (Zaremba et al. 1990).

The contribution of the mosses to the "essential character" of the Pinelands is unique. The
lack of diversity of the group as a whole stands in contrast with the extremely high diversity
of the one genus, Sphagnum. The closer affinity with the northern than with the southern
moss flora, as contrasted with the more southerly vascular plant affinity, suggests that mosses
are responding to different elements of the ecosystem and/or landscape as a whole. To
approach the goal of maintaining the "essential character" of the Pinelands, the reasons for
this difference should be further explored. In addition, the reactions of Sphagnum in
responding to disturbance are clearly important. Both of these are compelling reasons for
management to pay more attention to updating the moss flora and ecology of the Pinelands.

Finally, we need to be reminded of the importance of the human element in the Pinelands,
both in the present and the past. The Pinelands is not just a natural region, but it also has
a rich cultural heritage. While many associate it with the "Pineys," the history of other
cultural and ethnic groups contribute to this heritage. Research on such individuals as James
Still adds immeasurably to our appreciation of the kinds of lives people lived in the past, and
ways in which outstanding individuals could rise above their humble beginnings.
Management of the Pinelands should include interpretation of communities and individuals
who contributed to the unique cultural heritage of the region.

The actions required to manage a landscape or anything else are never clearcut.- The
ecologists cannot say with certainty that if you burn a specific site once every 10 years with
a crown fire you will get the desired plant and animal communities. The actual decisions
are subjective. A quotation from Paul Sears, written in 1935 in Deserts on the March,

PINE BARRENS RESEARCH AND MANAGEMENT OF

makes this point more clearly:

Science has the power to illuminate, but not to solve, the deeper problems of mankind. For always after
knowledge come choice and action, both of them intensely personal and individual. Science is like those services
which supply a battle commander with information of the enemy and technical advice on physical problems, but
which cannot relieve him of the task of decision. For the last his own intuition, perhaps even the toss of a coin,
must serve. The inescapable character of this dilemma is recognized in the old military maxim that action on an
inferior decision is far better than no action at all. (Sears 1988, p. 219)

The manager trying to use the results of research is in the position of the military
commander described by Sears. He receives information and technical details from the
researchers, and has overall goals to achieve, but how he uses the information to reach the
goals has to be a subjective decision. Sears does not discuss here the importance of setting
the goals, but that is clearly a part of the decision-making process. Research presented at
this symposium can be incorporated into management strategies in the Pinelands National
Reserve, and I think that all of it relates to the objectives for which the Reserve was set up.
The specific applications and the encouragement of future research will depend on effective
communication between those making the decisions and those doing the research.

LITERATURE CITED
BARTLETT, M. ET AL. 1978. A plan for a Pinelands National Preserve. Center for Coastal and Environmental
Studies, New Brunswick.
COLLINS, B. R. AND E. W. B. RUSSELL, EDS. 1989. peestraa the New Jersey Pinelands. a new direction in land-
use management. Rutgers University Press, New Bruns
FAIRBROTHERS, D. E. 1979. Endangered, threatened, ry rare vascular plants of the Pine Barrens and their
eg tne a 395-405 in R. T. T. Forman, ed. Pine Barrens. ecosystem and landscape. Academic Press,

aa : i cl ED. 1979, Pine Barrens: ecosystem and landscape. New York: Academic Press, Inc.

FORMAN, R. T. T. AND R. E. BOERNER. 1981. Fire frequency and the Pine Barrens of New Jersey. Bull. Torrey Bot.
Club 108:34-50.

FRANKLIN, J. F. AND R. T. T. FORMAN. 1987. Creating landscape patterns by logging: ecological consequences
and sag ag Landscape Ecology 1:5-18.

GIMINGHAM, C. H. 1970. British heathland ecosystems: the outcome of many years of management by fire.
Proceedings - the 10th Tall Timbers Fire Conference:293-321.

GLACKEN, C. 1967. Traces on the Rhodian Shore. University of California Press, Berkeley

Goop, R. E. 1982. pepe a to environmental management concerns in the vieclanil National Reserve.
Center for Coastal and E nmental Studies, New Brunswick.

HARSHBERGER, J. W. 1970 ‘lly 1916). The vegetation of the New Jersey Pine-Barrens. Dover Publications,
Inc, New York.

HouGH, F. B. 1882. Report on forestry. Government Printing Office (USDA), Washington.

LANDERS, J. L. AND B. S. MUELLER. 1986. Bobwhite quail management: a habitat approach. Tall Timbers Research
Station Miscellaneous Publication #6.

LITTLE, S. 1979. Fire and plant succession in the New Jersey Pine Barrens. Pages 297-314 in R. T. T. Forman, ed.
Pine Barrens: ecosystem and landscape, Academic Press, Inc, New Yor!

MEIER, F.R. 1902. Forest fires in New Jersey during ne Pees Sisvey of New Jersey. Annual Rep. State
Geologist:96-107. The John L. Murphy Publ. Co.,

MEIER, F, R. 3. Forest fires in New Jersey in are ‘esl Survey of New Jersey. Annual Rep. State
Geologist:45-54. The John L. Murphy Publ. Co, Trento

MEIER, F. R . Forest fires in New Jersey in 1904. Geologie Survey of New Jersey. Annual Rep. State
Geologist: 275-289. The John L. Murphy Publ. Co., Trento

OBERLE, M. 1969. Forest fires: suppression policy has its Sabai drawbacks. Science 165: 568-571.

PINELANDS COMMISSION. 1980. Comprehensive management plan for the Pinelands National Reserve. Pinelands

bes

Commission, New Lisbon. ‘
SEARS, P. B. 1988 (reprint of 1935 ed.). Deserts on the march. Island Press, Washington.
STODDARD, H. L. 1936. The Bobwhite Quail: its habits, preservation and increase. Charles Scribner’s Sons, New

28 BARTONIA

York.

STONE, W. 1911. The plants of southern New Jersey, with especial reference to the flora of the Pine Barrens and
the geographical sig of the species. Annual Rep. New Jersey State Mus., 1910, Part I1:23-828.

TUXEN, R. 19 use of fire in nature conservation. Proceedings of the 11th Tall Timbers Fire Ecology
Conference. 99- 120.

UNESCO. 1984. Action plan for Biosphere sey Nature and Resources 20(1):1-1

ZAREMBA, R. E., D. HUNT AND A. LESTER. 1990. Albany Pine Bush fire pbc aninee plan. Rep. to the Albany
Pine Bush Commission, The Nature Conservancy, New York Field Office, Albany.

Bartonia No. 58: 29-68, 1994

Taxonomy of Scirpus, Trichophorum, and
Schoenoplectus (Cyperaceae) in Virginia

MARK T. STRONG
Department of Botany, Smithsonian Institution, NHB/166, Washington, DC 20560

In the last 30 years, embryological, anatomical, and cladistical studies in the subfamily
Cyperoideae have lent supporting evidence to the premise that Scirpus sensu lato is a
heterogeneous group. Van der Veken (1965) studied embryo morphology of species in the
subfamily Cyperoideae. He classified the mature embryos into six types. All six of these
types were found to be represented in Scirpus sensu lato. Van der Veken felt that several
genera could have the same embryo type, but no genus should have more than one embryo
type represented in it. The embryos of Virginia Scirpus sensu lato fall into three of these
types (Table 1). This indicates that Virginia Scirpus sensu lato should be represented at least
by three different genera.

Studying anatomical and morphological characteristics as well as habitat types of natural
groups occurring in Scirpus sensu lato, Schuyler (197la, 1971b, 1972b) developed a
preliminary basis for taxonomically delineating these groups. The results of his studies are
incorporated into Table 2 which, in addition, includes a summary of other significant
characters that these groups have in common or separate them from each other. Schoeno-
plectus can be distinguished from Scirpus sensu stricto and Trichophorum by differences in
embryo type, leaf anatomy, inflorescence type, size and epidermal structure of achenes, style
morphology, anther morphology, and habitat. Trichophorum can be distinguished from
Scirpus sensu stricto by differences in embryo type, leaf anatomy, inflorescence type,
epidermal structure of achenes, and habitat.

Cladistical analysis of the subfamily Cyperoideae by Goetghebeur (1986) indicates that
Schoenoplectus is separate from the tribe Scirpeae and belongs in the tribe Fuireneae, which
also includes the genera Actinoscirpus, Pseudoschoenus, and Fuirena. This conclusion is
based primarily on embryo-type and morphology of the perianth bristles. In his cladistical
analysis of the tribe Fuireneae, Goetghebeur (1986) recognizes the genus Bolboschoenus as
have other recent workers (e.g., Raynal 1973; Oteng-Yeboah 1974; Wilson 1981; and
Koyama 1985, 1990), however, he seems to dismiss the similarities in embryo shape, floral
morphology (anthers, styles, perianth bristles, and achenes), leaf anatomy, inflorescence, and
habitat between Schoenoplectus and Bolboschoenus. Goetghebeur & Simpson (1991)
separate both Bolboschoenus and the monotypic Asian genus Actinoscirpus from
Schoenoplectus primarily on the basis of well-developed leaf blades and involucral bracts,
and terminal inflorescence of Bolboschoenus and Actinoscirpus vs. reduced leaf blades and
involucral bracts, and pseudolateral inflorescence of Schoenoplectus. However, these
characters are not consistent within Schoenoplectus and Bolboschoenus. There are a number
of species in Schoenoplectus (e.g., Sch. etuberculatus, Sch. subterminalis, Sch. torreyi, and

29

30 BARTONIA

TABLE 1. Embryo types of Virginia Scirpus sensu lato and their classification. From Van der Veken (1965) and
Wilson (1981).

CAREX-TYPE FIMBRISTYLIS-TYPE SCHOENOPLECTUS-TYPE
rpus embryo of Van der
Veken n)

EMBRYO SHAPE top-shaped top-shaped mushroom-shaped
LEOPTYLE lateral basal basal

(final position)

RADICLE basal lateral lateral

(final position)

FIRST LEAF perpendicular perpendicular parallel

(orientation of

germination pore
relative to first leaf)

GENERA IN TRICHOPHORUM SCIRPUS SCHOENOPLECTUS
VIRGINIA

Sch, pungens) that have well developed leaves, although they are confined primarily to the
lower portion of the culm. Schuyler (1971a) found anatomical evidence that the well-devel-
oped leaf blades of tuberous bulrushes (Bolboschoenus) were homologous to those found in
one species of Schoenoplectus (Sch. americanus). The leaf blades of this species and those
of Bolboschoenus have chlorenchyma occurring in about equal abundance on both the
adaxial and abaxial sides of the leaf blade, as well as stomates occurring in equal abundance
in the adaxial and abaxial epidermis. In further anatomical studies done on Schoenoplectus,
Schuyler (1972b) found two other species (Sch. etuberculatus and Sch. subterminalis) that
have chlorenchyma on both the adaxial and abaxial sides of the leaf blade.

Although the inflorescences of Schoenoplectus (as circumscribed by Goetghebeur and
Simpson 1991) are primarily pseudolateral and subtended by an erect bract appearing as a
continuation of the culm, there are species in Schoenoplectus (particularly those that have
elongate inflorescence rays) that have 1-3 additional inflorescence bracts subtending the
inflorescence (e.g., Sch. validus, Sch. acutus, Sch. californicus, Sch. torreyi, and Sch.
etuberculatus). In Bolboschoenus, the lowest involucral bract is leaf-like above the
inflorescence, but the inflorescence typically appears pseudolateral at anthesis and in reduced
forms. Only at maturity do the elongate involucral bracts spread out, giving the inflores-
cence a terminal appearance. Furthermore, the Asian species Bolboschoenus planiculmis has
a pseudolateral inflorescence, the lowest involucral bract appearing as a continuation of the
culm as in Schoenoplectus, but the floral morphology is that of Bolboschoenus. Therefore,
because of the similarities between embryo shape, leaf anatomy, floral morphology, and
inflorescence between Schoenoplectus and Bolboschoenus, | cannot justify the separation of
these plants into distinct genera.

This study examines plants of the genus Scirpus sensu lato that occur in Virginia. Scirpus
sensu lato has been treated in North Carolina (Cappell 1954) and species mapped by county
in Virginia (Harvill et al. 1992), but no formal treatment has been made for Virginia. Notes
and studies on species of Scirpus sensu lato that occur in Virginia are scattered in the
literature and many species have been poorly defined and disregarded by workers in
Virginia. A clarification of the taxonomic position of species of Scirpus sensu lato is thus
needed for Virginia.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS oi

TABLE 2. Summary of the main characteristics of the three segregate genera of Scirpus sensu lato in Virginia.

TRICHOPHORUM
1. Carex embryo-type

2. Leaf blades well developed or
reduced (keel-shaped or crescent-
form-shaped in cross section),

at least some of the vascular =
dles

3. Inflorescence usually of a sin-
gle termnal spikelet, the involucral
bract scales

4. Achenes with non-wavy inner

cell walls, lacking buttresses and

pgp apical nodules which give
achene a minutely papillate

e

5. Styles subulate, the branches
with minute rounded protrusions
(seen under high magnification)

6. Anthers apiculate at apex
7. Grow in dry to hak a
and

primarily in upland fore
deciduous woods

SCIRPUS
Fimbristylis embryo-type

Leaf blades well- -developed (keel-
shaped in cross section
chlorenchyma cells more devel-
oped on adaxial side than on ab-
axial side; stomates primarily
restricted to abaxial epidermis

Infloresence generally

branched, terminal, su abe by
a leafy involucre; spikelets small,
numerous

Achenes with wavy inner cell
eae buttresses and apical nod-

which give the achene a mi-
aah papillate texture, mostly
less than 1.5 mm long

Styles subulate, the branches with
minute rounded protrusions (seen
under high magnification)

Anthers apiculate at apex

ony in seasonally wet substrates

as swamps, marshes, shallow
nn culm bases not submerged
or only partially so

SCHOENOPLECTUS
Schoenoplectus embryo-type

Leaves bladeless or with well-de-
veloped, elongate blades (cres-

present in both abaxial and adaxi-
al epidermis

Inflorescence compact or few-
ched, pseudolateral, with an
€ 0 r

pearing terminal at maturity

Achenes smooth, pitted or rugu-
lose, epidermal cell structure di-
verse, mostly more than 1.5 mm
long

Styles sub-flattened to strap-like,
the branches scaly or furrowed,
glabrous or with scattered hairs

Anthers with a Te or prickly
appendage at ape

Grow primarily in water along
large river systems, and in estuar-
ies, lakes and ponds; culm bases
usually submerged

This study endeavors to provide a concise, comprehensive treatment of the species of

Scirpus sensu lato that occur in Virginia. With the increasing concern over our dwindling
natural resources in Virginia, particularly wetland areas in which species of Scirpus sensu
lato play an important role in the ecology of these habitats, it is important that a treatment
be made available to workers doing field work in Virginia. The resulting treatment will
facilitate the identification of species as well as clarify taxonomically the classification of
these plants.

The treatment contains: (1) a key to the genera of Scirpus sensu lato in Virginia and keys
to species in each genera; (2) a SEM photomicrograph of the fruit (achene) of each species;
(3) a drawing of a representative species in each genus and selected species as an aid, along
with the dichotomous keys, to identifying closely related species, and (4) a detailed
description of each species which includes flowering and fruiting times, distribution, habitat,
distinguishing features, and in some species, clarification of nomenclatural changes or
differences between related species are discussed.

32 BARTONIA

METHODS

Studies of Virginia Scirpus sensu lato were based on herbarium material and my own
personal collections. Morphological characteristics of each species were examined. These
included habit, rootstock, leaves, culms, inflorescence, flowers, and fruit. A taxonomic
treatment was then prepared based on morphological characteristics as well as evidence from
embryological, anatomical, and cladistical studies from recent literature. Specimens were
examined from the following herbaria: Longwood College, Farmville, Virginia (FARM);
George Mason University, Fairfax, Virginia (GMUF); New York Botanical Garden, Bronx,
New York (NY); Academy of Natural Sciences, Philadelphia, Pennsylvania (PH); United
States National Herbarium, Washington, D.C. (US); Virginia Polytechnic Institute and State
University, Blacksburg, Virginia (VPI); and The College of William and Mary, Williams-
burg, Virginia (WILLI). Searches for chromosome counts recorded in the literature were
made for each species. These are summarized in Table 3. Specimens cited are available
upon request. All specimens collected by the author are deposited at GMUF. This study
is based on the author’s MLS. thesis.

MORPHOLOGICAL COMPARISON OF THE THREE GENERA

RHIZOMES. The rhizomes of Scirpus are typically short, hardened and knotty, and
frequently bear two or more culms. In Trichophorum they are small, short and knotty, and
bear dense tufts of leaves and culms, or in some species, they are elongate and slender and
bear culms singly along the rhizome. In Schoenoplectus, they are elongate and bear culms
singly along the rhizome. Culms may either arise spaced at intervals along the rhizome as
in Sch. pungens and Sch. americanus or in a row as in Sch. validus and Sch. acutus. A few
closely related species of Schoenoplectus (e.g., Sch. robustus, Sch. fluviatilis, and Sch.
novae-angliae) have hard, tuberous enlargements along the rhizomes below culm bases.
These may function either as both supporting structures in the flooded habitat they typically
grow in or as tubers for vegetative reproduction.

CULMS. The culms of Scirpus are trigonous or obtusely trigonous, stout, and are rarely
less than 5 dm tall. Mature culms of Scirpus divaricatus frequently proliferate from the leaf
axils, the reclining culms eventually touching the ground and rooting. The culms of
Trichophorum are trigonous or terete, slender, and typically less than 4 dm tall. In
Schoenoplectus they are terete or trigonous, stout, and usually more than 5 dm tall. The
culms of many species of Schoenoplectus (e.g., Sch. validus, Sch. acutus, Sch. americanus,
and Sch. pungens) develop air cavities, presumably as an adaptation to aquatic environments.

LEAVES. Leaves of Scirpus are both basal and cauline. The sheath is usually
short-ligulate at the adaxial junction of sheath and blade and has a membranous inner band
with a U-shaped or truncate to convex orifice which may split as the plant increases in
circumference. The blades are typically flattened and have numerous veins with the midvein
being most prominent. The margins and the midvein beneath are usually scabrous.
Anatomically, the flattened, keeled, V-shaped blades of Scirpus (as seen in cross section)
have chlorenchyma more developed on the adaxial side of the blade than on the abaxial side.
Stomates are primarily restricted to the abaxial epidermis (Schuyler 1971a). The leaves of
Trichophorum are typically basal, bladeless, or the uppermost blade-bearing. Sheaths bearing
blades are short-ligulate adaxially at junction of sheath and blade and have a scarious inner
band. Blades are typically subflattened to flattened, narrow, usually less than 2 mm wide,

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 33

TABLE 3. Haploid chromosome numbers of Virginia Scirpus sensu lato.

TAXON a unite REFERENCE

Scirpus divaricatus 14 Schuyler 1963, 1967a

S. polyphyllus 29 Schuyler 1967a, 1975b

S. expansus 32 Schuyler 1967a

S. ancistrochaetus 27 Schuyler 1963, 1967a

S. georgianus 24: 25, 26. 27 Harriman 1981; Schuyler 1967a, 1975b

S. flaccidifolius 27 Schuyler 1967a

S. atrovirens 25: 28; 3] Harriman 1981; Schuyler 1963, 1975b, A. & D.
Léve 1981; Harriman 1981

S. hattorianus 28 Schuyler 1967a

S. cyperinus 50,32; 33;.35 Harriman 1981; Hicks 1928, Schuyler 1967a;
Harriman 1981

S. lineatus 18 Schuyler 1963, 1967a

S. pendulus 20 Schuyler 1963, 1967a, Harriman 1981

Trichophorum planifolium 46 Schuyler 1971a

Schoenoplectus smithii 20 Schuyler 1972a

Sch. purshianus 19 Schuyler 1972a

Sch. subterminalis 37 Schuyler 1975b

Sch. torreyi 21; 35 A. & D. Léve 1981; Schuyler 1975b

Sch. americanus 39 Hicks 1928, Schuyler 1975b

Sch. pungens 37; 38; 39 Harriman 1981; Hicks 1928; Schuyler 1975b

Sch. validus 21 Hicks 1928, pia 1975b, A. & D. Love
1981, Harriman

Sch. acutus 18. 19) 2) Harriman 1981; Schuyler 1975b; A. & D. Love
1981

Sch. etuberculatus a ase

Sch. fluviatilis AT: 52; 55 Harriman 1981; A. & D. Love 1981; Hicks 1928

Sch. robustus 53, 54, 55 Hicks 1928

Sch. novae-angliae

keel-shaped or crescentform in cross section and typically have sclerenchyma girders either
between the vascular bundles and the abaxial epidermis or between the vascular bundles and
epidermis on both abaxial and adaxial sides of the blade. Bulliform cells are also present

34 BARTONIA

in some species (Schuyler 1971a). In some species of Schoenoplectus (e.g., Sch. purshianus
and Sch. validus) the leaves are reduced to short protrusions from the bladeless sheaths. In
other species (e.g., Sch. americanus, Sch. pungens, Sch. torreyi, Sch. etuberculatus, and Sch.
subterminalis), they have short to elongate blades borne only near the base or they can be
both basal and cauline. The sheaths of some species of Schoenoplectus are eligulate while
others bear a short ligule at the adaxial junction of sheath and blade. The inner band is
membranous in some species and has a V-shaped to truncate orifice that may split with age,
while in others it is firm and has a concave to convex scarious-tipped orifice. Blades of
Schoenoplectus vary greatly in shape as seen in cross section. They may be dorsally
flattened at base becoming laterally flattened to sub-trigonous near the apex as in Sch.
pungens and Sch. americanus, triangular-channeled as in Sch. etuberculatus and Sch. torreyi,
crescentform-capillary as in Sch. subterminalis, or V-shaped to dorsally flattened with a
prominent midvein and indistinct lateral veins usually scabrous beneath as in Sch. robustus,
Sch. fluviatilis, and Sch. novae-angliae.

Anatomically, chlorenchyma is generally restricted to the abaxial side of the blade below
the air cavities, but in Schoenoplectus americanus, Sch. etuberculatus, Sch. subterminalis,
Sch. robustus, Sch. fluviatilis, and Sch. novae-angliae, chlorenchyma is on both abaxial and
adaxial sides of the blade, although in Sch. etuberculatus and Sch. subterminalis, the adaxial
chlorenchyma is not as well developed as the abaxial chlorenchyma. The margins of the
blades of Sch. americanus have chlorenchyma equally developed adaxially and abaxially.
Frequently, thin diaphragms develop across air cavities. These may serve a particular
function in regards to adaptation to an aquatic environment (Schuyler 1971la, 1972b).
Stomates are present in both the adaxial and abaxial epidermis of some species (e.g., ae
americanus, Sch. etuberculatus, Sch. robustus, Sch. fluviatilis, and Sch. novae-angliae). |
other species, e.g., Sch. torreyi and Sch. subterminalis, stomates are restricted to the cee
epidermis and only a portion of the adaxial epidermis near the margin of the blade (Schuyler
1971a, 1972b).

In marsh habitats along tidal rivers where species of Schoenoplectus co-occur, I have
observed the species with tubers, leaves with well-developed blades which extend above the
mid portion of the culm, and leaf-like involucral bracts (e.g., Sch. novae-angliae and Sch.
robustus) growing in the more densely vegetated portions of the marsh, back from open
water, while those species with reduced leaves (e.g., Sch. pungens and Sch. validus) tend to
be concentrated on the outer edges of the marsh (facing open water) where the density of
vegetation is much less. In portions of the marsh where vegetation is dense, there should
be increased competition for resources, space, and sunlight. Those plants with elongate leaf
blades and involucral bracts, as well as starch-storing tubers would have an advantage over
those having reduced leaves and involucral bracts (having only the stem and 1-few basal
leaves capable of photosynthetic activity) and non-tuberous rhizomes. Schuyler (1971a)
suggested that the well-developed leaf blades of some species of Schoenoplectus may repre-
sent an evolutionary modification of other species in this genus which have reduced leaf
blades

INFLORESCENCE. Inflorescences of Scirpus are typically broad, with an umbelliform
or corymbose terminal panicle of many rays. The rays are usually elongate, stiff, ascending
to divaricate as in species characteristic of S. georgianus, or flexuous and pendulous as in
species characteristic of S. cyperinus. The rays are terete and ribbed and may be smooth,
or upwardly scabrous in some species. Spikelets are borne singly at ray tips, or in

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 35

glomerules of 2-many. Involucral bracts are typically leaf-like with well-developed blades
and in some species greatly exceed the inflorescence, e.g., as in S. cyperinus, or are reduced
and shorter than the inflorescence as in S. pendulus. Trichophorum typically has a single
spikelet at the summit of the culm, subtended at the base by a single, scale-like involucral
bract which is thickened at the apex. Inflorescences of Schoenoplectus are typically a
A aeotien corymb or head, the lowest involucral bract appearing as a continuation of the

. The pseudolateral appearance of the inflorescence in some species is less pronounced,
ete han in those species having elongate, flattened, leaf-like involucral bracts. Species
such as Sch. robustus or Sch. novae-angliae have a number of well-developed, elongate,
involucral bracts subtending the inflorescence that at maturity spread out, giving the
inflorescence a terminal appearance. However, the lowest involucral bract is usually longer
than those above, and at anthesis is erect, giving the inflorescence a pseudolateral
appearance. In some species of Schoenoplectus, particularly those that have somewhat open,
anthelate inflorescences (e.g., Sch. validus, Sch. acutus, and the western North American
species Sch. californicus), the lowest involucral bract is erect and culm-like, but there is also
present one or two additional small involucral bracts above.

Inflorescence rays of Schoenoplectus are smooth, angled, terete or somewhat flattened,
either elongate as in Sch. validus and Sch. fluviatilis, or reduced and inconspicuous as in Sch.
americanus and Sch. pungens. Spikelets are borne singly at ray tips or in glomerules of
2-many. Involucral bracts are usually | (-3) in those species with the lowest bract appearing
as a continuation of the culm, or 3-6 in those species such as Sch. fluviatilis which have
elongate, leaf-like involucral bracts.

SPIKELETS. The spikelets of Scirpus, Trichophorum, and Schoenoplectus consist of
spirally imbricated scales that each subtend a flower. In Scirpus, the spikelets are generally
ovoid and range from 2-6 (-8) mm long and 1-3 mm wide, but in some species (e.g., S.
divaricatus, S. lineatus, and S. pendulus) they are ovoid to lance-ovoid or linear-cylindric
and elongate up to 1.5 cm. In Trichophorum the spikelets are ovoid to ovoid-ellipsoidal,
generally small, and range from 3-8 mm long and 2-4 mm wide. Spikelets of Schoenoplect-
us are generally ovoid to oblong-ovoid and range in size from 0.4-6 cm long and 3-12 mm
wide. In Sch. novae-angliae and Sch. robustus f. protrusus, the spikelets are elongate and
range from 2-6 cm long.

FLOWERS. The typical flower of Scirpus, Trichophorum, and Schoenoplectus is perfect.
The ovary is typically 3-angled and crowned with a 3-branched, caducous style continuous
with the ovary, but in Schoenoplectus smithii and Sch. purshianus, the ovary is 2-sided and
crowned with a 2-branched style. Style branches of Scirpus and Trichophorum are subulate
and have minute, rounded protrusions covering them. Style branches of Schoenoplectus are
furrowed or minutely scaly and sometimes have ascending to divergent hairs. Bristles are
typically six in all three genera, but in some species and forms they are abortive (e.g.,
Scirpus georgianus and Schoenoplectus smithii f. smithii). Bristles are retrorsely barbed in
many species, but in Trichophorum they have spinulose hairs, or vestiture may be lacking
altogether as in Scirpus pendulus. Stamens are typically 3, but 1 or 2 may be abortive in
some individuals which only have | or 2 functional. Anthers are narrowly oblong and
basifixed, and the thecae are parallel and longitudinally dehiscent. The apex may be
apiculate as in Scirpus and Trichophorum or have a glabrous, barbed or prickly, triangular
to acute appendage as in Schoenoplectus. Pollen of all three genera are typically
uniaperturate, psilate, binucleate or trinucleate, subspheroidal in polar view, and triangular

36 BARTONIA

IG. 1. Achenes of Scirpus, bar = 400 um. (a) S. atrovirens 7 14877, FARM, PH). (b) 5 hattorianus
callard 11493, US). (c) S. ae (Harvill 28935, FARM, PH). (d) S. flaccidifolius (Fernald = Long 10140,
US). (e) S. expansus (Small s.n., NY, PH, US). (f) S. uae (Wieboldt et al. 5595, PH).

to obovoidal in equatorial view (Tucker 1989).

ACHENES. Achenes of Scirpus (Figs. la-1f and 2a, 2c-2e) are typically plano-convex
or subtrigonous as in S. atrovirens, but may be sharply trigonous as in S. divaricatus. They
are usually less than 1.5 mm long and have thin, delicate pericarps (Schuyler 1971a). The
surface is typically minutely papillate, or faintly so. Each epidermal cell has a convex body
and apical nodules which give the surface a papillate texture (as seen under high
magnification). The inner cell walls have a wavy configuration. These undulating walls
may be high in some species or low in others. Buttresses frequently occur between the cell
walls and the achene body. These vary in size and may be well developed in some species
or very small in others. As mentioned previously, cells have a conical or convex body with
a single apical nodule or a series of compactly arranged apical nodules (Schuyler 1971a,
1971b). Achenes of Trichophorum (Fig. 2b) are typically trigonous. The surface is minutely
papillate (as seen under high magnification). Achene epidermal cells have a conical body,
apical nodules, and non-wavy inner cell walls that lack ridge-like buttresses between the
lateral cell walls and the body (Schuyler 1971a, 1971b). Achenes of Schoenoplectus ses
3 and 4) may be trigonous as in Sch. torreyi or Sch. fluviatilis, plano-convex as in

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 37

G. 2. Achenes ee Scirpus s.s. and Trichophorum, bar = m except for S. cyperinus (c) which = 200um

f peels (Harvill 14378, FARM, PH). (b) 7. dae sion 6537, US). (c) S. cyperinus (Egler 40- 257,
US). (d) S. lineatus poe Id & Moore 15045, NY,PH,US). (e) S. divaricatus (Kearney 1633, US). (f) S.
polyphyllus Posters 43092, US).

pungens or Sch. novae-angliae, or somewhat lenticular as in Sch. purshianus. They are
usually more than 1.5 mm long, typically are smooth with comparitively hard walls, and at
maturity are dark brown to blackish. In some species, e.g., Schoenoplectus purshianus, they
may be shallowly pitted, or in some species occurring out of the range of this treatment, they
may be transversely rugulose. Achene epidermal cell structure is very diverse among species
in this group. Some species have a great deal of internal differentiation, i.e. nodular peaks,
buttresses, etc., while others have very little. In some species, particularly the smaller
annuals, epidermal cells are vertically elongate and have little internal differentiation.
Others, such as Schoenoplectus validus, have shorter cells and a great deal of internal
differentiation (Schuyler 1971a, 1971b).

38 BARTONIA

Fic. 3. Achenes of Schoenoplectus s.l., bar = 400um. (a) Sch. validus (A. Ruth s.n., US). (b) Sch. acutus
mage & Ekvall 43092, US). (c) Sch. americanus (Fernald & Long 3777, PH). (d) - pungens (McCarthy
e) Sch. smithii (Shull 204, US). (f) Sch. eae (Harvill 20394, FARM,P

TAXONOMIC TREATMENT

Key to Genera

A. Involucral bract one, scale-like; spikelet solitary, terminal. Plants (in Virginia) of dry
upland forests and deciduous woods

an ule wh whe nw eg re ele oe ae ee en ete tans i eg Ohtani ek ea 2. Trichophorum

A. Involucral bracts | to many, not scale-like; spikelets numerous, rarely solitary. Plants

growing in water along rivers, estuaries, pond margins or in seasonally wet substrates
such as swamps, marshes, or shallow pools (B)
B. Involucral bracts (1-) 2-6, leaf-like, flattened; culms leafy (C)
C. Spikelets less than | cm long; achenes usually less than 1.5 mm long
tek awe ko SEA le Cae ee Oe eee te ae eR ee ee ee oe 1. Scirpus
C. Spikelets more than 1 cm long; achenes more than 1.5 mm long
CGAP RS ERE © REE ey ee ey ee ee eS 3. Schoenoplectus
B. Involucral bracts 1 (-3), the lowest elongate, appearing as a continuation of the
culm, terete or triangular; culms naked or leafy only towards base
ENG Ga Seek See wales ew ey ee lee Se ee ee 3. Schoenoplectus

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 39

FIG. 4. Achenes of Schoenoplectus s.\., bar = 400m except for Sch. fluviatilis (d) and Sch. novae-angliae (b)
which = 1 mm. (a) Sch. robustus (L. B. Smith 5523, US). (b) Sch. novae-angliae (Hotchkiss & Uhler 7143, US).
. a etuberculatus (Canby s.n., US). (d) Sch. fluviatilis (Ward s.n., US). (€) Sch. subterminalis (Nelson 1935,

(f) Sch. torreyi (Harvill 26411, FARM,GMUF,PRH).

1. Scirpus L., Sp. Pl. 47. 1753.
Type species: S. sylvaticus L.

Rhizomatous perennials with fibrous roots. Culms 0.3-2 m tall, trigonous. Leaves
well-developed, sheathed at base, the sheaths glabrous, with a hyaline, purple-dotted,
U-shaped (or convex to truncate on upper culm leaves) orifice. Inflorescences terminal and
occasionally in upper leaf axils, umbel-like corymbs subtended by (1-) 2-6 leaf-like
involucral bracts, many-rayed, the rays subtended by small bracts enclosed by tubular
prophylls at their base; spikelets in glomerules or solitary at tips of pedicels; scales of the
spikelet spirally arranged, each subtending a single hypogynous flower; stamens 3 or fewer;
anthers with apiculate apex, rounded at base; style 3- or 2-branched with minute rounded
protrusions. Achenes usually less than 1.5 mm long, minutely papillate; epidermal cell
structure of wavy inner cell walls with buttresses and apical nodules; bristles 6 or fewer at
base of achene, retrorsely barbed or smooth.

40 BARTONIA

Key to Species of Scirpus

A. Bristles retrorsely barbed, nearly straight (curled and crisped in No. 2) or wanting (B)
B. Spikelets solitary on the branches of the inflorescence, rarely in clusters of 2 or 3,
the lateral ones of each group long-pedicelled; mature scales with broad green
SENN oct Sg ela ok es eta heel kw ORE Ri 1. S. divaricatus
Spikelets clustered in glomerules on the branches of the inflorescence, rarely
solitary; midrib of mature scales narrow, inconspicuous (C)
C. Leaves 10-20 per culm, crowded; spikelet scales rounded, about as long as
wide; bristles curled and crisped, to over twice as long as achene when fully
RN a a a ny ele ee ee eee Wh ce eee 2. S. polyphyllus
Leaves fewer than 10 per culm, internodes prolonged, spikelet scales ovate to
elliptic-ovate; bristles at most 1-1/4 as long as achene or wanting. (D)
D. Bristles coarsely barbed almost to the base; rays of the inflorescence
antrorsely scabrous their entire length (E)
E. Leaf sheaths reddish near base; blades 10-25 mm wide; achenes
subtrigonous, 0.9-1 mm wide; bristles frequently caducous.
Sls We HOE AGEN ae nd ny eae aed ore ew ge 3. S. expansus
E. Leaf sheaths green throughout; blades 2-8(-10) mm wide; achenes
plano-convex, 0.6-0.7 mm wide; bristles persistent.
stg hej ele ane GE iad racy Shag gle seca ote nee NC Inlay Ae 4. S. ancistrochaetus
D. Bristles barbed only above the middle; rays of the inflorescence smooth or
only slightly scabrous near the apex (F)
F. Bristles 0 (-3), shorter than the achene; barbs if present concentrated
near the tips of the Gristlés <5 6665 eh Secs Ss x 5. S. georgianus
F. Bristles usually 5 or 6, shorter than to slightly longer than the achenes;
barbs extending downward from the tips of all or at least some of the

bristles (G)

G. Mature culms reclining with inflorescences bending over to (or near-
ly to) the ground; lower leaf sheaths and blades inconspicuously
septate-nodulose, nearly smooth; glomerules frequently with fewer
than 15 spikelets; lower scales blackish, slightly mucronate.

ee ee oe rt Se ee Se ree ee 6. S. flaccidifolius

G. Mature culms ascending or nearly so; lower leaf sheaths and blades
conspicuously septate-nodulose or nearly smooth; glomerules
frequently with more than 15 spikelets; scales blackish to brownish,
mucronate (H).

H. Lower leaf blades and sheaths septate-nodulose; spikelets (2-)

.5-5 (-8) mm long; scales brownish; longer bristles exceeding

MINE SS a ee aes Ca ee a 7. S. atrovirens

Lower leaf blades and sheaths inconspicuously septate-nodulose,

nearly smooth; spikelets 2-3.5 mm long; scales blackish; longer

bristles shorter than to equaling achenes ... 8. S. hattorianus
A. Bristles smooth, curled and crisped, elongated (I)

I. Mature bristles greatly elongated, strongly curled, exceeding the scales and

giving the spikelets a woolly appearance; spikelets rarely solitary; scales with

inconspicuous midribs; achenes less than | mm long ...... 9. S. cyperinus

w

0

=

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 41

—

Mature bristles shorter, curled, rarely exceeding the scales; spikelets predomi-
nantly solitary or in glomerules of 2 (-3); scales with conspicuous keeled, green
midribs; achenes more than | mm long (J)
Leaves (5-) 8-15 mm wide; ultimate rays of the inflorescence and pedicels
scabrous, with axillary bulblets; mature culms reclining with the inflor-
escences bending over to (or nearly to) the ground with 2 to 3 lateral
inflorescences in addition to the terminal inflorescence .. . 10. S. lineatus
Leaves 3-8 (-10) mm wide; ultimate rays of the inflorescence and pedicels
smooth, lacking axillary bulblets; mature culms ascending with terminal
inflorescence only or rarely with | or 2 lateral inflorescences

LT EEE TON Te et Cee ee eee eee 11. S. pendulus

=

01. Scirpus divaricatus Ell., Sketch Bot. S. Carolina 1: 88. 1816. Fig. 2e.
Isolepis divaricata (Ell.) A. Dietr., Sp. Pl. 2: 131. 1832.

DESCRIPTION. Rhizomes short, obscure. Culms at maturity weak, reclining, obtusely
angled, leafy, the nodes proliferating. Leaves 10-20 per culm; sheaths eligulate; blades 1-8
mm wide, crowded, smooth to weakly scabrous along margins and midvein beneath.
Involucral bracts 3-5, reduced, the lowermost leaf-like; rays firm, upwardly scabrous along
the ribs, elongate, at maturity becoming divaricate to sub-pendulous, frequently proliferating
in the axils; spikelets lance-ovoid to linear cylindric, 3-15 mm long, 1.5-2 mm wide, centra
spikelet of each group sessile, the lateral ones long pedicelled, predominantly solitary at ray
tips or rarely in clusters of 2-3; scales cucullate-ovate, 1-1.8 mm long, 1-1.3 mm wide, the
sides light brown to reddish, scarious margined, with broad green midrib ca. 0.6-0.7 mm
wide, rounded to short mucronate at the apex; stamens up to 3; anthers 0.8-0.9 mm long;
style 3-branched. Achene trigonous with protruding angles and concave sides, ellipsoid to
obovoid, somewhat stipitate at base, short-beaked, 0.7-1 mm long, 0.7-0.8 mm wide, surface
smooth, yellowish brown to brown; bristles 6, minutely barbellate at the apex, reddish,
shorter than to equaling the achene.

FLOWERING AND FRUITING. Late May-June; June-Aug.

DISTRIBUTION. Occurs in Chesapeake, Dinwiddie, Greensville, Nansemond, Prince
George, Southampton, and Sussex counties in southeastern Virginia (see Harvill et al. 1992);
Florida to Louisiana, north to southeastern Virginia, southern Tennessee, and southeastern
Missouri.

HABITAT IN VIRGINIA. Lowland swamps and wooded bottomlands along rivers (often
associated with Taxodium).

DISTINGUISHING FEATURES. This species is distinguished by its sharply trigonous
achenes; scales with broad, green midrib (ca. 0.6-0.7 mm wide); and at maturity, its long,
divaricately branched inflorescence with bulblets developing at the nodes.

02. Scirpus polyphyllus Vahl, Enum. Pl. 2: 274. 1806. Figs. 2f, 5. (See Schuyler in Proc.
Acad. Nat. Sci. Philadelphia 119: 312, 1967, for a complete list of synonyms.)

DESCRIPTION. Rhizomes short, tough, fibrous, 4-6 mm thick. Culms obtusely angled.
Leaves 10-20 per culm; sheaths short ligulate at adaxial junction of sheath and blade; blades
3-10 mm wide, to 3 dm long, scabrous on margins and midvein beneath. Involucral bracts
3-4, the lowermost leaf-like; rays firm, smooth, ascending to divaricate, frequently
proliferating inthe axils; spikelets ovoid, rufescent, 3-4 mm long, 2-2.5 mm wide, in

BARTONIA

42

FIG. 5. Scirpus polyphyllus. (a) habit. (b) spikelet scale.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 43

glomerate clusters or 3-8 at ray tips; scales rounded, 1-1.5 mm long, about as wide, reddish

brown to brownish black, the greenish midrib short mucronate at apex; stamens 3; anthers

0.9-1 mm long; style 3-branched. Achene plano-convex, obovate or nearly obtriangular,

1.1-1.3. mm long, 0.8-1 mm wide, smooth, pale brown; bristles 6, curled and crisped,

retrorsely barbed above the middle, to over twice as long as the achene when fully extended.
L ING AND FRUITING. June-early July; July-Sept.

DISTRIBUTION. Widely distributed in the piedmont and mountains, but absent from
southeastern Virginia and the outer coastal plain (see Harvill et al. 1992); additional counties
not recorded by Harvill et al. 1992: Caroline Co., Strong, Kelloff & Bradley 677 (GMUF)
and Roanoke Co., Woods 743 (PH); New Hampshire to Missouri, south to Georgia,
Tennessee, and Alabama.

HABITAT IN VIRGINA. Generally shaded areas along stream edges, roadside ditches,
magnolia swamps, springy ground in woodland, wet meadows, swampy bottomland,
sphagnous swales, wet clearings, and swampy streamsides.

DISTINGUISHING FEATURES. This species can be distinguished by its many-noded
culms with 10-20 leaves; ovoid to rounded-ovoid spikelets; rounded, dark brown to brownish
black mucronate scales; and obovate achenes bearing 6 curled and crisped bristles, retorsely
barbed above the middle, to over twice as long as the achene when fully extended.

03. Scirpus expansus Fern., Rhodora 45: 293. 1943. Fig. le.

DESCRIPTION. Rhizomes reddish, elongate, 4-5 mm thick. Culms obtusely angled, stout,
up to 1.5 cm wide near base. Leaves 6-9 per culm; sheaths reddish near their bases,
septate-nodulose, short-ligulate at adaxial junction of sheath and blade; blades 1-2.5 cm
wide, to 6 dm long, septate-nodulose, scabrous on margins and midvein beneath. Involucral
bracts 3-8, leaf-like, the lowermost overtopping the inflorescence; rays firm, ascending to
divaricate, antrorsely scabrous along the ribs. spikelets ovoid, 2-6 mm long, 1-2.5 mm wide,
in glomerate clusters of 3-12 at ray tips; scales ovate, keeled, 1.3-2.0 mm long, 1-1.2 mm
wide, brown to brownish black, the green midrib extending as a short mucro at apex;
stamens 3; anthers ca. | mm long; style 3-branched. Achene subtrigonous to plano-convex,
broadly ellipsoid, 1-1.2 mm long, 0.9-1 mm wide, yellowish brown; bristles 6, retrorsely
barbed to base, frequently disarticulating from the achene and remaining within the scales,
shorter than to equaling or exceeding the achene.

FLOWERING AND FRUITING. Mid June-early July; July-Aug.

DISTRIBUTION. Primarily in the mountains of midwestern and southwestern Virginia
with several stations on the inner coastal plain (see Harvill et al. 1992); Maine to Michigan,
south to Georgia.

HABITAT IN VIRGINIA. Creek edges, springheads, meadows, mountain streams, and
seepage slopes.

DISTINGUISHING FEATURES. S. expansus is similar to S. ancistrochaetus in that the
rays of the inflorescence are antrorsely scabrous their entire length and the hypogynous
bristles are almost identical, but can be distinguished from S. ancistrochaetus by a number
of other characters. The septate-nodulose sheaths of S. expansus are red-tinged near their
bases and have wide, septate-nodulose blades that range in size from 10-25 mm wide, while
those of S. ancistrochaetus are green near their bases with blades ranging from 2-8(-10) mm
wide. Culms of S. expansus are much stouter at the base than those of S. ancistrochaetus,
ranging up to 15 mm in width. The inflorescence of S. expansus is much more branched

44 BARTONIA

than that of S. ancistrochaetus. The achenes of S. expansus are subtrigonous, wider (0.9-1
mm), and subtended by brownish black scales, while those of S. ancistrochaetus are
plano-convex, narrower (0.6-0.7 mm), and subtended by reddish brown scales.

04. Scirpus ancistrochaetus Schuyler, Rhodora 64: 44. 1962. Fig. If.

DESCRIPTION. Rhizomes short, woody. Culms obtusely angled, to 1 cm wide near base.
Leaves 5-9 per culm, septate-nodulose at base, scabrous on margins and midvein beneath;
sheaths short ligulate at adaxial junction of sheath and blade; blades 2-8 (-10) mm wide, up
to 5 dm long. Involucral bracts 4-5, the lowermost leaf-like; rays firm, ascending, arching
at maturity, antrorsely scabrous along the ribs; spikelets ovoid, 3-4 mm long, 2-3 mm wide,
in glomerate clusters at ray tips; scales ovate-elliptic, 1.5-2 mm long, 1-1.2 mm wide,
reddish brown, the green midrib extending as a short mucro at apex; stamens 0-3; anthers
1-1.1 mm long; style, 3-branched. Achene plano-convex, elliptic-obovate, 1-1.6 mm long,
0.6-0.7 mm wide, yellow-brown; bristles 6, retrorsely barbed nearly to the base, equaling
to slightly longer than the achene.

FLOWERING AND FRUITING: Late May-June; June-July.

DISTRIBUTION. Rare in Alleghany, Bath, and Rockingham counties; New Hampshire,
Vermont, Massachusetts, New York, Pennsylvania, West Virginia, and Maryland. This
species is listed as LE (Endangered) by the United States Fish and Wildlife Service (1991),
and LE by the Virginia Department of Conservation and Recreation (1993).

HABITAT IN VIRGINIA. Mountain sinkhole ponds.

DISTINGUISHING FEATURES. Scirpus ancistrochaetus is similar vegetativiely to other
species in the genus Scirpus that have stiff, ascending to divergent primary inflorescence
rays, the ultimate rays tipped by l-many glomerules of spikelets (e.g. S. atrovirens, S.
expansus, S. flaccidifolius, S. georgianus, S. hattorianus and S. polyphyllus). Frequently
confused with S. atrovirens, it differs in a number of characteristics. The inflorescence rays
of S. ancistrochaetus are antrorsely scabrous their entire length, while in S. atrovirens and
other species with similar inflorescence type (with the exception of S. expansus), they are
smooth or only slightly antrorsely scabrous near the tip. The achenes of S. ancistrochaetus
are generally longer than those of S. atrovirens (1-1.6 mm long in the former, 1-1.3 mm long
in the latter), and the bristles of S. ancistrochaetus are rigid with sharp-pointed, retrorse teeth
that extend nearly to the base, while those of S. atrovirens are frequently wrinkled with
round-tipped, retrorse teeth that are concentrated towards the tip of the bristle (Schuyler
1962).

05. Scirpus georgianus Harper, Bull. Torrey Bot. Club 27: 331. 1900. Fig. Ic.
Scirpus atrovirens var. georgianus (Harper) Fern., Rhodora 23: 134. 1921.

DESCRIPTION. Rhizomes short, stout, 5-8 mm thick. Culms obtusely angled. Leaves
firm, 4-9, the lower leaves and sheaths with few septate partitions between the veins; sheaths
short ligulate at adaxial junction of sheath and blade; blades 0.2-1.5 cm wide, to 5 dm long,
scabrous on margins and midvein beneath; Involucral bracts 2-4, the lowermost leaf-like;
rays firm, smooth, ascending to divaricate; spikelets ovoid, 2-4 mm long, 1-2 mm wide, in
glomerate clusters at ray tips, these usually with 15 or more spikelets; scales rounded-ovate,
1.2-1.8 mm long, 0.8-1 mm wide, brownish to brownish black, the green midrib short
mucronate at apex; stamens 2 or 3; anthers ca. 1 mm long; style 3-branched. Achene
plano-convex, elliptic, 0.7-1.1 mm long, 0.5-0.6 mm wide, smooth, pale brown; bristles

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 45

Scirpus georgianus ; .
Scirpus atrovirens

Joh/e\
y7 aS
é Cust,
a

So eI Jee

FIG. 6. Distribution of Scirpus atrovirens and Scirpus georgianus in Virginia.

ose
wea
Paes

usually absent or rarely represented by 1-3 short ones, less than 3/4 the length of the achene.
FLOWERING AND FRUITING. Late May-June; June-Aug., sometimes into Sept. and

DISTRIBUTION. Widely distributed on the piedmont and coastal plain of Virginia (Fig.
6); Local on Prince Edward Island and in Wisconsin, west to Nebraska and eastern Texas,
south to Georgia (more common in southeastern United States).

HABITAT IN VIRGINIA. Damp soils in swales, marshes, wet meadows, roadside ditches,
borders of beaver ponds, and along streams and creeks.

DISTINGUISHING FEATURES. See S. atrovirens.

06. Scirpus flaccidifolius (Fern.) Schuyler, Rhodora 69: 198. 1967. Fig. 1d.
Scirpus atrovirens var. flaccidifolius Fern., Rhodora 40: 396. 1938.

DESCRIPTION. Rhizomes stout, 4-7 mm thick. Culms obtusely angled, weak, at maturity
arching or reclining, the inflorescences frequently bending over to the ground. Leaves 8-12,
the lower leaves and sheaths with few septate partitions between the veins; sheaths short
ligulate at adaxial junction of sheath and blade; blades 0.6-1.2 cm wide, up to 6 dm long,
scabrous along margins and midvein beneath. Involucral bracts 2-4, the lowermost leaf-like;
rays firm, smooth, ascending to divaricate; spikelets ovoid, 3-3.2 mm long, 2-3 mm wide,
in glomerate clusters of fewer than 15 at ray tips; scales elliptic to elliptic-obovate, 1.3-1.9
mm long, 1-1.1 mm wide, brown to brownish black, the green midrib prolonged as a short
mucro at apex; stamens 2 or 3; anthers ca. | mm long; style 3-branched. Achene
plano-convex, elliptic, 1-1.2 mm long, 0.6-0.7 mm wide, pale brown; bristles 6, retrorsely
barbed above the middle, shorter than to equaling the achene.

FLOWERING AND FRUITING. Late May-early June; June-July.

DISTRIBUTION. Known only from southeastern Virginia in Greensville, Southampton,
and Sussex counties and northeastern North Carolina. This species is listed as S1 (extremely
rare in the state) by the Virginia Department of Conservation and Recreation (1993).

HABITAT IN VIRGINIA. Wooded bottomlands.

DISTINGUISHING FEATURES. See S. atrovirens.

07. Scirpus atrovirens Willd., Enum. Pl. 1: 79. 1809. Fig. la. (See Schuyler in Proc. Acad.
Nat. Sci. Philadelphia 119: 310, 1967, for a complete list of synonyms.)

DESCRIPTION. Rhizomes stout, 5-8 mm thick. Culms obtusely angled. Leaves firm,
4-9; sheaths short ligulate at adaxial junction of sheath and blade, the lowermost

46 BARTONIA

septate-nodulose; blades 0.2-1.8 cm wide, to 5 dm long, scabrous along margins and midvein
beneath, the lower septate-nodulose. Involucral bracts 2-5, the lowermost leaf-like; rays
firm, smooth, ascending to divaricate; spikelets ovoid, (2-) 3.5-5 (-8) mm long, 1-2 mm
wide, in glomerate clusters at ray tips, these usually with 15 or more spikelets; scales
rounded-ovate, 1.4-2.1 mm long, 0.9-1.1 mm wide, brownish, the green midrib short
mucronate at apex; stamens 2 or 3; anthers ca. | mm long; style 3-branched. Achene
plano-convex, elliptic, 1-1.3 mm long, 0.5-0.6 mm wide, pale brown; bristles 6, barbed on
upper 2/3, shorter than to slightly exceeding the achene.

FLOWERING AND FRUITING. Late May-June; June-Aug.

DISTRIBUTION. Primarily in the foothills and mountains of northern and western
Virginia with several stations on the inner coastal plain (Fig. 6); Newfoundland south to
Georgia and eastern Texas, disjunct occurrences in Minnesota (more common in northeastern
United States).

HABITAT IN VIRGINIA. Damp soils in swales, marshes, wet meadows, marshy
seepages, roadside ditches, and stream edges.

DISTINGUISHING FEATURES. Scirpus atrovirens is closely related to S. flaccidifolius,
S. georgianus, and S. hattorianus. They are all similar in habit, but differ consistently from
each other in a number of characteristics (also see discussion below). Scirpus atrovirens can
be distinguished from other species in this group by its coarse, elongate, retrorsely barbed
bristles which frequently exceed the achene; septate-nodulose sheaths and leaf blades; and
longer spikelets (2-) 3.5-5 (-8) mm long (vs. 2-4 mm long in other species). Scirpus
flaccidifolius has elongate, lax, and reclining culms that bend over to the ground, in contrast
to the shorter, firm and upright culms of other species in this group; longer primary
inflorescence rays which rarely contain more than 15 spikelets per glomerule; relatively
smooth to inconspicuously septate-nodulose lower leaf sheaths and blades; and short, slightly
mucronate tips of lower scales (Schuyler 1967c). Scirpus georgianus can be readily
distinguished from other members of this group by its lack of bristles or one, two, or rarely
three short to rudimentary ones. Scirpus hattorianus has delicate, retrorsely barbed,
hypogynous bristles which are shorter than to barely reaching the summit of the achene;
shorter spikelets than Scirpus atrovirens (2-3.5 mm long in S. hattorianus vs. (2-) 3.5-5 (-8)
mm long in S. atrovirens); and dark brown to black scales (Schuyler, 1967b).

DISCUSSION. The "Scirpus atrovirens group," as here specified, consists of four species
(S. atrovirens, S. georgianus, S. hattorianus, and S. flaccidifolius). These were all treated
by Fernald (1950) as S. atrovirens. Schuyler (1967a), showed that these four species can be
distinguished morphologically, and that the characters distinguishing them from each other
were consistent over their entire range. He also found that the four species could be
separated geographically as well as phenologically.

Scirpus atrovirens has a more northern range in the eastern United States, occurring
northward to southern Canada. In Virginia, it is primarily confined to upland habitats (Fig.
6). S. georgianus has a more southern range in the eastern United States, occurring only in
disjunct populations northward. In Virginia, it is primarily confined to lowland areas on the
piedmont and coastal plain (Fig. 6). Scirpus hattorianus is sympatric with S. atrovirens
northward, but it is more common in New England, Quebec, and the Maritime Provinces.
It occurs only in disjunct populations southward (Maryland, Virginia, North Carolina,
Tennessee, Alabama, and Indiana). In northern populations of S. atrovirens and S.
hattorianus where their ranges overlap, S. hattorianus is primarily found in upland meadows

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 47

and ditches, while S. atrovirens is primarily found in lowland marshes. Scirpus flaccidifolius
is a rare species known only from wooded bottomlands in southeastern Virginia and
northeastern North Carolina.

08. Scirpus hattorianus Makino, J. Jap. Bot. 8: 44. 1933. Fig. 1b.

DESCRIPTION. Rhizomes stout. Culms obtusely angled. Leaves firm, 10-18, the lower
leaves and sheaths with few septate partitions between the veins; sheaths short ligulate at
adaxial junction of sheath and blade; blades 2-10 mm wide, to 5 dm long. Involucral bracts
2-4, the lowermost leaf-like; rays smooth, ascending to divaricate; spikelets oblong-ovoid,
2-3.5 mm long, 1.5-2.5 mm wide, in glomerate clusters at ray tips, these usually with 15 or
more spikelets; scales rounded-ovate, 1.2-2 mm long, 0.9-1.1 mm wide, brownish black to
blackish, the green midrib short-mucronate at apex; stamens 2-3; anthers 0.9-1 mm long;
style 3-branched. Achene plano-convex, elliptic-obovate, 0.8-1.1 mm long, 0.6-0.8 mm wide,
pale brown; bristles 5 or 6, retrorsely barbed above the middle, shorter than to equaling the
achene.

FLOWERING AND FRUITING. Early June-July; late July-Aug.

DISTRIBUTION. Known only from a single collection: Highland County, low, wet field
just south of Monterrey, B. Mikula 6797, 23 July 1950 (FARM, WILLI); Newfoundland to
western Ontario, south to Maryland, Virginia, West Virginia, Tennessee, North Carolina,
Alabama, Ohio, Indiana, and Wisconsin (more common in northeastern United States).

DISTINGUISHING FEATURES. See S. atrovirens

09. Scirpus cyperinus (L.) Kunth, Enum. Pl. 2: 170. 1837. Fig. 2c.
Eriophorum cyperinum L., Sp. Pl., ed. 2, 1:77. 1762.
Scirpus rubricosus Fern., Rhodora 47: 124. 1945. (See Schuyler in Proc. Acad. Nat. Sci.
Philadelphia 119: 319, 1967, for a complete list of synonyms.)

DESCRIPTION. Rhizomes short, stout, up to 2 cm thick at culm bases, branching; roots
minutely scabrous-papillate, or hirsute with pale brown hairs. Culms nearly terete. Leaves
many, rigid, crowded at base, curving; sheaths reddish, with short ligule at adaxial junction
of sheath and blade; blades narrowly linear, 3-10 mm wide, to 6 dm long, harshly scabrous
along margins and midveins beneath. Inflorescence pendulous, drooping at maturity;
involucral bracts 3-5, elongate, leaf-like, with reddish brown to brownish black bases; rays
flexuous, antrorsely scabrous, involucels at base reddish brown; spikelets ovoid to broadly
ovoid, 4-6 mm long, 2-3 mm wide, in glomerules of 2-7, or sometimes in dense glomerules,
rarely solitary; scales ovate to elliptic, 1.1-2.1 mm long, ca. | mm wide, reddish brown to
brownish black, the green midrib obtuse to mucronate at apex; stamens | or 2; anthers
0.9-1 mm long; style 3-branched. Achene subtrigonous to plano-convex, ellipsoid, 0.6-0.9
mm long, 0.4-0.5 mm wide, pale-brown; bristles 6, elongate, contorted, exceeding the scales
at maturity, barbs lacking.

FLOWERING AND FRUITING. July (later in mountains); Aug.-Oct.

DISTRIBUTION. Common throughout the state (see Harvill et al. 1992); Newfoundland
to Nova Scotia, south to Florida and Texas, westwards to West Virginia, Ohio, Indiana,
Illinois, Minnesota, lowa, Oklahoma, and Mexico.

HABITAT IN VIRGINIA. Open areas of peaty depressions, marshes, meadows, pond
borders, creeks, roadside ditches, and disturbed areas.

DISTINGUISHING FEATURES. This species is readily distinguished by its nearly terete
culms; elongate, narrow, rigid, curving, scabrous-margined leaves and involucral bracts; and

48 BARTONIA
large, pendulus, paniculate-corymbose inflorescence.

10. Scirpus lineatus Michaux, Fl. Bor.-Amer. 1: 32. 1803. Fig. 2d.
Scirpus fontinalis Harper, Bull. Torrey Bot. Club 30: 322, 1903. (See Schuyler in Proc.
Acad. Nat. Sci. Philadelphia 119: 306, 1967, for a complete list of synonyms.)

DESCRIPTION. Rhizomes horizontal, stout, 5-8 mm thick. Culms obtusely angled, at
maturity weak, reclining. Leaves 4-8; sheaths eligulate; blades (5-) 8-15 mm wide, scabrous
along margins and midvein beneath. Inflorescence terminal and axillary from 2 or 3 of the
upper leaves, pendulous, divaricately branching; involucral bracts 3-5 on terminal
inflorescence, reduced, the lowermost leaf-like, 1-2 short and subulate or wanting on lateral
inflorescences; rays flexuous, upwardly scabrous along ribs, frequently developing axillary
bulblets; spikelets ovoid to lance-ovoid, 4-7 (-10) mm long, 2-3 mm wide, predominantly
solitary at ray tips or rarely in clusters of 2-3; scales ovate to narrowly elliptic, 1.8-2.5 mm
long, 2-2.2 mm wide, lustrous with brown to reddish brown sides, the green midrib cuspidate
to mucronate at apex; stamens 3; anthers 1-1.2 mm long; style 3-branched. Achene
trigonous to plano-convex, ellipsoid, beaked, with stipitate base, 1.2-1.4 mm long, 0.6-0.7
mm wide, minutely papillate, surface pale brown; bristles 6, yellowish translucent, curled
and wrinkled, shorter than to slightly exceeding the achene, barbs lacking.

FLOWERING AND FRUITING. Apr.-May; late May-July.

DISTRIBUTION. Occurs only on the inner coastal plain in Gloucester, James City, Surry,
and York counties and the City of Hampton (see Harvill et al. 1992); Florida to Louisiana
north on coastal plain to southeastern Virginia.

HABITAT IN VIRGINIA. Moist shaded places in wooded calcareous spring-heads,
meadows, swampy woods, bluffs, gullies, and swales.

DISTINGUISHING FEATURES. Scirpus lineatus is closely related to Scirpus pendulus
but can be distinguished from the latter by its 2-3 lateral inflorescences in addition to the
terminal inflorescence; scabrous rays with axillary bulblets; wider leaves; and weak, reclining
culms, whereas Scirpus pendulus has a terminal inflorescence only, or one or rarely two
lateral inflorescences; smooth rays lacking axillary bulblets; and rigid, upright culms.

NOMENCLATURE. This species has been known as Scirpus fontinalis Harper in many
North American floras including Small (1933), Beetle (1947), Fernald (1950), Gleason
(1952), and Radford et al. (1968). Schuyler (1966), upon examining the type specimens of
S. fontinalis and Scirpus lineatus Michx., discovered that they were conspecific, and that
plants being treated as S. Jineatus Michx. (by authors of the above-mentioned floras), were
actually the species described by Muhlenberg in 1813 as S. pendulus. Since S. lineatus was
described earlier than S. fontinalis, plants previously recognized as S. fontinalis should be
correctly treated as S. /ineatus, and those plants previously recognized as S. lineatus should
be correctly treated as S. pendulus.

11. Scirpus pendulus Muhl., Cat. Pl. Amer. Sept. 7. 1813. Fig. 2a. (See Schuyler in Proc.
Acad. Nat. Sci. Philadelphia 119: 307, 1967, for a complete list of synonyms.)

DESCRIPTION. Rhizomes stout, 0.5-1 cm thick. Culms obtusely angled, rigid, upright.
Leaves crowded at base, less than 10 per culm; sheaths eligulate; blades 3-8 (-10) mm
wide, to 3 dm long, scabrous along margins and midvein beneath. Inflorescence terminal or
rarely with one or two lateral inflorescences from the upper leaf axils, pendulous; involucral
bracts 3-5 on terminal inflorescence, reduced, the lowermost leaf-like, 1-2 short and subulate

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 49

or wanting on lateral inflorescences; rays flexuous, smooth, lacking axillary bulblets;
spikelets ovoid to sub-cylindric, 6-12 mm long, 2-3 mm wide, predominantly solitary at ray
tips; scales ovate, 2 mm long, 0.8-1.1 mm wide, lustrous, reddish brown or brownish, the
greenish midrib ending ina short, recurved cusp at apex; stamens 3; anthers ca. | mm long;
style 3-branched. Achene at maturity trigonous to plano-convex, ellipsoid, beaked with
stipitate base, 1-1.2 mm long, 0.6-0.7 mm wide, lustrous pale brown to grayish brown,
minutely papillate; bristles 6, yellow- to reddish translucent, curling, exceeding the achene.
ING AND FRUITING. May-June; late May-July.

DISTRIBUTION. Primarily occurs in the foothills and mountains of northern and western
Virginia with several stations in the piedmont and inner coastal plain (see Harvill et al.
1992); additional counties not recorded by Harvill et al. 1992: Albemarle Co., Stevens 5447
(FARM) and Smyth Co., Small s.n. (NY,PH); Maine, west to southern Canada, Iowa, and
eastern Colorado, south to Florida and southwest to New Mexico and northern Mexico. In
the western United States, Scirpus pendulus occurs in disjunct populations in Oregon and
California.

HABITAT IN VIRGINIA. Moist open places in swales, along streambanks, open
meadows, and roadside ditches.

DISTINGUISHING FEATURES. See S. /ineatus.

NOMENCLATURE. See S. /ineatus.

2. Trichophorum Persoon, Syn. Pl. 1: 69. 1805. nom. cons.
Type species: T. alpinum (L.) Persoon

Caespitose perennials with fibrous roots, frequently forming tussocks. Culms trigonous
or terete. Leaves basal, bladeless, or the uppermost sheaths blade-bearing and ligulate, with
closed, tubular sheaths. Inflorescence a single, terminal, erect spikelet (In North American
species); involucral bract scale-like, simulating a fertile scale of the spikelet, the apex
thickened, blunt; spikelets ovate to ovate-elliptic; scales spirally imbricate, 3-5 nerved, each
subtending a single flower; stamens 3; anthers with apiculate apex; stigmas 3, with minute
rounded protrusions. Achene trigonous, smooth to papillate; epidermal cell structure of
non-wavy inner cell walls lacking buttresses and apical nodules; bristles 0-6, filiform,
scaberulous, silky or with setulose hairs.

1. Trichophorum planifolium (Sprengel) Palla, Oesterr. Bot. Z. 63: 402. 1913. Figs. 2b,
7

Isolepis planifolia Sprengel, Neue Ent. 3: 10. 1822.

Scirpus verecundus Fernald, Rhodora 50: 284. 1948, nom. nov. for Scirpus planifolius
Muhlenberg, Descr. Gram. 32. 1817, non Scirpus planifolius Grimm, in Nova Acta
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 3: App. 259. 1767.

Eleogiton planifolia (Sprengel) A. Dietrich, Sp. Pl. 2: 98. 1833.

Eleocharis planifolia (Sprengel) Nees, Linnaea 9: 294. 1834.

Baeothryon verecundum (Fernald) A. & D. Léve, Univ. Colorado Stud. Biol. Ser. No. Ws
15. 1965.

Baeothryon planifolium (Sprengel) Sojak, Cas. Nar. Mus. Odd. Pfir. 148: 193. 1979.

DESCRIPTION. Densely caespitose perennial with fibrous roots. Culms trigonous,
upwardly scabrous on the angles, 1-4 dm tall. Leaves borne from base up to 1/4 the length

50 BARTONIA

Mark T Sting 1
FIG. 7. Trichophorum planifolium. (a) habit. (b) spikelet scale. (c) spikelet.

VIRGINIA SC/IRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 5

of the culms, the upper elongate, shorter than to equaling or exceeding the culms; sheaths
glabrous, with short firm to scarious ligule borne at adaxial junction of sheath and blade, the
lower open, whitish to pale brown, red-dotted, sub-truncate at summit, with reddish brown
to pale brown scarious margins, the upper closed, with U-shaped orifice, green, the inner
band white, scarious; blades flat, 1-1.5 mm wide, with rounded cartilaginous tip, scabrous
on margins, the lower short, subulate-thickened, the upper with elongate blades. Inflores-
cence a single, erect, terminal spikelet; involucral bract 1, scale-like, 1-6 mm long; spikelet
ovate, 5-7 mm long, 2 mm wide; mature scales ovate, 3.5-4.5 mm long, 1.5-2 mm wide,
sagittate at base, green to pale brown, the midrib prolonged into an awn up to 0.8 mm long
on scales near base of spikelet, shorter on upper scales; anthers 1-1.2 mm long; style
subulate, 3-branched, the branches with minute, rounded protrusions. Achene trigonous,
ellipsoid, 2 mm long, 1-1.2 mm wide, pale brown to light grayish brown, minutely papillate;
bristles 6, terete to flattened, up to 0.1 mm wide with ascending, setulose hairs, shorter than
to equaling the achene.

FLOWERING AND FRUITING. April; late April-May.

DISTRIBUTION. Mountains and piedmont of Virginia, absent from the coastal plain (see
Harvill et al. 1992); Maine to Ohio, south to southern New England, Delaware, Maryland,
District of Columbia, and primarily uplands of Virginia, West Virginia, Kentucky, and
Missouri.

HABITAT IN VIRGINIA. Dry upland forests, deciduous woods.

DISTINGUISHING FEATURES. T. planifolium can be readily distinguished by its
solitary spikelet at the tip of the culm; small scale-like involucral bract; narrow leaf blades
(1-1.5 mm wide); trigonous, papillate achenes; and hypogynous bristles with setulose hairs
that are shorter than to equaling the achene.

DISCUSSION. Both the names Trichophorum Pers. and Baeothryon Dietr. have been
applied to this species. Salmenkallio and Kukkonen (1989), studying the typification of both
of these names, found that Baeothryon at present is only a synonym of the genus Eleocharis
and that of the two uses of the name 7richophorum (as a section of Scirpus sensu stricto and
traditionally, for the group of species related to 7. alpinum) the traditional and correct use
of the genus should be maintained.

3. Schoenoplectus (Rchb.) Palla, nom. cons., Verh. K.K. Zool. Bot. Ges. Wien 38, Sitzb.:
49. 1888.

Type species: Schoenoplectus lacustris (L.) Palla (Scirpus lacustris L.)

Emergent aquatics; perennials or annuals, with elongate, horizontal rhizomes or tufted.
Culms trigonous or terete, glabrous, air cavities often present, leafy or leaves reduced to
short protrusions from the bladeless sheaths. Inflorescences congested and head-like or
anthelate with elongate rays, pseudolateral or pseudolateral at anthesis and appearing terminal
at maturity; involucral bracts 1-2 (-3), the lowest appearing as a continuation of the culm,
or (1-) 2-6, flattened and leaf-like; rays with tubular prophylls at base; spikelets ovoid,
ellipsoid or cylindrical; stamens 2-3; anthers with prickly or barbed appendage at apex,
Sagittate at base; style sub-flattened to strap like; stigmas 2 or 3. Achenes plano-convex,
trigonous or lenticular, brown or blackish at maturity; epidermal cell structure diverse,
bristles 0-6, when present strap- or needle-like, retrorsely barbed with straight or recurved
barbs.

BARTONIA

Key to Species of Schoenoplectus

A. Involucral bracts 1 (-3), the lowest elongate, terete or triangular, appearing as a
continuation of the culm; culms naked or leafy only towards base; scales glabrous or
with prickle-like hairs; rhizomes usually etuberous (B)

Inflorescence simple, spikelets sessile, in glomerules or solitary (C)
C. Plants lacking rhizomes; culms slender, densely tufted; achenes 1.5-2 mm long,
blackish at maturity; anthers 1 mm long or less (D)

ED.

Mature achene cuneate-obovate, plano-convex, smooth to obscurely
rugulose; bristles wanting or rudimentary, or when present, slender.
Cas cel ok oh sarin oe Mactig sales oe ie carne ae A ae a tes a ee 1. Sch. smithii

D. Mature achene rounded-obovate, unequally biconvex, prominently pitted;

PWistigs Wiles Preset Sit. og kg kk ew a he wwe o's 2. Sch. purshianus

C. Plants with elongate, horizontal rhizomes; culms mostly solitary or scattered;
achenes 2-5 mm long, whitish, drab, olivaceous or brown at maturity; anthers
2-4 mm long (E)

E.

m7

Achenes distinctly trigonous, with a beak 0.5-0.7 mm long at apex; culms
0.5-5 mm wide; scales entire (F)
F. Spikelet solitary; culms terete, filiform; leaves numerous, blades

CHCOCENIIOMI-CUAAEY ns kv ke bk ba eRe 3. Sch. subterminalis
F. Spikelets 1-4; culms trigonous, stout with concave sides; leaves 2 or 3,
blades terete or triangular-channeled ............. 4. Sch. torreyi

Achenes plano-convex, with a beak 0.4 mm long or less at apex; culms 1-10

mm wide; scales with erose or ciliate margins (G).

G. Mature culms wing-angled, 4-10 mm wide at upper sheath; involucral
bracts 1-2 (-3.5) cm long; awns shorter than to equaling the shallowly
cleft apex of scales; achenes 2-2.7 mm long; bristles flattened, with
downcurved barbs. Plants restricted to coastal plain, primarily of
brackish to saline habitats ................. 6. Sch. americanus

. Mature culms not wing-angled, relatively slender, 1-6 mm wide at upper
sheath; involucral bracts 2-15 cm long; awns exceeding the deeply
emarginate apex of scales; achenes 2.5-3.0 (-3.2) mm long; bristles
narrow, retrorsely barbed. Plants widespread in fresh to saline habitats.

NCA oe wk hie ror sek ne aa s al eA ae 7. Sch. pungens

Q)

B. Inflorescence compound, usually with elongate branches (H).
H. Culms terete; achenes 1.7-2.8 mm long, 1.3-2.0 mm wide, the subulate beak
0.1-0.3 mm long (I).
I

—

Mature scales 2.5-3.2 mm long, the apex with a short, straight awn;
spikelets 5-7 (-9) mm long, solitary or in clusters of 2-3 (-5) at ray tips;
culms soit, easily compuessed:... |.u 24.46 454 sees Saas 8. Sch. validus
Mature scales 3.5-4 mm long, the apex with a prominent, usually contorted
awn; spikelets 6-15 mm long, occasionally solitary, but mostly clustered in
glomerules of 3-7 at ray tips; culms firm ............ 9. Sch. acutus

H. Culms trigonous; achenes 3-4 mm long, 1.9-2.2 mm wide, the stout beak 0.8-1
NG os ee er hs og ae ee aes 5. Sch. etuberculatus

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 53

A. Involucral bracts (1-) 2-6, leafy, of two or more flat leaves; culms leafy; scales
pubescent; rhizomes tuberous (J)

Mature achenes trigonous, the acute angles forming an equilateral triangle in cross

section; bristles persistent, exceeding the achene; spikelets cuneate at base; scales

pale brown; plants of fresh habitats... 6. . 25 0. 602% oa 10. Sch. fluviatilis

J. Mature achenes plano-convex, dorsiventrally compressed; bristles few or wanting

or if present, shorter than to equaling the achene; spikelets rounded to truncate at

base; scales chestnut brown to rufescent; plants of brackish to saline habitats (K)

K. Mature achenes reddish brown or dark brown, lustrous, the summit truncate,

abruptly differentiated into the beak; bristles mostly caducous; plants primarily

of saline shores: and tidal marshes 2. 2..6.26 2042.44 4% 11. Sch. robustus

. Mature achenes whitish to olive brown, dull, the summit rounded, gradually

tapering into the beak; bristles persistent; plants primarily growing in marshes

within the brackish zone between fresh and saline waters of tidal river

WI: 65k vee Se ee ae 12. Sch. novae-angliae

nw

01. Schoenoplectus smithii (A. Gray) Sojak, Cas. Nar. Muz. Odd. P¥ir. 141(1-2): 62. 1972.
ig. 3e.

Scirpus smithii A. Gray, Manual Bot. Ed. 5. 563. 1867.

DESCRIPTION. Caespitose. Culms subterete to somewhat angled, septate, 1-4 dm long,
0.9-1.5 mm wide. Leaves bladeless, the upper sheaths with a few, short, callous-tipped
blades, 2-10 cm long, or 1-2 elongate blades on upper sheaths; sheaths confined to basal
portion of culm, acute, whitish to pale brown, with reddish-brown to pale brown scarious
margins. Inflorescence pseudolateral, spikelets in a single, sessile glomerule; involucral
bract 1, appearing as a continuation of culm, septate, 3/4 to 1/3 as long as the culm,
callous-tipped; spikelets 1-9, ovoid, subacute, 0.4-1 cm long, 3-5 mm _ wide; scales
oblong-ovate, 2.5-3.5 mm long, 2 mm wide, green to pale brown with scarious margins, the
midrib prolonged into a short mucro; stamens 3; anthers narrowly oblong, 0.5 mm long,
with a small, brown, triangular, prickly tip; style 2-branched, the branches smooth or with
minute scales. Achene plano-convex, cuneate-obovate, 1.5-2 mm long, 1.2-1.5 mm wide,
smooth to obscurely rugulose, lustrous, black; bristles absent (forma smithii), represented
by minute rudiments at base, sometimes with 1-3 bristles and 1-3 minute rudiments, the
bristles retrorsely barbed only at the very tip (forma /evisetus), or 4-5 and well-developed
with retrorse barbs (forma sefosus).

FLOWERING AND FRUITING. July; late July-Aug.

DISTRIBUTION. Historically known from only a single locality: Fairfax County, along
gravelly beach between tides, Hunting Creek, 3/4 mi. SSW of Alexandria, G.H. Shull 214,
12 Aug. 1902 (NY,US). New Brunswick and Quebec to Minnesota, south to southern New
England, New Jersey, Delaware, northeastern Virginia, northwestern and southeastern
Pennsylvania, northern Ohio, northern Indiana, and Illinois.

DISTINGUISHING FEATURES. Schoenoplectus smithii is closely related to Sch.
purshianus but can be distinguished from the latter by its smooth to faintly rugulose,
cuneate-obovate achenes and slender bristles which are either well-developed, short to
rudimentary, or wanting, whereas Sch. purshianus has pitted, rounded-obovate, unequally
biconvex achenes and 6 stout, retrorsely barbed bristles that exceed the achene.

DISCUSSION. Taxonomically, Sch. smithii has been confused with Sch. purshianus, and

54 BARTONIA

some authors, e.g., Beetle (1942) and Gleason and Cronquist (1963), have treated it as an
"aquatic extreme" of Sch. purshianus. The pattern of infraspecific variation is very similar
in both species and may have been the main cause of this confusion. The ecologically
differentiated forms of each species were delineated by Fernald (1950). He described Sch.
smithii as having three forms: f. smithii (bristles absent of rudimentary), f. setosus (bristles
well developed with retrorse barbs), and f. /evisetus (bristles nearly smooth). Sch.
purshianus he described as having two forms: f. purshianus (bristles well developed with
retrorse barbs) and f. williamsii (bristles absent or rudimentary). The forms of Sch. smithii
are primarily associated with alkaline substrates and frequently occur in tidal estuarine
habitats, while those of Sch. purshianus are primarily associated with more acid substrates
and are never found in tidal estuarine habitats (Schuyler 1972a). The achenes of the
specimen collected along a tidal shoreline of the Potomac River have nearly smooth bristles
which are barbed only at the apex (Fig. 3e) and can be referred to f. /evisetus. Whether a
form of either species has retrorsely barbed bristles, bristles that lack barbules, or bristles are
absent or rudimentary, seems to depend on the habitat they are restricted to. The typical
forms of both species which have retrorsely barbed bristles occur in habitats that have little
or moderate water fluctuation, while those forms with bristles lacking barbs or the bristles
are absent or rudimentary, occur primarily in habitats where there is substantial water level
fluctuation (e.g. tidal shorelines). The absence of bristles, or the absence of barbs on bristles
appears to be an adaptation to the periodic water level fluctuations that occur in estuarine
environments (Schuyler, 1972a). Tidal emergent forms usually have septate-nodulose culms,
sheaths, and leaves, while those of non-tidal habitats have septate-nodulose sheaths only.

02. Schoenoplectus purshianus (Fernald) M. T. Strong, Novon 3(2): 202. 1993. Fig. 3f.
Scirpus purshianus Fernald, Rhodora 44: 479. 1942.
Scirpus debilis Pursh, Fl. Amer. Sept. 55. 1813, non Lam. 1791.
Schoenoplectus juncoides (Roxb.) Palla ssp. purshianus (Fern.) Sojak, Cas. Nar. Muz.
Odd. Prir. 141(1-2): 62. 1972.

DESCRIPTION. Caespitose. Culms glabrous, trigonous, not septate, 3-7 (-10) dm tall,
1.5-3 mm wide. Leaves bladeless, or with a few, short, callous-tipped blades on upper
sheaths; sheaths confined to basal portion of culm, closed, septate, glabrous, the summit
acute, short mucronate, the inner band scarious, red-dotted. Inflorescence pseudolateral,
spikelets in a single, sessile glomerule; involucral bract 1, appearing as a continuation of
the culm, 0.6-1.2 dm long, callous-tipped; spikelets ovoid to cylindric, (1-) 3-8 (-16), 0.5-1
cm long, 3.5-5 mm wide; seales orbicular to rounded-ovate, 2.5-3.5 mm long, 2.5-3 mm
wide, lustrous, many-nerved, greenish to stramineous or brownish, the indistinct midrib
prolonged into a short, acute mucro at the dark brown apex; stamens 2; anthers narrowly
oblong, ca. 1 mm long, with a small, brown, triangular, prickly tip; style 2-branched, scaly,
with ascending to divergent hairs. Achene unequally biconvex, rounded-obovate, 1.7-2.0 mm
long, 1-1.5 mm wide, blackish brown, lustrous, pitted; bristles 6, light brown, stout,
retrorsely barbed the entire length, equalling to exceeding the achene.

FLOWERING AND FRUITING. Late June-July (later in mountains); July-Oct.

DISTRIBUTION. Scattered localities throughout the state, but more frequent in the
piedmont (see Harvill et al. 1992); additional county not recorded by Harvill et al. 1992:
Bedford Co., Curtiss 3702 (US); Southwestern Maine, New Hampshire, Vermont,
southeastern New York, eastern Pennsylvania, West Virginia, southern Michigan, Wisconsin,

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 55

Minnesota, south to Georgia, Alabama and Mississippi.
HABITAT IN VIRGINIA. Open swales, pond borders, reservoirs, marshes, springy
ditches, and disturbed areas.
ISTINGUISHING FEATURES. See Sch. smithii.
DISCUSSION. See Sch. smithii.

03. Schoenoplectus subterminalis (Torrey) Sojak, Cas. Nar. Muz. Odd. P¥ir. 140(3-4); 127.
1972. Fig. 4e.
Scirpus subterminalis Torrey, Fl. N. Middle United States 47. 1824.

DESCRIPTION. Submersed or emersed rhizomatous perennial. Rhizomes soft, slender,
sometimes bearing small tubers. Culms terete, filiform, septate, 0.2-1.4 m long. Leaves
numerous, elongate and flaccid in submersed plants, few, short or wanting, and stiff in
emersed plants; sheaths open, whitish, friable, the adaxial summit with a short ligule, 0.4-0.5
mm long; blades crescentform-capillary or conduplicate, septate, 0.5-1 mm wide, to 5 dm
long. Inflorescence pseudolateral, with a single, sessile spikelet; involucral bract 1,
channeled, 1-6 cm long, callous-tipped, appearing as a continuation of the culm; mature
spikelet ovoid to lanceolate, 5-15 mm long, 4-7 mm wide; scales ovate-lanceolate, acute,
membranaceous, 4-6 mm long, 2-2.2 mm wide, green to pale brown with scarious margins,
the green midrib mucronate at apex; stamens 3; anthers narrowly oblong, 3-4 mm long,
with a rounded to bulbous, thickened appendage at tip; style somewhat flattened, 3-branched,
the branches scaly, with ascending to divergent hairs. Achene trigonous-obovoid, 3-3.5 mm
long, 2 mm wide, smooth, lustrous, becoming yellowish to olive-brown, with a 0.5-0.7 mm
long beak at apex; bristles 6, reddish, retrorsely barbed, unequal, shorter than to equaling
or slightly exceeding on be beak.

FLOWERING AN UITING. June; July and Aug.

DISTRIBUTION. bn in Augusta and Caroline counties and the City of Virginia Beach
(see Harvill et al. 1992); Newfoundland to Ontario, south to Virginia, South Carolina, and
eastern Georgia; westward to northern Ohio, northern Indiana, northern Illinois, Missouri,
southern Alaska to southern Oregon, northwestern Montana, Wyoming, northern Idaho, and
California. An isolated station occurs in Utah. This species is currently listed as S1
i ge rare in the state) by the Virginia Department of Conservation and Recreation
(199

ha IN VIRGINIA. Pond margins and fresh to brackish marshes. Two growth
forms exist: submersed or stranded (leaves filiform and flaccid) and emersed (leaves
conduplicate and stiff).

DISTINGUISHING FEATURES. This species is readily distinguished by its single, sessile
spikelet subtended by an elongate involucral bract appearing like a continuation of the culm;
terete culms; open sheaths; and crescentform-capillary leaf blades.

DISCUSSION. Fernald (1950) described Schoenoplectus subterminalis as having two
orms: f. subterminalis (plants submersed or stranded, the leaves capillary and flaccid with
elongate blades) and f. terrestris (plants emersed, the leaves firm and upright, few, short or
wanting).

04. Schoenoplectus torreyi (Olney) Palla, Allg. Bot. Z. Syst. 17, Beilage 10/12: 3. 1912.
Fig. 4f.
Scirpus torreyi Olney, Proc. Prov. Frankl. Soc. 1: 32. 1847.

56 BARTONIA

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate, flaccid, brownish, 2-4
mm thick. Culms 0.4-1.5 m tall, 3-5 mm wide, trigonous, with concave sides, usually
solitary. Leaves 2-3, elongate, confined to basal portion of culm; sheaths septate, inner ban
scarious, disintegrating with age, the prominant mid and laterally branching veins remaining,
the summit short ligulate at adaxial junction of sheath and blade; blades 1.5-4 mm wide, to
1.2 m long, septate, terete or triangular-channeled, rigid or somewhat flaccid in submersed
habitats. Inflorescence pseudolateral, with a single glomerule of 1-4 spikelets; involucral
bract 1, erect, appearing as a continuation of the culm, 3-16 cm long; spikelets ovoid to
narrowly cylindric, cuneate at base, 1-1.8 cm long, 4-7 mm wide; scales ovate, 4-7 mm long,
2-3 mm wide, smooth, lustrous, pale brown to golden brown, the green midrib short
mucronate at apex; stamens 3; anthers narrowly oblong to linear, 2.5-4 mm long, with a
rounded to bulbous, thickened appendage at tip; style somewhat flattened, 3-branched, the
branches glabrous. Achene trigonous-obovoid, 3.4-5 mm long, 2-2.2 mm wide, smooth,
yellowish to olive-brown, with a 0.5-0.7 mm long beak at apex; bristles 6, retrorsely barbed,
slightly shorter than to exceeding the achene.

FLOWERING AND FRUITING. Late June-July; late July-Sept.

DISTRIBUTION. Known only from Agusta and Rockingham counties in Virginia (see
Harvill et al. 1992); New Brunswick to Manitoba, south to southern New England, Long
Island, Pennsylvania, Virginia, Kentucky, Illinois, Missouri, and Nebraska. This species is
currently listed as a candidate for listing as threatened or endangered by the Virginia
Department of Conservation and Recreation (1993).

HABITAT IN VIRGINIA. Mountain ponds.

DISTINGUISHING FEATURES. Sch. torreyi can be distinguished by its 1-4, sessile, pale
brown to golden brown, lustrous spikelets which are cuneate at base; sheaths with scarious
inner band, which at maturity disintegrates, the prominent mid and laterally branching veins
remaining; and elongate, triangular-channeled leaf blades.

05. Schoenoplectus etuberculatus (Steud.) Sojak, Cas. Nar. Muz. Odd. Prir. 140(3-4): 127.
1972. Fig. 4c.
Rhynchospora etuberculata Steud., Syn. Pl. Glumac. 2: 142. 1855.
Scirpus etuberculatus (Steud.) Kuntze, Revis. Gen. Pl. 2: 758. 1891.

DESCRIPTION. Colonial perennial. Rhizomes horizontal, elongate, reddish, 1-3 mm
thick. Culms 1-2 m tall, trigonous, soft, easily compressed. Leaves 1-3, confined to basal
portion of culm; sheaths with narrow, scarious margins; blades 1-3, 2-12 mm wide, to 8 dm
long, septate, triangular-channeled or ribbon-like in submersed blades, excentric at tip.
Inflorescence pseudolateral, anthelate, with 4-16 spikelets; involucral bracts 1-3, the lowest
appearing as a continuation of the culm, triangular, 1-2.5 dm long, excentric at tip; rays 5-9,
somewhat flattened to obtusely trigonous, glabrous, scabrous along margins; spikelets
narrowly ovoid to cylindric, cuneate at base, 1-2.5 cm long, 4-8 mm wide; scales
oblong-ovate, acute to cuspidate, sagittate at base, 4.5-6 mm long, 2.2-3 mm wide, smooth,
green to pale brown, the green midrib ending in a short cusp at apex; stamens 3; anthers
narrowly oblong to linear, 2.5-3 mm long, tipped by a triangular-ovate appendage; style
somewhat flattened, 3-branched, the branches glabrous, with minute scales. Achene
plano-convex, obovate, 3-4 mm long, 1.9-2.2 mm wide, with a subulate beak 0.8-1 mm long,
smooth and shining, brownish; bristles 6, reddish, with antrorse to divergent barbs, unequal,
slightly shorter than to equaling or exceeding the achene.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 57

FLOWERING AND FRUITING. Late July-Aug.; Aug.-Sept.

DISTRIBUTION. Historically known from only one locality in Virginia: Princess Anne
County, shallow water, NW branch of Salt Pond, L.F. Randolph & F.R. Randolph 462, 29
June 1922 (GH); Florida to Louisiana, north to Delaware and Missouri, disjunct to Rhode
Island. This species is currently listed as S1 (extremely rare in the state) by the Virginia
Department of Conservation and Recreation (1993).

DISTINGUISHING FEATURES. Sch. etuberculatus can be distinguished by its
triangular-channeled, emergent leaf blades, and involucral bracts with excentric apex; flaccid,
ribbon-like submersed leaf blades; and open, anthelate, pseudolateral inflorescence of 4-16
large (1-2.5 cm long, 4-8 mm wide) spikelets.

06. Schoenoplectus americanus (Pers.) Volkart ex Schinz & Keller, Fl. Schweiz Ed. 2: 75.
1905. Figs. 3c, 8c and d.
Scirpus americanus Pers. Syn. Pl. 1: 68. 1805.
Scirpus Olneyi A. Gray ex Engelm. & A. Gray, Boston J. Nat. Hist. 5: 238. 1845.
Schoenoplectus olneyi (A. Gray) Palla, Bot. Jahrb. Syst. 10: 299. 1888.

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate, 3.5-5 mm thick,
reddish. Culms 0.3-2 m tall, 4-10 mm wide, sharply trigonous, wing-angled, with deeply
concave sides, red to red-tinged near base. Leaves 1-2, blade-bearing or the lower bladeless,
confined to basal portion of culm; sheaths septate, with a short ligule ca. 1 mm long at
adaxial junction of sheath and blade, the inner band firm with a U- or V-shaped notch at
orifice; blades septate, 1-15 mm long, 1-4 mm wide. Inflorescence pseudolateral, with a
single glomerate cluster of (3-) 5-12 spikelets; involucral bract 1, lance-triangular, obtuse
to subacute, 1-2 (-3.5) cm long, appearing as a continuation of the culm, glabrous; spikelets
ovoid, narrowly ovoid or ellipsoid, obtuse to subacute at apex, 5-10 (-20) mm long, 4-7 mm
wide; scales broadly ovate to orbicular, 2.5-3.2 mm long, 2-2.5 mm wide, light-brown to
reddish brown, scarious-margined, the green to yellowish midrib prolonged as a short awn,
shorter than to equaling the shallowly emarginate apex; stamens 3; anthers narrowly oblong,
2-2.5 mm long, with very short, blunt, minutely bearded appendage at tip; style straplike,
2- (sometimes 3-) branched, the branches glabrous, minutely scaly. Achene plano-convex,
suborbicular to broadly obovate, 2-2.5 mm long, 1.6-2.2 mm wide, smooth, brownish to
grayish brown; bristles 1-6, reddish, with downcurved-retrorse barbs, shorter than to
equaling the achene.

FLOWERING AND FRUITING. Late May-June; late June-Sept.

DISTRIBUTION. Restricted to the coastal plain of Virginia (see Harvill et al. 1992);
Florida to Mexico north to Nova Scotia, northern Ohio, southern Michigan, western Nova
Scotia, Missouri, Idaho, and Oregon; West Indies, Central America & South America.

HABITAT IN VIRGINIA. Brackish to saline marshes.

DISTINGUISHING FEATURES. This species is closely related to Sch. pungens but can
be distinguished from it by its stout, sharply trigonous, wing-angled culms with deeply
concave sides (4-10 mm wide); short involucral bract (1-2 (-3.5) cm long); shorter leaf
blades (1-15 mm long); scales with awns shorter than to equaling the shallowly cleft apex;
Shorter achenes (2-2.7 mm long); and broader, strap-like bristles with downcurved teeth.

Schoenoplectus pungens differs in having narrower culms (1-5 mm wide); longer
involucral bracts (2-15 cm long); longer leaf blades (0.5-4 dm long); scales with awns

58 BARTONIA

FIG. 8. a and b, Schoenoplectus pungens. (a) habit. (b) spikelet scale. c and d, Schoenoplectus americanus.
(c) spikelet scale. (d) habit.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 59

exceeding the deeply cleft, emarginate apex; longer achenes (2.5-3.0 (-3.2) mm long); and
narrower hypogynous bristles with retrorse teeth.

NOMENCLATURE. This species has been treated in many North American floras as
Scirpus olneyi Gray, e.g., Small (1933), Beetle (1947), Fernald (1950), Gleason (1952),
Gleason and Cronquist (1963), and Radford et al. (1968). Schuyler (1974), found that the
type specimen of Scirpus americanus Persoon was conspecific with plants usually treated as
Scirpus olneyi and the type specimen of Scirpus pungens Vahl was conspecific with plants
usually treated as Scirpus americanus. Thus the correct name for plants previously treated
as Scirpus olneyi (Schoenoplectus olneyi) is Scirpus americanus (Schoenoplectus
americanus), and the correct name for plants previously treated as Scirpus americanus
(Schoenoplectus americanus) is Scirpus pungens (Schoenoplectus pungens).

DISCUSSION. In Virginia, Schoenoplectus americanus occurs only on the coastal plain
and is primarily restricted to tidal saline marshes in the lower Chesapeake Bay and outer
coastal plain, whereas Schoenoplectus pungens is more widespread and occurs in a variety
of fresh to saline habitats, most notably on wet or inundated sandy shores of inland and tidal
rivers, creeks, and beaches. Generally these two species are isolated ecologically, but
apparent hybrids occasionally occur in intermediate habitats where they are sometimes
abundant (Schuyler pers. comm., 1991). The resulting hybrids tend to be intermediate in
characters.

07. Schoenoplectus gtk (Vahl) Palla, Bot. Jahrb. Syst. 10: 299. 1888. Figs. 3d, 8a, b.
Scirpus pungens Vahl, Enum Pl. 2: 255. 1805 (see Schuyler, Rhodora 76: 51-52. 1974,
Scirpus americanus ae misapplied).

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate, stout, 4-10 mm thick.
Culms sharply trigonous, frequently with concave sides above, 0.3-1.5 m tall, 1-5 mm wide.
Leaves 2-4, confined to basal portion of culm; sheaths short ligulate at adaxial junction of
sheath and blade, septate, the inner band firm with a scarious, red-dotted orifice, concave at
apex; blades linear-elongate, sharp-pointed, 0.5-4 dm long, 2-9 mm wide, dorsally to
laterally compressed, becoming sub-trigonous near apex. Inflorescence pseudolateral, a
single glomerate cluster of 1-6 (-8) spikelets; involucral bract 1, acuminate, 2-15 cm long,
appearing as a continuation of the culm; spikelets ovoid to subcylindric, 0.5-2 cm long, 3-5
mm wide; scales ovate, 4-6 mm long, 2-2.5 mm wide, reddish-brown, the green midrib
prolonged into a short, scabrous awn which exceeds the deeply emarginate apex; stamens
3; anthers narrowly oblong to linear, ca. 3 mm long, with fringed appendage at tip; style
straplike, 2- (sometimes 3-) branched, the branches glabrous, minutely scaly. Achene
plano-convex, obovate, 2.5-3.0 (-3.2) mm long, 1.5-2 mm wide, smooth, olivaceous to
brown; bristles 3-6, retrorsely barbed, reddish, sometimes rudimentary, less than 3/4 the
length of the achene.

FLOWERING AND FRUITING. Mid May-June; June-Sept.

DISTRIBUTION. Scattered localities throughout the state, but more frequent on the
coastal plain (see Harvill et al. 1992); additional county not recorded by Harvill et al. 1992:
Loudoun Co., Smith 102 (US); Florida to Texas, north to Newfoundland, Quebec, southern
Ontario, Michigan, Wisconsin, Minnesota and Nebraska; South America, Europe, Australia,
and New Zealan

HABITAT IN VIRGINIA. Tidal shores along rivers, sandy river banks and shores, and
creeks.

60 BARTONIA

DISTINGUISHING FEATURES. See Sch. americanus.
NOMENCLATURE. See Sch. americanus.
DISCUSSION. See Sch. americanus.

08. Schoenoplectus validus (Vahl) A. & D. Love, Bull. Torrey Bot. Club 81: 33. 1954.
Figs. 3a, 9a-c.
Scirpus validus Vahl, Enum. Pl. 2: 268. 1805.
Scirpus validus Vahl var. creber Fern., Rhodora 45: 282. 1943.
Scirpus lacustris L. ssp. validus (Vahl) Koyama, Canad. J. Bot. 40: 927. 1962.
Schoenoplectus lacustris L. ssp. creber (Fern.) A. & D. Love, Taxon 30(4): 849. 1981.

DESCRIPTION. Robust perennial. Rhizomes stout, scaly, horizontal, elongate, 0.7-1.5 cm
thick, reddish. Culms terete, easily compressed, 0.5-3 m tall, 0.3-2 cm wide at base, forming
in a row along the rhizome. Leaves 3-5 bladeless sheaths, short mucronate or the uppermost
with short blades, confined to basal portion of culm; sheaths with a short, firm ligule, ca.
1.5 mm long, borne at adaxial junction of sheath and blade, the margins of the sheath
scarious above, gradually narrowing towards base to V-shaped orifice; blades 1-2 on upper
sheaths, 2-10 cm long, dorsally flattened, thickened towards summit with cartilaginous tip.
Inflorescence pseudolateral, anthelate, compound to partially decompound, pendulous, the
numerous spikelets solitary or in small glomerules of 2-3 (-5) at ray tips; involucral bracts
2-3, the lowest one erect, appearing as a continuation of the culm, the others membrana-
ceous, scale-like; rays flat, to somewhat obtusely angled, glabrous, scabrous on margins,
arching to pendulous; spikelets ovoid to ovoid-ellipsoid, acute, 5-7 (-9) mm long, 3-4 mm
wide; scales ovate to elliptic, 2.5-3.2 mm long, 1.8-2.2 mm wide, reddish-brown,
erose-ciliate, with scarious margins, the green midrib prolonged as a short, straight, scabrous
awn at the emarginate apex; stamens 3, marcescent; anthers narrowly oblong, 1-2 mm long,
tipped by a triangular-ovate appendage; style straplike, 2 or 3 branched, the branches
glabrous, with minute scales. Achene plano-convex, broadly ovate, 1.7-2.3 mm long, 1.3-1.5
mm wide, smooth, grayish to drak-brown; bristles 6, slender, stiff, retrorsely barbed,
red-brown, shorter than to equaling or slightly exceeding the achene.

FLOWERING AND FRUITING. Late May-June; June-Sept.

DISTRIBUTION. Widespread throughout the state (see Harvill et al. 1992); additional
counties not recorded by Harvill et al. 1992: Alleghany Co., Stevens 390 (FARM), Patrick
Co., Harvill & Harvill 29863 (FARM), and Northumberland Co., Ruth s.n. (US);
Newfoundland to central Alaska, south to Florida and California; Mexico, Central America,
West Indies, South America, Eurasia, Australia, New Zealand, and Polynesia.

HABITAT IN VIRGINIA. Tidal shores of rivers, marshy meadows, swales, pond borders,
and bottomland swamps.

DISTINGUISHING FEATURES. Sch. validus is closely related to Sch. acutus but can be
distinguished from the latter by its shorter spikelets (5-7 (-9) mm long) that are solitary or
in clusters of 2-3 at ray tips and shorter scales (2.5-3.2 mm long) with short, nearly straight
awns, whereas Sch. acutus has longer spikelets (6-15 mm long) which are in glomerules of
3-7 at ray tips and longer scales (3.5-4 mm long) with an elongate, usually contorted,
scabrous awn.

Smith (1969) found that these two species could be separated by the number and size of
internal air spaces in the upper 1/4 of the culm. For Sch. validus, there were 2 to 4 internal
air spaces in a cross section diameter, these averaging 1.0-2.5 mm wide, whereas for Sch.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 61

FIG. 9. a-c, Schoenoplectus validus. (a) rhizome and lower portion of culm. (b) inflorescence. (c) spikelet
scale. d and e, Schoenoplectus acutus. (d) inflorescence. (e) spikelet scale.

62 BARTONIA

acutus, there were 9-14 spaces averaging 0.3-0.5 mm wide.

DISCUSSION. Hybridization between these two species in northern Michigan, Wisconsin,
Iowa, and North Dakota primarily occurs in areas of human or natural disturbance. Those
populations in undisturbed sites have different ecological tolerance ranges and as a result of
this physical isolation, hybridization does not occur. However, where tolerance ranges
overlap and mixed populations occur, particularly in parts of western North America, the
species may be isolated by other factors or extensive hybridization may obscure the
morphological boundaries between the species (Smith, 1969).

09. Schoenoplectus acutus (Muhl. ex Bigelow) A. & D. Léve, Bull. Torrey Bot. Club, 81:
33. 1954. Figs. 3b, 9d and e.
Scirpus acutus Muhl. ex Bigelow, Fl. Boston 15. 1814.
Schoenoplectus lacustris ssp. acutus (Muhl. ex Bigelow) A. & D. Love, Taxon 30(4): 849.
1981

DESCRIPTION. Robust perennial. Rhizomes stout, scaly, horizontal, elongate, 0.5-1.5 cm
thick, pale brown to brown. Culms firm, terete, 1-3 m tall, to 2 cm wide at base. Leaves 3-5
short-mucronate, bladeless sheaths or the uppermost with short blades, confined to the basal
portion of the culm; sheaths lustrous brown near base, septate, with a short, scarious ligule
borne at adaxial junction of sheath and blade, the ventral band of the uppermost leaves
scarious, veiny, with divergently branching midvein, forming a truncate to concave orifice
at apex; blades 1-2 on upper sheaths, 2-15 cm long. Inflorescence pseudolateral, anthelate,
the spikelets rarely solitary, usually in glomerules of 3-7 at ray tips; involucral bracts 2-3,
the lowest erect, appearing as a continuation or the culm, the others membranous, scale-like;
rays flat to somewhat obtusely angled, glabrous, scabrous on margins, divergent to
subpendulous; spikelets oblong-ovoid, 6-15 mm long, 2-5 mm wide, acute; scales 3.5-4 mm
long, 2 mm wide, brown to reddish-brown, erose-ciliate on margins, the green midrib often
prolonged into a contorted, scabrous awn at apex; stamens 3; anthers narrowly oblong, 1-2
mm long, tipped by a triangular-ovate appendage; style straplike, 2 or 3 branched, the
branches glabrous, minutely scaly. Achene plano-convex, ovate to ovate-elliptic, 2-2.8 mm
long, 1.5-2 mm wide, smooth, pale brown to grayish-brown or deep brown; bristles 2-6,
slender, stiff, retrorsely barbed, reddish, shorter than to equaling the achene.

FLOWERING AND FRUITING. June to early Aug.; Aug.-Oct.

DISTRIBUTION. Known only from Augusta and Arlington counties in Virginia;
Newfoundland to British Columbia, south to Nova Scotia, New England, northern New
Jersey, Virginia, North Carolina, Pennsylvania, Ohio, Illinois, Tennessee, Missouri,
Oklahoma, Texas, New Mexico, Arizona, and California. This species is listed as S2 (very
rare in the state) by the Virginia Department of Conservation and Recreation (1993).

HABITAT IN VIRGINIA. Calcareous wet meadows in the mountains and fresh to
brackish tidal marshes on the coastal plain

DISTINGUISHING FEATURES. See Sch. validus.

DISCUSSION. See Sch. validus.

10. Schoenoplectus fluviatilis (Torrey) M. T. Strong, Novon 3(2): 203. 1993. Figs. 4d, 10c.
Scirpus maritimus L. B ? fluviatilis Torrey, Ann. Lyceum Nat. Hist. New York 3: 324.

Scirpus fluviatilis (Torrey) A. Gray, Manual Bot. N. United States: 527. 1848.

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 63

Bolboschoenus fluviatilis (Torrey) Sojak, Cas. Nar. Muz. Odd. P¥ir. 141: 62. 1972.

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate, 4-7 mm wide. Culms
sharply trigonous, 0.7-2 m tall, to 1.5 cm wide at base, arising from hard, swollen, tuberous
enlargements to 2 cm thick. Leaves 5-9, nodes extending up culm to above middle; sheaths
firm, faintly septate, eligulate, the orifice with a U-shaped, rounded or truncate contraligule
with short-scarious apex; blades 7-20 mm wide, to 8 dm long, smooth to weakly scabrous
on margins and midvein beneath. Inflorescence a spreading, compound, umbel-like corymb,
anthelate, the lowest leaf-like bract erect at anthesis, the inflorescence appearing pseudo-
lateral, at maturity spreading and appearing terminal; involucral bracts 5-6, elongate,
leaf-like, much exceeding the inflorescence; rays 5-12, elongate, up to 12 cm long, glabrous,
somewhat flattened to obtusely angled or subterete, spreading, scabrous on margins, recurved
at maturity, the spikelets solitary or in glomerules of 2-5 at ray tips; spikelets oblong-ovoid
to ellipsoid, cuneate at base, 1.5-2.5 (-4) cm long, 0.7-1 cm wide; scales oblong-ovate to
elliptic, 0.7-1 cm long (excluding awn), 3.5-6 mm wide, pale brown, puberulent, with erose
to slightly lacerate margins, the pale green midrib prolonged as a recurved awn 2-4 mm long,
exceeding the notched apex; stamens 3; anthers narrowly oblong to linear, 3-6 mm long,
with an antrorsely barbed, linear appendaged tip; style angled to somewhat flattened,
3-branched, the branches glabrous, minutely scaly. Achene trigonous, pyriform to obovoid,
4-5 mm long, 2.2-2.5 mm wide, prominently beaked, pale brown; bristles 6, persistent,
retrorsely barbed, about equaling or slightly exceeding the achene.

FLOWERING AND FRUITING. June to early July; July-Aug.

DISTRIBUTION. Occurs in Fairfax, Giles, and King George counties in Virginia; Western
New Brunswick to Saskatchewan and Washington, south to New England, Delaware,
Maryland, Virginia, Pennsylvania, Ohio, Indiana, Illinois, Missouri, Kansas, New Mexico,
and California; Asia, Australia, New Zealand, and Japan. This species is listed as S1
(extremely rare in the state) by the Virginia Department of Conservation and Recreation
(1993).

HABITAT IN VIRGINIA. Fresh water tidal shores and marshes of large river systems on
the coastal plain, occurrring infrequently along shorelines of inland rivers.

DISTINGUISHING FEATURES. Schoenoplectus fluviatilis can be distinguished from the
related Sch. robustus and Sch. novae-angliae by its sharply trigonous, pyriform to obovoid
achenes which taper at the apex to a prominent beak; 6 persistent bristles that equal or
slightly exceed the achene; open, spreading inflorescence with elongate primary rays; cuneate
base of the spikelets; and pale brown scales. Schoenoplectus robustus has lenticular or
plano-convex, obovate, smooth, lustrous, blackish brown achenes which are truncate at apex,
abruptly differentiated into a short beak; caducous bristles; congested inflorescence with
short primary rays; and ovoid spikelets with truncate base, 1.3-3 cm long, or in forma
protrusus Fernald, 3-6 cm long. Schoenoplectus novae-angliae differs in having plano-con-
vex, obovate, whitish or olive brown achenes which are rounded at the summit, gradually
tapering into a prominent beak; 1-6 persistent bristles which are shorter than to equaling the
achene; and elongate spikelets ((1.5-) 2-6 cm long).

11. Schoenoplectus robustus (Pursh) M. T. Strong, Novon 3(2): 203. 1993. Figs. 4a, 10a.

Scirpus robustus Pursh, Fl. Amer. Sept. 1: 56. 1816. (based on Scirpus maritimus 8

macrostachyos Michaux, Fl. Amer. 1: 32. 1803, non Scirpus macrostachyos Lam., Tabl.
Encycl. 1: 142. 1791)

64 BARTONIA
Scirpus maritimus var. robustus (Pursh) Kiik., Repert. Spec. Nov. Regni. Veg. 23: 200.

26.
Bolboschoenus robustus (Pursh) Sojak, Cas. Nar. Muz. Odd. Prir. 141: 63. 1972.

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate, 3-6 mm thick. Culms
sharply trigonous, 0.7-1.5 m tall, 5-9 mm wide near base, arising from hard, swollen,
tuberous enlargements. Leaves 4-9, the nodes extending above mid portion of culm; sheaths
septate, eligulate, the orifice with a short, subscarious, acute to rounded or on lower sheaths,
convex contraligule, prominently veined below; blades 4-12 mm wide, up to 7 dm long,
smooth to weakly scabrous on margins. Inflorescence a congested, compound, umbel-like
corymb, anthelate, the lowest leaf-like bract erect at anthesis, the inflorescence appearing
pseudolateral, at maturity spreading and appearing terminal; involucral bracts 3-5, to 4 dm
long, leaf-like; rays short, terete to obtusely angled, smooth, the spikelets in a basal mass,
solitary, or 2-5 at ray tips; spikelets (1-) 3-11 (-20), ovoid to narrowly ovoid, or rarely
cylindrical, blunt to rounded at tip, truncate at base, 1.3-3 (-6) cm long, 0.8-1.2 cm wide;
scales oblong-ovate to elliptic, 5-7 mm long (excluding awn), 3-4 mm wide, reddish-brown
to rufescent, puberulent, the narrow, pale brown midrib prolonged as a recurved awn,
exceeding the notched apex, 1-3 mm long; stamens 3; anthers narrowly oblong to linear,
2-3.5 mm long, with acute, antrorsely barbed appendage at tip; style straplike, 3-branched,
the branches glabrous. Achene lenticular or plano-convex, obovate, truncate at apex, abruptly
short-beaked, 3-3.5 mm long, 2-3 mm wide, reddish brown to blackish brown at maturity,
lustrous; bristles none or caducous, or occasionally 1-6 persist at base of achene, retrorsely
barbed, 1/3 to 3/4 the length of the achene.

FLOWERING AND FRUITING. Late May-June; late June-Sept.

DISTRIBUTION. Primarily restricted to the coastal plain in Virginia (see Harvill et al.
1992); Florida to eastern Mexico, northern Georgia, South Carolina, North Carolina,
Virginia, Maryland, New Jersey, Pennsylvania, Long Island, New York, and Massachusetts,
ss to California.

ABITAT IN VIRGINIA. Brackish to saline tidal marshes, salt springs.

DE NGRISHING FEATURES. see Sch. fluviatilis.

12. Schoenoplectus novae-angliae (Britt.) M. T. Strong, Novon 3(2): 203. 1993. Figs. 4b,
10b.

Scirpus novae-angliae Britt. Ill. Fl. N. U.S. 3: 509. 1898.

Scirpus maritimus y cylindricus Torr., Ann. Lyceum Nat. Hist. New York 3: 325. 1836.

Scirpus cylindricus (Torr.) Brit., nom. illeg. Trans. N.Y. Acad. Sci. 11: 79. 1892. non
Scirpus cylindricus (Vahl) Lam., Encyl. Method. Vol. 5 (suppl.): 101. 1817.

Scirpus campestris var. novae-angliae (Britt.) Fern., Rhodora 8: 136. 1906.

Scirpus robustus var. novae-angliae (Britt.) Beetle, Amer. J. Bot. 29: 82. 1942.

Scirpus subterminalis var. cylindricus (Torr.) Koyama, Canad. J. Bot. 40: 930. 1962.

DESCRIPTION. Robust perennial. Rhizomes horizontal, elongate. Culms 0.7-1.5 m tall,
to 9 mm wide at base, smooth, arising from hard, tuberous enlargements. Leaves 4-7, nodes
extending above mid portion of culm; sheaths glabrous, eligulate, the orifice with a short,
scarious, convex to truncate or concave-tipped contraligule; blades 6-15 mm wide, to 7 dm
long, smooth to weakly scabrous on margins and midvein beneath. Inflorescence a congested
to semi-open, compound, umbel-like corymb, anthelate, the lowest leaf-like bract erect at
anthesis, the inflorescence appearing pseudolateral, at maturity spreading and appearing

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 65

Mart. T sing 9

A ace
FIG. 10. (a) Sch. robustus (habit). (b) Sch. novae-angliae (inflorescence). (c) Sch. fluviatilis (inflorescence).

66 BARTONIA

terminal; involucral bracts 3-5, leaf-like, up to 3 dm long; rays short, or few elongate,
smooth, terete to somewhat flattened or obtusely angled, the spikelets solitary or 2-5 at ray
tips; spikelets 15-40, narrowly oblong, elongate, (1.5-) 2-5 cm long, 5-8 mm wide; scales
7-8 mm long, 3.5-5 mm wide, pale brown to reddish brown, puberulent, the narrow, pale
brown midrib prolonged as a recurved awn, 1-4 mm long, exceeding the notched apex;
stamens 3; anthers narrowly oblong to linear, 3-3.5 mm long, with acute, antrorsely barbed
appendage at tip; style straplike, 3-branched, the branches scaly with ascending to divergent
hairs. Achene plano-convex, obovate, rounded at summit, gradually tapering into a beak,
3.5-4 mm long, 2.7-3.3 mm wide, whitish to olive brown; bristles 1-6, persistent, retrorsely
barbed to base, shorter than to equaling the achene.

FLOWERING AND FRUITING. Late June-July; July-Sept.

DISTRIBUTION. Occurs in Charles City, James City, New Kent, Prince William,
Stafford, and Surry counties on the coastal plain of Virginia; coastal plain of eastern North
America from Maine to Georgia.

HABITAT IN VIRGINIA. Fresh to brackish tidal marshes.

DISTINGUISHING FEATURES. See Sch. fluviatilis.

NOMENCLATURE. In North American floras, Sch. novae-angliae has gone under a
variety of names. Beetle (1947) treated it as Scirpus robustus var. novae-angliae. Fernald
(1943) considered it to be a variety of typical European Scirpus maritimus. He cited Scirpus
novae-angliae as a synonym, but this synonym was misapplied, the type of his variety being
conspecific with Scirpus paludosus Nels. (Schuyler 1975a). Schuyler (1975a) found S.
novae-angliae to be a distinct North American species citing both morphological as well as
ecological evidence (see discussion below), but the name he applied to it (S. cylindricus
(Torr.) Britt., (1898)) is a later homonym of S. cylindricus (Vahl) Lam. (1817). The earliest
name for this plant that can be retained is S. novae-angliae Britton.

DISCUSSION. Schoenoplectus novae-angliae is isolated ecologically from other species
of tuberous bulrushes in North America. It is restricted to brackish tidal marshes and shores
along the Atlantic coast. It is primarily found in the brackish transition zone between the
upper fresh waters (where Sch. fluviatilis occurs) and the lower saline waters (where Sch.
robustus usually occurs) of tidal river systems. This upstream-downstream zonation pattern
has been observed by Schuyler (1975a) on the Delaware River in Delaware, Pennsylvania,
and New Jersey, and on the Kennebec River in Maine, and by me on the Potomac River in
Virginia.

EXCLUDED SPECIES

Lipocarpha rehmanii (Ridley) Goetghebeur (Scirpus rehmanii Ridley). This, a species of
tropical Africa, ranges from Kenya to South Africa and Madagascar (Goetghebeur and
Vanden Borre 1989). Reed (1964) found this plant growing in wet areas on chrome ore piles
in Newport News (Reed 44058, 7 Aug. 1959 (US)). Apparently, the ore piles are now gone
(Reed pers. comm. 1991). Since no recent collections have been made in the state, these
plants are doubtfully persisting elsewhere, and at present cannot be considered an established
element in Virginia’s flora.

ACKNOWLEDGMENTS

I thank the following people for providing specimens and assistance regarding this study:
Ted R. Bradley (GMUF), Alton M. Harvill (FARM), J. Christopher Ludwig (Virginia

VIRGINIA SCIRPUS, TRICHOPHORUM, AND SCHOENOPLECTUS 67

Department of Conservation and Recreation), Dan H. Nicolson (US), S. Galen Smith
(University of Wisconsin, Whitewater), G.C. Tucker (NYS), Donna M.E. Ware (WILLI), and
Thomas F. Wieboldt (VPI). Special appreciation goes to Alfred E. Schuyler (PH) for all of
his assistance throughout this study, particularly in regards to availability of specimens and
manuscript review and preparation, and Carol L. Kelloff for taking the SEM photomicro-
graphs, assisting with manuscript preparation, and her patience. Further thanks are extended
to Dorothy Strong and the late Benjamin Strong for their support during this study.

LITERATURE CITED

BEETLE, A. A. 1942. Studies in the genus Scirpus L., V. Notes on the section Actaeogeton. Amer. J. Bot. 29:
653-656.

———. 1947. Cyperaceae: Scirpeae. Scirpus. North American Flora 18: 479-504.

CaPPEL, E. D. 1954. The genus Scirpus in North Carolina. J. Elisha Mitchell Sci. Soc. 70: 85-91.

FERNALD, M. L. 1943. Contributions from the Gray Herbarium of Harvard University. No. CXLVIII. V. Studies
on North American species of Scirpus. Rhodora 45: 279-296

1950. pe Manual of Botany. 8th ed. American Book Co., New York.

GLEASON, H. A. 1952. The new Britton and Brown illustrated flora of the northeastern United States. New York

GLEASON, H. A. AND CRONQUIST A. 1963. Manual of the vascular plants of northeastern United States and
adjacent Canada. Van Nostrand, seaineis NJ:

GOETGHEBEUR, P. 1986. Genera C m. Een bijdrage tot de kennis van de mag ig systematiek en
seis van de revue valance ottiy power sini Dr: - = ae Sinversey,

———. 1989. Studies in Cyperaceae 8. A revision ip
Agric. Univ. Papers ve

d Rikliella Wageningen

GOETGHEBEUR, P. & D A. SIMPSON. 1991. Critical ‘i 7 Actinoscirpus, Bolboschoenus, Isolepis, Phylloscirpus
and ae gen (Cyperaceae. Kew Bull. 46(1): 169-178.
HARRI In: Chromosome number toa LXXI. Taxon 30: 517.

eae nm oy T. ‘e ek C. E. STEVENS, T. F. WIEBOLDT, D. M. E. WARE, D. W. OGLE, G. W. RAMSEY,
& G. P. FLEMING. 1992. Atlas of the Virginia Flora III. Vicehde Botanical Associates, Farmville, Virginia.
HICKs, G.C. 1928. Chromosome studies in the Cyperaceae, with special reference to Scirpus. Bot. Gaz. 86:
295-317.
KOYAMA, T. 1985. romney Pages 125-405 in M. D. Dassanayake and F. R. Fosberg, eds. A revised handbook
of the Flora of Ceylon, vol. 5. Balkema, Rotterdam
90. ecient Pass 1381-1436 in W. L. Wagner, D. - | and S. H. Sohmer. Manual of the
flowering plants of Haw Vol. 2. Univ. of Hawai’i Press, ‘Hon
OVE, A. & D. LOVE. a. Sart number reports LXXIII. panel 30(4): 829-861.
OTENG-YEBOAH, A. A. 1974. Taxonomic studies in Cyperaceae-Cyperoideae. Notes Roy. Bot. Gard. Edinburgh
6

RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the vascular flora of the Carolinas. University
of North Carolina Press, Chapel Hill.

RAYNAL, J. 1973. ge ne nerre 19. Contribution a la classification de la sous-famille des Cyperoideae.
Adansonia 13: 145-1

REED, C. F. 1964. A es of the chrome and maganese ore piles at Canton, in the Port of Baltimore, Marylan and
and at Newport News, Virginia, with descriptions of genera and species new to the flora of the eastern United
States. Phytologia 10(5): 321-406.

SALMENKALLIO, M. & I. KUKKONEN. 1989. (930) Proposal to conserve 466a Trichophorum (Cyperaceae) with a
conserved type. Taxon:38(2): 313-316.

SCHUYLER, A. E, 1962. A new species of Scirpus in the Northeastern United States. ns 64: 43-49.

——. 1963. Notes on five species of Scirpus in eastern North America. Bartonia

——. i The taxonomic delineation of Scirpus lineatus and Scirpus pendulus. Not. Nat 390:

—.. . A taxonomic revision of the North American leafy species of Scirpus. Proc. pie *Nat Sci.
Piaget 119: 295-323.

———._ 1967b. Scirpus hattorianus in North America. Not. Nat. 398: 1-5.

———. 1967c. A new status for an eastern North American Scirpus. Rhodora 69: 198-202.

68 BARTONIA

. 1971a. Some relationships in Scirpeae bearing on the delineation of genera. Mitt. Bot. Staatssaml. Miinchen
10: 577-585.

. 1971b. Scanning electron microscopy of achene epidermis in species of Scirpus (Cyperaceae). Proc. Acad.
Nat. Sci. Philadelphia 123: 29-52.
. 1972a. Chromosome numbers of Scirpus purshianus and S. smithii. Rhodora 74: 398-406.
. 1972b. Morphological ia anatomical differences in leaf blades of three North American aquatic bulrushes
(Cypecacese: Scirpus). Bartonia 41: 5
——. 1974. neice and eB of the names Scirpus americanus Pers., S. olneyi Gray, and S. pungens
6: 805.

Vahl. Rhodor:
be Sa. pleat cylindricus: An ecologically restricted Eastern North American tuberous bulrush. Bartonia

’ th Chromosome numbers of some eastern North American species of Scirpus. Bartonia 44: "eas

SMALL, J. K. 1933. Manual of the southeastern flora. The University of North Carolina Press, Chapel H

SMITH, S. G. 1969. Natural hybridization in the Scirpus lacustris ie in north central United ra —_
175-200 in J. G. Gunckel, ed. Current topics in plant science. New York.

STRONG, M. T. 1991. aariae! of the genera Scirpus sensu stricto, Tchaphonun, and Schoenoplectus sensu lato
sce ges sin in Virginia. Unpublished M.S. Thesis, George Mason Signs — Virginia.

1993. w Combinations in Schoenoplectus (Cyperaceae). Novon 3(2): 2

TUCKER, G. C. pe Scirpus (Cyperaceae) in Connecticut. New York State sk pee N.Y. (unpublished
manuscript).

UNITED STATES FISH AND WILDLIFE SERVICE. cake an and threatened wildlife and plants; determination
of endangered status for Scirpus ancistrochaetus (northeastern bulrush). Fed. Register 56: 21091-21096.

VAN DER VEKEN, P. 1965. Contribution a 1’embryographie i a des Cyperaceae-Cyperoideae. Bull. Jard.
Bot. Etat 35: 285-354.

VIRGINIA DEPARTMENT OF CONSERVATION AND RECREATION. 1993. Natural heritage resources of Virginia: Rare
vascular plant species. Virginia Department of Conservation and Recreation, Richmond, Virginia.

WILSON, K. L. 1981. A synopsis of the genus Scirpus sensu lato (Cyperaceae) in Australia. Telopea 2: 153-172.

Bartonia No. 58: 69-73, 1994
Noteworthy Plant Collections from Pennsylvania

ALLISON W. CUSICK
Division of Natural Areas and Preserves, Ohio Department of Natural Resources
Fountain Square, Columbus, OH 43224

SUE A. THOMPSON
Section of Botany, Carnegie Museum of Natural History
4400 Forbes Avenue, Pittsburgh, PA 15213

New Pennsylvania county records not mapped in the Atlas of the Flora of Pennsylvania
(Wherry et al. 1979) were reported in Thompson et al. (1989) for over 1200 plant taxa.
Recent collecting by both authors as well as other additions to the Carnegie Museum of
Natural History herbarium (CM) have yielded further new distribution records and taxa
previously unreported in Pennsylvania. This paper cites specimens for 149 new county
records for 110 taxa in 45 plant families. Three taxa are reported as new to the Pennsylvania
flora: Dyssodia papposa (Asteraceae), Spiranthes casei (Orchidaceae), and Vitis cinerea var.
baileyana (Vitaceae).

This paper also includes specimen data for six species not listed in recent works as
occurring in Pennsylvania, but which were included in John Miller’s (1923) "Flora of Erie
County, Pennsylvania." Miller’s herbarium was donated to the Carnegie Museum of Natural
History in 1988 by the Erie Historical Society and Planetarium. Previous to that, these
specimens and Miller’s publication had been generally overlooked. Four of these species
are apparent waifs, which have not been recollected: Tidestromia lanuginosa (Amarantha-
ceae), Cirsium undulatum (Asteraceae), Ratibida columnifera (Asteraceae), and Jacquemontia
tamnifolia (Convolvulaceae). The other two species, Aster borealis (Asteraceae) and
Gentianopsis procera (Gentianaceae), are apparently native to Pennsylvania.

Three additional taxa have now been collected in all Pennsylvania counties bringing that
total to 52 taxa (see previous list in Thompson et al. 1989): Juncus tenuis (Juncaceae),
Rumex obtusifolius (Polygonaceae), and Fragaria virginiana (Rosaceae).

The following list is arranged alphabetically by family and taxon with nomenclature
generally following that of Wherry et al. (1979). Voucher specimens are deposited at CM.

ACANTHACEAE: Ruellia strepens—GREENE: Wind Ridge, Davis & Davis 5405 (Jun 1942), along roadside
on bank.

ACERACEAE: Acer platanoides—PERRY: NW of Dromgold on PA 850, eg cog (Sep 1988), along
Sherman Creek; SOMERSET: Somerset, Cusick & Shelton 28939 (Jun 1990), weedy

AMARANTHACEAE: Tidestromia lanuginosa—ERIE: Car Works Yard, Kuhn s.n. "Oak oe from the Miller
Herbarium.

ARACEAE: orus calamus—ADAMS: 2.2 mi WNW of Gettysburg on US 30, Thompson & York 6435 (Jun
1989), roadside a FULTON: 1 mi W of Harrisonville on US 30, Thompson & York 6438 (Jun 1989), ditch
through open field. Arisaema dracontium—LAWRENCE: ca 0.75 mi S of Pulaski, /saac & Isaac 3092 (May 1990),
floodplain woods.

ARISTOLOCHIACEAE: 4ristolochia macrophylla (A. durior}—FAYETTE: ca 0.2 mi N of New Geneva, /saac

3677 (Sep 1991), along old road grade in forest.

ASCLEPIADACEAE: Cynanchum nigrum—CUMBERLAND: 8 km S of Carlisle, Zanol 75 (Jul 1990).

69

70 BARTONIA

ASPLENIACEAE: sonnei montanum—INDIANA: ca 1.5 mi SE of Boltz, Isaac & Isaac 3088 (May 1990),
on sandstone outcroppin
ASTERACEAE: pare vulgaris—1.8 mi E of ne Isaac 3777 (Sep 1991), roadside. ipa borealis (=
A. = pala: listed in Miller (1923) as A. junceus)—ERI amarack Swamp, Miller s.n. (Aug 1903); apparently
first PA record, from the Miller Herbarium, mene Erie cou populations recently found at oF Pleasant and
Eanboo ees fen by James Bissell of the eland Museum of Natural History (pers. comm.). Aster firmus (=
. lucidulus)—FRANKLIN: Michaux State Forest, NW a Old Forge, Cusick pone (Sep 1989), streambank.
matic undulatum—ERIE: E&P Dock, Mie s.n. (Jul 1911); only known PA record, from the Miller Herbarium.
Dyssodia papposa—FRANKLIN: SW of Newbridge on Pennsylvania Turnpike, Cusick 27850 (Sep 1988), road
rm; first confirmed PA report, Strother (1969) includes PA in generalized distribution map but cites no specimens,
well established along interstate highways in Ohio. Eupatorium rotundifolium—FRANKLIN: 1.5 km NE of Pond
Bank, Thompson 8004 (Sep 1990), pine woodland. Hypochoeris radicata—FOREST: ca | mi S of Tionesta, /saac
3794 (Oct 1991), along highway. Lapsana communis—LAWRENCE: Rose Point, Isaac & Isaac 3512 (Jul 1991).
Matricaria matricarioides—GREENE: ca 5 km E of Aleppo, Thompson & Walck 71 7 (May 1990), ane road.
Ratibida pikes opceaay Elevator a iad s.n. (Jul 1908); listed in Miller (1923) as Lepachys columnaris.
Solidago arguta ssp. arguta—JUNIATA a Mountain, NW of Ickesburg, Cusick 27821 (Sep 1988), f ocky
ground. Solidago purshit_FAYETTE: Ses mi “ or Farmi iain , Thompson 1889 (Aug 1984), Markleysburg Bog.
Solidago be a—CLINTON: 4.2 km NNE of Loganton, Bier s.n. (Sep 1989), sandy road bank; reap NW of
Dromgold on PA 850, Cusick 27831 (Sep 1988), along Sherman Creek. Solidago squarrosa—PERRY: SE of New
Ponatticit on Be Hill Road, Cusick 27829 (Sep 1988), road bank. Tussilago farfara em aise near
Hunters Run at edge of Michaux State Forest, Isaac & Isaac 3338 (Mar 1991), roadside gravel.
BETULACEAE: Alnus glutinosca—SOMERSET: Barronvale, Cusick & Shelton 29789 (Aug 1991), along Laurel
Hill Creek; apparently first report of this taxon from western
BRASSICACEAE: Alliaria petiolata—LAWRENCE: ca 0. 25, mi SE of New sna Isaac & Isaac 3093 (May
1990), along edge of building. Brassica rapa—CUMBERLAND: 8 km S of Carlisle, Zano/ 51 (Jul 1990).
Cardamine hirsuia—ARMESTRONG. 3 mi SSW of Shay, Thompson & anaes 3633 (Apr 1987), along road.
Cardamine impatiens—CENTRE: SW of Bellefonte, ai 28407 (Jul 1989), along Spring Creek Road; CL «came mee
just SE of East Brady, Reznicek & Romme 8129 (May 1988), disturbed open hardwoods, scarce (specim
herbarium of University of Michigan); LAWRENCE: atin Isaac & Isaac 3352 (Mar 1991), along seb
tracks. Thlaspi arvense—FULTON: | mi W of Harrisonville, Thompson & York 6437 (Jun 1989), ditch through
open field.
CAPRIFOLIACEAE: Lonicera maackii—-WESTMORELAND: SE of New Stanton on PA Turnpike, are
war fh 1990), weedy thickets. Lonicera morrowi—LAWRENCE: Rose Point, Isaac & Isaac 3410 (May 1991).
OPHYLLACEAE: Cerastium nutans—CAMERON: 7 mi NW of Driftwood, Buker s.n. (May 1966), rocky
Pcie inilniers identified as C. viscosum. Cerastium semidecandrum—ERIE: Presque Isle State Park, Grisez
964 (May 1986), edge of beach and woods in sand; first record from western PA, previously reported in the area
of Philadelphia and Reading (R. Rabeler pers. comm.). Scle felon ee PA Furnace Road
at summit of Tussey Mountain, Cusick 28422 (Jul 1989), road berm; 6 km S of Huntingdon, Thompson & Rawlins
coe — 1991), gravel parking area. Silene nocti er oaeipeeldeen ede 8 km S of Carlisle, Zanol 57 (Jul
199

ELASTRACEAE: Celastrus orbiculata—BEDFORD: S of Breezewood on I-70, Cusick — (Sep e's
CUMBERLAND: NE of Carlisle Springs on PA 34, Cusick 27813 (Sep 1988); FAYETTE: NW of Perryopolis
PA 51, Cusick & Shelton 27886 (Oct 1988); FULTON: S of Crystal Spri 8 on I-70, Cusick ee (Sep ia:
SOMERSET: near entrance to Kooser emis Park, Cusick & Shelton 29791 ee ug lag Pg of woods;
WASHINGTON: E of Bentleyville on PA Turnpike, Cusick 28432 (Jul 1989); WESTMORE : NE of Belle
Vernon on PA Turnpike, Cusick pritin (Jul 1989); widely planted, it has escaped and spread, pia Feces
natural vegetation a et al.
CHENOPODIA poses patula—PERRY: Juniata River S of Millerstown, Cusick 27825 (Sep 1988),
fishing access site. Fins opodium glaucum—BEDFORD: NE of Kegg on PA Turnpike, Cusick 28435 (Jul 1989),
road berm; ERIE: gy of a 20 and PA 18, W of Girard, Cusick 29152 (Aug 1990). Kochia scoparia vat.
ee aaa RLAND: NE of Newburg on PA Turnpike, Cusick 27854 (Sep 1988), road berm
birt esaaieia: Jauenon tamnifolia—ERIE: Car Works Yard, Miller s.n. (Sep 1914): from Miller
Preis, only western ord.
CRASS ULACEAE. Sem cirnliekdc aeemuiens 4 mi W of New Eagle, Thompson 3707 (Jun 1987),
naturalized near small trash dum
CYPERACEAE: Gas gibi var. emmonsii (= C. emmonsii)—TIOGA: 1.5 mi S of US 6 on road to Colton
State Park, Rettig 1330 (Jun 1985); WARREN: ca 5 mi E of Warren, Rettig 1343 (Jun 1985), slope above road.

NOTEWORTHY PENNSYLVANIA PLANTS 71

Carex bushii—LAWRENCE: Rose Point, Isaac & Isaac 3433 (May 1991), dry open field. Carex bromoides—
MERCER: Mercer, Brandy Springs Park, /saac 3467 (Jun 1991), wet hummock; NW edge of Grove City, Tamarack
Swamp, /saac 3485 (Jun 1991), swampy woods. cays lacustris—LAWRENCE: Rose Point, Isaac & Isaac 3416
(May 1991). Carex lucorum—McKEAN: ca 10 m of Bradford, Rettig 1352 (Jun 1985), along trail. Carex
ASF se ahpinpennc : ca 10 mi W of B altel pain 135], 1353 (Jun 1985), along trail. Carex seorsa—
AW CE: 2 mi NW of Harlansburg, Jsaac 3505 (Jun 1991), pastured — on peat mat. Carex tenera—ERIE:
Presque Isle — Park, ene 1985:101 (May 1985), local on moist flats. rex willdenowii—HUNTINGDON:
Rothrock State Forest, 1.6 mi from Whipple Dam State Park, Rettig i "dun 1985), dry slope near top of
mountain; RREN: ca 5 mi E of Warren, Rettig 1344 (Jun 1985), slope above road. Scirpus atro-
cinctus—PERRY: New Bloomfield, Gress s.n. (Jul 1920); originally identified as S. cyperinus, recently redetermined
as S. atrocinctus by A. E. Schuyler. Scirpus atrovirens—MERCER: Mercer, Brandy Springs Park, /saac 346] (Jun
1991), swale below pond. rsa hattorianus—CENTRE: N of Farmers Mills on Penns Cave Road, Cusick 29072
(Jul 1990), moist field.
DIPSACACEAE: Dipsacus ca se ace E of Alexander on US 40, Cusick 28431 (Jul 1989).
This is the first record of this European species from w
DRYOPTERIDACEAE: Dryopteris parrsaieeres ca 0. 1 mi S of Hope Mills, Isaac 3669 (Sep 1991),
along old railroad grade.
ELAEAGNACEAE: Elaeagnus umbellata—BUTLER: Lake Arthur, Badin tt aye sess oe GREENE:
2 km SW of Garards Fort, Thompson & Rawlins 8321, (May 1991); WESTMORE Powdermill Nature
Reserve, Utech 91-51 (May aie: dry rocky slope; specimen from Butler County pe in Thompson et al. (1989)
Ea bes ifolia is an incorrect determination, there are apparently no specimens from Butler County.
FABACEAE: punlions 5 latflus—MERCER 0.2 mi N of Hope Mills, Jsaac 3668 (Sep 1991), nis old railroad
grade. peace cun LAND: Waltz Mill Station, Utech 91-1669 (Sep 1991), open sandy area.
Lespedeza ona ec IN: N of Pogue, Cusick 27844 (Sep 1988), S-facing shale banks along Aughwick
Creek. Lotus corniculatus—ERIE: junction of US 20 and PA 18, “ of Girard, Cusick 29154 (Aug 1990).
TIANACEAE: Géentianopsis procera—ERIE: lake shore west, Miller s.n. (Oct 1900), clay cliffs; ranges
from western New York and southern Ontario to the upper midwestern states (Mason and Iltis, 1965), Miller’s
collection apparently the only specimen from PA.
je egp uraean Elodea canadensis—ALLEGHENY: NW end of North Park Lake, Isaac et al. 3509

i! ey pe shallow w
CEAE: pian articulatus WESTMORELAND: SE of New Stanton on PA Turnpike, Cusick 29074
a ey wet ditch. Juncus bufonius—HUNTINGDON: junction of US 22 and PA 45 at Water Street, Cusick
29082 (Jul 1990). Juncus dudleyi—FRANKLIN: Blue Ridge Summit, Cusick 28615 (Sep 1989), wet field. Juncus
gerardii—SOMERSET: NE of Somerset on PA Turnpike, Cusick & Shelton 28942 (Jun 1990), moist, weedy ground;
first report from western PA, Stuckey (1980) describes spread westward from east coast through Great Lakes region,
tolerance if tt allows it to flourish in man-made saline habitat such as road berms. Juncus tenuis—SUSQUE-
HANNA: Ararat, Thompson & Macdonald 3377 (Jun 1986), along old railroad bed; previously reported from all
counties in on except Susquehanna (Wherry et al. 1979).
YCOPODIACEAE: Lycopodium clavatum—CUMBERLAND: near Hunters Run at edge of Michaux State
Forest, Isaac & Isaac 3337 (Mar 1991), along roadside bank. Lycopodium tristachyum—CUMBERLAND: near
unters Run at edge of Michaux State Forest, orn & Isaac 3336 (Mar 1991), along roadside ban
LYTHRACEAE: Lythrum salicaria—JUNIATA: Thompsontown, Cusick 27823A (Sep 1988), slong Delaware
Creek.
MORACEAE: Humulus lupulus—MERCER: ca 1 mi N of Hope Mills, /saac 3663 (Sep 1991), along old railroad

grade.
Pe cpaaepatan Nyssa sylvatica—LAWRENCE: ca 0.5 mi SE of New Bedford, /saac & Isaac 3446 (May 1991),

ef LEACEAE: Chionanthus virginicus—ALLEGHENY: 0.5 mi N of Homestead, /saac 3643 (Aug 1991), on bluff.

ORCHIDACEAE: Spiranthes casei—LYCOMING: Cedar Run Bog, Gress et al. s.n. (Sep 1920). McKEAN:
S of Cyclone, Jennings s.n. (Sep 1937); POTTER: W of Mills, Graham s.n. ie 1926); specimens originally
identified as Spiranthes cernua were redetermined’ by Paul Catling, co-author of S. casei, other than Catling’s
unpublished oa (pers. comm.) not reported for P

AGRACEAE: Ludwigia at oe NE end of North Park Lake, Isaac et al. 3510 (Jul

1991), in a water.

OXALIDACEAE: Oxalis corniculata—SOMERSET: PA Turnpike at Somerset, Cusick 27856 (Sep 1988), motel
area,

POACEAE: Calamagrostis porteri—JUNIATA: Tuscarora Mountain, NW of Ickesburg, Cusick 27824 (Sep

42 BARTONIA

1988); PERRY: Tuscarora State Forest, SE of East Waterford, Cusick 27801 (Sep 1988). Digitaria san-
pine LAWRENCE: ca 0.25 mi SE of New Bedford, <0 & Isaac 3298 (Sep 1990), garden weed. Diplachne
acuminata (= Leptochloa acuminata)—CUMBERLAND: NE of Carlisle on PA Tum mpike, Cusick 27798 (Sep 1988),
road berm; FRANKLIN: SE of Willow Hill on PA ser fetal sin (Sep 1988), road berm; first collected
in PA in 1984 ob pedi 1988), has spread — along salted ro s and now known from eight counties in
western PA. Hordeum jubatum—WESTMORELAND: ca 2 mi ae ms Latrobe, Utech 91-704, (Jul 1991), open
erase secre meadow. eae perenne—LAWRENCE: just E of Neshannock Falls, Isaac 3735 (Sep 1991),
oadside. Microstegium ineum (= Eulalia viminea)\—CUMBERLAND: Colonel Denning State Park, SW of
snes g Gap, Cusick elt :. (Sep ae picnic area, HUNTINGDON: N of Pogue, Cusick 27843 (Sep 1988), along
Aughwick Creek; JUNIATA: oo ontown, Cusick 27822A (Sep 1988), along Delaware Creek; PERRY:
Tuscarora State Forest, S of E sane ne Cusick 27807 (Sep 1988), along streams. Panicum agro-
stoides—PERRY: S of Let etn Cusick 27824A (Sep 1988), along Juniata River. Panicum aan
florum—LAWRENCE: ca 0.25 mi SE of New Bedford, Isaac & Isaac 3301 (Sep 1990), garden weed. Phragm
australisa—WESTMORELAND: ca 5 mi ENE of Waterford, Utech 91-1989 (Sep 1991), wet areas along pier
ernesosnaes distans—CENTRE: Buffalo Run, Cusick 28411 (Jul shy subise ground; CLARION: W of Wentlings
Corners on I-80, Cusick 28396 Gul 1989), road berm; CLEARFIE Clearfield on I-80, Cusick 28395 (Jul
1989), pee rm; ERIE: junction of US 20 and PA 18, W of se pits es (Aug 1990); pane junction
of I-79 and I- 80, SE of Mercer, Cusick 28398 (Jul 1989); spread along salted roadways throughou tern PA since
mid-1980’s (Cusick 1988). Setaria las LEGHENY: 4 km N of Tarentum, Zanol 18] a 1900) BEAVER:
between Koppel and Homewood, Utech 87-803 (Aug 1987), open wet ys Fae slope; INDIANA: ca 4 mi N of
Blairsville, Utech 90-1226 — 350 fields and meadows; SOMERSET: 2.25 mi NNE of Thomas Mills, Ross s.n.,
ry roadside; WESTMO D: ca 3 mi SW of Rector, Utech es sos oul ca ruderal meadow. Sefaria
glauca—LAWRENCE: rm ae of Neshannock Falls, Isaac 3713 (Sep 1991), al eek. Torreyochloa pallida (=
ip ee ose, Serge RD: S of Pennline pene s Road, Svan Sor Onl 1990), along Pymatuning Lake
INACEAE: Polygonum aviculare—LAWRENCE: just E of Neshannock Falls, Jsaac 3710 (Sep 1991),
song pois Polygonum cpm FAYETTE between Indian Head and Nebo, Utech 9]-1338 (Aug 1991),
andy soil; FOREST: ca i S of Tionesta, Isaac 3796 ea 1991), along highway; LAWRENCE: just E of
Meta Falls, /saac pie (Sep 1991), along creek. SOMERSET: Forbes State Forest along Jones Mill Run,
Utech 91-1319 (Aug 1991), sandy soil. Polygonum ee reccang nets NW of Marshall on Cold Springs Road,
Cusick 28611 (Sep 1989), weedy thicket; population reported here very lea has the potential to become major

1343 (Aug 1991), wet sandy soil; now collected from all PA counties. priced orbiculatus—ERIE: Presque Isle,
Jennings s.n. (Aug 1905); previously identified as R. verticillatus, recently redetermined as R. orbiculatus by James
Bissell.

PRIMULACEAE: Lysimachia thyrsiflora—LAWRENCE: Rose Point, Isaac & Isaac 3442 (May 1991), swamp.

SAXIFRAGACEAE: Ribes glandulosum—FOREST: 6 km NNW of Pigeon, Thompson & Rawlins 8309 (Apt
sg on large boulders

NUNCULA LACEAE: Ranunculus bulbosus—LAWRENCE: near Neshannock Falls, Isaac & Isaac 3398 (May

on edge of wet deciduous woods. sine repens—LAWRENCE: Rose Point, Isaac & Isaac 3366 (Apr
1991), swampy deciduous woods along gravel ro

RHAMNACEAE: Rhamnus cnjolio— MERCER 1.3 km W of Sandy Lake, Bier s.n. (Sep 1989), edge of
wooded eee: and shru

OSACE

| Se

REN
road; now collected in all counties in PA. ‘Giitiatod Pessintebig LEGHENY: Pittsburgh, Thaapson'é 30 (M
1983), ee along banks of ravine. Rosa multiflora—LAWRENCE: ca 0.5 mi SE of New Bedford, Isaac %
Isaac 3451 (May 1991), overgrown field.
RUBIACEAE: — concinnum—WESTMORELAND: ca 2 mi SSE of Latrobe, Utech 91-690 (Jul 1991),
sandy roadside meadow:
SALICA iia ale x gileadensis—CRAWFORD: NW of Shermanville on US 6, Cusick 29024 (Jul 1990),

moist groun

sc el eee Chaenorrhinum minus—CUMBERLAND: Blue Mountain, SW of Doubling Gap,
Cusick 27817 (Sep 1988), road berm. Lindernia Gaus Foca nee ea Creek, SE of Barnes Gap, Cusick
27704 (Aug 1988). Paulownia tomentosa—ALLEGHENY: Pittsburgh, near Highland Park, Zanol 3 371 (Sep 1990),
persisting along oe

SPARGANIACEAE: Sparganium eurycarpum—SUSQUEHANNA: 0.5 mi E of I-81 on PA 171, Thompson &
Macdonald 3383 ya 1986), roadside pond.

VERBENACEAE: Phyla caiabay (= Lippia lanceolata)—MERCER: Shenango Reservoir, Lochner s.n. (Sep

NOTEWORTHY PENNSYLVANIA PLANTS 73

— on south shore
ITACEAE: Vitis cinerea var. baileyana (= V. a 1 mi W of Glenshaw, Jennings s.n.
pe 1919), along west branch of Little Pine Creek; FAYETTE: near South Connellsville, Graham s.n. (May 1934),
above Youghiogheny River; FULTON: 0.5 mi N of Maryland bore Buker s.n. (May 1963), along see Tonoloway
Creek; GREENE: 2 mi S of Ryerson, cng & Beer s.n. (Aug 1951); ca 5 mi above mouth of Dunkar dC Creek,
Graham s.n. (Jun 1934); WASHINGTON: Riverview, Gress s.n. (Jun 1919); WESTMORELAND: ca 3 mi above
mouth of Jacob’s Creek, Graham s.n. (Jun 1934); previously not known from PA, these prord jot
identified as several different species were redetermined by Michael Moore.

ACKNOWLEDGMENTS
We thank Charles Bier, James Bissell, Bonnie and Joe Isaac, Fred Lochner, Jeff Rettig,
Fred Utech, Walter Zanol, and the Erie Historical Museum and Planetarium for depositing
specimens at CM and making these records available. We also thank James Bissell, Paul
Catling, and Michael Moore for sharing unpublished data.

REFERENCES

Cusick, A. W. 1987. Polygonum perfoliatum L. (Polygonaceae): A significant new weed in the Mississippi drainage.
Sida 12: 246-249.
———. 1988. agg collections. Castanea 53: 311-

Dreyer, G. D., L. M. Baird, and C. Fi one » 1987. ae sation and Celastrus orbiculatus: Comparisons of
reproductive potential pamades and an introduced woody vine. Bull. Torrey Bot. Club 114: 260-264.
and H. Iltis. 1965, gai reports on the flora of Wisconsin No. 53. Gentianaceae and

Menyanthaceae. Trans. Wis. Acad. Sci., Arts & Letters 54: 295-329

Miller, J. 1923. Flora of Erie County, Pennsylvania. Erie Woman’s Club, er Pennsylvania.

Strother, J. L. 1969. Systematics of Dyssodia. Univ. Calif. Publ. Bot. 4

Stuckey, R. S. 1980. bine Precti and establishment of Juncus foe (Juncaceae) in the interior of North
America. Sida = 334

Thompson, S. A., W. E. oe and M. Macdonald. 1989. Notes on the distribution of Pennsylvania plants based
on specimens in the Carnegie Museum of Natural History herbarium. Special Publ. Carnegie Mus. Nat. Hist. 14:
1-55.

Wherry, E. T., J. M. Fogg, Jr., and H. A. Wahl. 1979. Atlas of the flora of Pennsylvania. Morris Arboretum of the
University of Pennsylvania, Philadelphia.

Bartonia No. 58: 75-77, 1994
Magnolia tripetala in Pennsylvania

ANN F. RHOADS
Morris Arboretum of the University of Pennsylvania
9414 Meadowbrook Avenue, Philadelphia, PA 19118

Magnolia tripetala L., Umbrella tree, is a sub-canopy tree of low elevation, deciduous
forests of the southern Appalachians from Georgia and Arkanasa north to Pennsylvania, West
Virginia, Ohio, Kentucky and Missouri (Little 1977). In Pennsylvania it is found in the
wooded ravines along the lower reaches of the Susquehanna River where it occurs with other
components of a diverse, mixed deciduous forest dominated by Tuliptree (Liriodendron
tulipifera), another member of the family Magnoliaceae. The species is also found at
scattered sites in central and southwestern counties (Wherry et al. 1979) within the mixed
mesophytic forest zone described by Braun (1950).

Magnolia tripetala is also found in several counties of southeastern Pennsylvania.
However, the fact that it is not mentioned in the early floras of Philadelphia, Bucks or
Chester Counties is curious (Barton 1818, Darlington 1837, Porter 1903, Small 1913, Benner
1933, Stone 1945). Keller and Brown (1906) report the species as cultivated at Bartram’s
Garden and the Zoological Garden in Philadelphia as well as occurring naturally in woods
along the Susquehanna River in Lancaster County. A tree with leaves 18 to 24 inches long
is not easily overlooked, and surely would have been noticed by the early botanists had it
been a component of the forests in the Philadelphia area. One has to question whether
Magnolia tripetala is a recent arrival in southeastern Pennsylvania.

An examination of the herbarium specimens at the Academy of Natural Sciences is
revealing. The earliest specimen of Magnolia tripetala from southeastern PA, other than the
one planted at the Westtown School in Chester County in 1906, was collected in 1921 in
Montgomery County. A note on the specimen says "single tree, woods, Militia Hill." There
are hundreds of individual trees of all ages at that site today which is part of Fort
Washington State Park.

In 1922 Magnolia tripetala was noted as being "abundantly naturalized" in woods along
Tacony Creek in Montgomery County adjacent to northeast Philadelphia and by 1924 it was
reported from the Wissahickon Ravine in the city. Its spread in the Wissahickon is charted
by specimens collected in 1937, 1949 and 1964 when it was reported as "well established."
Today the species is abundant on slopes along the Wissahickon near Rittenhouse Village.

The earliest record from Bucks County (north of Philadelphia) is from 1937, the specimen
carries the note "established in thicket in the vicinity of Newportville.". By 1970 it had
moved north to the area around Bristol, and by 1989 additional populations had been located
even further north. At one site near Emilie, several hundred plants of all ages were scattered
throughout a woodland dominated by Liriodendron tulipifera. In addition the species had
become very abundant in the Delhaas Woods near Bristol, where an estimated 500 or more
trees, comprising all age classes, dominate the sub-canopy in several acres of low, moist,
Coastal Plain woods.

In Delaware County, in the western suburbs of Philadelphia, Magnolia tripetala as
collected three-quarters of a mile west of Haverford in 1939 and noted as "an escape now

7

16 BARTONIA

abundant in thin woods." By 1946 it was reported as "naturalized in woods near the Tyler
Arboretum" 2 miles northwest of Media.

Other than the record of a planted specimen at the Westtown School in 1906 there were
no collections of Magnolia tripetala from Chester County until very recently. However, in
1989 it was collected near Oxford where several hundred plants were noted in a remnant
woodland along Route One. Another smaller population was noted in woodlands adjacent
to the Nottingham Serpentine Barrens in the same year (Rhoads and Klein).

Magnolia tripetala is classified as rare by PA DER (Pennsylvania Department of
Environmental Resources 1987) but is not considered globally endangered because of its
more frequent occurrence outside the state (Pennsylvania Natural Diversity Inventory). It
is apparent from the herbarium records cited above and the absence of any reference to
native occurrence of the species in the published floras of southeastern Pennsylvania (even
as late as 1945), that Magnolia tripetala is a fairly recent arrival in woodlands in this part
of the state.

In addition to the 1906 reference by Keller and ras to M. tripetala at Bartram’s Garden
and the Zoo, it is known that the species was n at the nursery and botanical —n
established by Humphrey Marshall in Chester eelly PA 1773—1813 (Gutowski 1988). B
the mid to late 1800s prominent American landscape architects such as A. J. Downing =
Samuel Parsons were extolling the virtues of the tree for use in estate gardens (Downing
1852, Scott 1886, Parsons 1895) and Hedrick (1988) mentions that M. tripetala was also
used for understock for more tender species of magnolia

Whether its sudden appearance in the Philadelphia area in the 1920s represents spread
from cultivated sources or natural dispersal events is debatable. However, the spread of the
species in the counties around the city, as indicated from the herbarium records, seems to
radiate from Philadelphia, not from the known native occurrences in Lancaster Co. What
is clear is that Magnolia tripetala is actively expanding its range in the state, and developing
sizeable populations at a number of woodland sites in the Piedmont and Coastal Plain
provinces.

LITERATURE CITED

Sy W. as C. 1818. Compendium florae Philadelphicae. M. Carey and Son, Philadelphia.
ER, W. M. 1932. The flora of Bucks County, Pennsylvania. Ph.D. thesis, University of Pennsylvania,
Philadelphia.
BRAUN, E. L. 1905. Deciduous forests of eastern North America. The Blakiston co Philadelphia.
DARLINGTON, W. M. D. 1837. Flora cestrica. Published by the author, West Ches
DOWNING, A. J. 1852. A treatise on the theory and practice of landscape a adapted to North America.
George P. Putnam, New York.
GUTOWSKI, R. R. 1988. Humphry Marshall’s botanic garden: Living collections 1773-1813. Master’s Thesis,
University of Delaware, Newark
HEDRICK, U. P. 1988. A history of horticulture in sai to 1860, with an addendum of books published from
1861—1920 by Elizabeth Woodburn. Timber Press,
KELLER, I. A. AND S. BROWN. 1905. Flora of Philadelphia and waciaaey. racic Botanical wae dione!
i of

PENNSYLVANIA DEPARTMENT OF ENVIRONMENTAL RESOURCES. 1987, Tite 25, Chapter 82. Conservation of
Pennsylvania native wild plants. Pennsylvania Bulletin 17-5027.

PENNSYLVANIA NATURAL DIVERSITY INVENTORY. Unpublished. PNDI database. The Nature Conservancy, The
Western Pennsylvania Conservancy and DER Bureau of Forestry, Middletown.

PorTER, T. C. 1903. Flora of Pennsylvania. Ginn and Company, Publishers, Boston.

MAGNOLIA TRIPETALA IN PENNSYLVANIA TT

RHOADS, A. F. AND W. M. KLEIN. eo Flora of Pennsylvania database. Morris Arboretum of the
University of Pennsylvania, Philadelp

ScoTT, F. J. 1886. The art of beating a home grounds. John B. Alden, Publisher, New ay

SMALL, J. K. AND J. J. CARTER. 1913. Flora of Lancaster County. Published by the authors, New

STONE, HUGH E. 1945. A flora of si ae Pennsylvania. The Academy of Natural Sciences, se aciaidie
HERRY, W., J. M. FOGG AND H. A. WAHL. 1979. Atlas of the flora of Pennsylvania. Morris Arboretum of the

University of Pennsylvania, Philadelphia.

Bartonia No. 58: 79-96, 1994

Additions, Range Extensions, Reinstatements, and
Relocations in the New Jersey Flora

DAVID B. SNYDER
New Jersey Department of Environmental Protection and Energy
Division of Parks and Forestry, Office of Natural Lands Management
New Jersey Natural Heritage Program, CN 404, Trenton, NJ 08625

This is the fourth in a continuing series of papers for Bartonia reporting on recent
discoveries of rare native New Jersey plant species. In this paper four species (Atriplex
subspicata, Cardamine maxima, Carex formosa, Carex joorii) are reported as apparent
additions to New Jersey’s flora. All but one (Cardamine maxima) of these species are range
extensions, as is also Carex louisianica. The remaining taxa are all reinstatements of
historical or extirpated species. The majority of these reinstatements are of taxa which
apparently have not been collected in New Jersey for more than 50 years, or in a few cases,
not collected since the last century. While most were discovered at new locations, several
of these taxa were relocated at the same location from which they had been collected over
25, 50, or 70 years prior. Such relocations underscore the longevity that rare plant species
often demonstrate, and such lapses of time frequently reflect only a lack of active fieldwork,
and are not, therefore, reliable as evidence that a species has been truly extirpated from New
Jersey.

Nomenclature, unless noted otherwise, follows Kartesz and Kartesz (1980) or Gleason and
Cronquist (1991). Species are arranged alphabetically within phylogenetical sequence
according to Cronquist in Gleason and Cronquist (1991). Duplicate specimens of all taxa
discussed will be placed in CHRB, with some additional duplicates deposited at PH.

Quercus imbricaria

A small grove of about 20 twenty trees of Quercus imbricaria was located on 10
September 1989 on the wooded slope of a greensand marl (glauconite) ravine along Mantua
Creek at Wenonah, Gloucester County. This is a relocation of Robert Hirst’s 1962 station
(along railroad, south edge of Wenonah, 25 Aug 1962, B. Hirst s. n., PH), which in turn,
apparently is a relocation of a 1904 collection from Wenonah (Wenonah, 25 September
1904, G. T. Hastings s. n. NY). It was growing in association with Pinus virginiana,
Quercus alba, Q. montana, Liriodendron tulipifera, Carya sp., Plantanus occidentalis, Acer
rubrum, Juglans nigra, Cornus florida, and Sassafras albidum. Some of the Q. imbricaria
trees were estimated to be 50 feet high. There is no strong evidence suggesting that these
trees were planted or derived from cultivated trees, although the clearing of some woodland
to create a ballfield, has now left a few trees standing in a mowed lawn.

Quercus imbricaria is much more common west of the Appalachians, and reaches its
greatest abundance in the Central Lowland Province (Hunt 1990). The distribution of Q.
imbricaria in New Jersey has been reported as scattered throughout most of the Coastal Plain
counties, from Mercer south to Cape May (Fairbrothers and Hough 1973; Snyder and Vivian
1981; Hough 1983), but this is largely erroneous and is based primarily on misidentified

79

80 BARTONIA

specimens at CHRB and PH. Some of the counties included in Hough (1983) are based on
specimens collected from cultivated trees. Most of the remaining specimens are apparently
hybrids involving Q. phellos and other species. New Jersey’s most infamous imposter
imbricaria are the trees along Highs Beach Road on the Cape May bayshore. These
problematic trees have been the source of debate for more than 30 years, with opinions and
annotations running from hybrid taxa to Q. /aurifolia, and most recently, Q. imbricaria.
These specimens (CHRB) have now been identified as Q. phellos x Q. marilandica (Hunt
1990). Hunt (pers. comm.) has likewise disposed of all of the New Jersey specimens filed
as Q. imbricaria at PH as misidentifications, with the exception of the Hirst specimen.
Apparently Wenonah is the only native site documented in New Jersey.

The Wenonah trees had a very densely tomentose, almost velvety, surface on the
undersides of the leaves. This character is absent from the CHRB and PH hybrid specimens,
as well as from other material I have collected elsewhere on the Coastal Plain and incorrectly
identified as Q. imbricaria. The leaves of all the latter material are essentially glabrous
underneath, with the exception of a few scattered hairs, or tufts of hairs, in the vein-axils.
The only other Quercus in New Jersey that might possibly be confused with Q. imbricaria,
would be exceptionally large leaved Q. phellos, but the leaves of this species also lacks the
permanently tomentose undersides.

Atriplex subspicata

Plants of a puzzling Atriplex were collected on 21 October 1988 in a salt marsh along
Wigwam Creek, east of Port Republic, Atlantic County. Unlike the deep green and strongly
hastate leaved A. prostrata, these plants were a peculiar, almost silvery, lighter green, and
the upper leaves (most of the lower leaves had dropped) were lanceolate without the slightest
hint of basal lobing. With the standard northeastern manuals (Fernald 1950; Gleason 1952;
Gleason and Cronquist 1963) specimens keyed to Atriplex patula var. littoralis, but they did
not compare well to other specimens identified by Taschereau as A. littoralis at NY. Using
Taschereau’s (1972) treatment, the mysterious Atriplex keyed to A. subspicata. This species
differs from other nonhastate, narrow leaved Atriplex species in its seeds, which are wider
than high, and its radicle, which is median. Atriplex glabriuscula is the only other species
of Atriplex occurring in northeastern North America that has a median radicle, but it also has
an inflorescence that is leafy bracted throughout (A. subspicata’s inflorescence is leafy
bracted only at the base) and it is not known to occur in New Jersey.

Atriplex subspicata was described in 1818 by Nuttall and occurs in coastal areas of eastern
Canada south to North Carolina, and in saline habitats in western Canada and in the western
and central United States (Taschereau 1972; Bassett, et al. 1983). Most eastern collections
of A. subspicata have been identified as varieties of A. patula, primarily because all of our
eastern manuals follow Gray’s 1868 treatment of the genus (Taschereau 1972).

In 1990 I returned to the Atlantic County site to collect additional specimens, but after
careful searching, no plants were found. The site is located within the Forsythe (formerly
Brigantine) National Wildlife Refuge in which some recent wildlife enhancement activities
(digging ponds for waterfowl) have taken place. I don’t think this is the cause of the
seeming disappearance of A. subspicata. Being an annual species growing in a tidal habitat,
it’s likely that seeds do not always fall and germinate in the same spot year after year.

Potential unexplored habitat for this species is extensive. It grew in a salt marsh
dominated by a thick carpet of Spartina patens with scattered areas of open muddy patches
with the occasional Limonium carolinianum and the usual assortment of Salicornia and
Suaeda. This type of habitat is abundant throughout coastal New Jersey.

NEW JERSEY FLORA 81

Chenopodium berlandieri var. macrocalycium

Chenopodium berlandieri var. macrocalycium was described as a species by Aellen in
1929 (Wahl 1954). It is treated as a variety of C. berlandieri by Cronquist (Gleason and
Cronquist 1991). In gross morphology var. macrocalycium resembles C. album, and C.
berlandieri and its vars. bushianum and zschackei. Chenopodium berlandieri var.
macrocalycium is easily distinguished from the C. album complex by its obviously
reticulated pericarp. Chenopodium berlandieri var. bushianum and var. zschackei both have
reticulated pericarps but the latter variety has a light yellowish area at the base of the style
and the leaves are white-mealy (unlike either vars. bushianum or macrocalycium). It is also
mostly western in distribution, and is not known to occur in New Jersey (Wahl 1954;
Gleason and Cronquist 1991). The var. bushianum has larger seeds then does var.
macrocalycium (1.5—2.3 vs. 1.3—1.7) its inflorescence typically droops, and it mostly occurs
as a weed in cultivated fields (Koster D/1-20-1, PH) and waste places (Brown s. n.,

Chenopodium berlandieri var. macrocalycium is primarily coastal in distribution occurring
from Newfoundland south to North Carolina and also inland in the upper Mississippi
drainage and the Great Lakes (Wahl 1954; Gleason and Cronquist 1991). The majority of
New Jersey collections were made from sea beaches in Monmouth, Ocean, Atlantic, and
Cape May counties, and from the Delaware River Bayshore in Cape May County (see Wahl
1954 for specimen citations). The most recent specimen contained in our local herbaria
(CHRB, NY, PH ) was collected 16 October 1937 (Ludwig 643 PH). Apparently the only
recent records for C. berlandieri var. macrocalycium are those I collected at Sandy Hook,
Monmouth County on 21 September 1986 and from the Tuckerton Marshes, Ocean County,
on 14 September 1986 and 21 October 1988. At both locations the plants were growing in
damp sand at the upper limits of a bayside salt marsh. What led me to these locations were
some recent unvouched reports of C. rubrum, but as I far as I can tell, C. rubrum is not
present at either site. The superficial resemblance of the two taxa may account for some of
the unsubstantiated reports of C. rubrum in New Jersey.

Chenopodium rubrum

Chenopodium rubrum has been reported as a native New Jersey plant in both the early
literature (Knieskern 1856; Britton 1889; Britton and Brown 1913; Stone 1912; Taylor 1915)
and in recent manuals (Fernald 1950; Gleason 1952; Gleason and Cronquist 1963) and state
checklists (Anderson 1989; Hough 1983; Schmid 1990). I have been unable to locate
specimens at BK, CHRB, CU, GH, NY, or PH that substantiate these reports. On the
contrary, the specimens that I have seen at these institutions all seem to support a nonnative
status: CAMDEN CO.: Ballast, Camden, 23 Sep 1877, C. F. Parker s. n., CHRB; Ballast,
Camden, 15 Aug 1879, C. A. Boice s. n., PH; Ballast, Camden, 30 Aug 1879, C. F. Parker,
s. n., GH; Kaighn’s Point, 1898, A. Jahn, s. n., PH; On ballast, Kaighn’s Point, [not dated]
I. Burk, s. n., PH. HUDSON CO.: Hoboken, [undated and without collector’s name] NY.

Stone (1912) apparently also had difficulty in verifying these early reports of C. rubrum
as evidenced by his statement, "It is true that it has been mentioned in all the lists, but
apparently without any definite knowledge of its occurrence, and I have been unable to find
any specimens in the herbaria that I have examined except those collected by Mr. Bayard
Long on Barrel Island, near Tuckerton, September 11, 1908." I have been unable to locate
Long’s specimens cited by Stone. Stone missed the Burk and Parker Camden specimens
cited above even though he acknowledged examining their herbaria (Stone 1912).

More recently C. rubrum was reported, but not collected, from Secaucus, South Amboy,
Sandy Hook, Island Beach, Barnegat, Tuckerton Marsh, and Moore’s Beach by Vincent

82 BARTONIA

Abraitys (1957-1983) and from Seven Bridges, near Tuckerton by Gilbert Cavileer (pers.
comm.). This last location is about four miles southwest of where Long reportedly collected
it and may be also the same as Abraitys’ Tuckerton Marsh record. I have searched this site,
as well as Abraitys’ Sandy Hook and Island Beach locations (plus many other seemingly
suitable areas along the coast), but without success. It is possible that at least some of these
reports may be based on misidentified C. berlandieri var. macrocalycium (see discussion
above). In 1978, Joe Arsenault collected C. rubrum northeast of Smithville (about 6 miles
east of where Cavileer reported it) and deposited a specimen in the herbarium of Rutgers’
Camden campus. No plants were located at this site in 1988 when Joe, Ted Gordon, and I
made a careful search. Joe also told me about another specimen of C. rubrum that had been
collected along the Tuckahoe River in Atlantic County on 23 September 1982 by Leo Snead,
Jr. I examined this specimen, now at CHRB, in late 1989 and got very precise directions
to the location. It is on the north bank of the Tuckahoe River, Atlantic County, approximate-
ly 2.5 miles northeast of Middletown. On 12 October 1990 I found a patch of 28 plants here
in a dense stand of Spartina cynosuroides, growing with Limonium carolinianum,
Kosteletzkya virginica, Teucrium canadense, and Pluchea odorata. The site is remote and
accessible only by boat, and introduction by any but natural means, seems unlikely. In 1991,
Stephen Field found a population along Great Cedar Swamp Creek, Cape May County, not
too far from the above site. Thus doubt and rumor can be put to rest—C. rubrum does occur
in New Jersey as a native species.

Lechea tenuifolia

The only documented location for Lechea tenuifolia in New Jersey is Ringwood Junction,
Passaic County, collected by K. K. Mackenzie in 1916 and 1917 (Dry sunny bank, local, 7
Jul 1916, 7/45, NY, PH; Dry fields, Ringwood Junction, 20 May 1917, 7608, CHRB, NY,
PH). By the late 1920s, Ringwood Junction existed no more, having been flooded to create
Wanaque Reservoir. Curious as to whether some L. tenuifolia might have survived at higher
elevations adjacent to the reservoir, I made a search on 18 September 1990 and located a
population of 58 plants (Snyder 1993). They were ina clearing on a rock ledge in very dry
soil. Associated species included Aster liniariifolius, Solidago bicolor, Hieracium
paniculatum, H. venosum, Lysimachia quadriflora, Potentilla canadensis, Quercus alba, Q.
rubra, Fraxinus americana, and Acer saccharum.

The habitat seemed very ordinary and similar to hundreds of other places in northern New
Jersey. If habitat for this species is so abundant, why is Lechea tenuifolia so rare in New
Jersey? Although it may be undercollected in the state, I suspect that this species will
remain extremely rare in New Jersey. Although much more abundant in the western and
southern portions of its range, L. tenuifolia can hardly be considered peripheral in New
Jersey, ranging as it does from Maine, south to South Carolina, and interuptedly west to
Minnesota and Texas (Fernald 1950, Gleason and Cronquist 1991). The species apparently
is of spotty and local distribution throughout the Northeast, and eight states (Maine, New
Hampshire, New York State, Ohio, New Jersey, Delaware, Maryland, West Virginia) track
it at some level of rarity (The Nature Conservancy’s Biological and Conservation Database).
Whatever the factors may be that are limiting the distribution of L. tenuifolia in the
northeast, it seems unlikely that it is a result of a lack of suitable habitat.

Cardamine maxima

Cardamine maxima is a plant of northern affinity, ranging from New Brunswick, south

through New England to northern New Jersey, Pennsylvania, West Virginia and Kentucky,

NEW JERSEY FLORA 83

and west to Ontario, Michigan, and Wisconsin (modified from Montgomery 1955, Gleason
and Cronquist 1991). It is local to rare in most areas of its range with its main distribution
in New York State, and southeastern Canada (Montgomery 1955).

In morphological characters it is intermediate between Cardamine concatenata and C.
diphylla, differing primarily in its rhizome, which is alternately enlarged and only slightly
constricted along its length. In C. concatenata the rhizome is formed of a series of fragile,
tuber-like segments, while C. diphylla’s rhizome is without any constrictions, maintaining
a constant diameter throughout its length. Cardamine maxima can be further distinguished
from C. diphylla by its leaflets which bear along their margins spreading trichomes 0.2-0.3
mm long; those of C. diphylla are 0.1 mm and appressed. The number, shape, and position
of cauline leaves are variable (Bissell 1903; Montgomery 1955), and are unreliable
morphological characters. Cardamine maxima frequently has three alternate cauline leaves,
but varies between one to four or more (Bissell 1903; Montgomery 1955). Cardamine
diphylla typically has two alternate or opposite cauline leaves and C. concatenata frequently
has three opposite to subopposite leaves; but again this is variable, particularly in C.
concatenata (Fernald 1950; Montgomery 1 ,

Its variability, and seemingly intermediate morphological characters, has led some workers
to consider C. maxima to be of hybrid origin (Montgomery 1955; Voss 1985; Gleason and
Cronquist 1991). Wiegand and Eames (Montgomery 1955) suggested that C. maxima was
a hybrid between C. concatenata and C. diphylla. Montgomery (1955) has shown that both
these taxa and C. maxima are sexually sterile and that the chromosome number of C. maxima
also does not show it to be a direct hybrid between these species. He concluded that C.
maxima "is amember of the agamic complex that originated before complete sexual sterility
was effected in the members of the complex. It may have been derived during some
interglacial period of the Pleistocene glaciation" (Montgomery 1955). And although
polymorphic, C. maxima is no less variable than its sexually sterile near relatives, C.
concatenata and C. diphylla. There would seem then, little taxonomic consistency in treating
only one of the taxa in the concatenata-diphylla-maxima complex as a hybrid. I prefer to
follow Montgomery (1955) and Rollins (in Kartesz and Kartesz 1994, J. Kartesz pers.
comm.) and recognize C. maxima as a valid species.

The New Jersey record is based on a single station located in a ravine along the Shimmers
Brook in Millbrook, Sussex County. In 1985, I collected specimens on 28 April and 4 May
that were variable in height, and number and shape of the leaves. On 26 April 1991, I found
hundreds of plants in scattered, dense colonies that were interspersed among colonies of C.
diphylla. Also present, were plants of what I had mistakenly identified as C. concatenata
during my 1985 surveys. These plants are much smaller than either C. diphylla or C.
maxima. The mostly two, opposite to alternate cauline leaves, are much more divided, with
the divisions narrower, and with fewer, but sharper incisions. The foliage is blue-green,
unlike the yellow-green foliage of C. diphylla and C. maxima. Although the root stock of
an occasional plant was more obviously constricted than either of these taxa, root stocks of
the majority of plants were identical to C. maxima. In addition to the nearly perpendicular
coarse trichomes along the leaf margins, some of the plants also had few to many similar
trichomes on the stems and on one or both surfaces of the leaves. These plants key to what
has been described as Cardamine (Dentaria) anomala (Eames 1903; Gleason 1952; Gleason
and Cronquist 1991) or C. x anomala (Fernald 1950; Kartesz and Kartesz 1994). But given
the sexual sterility and the polymorphic nature of this complex, I prefer to recognize C.
anomala as an extreme expression of the C. maxima, to which it most closely resembles.

84 BARTONIA

Both C. maxima and C. diphylla grew either along the banks of a small stream or on the
adjacent narrow floodplain. It is a mixed deciduous forest with Acer saccharum, Plantanus
occidentalis, and Tsuga canadensis the common canopy species. Associated herbs include
Arisaema triphyllum, Symplocarpus foetidus, Allium canadense, Erythronium americanum,
Smilacina racemosa, Uvularia perfoliata, and the ubiquitous Alliaria petiolata. Cardamine
maxima (including "C. anomala") was both in flower and in bud, while C. diphylla had more
buds than open flowers.

Pyrola chlorantha

Pyrola chlorantha is circumboreal in distribution, occurring in the northeastern United
States south to Maryland and West Virginia (Gleason and Cronquist 1991). In New Jersey
in has been collected predominately from sites on the Coastal Plain. Its distribution on the
Coastal Plain, while scattered (Monmouth, Atlantic, Burlington, Camden, Gloucester, and
Cumberland counties), is more concentrated in the Delaware River drainage, with pe
majority of collections having been made in Burlington and Camden counties. There ar
only a few sites for this species that are not Coastal Plain, these being single locations in
each of Sussex, Passaic, and Morris counties

My relocation of Pyrola chlorantha just south of the New York state line, on 15 June
1989, apparently remains the only confirmed extant occurrence of this species in New Jersey.
In our local herbaria there are least 52 specimens collected from approximately 33 separate
locations in nine counties. For more then a decade I searched these historical locations
before finding what I presume to be Wherry’s 1948 site: Sussex County: Under pines across
road from Rockview Hotel, southwest of Duttonville, s. 7., PH. Sixteen plants occurred
scattered over a few feet of dry ground under Pinus strobus. Of these, only three produced
flowers. This site, which is located in Montague Township, is the most recent New Jersey
collection and it also may be the same as H. H. Rusby and N. L. Britton’s 1884 collection
from "Montague Township" (CHRB) and Mackenzie’s 1910 collection from "pine woods
along Delaware River, Montague Township" (NY).

Despite the numerous historical locations for P. chlorantha, it apparently never was
abundant in the state. Stone (1912) describes it as "everywhere local and not abundant."
Still, with so many historical locations for this species in the state, I have to wonder why no
new populations have been discovered or why no additional historical sites have been
relocated. Part of reason may be a combination of lack of recent fieldwork and the
inconspicuous appearance of the species, particularly when not in flower. Another part of
the reason is that several historical sites have been destroyed, especially those located in
Burlington and Camden counties where expanding cities and sprawling suburbs have covered
much of the species former habitat. With the strength of New Jersey’s wetland laws, upland
habitats are more vulnerable than ever. It may be too, that like some other northern taxa that
are disjunct, or at the southern limits of their ranges, deteriorating climatic and habitat
conditions and genetic isolation has led to inbreeding depression, and populations simply can
no longer maintain themselves. We may be witnessing the continued northward retreat of
a species that may be a relict from a time when New Jersey’s Coastal Plain was dominated
by taiga, tundra, muskegs, and mastodons (Wolfe 1977; Heusser 1979; Pielou 1991).
Cercis canadensis

While popularity of Cercis canadensis is great for the nursery trade, from the standpoint
of the biogeographer, it presents a difficult question: How do you tell the native populations
from those that may have originated from cultivated plants? In states where Cercis is a

NEW JERSEY FLORA 85

frequent component of the native vegetation, this question is relatively trivial in understand-
ing its overall distribution. But when the species is rare, or at the geographical limit of its
range in a particular state, as is Cercis in nee Jersey, the question becomes critical to the
conservation and legal protection of the s

The species has a broad geographical ck ea ranging from southern Connecticut,
southeastern New York State, central New Jersey, then south to northern Florida, west
through southern Ontario, Michigan, and Wisconsin, and south to Texas and northeastern
Mexico (modified from Hopkins 1942; Fernald 1950; Gleason and Cronquist 1991). While
there is sufficient documentation that there are at least some native populations of Cercis in
New Jersey, (Torrey 1819; Britton 1889; Stone 1912, Taylor 1915; Sargent 1922), the reports
of the species as being native in Connecticut and New York State have been questioned. '

The best evidence that Cercis occurs in New Jersey as a native, comes from the Delaware
River drainage where it was first collected in 1869 (Camden Co.: Banks of the Delaware
below Kaighn’s Point, Camden. 5 May 1869, C. F. Parker s. n., CHRB). This location is
listed by Britton (1889) as "Camden: In damp woods on the bank of the Delaware, between
Camden and Gloucester—Parker." As unlikely as Camden may seem as a site for a native
population of Cercis, the woods and swamps at Kaighn’s Point supported a number of rare
New Jersey plant species (Stone 1912). Other sites for Cercis along, or near, the Delaware
River are Bordentown (A. C. Stokes in Britton 1889, no specimen found), Burlington
County; Stockton (1 Aug 1895, H. L. Fisher s. n., PH), Rosemont (G. N. Best in Britton
1889, no specimen found), Hunterdon County; and banks of the Delaware, north of Trenton
(A. C. Apgar in Britton (1889) no specimen found), Moore’s Station (6 May 1884, 4. C.
Apgar s. n., CHRB), Titusville (10 May 1901, without collector’s name or number, Trenton
State Museum), all Mercer County. I believe that all the Mercer County records refer to the
same general locality, if not actually the same site. This site, which I relocated 30 April
1989, is on the western slope of Strawberry Hill (locally known as "Kuser Mountain"), 0.5
mile south of Moore’s Station, one mile northwest of Titusville, and about eight miles north
of Trenton. The habitat is a rich, mixed deciduous woods on a steep, boulder strewn diabase
hillside. On subsequent visits made 3 May, 30 August 1989, and 13 August 1990, 11 trees
were located scattered from the foot of the slope to its crest. Although no trees were found
that produced fruits, numerous young saplings were observed near two of the largest trees.
Growing with, or nearby Cercis are several other rare New Jersey plant species: Agastache
nepetoides, Agrimonia microcarpa, Chaerophyllum procumbens, Cynoglossum virginianum,
Jeffersonia diphylla, Hybanthus concolor, Panax quinquefolius, Stachys hispida, and
Tradescantia ohiensis. Most of these species are characteristic of rich (often calcareous)
woods, and Cercis is hardly out of place in this association.

Further evidence that Cercis is native in the woods at this site (and probably elsewhere
in the Delaware Valley) is that native populations of Cercis occur in adjacent Pennsylvania

'The record for Connecticut for Cercis as a native species is based on a single clump of trees located within New

Haven County Park (Nichols 1923; Reeder 1951). Although this record is accepted as a natural occurrence by
Hopkins (1942), Fernald (1950), and Dowhan and Craig (1976), Les Mehrhoff (pers. comm.) remains skeptical,
since the collection locality is known to have planted occurrences of some other rare Connecticut taxa. Early
editions of Gray’s Manual, as well as the eighth edition (Fernald 1950) have included southern New York State
within the native range of Cercis. Richard Mitchell (pers. comm.) is of the opinion that all records from New York
State are based on trees that have been planted or have escaped from cultivation.

86 BARTONIA

(Wherry et al. 1979), growing in habitats similar to those found in New Jersey. It seems
improbable to me that the distribution of native populations of Cercis stops abruptly on the
Pennsylvania side of the Delaware.

arther away you move from the Delaware River Valley, the more difficult it is to
distinguish native populations from the adventive or planted. While Cercis is abundant as
an ornamental planting statewide, I see no evidence indicating that the species successfully
escapes from cultivation. Planted trees may persist long after cultivation and may even
spread locally through suckering, but cultivated plants do not appear to readily colonize
suitable natural habitat. In a rather isolated area of Sussex County I found a single, very
large tree near an old foundation, growing with Syringa and other obvious cultivars.
Although the yard in which it had been planted had succeeded to woodland, there was no
evidence of sexual or asexual reproduction. There are also a few trees growing in a wooded
ravine on the Busch Campus of Rutger’s University, Middlesex County, which look native
enough, but according to David Fairbrothers (pers. comm.), they have spread from trees
planted around the College President’s residency which overlooks the ravine. Not too far
from this ravine is a single large redbud growing on the flood plain of the Raritan River.
Here it grows with native species like Carex grayii, Florekea proserpinaciodes, and
Mertensia virginica, but also with a planted species of spruce, and a suspiciously out of
place Kalmia latifolia.

The only other location that I am convinced Cercis is native is a Somerset County site,
first collected by M. A. Chrysler 8 May 1935 on the east slope of the First Watchung
Mountain, 1.3 miles west of Dunellen (CHRB). Chrysler noted it was "frequent (locally?)"
and that it "may be native." On 14 June 1989, I drove the designated 1.3 miles west of
Dunellen along Route 22 at the base of the Watchung Mountains, and ended up in the
middle of a fairly recent housing development where Cercis grew abundantly in roadside
woods. In subsequent visits, I have found Cercis to be exceedingly common here, where in
places, it is the dominant species in the subcanopy, and occurs over a mile or so stretch of
a steep, wooded, trap rock hillside.

There are other New Jersey specimens for Cercis, some of them clearly collected from
cultivation, others possibly native, but these all have vague location data and lack any
meaningful habitat descriptions.

Ammannia latifolia

Ammannia latifolia is a southern species ranging from northeastern New Jersey, south to
Florida, west to Texas, and south through the Caribbean to South America (modified from
Graham 1985). Plants from the eastern United States have long been known as A. koehnei
or A. teres, but both names are now reduced to synonymy: A. koehnei under A. latifolia, and
A. teres under A. coccinea (Graham 1985). Britton (1891) maintained that the petalous
plants from the eastern United States (i.e., A. koehnei) were distinct from the more southern
apetalous A. /Jatifolia. Graham (1985) has shown this character to be variable and noted that
petalous and apetalous plants are found in the same population, sometimes even on the same
plant.

The type locality of Britton’s A. koehnei has been given as the Hackensack Marshes, New
Jersey (Britton 1891; Stone 1912; Taylor 1915; Fernald 1936) but the actual notation on the
label of the type specimen (Leggett s. n., NY) appears to be Hackensack "Flat" or "Flats."
In any case, the locality data is so vague that the exact county remains uncertain. The

NEW JERSEY FLORA 87

Hackensack Meadowlands (as the area is now known) is extensive, and occurs along the
Hackensack River in both Bergen and Hudson counties. The extensive Atlantic white cedar
wetland that once occurred here was largely gone by 1896 as a result of ditching, diking,
logging, burning, and more recently, through filling and salt water intrusion (Sipple 1972;
Schmid 1987). Today it is hard to imagine that these wetlands dominated by Phragmites
australis, punctuated here and there by landfills, oil refineries, and warehouses, once
supported Ammannia latifolia and the botanical diversity described by Harshberger and
Burns (1919). Until Ammannia latifolia’s rediscovery in 1990, the Hackensack Meadow-
lands was its only documented New Jersey location where it was last collected there on 28
July 1868.

On 12 October 1990, I discovered a small colony of Ammannia latifolia in a brackish tidal
marsh along the south side of the Tuckahoe River, ca. 2 miles north of Middletown, Cape
May County. On 16 October 1990, a total of 620 plants were counted scattered over a
distance of nearly a quarter of a mile. The largest concentration was of 313 plants growing
in a somewhat circular open flat having patches of bare organic mud covered with one or
two inches of water. The flat was bordered by a zone of Cyperus sp. and Scirpus pungens
which graded into a dense zone of Spartina cynosuroides on the river side, and a zone of
Typha angustifolia and Phragmites australis on the upland side. Other associated species
included Atriplex prostrata, Kosteletzkya virginica, Samolus valerandi, Lythrum lineare,
Aster tenuifolius, Pluchea odorata, and Solidago sempervirens. Although | found Ammannia
growing in areas of much denser vegetation (i.e., pure stands of Phragmites), it was most
abundant in areas of open, low vegetation, particularly on, and around muskrat middens and
runs. Plants were found in all stages of development from very young plants, to plants in
bud, flower (petals pale lavender and falling with the slightest touch), and with mature fruit
capsules.

In 1991, Stephen Field informed me that he had found Rotala ramosior in a salt marsh
in the Tuckahoe River drainage, Cape May County. My suspicion that this plant was more
likely Ammannia latifolia (Rotala is not a salt marsh species) was confirmed on 18 July 1991
at Stephen’s site along Cedar Swamp Creek, a major tributary of the Tuckahoe River, which
is about three miles southeast of my location. Later Stephen told me of another population
of Ammannia that he had found on 10 August 1991 at a site a little north of his first
location. Apparently, Ammannia latifolia is well established in the marshes along the lower
Tuckahoe River.

Fernald (1936) stated that "North of Florida typical Ammannia [latifolia] is a very
localized plant." Although more locations are known today then were known in Fernald’s
day, his statement is still an accurate assessment. It is known from five or fewer extant
occurrences in both New Jersey and Maryland, from between five and twenty locations in
North Carolina (Alan Weakley pers. comm.), and apparently absent from Delaware (Tatnall
1946) and South Carolina (Radford et al. 1968). It seems to be most abundant in Virginia
where its recorded from twelve counties (Harvill et al. 1986).

Rhexia mariana var. ventricosa son leviaes

The relationships of this taxon are obscure, and its distribution north of Virginia is poorly
known. Originally described as a species by Fernald and Griscom (1935), and so recognized
by James (1956), Kral and Bostick (1969) reduced it to a variety of R. mariana. More
recently Cronquist (Gleason and Cronquist 1991) synonymized it with R. interior.
Regardless of nomenclatural preferences, the taxon is distinctive, and is easily recognized
in the field. Its general morphology suggests an intermediate between R. mariana var.

88 BARTONIA

mariana and R. virginica. Its stem is square with all faces nearly equal in width like that
of R. virginica, but essentially wingless like R. mariana var. mariana. It has the long,
horizontally spreading rhizomes of R. mariana var. mariana, but lacks the tuberous
thickenings of R. virginica. Mature hypanthiums are longer necked than R. virginica, and
are more like those of R. mariana var. mariana. The seeds of R. mariana var. ventricosa
have concentric rows of close-set papillae, which are unlike either R. virginica or R. mariana
var. mariana (Fernald 1935; James 1956; Kral and Bostick 1969).

hexia mariana var. ventricosa is restricted to the eastern United States, occurring from
southern New Jersey, south, mostly along the Coastal Plain, to South Carolina (James 1956;
Kral and Bostick 1969). It is apparently rare in New Jersey, Delaware, and Maryland, but
field work may show otherwise. Part of the reason that R. mariana var. ventricosa may be
overlooked north of Virginia is that until recently it has not been widely known that it is a
part of the flora of these states. Fernald (1950) listed it as occurring north only to Virginia,
Gleason (1952) synonymized it under R. mariana, and none of the checklists of New
Jersey’s flora have included it (Hough 1983; Anderson 1989; Schmid 1990).

On 15 October 1991, Andy Windisch and I found R. mariana var. ventricosa at an
abandoned sand pit at Pierces, just north of Kay Pond in Cape May County. There were
hundreds of plants growing in open, damp sand of an oak/pine woodland. Scattered here and
there in the colony were plants of R. virginica, but unlike the plants of R. mariana var.
ventricosa which was either still green or beginning to turn a brilliant yellow-orange, R.
virginica had already turned deep red and the majority of plants had already dropped their
leaves. In contrast, R. mariana var. ventricosa was still in good shape and even a few
flowering plants were located. Not too far away, we found an extensive colony of R.
mariana var. mariana which was in about the same condition as var. ventricosa. Comparing
the three taxa together in the field, the siren between them were apparent; something
that is not always as easy to do in the herbari

Prior to this relocation, R. mariana var. ventricosa was last collected in New Jersey on
18 September 1909 by K. K. Mackenzie, which is the basis for the New Jersey record cited
in James (1956) and Kral and Bostick (1969).

Cornus amomum var. schuetzeana

On 10 September 1990, while searching for a rare Cuscuta that had been collected from
the floodplain along the Delaware River, northwest of Columbia, Warren County, I found
a tangled thicket consisting mostly of a shrubby, unknown species of Cornus. It turned out
to be C. amomum var. schuetzeana. | later discovered that it had been collected 42 years
earlier about nine or ten miles south of my location (Warren Co.: alluvium, 1-2 mi SW of
Belvidere, 9 Jul 1948, R. L. Schaeffer, Jr. s.n., PH). This taxon has missed being recorded
as occurring in New Jersey in the recent checklists of the state’s flora (Anderson 1989;
Hough 1983, Schmid 1990). It is recorded for New Jersey by (Fernald 1950), possibly based
on the PH specimen.

The nomenclature for this species is complicated with four different names having been
applied to it within this century: C. amomum ssp. obliqua (Wilson 1965; Kartesz and Kartesz
1994), C. amomum var. schuetzeana (Rickett 1934; Gleason and Cronquist 1991), C. obliqua
(Fernald 1950), and C. purpusii (Rickett 1945; Gleason 1952). Despite the nomenclatural
problems, there is general agreement that the taxon is taxonomically significant and distinct,
except for some probable hybrids involving C . amomum var. amomum where their ranges
overlap.

NEW JERSEY FLORA 89

The var. schuetzeana is more northern and western in distribution, occurring from New
Brunswick south to New Jersey and Virginia (Harvill et al. 1986), then west and south to
North Dakota, Kentucky, Arkansas, and Oklahoma (modified from Fernald 1950; Wilson
1965). The var. amomum is more southeastern in its distribution, occurring from Georgia
west to Mississippi, northeast to southern Maine, and then west to southern Ohio and Indiana
(Fernald 1950; Gleason and Cronquist 1991). The New Jersey specimens that I collected are
unambiguous and show none of the transitional characters reported to sometimes occur in
eastern specimens (Rehder 1910; Rickett 1945; Wilson 1965). The Atlas of the Flora of
Pennsylvania (Wherry et al. 1979) shows a location dot along the Delaware River in Monroe
County that is nearly opposite of where I made my collection. With additional searching it
is possible that new locations for C. amomum var. schuetzeana will be found along this
portion of the Delaware River.

Linum intercursum

On 17 August 1990, Ted Gordon and I relocated what I take to be Stone’s 1913 location
for Linum intercursum: Burlington Co.: Symms Place, 3 mi W of Cedar (Warren) Grove, 29
Jul 1913, W. Stone 14525, PH (Snyder 1993). It was locally abundant at this site and grew
mostly in the drier portions of a recently burned pitch pine lowland forest. Nearby, we
found scattered plants growing in an open meadow, which had been several years ago, a
cultivated blueberry field. Linum intercursum was last collected at this site in 1939 (11 Aug
1939 B. Long 53083, PH) which apparently is also the last time it had been collected in the
state.

The species is distributed along the Atlantic Coastal Plain from Massachusetts south to
North Carolina then inland to Tennessee, with disjunct occurrences in Indiana (Rogers 1963).
Although commonly occurring in moist soil of oak or pine woodlands (Rogers 1963) it also
occurs in dry soil. In Pennsylvania, it has been collected on serpentine barrens (John
Kunsman pers. comm.). Its distribution in New Jersey is predominantly Coastal Plain where
it has been collected from widely scattered locations in most of the Coastal Plain counties.
Many of these collections record the habitat as dry sandy woods or dry oak and pine scrub,
but a few collections have been from wetland habitats, the most unusual being from the
sandy, clayey border of an intermittent pond (Long 47083 PH).

Despite its broad range and an abundance of seemingly suitable habitat, the species is
apparently rare throughout much of its range, with it currently being tracked at some level
of rarity in Massachusetts, Connecticut, Rhode Island, New York State, Indiana, Pennsylva-
nia, New Jersey, Delaware, Maryland, and North Carolina (The Nature Conservancy’s
Biological and Conservation Database). While populations certainly have been lost through
the development of its habitat, fire suppression has also been suggested as a significant factor
contributing to the species current rarity throughout most of the eastern part of its range
(Bruce Sorrie pers. comm.; Alan Weakley pers. comm.).

Pycnanthemum clinopodioides

Pycnanthemum clinopodioides has the most restricted distribution of any Pycnanthemum
that occurs in the northeastern United States. Grant and Epling (1943) record it from 14
counties in eight states (Massachusetts, Connecticut, New Jersey, New York, Pennsylvania,
Maryland, West Virginia, North Carolina) and the District of Columbia. At NY there is an
undated specimen collected from Green County, Indiana (A. Wood, s. n.), which, if it is

90 BARTONIA

correctly labeled, is a remarkable disjunct. It also has been reported from Delaware’.
Although some additional counties have been subsequently added to those reported by Grant
and Epling (1943), the species remains extremely local or rare throughout its range (The
Nature Conservancy’s Biological and Conservation Database). The species was first
described by Torrey and Gray in 1842 based on specimens collected in New Jersey and from
the vicinity of New York State (Grant and Epling 1943)

In general aspect this species suggests a narrow leaved Pycnanthemum incanum. It is
variable in habit, indument, amount of branching in the inflorescence, and the size and the
degree of unequalness of calyces, but constant in leaf shape and the presence of short,
hooked hairs on the stems. Leaf shape is lanceolate, often narrowly so, with median leaves
predominantly under 2 cm, rarely as wide as 2.7 cm. Both surfaces of the leaves are green,
except for the leaf-like bracts subtending the inflorescence, which may be either green, or
less frequently, canescent on the upper surface, especially near the base. By contrast, the
leaves of P. incanum are ovate, often broadly so, to oblong-ovate, and up to 5.5 cm wide
(fide Fernald 1950), with many of the upper leaves strongly canescent above. Pycnanthem-
um clinopodioides can be separated from the narrow leaved species in the genus by its more
open, distinctly branched inflorescence, bilabiate calyces, which are typically margined with
few to several long, multiicelled trichomes, and the presence on the stem of both long,
spreading hairs, and minute, strongly downwardly or upwardly curved hairs, many of which
have the tips pointing down toward the face of the stem, thus looking like a tiny inverted
7" ot.

Gray (1848) considered P. clinopodioides to be intermediate between P. incanum and P.
torrei. Grant and Epling (1943) saw P. clinopodioides as the species connecting what they
recognized as the Incanum Phylad with the Virginianum Phylad. And, although they
recognized P. clinopodioides as a valid species, they did note that the species "is rather
variable in habit and suggests strongly a hybrid origin between a member of the incanum
group and one of the narrow leaved species, such as P. verticillatum" (Grant and Epling
1943). This statement apparently serves as the point of departure for those authors since
Grant and Epling who have dismissed P. clinopodioides as being of hybrid origin (Gleason
1952; Gleason and Cronquist 1963), or have simply reduced it to synonymy (Radford et al.
1968). Current treatments, such as Kartesz and Kartesz (1994) and Cronquist (Gleason and
Cronquist 1991) recognize P. clinopodioides as a species.

In recent years, I have come to the opinion that P. clinopodioides is a distinct species and
I have spent many hours in the field trying to find it. Most of my efforts were concentrated
around six historical sites in Passaic, Bergen, Hunterdon, Mercer, and Gloucester counties
where the most recent specimen is: Gloucester Co.: Jennings. 2 Aug 1891, C. D. Lippencott

n. CHRB). This past winter, as I was going through my specimens of Pycnanthemum
hei for misidentifications, I found that I had collected P. clinopodioides on 20 July

hegre (1946) cites two records for Delaware. The first is a specimen at PH that was collected 18 ee, 1876
y A. Commons from Wills Rock, Wilmington, New Castle County. Since this specimen is not now

claps folder at PH and because Delaware is not included in the range given by Grant and pling (1943),
it may have been reidentified as something else. The second record is based on a specimen, which according to
ee tN was collected by Canby in 1872 "near Swayne Station, W. & N. R. R.". What I take to be this specimen

NY: New Castle County: Dry hills near Center Station, Aug 1872, W. M. Canby s. n. Originally identified as
P. clinopodioides, this specimen lacks the characteristic hooked pubescence of that species and I have identified it
as P. torrei.

NEW JERSEY FLORA 9]

1980 from open woods on a limestone ridge at Johnsonburg, Warren County, but had
identified it as "P. verticillatum." When | revisited this site, the once open ledge had
succeeded to a shrubby thicket, and I could not find P. clinopodioides. However on 9
October 1992, Tom Breden and I discovered a colony of 67 plants in full sun in a limestone
glade at Sterling Hill Mine, Ogdensburg, Sussex County.

Pycnanthemum torrei

The critical couplets in keys in Gleason (1952), Gleason and Cronquist (1963), and

Gleason and Cronquist (1991) all break on the same variable character: whether or not the

upper surfaces of the leaves and bracts are glabrous or nearly glabrous. Specimens of P.
torrei collected throughout its range at CHRB, NY, and PH, illustrate this variable character.
The upper surface of the stem leaves below the inflorescence of P. torrei are glabrous,
except for occasional scattered hairs. The leaf-like bracts subtending the inflorescence are
commonly minutely hairy over the entire upper surface, or less frequently, with hairs largely
confined to the base of the leaves. In some specimens, such as the isotype (New Jersey:
Mercer County: Princeton Jul 1831, J. Torrey s. n., NY) and Mackenzie’s number 266 (New
Jersey: Essex County: Milburn, 12 Jul 1903, NY), the bracts subtending the inflorescence,
as well as several of the leaves below these, are densely short hairy over the upper surfaces.
The inner faces of the involucre bracts, both the broader outer, and the narrower inner, are
also variable in pubescence. Frequently the inner faces of the bracts are essentially glabrous
as in the isotype, but hairs can be also just on the upper half (Delaware: Kent Co: Felton,
July 1874, W. M. Canby s. n., PH), the top, bottom, and midvein (Virginia: South Hampton
Co.: Near Raccoon Creek, north of Mill Neck Church, 9 Jul 1940, M. L. Fernald and B.
Long 12462, PH), or even hairy throughout, as in the above cited Mackenzie specimen. The
outer faces of the bracts are always hairy, sometimes densely so. Bentham’s (1832) type
description makes no distinction between the inner or outer faces, "Calyces et bracteae
molliter villosi."

Using the keys in the above manuals, some or many specimens (depending on how you
interpret modifiers like "evidently," "nearly," or "few") will run to P. verticillatum. The
most reliable characters by which to distinguish P. torrei from P. verticillatum are the shape
and size of the calyx lobes. In P. torrei, the lobes are lanceolate, commonly attenuate to a
Slender point, and 1 mm to 1.5 mm in length (occasionally a little longer). The calyces of
P. verticillatum are narrowly deltoid and 0.7 mm to 1 mm in length. Depauperate, or
immature specimens of P. clinopodioides can be similar to P. torrei, but these can be readily
identified by the distinctive pubescence of their stems (described previously). The stems of
P. torrei are commonly sparsely hairy on the faces with very short hairs, which are either
nearly perpendicular to the stem or downwardly or upwardly directed, being slightly bent,
mostly at, or above the middle (The short hairs of P. clinopodioides are mostly bent at or
below the middle). Longer hairs occur primarily along the edges of P. torrei’s stem, but
their length and abundance vary, especially towards the western and southern portions of its
range. These more densely hairy phases have been described as either P. leptodon Gray or
P. torrei Benth. var. leptodon (Gray) Boomhour, but as with many of the taxa of
Pycnanthemum, pubescence is quite variable, and shows little geographical or taxonomical
consistency.

The distribution of P. torrei is predominantly eastern, like that of P. clinopodioides, but
extends farther west and south then does that species. Typical material has been collect
or reported from southern New Hampshire (Hodgdon and Steele 1972), southeastern

92 BARTONIA

Connecticut (NY), southeastern New York State (PH; NY), eastern Pennsylvania (NY; PH),
New Jersey (BK; CU; CHRB; GH NY; PH), Delaware (NY; PH), Maryland (Grant and
Epling 1943), District of Columbia (Grant and Epling 1943), southeastern Virginia (NY;
PH); West Virginia (Grant and Epling 1943), North Carolina (PH), South Carolina (Grant
and Epling 1943), to northern Georgia (NY; PH). Fernald (1950) reports it from Ohio and
Arkansas, but these states are not included in Grant and Epling (1943).

Although considered by Grant and Epling (1943) to be a distinctive and fairly uniform
species, there are specimens of P. torreyi from the Midwest (Illinois, NY, Missouri, NY,
Kansas, NY, and Indiana, IND, and occasionally elsewhere (New Jersey, PH, Pennsylvania,
NY, PH), which seem to be intermediate between P. torrei and some of the es narrow-
leaved species. Frequently the calyx lobes of these intermediate specimens + deltoid
shaped (like P. verticillatum or virginianum) but with acuminately to ian prolonged
tips (like P. torrei). Although these specimens will key to P. torrei, they are at best,
atypical. While many of these specimens have been annotated as P. torrei by Grant or
Epling, they may be, as suggested by these authors (1943), of hybrid origin, possibly
involving P. flexuosum, P. virginianum, and P. verticillatum. Clearly more study is needed
to resolve the identity of these problematical specimens.

Pycnanthemum torrei is rare throughout its range with nearly every state Natural Heritage
Program tracking it as a conservation priority (The Nature Conservancy’s Biological and
Conservation Database). The greatest concentration of collections apparently are from the
area of southern New York State, northern New Jersey, and eastern Pennsylvania. At the
herbaria examined for New Jersey specimens there are 25 specimens representing at least 12
distinct localities in nine counties (Passaic, Bergen, Essex, Morris, Hunterdon, Middlesex,
Mercer, Monmouth, Gloucester). All but seven of these specimens were collected during the
latter part of the last century. The most recent collections were both collected in 1931:
Passaic Co.: High Mountain, Haledon, 13 June 1931, H. K. Svenson 4530, GH, NY;
Hunterdon Co.: 1 mi north of Byram, G. A. Loughridge and M. A. Chrysler s. n., CHRB.
On 15 August 1990 I relocated Svenson’s High Mountain site and on 17 July 1991 1
discovered a new site on the Ramapo Mountains, northwest of Darlington, Bergen County.
At both sites P. torrei was growing in open, dry, rocky glades in oak/hickory forests on
slopes near the summits of south facing hillsides. The substrate at the Bergen County site
is granitoid gneiss and at the Passaic County site it is trap rock. Clearly in the northeastern
portion of the range (and apparently elsewhere) P. torrei often grows in substrates rich in
iron and magnesium such is found in mafic formations like trap rock or diabase. Half of the
12 New Jersey locations are on trap rock or diabase, and the species also has been collected
on similar substrates in Connecticut (Les Mehrhoff pers. comm.), New York State,
Pennsylvania, Maryland (Dan Boone pers. comm.), and North Carolina (Alan Weakley pers.
comm.).

Carex amphibola var. amphibola (sensu Fernald)

On 3 July 1991, I found numerous patches of this taxon on the slope of a wooded diabase
hillside, approximately 0.4 mile southeast of Brookville, Hunterdon County. Specimens were
sent to Robert Naczi, who confirmed the identification.

Fernald (1950) recognizes four varieties under Carex amphibola: var. amphibola, vat.
globosa, var. rigida, and var. turgida (= C. grisea). All except var. globosa occur in New
Jersey. The differences between these varieties, seemingly distinct on paper, are sometimes
difficult to apply with consistency in living or dried specimens, particularly between the vars.
amphibola and rigida. Consequentially, many of the New Jersey specimens that I have

NEW JERSEY FLORA 93

examined have not been identified to variety. The distribution of the varieties within the state
is therefore difficult assess at present. In my experience, C. amphibola var. turgida is the
most abundant and widely distributed variety. It is distinctive in its inflated perigynia and
wider leaves (up to 10 mm). It is common in floodplain forests, and is found in most of the
northern counties and at scattered sites on the Inner Coastal Plain. It is especially abundant
along the Delaware River and its tributaries. The var. rigida differs in its scarcely inflated
perigynia and narrower leaf blades (usually under 10 mm). It seems to occur in more upland
habitats than does var. turgida. I have found it with some regularity in the rich limestone
woodlands of Sussex and Warren counties.

Clearly the rarest taxon we have is var. amphibola. Its leaves are typically much more
narrow (usually under 4 mm) and the base of the plant and its rhizomes are reddish purple.
The base and rhizomes of var. rigida are typically brownish, but exceptional specimens,
similar to those I collected from Andover Iron Mine, Sussex County, New Jersey, have deep
red bases (Robert Naczi pers. comm.). According to Robert Naczi (pers. comm.), typical C.
amphibola is known from only one other New Jersey collection: Dry wooded bluff,
Lambertville, Hunterdon County, 28 May 1921, K. K. Mackenzie s.n., NY). My site is
located about 1.5 mile north of Mackenzie’s locality. It was growing in association with C.
hirtifolia and with C. hitchcockiana, another rare member of section Oligocarpae,
particularly in Hunterdon County. Typical C. amphibola reaches the northeastern limit of
its range in New Jersey and adjacent eastern Pennsylvania (Robert Naczi pers. comm.).

Carex formosa

On 7 June 1990 I discovered a small population of C. formosa about 1 mile northeast of
Brighton, Sussex County. The plants grew in an ecotonal area located between a fairly open
wooded seepage swamp and the base of a low, wooded limestone ridge. In the swamp, Acer
rubrum, Fraxinus nigra, Quercus bicolor, Vaccinium corymbosum, and Osmunda
cinnomomea are the characteristic species with Acer saccharum, Ulmus sp., Hamamelis
virginiana, Carpinus caroliniana, Juniperus virginiana, Symplocarpus foetidus, Uvularia
perfoliata, Carex hirtifolia, and Cystopteris bulbifera frequent along its margin. I counted
19 plants with 55 vegetative shoots and nine fruiting culms scattered over an area of several
feet. Additional searches of this site were made on 11 and 12 June 1990 but no new
populations were found.

Carex formosa is primarily northeastern in distribution, ranging from southwestern
Quebec, south through Vermont, New York State, Massachusetts, Connecticut, to New
Jersey, Pennsylvania, and Ohio, then west through southern Ontario, Michigan, Wisconsin,
Minnesota, and reaching its western limit in North Dakota (modified from Fernald 1950;
Gleason and Cronquist 1991). The species is very uncommon to extremely rare throughout
its range, and apparently reaches its greatest abundance in Vermont, where there are 31 total
occurrences, all confirmed extant (Bob Popp pers. comm.).

Carex joorii

Fifty clumps of Carex joorii with a total of 203 fruiting culms were discovered by Andy
Windisch and me, while we searching for intermittent ponds south of the Bennett Bogs
Preserve, Cape May County. The pond we were searching for had been destroyed—converted
into a detention basin—and the woodland stream draining into had been ditched. A series of
wet swales adjacent to stream had also been ditched. It was in the ditch through the former
swales that we found C. joorii on 10 October 1991. The surrounding woodland is dominated
by Acer rubrum, Liquidambar styraciflua, Nyssa sylvatica, and Quercus phellos, with

94 BARTONIA

Clethra alnifolia, Ilex opaca, Aronia arbutiflora, and Vaccinium corymbosum _ the
conspicuous understory species. Associated herbs included Dulichium arundinaceum,
Scirpus cyperinus, Proserpinaca palustris, and Triadenum virginicum.

This large and attractive sedge is an addition to New Jersey’s flora and is the only
member of section Pendulinae known to occur in the state. It is a southern sedge that is
apparently extending its range northward along the Coastal Plain. Tatnall (1946) cited four
Maryland locations for the Delmarva peninsula, and Hirst (1983) reported a fifth Maryland
location on Delmarva. In 1984, Hirst (pers. comm.) discovered C. joorii in Delaware, and
currently there are six extant occurrences in Delaware (Keith Clancy pers. comm.). If C.
joorii is a relative newcomer in New Jersey, it would explain why such a conspicuous sedge
has not been previously discovered in an area so well botanized as Bennett Bogs.

Carex louisianica

On 6 September 1991, I found a small colony of Carex louisianica growing in Great Piece
Meadows on the north side of the Passaic River about 0.7 mile northwest of Two Bridges,
Morris County. W. D. Miller collected C. /ouisianica at its only other documented New
Jersey location at Islin, Middlesex County (15 Jul 1916, 629, CHRB, NY, PH). Miller’s
locality is on the southernmost edge of the Piedmont, just north of the Coastal Plain, and
immediately north of the terminal moraine (Wisconsin Stage). It was last collected there in
1918 by Mackenzie (28 Jul 1918, s.n., NY). This site appears to have been destroyed by
the intensive urbanization that has taken place in this part of central New Jersey.

Carex louisianica is of southern affinities occurring predominantly on the Atlantic Coastal
Plain from Maryland, south to Florida, then west to Texas, and northward in the Mississippi
drainage to Illinois, Indiana, and Missouri (Gleason and Cronquist 1991, Goodfrey an
Wooten 1979). It is disjunct in glaciated northeastern Ohio (Braun 1967; T. Reznicek pers.
comm.) and northern New Jersey, being apparently absent (or at least, undocumented) from
the rest of Ohio, Delaware, and the southern half of New Jersey. The Two Rivers, New
Jersey population is about 25 miles north Miller’s Islin site, and is nearly 200 miles
northwest of the northernmost Maryland record in Harford County (Gene Cooley pers
comm. ).

The colony appears to have been derived from a single clone, growing in a patch about
five by five feet. I counted thirty-five fruiting culms and many more vegetative plants. The
habitat is a broad floodplain forest dominated by an open canopy of Quercus palustris,
Liquidambar styraciflua, Fraxinus americana, and Acer rubrum. Carex lupilina and C.
typhina grew immediately adjacent to C. louisianica, with C. typhina forming extensive
colonies with thousands of plants, scattered throughout the flood plain. Isolated clumps of
C. grayi grew here and there. Nearby was large linear swale dominated by Zizania aquatica
and Bidens laevis.

These woods are reminiscent of many Coastal Plain lowland forests that I’ve been to in
southern New Jersey. Places like Riddleton and Muttontown Woods in Salem County,
Lannings Wharf in Cumberland County, or even the Pigeon Swamp in Middlesex County.
These are places that I have looked for C. /ouisianica, and these are places that I’d expect
to find it—not along the Passaic River in the ancient remains of Glacial Lake Passaic.

ACKNOWLEDGMENTS

I thank curators at BK, CHRB, CU, GH, IND, NY, and PH, particularly Patricia
Holmgren, Lewis Johnson, Janice Meyer, and Emie Schuyler for arranging loans. The late

NEW JERSEY FLORA 95

Arthur Cronquist, Bob Kral, Rob Naczi, and Tony Reznicek confirmed some of my
identifications. David Fairbrothers made available some difficult to obtain references. J

artesz helped with several nomenclatural matters. Larry Morse and Christa Russell
provided information from The Nature Conservancy’s central databases. Tom Breden, Leo
Bruederle, Roger Johnson, Jim Stasz, and Howard Wildman transcribed much of the herbaria
label data. Joe Arsenault, Dan Boone, the late Gil Cavileer, Keith Clancy, Steve Clemants,
Gene Cooley, Tom Cooperrider, Allison Cusick, Andy Cutko, Stephen Field, Ted Gordon,
Tom Halliwell, Frank Hirst, Mike Homoya, David Hunt, John Kunsman, Les Mehrhoff,
Richard Micthell, Mike Penskar, Bob Popp, Rick Radis, Bruce Sorrie, and Alan Weakley all
helped with various aspects of this paper.

LITERATURE CITED

ABRAITYS, V. A. 1957-1983. Unpublished field notes on New Jersey plants.

ANDERSON, K. 1989. A check list of the plants of New Jersey. New Jersey Audubon, Rancocas Nature Center,
Mount Holly.

BASSETT, I. J., C. W. COMPTON, J. MCNEILL, P. M. TASCHEREAU. 1983. The genus Atriplex (Chenopodiaceae)
in Canada. Agr. Canada Monogr. 3

BENTHAM, G. 1832-1836. Labiatarum genera et species. James Ridgeway and Sons, London.

BISSELL, ip H. 1903. A new station for Dentaria maxima. Rhodora 5: 168-169.

B , E. L. 1967. The vascular flora of Ohio, vol. 1: The nacigeat gba Ohio State Univ. min Columbus.
BRITTON, me L. 1889. Catalogue of plants found i in New Jersey. Geol. Surv. N. J., Final Rep. State Geol. 2: or
1891. pita or noteworthy North American Phanerograms--IV. "Ba ll. Torrey Bot. Club - 265-27.

BRITTON, N. L. AND A. BROWN. 1913. An renee flora of the United States, Canada and the British et

Charles aa s Sons, New York.
WHA AND R. J. CRAIG. 1976. Rare and gs species of Connecticut = their habitats. Rep. of
investigations 6, State Geol. Natural Hist. sie CE; of Environmental Protect
AMES, E. H. 1903. The Dentarias th cage cut. Sout 5: 213-219.
FAIRBROTHERS, D. E. AND M. Y. Hou “973 pare 1975). Rare or endangered vascular plants of New Jersey.
Science Notes No. 14, N. J. State enton.
wie ng piss L. 1936. Plants from the outer are Plain of il as re 38: 414-452.
aie Gray’s manual of botany. 8th ed. American Book Co. New Yor
a L. GRISCOM. 1935. Three days of botanizing in pacha Pera Rhodora 37: 167-189.
eso - A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent
anada. Hafner Press, New York.
AND A eye a 1963. Manual of vascular plants of northeastern United States and adjacent Canada. D.
Van Nostrand Co., New York.
. 1991. Manual a vascular plants of northeastern United States and adjacent Canada. 2nd ed. The N. Y. Bot.
Gard., New York.
GODFREY, R. K. AND J. W. WOOTEN. — os and wetland plants of southeastern United States,
Monocotyledons. The Univ. of Georgia, Ath
GRAHAM, S. A. 1985. A revision of Ammannia (Lstlencene) i in the Western Hemisphere. J. Arnold Arbor. 66: 395-
9.

41

GRANT, E. AND C. EPLING. 1943. A study of Pycnanthemum. Univ. Calif. Publ. Bot. 20: saagea

GRAY, A. 1848. A manual of the a of the northern United States. James Monroe and Co.,

HARSHBERGER, J. W. AND V. G. BURNS. 1919. The vegetation of the Hackensack Marsh: a — neem fen.
Trans. Wagner Free Inst. Sci. corer 9: 1-35.

HARVILL, A. M., JR. T. R. BRADELY, C. E. STEVENS, T. F. WIEBOLT, D. M. E. WARE, AND D. W. OGLE. 1986.
Atlas of the Virginia Flora, 2nd ed. Virginia Botanical Associates, Farmville.

— C. J. 1979. Vegetational history of the New Jersey Pine Barrens. Jn R. T. T. Forman, ed. Pine Barrens:

Osytem and landscape. Academic Press, New York.

bine F. 1983. Field cps on the Delmarva flora. Bartonia 49: 59-68.

Hopapon, A. R. AND F. L. STEELE. 1972. Recent a in the New Hampshire flora. Rhodora 74: 291-294.

HopkKINs, M. 1942. Cercis in North America. Rhodor: -211.

HOuGH, M. Y. 1983. New Jersey wild plants. sass eee Harmony.

96 BARTONIA

Hunt, D. 1990. A greaang review of Quercus series Laurifolia, Marylandicae, and Nigrae. M.S. dissertation,
Univ. of Georgia, Athen

JAMES, C. W. 1956. A revision of Rhexia (Melastomataceae). Brittonia 8: 201-230.

KARTESZ, J. T. AND R. KARTESZ. 1980. A synonymized checklist of the vascular flora of the United States, Canada,
and Greenland. Univ. of sani Carolina Press, Chapel Hill.

. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Timber

Press, Portland.

KNIESKERN, P. D. 1856. A catalogue of plants growing without cultivation in the counties of Monmouth and Ocean.
Ann. Rep. Geol. Surv. N.J. Suppl., Trenton.

KRAL, R. AND P. E. BOSTICK. 1969. The genus Rhexia (Melastomataceae) Sida 3: 387-440.

MITCHELL, R. S. 1986. A checklist of New York State plants. N. Y. State Mus. Bull. 458.

MONTGOMERY, F. H. 1955. Preliminary studies in the genus Dentaria in eastern United States. Rhodora 57: 161-

173.

NICHOLS, G. E. 1923. Cercis oe in Connecticut. Rhodora 25: 203-204.

PIELOU, E.C. 1991. After the age: the return of life to glaciated North America. Univ. Chicago Press, ann

RADFORD, A. E., H. E AML, AND C. R. BELL. 1968. Manual of the vascular flora of the Carolinas. Univ. of North
Carolina Press, Chapel H

REEDER, J. A. 1951. Lite note on Cercis canadensis in Connecticut. Rhodora 53: 246-247.

REHDER, A. 1910. A new hybrid Cornus (Cornus rugosa x stolonifera). Rhodora 12: 121-124.

RICKETT, H. W. 1934. Cornus amomum and Cornus candidissima. Rhodora 36: 269-274.

. 1945. Cicaeen. North American Flora 28B: 299-311.

ROGERS, C. M. 1963. Yellow flowered species of Linum in eastern North America. Brittonia 15: 97-122.

SCHMID, J. A. 1987. Atlantic white cedar in the Hackensack Meadowlands, New Jersey: its historic extirpation and
future a, In A. D. Laderman, ed. Atlantic white cedar wetlands. Westview Press, Boulder.

mh w Jersey higher plants. vol. I: desk manual. Schmid and Compan

SARGENT, c s. 1922, Manual of the trees of North America. Houghton Mifflin Co., —

ahaa W. S. 1972. The past and present flora of the Hackensack Marshes. Bartonia 41: 456

SNYDER, D. B. 1993. Extinct, extant, exitipated, or isannet? Batons 37 aries 50-5

ge V.E. VIVIAN. 1981.R vascular p p ew Jersey. ai and Wildlife Service,

U. S. Dept. of the Interior, Washington.

ina W. 1912. The plants of southern New Jersey, with especial reference to the flora of the Pine Barrens. Ann.
Rep. N. J. eo Mus., 1910, part II: 21-828.

TASCHEREAU, P. M. 1972. Taxonomy and distribution of Atriplex species in Nova Scotia. Canad. J. Bot. 50: 1571-

4.

1
TATNALL, R. R. 1946. Flora of Delaware and the Eastern Shore. The Society of Natural History of Delaware,
Wilm

ington.
TAYLOR, = 1915. Flora in the vicinity of New York. Mem. N. Y. Bot. Gard. vol. 5.
TorREY, J. 1819. A catalogue of plants growing spontaneously within thirty miles of the city of New York. Lyceum
of Natural History of N. Y., Alban

any
i ri G. 1985. Michigan Flora: Part II. Cranbrook Inst. Sci. Bull. 59 and Univ. Michigan Herbarium, Ann

as a A. ape A preliminary study of = 4 eee in North America. Bartonia 27: 1-46.
WHERRY, E. T., J. M. FoGG, JR., AND H. A. WAHL. 1979. Atlas of the flora of Pennsylvania. The Morris
Arboretum, hadi,
sui a - 965. Mea three taxonomic complexes of the genus Cornus in the eastern United States.
Trans. as Acad. Sci. 67: 747-817.
WOLFE, P. 5 1977 The ast and landscapes of New Jersey. Crane, Russak and Co., New York.

Bartonia No. 58: 97-100, 1994
The Vegetation of Little Beach Island, New Jersey

RICHARD STALTER
Department of Biological Sciences, St. John’s University, 8000 Utopia Parkway Jamaica, NY 11439

Little Beach Island, Atlantic County, New Jersey, is comprised of approximately 283 ha,
and is located in southern New Jersey just north of Atlantic City at 39° 28’ N, 74° 34’ W.
It is part of the Edwin B. Forsyth National Wildlife Refuge and includes land known as Dog,
Pullen and Little Beach islands. Human habitation of the island during the 20th Century was
limited to a fishing village and Coast Guard Station. The village was abandoned around
1940 and the Coast Guard Station was decommissioned in 1942.

Little Beach Island is among the most pristine coastal islands in maritime New Jersey.
Since this island has been uninhabited for nearly fifty years, and rarely visited, it may serve
as a bench mark to measure the effect of man’s impact on barrier islands in southern New
Jersey. The objective of the present study is to obtain a record of the vegetation of Little
Beach Island.

Beach erosion is especially severe on the western and central portion of the island. Coastal
storms erode the protective primary dunes and deposit sand on the inland salt marshes,
especially on the western and southern portions of the island. After storm disturbances, the
dunes are rebuilt by natural processes and are colonized and stabilized by Ammophila
breviligulata.

METHODS

The vegetation of Little Beach Island was sampled once every three weeks beginning May
3, 1989, terminating on November 6, 1989. Herbarium voucher specimens of each taxon
were prepared and deposited at Edwin B. Forsyth National Wildlife Refuge. The species
checklist contains an inventory of the vascular plants of Little Beach Island including native
taxa along with naturalized and adventive weeds. All non native species are marked by an
asterisk (*). The checklist is divided into three categories: Pinophyta, Magnoliophyta:
Magnoliopsida, and Magnoliophyta: Liliopsida. Families and species are alphabetical.
Nomenclature follows that of Gleason and Cronquist (1991).

RESULTS

The current vascular flora of Little Beach Island consists of 33 families, 74 genera and
84 species of which 69 are native (Table 1). The Asteraceae with 14 genera and 14 species
and Poaceae with 12 genera and 13 species are the largest families. Together they comprise
35% of all genera and 32% of all species. Other large families are the Chenopodiaceae (9
species) and Rosaceae (6 species). Eighteen percent (15 species) of the flora are not native
to the United States (Table 1).

97

98 BARTONIA

Table 1. A statistical summary of the vascular flora of Little Beach Island, New Jersey.

Pinophyta Magnoliopsida Liliopsida Total

Families 26 6 33

1 54 19 74

Species l 59 24 84

Native species 1 45 Za 69

Introduced species 0 14 l 15
DISCUSSION

The vegetation of Little Beach Island can be classified into three general plant
communities: dune community, salt marsh, and immature maritime forest.

DUNE COMMUNITY. Drift and areas of occasional overwash are present on the northern
portion of the island. This area is sparsely vegetated by annual plant species most notably
Salsola kali, Cakile edentula, Sesuvium maritimum, Atriplex arenaria, and Chamaesyce
polygonifolia. Characteristic perennials include Honkenya peploides ssp. robusta and
Solidago sempervirens. Slightly higher rarely flooded portions of the beach are vegetated
by the same species and by Ammophila breviligulata, Panicum amarum, and Solidago
sempervirens.

The ocean and bay bordering dunes at Little Beach Island are currently undergoing severe
erosion. These dunes are currently populated by Ammophila breviligulata, Lechea maritima,
and Solidago sempervirens. The sheltered landward side of these dunes is generally
vegetated by the same species along with Toxicodendron radicans, Myrica pensylvanica, and
Prunus serotina, On sheltered inland dunes in the northeast portion of the island Hudsonia
tomentosa, Schizachyrium scoparium, Lechea maritima, Polygonella articulata, Toxicoden-
dron radicans, Myrica pensylvanica, and Prunus serotina thrive. The habitat supporting
Hudsonia is extremely small covering only a few hectares.

MARITIME FOREST COMMUNITY. The maritime forest at Little Beach Island consists of an
almost pure stand of Prunus serotina, which reaches its best development on the large inland
stable dunes. These dunes have been in existance for 300 years (Beall personal communica-
tion). Prunus serotina, the dominant tree on the secondary dunes at Little Beach Island, has
been recognized as a successional species at Island Beach State Park, New Jersey (Martin
1959), at Sandy Hook, New Jersey (Stalter 1974), and at Fire Island, New York (Art 1976;
Schulte 1965; and Stalter et al. 1986). The trees near the summit of the dunes are generally
short and pruned by salt spray; those occupying the sheltered sides of the dunes where salt
spray deposition is minimal are taller, some as large as 10 meters tall. The dune soils are
well drained and consist of fine sand with a slight accumulation of organic matter. There
are occasional individuals of Amelanchier canadensis and Juniperus virginiana. Ilex opaca
is extremely rare; only 7 individuals were observed on the whole island during the growing
season of 1989.

Ilex opaca is the dominant tree at the Bayside Holly Forest, Sandy Hook, New Jersey
(Stalter 1979) and at the Sunken Forest Fire Island New York (Art 1976; Stalter et al. 1986).
The scarcity of American holly at Little Beach Island may be due to the young age of the
large stable inland sand dunes as Art (1976) concluded that Jlex opaca is the dominant

LITTLE BEACH ISLAND 99

species on portions of Fire Island that have been undisturbed for 200 to 300 years.

SALT MARSH COMMUNITY. The salt marsh community at Little Beach Island occurs along
the sheltered inland portions of the island. The vegetation of the low salt marsh is
dominated by Spartina alterniflora. The higher infrequently flooded salt marsh is populated
by Distichlis spicata, Spartina patens, Salicornia virginica, S. europaea, Spartina
alterniflora, Plantago maritima, Aster tenuifolius, and Limonium carolinianum. At the upper
marsh boundary, /va frutescens is generally dominant. Baccharis halimifolia forms the
interface between salt marsh and upland vegetation.

The rarest plants observed at Little Beach Island are Plantago maritima, Sesuvium
maritimum, and Honkenya peploides. Their rankings on the New Jersey Natural Heritage
Program list (Synder 1992) are: Sesuvium maritimum — S2 (imperiled in the state), Honkenya
peploides — S2 (imperiled in the state), and Plantago maritima — S2 (imperiled in the state).
Honkenya is the dominant plant on the overwash portions of northeast Little Beach Island.
Several hundred thousand Plantago maritima plants occupy the upper infrequently flooded
salt marsh in the southwest portion of the island.

The vegetation of Little Beach Island is similar to that found on other barrier islands in
New Jersey especially in the salt marsh and dune communities (Martin 1959). Species
diversity here is not as great as that at Island Beach State Park, NJ (Martin 1959), Sandy
Hook, NJ (Stalter 1979), Cape May Point State Park, NJ (Sutton et al. 1990), or Fire Island,
NY (Stalter et al. 1986), although there is a greater percentage of native taxa here than at the
aforementioned areas. The high percentage of native species at Little Beach Island is
probably the result of its isolation with respect to man’s activities.

CHECKLIST

PINOPHYTA
CUPRESSACEAE: Juniperus virginiana.

MAGNOLIOPHYTA —MAGNOLIOPSIDA
AIZOACEAE: Sesuvium maritimum (S2).
ANACARDIACEAE: Rhus copallinum; Toxicodendron radicans.
OCYNACEAE: Apocynum sep eerie
eas Ilex opaca.
RACEAE: Achillea millefolium*; Ambrosia artemisiifolia; Aster tenuifolius, Baccharis sion Conyza
reich Erigeron strigosus, Eupatorium sity Gnaphalium obtusifolium, Hypoc chlor Iva
Srutescens; Lactuca canadensis; Solidago sempervirens; Taraxacum officinale*,; Xanthium strumari

is.
ARYOPHYLLACEAE: Honkenya peploides (S2); Spergularia mariana, Stellaria media*.
CHENOPODIACEAE: Atriplex arenaria, patula; Bassia hirsuta; Chenopodium album*, ambrosiodes*, Salicornia
europaea, virginica; Salsola kali*; Suaeda linearis.
ISTACEAE: Hudsonia tomentosa; Lechea maritima.
CONVOLVULACEAE: Calystegia sepium.
EUPHORBIACEAE Papuan cuneate

PHYTOLACCACEAE: Phytolacca americana
PLANTAGINACEAE: Plantago maritima ssp. _juncoides (S82).
PLUMBAGINACEAE: Limonium carolinianum.

100 BARTONIA

POLYGONACEAE: Polygonum cespitosum*; Rumex acetosella*, crispus

ROSACEAE: Amelanchier canadensis; Prunus serotina; Rosa ba eng a allegheniensis, flagellaris.
RUBIACEAE: Galium aparin

SCROPHULARIACEAE: hast vulgaris*, Verbascum thapsus*.

SOLANACEAE: Solanum nigrum.

VITACEAE: seen quinquefolia.

MAGNOLIOPHYTA —LILIOPSIDA

CYPERACEAE: Carex poe San polystachyos, silicea; Cyperus strigosus; Scirpus pungens, robustus.

JUNCACEAE: Juncu.

POACEAE: pone “rv Cenchrus tribuloides; Digitaria sanguinalis*; Distichlis spicata; Elytrigia
repens; Eragrostis spectabilis; Fes. rubra; Panicum lancearium, virgatum; Phragmites australis; Schizachyrium
scoparium; Spartina alterniflora, mod Triplasis pupurea.

RUPPIACEAE: Ruppia maritima.

oe Smilax pone ifolia.

TERACEAE: Zostera marina.

ACKNOWLEDGEMENTS

I thank David Beall, Refuge Manager, for permitting me to collect and identify the
vascular plants at Little Beach Island, for providing information on the history of the island,
and for providing access to the Ranger office; Bob Lamoy, Barbara Hudac, and Mark Steffen
for providing me with boat transportation to and from Little Beach Island; Robert C. Meyer
for assistance in identifying grasses; Chris Bae and Donato Tramontozzi for assistance in
preparing herbarium specimens; Karl Anderson for providing information on rare an
endangered plant species in New Jersey; Karl Anderson and Eric Lamont for critically
reviewing the manuscript; and to St. John’s University for purchasing herbarium supplies.

LITERATURE CITED
ArT, H. W. 1976. Ecological studies of the Sunken Forest, Fire Island National Seashore New York. National

be

GLEASON, H. A. AND A. QUIST. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent
Canada. 2nd on New York Botanical Garden, B

MARTIN, W. E. 1959. The vegetation of Island Beach ee Park, New Jersey. Ecol. Monogr. 29:

SCHULTE, E. 1965. A study of the plants in the Sunken Forest, Fire Island, New York. Master’s cant e W. Post

ollege.

SNYDER, D. B. 1992. Special plants of New Jersey. New Jersey Department of Environmental Protection and
Energy, Trenton

STALTER, R. 1979. Some ecological observations on Jlex forest, Sandy Hook, New Jersey. Castanea 44: 202-207.

. LA

STALTER, R., E. E. LAMONT, AND J. NORTHUP. 1986. Vegetation of Fire Island, New York. Bull. Torrey Bot.
Club 113: 298-306.
SUTTON, P., R. MEYER AND R. STALTER. 1990. The vascular plants of Cape May Point State Park, Cape May

County, Mew Jersey. a Torrey Bot. Club 117: 294-300.

Bartonia No. 58: 101-113, 1994

The Vegetation of the Glades Region,
Cumberland County, New Jersey

STEVENS HECKSCHER
Natural Lands Trust, Inc., 10 Ridley Drive, Wallingford, PA 19086

Since about 1964 the Natural Lands Trust has been acquiring land in Downe Township,
Cumberland County, New Jersey, for inclusion in the Fortescue Glades Refuge and the
Florence Jones Reineman Wildlife Sanctuary, the former a property of the Natural Lands
Trust and the latter a property of the Erdman Trust under management by the Natural Lands
Trust. These privately-owned and managed wildlife refuges together comprise an area of
approximately 2,000 hectares (approximately 5,000 acres). With ongoing acquisition of
contiguous and nearby natural areas by the Natural Lands Trust, the Fortescue Glades Refuge
is continuing to increase in size.

The region covered by the present study is an area of 72.6 square kilometers, lying to the
east and northeast of the Delaware Bayshore town of Fortescue, and including but not
limited to the Fortescue Glades Refuge and the Reineman Sanctuary. It is referred to as the
"Glades Region," and is the total shaded area (in either dark or light gray) in Fig. 1.
Excluded from the study area are all agricultural and cultural areas, including farm fields
currently in use, lawns, built-up industrial, residential, and business areas, and barren piles
of mining spoils. However, railroad beds throughout our area as defined by the map are to
be included in our study area. Plants known to have been planted in our area for any
purposes, such as ornamentation, restoration, or dune-stabilization, and not occurring
naturally, e.g. Pinus strobus which has been planted extensively around mining pits, will not
be further considered in this study.

On current USGS maps, Bear Swamp is not correctly indicated. The section marked
"Bear Swamp" is a portion of that which is now known as Bear Swamp West. As
understood by residents of the area and my colleagues, Bear Swamp West is the entire
section bounded on the south by Methodist Road (County Route 553), a line from Watson’s
Corner to Frames Corner on the west, Baptist Road and the Central Railroad of New Jersey
on the north, and a line from Paynter’s Crossing to Turkey Point Corner on the east; Bear
Swamp West thus lies entirely within our study area, the Glades Region.

To the east of the Glades Region lies the other half of Bear Swamp, known as Bear
Swamp East, now constituting the Bear Swamp East Natural Area, held by the state of New
Jersey, and lying, roughly, east of Dividing Creek Road (County Route 555), south of the
Central Railroad of New Jersey, west of Whitehead Road, and north of Haleyville Road.
Plant communities of Bear Swamp East (for details see Cartica et al. 1988) are similar in
most respects to those of Bear Swamp West in the Glades Region. In particular, Bear
Swamp East contains an old-growth forest similar in size and species composition to the one
in Bear Swamp West but differs from the latter by having a stand of immense trees of
Liriodendron tulipifera. \ know of only one such tree in Bear Swamp West. Moreover,
before the gypsy moth kill of the early- to mid-1980s, a number of sizeable trees of Quercus
michauxii (most or all now dead) had been located in Bear Swamp East. Less sizeable

101

102 BARTONIA

To
Philadelphia o

k%& Bridgeton

Greenwich

Area of
Vegetation Map
Coverage

“— Strand
ff Community

Fig. 1. Location of the Glades Region.
individuals of Q. michauxii were found by myself and others in Bear Swamp West before

In all plant listings in this paper, names of plants considered by Moore (1989) as not
native to Cumberland County will be enclosed in brackets. Nomenclature follows that of
Kartesz and Kartesz (1980), except in a very small number of cases. When a name differing
from that given in Kartesz and Kartesz is used, the name according to those authors is listed
parenthetically.

This study is a qualitative report on a diverse area whose vascular flora has not been the
subject of thorough systematic botanical exploration. Especially in the old-growth forest
there is need for extensive quantitative studies.

METHODS

I began exploration of the Glades Region in the autumn of 1980, and continued it through
1990. Scattered sites at many points were visited, and their exact locations recorded on
USGS maps. At these sites plant species present, especially woody plants, were noted.
Specimens of unknown plants were taken for identification and usually deposited in the
Herbarium of the Academy of Natural Sciences, Philadelphia, and individual plants
belonging to species new to the study or new to the immediate vicinity were photographed.
I refer to these sites as "plant data sites." All information for each plant data site was
entered in a database, using the dBASE II system: date, site number, species name for each
species present, family, and whether or not that species was photographed at that site. At

VEGETATION IN GLADES REGION 103

a later time the soil type for that site as determined from the county soil survey (U.S.
Department of Agriculture 1978) was also entered.

A computer program was written on this system, enabling me to retrieve the stored data
in a manner most useful for vegetation mapping, which was begun in the summer of 1989.
Two aerial photographs of the region were used for remote sensing, a 1:400 black-and-white,
taken in 1987, and an infrared aerial photograph, obtained from the U.S. Geological Survey
(NNAP-80 series, 1:24,000 scale paper print, Wilmington Quadrangle, #441-44, photo-
graphed April 1981). The latter was by far the more useful of the two aerial photographs,
because on the black-and-white aerial photograph differences among various plant
communities were scarcely evident, while on the infrared each community had a distinctive
signature.

The procedure used in constructing the vegetation map accompanying this article was
basically that outlined by Anderson, Wentz, and Treadwell (1980). Beginning with the
broadleaf swamp forest, a site was assigned to mature or old-growth portions of it if Acer
rubrum, Nyssa sylvatica, and Magnolia virginiana were all present. The additional presence
of Liquidambar styraciflua or Osmunda cinnamomea or both was regarded as confirmation
of the occurrence of this forest type. Moreover, the presence of immense trees in the old-
growth area, which I had explored on foot in earlier years, was readily discernible on the
infrared, which in many cases clearly showed individual trees, so that old-growth forest
could be distinguished from secondary mature forest. Points on the infrared aerial
corresponding to plant data sites in these two communities were examined until I could
recognize these communities elsewhere on the infrared.

Each of these communities (mature secondary, and old-growth, broadleaf swamp forest)
possessed a characteristic appearance, or "signature," which was recognizable elsewhere on
the infrared photograph. Species occurring at plant data sites located within these signature
areas were checked (using the database and sometimes photographs taken at the site) and in
every case I was able to confirm that the sites were located within the predicted community
type. The boundaries of these two communities were plotted on the vegetation map, as
indicated on the infrared. Ground truthing in the form of onsite checks at several easily-
accessible locations confirmed that the community types were correctly determined.

This procedure was applied with variable success to each of the remaining plant
communities which had been identified in the previous decade’s field work. One difficulty
was that spatial transitions between communities were often broad and gradual, particularly
between hydric and mesic forests. On the infrared photograph, sharply definable boundaries
between these two communities were almost always unrecognizable, and data from field
observations seldom provided confident discrimination. For this reason it was decided to add
to the vegetation map a category, ‘ransition zone, to be applied where sharp boundaries
between one community and another were indeterminable. In some cases soil data from the
Cumberland County Soil Survey (U.S. Department of Agriculture 1978) were used to
discriminate between hydric and mesic plant communities, particularly if the infrared map
or plant data collected on the ground corroborated such discriminations. (Wet areas usually
had a purple or blue cast on the infrared aerial photograph.) In attempting to draw such
boundaries on the vegetation map, the following rules were helpful: (1) Muck soils, of the
Muck-Atsion-Berryland association, were taken as supporting a predominantly hydric
community unless the infrared photograph or ground vegetation data showed that artificial
drainage had led to the replacement, on muck, of a mesic community or regime. (2) Soils
of the Hammonton-Fallsington-Pocomoke association, which the Soil Survey describes as

104 BARTONIA

"moderately well drained to very poorly drained, loamy and sandy soils; on uplands and
lowlands," were taken as supporting a predominantly mesic community at a particular point
if corroborating evidence for a mesic regime was present in the form of a reddish cast on the
infrared, at that point, and provided that plant species recorded at that point, if any, were
consistent with such a determination. Frequently a discrimination between predominantly
hydric and predominantly mesic communities was difficult or impossible, and "transition
zones" were used on the vegetation mai

Another difficulty which was encountered in the construction of the vegetation map was
an initial inability reliably to recognize pine-dominated patches on the infrared aerial
photograph. A late-winter survey by automobile along back roads in the study area revealed
that I had mistaken pine-dominated stands for hardwood stands. The locations of a number
of pine-dominated stands were noted and compared with the appearances of these stands on
the infrared, with the result that I could recognize the signature of such stands on the
infrared. After these were mapped, I inspected some of them from an airplane and from the
ground. The result was a high degree of confidence that the vegetation map displays the
pine-dominated stands with high accuracy.

In naming the plant communities, I have tried to follow as closely as possible the
nomenclature used by McCormick (1979) for the New Jersey Pine Barrens. A number of
communities identified in the course of this study do not occur in the Pine Barrens and for
these I have tried to employ names in common usage. During the preparation of this paper
a preliminary classification of New Jersey plant communities by Thomas F. Breden (1989)
appeared, and I have wherever possible inserted into the following text Breden’s names for
the communities under discussion. In many cases his names are similar to, if not identical
with, the names I use and display on the accompanying vegetation map. A list of plants
from the study area is being maintained and I invite readers to let me know what they find
there so I can make the list more comprehensive and eventually suitable for publication.

This paper and its accompanying vegetation map describe qualitatively a rich and complex
mosaic of community types. Because of the complexity of this system, this article is
presented as a set of working hypotheses, proposing a classification of the plant communities
found to be present, and including a report on their structure, compositions, and distributions.
It is hoped that further work in the Glades Region will result in the augmentation,
refinement, and modification of the results here presented.

DESCRIPTION OF COMMUNITIES

The Glades Region consists of two landscape types, tidal wetlands and a forested mosaic.
The first contains bayshore beach and tidal marsh; the second contains two complexes,
hydric or lowland forest and mesic or upland forest. Within these two forest complexes,
various types of non-tidal wetlands are present.

The following plant communities have been identified in the Glades Region and are shown
on the accompanying vegetation map
I. Tidal wetlands landscape type: (1) Strand community. (2) Tidal marsh, including tidal
creeks and ponds.

Il. Forested mosaic landscape type, hydric forest complex: (3) Non-tidal freshwater marsh.
(4) Non-tidal freshwater pond. (5) Herbaceous wetland primary succession. (6) Early
broadleaf swamp forest. (7) Pitch pine lowland (hydric) forest. (8) Broadleaf swamp forest,
including some old-growth.

Ill. Forested mosaic landscape type, mesic forest complex: (9) Early upland (mesic)

VEGETATION IN GLADES REGION 105

broadleaf forest. (10) Upland ona pine forest. (11) Upland (mesic) oak forest. (12)
Beech monodominant forest. (13) Heath-shrub community.

(1) STRAND COMMUNITY. Because ee limitations in space, the strand community or sand
beach community is not shown on the large vegetation map. However, it is indicated on the
accompanying location map. It extends from intertidal beach, over shifting dunes (which
may be covered by extremely high tides or storm washover), to the edge of the saltmarsh.

The dominant plant of the strand community is Ammophila breviligulata, which covers
and stabilizes the dunes. This grass is native to the area and may also have been planted in
places to control erosion. Other important members of the strand community are Amorpha
fruticosa, Baccharis halimifolia, Cakile edentula, Cenchrus tribuloides, Chamaesyce
polygonifolia, Cyperus esculentus, Iva frutescens, Juniperus virginiana, Myrica cerifera,
Phragmites australis, Rhus copallina, [Salsola kali], Solidago sempervirens, Spartina
alterniflora, and Toxicodendron radicans.

With the slow rise in sea-level the beach is gradually being pushed inland into the tidal
marsh, and the strand community stands on what is known as washover barrier beach. In
places, stands of Spartina alterniflora may extend into the bay and from year to year one can
observe their rapid diminution and eventual disappearance. Between the dunes and the
Shoreline, there are places where beach erosion has exposed the long-dead roots of Iva
frutescens, a shrub which occurs on the boundary between the tidal marsh and the beach, and
which sometimes persists among the advancing dunes, marking the line where, years earlier,
the beach ended and the marsh began

Breden (1989) does not appear to list an estuarine (as opposed to marine) strand
community, but in floristic composition our strand community appears close to his marine
sand beach commu

(2) TIDAL MARSH, CREEKS, AND PONDS. Approximately sixty percent of the Glades
Region is covered by tidal marsh, including salt or brackish creeks and ponds. Distichlis
spicata, Spartina alterniflora, and S. patens are the dominant plants of the tidal marsh. S.
alterniflora occurs along the edges of creeks and ponds, where it occurs with high vigor or
robustness, and also farther from open water, where it is less robust. Limonium carolini-
anum and Aster tenuifolius are conspicuous when flowering in the open salt marsh.
Baccharis halimifolia, Hibiscus moscheutos, Iva frutescens, Juniperus virginiana, and
Phragmites australis are also important components of the tidal marsh, but occur mainly or
exclusively on the boundaries between the marsh and the strand community or between the
marsh and the forest.

Phragmites australis requires moist, ecologically disturbed areas free from flooding, where
it may become an aggressive pest. Its occurrence in the Glades Region is probably limited
to areas of the tidal marsh which have a history of ditching, dumping, digging, or landfill,
or which have been compromised by anthropogenic disturbance to contiguous upland.
Whether it is spreading into little-disturbed or pristine portions of the tidal marsh is not
immediately apparent.

All tidal marshes within our study area belong to Breden’s (1989) salt marsh complex,
except that detailed studies may show that tidal marsh north of Route 553, where Oranoaken
Creek flows out of Bear Swamp West, belongs to his brackish tidal marsh complex.

(3) NON-TIDAL FRESHWATER MARSH. In the western portion of Bear Swamp West, due
north and northwest of Beaver Dam, there are several small fragments of freshwater marsh.
There is historical evidence that portions of this section once were under cultivation as
cranberry bogs (Daniel O’Connor pers. comm.), and the cranberries which occur there now

106 BARTONIA

may be in part descended from cultivated plants. These marshes are now remote and
inaccessible. I visited one of them on 16 October 1983 and found Chamaedaphne
calyculata, Cladium mariscoides, Decodon verticillatus, Dulichium arundinaceum, Juncus
canadensis, Proserpinaca palustris, and Vaccinium macrocarpon. The presence of
Chamaedaphne suggests a successional process taking place. Further study of this and
similar communities may reveal that they are referable to Breden’s Pine Barren shrub swamp
community.

(4) NON-TIDAL FRESHWATER PONDS. Within the western half of Bear Swamp West,
remote sensing reveals several small freshwater ponds. These bodies of water are probably
not natural and most likely are remnants of excavations made early in the present century
or earlier, as part of the putative cranberry-growing operation mentioned above. To the best
of my knowledge, none of these ponds, which are remote and difficult to find on foot, have
been studied botanically; indeed, the same is true of most of the surrounding forests. Further
conjectures concerning species which may occur there will be made below in the section on
pitch pine lowland forest.

A few old farm ponds can be found in places in the Glades Region. In some cases the
farms were abandoned long ago and woodland has become established around the ponds.
Acer rubrum var. trilobum and Magnolia virginiana are probably the most abundant trees
in such wooded stands.

Flooded borrow pits in Bear Swamp West are discussed in the next community type

(5) HERBACEOUS WETLAND PRIMARY SUCCESSION. In the early decades of the present
century, extensive glass-sand mining activities were undertaken in the eastern half of Bear
Swamp West. Three large borrow pits were dug, which filled immediately with fresh water.
From them a mixture of water and sand was pumped to a processing plant where extraction
of the glass-sand took place. Activities at two of these ponds were completed in about 1960,
and activities at the third have been completed, or nearly completed, within the past ten

ars.

Upon abandonment, the water in these ponds has become clear, and a
successional process has begun on their bare, sandy shores. Many species of herbaceous
plants found on the shores of these ponds include Andropogon glomeratus, Calamagrostis
cinnoides, Carex crinita var. crinita, C. debilis, C. intumescens, Cladium mariscoides,
Decodon verticillatus, Dichanthelium acuminatum (vars. acuminatum, densiflorum,
implicatum), D. scoparium, Drosera intermedia, Dulichium arundinaceum, Eleocharis
microcarpa, E. olivacea, E. tenuis, E. tuberculosa, Glyceria obtusa, Juncus acuminatus, J.
dichotomus, J. pelocarpus, J. scirpoides, J. subcaudatus, J. effusus (including var.
compactus), Lobelia nuttallii, Lycopodium appressum, L. carolinianum, L. complanatum vat.
flabelliforme, Myriophyllum humile, Nymphaea odorata, Panicum longifolium, Phragmites
australis, Pogonia ophioglossoides, Polygala nuttallii, Potamogeton confervoides, P.
oakesianus, Rhexia virginica, Rhynchospora alba, R. capitellata, R. macrostachya, Sagittaria
engelmanniana, Scirpus cyperinus, S. pungens, S. subterminalis, Spiranthes cernua,
Triadenum virginicum, Utricularia cornuta, U. fibrosa, U. geminiscapa, U. purpurea, U.
subulata, U. vulgaris, Vaccinium macrocarpon, and Xyris difformis var. difformis.

Bog formation has not occurred along these shorelines. However, Decodon verticillatus,
a well-known bog-builder, is common here.

Phragmites australis has become well established at several places along the shores of
these ponds, although it does not seem to be spreading at the moment.

Breden (1989) does not treat "open canopied communities which have formed as a result

VEGETATION IN GLADES REGION 107

of anthropogenic ground disturbances such as agriculture or mining," so the early-
successional shores of these abandoned borrow pits have yet to be classified under his
system. However, McAvoy (1993) describes a coastal strand and barrier island community-
type in Delaware, interdunal wetlands, whose floristic composition is remarkably similar to
that of the early-successional pond shores under discussion. His list includes Cladium
mariscoides, Drosera intermedia, Euthamia tenuifolia, Juncus dichotomus, J. scirpoides,
Lycopodium appressum, Scirpus pungens, Xyris difformis var. difformis, and Vaccinium
macrocarpon, which are also found along the shores of the abandoned borrow pits of Bear
amp West. McAvoy also lists the following mesic species as belonging to the edges of
the Delaware interdunal wetland community: Myrica cerifera, Vaccinium corymbosum,
Aronia arbutifolia, Acer rubrum, and Liquidambar styraciflua. These species also occur on
the landward edges of the shorelines of Bear Swamp West’s borrow pits or occur nearby.

In this rough comparison of signature plant species, the Bear Swamp West shoreline
community is similar to interdunal wetland communities in Delaware. Breden’s (1989)
description of interdunal wetland communities (coastal interdunal marsh) in New Jersey
differs from the pond shoreline community described here and from interdunal wetland
communities in Delaware as described by McAvoy. Further study is needed here.

(6) EARLY BROADLEAF SWAMP FOREST. When fields or other open areas on poorly-
drained soils are released to natural succession, the initial woody plant community is a forest
of young trees often dominated by Acer rubrum var. trilobum, which may occur in nearly
pure stands. Liquidambar styraciflua is also often common in such communities. As these
forests mature, many additional woody plant species are recruited. Our findings thus agree
generally with those of Hanks (1971) on early hydric woody succession on the Inner Coastal
Plain in west-central New Jersey.

(7) PITCH PINE LOWLAND FOREST. Because within the Glades Region no elevations exceed
about five meters above mean sea level, the water table is never far below the surface.
Wherever the water table is very near or at the surface and forest is present, that forest is by
definition hydric, and there are broad transition zones between hydric forests and other
vegetation types. Hydric forest in the Glades Region is either broadleaf forest or pine-
dominated forest.

In the Glades Region a large number of hydric sites support patches of forest dominated
by Pinus rigida. | am proposing to classify these patches as pitch pine lowland forest. They
are probably very similar in etiology and composition to the pitch pine lowland forests of
the Pine Barrens described by McCormick (1979). Breden (1989) uses the identical term,
pitch pine lowland forest, for the same community. However, Xerophyllum asphodeloides
and Gaultheria procumbens, mentioned by McCormick as understory components of pitch
pine lowland forests, are apparently absent from our study area, and Chamaedaphne
calyculata has not been noted by this observer in such situations, although it is present
elsewhere in the Glades Region.

In the vicinity of our study area a number of stands of Pinus serotina have been reported
by many observers. This southern (supposed) species has a disjunct population in
Cumberland and Cape May counties in New Jersey. Its taxonomic status is poorly
understood, and it may be conspecific with P. rigida. In south-central Bear Swamp West,
there are trees referable to P. serotina and on the Glades I have seen one tree at least
Suggestive of this species. Remote sensing and direct observation from the air indicate that
patches of pitch pine lowland forest are numerous in the potentially very rich western portion
of Bear Swamp West, so additional occurrences of trees identifiable as P. serotina should

108 BARTONIA

be expected there, as well as elsewhere in pitch pine lowland forests of the Glades Region.

Pinus taeda is another rare tree in Cumberland County. There are scattered individuals
or small stands of this species not far from our study area, south and east of the town of
Dividing Creek, and Dale Schweitzer (pers. comm.) has found a small stand within the
Glades Refuge. This tree is often associated with P. serotina, so it too should be looked for
in the extensive but patchy pitch pine lowland forests of the Glades Region, especially in the
western portion of Bear Swamp West where trees resembling P. serotina occur.

(8) BROADLEAF SWAMP FOREST. In the Glades Region’s hydric forests standing water is
often present for at least part of any year with normal rainfall. In our study area, mature
forests over such a substrate, when not pine-dominated, fall into the category of broadleaf
swamp forest, which Breden (1989) denotes by the term Cape May lowland forest.

Acer rubrum var. trilobum, Magnolia virginiana, and Nyssa sylvatica co-dominate the
mature and old-growth broadleaf swamp forests. Wherever these three species occur together
in the Glades Region, the forest type is almost certainly of the present type, and there
Osmunda cinnamomea will almost always be found. Scattered individuals of Pinus rigida
and occasional specimens of Liriodendron tulipifera occur in some places in this forest.

To the east of the North-south Trail in Bear Swamp West and extending to the shores of
the freshwater ponds there is a remnant patch of very old broadleaf swamp forest of
approximately 36 ha in area. Here trees attain heights of up to approximately 40 m, and
there are at least three strata of woody plants, a closed-canopy "A" story, a "B" story with
semi-closed canopy, and a shrub and small-tree layer or "C" story. Ferns, sparse sedges,
very sparse forbs, and Sphagnum, often with many tree and shrub seedlings, constitute the
field layer of this forest. Photographs taken from the opposite shore of one of the ponds
before leafing-out in early spring reveal this stratification very well. The complex
physiognomy of the forest, the size (and by inference the age) of its largest trees, and the
diversity of its woody plants, all suggest strongly that the forest has never been cut, or at
most that it has been cut only minimally and selectively. I am provisionally labelling this
as an "old-growth" tract of broadleaf swamp forest. Final determination of the age and
history of this forest awaits core-sampling studies.

In this old-growth forest, the following estimates of size and demographic relationships
among the three co-dominant species should be verified by quantitative studies. Nyssa
sylvatica is the tallest of these trees and is very abundant; Acer rubrum var. trilobum appears
of equal or greater abundance, but although it is an important constituent of the canopy, it
seldom if every attains the heights of the taller Nyssa. Magnolia virginiana is probably
somewhat less abundant than N. sylvatica, and attains reproductive maturity at considerably
less than sub-canopy height. Here it occurs as a large tree, to heights of approximately 30
m (approximately 100 ft), entering the sub-canopy, although in our area it seldom exceeds
10 m in height outside of the old-growth stand. A living specimen of M. virginiana had a
circumference at breast height (Cbh) of 157.5 cm (5 ft 2 in) in 1982. The previous state
record for this species was 4 ft 8 in (Daniel O’Connor pers. comm.). A larger, wind-thrown
but still-living, specimen in the old-growth of Bear Swamp West measured 186.7 cm (6 ft,
1.5 in) Cbh in 1986.

In old growth, wherever there is slightly better drainage, Liguidambar styraciflua often
occurs as a fourth co-dominant with N. sylvatica, A. rubrum, and M. virginiana, even tending
to replace N. sylvatica; these marginally higher and drier patches are evident to an observer
on the ground and show up well on the infrared aerial photograph of the forest, and in these
patches Osmunda cinnamomea appears to be absent. On the vegetation map these drier

VEGETATION IN GLADES REGION 109

patches have been labeled, rather unsatisfactorily, as a transition zone, "broadleaf
swamp/upland broadleaf old-growth," even though no mesic old-growth is present in or near
the study area. Possibly such patches should be regarded as better-drained portions of the
hydric forest, but because on the infrared aerial and on the ground they are so readily
distinguishable from the surrounding forest, they have tentatively been considered as a
separate category, a transition zone. Broadleaf swamp forest in our regions seems to occur
mainly if not exclusively over muck (Muck-Atsion-Berryland association); the "transitional"
patches in question may be indicators of inclusions of another soil type, most likely
Fallsington sandy loam (Hammonton-Fallsington-Berryland association) (U.S. Department
of Agriculture 1978). These patches appear to resemble Breden’s Liquidambar/Acer
hardwood swamp, concerning which, Breden says, the "classification needs work." The
status of these patches and their substrate type needs further investigation.

Fagus grandifolia, Quercus bicolor, and Q. michauxii are three other species found
occasionally in old-growth. Quercus michauxii, a tree of southern distribution which occurs
sporadically in Cumberland and Cape May counties, suffered heavily during the gypsy moth
infestation of the mid-1980s, and may now be extremely rare in the Glades Region.

Understory trees and shrubs in mature or old-growth broadleaf forest include Clethra
alnifolia, Gaylussacia baccata, G. frondosa, Ilex laevigata, I. opaca, I. verticillata, Kalmia
latifolia, Lindera benzoin, Rhododendron viscosum, Sassafras albidum, Vaccinium
corymbosum, V. vacillans (V. pallidum), and Viburnum nudum. A stand of Chamaecyparis
thyoides, containing only a few small individuals, exists within hydric (but not old-growth)
broadleaf forest in Bear Swamp West.

At a few stations within mature and old-growth broadleaf swamp forest, the mistletoe
Phoradendron serotinum is parasitic on Nyssa sylvatica. To the extent of my experience,
N. sylvatica is the only tree species parasitized by mistletoe in and around the Glades
Region, although in Delaware it can be found on a number of additional species, including
Acer rubrum, which are abundant in suitable habitat in the Glades Region.

A stand (of approximately twenty individual plants) of Listera australis occurs in the
broadleaf swamp forest of Bear Swamp West.

(9) EARLY UPLAND BROADLEAF FOREST. Juniperus virginiana is an important pioneer
woody plant on abandoned fields over better-drained soils in the study area. It persists
underneath the developing woody canopy, but is not present in the mature forest. Acer
rubrum var. trilobum is a frequent occupant of mesic sites at later seral stages; unlike J.
virginiana this species often persists indefinitely, being found as a large tree among the
dominant oaks characteristic of the upland oak forest. Scattered individuals of Pinus rigida
and P. virginiana also occur in this sere and persist into oak-dominance.

(10) UPLAND PINE FOREST. A patch of upland or mesic pine forest occurs near Beaver
Dam, standing on Fallsington sandy loam of the Hammonton-Fallsington-Pocomoke
association (U.S. Department of Agriculture 1978) which appears to be moderately well-

rained. Being dominated by Pinus rigida with P. virginiana usually present, this patch
resembles upland pine forests of the nearby New Jersey Pine Barrens except for its thick //ex
opaca understory. A few similar patches occur elsewhere in our study area. From the soil
survey (op. cit.) it appears that all such patches occur on Hammonton-Fallsington-Pocomoke
soils, which soils tend to be "moderately well drained to very poorly drained" (op. cit.).
Wherever the infrared aerial indicates moderate to good drainage, pine-dominated patches
on such soils have been classified as upland pine forest. Among Breden’s categories, the
most appropriate one for these patches appears to be his dry pine-oak forest.

110 BARTONIA

(11) UPLAND OAK FOREST. I have classified about half of the forested area in the Glades
Region as upland or mesic oak forest. Such forest is oak-dominated and stands on better-
drained soils. Quercus alba, Q. falcata, Q. montana, Q. phellos, Q. rubra, and Q. stellata
are important species. Scattered among these oaks are Acer rubrum var. trilobum,
Liquidambar styraciflua, Nyssa sylvatica, Pinus rigida, and P. virginiana; the understory is
often dominated by J/ex opaca which sometimes forms thick, insular stands which J have
nicknamed "holly islands." Shrubs in the Ericaceae also constitute an important component
of the understory of this forest, as they do in the Pine Barrens. Among these are Kalmia
latifolia and Clethra alnifolia (a heath relative). Gaylussacia spp. and at least occasionally
Vaccinium vacillans (V. pallidum) are among the important heaths in the understory of this
diverse forest.

Except for the presence of the scattered pines, these forests in our area seem to fit
Breden’s mesic Coastal Plain mixed oak forest, southern Coastal Plain mesic oak forest
subtype. All of the tree species which Breden (1989) lists as important for this subtype are
present: Quercus falcata, Q. phellos, Liquidambar styraciflua, Fagus grandifolia, Ilex opaca,
Cornus florida. Two additional species, which Breden says reach the northern limits of their
ranges, Q. michauxii and Q. nigra, are present within or close to our study area.

Gypsy moth damage was severe in the upland oak forests of the Glades Region in the
mid-1980s. Many oaks, especially in the white-oak group, died from this infestation, and
the result today is that there are sunlit openings in the forest, where no doubt succession is
occurring. Studies of this succession are needed for their intrinsic interest and their potential
importance to land-management.

(12) BEECH MONODOMINANT FOREST. On the edge of one of the abandoned borrow bits
in Bear Swamp West is an extensive stand of large trees apparently dominated by Fagus
grandifolia. The stand is, roughly, a rectangle, of dimensions approximately 360 m by 180
m. From its edge, from the air, and from the infrared aerial photograph, it appears that tall
beeches constitute over 50% of the forest canopy. Its etiology is not known. According to
Breden (1989), mesic Coastal Plain mixed oak forest may contain beech-dominated stands,
of which our beech-monodominant patch is presumably an example.

(13) HEATH-SHRUB COMMUNITY. In northern Bear Swamp West, due north of the ponds
and lying just south of the Central Railroad of New Jersey, is a small savannah, a patchy
strip of land with scattered trees and a low shrub layer with closed canopy, under one meter
in height, containing the following low shrubs of the Ericaceae: Gaylussacia baccata, Kalmia
angustifolia, Lyonia mariana, and Vaccinium vacillans (V. pallidum). Trees scattered
through this narrow strip include Quercus alba, Q. coccinea, Q. montana, and Pinus rigida.
This strip of savannah is restricted to a single soil type, Downer sandy loam of the Downer
series. Its southern boundary coincides almost exactly with the boundary between this soil
and an area of muck lying just to the south. Although there is no obvious sign of recent fire
in this community, nor is any other explanation known, it is possible that the community is
seral and owes its existence to fairly recent anthropogenic or natural disturbance, including
fire. The name heath-shrub community is tentative, and further study is needed.

DISTURBED AREAS. The present physiognomy and floristic composition of the mature
forests of the Glades Region reflect a recent history of minimal anthropogenic disturbance,
except where the gypsy moth infestation of the mid-1980s was severe. Yet there are
scattered areas comprising perhaps ten to twenty per cent of the forested landscape, where
indications of long-abandoned homesites or fairly recent agricultural activity are evident.
Occurring in both hydric and mesic situations, successional stages in these abandoned areas

VEGETATION IN GLADES REGION 111

range from oldfield to nearly mature forest. A number of weedy plants, some of them alien,
aggressive invasives, can be found where fairly recent disturbance has taken place.
Indicators of disturbance include Asplenium platyneuron, [Hemerocallis fulva], [Ligustrum
vulgare], [Lonicera japonica], Phragmites australis, and [Rosa multiflora]. Other native,
woody, successional plants in these seral areas include Acer rubrum var. trilobum,
Liquidambar styraciflua, and Prunus serotina.

e rare Polygala mariana appears here as an opportunist able to establish itself in a
disturbed area.

DISCUSSION

As it is to be expected in an area in which anthropogenic disturbance has been minimal,
the entire study area is a richly variegated, patchy mosaic, as the vegetation map reveals.
In many respects it is comparable to the New Jersey Pine Barrens, the nearest point of which
lies a few kilometers to the northeast of Bear Swamp West. McCormick (1979) considers
that the upland portions of the Glades Region belong to the "oak-pine fringe area" of the true
Pine Barrens. His description of the vegetation of the New Jersey Pine Barrens parallels in
several respects what I have found in the Glades Region. In the forested section of our study
area which lies inland from the narrow sandy bayshore beaches and the broad tidal marshes
I have found a hydric complex and a mesic complex corresponding to the "lowland and
upland complexes" identified by him in the Pine Barrens. I have placed these complexes in
my "forested mosaic landscape type," to which I have added a "tidal wetlands landscape
type" to include the bayshore and adjacent tidal wetlands, a type which of course is absent
from the Pine Barrens of the interior.

It is probable that fire in the Glades Region has historically been a major ecological factor
as it has in the Pine Barrens (McCormick 1979, Little 1979). I know of no published studies
of the ecological consequences of fire in and around the Glades Region paralleling similar
studies in the Pine Barrens carried out by Little, but by inference from the results of these
studies it is likely that frequent fire has a place in the etiology of patches of pine forest in
the Glades Region.

The large stands of Pinus rigida in the Glades Region are strongly reminiscent of similar
forests of the Pine Barrens, although a major difference in general aspect is that /lex opaca
occurs frequently in the understory of the pine forests of the Glades Region. McCormick
(1979) states that on the western margin of the Pine Barrens of New Jersey, P. virginiana
occurs as an old-field tree. In the Glades Region, I have seen that species as a pioneer in
disturbed sites, and also as larger specimens in patches of older forest where evidence of
recent disturbance is less clear. The scattered patches of forest dominated by P. virginiana
which I have seen in the study area may belong to Breden’s (1989) Virginia pine-oak forest.

In oak or oak-pine forests of the Glades Region, all of McCormick’s upland oaks are
present: Quercus alba, coccinea, falcata, marilandica, montana, stellata, and velutina. Ilex
opaca often dominates the understory, although heaths are often important. Of the
herbaceous members of upland forests in the Pine Barrens according to McCormick,
Tephrosia virginiana is apparently ney and Melampyrum lineare is common especially
along paths or in similar disturbed ar

McCormick (p. 236) suggests aa fi in en vertical structure, the mesic forests in the Pine
Barrens could be visualized as a heath forest under a pine or oak forest. In our study area
this description is applicable to both hydric and mesic forests except where Ilex opaca is a
true tree with lowest branches near the ground, rather than a shrub, so we may think of it

112 BARTONIA

as occurring above rather than in the heath layer, which seldom exceeds 3 m in height. In
old-growth the holly forms a portion of the lowest tree layer. Perhaps the optimal situation
in the oldest hydric and even mesic forests is broadleaf, over holly and other small trees,
over heath shrubs, except that where the holly forms a monospecific closed understory
canopy it creates deep, permanent shade and very little grows beneath it.

There are several differences between hydric communities in our study area and those of
the Pine Barrens as reported by McCormick (1979). Among communities dominated by
woody plants there are no cedar swamps within the Glades Region although small numbers
of Chamaecyparis thyoides are present and two cedar swamps do occur nearby. As with
McCormick, Acer rubrum (always var. trilobum), Nyssa sylvatica, Magnolia virginiana, and
sometimes Liguidambar styraciflua are the co-dominants of hydric broadleaf forests in our
area, although from visual evidence it is not clear here that, as McCormick suggests for the
Pine Barrens, Acer is the dominant and the others are associates. As in the Pine Barrens,
heaths and heath-allies in our study area include Vaccinium corymbosum, Clethra alnifolia,
Leucothoe racemosa, and Rhododendron viscosum. Unlike the Pine Barrens, Lindera
benzoin may form dense stands in the shrub layer of extremely poorly-drained hydric old-
growth, and J/ex opaca often dominates the understory of hydric forests with heaths
secondary in importance (as with mesic forest). As with McCormick, Pinus rigida may be
scattered through mesic and hydric forests, but Xerophyllum asphodeloides (listed as rare in
the county by Moore) and Chamaedaphne calyculata are completely absent (though
Chamaedaphne occurs in another habitat). Similarly to the Pine Barrens, Pteridium
aquilinum is common

To the best of my knowledge, Quercus palustris is uncommon in hydric forests of our
study area and Q. nigra is absent, the latter a rare species at the northern limit of its range
in Cumberland County (Moore 1989). Quercus phellos and michauxii are present, although,
as discussed earlier, Q. michauxii has been seriously reduced in numbers by the gypsy moth.

In herbaceous hydric communities within our region, Nuphar lutea and Eriocaulon spp.
are apparently absent. The small patches of freshwater marsh in western Bear Swamp West,
easily visible by remote sensing, have been scarcely studied, but may be found to contain
some of these species. The presence of Chamaedaphne calyculata suggests that these
patches of marsh may be "savannahs" (McCormick 1979) being replaced by thickets and
swamp forests of Acer rubrum var. trilobum, Nyssa sylvatica, Magnolia virginiana, an
Vaccinium corymbosum, all of which are abundant in the vicinity. As noted above,
communities in Bear Swamp West dominated by Chamaedaphne may belong to Breden’s
(1989) Pine Barren shrub swamp community type. The presence of V. macrocarpon in the
marsh community reinforces the local tradition that a cranberry-growing operation existed
in this western portion of Bear Swamp West long ago.

Similar forests to the beech monodominant community in Bear Swamp West are common
in the Piedmont and are not unknown in the New Jersey Coastal Plain. Of the Inner Coastal
Plain, Robichaud and Buell (1973, p. 210) report, "Near the Delaware River in Camden
County almost pure stands of beech trees have developed and here, as in North Jersey, the
beech appears to propagate itself by root sprouts." Another beech-dominated forest, similar
to that in Bear Swamp West, lies in Bear Swamp East. Cartica, Niles, and Petrongolo (1988)
describe this beech hardwood forest as follows and a map accompanies their description:

American beech dominates the canopy of approximately 125 acres which also commonly includes white oak,
sweet gum and red maple (Terrestrial Environmental Specialists, Inc., . . . ).... Field examination of this

e
O,
Ks 5

Tidal:
Marsh
-~...--~ Creeks and Ponds
Non-Tidal Freshwater:
FM Marsh
FP Ponds
BP Borrow Pits
PS — Hydric Herbaceous
HS — Heath-Shrub
Forests:
Hydric:
H Early Broadleaf
HP Pitch Pine Lowland
HB Broadleaf Swamp
B Beech Monodominant
Broadleaf Old Growth

ds — ae” 0, 8
SA
* end? a . ‘
oo one .
a < ° *.
Sa NEWPORT

M Early Broadleaf
MP Upland Pine
MB Upland Oak
" MBOG Broadleaf Old Growth
i HP/HB _ Transition Zones
| Agriculture
Disturbed
Developed

Bn WE Crete Oke
is Ci mea SN ae y. a eH aN ad Sh" ly
( 3 ; Van

=
4 £ A. oe
” ae pane
a ee ~~; = sates
ee ea”

ae \ “

VEGETATION IN GLADES REGION 113

forest type was not undertaken and the extent of domination by beech is not known. C. E. Maguire, Inc.
. indicates that recent selective cutting . . . occurred here

Daniel O’Connor (pers. comm.) has suggested to me that selective cutting may also have
taken place in the beech forest of Bear Swamp West, and may have played a part in its
etiology.

ACKNOWLEDGEMENTS

Part of the preparation of this paper was financed through grants to the Natural Lands
Trust by the Fund for New Jersey, the Geraldine R. Dodge Foundation, and the Dolfinger-
McMahon Foundation. I am grateful for their support. I thank Dr. A. E. Schuyler of the
Academy of Natural Sciences who contributed many suggestions during the entire project,

and Andrew Pitz of the Natural Lands Trust who supervised the preparation of the
Glades—Bear Swamp Master Plan, for which the research reported in this paper was carried

ut. Meg Thayer of the Natural Lands Trust gave constant advice and skillfully prepared
the vegetation map in which many of her suggestions were incorporated which improved its
final appearance; Steve Kuter, also of Natural Lands Trust, helped with final revisions of the
vegetation map. This research would never have been carried out at the present scale were
it not for constant assistance from Daniel O’Connor, who knows the entire study area
intimately and is a public advocate for its preservation. He guided me on numerous
occasions, especially in Bear Swamps East and West, and he frequently called my attention
to natural features deserving of study. Dale Schweitzer also contributed important
information. Thanks are due Vickie Kress who painstakingly typed the plant list.

LITERATURE CITED

, W.H., W. A. WENT: D B. D. TREADWELL. 1980. A Guide to Remote Sensing Information for
Wildlife Biologists. Chapter aa in n Wildlife Management Techniques Manual, 4th ed., rev. Wildlife Society,
Washington.
BREDEN, T. F. 1989. A Preliminary Natural Community Classification for New Jersey. Pages 157-191 in New
Jersey’s Rare and Endangered Plants and Animals. Institute for Environmental Studies, Ramapo College,
Mahwah.

CaRTICA, R. J., L. NILES, AND T. PETRONGOLO. 1988. Bear Swamp East Natural Area Management Plan. New
Jersey Department of Environmental Protection, Division of Parks and Forestry, Office of Natural Lands
Management, Trenton.

KS, J. P. 1971. Secondary succession and soils on the inner coastal plain of New Jersey. Bull. Torrey Bot.
Club 98(6): 315-321.

KARTESZ, J. AND R. KARTESZ. 1980. A Synonymized Checklist of the eas Flora of the United States, Canada,
and Greenland. The University of North Carolina Press, Chapel H

LITTLE, S. 1979. Fire and Plant Succession in the New Jersey Pine alee Chapter 17 in R. T. T. Forman, ed.
Pine Barrens: Ecosystem and Landscape. Academic Press, New York.

McAvoy, W. 1993. Characterization of Category I Non-tidal Wetland Communities in Delaware: Interdunal
Wetlands. Division of Parks and Recreation, Delaware Department of Natural Resources and Environmental
Control, Dover.

McCormick, J. 1979. The Vegetation of the New Jersey Pine Barrens. Chapter 13 in R. T. T. Forman, ed. Pine
Barrens: Ecosystem and Landscape. Academic Press, New York.

Moore, G. 1989. A checklist of = vascular plants of Cumberland County, New Jersey. Bartonia 55: 25-39.

ROBICHAUD, B. AND M. BUEL LL. 1973. Vegetation of New Jersey: A Study of Landscape Diversity. Rutgers
sing ay pe New Brun

STALTER, B),. OF een "pe. Ecological observations on Ilex opaca. Holly Society Journal 2: 1-10.

US. routes OF AGRICULTURE. 1978. Soil Survey of Cumberland County, New Jersey. Soil Conservation
Service in cooperation with New Jersey Agricultural Experiment Station and New Jersey Department of
Agriculture.

= i -
¥ =e, 7
i a Alin 3 6 : .
Ht
7 Z Pa] :
; - : an ee : : ; .

4
r
a

7 7 ee

=.
\ 7 -
j
+> a all Lar =
* 7 oe a a ? -
5 §
? i a a
-
La ‘ he

: ‘ :

eae Ty Be ae ro :
‘ 7 t it J pan ee Hs
- 7
7 ; ‘ Perea tien
:
: : 5 a ae
¢ 7 z 7 7 - ie ) 7 Snel
: ; , ee :
% at yee - - >
a i a Lam an ad
7 y a A 7
, cis ‘ .
oe. a yy : tet 7 a
ad ne mh Fos A sad + ee ee 2)
7 ee ca ee eee ee 5
; 27. mew oe ind erie Si cee
ce : a - ai! 7 , : a % a igh . >
ee . 4 ee al A Pe > ae iy
Pa i Win a : oan ~ a aa
: . ie MD, at SI. alk

“a

Bartonia No. 58: 115-119, 1994

Ecological Observations of Bear Swamp,
Cumberland County, New Jersey

RICHARD STAL
Department of Biological Sciences, St. John’s University, ee Utopia Parkway, Jamaica, NY 11439

DANIEL O’CONNOR
66 Main Street, Port Norris, NJ 18349

The objective of the present study was to determine the composition of the arborescent
vegetation of Bear Swamp East and Bear Swamp West. Bear Swamp, located in
Cumberland County, New Jersey, may contain the most extensive uncut forest in southern
New Jersey. Bear Swamp consists of two distinct segments: Bear Swamp East (820.4 ha)
and Bear Swamp West (747 ha). Bear Swamp East is located at the headwater of the
Dividing Creek watershed 39° 17’ N, 75° 05’ W; Bear Swamp West is located in the
Oranoakan Creek watershed 39° 17’ N, 75° 08’ W. The water table at both sites is at or
above the surface of the ground for long periods of time each year with Bear Swamp West
being wetter than Bear Swamp East. Bear Swamp is underlain by the Cape May and
Cohansey Formations, and is bordered to the north by the Bridgeton Formations. The soils
are wet and in many portions are water-covered for most of the year. The soils are acid with
a pH ranging from 4.0-5.5.

An 1843 description of Bear Swamp, which applied primarily to what is known today as
Bear Swamp West, stated that it was "a noted swamp of Downe t-ship [sic], Cumberland
Co., near Nantucket or Newport, through which flows the Oronoken [sic] Creek. The timber
upon it is chiefly oak and poplar" (Stalter and O’Connor 1984). Much later shee (1912)
wrote, "Silent inscrutable and alien lay the swamp, and untouched by hum ands .

I was now in the real swamp, the old uncut forest. It was a land of tree ee huge satis
poplar and swamp white oak so old that they had become solitary, their comrades having
fallen one by one; while some of them, unable to loose their grip upon the soil, which has
widened and tightened through the centuries, were still standing, though long since, dead."

METHODS

The quadrat method was used to sample trees (a tree being defined as having a straight
unbranched axis 6 m or more tall with a diameter at breast height of 7.6 cm or greater).
Twenty five 10 x 10 m quadrats were selected in the center of Bear Swamp East while
fourteen quadrats of the same size were selected to sample arborescent vegetation in the
center of Bear Swamp West. Quadrats at both sites were nested in groups of 2s and 3s.
Fewer plots were selected to sample trees in Bear Swamp West because this site contained
fewer tree species than Bear Swamp East. Frequency (percent quadrats occupied by each
species), density (average number of trees per quadrat), relative dominance (percent basal
area), and importance values (the sum of the relative frequency, relative density, and relative
dominance) were calculated for trees in Bear Swamp East and Bear Swamp West (Table 1).

115

116 BARTONIA

Table 1. Density (D), relative density (RD), frequency (F), relative frequency (RF), basal area alia peal
wet (RDo), mu importance values (IV), for trees in Bear Swamp, Cumberland County, New Jersey. Basal
values are in cm

Species D RD F RF BA RDo IV
Bear Swamp East
Acer rubrum 1.48 30 88 26 12,268.7 47 103
Ilex 92 18 64 18 1,274.8 41
Nyssa sylvatica 80 16 Ad 15 1,664.89 6 ST
Magnolia sees .80 16 44 15 Los) a 6 37
icha 36 7 36 10 4,228.8 16 33
Liriodendron aes 24 5 20 6 2,665.2 10 21
Fagus grandifolia 20 4 20 6 1,011.4 4 14
Pinus seroti 12 2 12 3 776.0 3 8
Sassafras albi 12 2 12 3 289.3 1 6
Liquidambar styraciflua 04 l 4 1 287.3 1 3
Bear Swamp West
Ilex opaca 2.64 46 86 30 3,965.9 11 87
Nyssa _sylvatic iA 13 43 15 16,723.4 45 73
Magnolia virginiana 1.0 18 64 23 3,081.0 8 49
79 14 50 18 5,595.6 15 47
Liquidambar styraciflua A3 8 28 10 6,055.4 16 34
Liriodendron tulipifera .07 1 7 3 Lay 3 7
Fagus grandifolia 07 1 7 3 127.7 J

Nomenclature follows Gleason and Cronquist (1991).

e moisture tolerance of the trees at Bear Swamp was determined using a surveyor’s
transit and stadia pole. Three sites were selected for study at Bear Swamp East; a tangle of
Smilax rotundifolia prevented us from conducting survey work at Bear Swamp West. The
most water tolerant tree species occupies the lowest land, the land where the water table is
highest relative to the surface of the ground. The elevation of the most flood tolerant
individual measured of this tree species serves as a reference point at 0.00 feet in elevation.
Additional individuals of this same flood tolerant species and other species were surveyed
to determine their distribution along a moisture gradient.

Historical information was obtained by a search of deeds, tax records, court house records,
and census information at the County Seat of Cumberland County, Bridgeton, New Jersey.
More detailed historical information of Bear Swamp and the surrounding area is presented
by Stalter and O’Connor (1984).

RESULTS

Acer rubrum is the dominant tree of Bear Swamp East (Table 1). It has the highest
density, frequency, relative dominance and importance values. Ilex opaca, which rarely
grows over 12 m tall, is the dominant subcanopy species and has the second highest
importance value of all the arborescent species sampled at Bear Swamp East. Ilex opaca has
the highest importance value at Bear Swamp West (Table 1) because it is the most numerous
species (density of 2.64 individuals/10 x 10 m quadrat) and attains the highest frequency
(86% of the quadrats). Nyssa sylvatica is the dominant canopy species at Bear Swamp West
and has the highest relative dominance value. Acer rubrum and Liquidambar styraciflua, are
associated with Nyssa sylvatica in the canopy. Magnolia virginiana ranks second in density

BEAR SWAMP 117

and frequency at Bear Swamp West and is more abundant here than in Bear Swamp East.
Species composition of Bear Swamp East and Bear Swamp West is similar (similarity
coefficient of 0.82).

DISCUSSION

Ilex opaca, Nyssa sylvatica, Magnolia virginiana, and Acer rubrum occupy the lowest
wettest portions of Bear Swamp. There is little variation in moisture tolerance for many
species in the swamp since [lex opaca, Acer rubrum, Nyssa sylvatica, Magnolia virginiana,
Fagus grandifolia, and Liquidambar styraciflua all are distributed within an elevation range
of less than 7.6 cm.

All trees in Bear Swamp can apparently tolerate flooding, although Acer rubrum, Nyssa
sylvatica, Ilex opaca, and Liquidambar styraciflua have higher density, frequency, relative
dominance and/or importance values than Fagus grandifolia, Pinus serotina, and Sassafras
albidum. Nyssa sylvatica and Magnolia virginiana appear to be restricted to the wettest sites
while Acer rubrum and Ilex opaca are distributed over a greater moisture gradient.

Bear Swamp East possibly contains the most extensive (ca. 83 ha) uncut forest in southern
New Jersey. Within this swamp are two state record trees: Pinus serotina (pond pine), and
Quercus michauxii (basket oak). Many large trees of Liriodendron tulipifera, Ilex opaca,
Acer rubrum, Magnolia virginiana, Liquidambar styraciflua, Nyssa sylvatica, and Fagus
grandfolia grow here. Bear Swamp West contains many old large trees including the state
record Magnolia virginiana. Both Bear Swamp East and Bear Swamp West contain the rare
Phoradendron serotinum (mistletoe), which last century was far more commonly distributed
in New Jersey (Stone 1912).

Tree seedlings and saplings (a sapling being defined as a tree with a DBH less than 7.6
cm) are represented by Ilex opaca, Liquidambar styraciflua, Acer rubrum, and Nyssa
sylvatica.

The most common shrubs in Bear Swamp are Clethra alnifolia and Toxicodendron
radicans. Other shrubs include: Vaccinium corymbosum, Rhododendron viscosum, Lyonia
mariana, Itea virginica, and Lindera benzoin. The most common liana is Smilax
rotundifolia. Smilax festoons the shrubs and trees; portions of Bear Swamp West are all but
impassible because of the profuse growth of S. rotundifolia.

The herbaceous stratum contains: Panicum spp., Eleocharis spp., Peltandra virginica,
Orontium aquaticum, Dulichium arundinaceum, Andropogon virginicus, A. virginicus vat.
abbreviatus, Utricularia fibrosa, U. purpurea, Proserpinaca palustris, Myriophyllum humile,
Eupatorium album, E. hyssopifolium, and Cladium mariscoides. Many of the aforementioned
species are characteristic of areas where the water table is above the ground for most of the
growing season. Ferns include Osmunda cinnamomea, Athyrium filix-femina, and
Woodwardia areolata. Sphagnum is common in wet depressions.

The moisture tolerance of Magnolia virginiana and Liriodendron tulipifera at Bear Swamp
differs from that reported by Gemborys and Hodgkins (1971) in southwestern Alabama.
They report that, "A. virginiana may be similar to Liriodendron tulipifera with respect to
tolerance for wetness." At Bear Swamp, Liriodendron was observed on the higher, drier
islands while Magnolia virginiana generally occurred on sites where the water table was at
or above the surface of the ground during a 10 year interval, 1974-1984, even during periods
of drought.

Bear Swamp is distinctly southern in character. With the exception of Taxodium, species
composition in Bear Swamp is similar to that of the Great Cypress Swamp or Burnt Swamp

118 BARTONIA

in southern Delaware and Maryland (Stalter 1981a). Acer rubrum was the dominant species
in Burnt Swamp with Liquidambar styraciflua, Nyssa biflora (reported by some as N.
sylvatica var. biflora), Ilex opaca, and Magnolia virginiana. All of the aforementioned
species are found in Bear Swamp. Bear Swamp contains five species that are structurally
important in Quaterman and Keever’s (1962) southern mixed hardwood forest: J/ex opaca,
Nyssa biflora, Liquidambar styraciflua, Quercus michauxii, and Fagus grandifolia.

A thorough search of the existing literature and an examination of deeds and court-house
records indicated that the wettest portions of Bear Swamp have never been logged. The first
piece of indirect evidence for this is a comparison of the existing vegetation in Bear Swamp
today with the giant trees described by Sharp in 1912. Sharp’s poetic description, even
accounting for poetic license, could be used to describe portions of Bear Swamp today, since
there are many trees in the swamp as large and as old as those described by Sharp 80 years
ago. Large trees today include two giant tulip poplars which are almost 5 ft (1.3 m) in
diameter. The state record basket oak and a 200 yr old pond pine would have been prime
lumber trees had the area been logged 100 yr ago. O’Connor recorded the DBH of a dead
L. tulipifera that was 5.94 m in circumference in 1976. This tree fell in late 1976 and had
completely decomposed by 1981

New Jersey State Forester Gordon T. Bamford described (pers. comm. 1982) the large oak
trees on the property of the Perry brothers near Pennsville, New Jersey. Bamford stated, "At
the time that a section was cut for New Jersey’s Tercentenary Commission in 1961, these
were identified as Quercus michauxii. As | recall the general stand was not uniformly old;
at the time of our first contact in 1948 there were a number of the larger 300+ year old
individuals that we assumed had been left when the area was horse-logged, prior to tractor
skidding equipment." The Perry property borders rt 49, a well travelled highway in southern
New Jersey. If this accessible area escaped logging, it is quite probable that a wild boggy
inaccessible area such as Bear Swamp could have also escaped cutting. The boggy nature
of Bear Swamp would have posed a formidable obstacle to those who wished to harvest its
trees. Horses would have had a difficult time pulling even 0.5 m or | m DBH logs from a
forest and would have had a far more difficult time with logs in Bear Swamp where a man
can sink to his knees in peat and ooze.

Many of the largest trees in Bear Swamp are hollow and unsuitable for lumber. Moreover
there are far more valuable lumber trees such as Atlantic white cedar (Ch }
thyoides), oaks (Quercus spp.), and pine (Pinus spp.) growing in abundance in south pe
Therefore it is reasonable to believe that portions of Bear Swamp have never been logged.
Bear Swamp may represent one of the last extensive areas of virgin woodland in New
Jersey.

ACKNOWLEDGEMENTS
We thank the American Holly Society for funds to conduct this study, Stevens Heckscher
for an unpublished list of species he identified in the swamp, N.J. State Forester Gordon
Bamford for information on the age and size of oak trees on the Perry Brothers property, and
Karl Anderson and Eric Lamont for reviewing the manuscript.

LITERATURE CITED

GEMBORYS, S. R. AND E. L. HODGKINS. 1971. Forests of small stream bottoms in the coastal plain of southwestern
Alabama. Ecology 52: 69-84.

BEAR SWAMP 119

GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of the foo plants of northeastern United States and

— Eee 2nd ed. The New York Botanical Garden, Bronx.
ND C. KEEVER. Southern mixed eh pas climax in the southeastern coastal plain,

as S.A A, Baal, ae 32: 167-1

SHARP, D. L. 1912. The spring of a year. Houghton-Mifflin, Bos

STALTER, R. 198la. Some ecological observations of Delaware eae Castanea 47: 105-1

———. 1981b. Some observations of American Holly, //lex opaca Aiton on the east coast of “ United States of
America. Proc. 57th mtg. Holly Soc. Amer. 57: 2-3

STALTER, R. AND D. O’CONNOR. 1984. Ecological observations on /lex opaca. Holly Soc. J. 2: 1-10.

STONE, W. 1912. The plants of southern New Jersey with special reference to the flora of the Pine Barrens. Ann.

ep. N.J. State Mus. 1910, part II

Rien

Bartonia No. 58: 121-124, 1994

Juncus caesariensis Coville (New Jersey Rush) in
Nova Scotia, Canada

RUTH E. NEWELL AND REG. B. NEWELL
Biology Department, Acadia University, Wolfville, Nova Scotia, Canada BOP 1X0

The first collection of Juncus caesariensis Cov. (New Jersey Rush, Juncaceae) in Canada
was made in 1951 by E. C. Smith et al. from a bog in Gracieville, Richmond County, Cape
Breton Island, Nova Scotia (45°35’N, 60°43°W). Prior to this time, J. caesariensis was
known only from the Atlantic coastal plain of New Jersey, Maryland and Virginia (Fig. 1).
The Nova Scotia collection although deposited in the E. C. Smith Herbarium of Acadia
University (ACAD), remained neglected until 1988. Neither The Flora of Nova Scotia
(Roland and Smith 1969) nor The Rare Vascular Plants of Nova Scotia (Maher et al. 1978)
report this species for the province. From 1989~1991 three field surveys were carried out,
initially to relocate the plant in the field, and subsequently to determine the range and status
of this species.

Currently, J. caesariensis is considered globally imperilled (The Nature Conservancy
1990), a ranking based on its known occurrence in the United States.

In New Jersey, J. caesariensis is reported from wet, sandy, peaty substrate dominated by
sphagnum. The sites are generally open to slightly shaded and are often in or near white
cedar swamps (Schuyler 1990). There are 20 extant occurrences with 35 additional historical
occurrences over a seven county area. In New Jersey, this species has been designated as
rare (Snyder and Vivian 1981). Ware and Wieboldt (1981) summarize the habitat in Virginia
as "very acidic, usually sphagnous, extremely wet, springy or seepy areas with perennially
reliable flow, but without standing water." More recently, Strong and Sheridan (1991)
classify the habitat of New Jersey Rush in Virginia as peat bog habitat, i.e. acidic, eevee
soil with a thick overlying layer of Sphagnum. Virginia has 13 extant populations and tw
historical stations in 5 counties. The plant is considered to be endangered (Porter oa in in
this state. Until very recently, this plant was known only from one historic station in
Maryland and was designated as extirpated (Reveal and Broome 1982). Strong and Sheridan
(1991) discovered three new sites in 1

To date, New Jersey Rush has been found in 16 sites (Fig. 1, inset) in southeastern Cape
Breton Island in an area extending a distance of 35 km along the coast from Gracieville
(Point Michaud) northeastward to Fourchu Bay and inland as far as Loch Lomond (sites
Occur in both Richmond and Cape Breton Counties). Many more potential sites for this
species remain to be investigated as this part of Cape Breton Island possesses extensive
wetland habitat.

Data collected so far suggest that in Nova Scotia, J. caesariensis is limited to sphagnous
edges of bogs and fens and/or throughout small bays and coves around the perimeter of
bogs/fens or small boggy openings in coniferous woods. Frequently associated species
include Picea mariana, Calamagrostis pickeringii, Muhlenbergia unifolia, Carex exilis, C.
michauxiana, Smilacina trifolia, Myrica gale, Alnus rugosa, Sarracenia purpurea, Aronia

121

122 BARTONIA

50.00

45.00

40.00

5CO km

Fig. 1. World distribution of Juncus caesariensis (inset represents its distribution in Nova Scotia, Canada).

sp., Rosa nitida, Ledum groenlandicum, Kalmia angustifolia, K. polifolia, Chamaedaphne
calyculata, Solidago uliginosa and Aster nemoralis. Both Juncus canadensis an
brevicaudatus may occur with or in the vicinity of J. caesariensis populations. Population
size estimates per site range from 3 to 1000 (individual clumps were counted).

The occurrence of New Jersey Rush along bog and fen margins immediately adjacent to
the shrubs and trees of the transitional zone between wetland habitat and surrounding
wooded areas suggests some shade tolerance. Clumps of New Jersey Rush were often found
in animal trails which skirt the margins of bogs and fens. In several localities, plants were
observed on moderately used all terrain vehicle (ATV) trails. These occurrences may
indicate a tolerance to or a dependence upon some degree of disturbance.

J. caesariensis is reported to be most sensitive to circumstances affecting the hydrologic
regime of its habitat (Schuyler 1990; Kral 1983; Ware and Wieboldt 1981) such as site
drainage or flooding. Although New Jersey Rush occurs ina relatively undeveloped, isolated
section of Nova Scotia, threats exist which could alter land drainage on two fronts. A major
concern exists over the extensive logging currently being undertaken in the immediate
vicinity of some populations of J. caesariensis. It is unknown whether this will have an
impact on the hydrologic regime of adjacent new Jersey Rush wetland habitat. Future
cottage development is imminent in at least the southern portion of its Nova Scotia range.
This may affect J. caesariensis through road construction allowing landowners access to
ocean frontage. Since this latter threat is likely going to threaten only a small portion of the
entire range of New Jersey Rush in the province, it remains of minor concern.

JUNCUS CAESARIENSIS IN NOVA SCOTIA 3

The current degree of ATV activity in some portions of its range appears to be beneficial
to J. caesariensis populations. An increase in usage of ATVs may be a potential threat to
this plant.

Although potential threats to J. caesariensis exist in parts of its Nova Scotia range, in
general the status of this plant here is relatively stable. Considering the global ranking of
J. caesariensis however, the Cape Breton populations should be monitored on a regular basis
to ensure that their status does not change. The full extent of the range of New Jersey Rush
in Nova Scotia and a complete tally of the number of populations remain to be documented.

This occurrence of J. caesariensis is noteworthy in terms of being the first documentation
of a disjunct, rare coastal plain species from Cape Breton Island. Its location makes it
distinct from the well known and much studied Atlantic coastal plain flora of southwestern
Nova Scotia.

The coastal plain species of southwestern Nova Scotia are believed to have migrated
northward over a land bridge existing between Cape Cod and the southwestern tip of the
province towards the end of the last glaciation (Keddy and Wisheu 1989). The presence of
New Jersey Rush in eastern Cape Breton Island supports the theory that along with a land
bridge, there was an off-shore refugium along the Atlantic coast during the Wisconsin
Glaciation when sea levels were about 100 m lower than they are presently (Roland 1982).
As parts of eastern Cape Breton have been noted to share some similarities of climate and
habitat with the other end of the province (Roland and Smith 1969) there remains the
possibility that New Jersey Rush may yet be found in southwestern Nova Scotia.

ACKNOWLEDGEMENTS

This study forms the basis of a COSEWIC Status Report prepared by the authors, with
funding provided by the World Wildlife Fund, Canada. We thank the Nova Scotia
Department of Natural Resources for allowing us access to their various resource materials.
Erich Haber kindly generated the distribution map.

LITERATURE CITED

KEDpy, P. A. AND I. C. WISHEU. 1989. Ecology, biogeography, and moet oven et “ore plain plants: some
general principles from the study of Nova Scotian wetlands. Rhodora 91(865):

KRAL, R. 1983. Juncaceae: Juncus caesariensis Cov. (Endangered species, laa pe of bogs, ravines and
seep a areas, New Jersey, Maryland, Virginia). USDA Forest Service, Southern Region, Technical Publication R8-
TP: 179-182.

Mauer, R. V., D. J. WHITE, G. W. ARGUS AND P. A. KEDDY, 1978. The rare vascular plants of Nova Scotia.
Syllogeus Series no. 18, National Museums of Canada, Ottawa.

PORTER, D. M. 1979. Rare and endangered vascular plant species in Virginia. Prepared by the Virginia
em Institute and State University in cooperation with the U.S. Fish and Wildlife Service.

REVEAL, J. L. AND C. R. BROOME. 1982. Comments on the proposed endangered and threatened vascular plant
antl of Maryland, U.S.A. Castanea 47(2): 191-200.

OLAND, A. E. 1982. Geological background and physiography of Nova Scotia. Nova Scotian Institute of Science,
Halifax

E. C. SMITH. 1969. The flora of Nova Scotia. Proceedings of the Nova Scotian Institute of Science
26: pert 277-743.
SCHUYLER, A. E. 1990. Element stewardship — for Juncus caesariensis Cov. (Juncaceae). New Jersey
Department of Environmental Protection, Trento
SNYDER, D. B. AND V. E. VIVIAN. 1981. Rare sad endangered vascular plant species in New Jersey. Prepared

124 BARTONIA
by the Conservation and Environmental Studies Center, Inc. in cooperation with the U.S. Fish and Wildlife
Service.
TRONG, M. T. AND P. M. SHERIDAN. 1991. Juncus caesariensis Coville (Juncaceae) in Virginia peat bogs.
Castanea 56(1): 65-69.
ARE, D. M. E. AND T. F. WIEBOLDT. 1981. Rediscovery of Juncus caesariensis Coville in Virginia. Jeffersonia
12(2): 6-10.

Bartonia No. 58: 125-129, 1994

OBITUARIES

Ida Kaplan Langman (1904-1991). A
member of the Philadelphia Botanical
Club since 1937, Ida Langman died at
Stapeley in Germantown health care cen-
ter, on 10 July 1991 of cardio-pulmonary
arrest after suffering from Parkinson’s
Disease for many years. Ida was a life
member of the club and substantially
contributed to its vitality. She had a deep
love for botany and was generous with
sharing her botanical knowledge. I re-
member Ida as a kind, flexible, and toler-
ant woman who had strong opinions and
a lot of determination.

Born on 7 February 1904 in the city of
Nezhin in the Ukraine, Ida came to Phila-
delphia as an infant a few months old.
She graduated from Sharswood Elementa-
ry School, South Philadelphia High
School, and Philadelphia Normal School.
At the age of 18 she began her first ca-
reer—35 years as a teacher assigned by the Board of Education to the Academy of Natural
Sciences of Philadelphia. While working as a teacher, Ida continued her education at the
University of Pennsylvania where she received her bachelor’s degree in Education in 1930
and her master’s degree in Botany in 1947. She married Oscar Langman in 1928, a profes-
sional violinist and photographer who died in 1987. In the 1930s, Ida did botanical field
work in Monroe County, Pennsylvania, where she found Juncus greenei for the first time in
the state. After spending the summer of 1938 in Mexico, she became fluent in Spanish and
developed an interest in Mexican botany that culminated in 1964 with the publication of A
Selected Guide to the Literature on the Flowering Plants of Mexico. Funds for preparation
and travel during the 20+ years she worked on the book were provided by the National
Science Foundation, American Philosophical Society, and U.S. State Department. The
American Library Association selected it as the outstanding bibliography of the year.
Subsequently Ida was elected a Corresponding Member of the Academia Nacional de
Ciencias de Mexico, and in 1972, the Botanical Society of Mexico awarded her a medal for
making a special contribution to Mexican botany. Mexican botanists named two plant
species after Ida: Lopezia langmanae Miranda and Stachys langmaniae Rzedowski &
Rzedowski.

In the latter 1960s Ida became the first leader of the Index Nominum Genericorum Project
at the Smithsonian Institution where her work was praised for scholarly excellence. In the
1970s Ida worked as a bibliographer and visited European libraries for the Hunt Institute for
Botanical Documentation of Carnegie-Mellon University where her reliable performance was
praiseworthy.

125

126 BARTONIA

Ida published about 50 articles in magazines and journals in addition to her book. She
also was an active member in the Women’s International League for Peace and Freedom, the
Philadelphia Teachers Union and the American Civil Liberties Union.

Survivors include a brother, Frank E. Kaplan, M.D., of Doylestown PA, a sister, Mae K.
Millstone of York PA, five nephews, and two nieces. ALFRED E. SCHUYLER.

Arthur Cronquist (1919-1992). There
was never a dull moment when travelling
with Art Cronquist. If we weren’t talking
botany, he would be telling jokes or sing-
ing; he could handle both difficult pieces
and plain doggerel with ease. At a res-
taurant he would carefully study the menu
and then order a hamburger, plain, and
water, "lots of water, bring a pitcher." He
loved to meet people and converse.

He had written profusely, and his
writings profoundly affected the course of
botanical thought. He wrote and spoke
with authority, as evident in his evolution
and classification book: "The book pres-
ents taxonomy as seen by Cronquist .. . I
make no pretense of equal time for oppos-
ing views. For other points of view, read
other authors" (Cronquist 1988).

Arthur Cronquist was born on 19
March 1919 in San Jose, California. His
parents divorced when he was 4 years old,
and he was raised by his mother whose
maiden name he kept for his own. As a
teenager he collected plants in Utah and
Idaho, and discovered several State re-
cords which led to his first scientific
publication in 1939: "New plant records in
Utah and Idaho" (Cronquist 1939).

Art began his undergraduate studies at
Idaho State University and majored in
range-management. His first field botany
course was taught by the late Dr. Ray J. Davis. One of the course requirements was the
preparation of a plant collection. Dr. Davis chose the two top students to study the two
largest families: the Poaceae and the Asteraceae. To determine who would be assigned each
family the two students flipped a coin and Art got the comps. He would eventually become
a world authority on the family, and among his many publications was the Asteraceae
treatment in Ray J. Davis’ Flora of Idaho (Cronquist 1952).

In 1938, at the age of 19, Art earned his B.S. degree from Utah State University where
he was influenced by the late Dr. Bassett Maguire—Art’s professed "professional father."

OBITUARIES 127

In 1940 Art married Mabel Allred, and during that same year he received his M.S. degree
from Utah State; his master’s thesis was a taxonomic revision of a group of asters (Cronquist
1943). A childhood injury left Art ineligible for military service during World War II; thus,
in the early 1940s he worked on his Ph.D. at the University of Minnesota. His major
professor was Dr. Carl O. Rosendahl, a former student of the German botanist Adolf Engler.
Rosendah] and Cronquist would later co-author "The goldenrods of Minnesota" (1945) and
"The asters of Minnesota" (1949).

In 1943, while still working on his doctorate, Art accepted a position at the New York
Botanical Garden. For one year he worked half-time on the tropical families Simaroubaceae
and Sapotaceae, and half-time on his dissertation, "Revision of the North American species
of Erigeron, north of Mexico" (Cronquist 1947). After defending his doctoral dissertation
in 1944, Art continued working another two years at the New York Botanical Garden.
However, he did not want to pursue a career in neotropical botany; thus, in 1946 he accepted
the position of Assistant Professor of botany at the University of Georgia.

Arthur Cronquist excelled at teaching. He maintained affiliation with academic
institutions throughout his career (Washington State University: Assistant Professor,
1948-1951, and Research Associate, 1953—1966; Columbia University: Adjunct Professor,
1964—1992; City University of New York: Adjunct Professor, 1968-1992), and he wrote
two college textbooks on general botany: J/ntroductory Botany (1961) and Basic Botany
(1973). Both books went through two editions and both appeared in Spanish, while
Introductory Botany also appeared in Italian. Many of his botany courses were field-
oriented, and some of his "field trips" covered over 5,000 miles by vehicle in three weeks
time. His door was always open to students (and anyone else), and he never hesitated to
assist students in the field. As my major professor, he and I traveled to Utah, Canada, New
England, and the southern Appalachian Mountains, largely in pursuit of eupatoriums but any
plant was fair game. I probably learned more botany in his pick-up truck than in any
graduate level course. When I discovered a new species of Eupatorium | assumed that my
major professor would co-author the diagnostic publication; but no, Art declined authorship
and as a result I learned-by-doing and grew as a botanist.

After teaching two years at Georgia, he accepted the position of Assistant Professor at
Washington State University. In 1951 he spent a year in Brussels, with his wife and two
children, working on the flora of the Belgian Congo for the national government of Belgium.
Then, after an absence of five years, Art returned to the New York Botanical Garden where
he remained for the next 40 years.

Arthur Cronquist probably contributed more to an understanding of the flora of temperate
North America than any other 20th century botanist. He authored or co-authored many of
the region’s major floristic works, including Vascular Plants of the Pacific Northwest
(1955-1969), Manual of Vascular Plants of Northeastern United States and Adjacent
Canada (1963, 1991), the Asteraceae for Vascular Flora of the Southeastern United States
(1980), and Flora of the Intermountain Region (1972—present). He was also involved in
varying degrees in the preparation of many other major floristic works.

Early in his career Art began studying the problems associated with the concept of a plant
Species. He was critical of the strictly reproductive species-concept, and formalized a
species-concept under which most plant taxonomists now work: "Species are the smallest
groups that are consistently and persistently distinct, and distinguishable by ordinary means"
(Cronquist 1978). The practical application of this concept was not always initially popular
with others. In 1945, while preparing a treatment of the Asteraceae for the New Britton &

128 BARTONIA

Brown Illustrated Flora (at the age of 26), Art published a paper on the genus Antennaria
in the journal Rhodora. At the time, over 30 species of Antennaria were generally
recognized from the northeastern states and adjacent Canada. Art recognized three species
and six additional infraspecific taxa poe 1945). Art’s treatment of the genus brought
a quick and biting response from Dr. tt Lyndon Fernald: "It is, then, at least surprising
to be told by one who blew in from . Wes so recently . . . that in all this diverse area we
have only three fairly well-marked species .... If his [Cronquist] treatment of Antennaria
is typical of what is to be expected for other stoups in the new Illustrated Flora, it would
seem that that work will be an abbreviated pocket-novel ..." (Fernald 1945). But Art was
undaunted by Fernald’s harsh review; throughout his career dat reserved a special passion
for taxonomically difficult and complex groups of flowering plants. When I asked him about
his treatment on the stemless blue violets (Gleason and Cronquist 1991), he replied: "If I
can’t tell the plants apart, I can’t expect someone else to." His taxonomic conclusions were
always based on extensive field, herbarium, and library study.

The general system of classification of flowering plants was in a moribund condition when
Art published his early taxonomic works. The popular but archaic system of Engler and
Prantl no longer reflected current evolutionary thought. In the 1950s Art began publishing
his rapes on a new system of angiosperm classification (Cronquist 1957, 1960, 1963,
1964, 1965). These sean culminated in three books on evolution and classification
(Cronquist 1968, 1981, 1988), including his magnum opus, An Integrated System of
Classification of Flowering Plants, currently one of the most widely used systems of
classification in North America, Australia, and China.

ta sikatapi service held at the New York Botanical Garden on 5 May 1992, Dr. Peter
Raven compared the botanical achievements of Arthur Cronquist with those of Carl
innaeus. ea other outstanding botanists of our day have likened him to Asa Gray. It
was a great privilege to have had Arthur Cronquist as a friend and mentor. He will be sorely
missed by all who knew him

LITERATURE CITED

CRONQUIST, A. 1939. New plant records in Utah and Idaho. Leafl. W. Bot. 2: 210-211.

———. 1943. Revision of the western North American species of Aster centering about Aster foliaceus Lindl.
Amer. Midl. Naturalist 29: 429-468.

—. 1945. Notes on the Compositae of the northeastern United States. I. Inuleae. Rhodora 47: 182-184.

——. 1947. Revision of the North American species of Erigeron, north of Mexico. Brittonia 6: 121-300.

1952. Compositae. Pages 660-800 in R. J. Davis. Flora of Idaho. W.C. Brown Publ. Co., Dubuque,

. 1952. Compositae. Pages 323-545 in H. A. Gleason. The new Britton and Brown illustrated flora of the
northeastern United States and adjacent Canada. Vol. 3. Hafner, New Yor

1957. Outline of a new system of families and orders of tyes. Bull. Jard. Bot. Brux. 27: 13-40.
—. 1960. The divisions of classes of plants. Bot. Review 26: 4

———. 1961. Introductory Botany. Harper & Row, New York.

———. 1963. The taxonomic significance of evolutionary parallelism. Sida 1: 109-116.

———. 1964. The old systematics. Jn C. Leone, ed. Taxonomic biochemistry and seriology. Ronald Press.
———. 1965. The status of the general system of classification of flowering plants. Ann. Mo. Bot. Gard. 52: 281-

——. 1968. The evolution and ae ibe flowering plants. Houghton Mifflin, New York.
——. 1973. Basic Botany. Harper & Row, New Yo
Once again, what is a hari* Beltsville Symposia in Agricultural Research, No. 2, pp. 3-20.
Biostematics in oe
1980. Vascular flora of the southeastern United States. Volume 1. Asteraceae. Univ. of North Carolina
Press, cua Hill.

OBITUARIES 129

1981. An integrated system of classification of flowering gue Columbia Univ. Press, New York.

. 1988. The evolution and classification of flowering plants, 2nd ed. N.Y. Botanical Garden, Bronx.
——, A. H. HOLMGREN, N. H. HOLMGREN . REVEAL, AND P. K. pseu niits . 1972—present. Intermountain
flora. Vascular plants of the Seianieertann West (6 vols.). Hafner Publ. Co.; Columbia Univ. Press; N.Y

Botanical Garden.
FERNALD, M. L. 1945. Contributions from the Gray Herbarium of Harvard University—No. CLVII. I. Key to
Antennaria of the "Manual Range." Rhodora 47: 221-235, 239-247.
GLEASON, H. A. AND A eS lteei 1963. Manual of vascular plants of northeastern United States and adjacent
Canada. Van Nostrand, Princet
1991. bine of vascular plants of northeastern United States and adjacent Canada, 2nd ed.
N.Y. Botanical Gasicn, Bronx.
HITCHCOCK, C. L., A. CRONQUIST, M. OWNBEY, AND J. W. THOMPSON. 1955—1969. Vascular plants of the Pacific
Northwest (5 vols). Univ. Wash. Press, Seattle.
ROSENDAHL, C. O. AND A. CRONQUIST. 1945. The goldenrods of Minnesota: a floristic study. Amer. Midl.
aie 33: 244-253.
———. 1949. The asters of Minnesota: a floristic study. Amer. Midl. aap 42: 502-512.
C E. LAMONT

Thomas J. Day (21 March 1938-2 July 1992). Of Mahanoy City, Pennsylvania, and a
long-time member of the Philadelphia Botanical Club, Tom had a great interest in botany
from his days as a Boy Scout. He spent many hours in local woodlands studying the flora
and kept extensive records of the plants he found. He was an avid hunter and fisherman as
well as gardener and amateur cook. Tom also assembled a large library of books on plants
and gardening. He attended the Pennsylvania State University and Lehigh Community
College, and was pursuing the bachelor of science degree at the time of his death. He is
survived by his wife Betsy and his son Daniel. ELIZABETH B. FARLEY

Bartonia No. 58: 130-137, 1994
REVIEWS

Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd
ed. by Henry A. Gleason and Arthur Cronquist. New York Botanical Garden, Bronx.
1991. Ixxv + 910 pp. $74.60 incl. shipping (cloth).

The Gleason and Cronquist manual has returned looking very similar in appearance to the
first edition. However, upon closer inspection the reader will notice that the manual has
changed considerably.

Changes are noted upon first opening the book. The front endpapers provide a map

indicating the region covered. With the senior author, Henry Gleason, having died in 1975,
Cronquist in the preface now takes full responsibility for the entire contents of the manual
(p. v).
The glossary is now much more complete with 967 entries, 378 of which are new.
Cronquist has done an excellent job providing succinct, accurate definitions that are
understandable to all. The glossary now includes many terms used but not defined in the
first edition, such as cuneate, ray, spikelet, and style. However, some terms defined in the
first edition are no longer defined in the new edition. Students would have been better
served if basal, rhizophore, seed, and stem had been retained in the glossary.

The general keys that follow the glossary are superb. Out of all the manuals I have used,
I have found the general keys in the Gleason and Cronquist manual to be the easiest to use.
The general keys are now strictly dichotomous, and this adds to their simplicity and ease of
use. There has been some realignment of the general sections, which now total eighteen.
However, a couple of things in the general keys may cause confusion for the beginning
botanist. In section 10, in referring to the perianth, the term cycle is used while in section
11 the term circle is used. The term cryptogam is used, although in a current standard
botany text (Raven et al. 1992) this term is referred to as "archaic."

New to this edition are the synoptical keys to the angiosperms which precede the general
keys. These keys are limited to the angiosperms, the taxa addressed in Cronquist’s
Integrated System (which the manual is now arranged on). They will certainly prove helpful
to taxonomy instructors in teaching the natural order of the angiosperms. However, there
is a problem with the type size used in the synoptical key headings. The heading of the key
to the subclasses of Magnoliopsida is in large type, followed by keys for orders and families
of each subclass in smaller type. The subsequent key to the subclasses, orders, and families
of Liliopsida is also headed in the smaller size type. This gives the misleading impression
that the synopses deal only with the Magnoliopsida.

The text follows the same format as edition one, but there are plenty of taxonomic
changes. Breweria is now Stylisma. Psilocarya and Dichromena are now in Rhynchospora.
Manisuris is now Coelorachis. Some species of local interest have been lost not to
bulldozers but to lumping. These include: Panicum hirstii, Rhynchospora microcephala,
Scleria nitida, and Scirpus ancistrochaetus.

The nomenclatural nightmare that exists in Xyris could have been better dealt with in the
manual. Xyris flexuosa of the first edition is now Xyris caroliniana in the new edition.
Xyris caroliniana of the first edition is now Xyris difformis in part and Xyris jupicai in part.
In the new edition, Cronquist mentioned X. flexuosa in synonymy under X. caroliniana but

130

REVIEWS 131

failed to mention the misapplication of the former X. caroliniana under X. difformis and X.
jupicai. This could give the misleading impression that X. flexuosa merely has been lumped
into the X. caroliniana of the first edition.

Sure to create controversy are the new treatments for Dichanthelium and the stemless blue
violets. For Dichanthelium, Cronquist claimed to have "drawn heavily" (p. v) from Gould
and Clark (1978). However, Gould and Clark recognized only 22 species for the manual’s
range and treated Dichanthelium as a genus, while Cronquist recognized 32 species and kept
Dichanthelium at the subgeneric level under Panicum. The most striking example of
lumping in this new treatment occurs with P. dichotomum, which now includes P. annulum,
P. lucidum, P. mattamuskeetense, P. nitidum, and P. roanokense of the first edition. For the
stemless blue violets, Cronquist as drawn on his "own observations in the field over several
decades" (p. v). Apparently his viewpoints over the past three decades have changed
drastically: in the new edition only 6 species are recognized compared to 24 in the first
edition. Most of the formerly recognized species have fallen into synonymy under either
Viola sororia or V. palmata, making these two species so broad that they are as difficult to
comprehend as the numerous cryptic species recognized in the first edition. For both
Dicanthelium and Viola the reader is given very little evidence to substantiate this radical
changes.

The distributional information is significantly improved from the first edition, but there
are still more errors than I would like to see. Maine is listed in the range of Monotropsis
odorata, although it should be Maryland. Andropogon ternarius, Listera smallii, Salix
serrisima, Utricularia inflata, and Xyris caroliniana all have been recorded from New Jersey,
but the manual does not include New Jersey in their ranges. Utricularia olivacea is not
included in the manual although it occurs in New Jersey. The manual has Helonias bullata
formerly occurring in New Jersey, yet New Jersey has more extant populations of Helonias
than any other state in the plant’s range. Information on the distribution of weeds could
have also used some updating. Describing Microstegium vimineum as only "sparingly
introduced" and Bidens polylepis as only "occasionally adventive eastward" may have been
accurate in the 1960s but it certainly is not now.

The index suffers from numerous typographical errors and errors of omission. Recognized
names for plants established in the range are listed in Roman type with all other names in
italic type. However, I have noticed numerous instances where the wrong type was used for
a taxon. Several synonyms used in the text are omitted in the index. The recognized taxa
Mollugo, Saxifraga rivularis, and Scirpus pungens are also omitted. Species for Carya,
Cuscuta, Hedyotis, Linum, Myriophyllum, and Sparganium are not indexed even though 10
Or more species are recognized for each genus. The reader is also referred to the wrong page
for Amianthium, Corema, ironweed, Jerusalem-artichoke, and Melampyrum.

Lastly, this new edition, like the first edition, fails in not providing a summary of the
number of families, genera, and species established in the range. Willing to wait no longer,
I have counted them as follows: 192 families, 1080 genera, and 4271 species.

I recommend this book to any field botanist, botany student, or taxonomist working near
or in the range of the manual. Despite the criticisms, it is the only recent manual for the
northeastern US available. Cronquist did us all a great service by getting this published
before his unfortunate death on 22 March 1992.

13Z BARTONIA
REFERENCES

GOULD, F. W. AND C. A. CLARK. 1978. Dichanthelium (Poaceae) in the United States and Canada. Ann. Missouri

Aes Gard. 65: 1088-1132.
RAVEN, P. H., R. F. EVERT, AND S. E. EICHHORN. 1992. Biology of plants. Worth Publishers, Inc., New York.
GERRY MOORE

New Jersey Ferns and Fern-Allies, by J. D. Montgomery and D. E. Fairbrothers. Rutgers
Press, New Brunswick. 1993. 300 pp., 100 maps, and 125+ line drawings. $45.00.

For those who are fortunate to own a copy, or who have access to a library copy of
Chrysler and Edwards’ 1947 Ferns of New Jersey (a very rare book, with supposedly only
700 copies printed, most of which went to libraries and colleagues and friends of the
authors), the eagerly awaited new New Jersey fern book, by Montgomery and Fairbrothers,
is a little like running into an old friend who has undergone a face lift. But unlike some face
lifts or revisions that obliterate, muddle, and deface, this one not only preserves much of
what worked in the 1947 fern book, but invigorates their book with the 46 years of
additional taxonomic research and distribution data that had passed-by and outdated much
of the Chrysler and Edwards book. This revision is much more than cosmetic, and
everything in this book, from text to maps to illustrations, has been rethought, rewritten, and
revised. The result is a concise, competent, and readable treatment, which makes this book
the new standard on the distribution, diversity, and identification of New Jersey ferns and
fern-allies.

One of the strong points of the Chrysler and Edwards book, was its format, which this
book largely adopts. This first five sections—Historical Introduction, Fern Structure and
Classification, Hybrids, keys, Ecology and Distribution—are similar in scope to those of the
1947 book. However, all these sections have been greatly expanded and contain much more
information. The Historical Introduction is particularly well done; being both informative
and entertaining, it nicely sets the context. For the new student of pteridophytes, the section
on Fern Structure and Classification is very helpful. In it, the technical terms used in the
book are introduced, clearly defined, and illustrated. And for those whose knowledge of
New Jersey is limited to car window views from the state’s major highways, the section
Ecology and Distribution provides an excellent overview of New Jersey’s geology,
geography, and generalized habitats. The listing of references is extensive and is almost
worth the price of the book alone.

A single page per taxon is used to describe each species or subspecific taxon. For each,
its latin name, major synonyms, and common name(s) head the page. This is followed by
brief notes on description, habitat, general distribution, distribution in New Jersey, and
general comments. The arrangement is fern-allies first, true ferns second, with the genera
grouped by family. The facing page is devoted to a distribution map and line drawings of
the taxon. Nearly all taxa discussed are illustrated and their distribution within New Jersey
mapped. This is also the format of the 1947 book, although not quite as assiduously adhered
to. The new fern book’s dimensions are virtually identical to the old fern book, and at 23.5
cm x 16 cm, it can be conveniently carried into the field.

A tremendous amount of new information is presented in the new book. For example,
Chrysler and Edwards recognized 8 families, 29 genera, 78 species, 16 subspecific taxa, and
16 hybrids, while Montgomery and Fairbrothers now recognize 15 families, 30 genera, 80
species, 10 subspecific taxa, and 32 hybrids as occurring in New Jersey. A total of 118 taxa

REVIEWS 133

(by my tally, not the authors’ 114 on p. 25) are included in the checklist—a remarkable
diversity for such a small, and densely populated state. Even more remarkable is the
statement on page 41, that 94 taxa of ferns and fern-allies have been documented from
Sussex County, nearly one quarter of total taxa reported from the continental United States.
The distribution maps are based on collection localities that are vouchered by herbarium
specimens and are plotted on a county outline map of the state. Three different sized dots
are used to indicate the relative age of the collections. An oversight of the book is that
herbaria examined are not acknowledged. According to Montgomery (pers. comm.) the
following collections were examined: Chrysler Herbarium, Rutgers University, New York
Botanical Garden, Academy of Natural Sciences of Philadelphia, and the Smithsonian
Institution were checked for records of all taxa, with selected genera or taxa examined at
Berlin, British Museum, Drew University, University of Michigan, Montclair State
University, Newark Museum, Rutgers-Camden, Staten Island Museum, and Upsala College.

The book does contain some omissions, errors, inconsistencies, and other problems. My
greatest criticism is the generally poor layout and reproduction of the illustrations. This is
a shame, because the line drawings greatly enhance the usefulness of the book, and, as an
aid in identification, frequently surpass the black and white photographs used in the 1947
book. However, may of the illustrations are too small, and the details are often difficult, or
in the case of some species of Equisetum, nearly impossible to discern. No scale is provided
for the size of the reproductions, and there is no consistency of scale between illustrations
or even within genera. There is a lot of unused, blank space in the book that could have
been better used by filling them with full page illustrations. The distribution maps could
have then been placed as insets within non-critical areas of the illustrations, or better yet,
moved to the text page. The distribution maps also suffer from being reproduced a little too
small, and, as a result, the smallest sized dots are sometimes obscured by county lines or
county names. Going to a slightly larger format may have solved the problem.

Some other comments, more or less from the front to back, are as follows. The keys are
awkwardly placed between the sections Hybrids and Ecology and Distribution, rather than
immediately preceding the section describing the taxa or the very beginning of the book
where they would have been a little more convenient to use. Table 2 on page 41 lists
Lycopodium inundatum as being restricted to the Ridge and Valley and Highlands Provinces,
whereas the distribution map (p. 73) for this species accurately shows occurrences in four
Coastal Plain counties. The illustrations for Ophioglossum pusillum and O. vulgatum (pp.
143 and 145) are reversed. The second sentence under comments on page 144 should read
"having no known extant occurrence." The Appalachian gametophyte (Trichomanes sp., p.
168) is neither keyed nor illustrated. Granted, a working key to it is probably hopeless
without including look-alike algae and mosses. An illustration, however, would have greatly
facilitated identification, since it is probable that only a handful of the book’s readers have
ever seen this most obscure of all of New Jersey’s ferns. After publication of this book, it
was described as a new species, Trichomanes intricatum Farrar (for a detailed description,
illustration, and distribution see Farrar 1992). The index is inconsistent in its inclusion of
people, places, and subjects. For example, St. Patrick (p. 1), Quaker Bridge (p. 152), and
snow (p. 35), are in, but Napoleon (p. 232), Blairstown (p. 194), and condoms (p. 56), are
not

Non-native taxa like the introduced (and presumably planted) Marsilea quadrifolia (p.
274) and the adventive Azolla caroliniana (p. 276) are keyed, described, illustrated, and
included in the checklist (p. 278), while Phyllitis scolopendrium, another established

134 BARTONIA

introduction, is not, and is only briefly discussed under Asplenium rhizophyllum on page 192.
Adding this species to the checklist would then bring the total of taxa of New Jersey ferns
to 119. It should be noted that there are additional taxa covered in the book that are not
included in the checklist or tallies. Some, like Polypodium appalachianum (p. 170),
Asplenium trichomanes subsp. quadrivalens (p. 198), an smunda cinnamomea var.
glandulosum (p. 147) are generally accepted as valid taxa, while others like the three
varieties of Botrychium simplex (p. 134) are no longer considered as taxonomically
significant. So at least 122 taxonomically valid taxa of New Jersey ferns and fern-allies are
discussed in this book—eight more than the 114 advertised.

I suppose that like any face lift, a certain amount of blemishes and wrinkles are bound to
be missed. Fortunately for the users of this new fern book, the imperfections are mostly
minor, and do not seriously detract from the value and usefulness of the book. As a
revision, it is an improvement over the old, and solidly advances our knowledge of New
Jersey’s ferns. I give it two fronds up.

REFERENCES
CHRYSLER, M. A. AND J. L. EDWARDS. 1947. The Ferns of New Jersey. Rutgers University Press, New
Brunswic
FARRAR, D. R. 1992. Trichomanes intricatum: The independent Trichomanes gametophyte in the eastern United
States. Amer. Fern J. 82: 68-74.
DAVID B. SNYDER

British Plant Communities. Volume 1. Woodlands and Scrub, edited by J. S. Rodwell.
Cambridge University Press, New York. 1991. x + 395 pp. $160.00 (cloth).

This book is the first in a five-volume series that will provide a comprehensive, uniform
description of the vegetation of Great Britain and adjacent islands, excluding Ireland.
Volume 1 begins with the history of the project and the methods of collecting, analyzing,
and presenting the data. This general introduction is followed by an overview of the
edaphic, climatic, and successional relationships of the five major groups of woodland and
scrub community types: "mixed deciduous oak-birch woodlands," "beech and yew
woodlands," "pine and juniper woodlands and montane willow scrub," "wet woodlands with
alder, birch and willows," and "scrub and underscrub communities." The overview
concludes with a key that distinguishes the community types and sub-types and describes
their intergradation and regional variation. The categories are based on physiognomy and
floristics and are named after one to three of the most frequent species—for example

"Quercus petraea-Betula pubescens-Dicranum woodland," "Salix lapponum-Luzula sylvatica
scrub," "Holcus lanatus sub-community."

Most of the volume consists of detailed accounts of each of the 25 community types. The
treatments all follow the same format, beginning with a list of synonyms for the community
type plus lists of its constant species and rare species. Next come descriptions of
physiognomy, sub-communities, habitat (including he climatic, biotic, and human
factors), zonation and succession, distribution in Great Britain, and affinities (i.e. correlations
with European phytosociological literature and distinctions from related community types).
The descriptions are followed by a floristic table with a species list of vascular plants,
bryophytes, and lichens; plus columns representing their presence in the respective sub-
community types and the community type as a whole. The species are grouped according
to stratum, frequency, and sub-community. Each species in each column is given a

REVIEWS 135

frequency value for the percentage of stands in which it occurs, and a range of abundance
according to the Domin scale of cover classes. The table concludes by summarizing the
altitude, slope, and species richness of the samples; and height and percent cover of their
four strata (tree, shrub, herb, and ground). Each accounting concludes with outline maps of
Great Britain containing dots for the community locations according to a grid system of 10

e final sections of the book are an index to synonyms of the community
types, an index to species, and a bibliography.

This series will be valuable to botanists and ecologists of northeastern North America.
Features of the methodology could be applied here, and the floras of this region and
northwestern Europe have many species in common. The series is appealing because of the
detail and integration of its content; the informational classification of community types (in
contrast to Braun-Blanquet nomenclature); the consistent use of scientific binomials; the
abundance of charts, tables, diagrams, and maps; and the lucid style of writing. The printing
and binding are of good quality. Besides the volume reviewed here, two of the other
volumes in this series are now available (at $195.00 each): Volume 2, Mires and Heaths
(1991); and Volume 3, Grasslands and Montane Communities (1992). Despite its high cost,
the entire series should be acquired by libraries of universities with programs in botany or
ecology. LARRY H. KLOTZ

Easter Island Earth Island, by P. Bahn and J. Flenley. Thames and Hudson Inc., New
York. 1992. 240 pp., fully illustrated with maps and photographs. $24.95.

The remoteness of Easter Island perhaps is best appreciated from aboard a jet airliner as
it begins its descent and starts its approach to the only landing strip available for thousands
of kilometers. Santiago, Chile, more than 3,750 km to the east, is some five hours away.
The nearest airport west of Easter Island is in Papeete, Tahiti, and is over 4,000 km and six
hours away. You pray that nothing goes wrong, and that the wheels find the tarmac, and not
the turbulent waters of the Pacific Ocean, which have been pounding and slowly eroding the
cliff-lined shores of this volcanic mote for its three million years of existence above the sea.
The plane, of course, is guided by twentieth century technology, but the small band of
polynesians that set sail from eastern Polynesia sometime in the first centuries AD, reached
Easter Island by using a combination of navigational skills developed over thousands of
years of colonization of the islands and atolls of Polynesia, as well as an extraordinary bit
of good luck.

Where these voyagers came from, why they came, the geology, climate, flora and fauna
of the island they settled, how they existed, how they carved, transported, and erected the
massive, monolithic moai (statues), European discovery of the island, and remarkable new
insight into the probable causes of the warfare and violence that ultimately brought the near
extinction of one of the world’s most remarkable prehistoric cultures, are explored in Easter
Island Earth Island. By using the most recent archaeological, ethnological, linguistic,
physical anthropology, and botanical data to support their theories, the authors minimize the
use of conjecture and speculation, which has been a common failing of many other popular
books on Easter Island.

Through the years the debate over the origin of the people of Easter Island has been the
Subject of much controversy. Some, like Erich Von Daniken’s theory that explained all the
mysteries of Easter Island through the intervention of benevolent ancient astronauts, can be
dismissed without need for further comment. Other theories, although equally implausible,

136 BARTONIA

have gained general acceptance from the lay public. Number one among these is Norwegian
explorer Thor Heyerdahl’s theory that Easter Island was first settled by Amerindians sailing
from Peru. Heyerdahl’s books promoting a South American origin for the culture of Easter
Island have been so popular, that the authors devote much space to countering many of
Heyerdahl’s arguments. For example, Heyerdahl has claimed that the totora (Schoenoplectus
californicus, but identified as Scirpus riparius in the book) used by the Peruvians to build
their reed sailing boats is identical with the Easter Island plant now growing on the shores
of the fresh water lakes in the calderas of the island’s volcanoes and, since the major method
of reproduction is by rhizomes, the species clearly had to be transported by humans to the
island. The authors counter this argument by stating that the variety of this species now
growing on Easter Island is distinct from varieties occurring in Peru and, since the species
obviously reproduces from seeds, totora could have easily been transported to the island by
birds, noting that Darwin washed the seeds of 52 plants from the feet of water birds. Recent
pollen analysis has shown that totora has been present on Easter Island for at least 30,000
years, so dispersal by humans, regardless of which way they sailed, is no longer tenable.
Like floating ducks at a carnival shooting gallery, Heyerdahl’s arguments are shot down one
by one, leading the authors to conclude, "Heyerdahl’s theory of a South American source for
Easter Island culture is indeed a tottering edifice precariously based on preconceptions,
extreme subjectivity, distortions and very little hard evidence. In putting it together over the
years, he has come to resemble someone who has painted himself into a corner with no
means of escape, but is loath to admit it."

For readers not particularly interested or amused by academic shenanigans, the overriding
issue presented in Easter Island Earth Island is an ecological one, centering on the dangers
of over-exploitation of limited natural resources and the globally relevant consequences of
that action. We are all familiar with the "canary in the coal mine" analogy used to generate
support for the preservation of species, fragile ecosystems, and overall biological diversity.
Or the one that compares the loss of endangered species to the loss of bits and pieces of a
jet airliner. Although still frequently invoked by defenders of diversity, both of these
analogies are really a couple of old chestnuts, and are about as viable as the American
species of that tree. Even if we lost every rare and endangered plant species currently extant
in New Jersey, it would be unlikely that this would be enough to set into motion a domino
effect that would ultimately unravel any given ecosystem, let alone bring about the collapse
of civilization. Our flora, and the ecosystems they are dependent on, are too diverse and
resilient to be taken down by the loss of such a relatively minor portion of the overall
biomass. However, if this scenario were to be played out on a different stage, one where
biological diversity is depauperate, where critical links between species and ecosystem are
real, where equilibrium between the two are tenuous at best, the loss of even a few species
could be enough to tip the balance. This is apparently what happened on Easter Island.

The story of Easter Island is, as the authors conclude, a cautionary tale that carries two
timely warnings. The first is that without wise management, all natural resources are finite.
Despite the obviousness of the statement, it is a warning that remains largely unheeded by
most people on our planet. The second warning comes in the form of a lesson. In the
author’s words, "Easter Island is small, and its ecosystem relatively simple. It has limited
topography, a rather stable climate, a limited range of rocks, soils, plants, and animals.
Whatever one did to alter that ecosystem, the results were reasonably predictable. One could
stand on the summit and see almost every point on the island. The person who felled the
last tree could see that it was the last tree. But he (or she) still felled it. This is what is so

REVIEWS 137

worrying. Humankind’s covetousness is boundless. Its selfishness appears to be genetically
inborn. Selfishness leads to survival. Altruism leads to death. The selfish gene wins. But
in a limited ecosystem, selfishness leads to increasing population imbalance, population
crash, and ultimately extinction."

With this new insight, many of the old riddles and mysteries of Easter Island no longer
seem to be quite so compelling. From this the most isolated inhabited flyspeck of
geography, a new mystery is born, global in its reach: Will humankind learn the lesson of
Easter Island? DAVID B. SNYDER

Bartonia No. 58: 138-140, 1994
NEWS AND NOTES

SUPPLEMENT TO NEW JERSEY WILD PLANTS. This 1992 supplement corrects the nomencla-
ture to that used in the 1991 second edition of Gleason and Cronquist’s Manual of Vascular
Plants of Northeastern United States and Adjacent Canada. This supplement, plus a 1989
list of species reported since the book was published in 1983, and some corrections, will be
sent postpaid for $1.00 to anyone wishing the information. Order from Mary Y. Hough, #1,
Route 94, Blairstown, NJ 07825. The book is also still available for $14.95 postpaid from
the same address, and would include the supplements. MARY Y. HOUGH

1991 AND 1992 NEW CASTLE COUNTY, DELAWARE, RARE PLANT SURVEY HIGHLIGHTS. The
1991 botanical field season proved, as in earlier years, to be both interesting and exciting for
us in the way of new discoveries and old locations of both historical and extant species in
the state. Beginning in the Brandywine drainage, some notable finds included the second
DE population of Smilax hispida, formerly known only from the Mt. Cuba area, a third DE
station for Carex davisii, several new populations of Panax quinquefolius, Aster pre-
nanthoides (a former SH species), and the second DE population of Heliopsis helianthoides
on the PA/DE state line. A major change was the discovery, partly through a discussion
with fern expert Donnell Redman, and confirmed in the field, was that our DE population
of Cystopteris bulbifera was actually C. tennesseensis. The most exciting find in the
drainage, though, and perhaps of the field season, was the confirmation of the existence of
a small population of Asplenium trichomanes, which had been ranked SX and had not been
seen in the state in nearly 90 years. Its current presence had first been noted in a Delaware
Nature Society publication. Near this fern also grew populations of other rare species,
including Hybanthus concolor, Poa sylvestris, Conopholis americana, and Cystopteris
protrusa, many growing on rich wooded slopes dominated by an enormous population of
naturalized Trillium grandiflorum.

The Red Clay valley did not yield quite as exciting finds as the Brandywine drainage, but
still produced a number of surprises, the best being a single plant of Aster laevis (former SH)
found growing on a serpentine roadcut. Other interesting sightings included a small
population of Caltha palustris, two colonies of Amianthium muscaetoxicum, Eupatorium
sessilifolium, Ostrya virginiana, and Dryopteris celsa, the last in a roadside seep.
Discoveries in the other Piedmont drainages included a small colony of Poa autumnalis in
the Middle Run valley, Lilium canadense in the White Clay, and the state’s largest current
population of Lobelia spicata in the Mill Creek drainage along with a small but dense colony
of Scleria pauciflora at its second DE station.

On the upper coastal plain and Fall Line region of New Castle County, including Iron and
Chestnut hills, occurred a mix of Piedmont and Coastal Plain rarities. Chestnut Hill yielded
Botrychium matricariaefolium and Ophioglossum pycnostichum, while Liparis lilifolia was
found on Iron Hill. A roadside and young woods near the latter’s base yielded Angelica
venenosa at its first confirmed recent sighting, along with Carex lanuginosa, more
Ophioglossum pycnostichum, and a single male plant of Thalictrum revolutum, a new DE
record. Lastly, a small population of the elusive Aster radula was found near a tributary of
the Christina River.

Farther south, our explorations in one of the Blackbird State Forest tracts yielded

138

NEWS AND NOTES 139

Lespedeza hirta, Agrimonia rostellata (former SH), Kalmia angustifolia, and Eupatorium
sessilifolium, among more than a dozen DE rarities. Clitoria mariana was also discovered
near another Blackbird Forest tract. Several localities for the rather local Lysimachia hybrida
were discovered in the Smyrna River and Cypress Creek drainages. Apocynum androsaemi-
folium, which we expected more likely to be found in the Piedmont, actually turned up in
great abundance along a disturbed roadside and in a powerline cut, the latter population
growing with or near a number of leguminous rarities such as Desmodium marilandicum,
laevigatum, and viridiflorum and Lespedeza hirta as well as Gentiana saponaria. A small
population of Hypericum gymnanthum turned up in a recently sprayed powerline cut, along
with Aletris farinosa in great abundance. Finally, near the end of the season, exploration of
a powerline cut along the Maryland border yielded a large population of Cacalia
atriplicifolia well away from its more Piedmont haunts along with the previous SH species
Lechea villosa.

In 1992 on the Piedmont, finds in the Brandywine drainage included the second DE
population for Corallorhiza odontorhiza, another station for Lobelia spicata, and two new
stations for Scirpus pendulus. The Red Clay produced Aplectrum hyemale (on the PA/DE
state line), Asclepias exaltata blooming along a much-traveled roadside, and two stations for
Lilium canadense. Further exploration of the serpentine outcrops near Hoopes Reservoir
with Bill McAvoy led to a second DE station for Carex bushii, and on the closely mown
grounds of an estate, more Cerastium arvense var. villosum in a population extension of a
previously known roadbank station along with a colony of Senecio anonymus, previously
thought extirpated in the state. Along a Piedmont roadside passed frequently by botanists,
we found Chamaelirium luteum and Sanicula marilandica in bloom, both SH species not
seen in the state for 50 years, along with Zizia aptera at its third DE station.

The Mill Run drainage provided the largest population of Dryopteris goldiana in the state.
In the White Clay valley, finds included thousands of plants of Corydalis flavula growing
along an old railroad bed on E. I. DuPont property, and the largest population yet for
Aplectrum hyemale in the state. Finally, on the Piedmont outliers of Iron Hill and Chestnut
Hill south of Newark, additional populations of $1 species Eupatorium sessilifolium and
Angelica venenosa were found on the former while the SH species Veronicastrum virginicum
was found on the latter on I-95 roadbank.

In the Fall Line zone and Coastal Plain of central and southern New Castle County, a
number of rare species were found to be much more common than previously thought. A
chance stop along a roadside led to the discovery of large populations of both Alopecurus
carolinianus and A. aequalis, at that time both S1 DE species, along with the alien Myosurus
minimus, a new state record. Further searches of the same habitats—wet mucky fallow
cornfields—revealed an abundance of both grasses along with several stations of a new state
record, Myosotus macrosperma (first discovered in the White Clay valley), causing some
concern about its nativity and whether or not the two grasses have beocme naturalized in this
habitat since Tatnall’s flora of 1946. A third grass previously thought quite rare but perhaps
more common than previously recorded, Glyceria acutiflora, was found in several Carolina
Bays and may be in many more. Deschampsia flexuosa, previously a DE SH species, was
rediscovered at Noxontown Pond and at several other stations in like habitat—steep slopes
under oaks with little or no leaf litter in firm soils.

An unusual upland Piedmont-like habitat under and near a powerline cut in the Sandy
Branch drainage, first noted at the end of the 1991 highlights, yielded Angelica venenosa
(one plant in flower), Caltha palustris, Ceanothus americanus, Deschampsia flexuosa,

140 BARTONIA

Helianthemum propinquum, Helianthus divaricatus, and Paronychia canadensis. This was
the first Coastal Plain record for H. divaricatus. A small population of Luzula acuminata,
just across the state line in MD, was also found. Solidago arguta was also discovered ina
similar but much smaller habitat along Drawyers Creek. A list of other Coastal Plain
highlights included Callitriche terrestris (DE SH) at a degraded pond south of Wilmington,
Aster infirmus (DE SH) and Agalinis tenuifolia (DE $1) in the Blackbird region (the latter
probably soon to be eradicated by housing), Cornus stricta (new state record) in a freshwater
marsh near the Appoquinomink where Equisetum fluviatile had recently been rediscovered
by Keith Clancy, Scutellaria galericulata and Juncus torreyi (both $1) in the marshes near
Deemer’s Beach, three stations for Hedyotis uniflora (along with Isoetes engelmannii at one
of them) that extends its range north to New Castle County, Malaxis unifolia near Cedar
Swamp, Matelea caroliniensis in fruit and flower on the DE/MD state line, and a station for
Hypericum drummondii (new state and second Delmarva record) near Glasgow on DuPont
property in central New Castle County. JANET EBERT AND JACK HOLT

Bartonia No. 58: 141-147, 1994

1991 FIELD TRIPS

April 26: Pine Barrens of Burlington and Ocean cos., NJ. This trip was a bus tour co-
sponsored by the Divison of Pinelands Research at Rutgers University and the club in
conjunction with our symposium of April 25, "Research Perspectives in the Pinelands." On
the previous evening, trip participants heard Ted Gordon’s slide-illustrated lecture, "Flora and
Habitats of the Pine Barrens," a preview of some of the selected sites. En route, David
Fairbrothers presented a capsuled history of botanists who explored the Pine Barrens since
the 18th century. At the Pinelands Biosphere Reserve Research Center near New Lisbon we
were met by Ralph Good and students, who presented an overview of their current and past
research. Our first stop was a mature upland oak-pine forest near the Lebanon Headquarters.
At the next stop at Shinn Branch, we compared three forest types: pitch pine lowland, white
cedar swamp, and pine-blackjack oak. We proceeded to the Lower (East) Plains where
possible causes of stunting of this pygmy forest were discussed. The root crown of the
blackjack oak (Quercus marilandica) was excavated to expose lignotubers, which were the
subject of considerable discussion. Our final stop was a lowland pitch pine forest at Atison.
Here we assessed the impact of a severe wildfire (1983) on species composition and rare
herbs. Leaders: David E. Fairbrothers, Alfred E. Schuyler, William Olson, Joseph Arsenault,
and Ted Gordon.

May 4: Mount Cydonia Ponds and Mountain Run Ponds, Franklin Co., PA. This trip
emphasized the seasonal wetlands that occur along the west flank of South Mountain. They
were filled with water on the date of the excursion. We examined two of the Mount
Cydonia Ponds, which are in the Michaux State Forest south of Caledonia State Park; and
three of the Mountain Run Ponds, which are on private land east of Scotland, Pa.
Noteworthy finds included two large stands of Woodwardia virginica, a spectacular floral
display of Orontium aquaticum, plus flowering populations of Aronia arbutifolia, Vaccinium
corymbosum, Nuphar lutea, and Carex vesicaria. The taxonomic difficulties in Aronia were
discussed. The surrounding forests also contained several species in bloom: Rhododendron
periclymenoides, Cornus florida, Viola fimbriatula, V. primulifolia, Cypripedium acaule, and
(in bud) Jsotria verticillata. In the late afternoon, we observed vegetative plants of /ris
verna in a pitch pine-scrub oak barren along the Appalachian Trail on "Big Flat" (the summit
of South Mountain). Subsequently, while returning to Caledonia, we were pleased to find
this species in bloom on a grassy roadside along Route 233. Leaders: Larry Klotz and
Jeffrey Walck.

May 5: Licking Creek, Franklin Co., PA. We began by visiting a mesic upland woods
on the limestone and calcareous shale of the Onondaga and Old Port formations. The forest
Overstory mostly consisted of Acer saccharum, Liriodendron, Tilia, Quercus spp. and Carya
spp. The herbaceous layer was luxuriant and had an unusual composition for southcentral
Pennsylvania. Notable herbs were Delphinium tricorne (including one plant with very pale
purple flowers), Ranunculus micranthus (as well as R. abortivus), Jeffersonia diphylla, Vicia
caroliniana, Chaerophyllum procumbens, Hydrophyllum macrophyllum (as well as H.
virginianum), Phacelia purshii, Phlox divaricata, Aster shortii, Carex jamesii, and 7; rillium
sessile. The unexpected find was a cluster of 15 shoots of Corallorhiza wisteriana. In this

141

142 BARTONIA

same woods was a shale outcrop with Juniperus virginiana, Viola pedata (both concolorous
and bicolorous corollas), Antennaria plantaginifolia, and several small-flowered herbs such
as Lithospermum arvense, Myosotis verna, and Veronica arvensis.

After lunch in Robert Hunter’s yard, we visited another upland woods, this time on the
Wills Creek Formation. Some of the notable species that we observed were Ranunculus
hispidus, Dodecatheon amethystinum, Matelea obliqua, Phacelia dubia, and Carex
platyphylla. Along the roadside we encountered Lithospermum canescens and Zizia aptera.

We then crossed Licking Creek into Maryland to observe a calcareous cliff. Here we saw
Asplenium trichomanes, A. rhizophyllum, A. resiliens, Pellaea atropurpurea, Galium boreale,
Aquilegia canadensis, and Sedum telephioides. Leaders: Jeffrey Walck and Larry Klotz.

June 8: Bowmans Hill State Wildflower Preserve, Bucks Co., PA. Among the flowering
plants seen were: Pyrularia pubera, Dicentra eximia, Carex grayi, Gaylussacia brachycera,
Lonicera sempervirens and Paxistima canbyi. Some of the fern species encountered include:
Diplazium pycnocarpon, Deparia acrostichoides, Matteuccia struthiopteris, Cystopteris
protusa, C. bulbifera, Dryopteris goldiana, D. cristata, D. clintoniana, Cheilanthes lanosa
and Polystichum braunii. In addition, a tour of the propagation area at Bowmans Hill was
given by propagator, Tom Stevenson. Leader: Bill Olson.

June 23-27: Glassboro, NJ. The 1991 Joint Field Meeting of the Northeastern Sectiuon
of the Botanical Society of America, the Torrey Botanical Club, and the Philadelphia
Botanical Coub was held at Glassboro State College in Glassboro, New Jersey.

Three days were taken up with field trips. The first day’s excursions went to sites in the
New Jersey Pine Barrens, including the Atsion ore beds, the East Plains, and to cedar bogs
and savannahs north of Pleasant Mills. On the following day, wetlands along the Menantico
Creek and Manumuskin River in New Jersey’s Cumberland County were visited, as were
saltmarshes along Delaware Bay. The last day’s trips went to Pennsylvania, to visit
Serpentine Barrens in West Chester and at Tyler Arboretum. Trip leaders were Ted Gordon,
Karl Anderson, James Stasz, Joseph Arsenault, Bill Olson, Gerry Moore, Heinrich Zoller,
Anne Newbold, Jack Holt, and Janet Ebert. Evening programs were presented by Ted
Gordon, who spoke about the New Jersey Pine Barrens; by Bill Olson, who discussed New
Jersey’s Wetland Habitats; by Nancy Kuntzleman, who spoke about Serpentine Barrens; and
by George Pierson, who presented a History of Forestry in New Jersey. Chairman: Ted
Gordon.

July 13: Oswego River, Burlington Co., NJ. A torrential downpour that in six hours
dumped 9 inches of rain on the area made driving almost impossible. Nonetheless, nine
seasoned veterans—four from out-of-state—convened at Jenkins. During a lull in the storm,
it was agreed to attempt botanizing at Martha. We had just recorded Lobelia nuttallii,
Cladium mariscoides, and Oxypolis rigidior var. longifolia, when curtains of rain an
deepening puddles convinced us to call it a day. Leader: Ted Gordon.

July 20: Palmyra, Burlington Co., NJ. The focus of this trip was on adventive plants that
were noted on a walk into an overgrown dredge spoil area along the Delaware River, south
of Route 73. Some that appeared thoroughly naturalized on the site included Cyperus
amuricus, Polygonum orientale, Sorghum halapense, and perhaps Datura innoxia and
Petunia violacea. Present, but probably only as short-lived artifacts of leaf-dumping, were

1991 FIELD TRIPS 143

such plants as Sorghum vulgare, Cleome spinosa, Ipomaea purpurea (Morning Glory),
Portulaca grandiflora (Moss-rose), Tagates sp. (Marigold), Curcurbita pepo (Pumpkin), and
Lycopersicon esculentum (Tomato). A few other plants seenon this site included Populus
deltoides which is the dominant tree on most of the area, Urtica dioica which is also
abundant, and eee palustris, Humulus japonicus, Digitaria (Leptoloma) cognatum,
Croton glandulosus, Mirabilis nyctaginea, Rumex verticillatus, Froehlichia gracilis,
Helianthus annuus, and Heterotheca subaxillaris. Geranium pusillum, small but attractive,
was found growing in a vacant lot along Route 73. Leader: Karl Anderson.

July 27-28: Tuckahoe River/Belleplain State Forest, Atlantic, Cape May, Cumberland cos.,
NJ. This trip began in the Peaslee Wildlife Management Area at Hunters Mill in
Cumberland County where along a gravel road the group saw large populations of the rare
spurge Crotonopsis elliptica. Along a powerline, the following species were recorded:
Amphicarpum purshii, Andropogon virginicus, A. glomeratus, Aster dumosus, Carex
complanata, C. striata, Desmodium ciliare, D. viridiflorum, Eleocharis palustris, E. tenuis,
Eupatorium album, E. hyssopifolium, E. pilosum, E. rotundifolium, Helianthemum canadense,
H. propinquum, Hypericum canadense, H. hypericoides, Lechea pulchella, Lespedeza
procumbens, Liatris graminifolia, Lycopodium alopecuroides, Muhlenbergia uniflora,
Panicum dichotomum, Pinus rigida, P. verrucosum, P. virgatum, Polygala cruciata, P. lutea,
P. nuttallii, Quercus falcata, Rhexia mariana, R. virginica, Rhynchospora alba, R.
capitellata, R. macrostachya, Schizachyrium scoparium, Solidago odora, Xyris difformis, X.
torta, and X. smalliana.

An intermittent pond just north of the powerline cut that was devoid of water in most
places, had saplings of Acer rubrum, Liquidambar styraciflua, and Pinus rigida scattered in
the pond. The herbaceous layer was dominated by Cladium mariscoides. A few patches of
Panicum hemitomon, rare in New Jersey, were noted. The rare Lobelia canbyi was present
along with Carex striata, Drosera intermedia, Erianthus giganteus, Juncus canadensis,
Panicum verrucosum, Proserpinaca pectinata, Rhexia virginica, Rhynchospora alba, R.
chalarocephala, Vaccinium corymbosum, V. macrocarpon, and Xyris difformis.

Along the path leading to an intermittent pond south of the powerline cut, we saw the rare
Hedyotis uniflora. This pond was inundated with water to depths of 3 to 6 inches. Rare
species found here were Coreopsis rosea and Panicum hemitomon. Brasenia schreberi,
Nymphaea odorata, Scirpus subterminalis (with emergent stems and fruit), and Vaccinium
corymbosum were also found here.

The group then entered Cape May County en route to yet another intermittent pond in the
Peaslee Wildlife Management Area. Rare species in the pond were Ludwigia linearis,
Rhynchospora inundata (large patches present in the center of the pond), Sclerolepis
uniflora, and Sphagnum macrophyllum. Other species included Carex striata, Cladium
mariscoides, Drosera intermedia, Erianthus giganteus, Eriocaulon compressum, Juncus
canadensis, J. pelocarpus, Panicum ensifolium, P. spretum, Proserpinaca pectinata, Rhexia
virginica, Xyris difformis, and X. smalliana. Upon leaving the pond, the leader got lost in
a woods dominated by Quercus alba, Q. coccinea, Q. falcata, and Pinus rigida, with a shrub
layer of Gaylussacia baccata, G. frondosa, and Vaccinium vacillans.

The second day of the field trip began at Belleplain State Forest in Cape May County
where the group visited an intermittent pond off of Route 550. This pond was devoid of
water and dominated by Muhlenbergia torreyana. Other rare species included Eleocharis
microcarpa, Hypericum denticulatum, Lobelia canbyi, Panicum wrightianum, Rhynchospora

144 BARTONIA

(Psilocarya) sp., Sceria reticularis, and Sphagnum macrophyllum. The group then visited
another pond in the State Forest, south of Steelmantown Road and west of Crossing Road.
Carex striata was throughout much of the pond with a few stands of Panicum hemitomon.
Other species included Dulichium arundinaceum, Erianthus giganteus, Eleocharis
microcarpa, Panicum spretum, Rhexia virginica, and Sphagnum macrophyllum

e group’s last stop of the day was a tidal marsh along the Tuckahoe cai in Tuckahoe
to see Ammannia latifolia, Liliaeopsis chinensis, and Lythrum lineare. Leader: Gerry Moore.

August 3: Paulinskill Creek area and Stokes State Forest, Sussex Co., NJ. A walk along
the Paulinskill Creek on an abandoned railroad right-of-way produced a plant list which
included: Hepatica nobilis var. obtusa, Actaea pachypoda, Epigaea repens, Helianthus
divaricatus, Heliopsis helianthoides, Aster cordifolius, Sanguinaria canadensis, Lobelia
syphilitica, Melanthium hybridum, Polypodium virginianum, Asplenium platyneuron, A.
trichomanes, A. ruta-muraria, Woodsia obtusa, and Pellaea x atropurpurea. Within
creek itself, the following species were encountered: Potamogeton natans, P. crispus, and
Heteranthera dubia. A short distance away, we explored selected areas in Stokes State
Forest. On the road to Sunrise Mountain, we encountered Sorbus americana, Betula
papyrifera, and Eupatorium sessilifolium. Our final stop at the Steam Mill area produced
the following species: Glyceria canadensis, Lindernia dubia, Penthorum sedoides,
Lysimachia ciliata, Silene stellata, and Lobelia cardinalis. This trip was a joint trip with the

orrey Botanical Club. Leader: Bill Olson.

August 10: Mad Horse Creek Fish and Wildlife Mangement Area, Lower Alloway Creek
Township, Salem Co., NJ. Members explored the southern corner of Salem County’s tidal
wetlands and adjacent upland. Mad Horse Creek is a State owned parcel located on the west
side of Stow Creek. This area is dominated by a Delaware Bay brackish tidal marsh with
small, beaded chains of upland islands scattered throughout the marsh. We began our trip
by the Artificial Island Nuclear Power Plant causeway. From this vantage point, we had
access to a well maintained dirt road through the wetlands

Along this route there was a dense ground cover of Carex debilis, C. swanii, C.
complanata, and Chasmanthium laxum under the upland forest canopy of Liguidambar
styraciflua, Quercus phellos, Q. falcata, and Acer rubrum var. trilobum. Pinus taeda was
identified as a forest associate throughout the Fish and Wildlife Management Area. It was
interesting to note the presence of typically tidal species such as Spartina alterniflora
(dominant), Hibiscus moscheutos, and Spartina cynosuroides intermixed with the non-tidal
wetland species along the edge of the access road. Other species found along this road
included Aralia spinosa in very dense thickets, and a dense ground cover of Cyperus
amuricus in recently disturbed areas under the powerlines.

After lunch, the group drove to the Pine Island Boat Launch at the end of Stow Creek
Neck Road, to investigate Pine Island and Stretch Point. We walked a path from the parking
lot to Stretch Point through a series of small upland islands surrounded by the salt marsh of
Stow Creek. The islands were dominated by Quercus alba, Q. stellata, Q. velutina, Q.
falcata, Pinus taeda, Liquidambar, Nyssa sylvatica, Sassafras albidum, and Diospyros
virginiana with a dense understory of Vaccinium corymbosum, Myrica pensylvanica, and M.
cerifera. Other species included Eupatorium fistulosum and Eleocharis palustris.

After a brief rest, we drove to the headwaters of Stow Creek, in the vicinity of the village
of Jericho, on the Salem-Cumberland County border. The area around the pond and the

1991 FIELD TRIPS 145

stream below the dam produced Tipularia discolor, Lobelia cardinalis, Chionanthus
virginicus, and Carpinus caroliniana. The trip was ended after a brief drive over Coffin
Hill. Leader: Joe Arsenault.

August 17: Bennett Bogs Preserve, Lower Township, Cape May Co., NJ. After being
almost continually flooded since 1988, these intermittent ponds were finally free of standing
water for this trip. Although Habenaria nivea was not encountered, a number of species not
seen for a few years reappeared. These included: Rhynchospora filifolia, Psilocarya nitens,
and Scleria reticularis. Other species included Eryngium aquaticum, Hydrocotyle umbellata,
Sium suave, Aster dumosus, Bidens connata, Boiltonia asteroides, Sclerolepis uniflora,
Viburnum nudum, Hypericum boreale, Proserpinaca pectinata, P. palustris, Lycopus
amplectans, Lindernia anagallidea, Carex vesicaria, Eleocharis melanocarpa, Juncus debilis,
Tipularia discolor, Coelorachis rugosa, Panicum wrightianum, and Xyris difformis. Leader:
Bill Olson

August 24: Delaware River tidal wetlands, New Castle Co., DE. We began the day at
Augustine Beach where we saw plants characteristic of brackish marshes such as Cyperus
filicinus, Iva frutescens, Kosteletzkya virginica, Lilaeopsis chinensis, Pluchea odorata,
Samolus parviflorus, Spartina alterniflora, S. cynosuroides, and S. patens. There also were
plants that extend into the freshwater portion of the Delaware Estuary such as Amaranthus
cannabinus, Cyperus odoratus, Echinochola walteri, Hibiscus moscheutos, Phragmites
australis, Polygonum punctatum, and Schoenoplectus pungens. Farther upstream at Deemers
Beach there were fewer brackish species although Lilaeopsis chinensis and Spartina
alterniflora were abundant. Freshwater species not seen at Augustine Beach included Bidens
bidentoides, B. laevis, Cyperus bipartitus, Juncus acuminatus, Helenium autumnale, Sium
suave, Peltandra virginica, and Pontederia cordata. Still farther upstream at Bellevue, we
Saw many more freshwater species including Cardamine pensylvanica, Eleocharis
erythropoda, E. palustris, Heteranthera reniformis, Lindernia dubia, Ludwigia palustris,
Sagittaria graminea, S. latifolia, Schoenoplectus fluviatilis, S. tabernaemontani, and Zizania
aquatica. Species seen here that are more characteristic of brackish conditions included
Pluchea odorata, Samolus parviflorus, Solidago sempervirens, and Spartina alterniflora.
Leader: Alfred E. Schuyler.

September 14: New Lisbon, Whitesbog, and Lebanon State Forest, Burlington and Ocean
cos., NJ. At the Experimental Station in the Lebanon State Forest, we saw Aster patens, A.
linariifolius, Agalinis setacea, and an extensive patch of leaves of Solidago tarda (S.
ludoviciana) with just two flowering stems. The latter population, which frequently lacks
flowering stems, has persisted here for at least 25 years. Cessation of annual prescribed
burns of this site has accelerated shrub growth and diminished the spectacle of fall flowers.
Along the banks of Mt. Misery Brook in New Lisbon, we paused to see a vigorously-fruiting
colony of Lygodium palmatum, a station of long standing. Near the old "Dory" Bogs, east
of Magnolia, were Cyperus strigosus, C. dentatus, Spiranthes cernua, Agalinis purpurea, and
Solidago elliottii. East of Retreat, we carefully examined scattered plants of Spiranthes
ochroleuca, comparing them to a specimen (collected earlier) of the very similiar S. cernua.
Near our lunch stop at Pakim Pond, a few plants of Gentiana autumnalis, growing in
association with Calamovilfa brevipilis, had escaped mowing. At Woodmansie, in a recently
clearcut mature pine/oak stand, there were impressive clumps of Scleria nitida and S.

146 BARTONIA

pauciflora scattered over about two acres—the largest population of these species known to
the leader. It appears that these rare species have benefited from disturbances related to the
harvest and/or post-harvest preparation for planting of pine seedlings. At the edge of an
abandoned orchard in Pasadena, we saw a cluster of 16 plants of Aster concolor and a fine
patch of A. undulatus. A short visit to Goose Pond produced Rhynchospora inundata,
Utricularia purpurea, U. subulata (including forma cleistogama), Xyris difformis, and
possibly Sagittaria graminea. A brief search for a small population of Narthecium
americanum, seen here by the leader several years ago, was unsuccessful. On a road
shoulder at Whitesbog, we noted an impressive stand of about 60 plants of Prenanthes
autumnalis is association with Liatris graminifolia, Calamovilfa brevipilis, Scleria
triglomerata, and Polygala cruciata. Leader: Ted Gordon.

September 15: Trenton/Hamilton Marshes, Mercer Co., NJ (joint trip with Torrey
Botanical Club). We visited cleared areas, lowland forest, marsh edges near Sturgeon Pond,
and tidal channel and marsh at Watson’s Creek near Robeling Park picnic area. Plant
species of interest were Aristida purpurascens, Arthraxon hispidus, Bartonia virginica,
Bidens connata, B. discoidea, B. laevis, Carex trichocarpa, Eupatorium altissima, Laportea
canadensis, Lespedeza cuneata, Lycopus rubellus, Ludwigia peploides, .Mikania scandens,
Lycopodium obscurum, Phellodendron lavallei, Polygonum amphibium, Potamogeton
pusillus, Rumex verticillatus, Selaginella apoda, Spiranthes cernua, Vallisneria americana,
and Zizania aquatica. The leaders thank Karl Anderson, Bill Olson, Naomi Dicker, Robert
Meyer, Jessie Harris, and others for various identifications. Leaders: Mary and Charlie Leck.

September 21: Sandy Hook National Recreation Area, Monmouth Co., NJ. Joined by
members of the New Jersey Audubon Society, we visted a variety of habitats, including sand
dunes, woodlands, and salt marsh. Plants of sand dunes and dune fields included Cyperus
filiculmis, Opuntia humifusa, Helianthus petiolaris, Artemisia stelleriana, the introduced and
invasive sedge Carex kobomugi, and several grasses, among them Triplasis purpurea,
Aristida tuberculosa, Cenchrus tribuloides, and both Panicum amarum var. amarum and var.
amarulum. Dune woodlands and thickets dominated by Celtis occidentalis, Ilex opaca, and
Myrica pensylvanica also yielded Aster laevis, Rhus aromatica, Smilacina stellata, Lechea
maritima, Baccharis halimifolia, and abundantly-fruiting Prunus maritima. The edge of a
small salt marsh had Afriplex arenaria, Sueda maritima, Chamaesyce polygonifolia, Salsola
kali, Salicornia europaea, S. virginica, Iva frutescens, and the expected grasses Spartina
alterniflora, S. patens, and Distichlis spicata. The shore of a dune pond produced Lycopus
europaeus, Scirpus pungens, S. validus, and Pluchea odorata. About a hundred species of
plants we looked at during the day. Leaders: Bill Olson and Karl Anderson.

September 28: Presque Isle State Park, Erie Co., PA. Species in the unusual plant
communities here included Juncus alpinus, J. brachycephalus, J. dudleyi, J. articulatus,
Hemicarpha micrantha, Carex aquatilis, C. garberi, C. viridula, C. bebbii, Eleocharis
quadrangulata, Scirpus smithii, S. acutus, Cyperus diandrus, C. engelmannii, C. schweinitzii,
C. x nieuwlandii (C. flavescens x C. bipartitus), Panicum tuckermanii, Potentilla paradoxa,
Megalodonta beckii, Potamogeton richardsonii, and Equisetum laevigatum. Many of these
species are found on the Pennsylvania List of Species of Special Concern. Leader: James
K. Bissell.

1991 FIELD TRIPS 147

October 5: Cumberland and Cape May cos., NJ. Thirteen species of asters and nine
species of goldenrods were found on this trip. These include Aster tenuifolius, A. subulatus,
A. pilosus, A. dumosus, A. vimineus, A. patens, A. undulatus, A. concolor, A. nemoralis, A.
spectabilis, A. novi-belgii, A. gracilis, A. linariifolius, Solidago sempervirens, S. erecta, S.
odora, S. nemoralis, S. rugosa, S. uliginosa, S. canadensis var. scabra, Butheamia
graminifolia, and E. tenuifolia, Other species of interest were Setaria magna, Scleria nitida,
and Eupatorium capillifolium. Near the end of the day, Larry Klotz showed the group Lake
Nummy in Belleplain State Forest. There we found Myriophyllum humile. Later after the
trip, Larry located Psilocarya scirpoides in the same area. Leader: Bill Olson.

Bartonia No. 58: 148-154, 1994

1992 FIELD TRIPS

April 12: Lebanon State Forest, Burlington Co., NJ. This trip was primarily for lichens.
Locations and habitats visited included oak woods near the State Forest office and at Deep
Hollow Pond; a cedar swamp along Shinn’s Branch; upland pine forest east of Shinn’s
Branch; and lowland pine forest along Cooper Road. 39 species of lichens were identified.
Foliose species included Jmshaugia aleurites, I. placorodia, Pyxine sorediata, Parmelia
sulcata, P. squarrosa, Phaeophyscia rubropulchra, Hypogymnia physodes, and others. In
the fruticose category, four species of Cladina—Cladina subtenuis, C. submitis, C.
rangiferina, and C. arbuscula—were noted within arm’s reach of each other. As expected,
many species in the genus Cladonia were seen; some of them were Cladonia atlantica, C.
bacillaris, C. clavulifera, C. floridana, C. pleurota, C. rappii, C. strepsilis, and C. cristatella.
A few additional species, mostly crustose, were identified only to genus. Among the few
flowering plants in bloom were Pyxidanthera barbulata, Draba verna, and Cardamine
hirsuta—the last two as "weeds" in the lawn of the Lebanon State Forest office. Leader: Karl
Anderson.

April 18: Shenks Ferry and Susquehanna River Glens, Lancaster Co., PA. We started the
trip at the Holtwood Dam overlook where we saw Pinus pungens, Woodsia obtusata,
Asplenium pinnatifidum, and Viola fimbriatula. From there we travelled south along the
river to an unnamed glen adjacent to the Route 372 bridge (east side of river) where we
searched to no avail for Asplenium bradleyi. We did, however, see a large patch of
Asplenium montanum. We travelled north along the river to Tucquan Glen where we did
see a small patch of Asplenium bradleyi. The glen also had a good population of Trillium
erectum and Claytonia virginica. The last stop was at Shenk’s Ferry Glen. At the glen we
saw numerous plants of Mertensia virginica, Hybanthus concolor, Trillium flexipes/erectum,
Erythronium americanum, and Claytonia virginica. We searched to no avail for Corallo-
rhiza wisteriana. The plants were seen at the location two weeks later. Leader: Mark
Larocque.

April 25: Inner Coastal Plain at Vincentown and Pine Barrens, Burlington Co., NJ. We
explored a limestone ravine of the Vincentown Formation bordering the South Branch of
Rancocas Creek from the village of Vincentown west to the Hatcher Farm. Well known to
geologists and paleontologists, this ravine, rich in foraminifera and bryozoan fossils,
apparently has been little investigated by botanists. Here, within one mile of the Pine
Barrens, lies an outstanding example of a mature palustrine forest of mixed Quercus, Carya,
Liquidambar styraciflua, Acer rubrum, A. negundo, Platanus occidentalis (13 ft, 2.5 in cbh),
Liriodendron tulipifera (8 ft, 9.5 in cbh), Ulmus americana (9 ft, 1 in cbh), Fagus
grandifolia, Betula nigra (7 ft, 4.5 in cbh), Tillia americana, Carpinus caroliniana, Cornus
florida, and Celtis occidentalis. Herb and shrub diversity is especially high and includes
Aquilegia canadensis, Erythronium americanum, Corylus americana, Polystichum
acrostichoides, Cystopteris tenuis, Viburnum acerifolium, Rhododendron periclymenoides,
Heuchera americana, Chelone glabra, Goodyera pubescens, Luzula multiflora, Euonymus
americanus, Carex comosa, Cornus amomum, and Geum canadense. Several species that
are infrequent or local on the inner coastal plain are here too: Panax trifolium, Phegopteris

148

1992 FIELD TRIPS 149

hexagonoptera, Staphylea trifolia, Polypodium virginianum, and Tsuga canadensis. (Stone
cites only 3 southern New Jersey stations for Broad Beech Fern and 7 for Hemlock.) Rare
species seen here are Cercis canadensis and Geum vernum. In the afternoon we vistied
sections of the Rancocas Cranberry Company at Ongs Hat to see Aletris farinosa,
Amianthium muscaetoxicum, Botrychium matricariaefolium, Carex vestita, C. barrattii,
Krigia biflora, and Helonias bullata. In a Lebanon Forest cedar swamp near Four Mile we
saw another population of the latter species, associated with Orontium aquaticum. Leaders:
Joseph Arsenault and Ted Gordon.

April 26: Salem and Cumberland cos., NJ. In the woods and ravines of Salem Creek near
Woodstown we saw Alisma subcordatum, Allium tricoccum, Arisaema triphyllum, Cardamine
concatenata, C. hirsuta, Dicentra cucullaria, Geum canadense, Hydrastis canadensis,
Hydrophyllum virginianum, Iris pseudacorus, Lysimachia nummularia, Osmorhiza longistylis,
Podophyllum peltatum, Polemonium reptans, Polygonatum biflorum, Ranunculus abortivus,
Sambucus canadensis, Sicyos angulatus, Sparganium americanum, Veratrum viride, and
Viola pubescens. A side trip was made to Bridgeton to see Helonias bullata along
Roadstown Road. Leader: Michael O’Brien.

May 9: Bowman’s Hill Wildflower Preserve, Bucks Co., PA. Various trails through the
preserve were visited. Mertensia virginica was in full bloom in the low moist woodlands.
In the trail guide for the area this species is listed as blooming in early April. Perhaps due
to the cool April weather, these plants, and many others, were retarded in their development
so that many were not in full bloom on this trip. But altogether, about 25 species of herbs
were found flowering in scattered locations. A few plants of Silene virginica were observed
in flull bloom while one specimen of Chrysogonum virginianum was noted. Handsome
specimens of Trillium grandiflorum and T. erectum were seen as was one luxurious bloom
of Caltha palustris. Plants that were past bloom on this trip included Erythronium
americanum, Epiagaea repens, Dicentra cucullaria, D. canadensis, Sanguinaria canadensis,
and Jeffesonia diphylla. Only four grasses were observed in bloom; these were Antho-
xanthum odoratum, Poa annua, P. pratensis, and Dactylis glomerata. Leader: Robert Meyer.

May 16: Wissahickon Valley, Philadelphia Co., PA. Along Bells Mill Road and
downstream along Wissahickon Creek, we encountered a mixture of native and non-native
flora. Native herbs included Actaea pachypoda, Arisaema triphyllum, Circaea lutetiana,
Claytonia virginica, Collinsonia canadensis, Erigeron philadelphicus, Erythronium
americanum, Medeola virginiana, Podophyllum peltatum, Polemonium reptans, Polygonatum
pubescens, Ranunculus abortivus, R. caricetorum, R. recurvatus, Viola affinis, V. cucullata,
V. pubescens, and V. sororia. Non-native herbs included Aegopodium podograria, Alliaria
petiolata, Allium vineale, Artemisia vulgaris, Cardamine impatiens, Duchesnea indica,
Glecoma hederacea, Hesperis matronalis, Lysimachia nummularia, Ornithogalum
umbellatum, Ranunculus bulbosus, R. ficaria, R. repens, Stellaria media, and Verbascum
Phlomoides. A similar mixture of native and non-native woody plants and vines were also
observed. Some of these that were non-native included Berberis thunbergii, Celastrus
orbiculatus, Cercidophyllum japonicum, Deutzia scabra, Euonymus alatus, E. europaeus,
Hedera helix, Ilex crenata, Ligustrum vulgare, Lonicera japonica, L. maackii, Rhodotypos
scandens, Viburnum plicatum, and V. sieboldii. Numerous other species were recorded to
compare with those in the unpublished checklist prepared by John Fogg in the 1950s.

150 BARTONIA
Leaders: Sioux Baldwin and Alfred E. Schuyler.

June 6: Iron Hill, New Castle Co., DE. We spent the day exploring this monadnock
named for its long-abandoned iron mines. Starting at the parking lot the party observed a
few plants of Liparis lilifolia in bloom. Further along in the paths, we encountered, among
other species, Oxalis violacea, Scutellaria elliptica, Angelica venenosa, and a large colony
of Aristolochia serpentaria, along with the featured species of the trip, large quantities of
Ophiglossum pycnostichum with tongues almost fully exserted. We then followed a trail
through mature oak-beech-tulip woods toward Route 896 on the estern edge of the hill,
noting on the way populations of Galearis spectabilis, Pyrola elliptica, and P. rotundifolia.
We then bushwhacked through the woods in an unsuccessful search for a population of
Melanthium hybridum reported a few years back, but did encounter Adiantum pedatum along
with Obolaria virginica which was a new discovery for Iron Hill, before returning via a
different trail back to the parking lot for lunch. After lunch, we explored a road cut along
I-95, where we encountered Spergularia marina along the road, and on the banks such
species as Veronicastrum virginicum, Quercus stellata, Sisyrinchium mucronatum (just past
full bloom), young plants of Scleria pauciflora, and Lobelia spicata (still in bud). Leaders:
Janet Ebert and Jack Holt.

June 21-25: Chambersburg, PA. The 1992 Joint Field meeting of the Philadelphia
Botanical Club, the Torrey Botanical Club, and the Northeastern Section of the Botanical
Society of America was housed at Wilson College in Chambersburg. Three days were taken
up with field trips. The first day’s excursions went to sites along the Appalachian Trail in
Cumberland County, including swamps, dry woods, and the floodplain of the Conodoguinet
Creek. On the following day, the Altona Marl Marsh in West Virginia, and the C & O
Canal near Sharpsburg, Maryland, were visited. On the last day, trips went to Caledonia
State Park, a beautiful area, and to intermittent ponds on South Mountain. Trip leaders were
Greg Edinger, Larry Klotz, John Kunsman, Bill Olson, Rodney Bartgis, Kathy Bilton, and
Robert and Fanny Johnsson. Evening programs were presented by Wallace Drexler, who
spoke about Pennsylvania Geology; Robert Johnsson, who gave a fine program about the C
& O Canal; Rodney Bartgis, who discussed endangered plants of Maryland; and Eugene
Wingert, who spoke about the fauna of the South Mountain ponds. Charman: Larry Klotz;
Treasurer: Karl Anderson.

July 11: Oswego River Savannah and Turfcut, Burlington Co., NJ. Near the confluence
of Buck Run and the Oswego River, we visited a quaking savannah that, during a severe dry
spell, was swept by a devastating fire on June 22, 1986, extending to the river’s edge. The
narrow band of Chamaecyparis thyoides saplings that ringed the savannah was consumed by
the fire; however, the cedars in the wettest sloughs survived. During the first two years after
the fire, the proliferation of such grasses as Andropogon glomeratus, A. virginicus,
Calamovilfa brevipilis, Panicum virgatum, Agrostis perennans var. elata, and Danthonia
epilis by now had diminished remarkably. No other enduring impacts of the fire were
observed at this time. Vigorous populations of Sarracenia purpurea were thriving in iron-
stained pools, overtopped by the white heads of Eriocaulon decangulare, replacing the
decomposing heads of E. compressum. Rising above lush carpets and hummocks of
sphagnum were the yellow racemes of Narthecium americanum, the fuzzy inflorescences of
Lophiola aurea, the bright yellow spikes of Utricularia cornuta, and the white flowers of

1992 FIELD TRIPS 151

Tofieldia racemosa. Among associates were Juncus caesariensis, J. canadensis, Carex
livida, C. exilis, C. folliculata, Cladium mariscoides, Sabatia difformis, Pogonia ophiogloss-
oides, Calopogon tuberosus, and Utricularia fibrosa. In a nearby turfcut downstream,
dominated by Drosera filiformis and Utricularia cornuta, we again saw Narthecium scattered
with Lophiola plus fine pockets of Schizaea pusilla and Lycopodium carolinianum. Bladder-
bearing plants of Utricularia purpurea were submersed in an adjacent canal. Since our visit,
the NJ Natural Areas Council has been apprised of the species richness of this savannah and
turfcut, and is considering its incorporation within the boundaries of the existing Oswego
Natural Area. Leader: Ted Gordon.

July 18, 19: Manumuskin River, Muskee Creek, and Tuckahoe River; Atlantic, Cape May,
and Cumberland cos., NJ. The trip began in the headwaters of the Manumuskin River in
Buena Vista Township where we visited a chain of vernal ponds. The first pond was devoid
of water and was dominated by two rare species, Coreopsis rosea and Panicum wrightianum.
Other species present in this pond included: Scleria reticularis, Viola lanceolata, and Xyris
difformis. In the second pond, the outer edges were dominated by zones of Carex striata,
Panicum spretum, and Woodwardia virginica. In the wetter inner areas of the pond,
Eleocharis robbinsii, Eriocaulon aquaticum, E. compressum, Nymphaea odorata, Sagittaria
englemanniana, Utricularia fibrosa, Xyris difformis, and X. smalliana were recorded. The
third and final pond was much less diverse, being dominated by Cephalanthus occidentalis
and Dulichium arundinaceum. After leaving the ponds, A/etris farinosa, Lechea minor,
mucronata, Plantago lanceolata, P. major, P. rugelii, and Scleria triglomerata were recorded
in an open, dry area adjacent to an abandoned building. Moving on to an open wetland
adjacent to a farm field near the railroad crossing along Landis Avenue, south of Richland,
we keyed out the following sedges: Carex barrattii, C. striata, Cyperus dentatus, Eleocharis
microcarpa, E. obtusa, E. olivacea, E. tenuis, E. tricostata, E. tuberculosa, Rhynchospora
capitellata, and R. fusca. Our keys were of little help for the large bed of Scirpus that
dominated one area of the wetland. No flowering or fruiting material was present, and we
could only wonder whether this was the rare S. /ongii, which has historically been reported
from here, or the common S. cyperinus. Also noted was Proserpinaca intermedia. The last
Stop of the day was a cedar swamp in the headwaters of the Muskee Creek in Maurice River
Township. The following herbs were noted on the corduroy road that led into the swamp:
Drosera intermedia, D. rotundifolia, Eriophorum virginicum, Lycopodium appressum,
Panicum mattamuskeetense, Pogonia ophioglossoides, and Polygala lutea. A very large
white cedar was noted just off the corduroy road. On the next day, we recorded a
circumference of approximately 91% ft at breast height for this tree, which makes it the lagest
living Atlantic white cedar for NJ. Also in this cedar swamp, we visited an open pond,
which had over 3 ft of standing water and was dominated by Scirpus subterminalis and
Utricularia fibrosa. Farther east in the Tuckahoe River watershed, we visited a vernal pond
in the Peaslee Wildlife Management area. In the inner wetter areas of the pond there were
large stands of the rare Rhynchospora inundata and Sclerolepis uniflora. A large stand of
Erianthus giganteus dominated the outer edges of the pond. The Rhexia that dominated
some areas of the pond appeared to be R. virginica with appressed leaves. Later these plants
were identified as R. aristosa by Robert Kral. They differ, however, from plants ee R.
aristosa at the type locality in Atlantic County in a number of characteristics (¢.g., leaf
width, leaf margin, sepal lobe position, seed surface). A careful systematic study of foi
Jersey’s Rhexia appears to be warranted. Our last stop was another vernal pond, north of

baz BARTONIA

the Head-of-River Cemetery. This pond was dominated by Carex striata, Eleocharis
robbinsii, Eriocaulon compressum, and Panicum spretum. A very small stand of
Rhynchospora inundata was also here. The woodland path that led to the pond from County
Route 666 was dominated in places by the state rare Crotonopsis elliptica; the rare violet,
Viola brittoniana, was also noted along this path. Leader: Gerry Moore

July 25: Whitesbog and Vicinity, Ocean and Burlington cos., NJ. The morning was spent
exploring damp, peaty roadsides in the vicinity of Whitesbog, a historic cranberry-blueberry
village that is now part of Lebanon State Forest. A large and diverse assemblage of plants
included Habenaria blephariglottis just coming into bloom, Calopogon tuberosus at the end
of its blooming season, Spiranthes gracilis, Polygala lutea, P. brevifolia, Galactia regularis,
Schwalbea americana, Drosera filiformis, D. intermedia, D. rotundifolia, Scleria
triglomerata, Juncus biflorus, some interesting oaks includin oe stellata, Q.
marilandica, and Q. prinoides, and numerous species of enh shrubs. A post-lunch
visit to Webb’s Mills produced Schizaea pusilla, Lycopodium carolinianum, Lophiola aurea,
Sarracenia purpurea, Triadenum virginicum, Sabatia difformis, Narthecium americanum, and
three species of Utricularia including U. fibrosa, U. cornuta, and U. juncea. The final stop
was a sandy swale along an old railroad bed near Roosevelt City. Here, we found that a
stand of Eupatorium resinosum, a globally-rare species, had been recently mowed, probably
by a utility line maintenance crew. Only a few plants remained standing. Leader: Karl
Anderson.

August 1: Goat Hill Serpentine Barrens, Chester Co., PA. The barrens had been burned
during a wildfire during the last winter so the plants were very vigorous. Ascelpias
verticillata was in full bloom with greater numbers of plants than seen there previously. We
also saw Asclepias viridiflora, Sporobolis heterolepis, Liparis loeselii (first known Chester
Co. record), Sabatia angularis, Helianthus giganteus, and Vernonia noveboracensis. After
lunch at Harold Sweetman’s house, we hiked into the southern end of the barrens to see the
Federally Endangered chickweed, Cerastium arvense var. villosissimum. We also saw
Arisaema dracontium and Campanula rotundifolia. Four hardy people plus myself ventured
to Nottingham Park to search for Spiranthes vernalis to end the day. Leader: Mark
Larocque.

August 15: Lehigh Pond, Wayne Co., PA. This trip started with a visit to an excellent
example of a glacial bog natural community at Lehigh Pond. Here we saw concentric
vegetation zones on a floating peat mat surrounding an open body of water. The shrub mat
was dominated by Chamaedaphne calyculata, Kalmia angustifolia, K. polifolia, Spiraea
latifolia, and S. tomentosa. Open peat areas were dominated by numerous sedges such as

arex canescens, C. lasiocarpa, C. limosa, C. paupercula, C. stricta, C. trisperma,
Dulichium arundinaceum, and Eriophorum virginicum. Insectivorous species were also
abundant including Drosera intermedia, D. rotundifolia, and Sarracenia purpurea. Other
plants of interest were Platanthera blephariglottis (in bloom), Andromeda polifolia, Ledum
groenlandicum, Menyanthes trifoliata, and Solidago uliginosa. After lunch at Tobyhanna
State Park, the group headed to State Game Lands No. 127 to see a large stand of Myrica
gale, growing with Nemopanthus mucronata and Rhododendron canadense. Leader: Greg
Edinger.

1992 FIELD TRIPS 153

August 29: Hutcheson Memorial Forest, Somerset Co., NJ. We explored the successional
fields (aged 3—30 years) and a part of the forest of this ecological research property. The
"old forest" has lost most of the large trees in recent decades due to drought, gypsy moth
damage, hurricanes, and natural mortality, but some older oaks were noted. One of the more
dramatic research projects showed the vegetation effects of deer and rodent exclosure fences.
Some of the plants observed were Pycnanthemum muticum, Quercus robur, a variety of
Solidago (S. nemoralis, S. juncea, S. rugosa, S. gigantea, S. altissima, S. canadensis),
Euthamia graminifolia, Helianthus giganteus, Cornus racemosa, and C. amommum. In mid-
afternoon we made an excursion to another property of Rutgers University, the Display
Garden and Helyar Woods at Cook College, New Brunswick. The public garden is noted
for trails of new varieties and its perennial collections (e.g. hollies, dogwoods, and other
ornamental trees). Helyar Woods was seen to be a healthy climax forest dominated by
beech, oaks, and hickories. Leaders: Mary and Charlie Leck.

September 12: Cecil, Gloucester Co., NJ. Two pine barren edge locations were explored
on the border of Monroe and Franklin townships for roadside fall composites and
successional conditions found in two sand and gravel pits. Along a stretch of Blue Bell
Road, between Malaga Road and Coles Mill Road, we found Solidago odora, S. rugosa, S.
puberula, S. erecta, S. nemoralis, S. ludoviciana, S. juncea, Aster lineariifolius, A. undulatus,
A. umbellatus, A. gracilis, A. spectabilis, A. novi-belgii, Liatris graminifolia, Lespedeza
repens, Desmodium rotundifolium, D. marilandicum, D. paniculatum, Danthonia spicata, and
Epigaea repens. This rich assemblage is found on the infrequently mowed roadside adjacent
to an oak-pine forest. We investigated an abandoned gravel pit on the north side of Blue
Bell Road that was in an early successional stage ten years after being abandoned.
Observations and comments were made on the continued bank erosion and movement of all
surface soils. The few plants that invaded this unstable, dry, open habitat included Aristida
longspica, Diodea teres, Digitaria filiformis, Andropogon virginicus, Panicum virgatum, and
Euthamia tenuifolia. After lunch, we inspected and compared the conditions of slopes of a
restored section of Arrowwood Nursery. This site was planted with oaks (black, white, post,
bear, chestnut, red), pitch pines, old field composites and grasses seven years ago.
Restoration definitively improves the success of plants becoming established on soils without
any nutrient retention. The slope soil surfaces were completely covered. The soil was
sealed with a lichen, algae, moss, and fungi surface crust which supported grasses and old
field composites common in dry fields in Southern New Jersey. Limiting foot traffic,
incorporating a carbon source into the soil, and eliminating stream flow erosion are keys to
Success. Native species found on the uplands included Solidago speciosa, S. ludoviciana,
Polygala polygama, and Aster spectabilis. Wetland habitats supported numerous sedges and
rushes, including Cyperus dentatus, C. iria, Xyris difformis, Spiranthes cernua, and Juncus
pelocarpus. Later at a gravel excavation in Winslow Fish and Wildlife Management Area
we found a colony of Eleocharis quadrangulata discovered earlier by Mr. Ben Gindville, the
only known recent record for this species in Camden County. Additional species discovered
included Scleria triglomerata, Carex barrattii, Rhynchospora gracilenta, R. fusca, R.
macrostachya, and Leiophyllum buxifolium. Leader: Joe Arsenault.

October 3: Craven’s Estate, Natural Lands Trust, near Sumneytown, Montgomery Co., PA.
The "Potato Patch,” a lichen covered field of diabase rock—the type also known as Ringing
Rocks—is home to Corydalis sempervirens, Arabis laevigata, and Polypodium virginanum.

154 BARTONIA

Across the dirt road is a wild flower field filled with Pycnanthemum tenuifolium, P.
virginianum, Heliopsis helianthoides, Cirsium muticum, Solidago speciosa, S. rigida,
Gentianopsis crinita, Gentiana andrewsii, Zanthoxylum americanum, Dirca palustris, etc.
Thirteen ferns were found on the Trust’s land. We started with a pre-recorded list of 314
plants to which we added 21 more. Jack Holt’s orchidean eye spotted Liparis lilifolia,
Goodyera repens, and Corallorhiza odontorhiza. In the afternoon we walked up to "Indian
Alleys," the cave area reported to be the home of some Lenni-Lenapes of yesteryear.
Leaders: Ann Newbold and Henrich Zoller.

r Bartonia No. 58: 155-159, 1994
MEMBERSHIP LIST 1993
' ALDHAM, ALBERT—27 Madison Avenue, New York NY 10010, 212-340-0507

ALLEN, FRANCIS—4234 Tudor St., Philadelphia PA 19136, 215-624-2294
AMOS, SANDRA—41 Laurel ie Clementon NJ 08021, alla Rie

ANDERSON, KARL H.—Rancocas Nature Center, 794 Rancocas Road, Mt. Holly NJ 08060, 609-261-2495
ANDET, ANNE-MARIE — pee St, Apt. 19A, Philadelphia PA 19106, 215-351-2833

ARCHIBALD, MARY E.—21 Wayne Gardens, Collingswood NJ 08108, 609-854-7058
| eas, JOSEPH & CATHY—1042 Clark Avenue, Franklinville NJ 08322, 609-697-6044
ATTARDI, VINCENT J.—P.O. Box 472, 34 S. Main St., Suite C, Allentown NJ 08501, 609-259-2423
BACON, JULIA—160 Kendal at Longwood, Kennett a PA 19348, 610-388-8656
BAKER, BRADLEY—P.O. Box 244, Landenberg PA 5
BALDWIN, SIOUX—634 W. Ellet St., pave a 19119, 215-242-5610
BALLAS, JOHN—P.O. Box 4265, Elwyn PA 19063, 610-566-7392
| BARR, CAMILLE—4418 Wainut St., Apt. 7 Philadelphia PA 19104
BART, ELLEN S.—527A S. 16th St., Philadelphia PA 19146, 215-735-0801
BASKIN, JERRY—School of Biological Science, University of Kentucky, Lexington KY 40506, 606-257-8770
BAUCHSPIES, JAMES T.—1205 Washington St., Easton PA 18042, 610-253-8498
BELLING, ALICE—74 Palisade Avenue, Jersey City NJ 0
BETANCOURT, RAUL JR.—608 W. Hartwell Lane, Philadelphia PA 19118
BIEN, WALTER F.—144 Summit Avenue, Langhorne PA 19047, 215-752-3762
BIER, CHARLES W.—372 Kepple Road, Sarver PA 16055, 412-288-2777
BOONIN, ROSE K.—3516 Lewis Road, Newtown Square PA 19073, 610-356-1073
Salons HELEIGH—2036 Spring Garden St., Philadelphia PA 19130, 215-422-3917
HILL WILDFLOWER PRESERVE—P.O. Box 103, Washington Crossing PA 18917, 215-862-2924
23 734

7)

BOYLE, E. MARLE—Unitarian-Universalist Home, 224 W. Tulpenhocken St., foe PA 19144, 215-844-0376
BRESSLER, CARL—264 W. Wolfert Station Road, Mickleton NJ 08056, 609-467-3642
BRINTON, JOAN Z.—896 Roundelay, West Chester PA 19382, 610-793-1582
BROADDUS, LYNN—I68 Clemson Road, Bryn Mawr PA 19010, 610-525-5406
BROTHERSON, ROBERT—Box 179, Revere PA 18953, 610-847-5074
’ BRUEDERLE, LEO P. —Biology Department, sasiagend of Michigan, Flint MI 48502-2186, 313- 762-3041
BUCK, WILLIAM R.—New York Botanical Garden, Bronx NY 10458, 212-220-8624
, BURCHICK, MARK—5210 Cochran Road, Beltsville MD 20705, 301-937-7045
BURGESON, DENNIS K.—236 Adams Avenue, Barrington NJ 08077
BURLINGHAM, JAMES P.—500 E. Marylyn Avenue, State College PA 16801, 814-231-8350
CALLENDAR, GIZELLA P.—225 N. Charlotte St., Lancaster PA 17603, 717-393-8438
CHADY, SUSAN—Highspire Farm, R.D. 2, Box 100, Glenmoore PA 19343
} CHILCOTT, THOMAS—118 N. Wycombe Avenue, Lansdowne PA 19050, 610-622-7607
| CHRISTIAN, PATRICIA H.—Box 24, Starlight PA 18461, 717-798-2433
CLANCY, KEITH—300 N, State St. #4, Dover DE 19901, 302-739-5285
COLLIER, MITCHELL N.—316 Gerard Avenue, Apt. A, Elkins Park PA 19117, 215-635-4414
Cooper, Ton! ANNE—1238 S. Sheridan St., Philadelphia PA 19147
COURTNEY, JOHN E. & MARILYN—439 Gladstone Avenue, Haddonfield NJ 08033, 609-429-4987
CRICHTON, OLIVER W.—94 Colorado Avenue, Wilmington DE 19083, 302-764-55
CROCKETT, LAWRENCE J.—240 E. Palisade sonnel ei Englewood NJ 07607

CROWLEY, MARY—34 Schappet Terrace, Lansdowne PA 19050
CUSICK, ALLISON W.—Division of Natural Areas tke Fountain Square, Columbus OH 43224, 614-265-6471
CUTTER, DAVID M.—210 Ardmore Avenue, Ardmore PA 19003, 610-642-7857

DABYDEEN, SIMON—Frostberg State University, Frostberg MD 21532, 301-689-4213
DAHLBURG, MADELINE B.—508 Kent Place, Berwyn PA 19312, 610-644-5738

DaRIGO, DOLLY V.—553 Mapleview Drive, ee City MO 63130, 314-863-7057
DARKE, RICK—Longwood Gardens, P.O. Box 501, Kennett Square PA 19340, 610-388-6741
DAvis, LINK—14 Cemetery Lane, Schwenksville os 19473, 610-287-6635

155

156 BARTONIA

DECASTRO, LINDA—1100 Concord Drive, Bridgewater NJ ma 908-658-4497
DENNY, GUY—1889 Fountain Square, Bldg. F, Columbus OH 43244
DICKER, NAOMI D.—309 W. 93rd St., 6A, New York NY 10025, 212-222-5179
DILL, NORMAN—Biology aad Natural Resources, Delaware State College, Dover DE 19901
Dix, EDWARD—26 S. 8th St., #1, Lemoyne PA 17043, 717-783-0392
DOLAN, THOMAS IV—721 Glengarry Road, Philadelphia PA 19118, 215-242-6136
DRAUDE, TIMOTHY—415 Poplar St., Lancaster PA 17603
EBERT, JANET—394 Smith Bridge Road, Chadds Ford PA 19317, 610-459-0585
EDINGER, GREG—6006 Carversville Road, Doylestown PA 18901, 215-862-7924
ESHERICK, HELEN K.—2346 Dublin Road, Orefield PA 18069, 610-395-5292
EVANS, JANET—Library, Pennsylvania Horticultural Society, 325 Walnut - Philadelphia PA 19106, 215-625-8261
EWAN, JOSEPH—Missouri Botanical Garden, P.O. Box 299, St. Louis MO 63166, 314-577-9505
FADRA, SHEELA—380 Parry Road, uenitenier PA 18974, 215-674-4
FARLEY, ELIZABETH B.—319 Bala Avenue, Bala Cynwyd PA 19004, 610-667-0625
FARRELL, MAUREEN—437 W. ee House Lane, Philadelphia PA 19144, 215-848-7529
FEENEY, JOHN & DARLENE—P.O. 2359, Vincentown NJ 08088
FELTON, ANNA—37 Crawford ae gre PA 19403, 610-666-5922
FIELD, STEPHEN R. & THERESA—S Evelyn Avenue, Vineland NJ 08360, a -5868
FINE, NORMAN—16 Overhill Road, East Brunswick NJ 08816, 908-257-2
FINGERUT, JOYCE—2106 Sine og Avenue, Ft. Washington PA shh ae 542-0153
FIORINO, RICHARD—32 Manor Drive, Red Bank NJ 07701, 908-842-5189
comes ‘Wecnps—8110 Eastern Avenue, Wyndmoor PA 19118, 215-233-3035
FOG. , KASIA—317 Roxborough Avenue, Philadelphia PA od 215-482-3835
FRANK, pm AN—2508 Pine = Philadelphia PA 19103, 215-732-928
FREYBURGER, HELEN R.—5258 34th Avenue North, St. Petersburg = 33710, 813-526-1579
FRICK, JULIA sep 213, 1400 Waverly Road, Gladwyne PA 19035, 215-645-8863
GALLAGHER, MARK—1638 Ramblewood Lane, Jamison PA 18929
GARAFFA, TAMARA—Birdsall Engineering, 1700 F St., Belmar NJ 07719, 908-681-1165
inane MARY E.—1789 Scattergood St., Philadelphia PA 19124, 215-535-1323

S, CLARENCE W.—Shell Point Village, Cameo 1133, Fort Myers FL 33908, 813-466-6407
na ” AMELIA—135 Washington Avenue, Pitman NJ 08071, 609-589-6435
GOFF, ELINOR—791 College Avenue, Apt. 1, Haverford PA 19041, 215-649-2933
GOHSLER, ANNE MARIE—3000 Moser St., Aston PA 19014, 215-494-3760
Goop, NORMA—745 Redman Avenue, Haddonfield NJ 08033
GORDON, THEODORE & PATRICIA—31 Burr’s Bill Road, Southampton NJ 08088, 609-859-3566
GREENLAND, CHRISTINE M.—790 E. Street Road, Warminster PA 18974, 215-322-4105

RIND 4
GUARNACCIA, JOHN—215 Overlook Road, Philadelphia PA src 215-482-2713
ane SHALL, MARK—P.O. Box 97, Oxford PA 19363, 215-932-

R, JOHN & JANET—Box 185, 243 Jessup Mill Road, Gk NJ ig 609-423-3889

aah THOMAS B.—19 Kings Road, Netcong NJ 07857, 201-347
bane pass GORY M.—Box 263, R.D. 3, Phillipsburg NJ 08865, hae 13
HARDESTY, GAIL B.—488 Big Oak Road, via PA 19067, 215-295-4734
HARRIS, pide M.—4401 W St. NW, Washington, DC 20007, 202-338-9083
HART, RoBIN—Sarasota County Natural Resources a P.O. Box 8, Sarasota FL 34230, 813-378-6113
HEILMAN, LYNN—1004 Cherry Circle, Lansdale PA 19446
HEISTER, RALPH D.—693 gears iat Wayne a 19087, 215-254-9942
HENRY, JOSEPHINE DEN.—801 Stony Lane, Box 7, Gladwyne PA 19035
HILL, Roy L.—180 W. Drexel pan Lansdowne PA 19050, 215-626-7743
HIRST, FRANKLIN S.—500 Little Mill Road, Stockton MD 21864, 301-436-9303
HOLINKA, JOHN A.—126 West 11th St., Bayonne NJ 07002, 201-436-9303

HOLLINS, MICHA —Envirens, 1927 York Road, Timonium MD
HOLT, sang J—30 Road, Norristown PA 19401, 215-584-5578
HORNBECK, JOHN A.—1650 Viade Luna, Apt. 2, Pensacola Beach FL 32356, 904-932-8591

HOUSE, echingyeres Healing Center, Inc., ie A pss St., Philadelphia PA 19139
HUNT, LYNN F.—P.O. Box 553, New Gretna NJ

A

MEMBERSHIP LIST 1993 157

ILNICKI, RICHARD D.—403 Georges Road, Dayton NJ 08810-1400, 201-932-9711
INGRAM, cong ges Cardinal Lane, West Chester PA 19382, 610-399-0192
ISAAC, BONNIE—R.D. 1, Pulaski PA 16143, 402-964-8770
JESS, ROBIN A.—55 eth Avenue, Edison NJ 08817, 908-572-5928
JOHNSON, ELIZABETH—The Nature Conservancy, P.O. Box 181, 17 Fairmont Road, Pottersville NJ 07979,
908-439-3007
JOHNSSON, ROBERT G. & FANNY M.—7422 Ridge Road, Frederick MD 21701, 301-371-5215
JOHNSTON, KAREN tees 1, Box 488F, Liverpool PA 17045, 610-444-3691
KAISER, GEOFFREY—P. O. Box 222, Sumneytown PA 18084, 215-234-8424
KAMPF, JENNIFER—226 ae Sania Square #3117, Philadelphia PA 19103, 215-545-7128
KARLIN, ERIC—Institute for Environmental Studies, Ramapo College, Mahwah NJ 07430
KELLER, ELIZABETH—114 Wyomissing Boulevard, bd peor PA 19610, 610-374-3458
KENDIG, JAMES W.—35 Arlington Drive, Pittsford NY 14534
KIRCHHOFFER, DONALD—100 Burnt House Road, bial NJ 08088, 609-268-2011
KLEIN, WILLIAM M. JrR.—11121 SW 62nd Avenue, Miami FL 33157, 305-667-1651
KLOTZ, LARRY H—Biology Department, Shippensburg University, Shippensburg PA 17257, 717-532-1402
KNEZICK, DON—323 Tstand Road, Columbus NJ 08022, 609-291-9486
KOERBER, WALTER A. JR.—1380 Valley Green Road, Etters PA 17319, 717-938-9618
KOLAGA, " VALERIE—186 Dilworthtown Road, West Chester PA 193
KRAFFT, CAIRN—57 Willow Road, Churchville PA 18966, 215-322-855
KRAIMAN, prio: Hills, 7 Chip Lane, Reading PA 19607, 610- aes
KUNTZLEMAN, NANCY—R.D. 1, Box 1191, Hilltop Road, Leesport PA 19533
— bas ao 8, Box 313, Coatesville PA 19320, 610-383-7542
LADEN, MILTON—334 Wellesley Road, Philadelphia PA 19119, 215-247-7616
LAIDIG, KIM—712 Wood Lane, Haddonfield NJ 08033
ea ERIC—586-14 Sound Shore Road, Riverhead NY 11901
LANSING, DoROTHY—20 Old Paoli Pike, Box 537, Paoli PA 19301, 610-644-1890
LAROCQUE, MARK—R.D. 2, Beacon Light Road, Coatesville PA 19320, 215-857-9623
LATHAM, ROGER—Box 57, Wallingford PA 19086, 610-565-8979
LATHROP, SUSAN H.—107 Blackthorn Road, Wallingford PA 19086
LAUER, DAVID M.—607 E. Thompson St., Philadelphia PA 19125, 215-423-948
LECK, oie Department, Rider College, 2083 Lawrenceville <a Lawrenceville NJ 08648,
609-896-5092
LEVIN, anaaah Mill Road, Havertown PA 19083
LIGHTY, RICHARD W.—Mt. Cuba Center, P.O. Box 3570, Greenville DE 19807, 302-239-4244
LITTLE, ROBIN—705 Thunderhill Road, Lincoln ee PA 19352
LIVE wea pou DAVID—46A Brainerd St., Mt. srgee
LLo WRENCE—R.D. 3084A, Mohnton PA |
“pf Saa g MICHAEL J. —2028 Fitzwater St., pilleas A PA 19146, 215-732-0849
LOEFFLER, CAROLE C.—Department of Biology, Dickinson College, Carlisle PA 17013
LOGAN, MICHAEL—1027 Morris St., Philadelphia PA 19148, 215-334-6151
NKER, MILTON L.—8704 Patton Road, Wyndmoor PA 19118, 215-233-4818
Lowry, JANET L.—23 N. Lincoln Avenue, Newtown PA 18940, 215-968-8683
MACLEAN, DONALD R.—9023 Queen Maria, Columbia MD 21045
MACIARELLO, MICHAEL J.—Delaware State College, Dover DE 19901, 302- cee
he picasa GEORGE—17 Hunters Lane, beara en NJ 08089, 609-768-0

ES, CAROL C.—Biology Department, Kutztown University, Kutztown PA tt ‘ena
‘aiseiks SIDNEY—600 W. Harvey St., Pind PA 19144, 215-848-2431
MARGOLIS, WENDY—204 Leedom m Way, Newtown PA 18940, 215-579-1621

MAURICE, KEITH R—186 Main St., Linfield Pa 19468, 610-495-7951

McAvoy, WILLIAM—35 Windflower Drive, Newark DE 19711, 302-734-3219

MCCABE, MARIANA F.—803 N. Franklin St. Pottstown PA 19464, 610-326-3082
McComBE, ROBERT W.—20 Sylvan Lake Avenue, Haddonfield NJ 08033, 609-429-4628
MCCONVILLE, LYNN—1835 Wynnewood Road, Philadelphia PA 19151, 215-877-4311
MCCULLEY, JAMES—158 Coo oopers Drive, Newark DE 19702, 302-456-9066

MCFADDEN, ANDREA—820 Church Road, Wayne PA 19087

MCGRATH, JAMES K.—304 Derwyn Road, Lansdowne PA 19050, 610-284-2594

158 BARTONIA

MCLAUGHLIN, WILLIAM—P.O. Box 645, Medford NJ 08055, 609-268-8010

MCLEAN, WILLIAM & ELIZABETH—139 Cherry Lane, Wynnewood PA 19016, 610-642-4196

MEADE, JOHN A.—16 Tall Oaks, co Brunswick NJ 08816

MEAGHER, WALTER L.—R.D. 3, Box 407A, Robbinsville NJ 08691

MEHRHOFF, LESLIE J ee Geological and Natural History Survey, University of Connecticut, Storrs, CT

MELLON, RICHARD—P.O. Box 63, Morrisville PA 19067

MEYER, ROBERT C. JR.—5 Railroad Lane, Whitehouse Station NJ 08889

MICKLE, ANN M.—Department of Biology, LaSalle University, pea PA 19141, 215-951-1254

MITCHELL, JOHN D.—48 Upper Creek Road, Stockton NJ 08559, 908-996-3744

MONTGOMERY, JAMES D.—Ecology III, R.D. 1, Berwick PA 18603, 717-542-2191

MOoBERRY, F. M.—106 Spottswood Lane, Kennett Square PA 19348, 610-444-5495

Moore, GERRY—Department of Biology, Vanderbilt seeps Nashville, TN 37235, 615-385-4577

Moore, JULIA E.—Molyneaux Road, Camden, ME 04843, 2 1-83

Morse, LARRY E.—The Nature Conservancy, 1815 N. Lynn St., Arlington VA 22209, 703-841-5361

Moss, MIRIAM—8120 Brookside Road, Elkins Park PA 19117, 215-635-0176

MULVEY, ra get Warner Lane, Frazer PA 19355

NAczI, ROBERT F. C.—Department of Biological Sciences, Northern Kentucky University, Highland Heights KY
41099, esos

NEWBOLD, ANN—Heinrich Zoller, 50 Renninger Road, Bechtelsville PA 19505, 610-754-7573

NEWSTEAD, EDWIN & CHARLOTTE—270 Roseland Avenue, ce Fells NJ 07021, ote 7651

NICHOLS, HORATIO R.—412 Federal City Road, Pennington NJ 08543, 609-737-744

NOBLE, WILFRED R.—P.O. Box 4331, Philadelphia PA 19118, ee

NYSTEDT, JOHN—227 Chestnut St., Audubon NJ 08106

pboseandid ieee Richards Avenue, Pine Hill NJ 08021, 609-783-1103

O’CONNOR, DANIEL J—59 Grove St., Bloomfield NJ 07003
O’DELL, enn = . Schuylkill Road, Pottstown PA 19464, 610-323-3971
OLSON, WILLIAM F.—11 Venderveer Road, Freehold NJ 07728, 908-409-3647

PARKS, JAMES aes a tort Road, Washington Boro PA 17582, 610-872-5206
PENMAN, W. R.—20 Old Paoli Pike, Box 537, Paoli PA 19301, 610-644-1890
PINGEL, PATRICIA A—R.D. 1, Box 324, Reading PA 19607, 610-777-8883
PIOMBINO, CHRISTINE A.—452 Greenhill Lane, Berwyn PA 19312, 610-644-9345
PITTAM, SHERRY—Division of Botany, NHB 166, Smithsonian Institution, Washington, DC 20560, 202-357-2545
PLYLER, DOROTHY D.—18 Bridle Path, Chadds Ford PA 19317, 610-459-3969
PORTER, JOHN R.—631 Folsom pale Folsom PA 19033, 610-461-6612
PRESSLER, BARBARA—R.D. 2, Box 2129, Orwigsburg PA 17961
IS, RICHARD P.—69 Ogden Avenue, a NJ 07866
RAYSER, JEANNE—Cedar Run Lake, Medford NJ 08055
RHOADS, ANN F.—3 Blythewood Road, Doylestown PA 18901, 215-348-8139
RHODES, CHARLES A. JR—107 Stony Creek Avenue, Lansdale PA 19446, 215-368-9591
RISKA, MICHAEL E.—Delaware yen Society, Box 700, Hockession DE 19707
RITTENHOUSE, WILBUR H.—Rt. 1, Box 13, Queen Anne MD 21657, 301-364-5731
ROBACK, HELEN M.—188 Redwood pain King of Prussia PA 19406, 610-265-1158
ROBERTS, MARVIN L.—Department of Biology, Salem State College, Salem, MA 01970, 508-741-6236
ROBERTS, WILLIAM H.—1992 Rittenhouse Square, Philadelphia PA 19103, 215-569-5632
ROWAN, JANE O.—257 Barren Road, Media PA 19063, 610-891-065
RUDNICK, i J—11 Cavendish Road, Ambler PA 19002

RupDyJ, Eric S.—915 Olive Branch Court, Egdgewood MD 21040, Scgled $782
RUSSELL, ay W. B.—Box 430, Mt. Tabor NJ 07878, 201-625-338
AN, NANCY E.—419 S. Carlisle St., Philadelphia PA 19146, 215- 789

SCHAEFFER, ROBERT L. JR—32 N. 8th St., Allentown PA 18101, 610-433-4780
SCHMIDT, DAVID E.—6 Bayview Drive, Strathmore NJ 08248, 609-263-8049

ScHoss, Jay—545 N. Edgemere Drive, West Allenhurst NJ 07711

SCHRENK, W. JUERGEN—11 Westbrooke Boulevard, Cranbury NJ 08512, 609-936-9652
SCHROT, EDITH FEUERSTEIN—402 Laurens St., Olean NY 14760, 716-372-6121

MEMBERSHIP LIST 1993 159

SCHULTZ, MONICA—Morris Arboretum, 9414 Meadowbrook Avenue, Philadelphia PA 19118

SCHUYLER, ALFRED E.—Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia PA 19103,
215-299-1193

SCOTT, CONNIE—139 Myrtle Avenue, Havertown PA 19083, 610-688-2603

Scott, JOHN D.—R.D. 1, Box 249D, Hertzog School Road, Mertztown PA 19539, 610-682-2809

SEAGER, KEITH A.—278 Brn Creek Road, Cape May NJ 08204, gs 884-8778

SEIP, ROBERT G.—R.D. ox 683, Alburtis PA 18011, 610-845-207

SELL, LAURA M.—118 os Valley Drive, Medford NJ 08055, pape 5174

SETTLEMYER, KENNETH T.—219 Maple St., Jersey Shore PA 17740, 717-398-2546

SHAEFER, CHARLOTTE—2976 Dorman Avenue, Broomall PA 19008, 610-356-0497

SIPPLE, ¢ Namen Red Bluff Court, Millersville MD 21108

SMITH, BARBARA—11 Cavendish Drive, Ambler PA 19002, 215-628-2277

SMITH, DOUGLAS—352 Fieldbrook Drive, Pittsburgh PA 15228, 412-561-7267

SNYDER, DAVID—12 Chesterfield-Georgetown Road, Trenton NJ 08620, 609-298-1555

SO, ANTHONY—8 Rex Avenue, #11, Philadelphia PA 19118

SOBOLOW, BERNADETTE LLOYD—433 Wadsworth Avenue, pers PA 19119

STAHL, HERBERT—22 Lincoln Avenue, Lansdale PA 19446, 215-855-9921

STAILEY, HELEN M.—8701 Macon St., Philadelphia PA ite 215-673-8163

STALTER, RICHARD—St. ig University, Grand Central & Utopia Parkway, Jamaica NY 11439, 718-990-5237

STASZ, igi L.—P.O. Box 71, North Beach MD 20714, 410-952-365

STEVENS, RLES E.—615 Kia Place, Charlottesville VA 22903, 804-293-8658

STORBECK, Caen ne Ravenwood Road, Exton PA 19341

STUCKEY, RONALD L.—1315 Kinnear Road, Columbus OH 43212

SWEETMAN, HAROLD—Jenkins Arboretum, 631 Berwyn Baptist Road, Devon PA 19333, 610-647-8870

TEES, GRACE M.—Rydal Park, Rydal PA 19046

THOMPSON, SUE—Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh PA 15213, 412-622-3295

Tiss, KENNETH A.—49 Princeton Road, Havertown PA 19083, 610-446-6746

TRASATTI, DANIEL—204 Edith Road, West Chester PA 19380, 610-692-2897

TREW, LESLIE D.—Delaware Natural Heritage Inventory, P.O. Box 1401, Dover DE 19903, 302-796-5825
CKER, ARTHUR O.—Delaware State College, Dover DE 19901, 302-739-5120

UDELL, VAL—2746 Yost Road, Perkiomenville PA 18074, 610-754-7163

UTECH, FREDERICK H.—Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh PA 15213,
412-622-3296

VIRKLER, ROBERT J—429 Elliger Avenue, Ft. Washington PA 19034

VIVIAN, V. EUGENE—284 Country Club Boulevard, Little Egg Harbor NJ 08087, 609-296-7479

VOLLMER, JOHN—42 Burrs Mill Road, Southampton NJ 08088, 609-859-28

WEINER, JACOB—Department of Biology, Swarthmore eg Swarthmore PA 19081, 610-328-8038

WEINSTEIGER, EILEEN—R.D. 1, Box 662, Alburtis PA 1

WENGER, CONSTANCE L.—1090 Street Road, Oxford PA fete 610-932-5361

WHEELER, STEPHEN A.—312 W. 6th S St., Boyertown PA 19512, 610-369-3768

WIEBOLDT, THOMAS F.—2488 Crab Creek Road, Christiansburg VA 24073

WILKENS, HANS—Lutheran Home, Topton PA 16684

WILLIAMS, CARL—165 W. Ridge Pike, Limerick PA 19468

WILLIAMS, DaviD L.—641 Coppermine Road, Princeton NJ 08540, 201-297-0642

WILLIAMS, JAMES W.—Box 222, Wycombe PA 18980, 215-357-6533

WILLIG, SARAH ANDERSEN—R.D. 2, Box 435, Phoenixville PA 19460, 610-933-3539

WILSON, RONALD—3740 Ridge Road, Snow Hill MD 21863, 410-632-3892

haa ANDREW—P.O. Box 312, Chatsworth NJ 08019, 609-726-9054

WOLFF, EMILY T.—295 E. Rose Tree Road, Media PA 19063, 610-566-4907

Wenee , JOHN—2640 Breezewood Drive, Lancaster PA 17601, 717-569-6955

WoLrr, PETER—79 Pennell Road, tana. PA 19037, 215-565-9973

Woop, Howarp P. Fe Darby Road C-802, Haverford PA 19041, 610-642-9963

WYGANT, NANCY—815 S. 49th St. #3, Philadelphia PA 1914

ZAREMBA, ROBERT E.—125 Circular St. #4, Saratoga Springs NY 12866, 518-463-6133

Program of Meetings 1991 and 1992

Date Subject Speaker
199]
24 Jan The Pause that Refreshes: 1791-1891-1991. Botany in Philadelphia and

Sebago rs a Sy Re ee a Joseph Ewan
28 Feb John. and William Bartram ©... ooo kc ids ok Oe ee ge es Elizabeth P. McLean

ORE, sponsored with the John Bartram Asso
28 Mar and Endangered Species Symposium. pesag ee by Ann F. Rhoads.

Jo an gers with the Morris Arboretum
25 & 26 Apr Research Perspectives in the Pinelands. Symposium coordinated b

4 i E. Fairbrothers & Ralph E. Good. Jointly sponsored with Rutgers

rsity

23 May peas of the Philadeiphia Awa). oe ss ee Peggy Bowditch

Jointly sponsored with the Pennsylvania Horticultural Society
26 Sep The Collector's Garden 2. 6 5 SUE nae ee eh ee ee ee ee
24 Oct Changes in Our Flora with a Comparison to Barton’s 1818 Flora ......... Alfred E. Schuyler
21 Nov Wetlands Flora: Diversity, Function, and Importance. Symposium

coordinated by Jane Rowan. Jointly sponosored with the Academy of

Natural Sciences, Stroud Water Research Center
19 Dec Changing Ecology: of one Blosphete . 22>. . ig. os aes eh eas Ruth Patrick
1992
23 Jan Hunting Dendrobium Orchids in New Zealand, Australia, New Guinea,

Bh i xe ee SE es i ke ee Howard P. Wood

27 Feb Rare Plants of the Piedmont of Chester County, Pennsylvania..............- Robert J. Holt
26 Mar American Lotus (Nelumbo lutea) in New Jersey and Pennsylvania ....... William S. Ettinger
23 Apr Asymmetric Competition in Plant Populations ........-..--- 5+ eee eee ees Jacob Weiner
28 May —— Guat ing Bogs, Turfcuts and Cedar Swamps of the New Jersey

Pine Barees Sou oo ee ee os ee ree Ted Gordon
24 Sep Slide eee Report of Summer Botanizing ........---.++-+-sse002- Club Members
22 Oct New Jersey Ferms and Fern Allies: 2. 52 oo. 0.3 55 ee ne eee James Montgomery
19 Nov Darlington’s a Erg Revisited (1854) =... ..5 ee Dorothy Lansing
17 Dec Cichlids of Net dation 5 a Mark Lar

;
e

IGUa:
ene:

ISSN 0198-7356

BARTONIA

JOURNAL OF THE
PHILADELPHIA BOTANICAL CLUB
No. 59
CONTENTS
Annotated Checklist of the Plants of the Wissahickon Valley ............ JOHN M. FoGG, JR. 1

The Identity of Anemone riparia (Ranunculaceae)
CARL S. KEENER, EDWARD T. DIx, AND BRYAN E. DUTTON 37

Distribution, Life Forms, Taxonomic Categories, and Habitats of the Endangered
JEFF!

Threatened Vascular Plants in Pennsylvania: A Summary ............ y L. WALCK 49
he Genus Niece lu Mew Jereey ae DAVID SNYDER 55
The Se cam Work of the Ruth Brothers of Bucks County, piace and its Significance

TONNE ek ee a ey Pe ay ELA WHITE AND ANN OADS §671

Additions to the Flora of Bradford County, Pennsylvania
ROBERT F. C. NACZI AND JOHN W. THIERET 81

samen a of Eriophorum gracile Koch (Cyperaceae) at its Southern ee Limit in
Penn AMILLE BARR 87

ES Status of Panicum hirstii Swallen .........-. 60-2 -25+. +s: ALFRED E. SCHUYLER 95
Soil Chemistry Data for Species of Special Concern in Pennsylvania ...... RICHARD MELLON 97
Soil Chemical and Physical Properties Associated with Two Pennsylvania feces Plant
Species (Juncus biflorus and — dumosus) at Crow’s Nest Farm, Chester County
HN T. LAROCCO, ANNE K. WEBSTER, AND R. nate WIEDER 107
Polypodium appalachianum, P. virginianum, and Their Hybrid in New gan ip Pennsylvania
JAMES D. MONTGOMERY 113
Observation of Polyembryony in Trillium and Smilacina .................- JOHN F. GYER
The Gold-Cone Tamarack (Larix laricina forma lutea, Pinaceae) in Pennsylvania
ROBERT F. NACZI AND JOHN W. THIERET 123
Paintings of Fungi by Lewis David von Schwinitz in the Archives of the engine " Natural
Sciences of PRMMMORI a ee ae ae es eee es A M. LYNcH 125

Contents continued on inside back cover

PUBLISHED BY THE CLUB
ACADEMY OF NATURAL SCIENCES, 1900 BENJAMIN FRANKLIN PARKWAY
PHILADELPHIA, PENNSYLVANIA 19103-1195
Issued 4 September 1996

The Philadelphia Botanical Club

Editor: Alfred E. Schuyler
Editorial Assistant: Elizabeth B. Farley

Editorial Board

DAVID E. FAIRBROTHERS CARL S. KEENER
JAMES D. MONTGOMERY RONALD L. STUCKEY

Officers of the Philadelphia Botanical Club for 1991-1993
President: TED GORDON Vice President: ROBERT J. HOLT

Recording Secretary: WILFRED R. NOBLE Treasurer: MILTON LADEN
Corresponding Secretary: ELIZABETH B. FARLEY Curator: JOSEPH ARSENAULT

Bartonia No. 59: 1-36, 1996

Annotated Checklist of the Plants of the
Wissahickon Valley

BOTANICAL

OCT1 4 1996

JOHN M. Foe, Jr.!
GARDEN LIBRARY

Interest in the plant-life of the Wissahickon Valley dates back more than two and a half
centuries. In 1694 the German Pietists, under the leadership of Kelpius, established in the
lower Wissahickon a garden for the growing of medicinal plants. Doubtless many of the
native plants of the Valley were cultivated by them for their real or supposed therapeutic
values

The region was well known to John Bartram, founder of another famous garden on the
banks of the Schuylkill River, and during the first half of the nineteenth century the Valley
was visited by Frederick Pursh and Thomas Nuttall, two of the most distinguished students
of the flora of eastern North America. Throughout the latter half of the last century the
Wissahickon was a botanical hunting ground for such eminent naturalists and botanists as
Joseph Leidy, Joseph T. Rothrock, Thomas C. Porter, and J. B. Brinton. Unfortunately,
these men, although well acquainted with the plants of the Valley, collected few if any
specimens, and left almost no record of the occurrence of individual species.

Between 1900 and 1925 the Wissahickon was the scene of considerable botanical activity.
Two botanists in particular, S. S. Van Pelt and Henry A. Lang, made numerous collections
in the Valley and their specimens are in the herbaria of the Philadelphia Academy of Natural
Sciences, and the University of Pennsylvania. Although Mr. Van Pelt was careful to note
on his labels the localities from which his plants were collected, Mr. Lang’s specimens are
usually designated merely “along the Wissahickon Creek” or “Wissahickon Creek Ravine,”
and we have no means of knowing the exact source of his material.

In 1937-38 the Friends of the Wissahickon established a fellowship to a graduate student
at the University of Pennsylvania for the purpose of making a botanical survey of the Valley.
The holder of the fellowship was Mr. Ernest H. Ludwig, then a student of the present author.
Mr. Ludwig performed a valuable service by going through the mounted collections at the
Academy and the University and listing all of the specimens from the Valley. He also did
some collecting of his own, but his work was interrupted by the war, and his specimens were
never labeled or mounted.

In 1953, at the instigation of the Hon. Harold D. Saylor, Vice President of the Friends of
the Wissahickon, the author undertook to revise and edit Mr. Ludwig’s compilation and to
augment it with his own observations based on many years’ acquaintance with and countless
walks in the valley.

‘This checklist is posthumously published through the urging of the Friends of the Wissahickon with the hope that
it will stimulate renewed interest in recording the present status of the Wissahickon flora. Current nomenclature has

added in parentheses to correspond with Rhoads and Klein (1993). Similarly, changes in family oo
have been indicated by placement of the current family name, where it differs from that used by Fogg,
parentheses. For information about Dr. Fogg see Hill, Farley, and Fogg 1983.

2 BARTONIA

The present list is admittedly incomplete. Many species doubtless still remain to be
discovered. Others reported as rare may prove to be abundant. Still others, included on the
basis of older collections, may no longer be members of the present-day flora. The list is
offered at this time in the hope that it may stimulate those who are interested in the plant-life
of the Valley to make and report their own observations to the end that a really comprehen-
sive and up-to-date checklist may be assembled.

The following list includes the more important and familiar of higher (vascular) plants
believed to occur in the Valley of the Wissahickon Creek from the Schuylkill River to
Northwestern Avenue in Chestnut Hill, including the extensions into the Cresheim Valley
and along Bells Mill Road. Omitted from the enumeration are several families which, while
of interest to the botanist, contain forms which are either so inconspicuous or so difficult to
classify, that their inclusion in a popular check-list seems inadvisable. Among these are the
grasses (Gramineae), sedges (Cyperaceae), and rushes (Juncaceae). Those desiring to study
these groups should consult one of our standard manuals or floras, such as Gray’s Manual
of Botany (Fernald 1950), The New Britton and Brown Illustrated Flora of the Northeastern
United States and Adjacent Canada (Gleason 1952), or Manual of Vascular Plants of
Northeastern United States and Adjacent Canada (Gleason & Cronquist 1991).

e arrangement of the families in this list conforms in general to that found in the above
works. Within each family, however, the sequence of genera and species is, for the sake of
convenience, alphabetical.

SUMMARY OF SPECIES AND VARIETIES

Native Introduced Total
36 0 36

Pteridophytes

Gymnosperms 5 2 7
Monocotyledons 37 12 49
Dicotyledons 417 145 562
TOTALS 495 159 654

LIST OF LOCALITIES PROCEEDING UPSTREAM

Mouth of Creek, Reading R. R. Bridge, Gipsy Lane, Picnic Grounds, Hermit’s Lane, Henry
Avenue, Paper Mill Run, Bluestone Bridge, Indian Profile Rock, Walnut Lane, Roxborough
Avenue, Kitchen’s Lane, Monastery, Leverington Street, Carpenter’s Woods, Mount Airy
Avenue, Gorgas Lane, Livezey Lane, Devil’s Pool, Cresheim Creek , McCallum Street, Bog
Garden, Lake Surprise, Valley Green, Springfield Avenue, Hartwell Lane, Wise’s Mill Road,
Summit Avenue, Paper Mill, Rex Avenue, Indian Rock Bridge, Tedyuscung Statue, Thomas
Mill Road, Red Covered Bridge, Bells Mill Road, (Thorps Lane), Barren Hill Road,
Northwestern Avenue, Harper’s Meadow, Germantown Avenue.

LITERATURE CITED

FERNALD, M.L. 1950. Gray’s Manual of Botany. American Book Company, New York.
GLEASON, H.A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent
Canada. New York Botanical Garden, Bronx.

WISSAHICKON VALLEY PLANTS 3

GLEASON, H.A. & A. CRONQUIST. 1991. Manual of en Plants of Northeastern United States and Adjacent
Canada. 2nd ed. The New York Botanical Garden,

HILL, R., E.B. FARLEY, & ~ FoGcG. 1983. Milton akg in (1898- 1982). Bartonia No. 49:

RHOADS, A.F. & W.M. KLEIN. 1993. Vascular Flora of Pennsylvania: Annotated Checklist a hae American
Philosophical Society, Phitibei

PTERIDOPHYTES
EQUISETACEAE, HORSETAIL FAMILY
Equisetum arvense. Common horsetail. Fairly frequent in low ground. Appearing early
spring. Walnut Lane, Valley Green, Rex Avenue.

LYCOPODIACEAE, CLUB-MOSS FAMILY
Lycopodium lucidulum (Huperzia lucidula). Shining club-moss. Rich woods, usually in deep
humus. Uncommon. Rex Avenue, Thomas Mill Roa
Lycopodium obscurum. Flat-branched ground-pine. Dry woods and thickets. Known from
only a few localities, e.g. above Hartwell Avenue.

SELAGINELLACEAE, SPIKE-MOSS FAMILY
Selaginella apoda. Meadow spike-moss. Small, inconspicuous plant suggesting a moss in
appearance. Found occasionally in meadows or grassy clearings. Gorgas Lane, Cresheim
Valley.

OPHIOGLOSSACEAE, ADDER’S TONGUE FAMILY

Botrychium dissectum. Lace-leaf grape-fern. Fairly frequent on moist to dry wooded slopes
of the ravine. Hartwell Avenue.

Botrychium obliquum (B. dissectum). Coarse-lobe grape-fern. Same occurrence, and often
with B. dissectum. Bells Mill Road.

Botrychium virginianum. — Rattlesnake-fern. Rich, deciduous woods. Fairly common
throughout Wissahickon and Cresheim valleys, especially northward. Shawmont Avenue,
Kitchen’s Lane.

OSMUNDACEAE, FLOWERING FERN FAMILY
Osmunda cinnamomea. Cinnamon fern. Low, damp ground, —— etc. Locally abundant,
especially in the upper portion. Rex Avenue, Thomas Mill
Osmunda claytoniana. Interrupted fern. In similar habitats, se less frequent than O.
cinnamomea. Valley Green, Thomas Mill Road.
Osmunda regalis. Royal fern, Wet woods and thickets. Rare. Known only from Bells Mill
Road west of creek.

pi Peng gted FERN FAMILY

(Adiantaceae)
Adiantum pedatum. Maiden-hair fern. Rich, oa woods. Occasionally met in the upper
Wissahickon. Rex Avenue

(Aspleniaceae)

Asplenium pinnatifidum. Lobed spleenwort. Crevices of rock outcrops. Rare. Not seen in
recent years.

Asplenium platyneuron. Ebony spleenwort, brownstem spleenwort. Open woods, wooded

4 BARTONIA

slopes, rock faces, etc. Only occasional throughout the ravine. Thomas Mill Road.
Asplenium trichomanes. Maiden-hair spleenwort. Damp, shaded crevices and rock ledges.
Infrequent, but known from three or four localities in the ravine. Thomas Mill Road.
ual g ied rhizophyllus aap rhizophyllum). Walking fern. Shaded rock ledges.
e. Thomas Mill Road

(Blechnaceae)
Woodwardia areolata. Broad-leaved chain fern. Wet woods, swamps, etc. Rare. Rex Avenue.

(Dennstaedtiaceae)

Dennstaedtia punctilobula. Hay-scented fern. Common throughout in open, shaded woods,
rock faces, damp slopes, etc. One of the most abundant ferns in the area. Henry Avenue,
Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Livezey Lane, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road, Thorps Lane.

Pteridium aquilinum. Bracken fern. A variable species represented in our area chiefly by the
variety /atiusculum. Dry woods, clearings. Occasional. Hartwell Avenue, along bridle path
south of Bells Mill Road.

(Dryopteridaceae)

Athyrium angustum, var. rubellum (A. filix-femina var. angustum). Northern lady fern. Moist
woods and thickets. Fairly common. Kitchen’s Lane, Valley Green, Rex Avenue, Bells
Mill Road

Athyrium asplenoides (A. filix-femina var. aspleniodes). Southern lady fern. Similar habitats,
but less frequent than A. angustum. Thomas Mill Road.

Athyrium thelypteroides (Deparia acrostichoides). Silvery spleenwort. Rich woods and
shaded slopes. Plentiful throughout. Henry Avenue, Walnut Lane, Livezey Lane, Cresheim
Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.

Cystopteris fragilis var. mackayii (C. tenuis). Fragile fern. Damp rocks, stony outcrops, open
woods, etc. Fairly common. Walnut Lane, Cresheim Valley, Thomas Mill Road.

Dryopteris cristata. Crested wood fern. Rich woods. Rare. Thomas Mill Road.

Dryopteris goldiana. Giant wood fern. Rich woods. Rare, known only from second ravine
above mouth of creek.

Dryopteris marginalis. Marginal wood fern. Rich woods, rocky slopes. Fairly frequent.
Henry Avenue, Walnut Lane, Livezey Lane, Cresheim Valley, Thomas Mill Road.

Dryopteris spinulosa (D. intermedia). Common wood fern. A variable species, represented
in our area by the variety intermedia. Dry to moist woods and thickets. Walnut Lane,
Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.

Onoclea sensibilis. Sensitive fern. Low woods and thickets, open ground along trails, etc.
Frequent throughout. Hermit’s Lane, Bluestone Bridge, Walnut Lane, Carpenter’s Woods,
Cresheim Valley, Thomas Mill Road.

Polystichum acrostichoides. Christmas fern. Woods, shaded and often rocky slopes. Common
throughout. Walnut Lane, Kitchen’s Lane, Livezey Lane, Cresheim Valley, Valley Green,
Rex Avenue, Thomas Mill Road, Bells Mill Road.

Pteretis pensylvanica (Matteuccia struthiopteris). Ostrich fern. Rich woods, bottomland, etc.
Rare. Known only from bog garden along Cresheim Creek, and below Harper’s Meadow.

Woodsia obtusa. Blunt-leaved cliff fern. Rock ledges, dry woods, etc. Infrequent, known
only from Wise’s Mill Road and above Thorps Lane.

WISSAHICKON VALLEY PLANTS §

(Polypodiaceae)
Polypodium virginianum. Rock fern. Rocks, crests of ledges, cliff faces, etc. Of very local
occurrence in steep, shaded areas of ravine. Thomas Mill Road, Bells Mill Road.

(Thelypteridaceae)

Thelypteris dryopteris. Oak fern. Dry woods. Rare. Known only from near Graver’s Lane.

Thelypteris hexagonoptera. Broad beech-fern. Rich to dry woods. Rare. Hartwell Avenue.

Thelypteris novaboracensis. New York fern. Dry to damp woods, slopes and thickets.

ommon throughout. Walnut Lane, Livezey Lane, Cresheim Valley, Rex Avenue, Thomas
Mill Road, Bells Mill Road.

Thelypteris palustris. Marsh fern. A wide-ranging species, represented in our area by the
variety pubescens. Swamps, meadows, wet thickets. Occasional in more open sections of
the valley and along Cresheim Creek.

Thelypteris phegopteris. Long beech-fern. Woods and thickets. A northern species found
many years ago near Thomas Mill Road; locality obliterated by Japanese honeysuckle.

GYMNOSPERMS
PINACEAE, PINE FAMILY
(Cupressaceae)
Juniperus virginiana. Juniper, red cedar. Dry slopes, clearings. Cccenicned in open woods,
especially on slopes and summits of ridges. Bluestone Bridge.

(Pinaceae)

Pinus nigra var. austriaca. Austrian black pine. Planted in certain areas, e.g: near Walnut
Lane Bridge

Pinus rigida. Pitch pine. Occasional on slopes and summits of ridges. Along bridle trail
above Thomas Mill Road.

Pinus strobus. Eastern white pine. Although white pine may originally have occured in the
valley, the trees growing here today are the result of recent plantings. A fine grove exists
along Hermit’s Lane near Hermit Street.

Pinus sylvestris. Scots pine. Planted in several areas and possibly naturalized.

Pinus virginiana. Scrub pine. Probably persisting from earlier plantings. Occasional on dry,
rocky slopes and crests; near mouth of creek, Carpenter’s Woods, below Bells Mill
Road.

Tsuga canadensis. Hemlock. One of the commonest and most handsome trees of the valley.
Often forming pure stands, especially on the west or north facing slopes, as at: Walnut
Lane, Valley Green, south of Rex Avenue, above Covered Bridge.

MONOCOTS
ALISMACEAE, WATER-PLANTAIN FAMILY
Sagittaria australis. Southern arrow-head. a water. Collected in the valley in 1908,
but no locality given. Perhaps no longer presen
Sagittaria latifolia. Common arrow-head. —— water and muddy shores. Infrequent.
Above Walnut Lane.

6 BARTONIA

ARACEAE, ARUM FAMILY
(Acoraceae)
Acorus calamus. Sweet flag. Marshes, wet meadows. Very local. Cresheim Valley, Harper’s
Meadow.

(Araceae)

Arisaema triphyllum. Sack-in-the-pulpit. Woods and thickets. Common. Henry Avenue,
Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Cresheim Creek, Mt.
Airy Avenue, Valley Green, Rex Avenue, Bells Mill Road, Harper’s Meadow.

Symplocarpus foetidus. Skunk cabbage. Wet meadows and thickets, swampy woods.
Common in bottomland. Henry Avenue, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue,
Cresheim Creek, Allen’s Lane, Thomas Mill Road, below Bells Mill Road.

COMMELINACEAE, DAYFLOWER FAMILY
Commelina communis. Dayflower. Low, moist ground, roadsides, etc. Fairly common
throughout the valley. Naturalized from Asia. Bluestone Bridge, Walnut Lane, Livezey
Lane, Valley Green, Thomas Mill Road, Bells Mill Road.
Tradescantia virginiana. Spiderwort. Woods, thickets, meadows. Occasional. Probably
naturalized from farther south. Thorps Lane.

PONTEDERIACEAE, PICKERELWEED FAMILY
Heteranthera reniformis. Mud plantain. Collected many years ago from shallow water in the
ower Wissahickon. Not seen recently and possibly extirpated.

LILIACEAE, LILY FAMILY

Aletris farinosa. Colicroot, star-grass. Dry sand and gravel. Rare. Gorgas Lane.

Allium tricoccum. Wild \eek. Rich woods and bottomland. Conspicuous in spring because
of its elliptic, fleshy leaves which soon die, leaving the flowers to appear on naked stalks
in early summer. Rare and local. Mt. Airy Avenue, Rex Avenue, Thomas Mill Road, Bells
Mill Road.

Allium vineale. Field garlic. A troublesome weed, naturalized from Europe and found
commonly along trails in grassy clearings, etc. Near mouth of creek, Walnut Lane,
Carpenter’s Woods, Valley Green.

Asparagus officinalis. Asparagus. Escaped from cultivation and often persistent in disturbed

ground.

Chamaelirium luteum. Devil’s bit. Rich to dry woods and thickets. Rare in upper valley; not
collected since 1910.

Convallaria majalis. Lily-of-the-valley. Occasional as an escape from gardens bordering the
valley.

Erythronium americanum. Trout-lily, dogtooth-violet. Rich to sandy woods and thickets,
bottomland. Fairly frequent throughout. Henry Avenue, Bluestone Bridge, Kitchen’s Lane,
Mt. Airy Avenue, Cresheim Creek, Valley Green, Rex Avenue, Thomas Mill Road, Bells
Mill Road.

Hemerocallis fulva. Day-lily. Frequently established. Kitchen’s Lane, Cresheim Creek,
Thomas Mill Road.

Hosta plantaginea. Plantain-lily. Sparingly escaped from cultivation. Below Mt. Airy
Avenue.

WISSAHICKON VALLEY PLANTS |

Hosta ventricosa. Plantain-lily. Frequent as an escape. Henry Avenue, Cresheim Creek,
Valley Green, Rex Avenue.

Maianthemum canadense. Wild lily-of-the-valley. Woods, thickets and dry clearings.
Frequent throughout. Above Henry Avenue, Bluestone Bridge, Walnut Lane, Mt. Airy
Avenue, Cresheim Creek, Bells Mill Road.

Medeola virginiana. Indian cucumber-root. Rich woods. Fairly common. Kitchen’s Lane,
Livezey Lane, Cresheim Creek, Valley Green, Rex Avenue, Thomas Mill Road.

Muscari botryoides. Grape-hyacinth. Introduced and naturalized from Europe. Occasional in
woods and clearings. Thomas Mi

Ornithogalum umbellatum. Star-of-Bethlehem. Introduced and naturalized from Eurasia.
Fairly common in woods, thickets, and bottomland. Henry Avenue, Bells Mill Road,
Kitchen’s Lane, Thomas Mill Road.

Polygonatum canaliculatum (P. biflorum var. commutatum). Great Solomon’s-seal. Woods,
thickets and rich bottomlands. Fairly frequent. Mt. Airy Avenue, Cresheim Valley,
Harper’s Meadow.

Polygonatum pubescens. Small Solomon’s-seal. Rich woods and rocky slopes. Occasional.
Henry Avenue, Carpenter’s Woods, Valley Green, Rex Avenue, Bells Mill Road.

Scilla sibirica. Siberian squill. Established in rich woods. Rare. Thomas Mill Road.

Smilacina racemosa. False Solomon’s-seal. Woods and clearings. Frequent throughout.
Henry Avenue, Bluestone Bridge, Walnut Lane, Carpenter’s Woods, Kitchen’s Lane, Mt.
Airy Avenue, Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill
Road, Harper’s Meadow.

Tricyrtis hirta. Toad-lily. Introduced and naturalized from Asia. An occasional escape from
gardens bordering the valley. Hartwell Avenue.

Trillium cernuum. Wake-robin, nodding trillium. Rich woods and thickets. Rare. Thomas
Mill Road.

Uvularia sessilifolia. Bellflower. Rich woods. Kitchen’s Lane, Thomas Mill Road.

(Smilacaceae)

Smilax glauca var. leurophylla. Sawbrier. Occasional in dry, rocky woods. Henry Avenue,
Walnut Lane, Carpenter’s Woods, Thomas Mill Road, Bells Mill Road.

Smilax herbacea. Carrion-flower. Woods and alluvial thickets. Infrequent. Walnut Lane,
Thomas Mill Road.

Smilax rotundifolia. Common greenbrier. Moist to dry woods and thickets. Known from
several localities in the ravine. Hermit’s Lane, Carpenter’s Woods, Valley Green, Thomas
Mill Road, Bells Mill Road.

AMARYLLIDACEAE, AMARYLLIS FAMILY
(Liliaceae
Hypoxis hirsuta. Yellow star-grass. Dry woods and clearings. Occasional. Cresheim Creek,
Valley Green, Bells Mill Road.
Narcissus pseudo-narcissus. Daffodil. Locally escaped from cultivation. Thomas Mill Road.

DIOSCOREACEAE, YAM FAMILY
Dioscorea villosa. Wild yam. Dry woods and thickets. Apparently rare. Known from
Carpenter’s Woods and from location one-quarter mile below Bells Mill Road.

8 BARTONIA

IRIDACEAE, IRIS FAMILY
Iris pseudoacorus. European yellow iris. Naturalized near Harper’s Meadow.
Sisyrinchium angustifolium (S. graminoides). Broad-leaved blue-eyed-grass. Dry woods and
thickets. Occasional in the ravine.
Sisyrinchium mucronatum. Narrow-leaved blue-eyed-grass. Similar habitat to S. angustifoli-
um. Rare. Known only from the upper Wissahickon.

ORCHIDACEAE, ORCHID FAMILY

Corallorhiza odontorhiza. Autumn coral-root. Rich to dry woods, often in deep humus. Rare.

Corallorhiza wisteriana. Coral-root. Rich woods. Collected somewhere in the valley in 1839.
Not seen in recent years and probably extirpated.

Cypripedium acaule. Pink moccasin-flower. Dry woods and clearings. Rare. Bells Mill Road.

Cypripedium parviflorum var. pubescens (C. calceolus var. pubescens). Mocassin-flower.
Rich woods. Collected in the valley many years ago. Not recently observed.

Goodyera pubescens. Downy rattlesnake-plantain. Moist to dry woods and thickets. Known
only from extreme upper portion of the valley. Thorps Lane, Chestnut Hill.

Isotria verticillata (Pogonia verticillata). Purple five-leaved orchid. Extremely rare. Known
only from slope above Bells Mill Road.

Liparis lilifolia. Twayblade. Rich to loamy woods. Rare. Hartwell Avenue.

Malaxis unifolia. Adder’s-tongue. Rich woods. Rare. Known only from near Devil’s Pool.

Orchis spectabilis (Galearis spectabilis). Showy orchid. Rich woods. Rare. Found in recent
years only below Livezey Lane.

Spiranthes cernua. Common a -tresses. Low, wet woods and thickets. Infrequent. Along
Cresheim Creek, Thorps Lan

Spiranthes gracilis (S. lacera var. ee Slender ladies’-tresses. Open woods and thickets.
Rar own only from Thomas Mill Road.

Triphora trianthophora. Three-birds. Collected in the 1860’s at the extreme upper end of the
valley, and not rediscovered in recent years. Probably obliterated.

DICOTS
SAURURACEAE, LIZARD’S TAIL FAMILY
Saururus cernuus. Lizard’s tail. Muddy shores. Occasional near mouth of creek. Not recently
seen probably due to pollution of river.

SALICACEAE, WILLOW FAMILY

Populus grandidentata, Large-toothed aspen. Dry woods. Infrequent. Carpenter’s Woods,
Cresheim Valley, Thomas Mill Road, below Bells Mill Road.

Populus tremuloides. Quaking aspen. Dry slopes. Apparently rare. Carpenter’s Woods.

Salix babylonica. Weeping willow. Occasional as an escape in low, damp ground. Near
Reading R.R. bridge, Bluestone Bridge, Carpenter's Woods, Lake Surprise, Harper's
Meadow.

Salix caprea. Goat willow. Introduced from Europe. Rare in the ravine.

Salix fragilis. Crack willow. Occasional along stream banks. Often a good-sized tree.

Salix nigra. Black willow. Usually a shrub. Wet thickets and stream banks. Infrequent.
Carpenter’s Woods, Mt. Airy Avenue, Cresheim Valley.

_—

—

WISSAHICKON VALLEY PLANTS 9

Salix purpurea. Basket willow. Introduced from Europe for basket making and occasionally
escaped. Known only from west bank above Reading R.R. bridge and near Bells Mill
Road.

Salix sericea. Silky willow. Damp thickets. Cresheim Valley.

MYRICACEAE, WAX-MYRTLE FAMILY
Comptonia peregrina. Sweet-fern. Occasional on dry, acid slopes in the upper Wissahickon,
Thomas Mill Road.

JUGLANDACEAE, WALNUT FAMILY

Carya alba (C. tomentosa). Mockernut. A tall, handsome tree, found usually on dry,
west-facing slopes.

Carya cordiformis. Bitternut, swamp hickory. Stream banks. Known only from lower valley,
where rare.

Carya glabra. Pignut. Dry woods and slopes. Not common. Near mouth of creek, Bluestone
Bridge, Gorgas Lane, above Bells Mill Road.

Carya ovata. Shag-bark hickory. Rich woods, bottomland. Sporadic throughout. Carpenter’s
Woods, Cresheim Creek.

Juglans cinerea. Butternut. Occasional in rich woods.

Juglans nigra. Black Walnut. Fine specimens occur in rich woods, e.g: Hermit’s Lane,
Walnut Lane, Cresheim Valley, Valley Green, Harper’s Meadow.

CORYLACEAE, HAZEL FAMILY
(Betulaceae)

Alnus glutinosa. European alder. Occasional as an escape from cultivation. Picnic grounds
below Henry Avenue.

Alnus serrulata. Common alder. Occasional along margin of streams and in low, wet ground.
Walnut Lane, Cresheim Valley

Betula lenta. Sweet, black, or cherry birch. Common throughout. Walnut Lane, Devil’s Pool,
Cresheim Valley, Valley Green, Thomas Mill Road, Bells Mill Road.

Betula nigra. River birch. Low ground, close to the stream. Infrequent. Along both sides of
the stream between mouth and Henry Avenue, Bluestone Bridge, Thomas Mill Road,
Harper’s Meadow.

Betula populifolia. Gray birch. Dry, sterile ground, fields, etc. Occasional. Upper
Wissahickon, Carpenter’s Woods, Cresheim Valley, above Thomas Mill Road.

Carpinus caroliniana. Hornbeam, Ironwood. Bottomland, usually close to stream. Fairly
frequent. Bluestone Bridge, Walnut Lane, Thomas Mill Road, Harper’s Meadow.

Corylus americana. Hazel-nut. Thickets. Infrequent. Above Henry Avenue.

Ostrya virginiana. Hop-hornbeam. Bottomland and lower slopes, especially in upper portion
of valley, but nowhere common. Thomas Mill Road.

AGACEAE, BEECH FAMILY

Castanea dentata. American chestnut. Formerly one of our finest trees; today known only

as regeneration shoots from old stumps, some of them old enough to bear flowers and

fruit, but apparently none resistant to blight. Bluestone Bridge, Thomas Mill Road, Bells
Mill Road, Harper’s Meadow

Fagus grandifolia. American bench Common throughout in low ground and wooded slopes;

10 BARTONIA

good specimens occur near Henry Avenue, Walnut Lane, Carpenter’s Woods, Thomas
Mill Road, Bells Mill Road, Harper’s Meadow.

Quercus alba. White oak. Common throughout in dry ground and on rocky slopes. Henry
Avenue, Walnut Lane, Carpenter’s Woods, Thomas Mill Road, Bells Mill Road, Harper’s
Meadow.

Quercus bicolor. Swamp white oak. Bottomland, stream borders. Occasional. Thomas Mill
Road.

Quercus coccinea. Scarlet oak. A handsome tree in the autumn. Occasional throughout.
Walnut Lane.

Quercus palustris. Pin oak. Occasional in low ground. North of Bells Mill Road.

Quercus prinus (Q. montana). Chestnut oak. Dry woods and rocky slopes. Fairly frequent
throughout. Henry Avenue, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Livezey

Lane.

Quercus rubra. Red oak. A variable tree as to size and shape of leaves. Common throughout
in low land and on rich slopes. Hermit’s Lane, Henry Avenue, Walnut Lane, Kitchen’s
Lane, Carpenter’s Woods, Cresheim Valley, Rex Avenue, Bells Mill Road.

Quercus velutina. Black oak. Dry woods. Occasional.

ULMACEAE, ELM FAMILY

Celtis occidentalis. Hackberry. Throughout in low ground and wooded slopes. Often
distinguished by the presence on the branches of witches-brooms. Near mouth of creek,
Hermit’s Lane, Walnut Lane, Livezey Lane.

Ulmus americana. Common elm. Rich soil, especially along streams. Occasional in lower
valley, especially between mouth of creek and Henry Avenue.

Ulmus rubra. Slippery elm. Usually distinguished from the preceding by its harsh upper leaf
surfaces. Common throughout in bottomland and on lower slopes. Mouth of creek, Henry
Avenue, Bluestone Bridge, Walnut Lane, Kitchens Lane, Cresheim Valley, Valley Green,
Thomas Mill Road, Bells Mill Road.

MORACEAE, MULBERRY FAMILY

Broussonetia papyrifera. Paper mulberry. Introduced from Asia and naturalized sparingly in
the valley. Above Henry Avenue.

Maclura pomifera. Osage-orange. Naturalized near mouth of creek.

Morus alba. White mulberry. Native of Asia. Escaped from cultivation and spreading
aggressively. Walnut Lane, Kitchen’s Lane, Livezey Lane, Cresheim Valley, Thomas Mill
Road, Harper’s Meadow.

Morus rubra. Red mulberry. Rich woods. Occasional. Hermit’s Lane, along west bank below
Henry Avenue, near Monastery, Carpenter’s Woods.

CANNABINACEAE, HEMP FAMILY
Humulus japonicus. Japanese hop. A comparatively recent introduction from Asia which is
rapidly becoming a troublesome weed. Hermit’s Lane, Henry Avenue, Kitchen’s Lane,
Cresheim Valley, Harper’s Meadow.
Humulus lupulus. Common hop. Rare and local.

URTICACEAE, NETTLE FAMILY
Boehmeria cylindrica. False nettle. Very abundant in low, moist habitats throughout. Henry

WISSAHICKON VALLEY PLANTS 11

Avenue, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, pase et Valley
Green, Rex Avenue, Thomas Mill Road, Bells Mill Road, Harper’s

Laportea canadensis. Wood nettle. Of frequent occurrence in low secatedl tae paths, etc.
Henry Avenue, Walnut Lane, Kitchens Lane, Mt. Airy Avenue, Livezey Lane, Valley
Green, Thomas Mill Road, Harper’s Meadow.

Parietaria pennsylvanica. Pellitory. Occasional on stone work of bridges, walls, etc.

Pilea pumila. Clearweed. Extremely common in moist ground along drives, bridle paths, etc.
Henry Avenue, Walnut Lane, Carpenter’s Woods, Cresheim Valley, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road.

Urtica gracilis (U. dioica ssp. gracilis). Nettle. Open woods, meadows, and clearings.
Occasional. Hermit’s Lane, Cresheim Valley, Harper’s Meadow.

ARISTOLOCHIACEAE, BIRTHWORT FAMILY
Aristolochia durior (A. macrophylla). Dutchman’s pipe. Rex Avenue, just below Seminole
Street
Asarum canadense. Wild ginger. Woods and shaded slopes. Locally abundant. Henry
Avenue, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Livezey Lane, Valley Green,
Rex Avenue, Thomas Mill Road, above and below Bells Mill Road.

POLYGONACEAE, BUCKWHEAT FAMILY

Polygonum arifolium. Halberd-leaved tearthumb. Moist thickets, infrequent. Walnut Lane.

Polygonum aviculare. Common knotweed. Frequent along trails, foot-paths, clearings, etc.

Polygonum cespitosum var. longisetum. Knotweed. A small, branching annual with spikes
of purple or roseate flowers. Comparatively recent introduction from southeast Asia. It has
spread with phenomenal success and now grows along practically every drive, bridle path,
and foot-trail in the valley. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s
Lane, Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road,
Harper’s Meadow.

Polygonum convolvulus. Black bindweed. An occasional weed in thickets and waste ground.
Cresheim Valley, above Bells Mill Road.

Polygonum cuspidatum. Japanese knotweed. Introduced from east Asia. Tenaciously
established at a number of localities, above Henry Avenue, Walnut Lane, Livezey Lane,
Carpenter’s Woods, Harper’s Meadow.

Polygonum lapathifolium. Dock-leaved smartweed. Wet thickets, damp clearings and fields.
Occasional. Mt. Airy Avenue, near Germantown Avenue.

Polygonum pensylvanicum. Smartweed. Damp thickets, muddy shores, etc. Bluestone Bridge,
Cresheim Valley.

Polygonum persicaria. Lady’s-thumb. A frequent weed in damp clearings, ditches, etc.
Kitchen’s Lane, Mt. Airy Avenue, Cresheim Valley, Harper’s Meadow

Polygonum punctatum. Water smartweed. Abundant in damp hollows, ditches: etc. Henry
Avenue, Walnut Lane, Cresheim Valley, Harper’s Meadow

Polygonum Sagittatum. Arrow-leaved tearthumb. Low, wet diickets: Occasional. Walnut
Lane, Mt. Airy Avenue, Cresheim Valley.

Polygonum scandens. Climbing false buckwheat. Occasional in damp thickets, waste ground,
etc. Walnut Lane, Cresheim Valley.

Polygonum tenue. Knotweed. Dry, open, often rocky slopes. Rare. Not recently found.

Rumex acetosella. Sheep sorrel. A common weed along drives, trails, and in disturbed soil

12 BARTONIA

of picnic areas, etc.
Rumex altissimus. Sorrel. Alluvial shores. West bank above Reading R.R. bridge.

Rumex crispus. Yellow dock. Occasional in clearings, waste ground, and along drives.
Rumex obtusifolius. Bitter dock, great dock. Occasional in low, wet ground or clearings.
Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim Valley.
Tovara virginiana (Polygonum virginianum). Virginia jumpseed. Extremely common on
moist, shaded banks, in bottomland, woods and thickets. Henry Avenue, Bluestone Bridge,
Walnut Lane, Kitchen’s Lane, i s Woods, Cresheim Creek, Valley Green, Rex

Avenue, Bells Mill Road.

CHENOPODIACEAE, GOOSEFOOT FAMILY

Chenopodium album. Lamb’s quarters. An occasional weed in fields, clearings, etc. Flowers
in June or July.

Chenopodium ambrosioides. Mexican-tea. Occasional as a weed in old fields, clearings, etc.

rmit’s Lane, Walnut Lane, Cresheim Valley.

Chenopodium missouriense (C. album var. missouriense). Goosefoot. Resembles C. album
in appearance and distribution, but flowers in August or September. Walnut Lane,
Cresheim Valley.

Chenopodium standleyanum (C. boscianum). Goosefoot. Dry woods. Occasional in lower
portion of valley.

AMARANTHACEAE, AMARANTH FAMILY
Amaranthus hybridus. Pigweed. A weed in clearings, disturbed soil, etc. Walnut Lane.

PHYTOLACCACEAE, POKEWEED FAMILY
Phytolacca americana. Pokeweed. Rich, low ground, clearings, etc. Common. Henry
Avenue, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Mt. Airy Avenue, Livezey
Lane, Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.

AIZOACEAE, CARPETWEED FAMILY
(Molluginaceae)
— verticillata. Carpetweed. Occasional as a weed along paths, in clearings, etc. Henry
venue, Mt. Airy Avenue.

CARYOPHYLLACEAE, PINK FAMILY

Arenaria serpyllifolia. Thyme-leaved sandwort. Dry fields, roadsides, etc. Bells Mill Road.

Cerastium nutans. Nodding mouse-ear chickweed. Rich woods, bottomland. Occasional.

Cerastium vulgatum (C. fontanum ssp. triviale). Common mouse-ear chickweed. Frequent
along trails, in clearings, etc. Cresheim Creek.

Dianthus armeria. Deptford pink. Frequent in grassy clearings and along drives. Valley
Green, Rex Avenue.

Paronychia canadensis. Forked chickweed. Occasional on dry, rocky slopes in upper portion
of valley. Thorps Lane.

Paronychia fastigiata. Whitlow-wort. Same habitat as P. canadensis. Rare. Thorps Lane.

Sagina procumbens. Pearlwort. Occasional along paths, in open spots in woods, and along
gravelly shores. Below Henry Avenue.

Saponaria officinalis. Soapwort, bouncing-bet. Clearings and along roadsides. Fairly

WISSAHICKON VALLEY PLANTS 13

frequent. Livezey Lane, Thomas Mill Road, (picnic ground), Bells Mill Road, Harper’s
Meadow.

Silene antirrhina. Sleepy catchfly. Clearings, waste places. Infrequent.

Silene cucubalus (S. vulgaris). Bladder campion. Damp woods and thickets. Not common.

Silene stellata. Starry campion. Woods and clearings. Bluestone Bridge, Thomas Mill Road,

Harper’s Meadow.

Stellaria alsine (S. uliginosa). Chickweed. Springheads, wet banks, etc. Occasional. Thomas
Mill Roa

Stellaria dintiniton. Common stitchwort. Grassland and clearings. Infrequent.

Stellaria Seid peer Long-leaved stitchwort. Deep thickets, meadows and clearings.
Infreque

Stellaria aie Common chickweed. Common along trails, in damp thickets, and cleared
ground. Henry Avenue, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Rex Avenue,
Thomas Mill Road.

Stellaria pubera. Great chickweed. Occasional in rich woods. Henry Avenue, Bluestone
Bridge, Kitchen’s Lane, Mt. Airy Avenue, Bells Mill Road.

PORTULACACEAE, PORTULACA FAMILY
Claytonia virginica. Spring beauty. Common throughout in rich woods, thickets,
clearings. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt.
Avenue, Cresheim Creek, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill
oad.
Portulaca oleraceae. Purslane. Cosmopolitan. Occasional as a weed along trails and in picnic
areas. Henry Avenue.

RANUNCULACEAE, BUTTERCUP FAMILY

Actaea hesohaeeandi (A. alba). White baneberry, doll’s-eyes. Rich woods. Infrequent. Below
Valley Green, above Rex Avenue.

Anemone quinquefolia. Wood anemone. Apparently rare and local. Above Bells Mill Road.

Anemone virginiana. Thimbleweed. Dry, open woods and thickets. Infrequent. Walnut Lane,
Hartwell Avenue

Anemonella thalictroides (Thalictrum thalictroides). Rue anemone. Rich to dry woods and
Shaded slopes. Fairly abundant throughout.

Aquilegia canadensis. Columbine. Less frequent than formerly, but still found on rocky
ledges, cliffs, etc. Thorps Lane.

Cimicifuga racemosa. Black cohosh. Rich woods. Locally abundant as at Henry Avenue,
Walnut Lane, Bells Mill Road, Rex Avenue.

Clematis paniculata (C. terniflora). Virgin’s bower. Occasionally escaped from cultivation.
Hermit’s Lane, Walnut Lane, Cresheim Valley.

Hepatica americana (H. nobilis var. obtusa). Liverleaf. Dry woods and thickets. Not
common. Below Rex Avenue, Thomas Mill Road.

Hydrastis canadensis. Goldenseal. Collected in lower Wissahickon around 1910. Not seen
in recent years.

Ranunculus abortivus. Kidney-leaved buttercup. Abundant in early spring in rich woods
throughout. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Carpenter’s
Woods, Cresheim Creek, Mt. Airy Avenue, Valley Green, Thomas Mill Road, Bells Mill
Road.

14 BARTONIA

Ranunculus acris. Tall buttercup. Occasional in clearings, along trails, etc. Not seen in recent
years.

Ranunculus bulbosus. Bulbous buttercup. Frequent along trails, in grassy clearings, etc.
Thomas Mill Road.

Ranunculus ficaria. Lesser celandine. Established in several localities. Most of ours is the
double-flowered variety. — Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy
Avenue, Rex Avenue, Thomas oad.

Ranunculus hispidus. Hairy feat Rich woods. Occasional. Walnut Lane.

Ranunculus recurvatus. Buttercup. Rich woods. Occasional. Kitchen’s Lane, Valley Green,
Rex Avenue, Bells Mill Road.

Thalictrum dioicum. Early rue. Scattered throughout in woods and thickets.

Thalictrum polygamum (T. pubescens). Meadow rue. Meadows and moist thickets.

Occasional. Cresheim Valley, Rex Avenue, Valley Green.

MAGNOLIACEAE, MAGNOLIA FAMILY

Liriodendron tulipifera. Tuliptree. Common throughout. Fine specimens occur near Henry
Avenue, Walnut Lane, Cresheim Valley, Valley Green, Rex Avenue, Bells Mill Road.

Magnolia acuminata. Cucumber-tree. Planted in a number of places and apparently
occasionally established and spreading, e.g: above Rex Avenue.

Magnolia tripetela. Umbrella tree. Planted throughout and occasionally spontaneous.
Hermit’s Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim Creek.

Magnolia virginiana. Swamp magnolia. Probably planted, but appearing as though native.
Carpenter’s Woods.

BERBERIDACEAE, BARBERRY FAMILY

Berberis thunbergii. Japanese barberry. The common Japanese barberry, much planted for
hedges, is a frequent escape to wooded slopes along the valley. Henry Avenue, Mt. Airy
Avenue, Cresheim Valley.

Caulophyllum thalictroides. Blue cohosh. Occasional in rich woods. Thomas Mill Road.

Podophyllum peltatum. Mayapple. Abundant throughout in woods, thickets, clearings, etc.

enry Avenue, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Valley Green, Rex

Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

LARDIZABALACEAE, LARDIZABALA FAMILY
Akebia quinata. Five-leaf akebia. Occasionally escaped from cultivation. Rex Avenue, just
below Seminole Street.

LAURACEAE, LAUREL FAMILY
Lindera benzoin. Spicebush. Common everywhere in woods, thickets, on rich slopes, and in
bottomlands. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Valley
Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.
Sassafras albidum. Sassafras. Frequent in dry woods and thickets. Bluestone Bridge, Walnut
ane, Carpenter’s Woods, Mt. Airy Avenue, Cresheim Valley, Valley Green, Thomas Mill
Road, Bells Mill Road.

FUMARIACEAE, FUMITORY FAMILY
Corydalis flavula. Yellow fumewort. Rocky slopes. Rare. Upper portion of valley.

WISSAHICKON VALLEY PLANTS 15

Dicentra cucullaria. Dutchman’s breeches. Rich woods. Rare. Thomas Mill Road, Mt. Airy
Avenue.

PAPAVERACEAE, POPPY FAMILY
Chelidonium majus. Celandine. Naturalized from Europe and common in rich, damp soil
throughout. Walnut Lane, Livezey Lane, Cresheim Valley, Valley Green.
Sanguinaria canadensis. Bloodroot. Apparently infrequent. More abundant in upper portion
of valley. Mt. Airy Avenue, Rex Avenue, Thomas Mill Road, Bells Mill Road.

CRUCIFERAE, MUSTARD FAMILY
(Brassicaceae)

Alliaria officinalis (A. petiolata). Garlic-mustard. Fairly common throughout valley in low
woods and thickets. Henry Avenue, Bluestone Bridge, Kitchen’s Lane, Livezey Lane,
Cresheim Creek, Valley Green, Thomas Mill Road, Harper’s Meadow.

Arabis canadensis. Sicklepod. Rich woods and rocky banks. Infrequent. Thorp’s Lane.

Arabis laevigata. Smooth Rock-cress, Rich woods and shaded slopes. Rare. Henry Avenue,
Bluestone Bridge, Mt. Airy Avenue, Kitchen’s Lane, Valley Green.

Arabis lyrata. Lyre-leaved rock-cress. Ledges, cliffs, rocky woods. Occasional.

Barbarea vulgaris. Winter cress. This is the plant most commonly known as mustard. It is
frequent throughout the valley in meadows, clearings, and along trails. Henry Avenue,
Kitchen’s Lane, Carpenter’s Woods, Valley Green, Rex Avenue, Bells Mill Road.

Brassica arvensis (Sinapis arvensis). Mustard. Occasional in clearings and disturbed soil.

Capseila bursa-pastoris. Shepherd’s-purse. Frequent as a weed in clearings, waste ground,
etc

Cardamine bulbosa. Spring cress. Occasional in ditches, along streams, about springs, etc.
Henry Avenue, Valley Green, Bells Mill Road.

Cardamine impatiens. Bitter-cress. Bottomland. Rare. Bells Mill Road, Thomas Mill Road.

Cardamine pensylvanica. Wood cress, Pennsylvania bitter-cress. Deep hollows in woods,
shaded ditches, etc. Local. Henry Avenue, Walnut Lane, Mt. Airy Avenue, Livezey Lane,
Valley Green, Rex Avenue, Bells Mill Road.

Dentaria heterophylla (Cardamine heterophylla). Pepper-root, toothwort. Woods and
thickets. Rare. Gorgas Lane.

Dentaria laciniata. (Cardamine concatenata). Cut-leaved toothwort. Rich woods. Scattered
and locally abundant. Henry Avenue, Kitchen’s Lane, Mt. Airy Avenue, Bells Mill
Road

spring.

Hesperis matronalis. Dame’ s-rocket. Established along trails and borders of woods. Both the
purple and white flowered forms occur. Livezey Lane, Rex Avenue, Thomas Mill Road.

Lepidium virginicum. Wild pepper-grass. Dry, open situations, weedy. Henry Avenue,
Carpenter’s Woods, Livezey House, Valley Green.

Rorripa islandica (R. palustris ssp. palustris). Yellow cress. Ditches, wet, seepy slopes.

enry Avenue, Walnut Lane, Livezey Lane, west side below Valley Green.

Rorripa sylvestris. Creeping yellow-cress. Muddy stream banks and damp meadows.
Occasional. Thorps Lane.

Sisymbrium altissimum. Tumble mustard. An occasional weed in waste places.

Sisymbrium officinale var. leiocarpum. Hedge mustard. An occasional weed in waste places.

oad.
Draba verna (Erophila verna). Whitlow-grass. Found sparingly in dry, open places in early
n

16 BARTONIA

Sisymbrium thalianum (Arabidopsis thalianum). Mouse-ear cress. Dry, open slopes and sandy
clearings; early spring.

CRASSULACEAE, STONE-CROP FAMILY
Sedum sarmentosum. Live-for-ever. Sparingly escaped from cultivation, below Walnut Lane.

(Saxifragaceae)
Penthorum sediodes. Ditch stonecrop. Ditches and wet, muddy shores. Infrequent. Above
Walnut Lane.

SAXIFRAGACEAE, SAXIFRAGE FAMILY
Chrysosplenium americanum. Golden saxifrage. Ditches, springheads, streams. Apparently
uncommon. Below Bells Mill Road.
Heuchera americana. Alum-root. Wooded, often rocky slopes. Occasional.
Mitella diphylla. Miterwort. Rich to dry woods and thickets. Infrequent. Thomas Mill Road.
Saxifraga pensylvanica. Swamp saxifrage. Wet slopes, boggy thickets. Infrequent. In upper
valley.

Saxifraga virginiensis. Early saxifrage. Dry to moist, rocky woods or shady slopes. Locally
abundant. Henry Avenue, Bluestone Bridge.

(Hydrangeaceae)

Deutzia scabra. Deutzia. Cultivated and occasionally escaped to thickets, stream banks, etc.
Cresheim Creek, Thomas Mill Road.

Hydrangea paniculata. Panicled hydrangea. Native of eastern Asia. Occasionally escaped to
thickets. Walnut Lane, Cresheim Valley.

pga quercifolia. Oak-leaved hydrangea. Native of southeastern states. Cultivated and

aringly escaped. Cresheim Valley.

Phitadelphus coronarius. Mock-orange. Occasional as an escape from cultivation. Henry

Avenue, Mt. Airy Avenue, Livezey Lane, Harper’s Meadow.

HAMAMELIDACEAE, WITCH-HAZEL FAMILY
Hamamelis virginiana. Fall-blooming witch-hazel. Common throughout. Bluestone Bridge,
Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim Valley, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.
Liquidambar styraciflua. Sweet-gum. A tree of the Atlantic Coastal Plain, planted for its
star-shaped leaves and brilliant autumn color. Hermit’s Lane, Paper Mill Run.

PLATANACEAE, PLANE-TREE FAMILY
Platanus occidentalis. Buttonwood, sycamore. Common throughout. Henry Avenue,
Bluestone Bridge, Walnut Lane, Mt. Airy Avenue, Cresheim Valley, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road.

ROSACEAE, ROSE FAMILY
Agrimonia gryposepala. Agrimony. Thickets and clearings. Frequent. Kitchen’s Lane,
Cresheim Valley, Harper’s Meadow.
Agrimonia parviflora. Agrimony. Damp thickets, meadows, clearings. Common. Mt. Airy
Avenue, Cresheim Valley, Bells Mill Road, Harper’s Meadow

WISSAHICKON VALLEY PLANTS iy

Agrimonia pubescens (A. mollis). Downy agrimony. Woods and thickets. Occasional. Below
Devil’s Pool.

Amelanchier arborea. Service-berry. Infrequent on dry, wooded slopes. Kitchen’s Lane,
Hartwell Lane.

Crataegus. Hawthorn. A number of species of Crataegus have been found in the valley.
However, this genus is extremely difficult, and no attempt is made here to differentiate
the various kinds.

Duchesnea indica. Indian strawberry. Grassy clearings, waste ground, etc. Bluestone Bridge,
Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim Valley, Bells Mill Road,
Thorps Lane, Harper’s Meadow.

Fragaria virginiana. Wild strawberry. Occasional on open slopes and in grassy clearings.

Geum canadense. Avens. Thickets and borders of woods. Cresheim Valley, Valley Green,
Hartwell Avenue, hae Avenue, Bells Mill Road, Harper’s Meadow

Gillenia trifoliata (Poteranthus trifoliatus). Bowman’ s-root. Collected in the valley around
1910 near Thorps Lane. Not seen in recent years. Should be sought for.

Physocarpus opulifolius. Ninebark. Thickets. Cresheim Valley, Hartwell Avenue.

Potentilla canadensis. Cinquefoil. Dry, open ground. Frequent. Henry Avenue, Bluestone
Bridge, Walnut Lane, Mt. Airy Avenue, Bells Mill Road.

Potentilla norvegica. Three-leaf cinquefoil. Clearings and waste ground. Fairly frequent.

Potentilla recta. Sulfur cinquefoil. Dry fields and roadsides. Occasional.

Potentilla simplex. Old-field cinquefoil. Dry fields, clearings, roadsides. Fairly common
throughout,especially along West Drive. Henry Avenue, Carpenter’s Woods, Valley Green.

Prunus avium. Bird cherry, sweet cherry. Occasional as an escape. Hermit’s Lane, Thomas
Mill Road.

Prunus persica. Peach. Occasional as an escape.

Prunus serotina. Wild black cherry. Fairly common throughout. Hermit’s Lane, Henry
Avenue, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Harper’s Meadow.

Pyrus arbutifolia (Aronia arbutifolia). Red-fruited chokeberry. Occasional on dry slopes and
in clearings.

Pyrus communis. Pear. Occasional as an escape.

Pyrus malus (Malus pumila). Apple. Occasional as an escape.

Pyrus melanocarpa engi melanocarpa). ila tried chokeberry. Dry clearings and
thickets. Occasional.

Rhodotypus soabada (R. scandens). Jetbead. Escape from cultivation. Carpenter’s Woods,
Cresheim Valley, Hartwell Avenue, Valley Green, Rex Avenue, Bells Mill Road.

Rosa carolina. Pasture Rose. Occasional in low thickets, borders of woods, etc. Springfield
Avenue.

Rubus allegheniensis. Blackberry. Frequent in clearings, borders of woods, etc. Hermit’s
Lane, Bluestone Bridge, Kitchen’s Lane, Devil’s Pool, Cresheim Valley, Harper’s
Meadow

Rubus baileyanus (R. flagellaris). Northern dewberry, trailing blackberry. Occasional in
grassy clearings, open slopes, etc.

Rubus hispidus. Trailing raspberry. Thickets and damp ground. Walnut L

Rubus laciniatus. Cut-leaved blackberry. Introduced from the Old World. ies Bluestone
Bridge, Mt. Airy Avenue, Thomas Mill Road.

Rubus occidentalis. Black raspberry. Fairly frequent in thickets and on dry ve aes Hermit’s
Lane, Cresheim Valley, Thomas Mill Road, Bells Mill Road, Harper’s Meadow

18 BARTONIA

Rubus odoratus. Purple-flowered raspberry. Scattered throughout. Thomas Mill Road.

Rubus phoenicolasius. Wineberry. An introduction from Asia. Fairly c on. Henry
Avenue, Walnut Lane, Carpenter’s Woods, Livezey Lane, Cresheim Valley, Rex sheet
Bells Mill Road, Harper’s Meadow.

Spiraea latifolia. Meadow-sweet. Occasional on dry slopes.

LEGUMINOSAE, PEA FAMILY
(Caesalpiniaceae)
Cassia hebecarpa (Senna hebacarpa). Wild senna. Alluvial thickets. Rare. South of Rex
Avenue, Thorps Lane
Cassia _ nictitans (Cheemciiea tise nictitans). Wild sensitive-plant. Dry, sandy soil. Rare.
Thomas Mill Road.

Cercis canadensis. Redbud, judas tree. Although originally native to the valley, most of the
specimens seen today are the result of recent plantings. Hermit’s Lane, Cresheim Creek,
Rex Avenue, Thomas Mill Road.

Gleditsia triacanthos. Honey-locust. Planted here and there throughout the valley and
occasionally naturalized.

Gymnocladus dioica. —— coffee-tree. Native further south; possibly escaped from
cultivation. Thomas Mill Roa

(Fabaceae)

Amorpha fruticosa. Lead plant, false indigo. Established at several localities. Hermit’s Lane,
Henry Avenue, Bluestone Bridge, Livezey house, Thomas Mill Road, Harper’s Meadow.

Amphicarpa bracteata, Hog-peanut. Abundant in thickets and trailing over low vegetation.
Bluestone Bridge, Walnut Lane, Mt. Airy Avenue, Livezey Lane, Cresheim Valley, Rex
Avenue, Bells Mill Road, Harper’s Meadow. Variety comosa, with more hairy stems, is
known from Thorps Lane.

Crotalaria sagittalis. Rattlebox. Occasional in dry, sandy soil.

Desmodium canadense. Beggar’s-tick. Thickets and dry, open ground. Occasional. Cresheim
Valley, Thomas Mill Road, Bells Mill Road.

Desmodium canescens. Tick-trefoil. Dry clearings and meadows. Occasional

Desmodium dillenii (D. perplexum). Tick-trefoil. Borders of dry wdiedes and_ thickets.
Infrequent. Walnut Lane, Cresheim Valley, Bells Mill Road.

Desmodium grandiflorum (D. cuspidatum). Tick-trefoil. Open woods, thickets and clearings.
Occasional.

Desmodium laevigatum. Tick-trefoil, smooth tick-clover. Dry, open situations. Rare.
Springfield Avenue.

Desmodium marilandicum. Tick-trefoil, Maryland tick-trefoil. Dry, open sandy woods and
clearings. Rare. Hartwell Avenue.

Desmodium nudiflorum. Tick-trefoil, naked-flowered tick-trefoil. Clearings and borders of
dry woods. Fairly frequent. Cresheim Valley, Thomas Mill Road, Bells Mill Road.

Desmodium paniculatum. Tick-trefoil. Dry thickets and clearings. Scattered throughout.
Cresheim Valley.

Desmodium —— (D. obtusum). Tick-trefoil. Dry banks and open, sandy places. Rare.
Thomas Mill Roa

Desmodium rotund flim Tick-trefoil. Occasional in open woods, thickets, and clearings.
Hartwell Avenue

WISSAHICKON VALLEY PLANTS 19

Desmodium viridiflorum. Tick-trefoil, velvety tick-trefoil. Borders of woods, thickets, and
ry, open situations. Infrequent. Springfield Avenue.

Lespedeza hirta. Bush-clover. Dry banks and clearings.Infrequent, Thomas Mill Road, Bells
Mill Road.

Lespedeza intermedia. Bush-clover. Dry, open woods and thickets. Occasional. Thorp’s Lane.

Lespedeza procumbens. Bush-clover. Dry woods and clearings. Occasional. Thorp’s Lane.

Lespedeza violacea. Bush-clover. Dry, open ground. Occasional. Below Bells Mill Road.

Medicago lupulina. Black medick. Roadsides, clearings, waste ground.

Medicago sativa. Alfalfa, lucerne. An occasional weed in open ground, along trails, and in
waste places.

Melilotus alba. White melilot. Occasional as a weed along trails and in clearings.

Melilotus officinalis. Yellow melilot. Occasional as a weed along trails and in waste ground.

Robinia pseudo-acacia. Black locust. Although not originally indigenous in this area, Black
locust has been much planted and is now widely scattered. Hermit’s Lane, Walnut Lane,
Cresheim Valley.

Strophostyles helvola. Wild bean. Dry woods and clearings. Rare.

Tephrosia virginiana. Goat’s-rue. Dry woods and clearings. Rare. Hartwell Avenue.

Trifolium agrarium (T. aureum). Yellow clover. Dry slopes and grassy clearings. Occasional.
Bells Mill Road.

Trifolium hybridum. Alsike clover. Grassy clearings, roadsides. Frequent.

Trifolium pratense. Purple clover, red clover. Open fields and meadows. Common. Cresheim
Valley.

Trifolium repens. White clover. Open fields and clearings. Common. Mt Airy Avenue.

Wisteria sinensis. Chinese wisteria. Native of Asia. Occasionally established, mouth of Paper
Mill Run, Bluestone Bridge.

LINACEAE, FLAX FAMILY
Linum virginianum. Flax, slender yellow flax. Dry, open woods, clearings, and slopes.
Apparently rare, seen only along bridle path south of Bells Mill Road.

OXALIDACEAE, WOOD SORREL FAMILY
Oxalis europaea (O. stricta). Yellow wood-sorrel. A common weed of dry, open slopes,
roadsides, clearings, etc. Bluestone Bridge, Walnut Lane, Mt. Airy Avenue, Cresheim
Valley, Rex Avenue.
Oxalis violacea. Violet wood-sorrel. Dry, wooded slopes. Rare. Not recently seen.

GERANIACEAE, CRANE’S BILL FAMILY
Erodium cicutarium. Storksbill, redstem filaree. Fields and waste places. Rare.
Geranium maculatum. Wild geranium, cranesbill. Common throughout in woods, thickets,
and meadows. Kitchen’s Lane, Cresheim Creek, Valley Green.

SIMARUBACEAE, QUASSIA FAMILY
Ailanthus altissima. Tree-of-heaven. Introduced from Asia and naturalized everywhere
throughout the area. Henry Avenue, Walnut lane, Cresheim Valley, Livezey Lane.

EUPHORBIACEAE, SPURGE FAMILY
Acalypha rhombiodea. Three-seeded mercury. Moist hollows, roadsides, etc. Frequent.

20 BARTONIA

Bluestone Bridge, Walnut Lane, Livezey Lane.

Acalypha virginica. Three-seeded mercury. Dry ground. Uncommon.

Euphorbia cyparissias. Cypress spurge. Introduced from Europe and naturalized in fields and
clearings, Livezey Lane, Thomas Mill Road, Thorps Lane, Bells Mill Road, Harper’s
Meadow

Euphorbia maculata (Chamaesyce maculata). Eyebane. Dry, open ground. Occasional.

Euphorbia supina (Chamaesyce maculata). Milk purslane. Roadsides, clearings, etc. Fairly
frequent, especially around parking and picnic areas.

CALLITRICHACEAE, WATER-STARWORT FAMILY
Callitriche stagnalis. Water-chickweed. Occasional in pools and ditches. Henry Avenue,
Walnut Lane, Cresheim Creek.

LIMNANTHACEAE, FALSE MERMAID FAMILY
Floerkea proserpinacoides. False-mermaid. Low, rich woods. Occasional. Early spring.
Kitchen’s Lane, Bells Mill Road, Harper’s Meadow.

ANACARDIACEAE, CASHEW FAMILY
Rhus glabra. Smooth sumac. Dry thickets and borders of woods. Cresheim Valley, Thomas
Mill Road.

Rhus radicans (Toxicodendron radicans). Poison-ivy.Thickets, open woods. Some forms
shrubby, others trailing or highclimbing. Common throughout. Bluestone Bridge, Walnut
Lane, Kitchen’s Lane, Cresheim Valley, Thomas Mill Road, Bells Mill Road.

Rhus typhina. Staghorn sumac. Thickets and dry clearings. Occasional.

CELASTRACEAE, STAFF-TREE FAMILY

Celastrus orbiculatus. Bittersweet. Introduced from Asia; cultivated and spreading rapidly
in our area. Bluestone Bridge, Livezey Lane, Devil’s Pool, Thomas Mill Road. (This
species has rounder leaves than the native bittersweet, which follows).

Celastrus scandens. Bittersweet. Woods, thickets, stream banks. Fairly common. Monastery,
Carpenter’s Woods, Cresheim Valley, Thomas Mill Road, Bells Mill Road.

Euonymus alatus. Spindle-tree. Introduced from Asia and occasional as an escape from
cultivation, as above Walnut Lane, and along Rex Avenue.

Euonymus americanus. Strawberry-bush. Rich woods and thickets. Apparently not common.
Henry Avenue, Cresheim Valley, Rex Avenue, above Thomas Mill Road, Bells Mill Road.

Euonymus europaeus. European spindle-tree. Introduced from Europe and sparingly escaped
from cultivation. Thorps Lane.

STAPHYLACEAE, BLADDERNUT FAMILY
Staphylea trifolia. Rich woods and thickets, mostly in bottomland. Fairly frequent
——— Henry Avenue, Walnut Lane, Thomas Mill Road, Bells Mill Road, Harper’s
Meado

ACERACEAE, MAPLE FAMILY
Acer negundo. Ash-leaved maple, box-elder. Steam banks and low ground. Common
throughout. Mouth, Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane,
Livezey Lane, Cresheim Valley, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

WISSAHICKON VALLEY PLANTS 21

Acer platanoides. Norway maple. Cultivated and occasionally escaped. Livezey Lane,
Cresheim Valley, Harper’s Meadow.

Acer pseudo-platanus. Sycamore maple. Sparingly escaped from cultivation. Walnut Lane,
Cresheim Valley.

Acer rubrum. Red maple, swamp maple. Low, usually moist ground. Fairly frequent
throughout. Walnut Lane, Carpenter’s Woods, Cresheim Valley, Valley Green, Rex
Avenue.

Acer saccharinum. Silver maple. Rich woods of bottomland. Occasional. Kitchen’s Lane,
Cresheim Valley, Harper’s Meadow.

Acer saccharum. Sugar maple. Dry slopes. Occasional. Hermit’s Lane.

Acer spicatum. Mountain maple. Dry, open woods and thickets. Apparently rare. Collected
in the valley in 1924, but no locality given.

SAPINDACEAE, SOAPBERRY FAMILY
Cardiospermum halicababum. Balloon-vine. Native of tropical America. Established around
spring on east bank, just below Bells Mill Road.

HIPPOCASTANACEAE, BUCKEYE FAMILY
Aesculus glabra. Ohio buckeye. Cultivated and occasionally escaped. Henry Avenue,
Carpenter’s Woods, south of Bells Mill Road, Harper’s Meadow.
Aesculus hippocastanum. Horse-chestnut. Native of Europe. Widely planted in our area and
sparingly escaped from cultivation. Near Leverington Street.

BALSAMINACEAE, TOUCH-ME-NOT FAMILY
Impatiens biflora (I. capensis). Orange jewelweed. Low, rich ground. Common throughout.
Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Cresheim Valley, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.
Impatiens pallida. Yellow jewelweed. Low, rich ground. Common throughout. Walnut Lane,
Devil’s Pool, Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill
Road, Harper’s Meadow.

RHAMNACEAE, BUCKTHORN FAMILY
Ceanothus americanus. New Jersey tea. Dry ground. Collected somewhere in the valley in
1889, but not seen in recent years.

VITACEAE, VINE FAMILY

Ampelopsis arborea. Pepper-vine. Swampy woods. Cultivated in our area and sparingly
escaped from gardens. ;

Parthenocissus quinquefolia. Virginia-creeper. Woods and thickets. Frequent. Hermit’s Lane,
Walnut Lane, Cresheim Valley, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill
Road, Harper’s Meadow.

Vitis aestivalis. Summer grape. Dry woods and thickets. Infrequent. Hermit’s Lane, Thomas
Mill Road, below Bells Mill Road.

Vitis labrusca. Fox grape. Woods and thickets. Infrequent. Cresheim Valley.

Vitis riparia. River-bank grape. Thickets and stream banks. Occasional throughout. Bluestone
Bridge, Walnut Lane, Rex Avenue.

Vitis vulpina. Frost grape. Thickets and bottomland. The commonest grape in the valley.

22 BARTONIA

Hermit’s Lane, Henry Avenue, Walnut Lane, Monastery, Mt. Airy Avenue, Cresheim
Valley, Bells Mill Road, Harper’s Meadow.

TILIACEAE, LINDEN FAMILY
Tilia americana. Basswood. Rich woods. Common. Livezey Lane, Cresheim Valley, Thomas
Mill Road, Bells Mill Road, Harper’s Meadow.

MALVACEAE, MALLOW FAMILY
Althaea rosea (Alcea rosea). Hollyhock. Escaped from cultivation and occasionally persistent
in waste ground.
Hibiscus syriacus. Rose-of-sharon. Native of Asia. Cultivated and sparingly escaped.
Carpenter’s Woods.
Malva neglecta (M. rotundifolia). Cheeses. Occasional as a weed near parking lots and picnic
areas.

HYPERICACEAE, ST.-JOHN’S-WORT FAMILY
(Clusiaceae)

Ascyrum hypericoides (Hypericum hypericoides). St.-Andrew’s- cross. Dry sandy soil. Rare;
known only from near Thomas Mill Road.

Hypericum gentianoides. Orange-grass, pineweed. Sandy or rocky slopes. Rare. Hartwell
Avenue.

Hypericum mutilum. St.-John’s-wort. Low, damp ground. Frequent in ditches along drive,
below Valley Green, near Germantown Avenue, Chestnut Hill.

Hypericum perforatum, Common St.-John’s-wort. Dry fields, clearings, etc. Fairly frequent
in Open, grassy situations.

Hypericum punctatum. St.-John’s-wort. Low, usually damp ground. Occasional. Above
Thomas Mill Road, Bells Mill Road.

CISTACEAE, ROCKROSE FAMILY
Helianthemum canadense. Frostweed, rock-rose. Dry, open woods. Rare. Bells Mill Road.
Lechea minor. Pinweed. Dry woods and open slopes. Rare. Springfield Avenue.
Lechea racemulosa. Pinweed. Open, sandy woods and slopes. Rare. Carpenter’s Woods,
Hartwell Avenue.

VIOLACEAE, VIOLET FAMILY
Viola blanda. Sweet white violet. Rich woods, damp thickets, and rocky slopes. Frequent.
Bluestone Bridge, Walnut Lane, Allens Lane.
Viola conspersa. Dog violet. Damp woods, thickets, and shaded slopes. Occasional. Allens

Lane.

Viola cucullata. Swamp blue violet. Springheads, seepy slopes, moist depressions. Frequent.
Cresheim Creek, Mt. Airy Avenue, Valley Green, Rex Avenue, Bells Mill Road.

Viola hirsutula. Hairy blue violet. Moist to dry wooded slopes. Infrequent. Allens Lane.

Viola en (V. blanda). Wet to dry woods, thickets, and clearings. Rare. Near Indian
Roc

Viola pallens (V. a, ee ssp. pallens). Wet or springy woods, thickets, or slopes.
Occasional. Thomas Mill Road.

Viola palmata. Early blue violet. Rich woods and slopes. Infrequent.

WISSAHICKON VALLEY PLANTS 23

Viola papilionacea (V. sororia var. sororia), Common blue violet. Meadows, clearings, open
banks, etc. Fairly common throughout. Henry Avenue, Walnut Lane, Cresheim Valley.
Viola pedata. Birds-foot violet. Dry, sterile soil. Rare, known only from along Bells Mill
Road, west of the creek. Ours is the variety lineariloba with all the petals lilac-purple.

Bells Mill Road.

Viola pensylvanica (V. eriocarpa). Smooth yellow violet. Rich woods and thickets. Common
throughout. Henry Avenue, Bluestone Bridge, Kitchen’s Lane, Bells Mill Road.

Viola primulifolia. Primrose-leaved white violet. Dry clearings. Carpenter’s Woods.

Viola pubescens. Hairy yellow violet. Rich woods and thickets. Somewhat restricted. Henry
Avenue.

Viola rotundifolia. Early yellow violet, round-leaved yellow violet. Extremely common on
moist to dry, wooded slopes, especially under hemlock. Walnut Lane, Mt. Airy Avenue,
Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.

Viola sagittata var. ovata (V. fimbriatula). Arrow-leaved violet. Dry woods and clearings.
Occasional. Bells Mill Road.

Viola sororia. Common blue violet. Open woods, moist slopes. Frequent throughout. Henry
Avenue, Walnut Lane, Rex Avenue.

Viola striata. Striped white violet. Rich woods and thickets. Occasional. Cresheim Creek.

Viola triloba (V. palmata). (Early blue violet). Open thickets, clearings, dry slopes, etc.
Infrequent. Near Devil’s Pool.

ONAGRACEAE, EVENING-PRIMROSE FAMILY

Circaea quadrisulcata var. canadensis. (C. lutetiana ssp. canadensis). Enchanter’s-
nightshade. Rich woods, thickets, and bottomland, extremely common throughout. Native.
Henry Avenue, Bluestone Bridge, Walnut Lane, Livezey Lane, Cresheim Valley, Valley
Green, Rex Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Epilobium angustifolium. Great willow-herb. fireweed. Recent clearings and waste grounds.
Infrequent. Hartwell Avenue.

Epilobium coloratum. Tawny willow-herb. Low, damp ground. Rare.

Epilobium densum (E. strictum). Willow-herb. Occasional on sandy and gravelly shores.

Gaura biennis. Biennial velvet-weed. Thickets, bottomlands, damp shores. Rare. Hartwell
Avenue.

Ludwigia alternifolia. Seed-box. Low, damp ground, muddy shores. Infrequent. Cresheim
Valley, Bells Mill Road.

Ludwigia palustris. Water-purslane. Wet, muddy, and gravelly shores. Common. Mouth of
creek, Henry Avenue, Walnut Lane, Mt. Airy Avenue, Livezey Lane, Thomas Mill Road.

Oenothera biennis. Common evening-primrose. Meadows, clearings, waste ground, disturbed
soils. Occasional. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Valley

Green, Harper’s Meadow
Ocenothera perennis (O. pumila). Sundrops. Damp to dry open ground. Infrequent.

ARALIACEAE, GINSENG FAMILY
Acanthopanax sieboldianus. Acanthopanax. Introduced from Asia and occasionally escaped
from cultivation, as near Harper’s Meadow.
Aralia chinensis. Chinese angelica or Hercules’-club. This large Asiatic shrub with prickly
stems and leaves is established and apparently spreading in the valley. It is especially
common near Walnut Lane Bridge, but occurs also near near Bluestone Bridge, near

24 BARTONIA

Valley Green, at Thomas Mill Road.
Aralia nudicaulis. Wild sarsparilla. Dry woods and shady slopes. Scattered throughout.
enry Avenue, Walnut Lane, Kitchen’s Lane, Cresheim Valley, Rex Avenue, Thomas
Mill Road.

Aralia racemosa. Spikenard. Rich woods and damp slopes. Very abundant at foot of wet
cliffs along Drive between Valley Green and Thomas Mill Road. Also Bluestone Bridge,
Walnut Lane, Kitchen’s Lane, Livezey lane, Cresheim Valley, Bells Mill Road.

Aralia spinosa, Hercules’-club. This is the native American species of Hercules’-club which
has leaves less prickly than A. chinensis. It is scattered locally in the upper valley.
Hermit’s Lane, Kitchen’s Lane, Thomas Mill Road.

Hedera helix. English Ivy. Abundantly planted on dwellings and stonework, and occasionally
spreading in shaded situations, often growing on tree trunks. Henry Avenue, Walnut Lane,
Cresheim Creek.

Panax trifolius. Dwarf American ginseng. Rich woods. Rare and local. Henry Avenue,
Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, ravine above Thomas Mill Road.

UMBELLIFERAE, PARSLEY FAMILY
(Apiaceae)

Aegopodium podagraria. Goutweed. Naturalized from Europe and abundantly established at
a number of localities. Along drive below Henry Avenue, Kitchen’s Lane, Mt. Airy
Avenue, Cresheim Creek, Valley Green, Thomas Mill Road.

Cicuta maculata. Spotted cowbane, water-hemlock. Swales, bottomlands. Occasional.
Cresheim Vall

Cryptotaenia canadensis. Honewort, wild chervil. Dominant in damp ground along drives
and trails throughout. Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Cresheim Valley,
Valley Green, Rex Avenue, Bells Mill Road, Harper’s Meadow.

Daucus carota. Wild carrot, Queen-Anne’s-lace. Meadows, grassy clearings, etc. Locally
abundant throughout. Livezey Lane, Cresheim Valley.

Heracleum maximum (H. lanatum). Cow-parsnip. Swales and low, damp woods. Occasional.
Walnut Lane, Valley Green, near Harper’s Meadow.

Hydrocotyle americana. Water penny-wort. Damp woods and wet depressions. Occasional
in ditches, e.g: below Bells Mill Road, Wise’s Mill Road.

Osmorhiza_ claytonii. Sweet-cicely, wild-licorice. Woods and shaded slopes. Frequent
throughout. Cresheim Valley, Thomas Mill Road.

Osmorhiza longistylis. Anise-root. Rich woods and alluvial thickets. Less abundant than O.
claytoni. Bluestone Bridge, Kitchen’s Lane, Valley Green, Bells Mill Road.

Oxypolis rigidior. Cowbane. Wet woods and swamps. Rare. Thorp’s Lane, near Harper’s
Meadow.

Pastinaca sativa. Parsnip. Clearings and waste ground. Sporadic. Near Harper’s Meadow.

Sanicula canadensis. Black snake-root. Dry, open woods and shaded slopes. Abundant
throughout. Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim
Valley, Thomas Mill Road, Harper’s Meadow.

Sanicula gregaria (S. odorata). Black snake-root. Rich woods. Infrequent. Mt. Airy Avenue.

Sanicula trifoliata, Black snake-root. Occasional in rich woods. Walnut Lane, Mt. Airy
Avenue, Gorgas Lane, Valley Green.

Thaspium barbinode. Meadow-parsnip. Abundant in rich woods and thickets. Walnut Lane,
Kitchen’s Lane, Rex Avenue, Thomas Mill Road, Bells Mill Road.

WISSAHICKON VALLEY PLANTS 25
Thaspium trifoliatum. Meadow-parsnip. Woods and shaded slopes. Rare. Thorps Lane.

CORNACEAE, DOGWOOD FAMILY

Cornus alternifolia. Alternate-leaved dogwood. Dry woods and rocky slopes. Rare and local.
Walnut Lane, Allens Lane, Valley Green.

Cornus amomum. Silky cornel. Moist to dry woods and thickets. Infrequent. Carpenter’s
Woods, Livezey Lane, Cresheim Valley, Harper’s Meadow.

Cornus florida. Common dogwood. Both planted and native. Frequent. Walnut Lane,
Carpenter’s Woods, Thomas Mill Road, Bells Mill Road.

Cornus racemosa. Panicled dogwood. Woods and thickets. Occasional. Henry Avenue,
Kitchen’s Lane.

(Nyssaceae)
Nyssa sylvatica. Sour-gum. Moist to dry woods. Fairly common. Hermit’s Lane, Walnut
Lane, Carpenter’s Woods, Livezey Lane, Valley Green, Thomas Mill Road, Bells Mill
oad.

PYROLACEAE, WINTERGREEN FAMILY
Chimaphila maculata. Striped pipsissewa, spotted wintergreen. Dry, usually acid woods and
slopes. Rare and local. Carpenter’s Woods, Valley Green, Bells Mill Road.
Pyrola elliptica. Shinleaf. Dry to moist woods. Occasional. Thorps Lane.
Pyrola secunda (Orthilia secunda). One-sided pyrola. Dry to moist woods. Rare.
Pyrola virens (P. chlorantha). Wintergreen. Dry, open woods and clearings. Rare.

(Monotropaceae)
Monotropa hypopithys. Pine-sap, false beechdrops. Woodland humus. Infrequent and
seasonal. Rex Avenue.
Monotropa uniflora. Indian-pipe. Woodland humus. Occasional. South of Rex Avenue,
Walnut Lane.

ERICACEAE, HEATH FAMILY

Epigaea repens. Trailing arbutus. Dry, usually acid woods and rocky slopes. Less common
than formerly, but still locally abundant, especially near the crests of the Valley. Walnut
Lane, Carpenter’s Woods, Cresheim Valley, Valley Green, Bells Mill Road.

Gaultheria procumbens. Wintergreen, teaberry, checkerberry. Dry woods and clearings.
Collected many years ago, with no locality given. Probably no longer present.

Gaylussacia baccata. Black huckleberry. Dry woods and thickets. Infrequent. Carpenter’s
Woods, Bells Mill Road.

Gaylussacia frondosa. Dangleberry. Dry woods, shaded slopes, and clearings. Infrequent.

Kalmia angustifolia. Sheep laurel, lambkill. Dry or moist, usually acid soil. Rare. Hartwell
Avenue.

Kalmia latifolia. Mountain laurel, calico bush. Both planted and native. Dry, acid woods and
Slopes throughout. Walnut Lane, Carpenter’s Woods, Hartwell Avenue, Rex Avenue,
below Bells Mill Road.

Lyonia ligustrina. Maleberry. Dry thickets and clearings. Rare. Cresheim Valley.

Lyonia mariana. Stagger-bush. Dry, sandy soil. Rare. Hartwell Avenue.

26 BARTONIA

peep maximum. Great-laurel, rosebay. Widely planted throughout the valley, but

as a native.

Sigedeareae: nudiflorum (R. poriciymensides). Pinxter-flower. Dry woods and shaded
slopes. Locally abundant. Carpenter’s Woods, Livezey Lane, Cresheim Valley, Valley
Green, Bells Mill Road.

Vaccinium angustifolium. Lowbush blueberry. Dry, sterile slopes. Infrequent. Thomas Mill
Road.

Vaccinium corymbosum. Highbush blueberry. Low woods and moist thickets. Occasional.
Carpenter’s Woods, Cresheim Valley, Bells Mill Road.

Vaccinium stamineum. Deerberry, squaw-huckleberry. Dry woods, thickets, and clearings.
Rare. Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Bells Mill Road

Vaccinium vacillans (V. pallidum). Early sweet blueberry. Dry, open woods, thickets, and
clearings. Rare. Hermit’s Lane, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods, Valley
Green, Thomas Mill Road, Bells Mill Road.

PRIMULACEAE, PRIMROSE FAMILY
Lysimachia ciliata. Fringed loosestrife. Low, moist ground. Occasional. Bluestone Bridge,
Livezey Lane.
Lysimachia nummularia. Moneywort. Damp roadsides, shores, and springheads. Fairly
common throughout. Livezey Lane.
Lysimachia quadrifolia. Whorled loosestrife. Dry, open woods, thickets, and clearings.
Frequent throughout, especially on upper slopes. Walnut Lane, Mt. Airy Avenue.

OLEACEAE, OLIVE FAMILY

Fraxinus americana. White ash. Rich woods. Common throughout. Henry Avenue, Walnut
Lane, Kitchen’s Lane, Carpenter’s Woods, Cresheim Valley, Rex Avenue, Thomas Mill
Road, Bells Mill Road. |

Fraxinus pennsylvanica. Green ash. Rich woods. Rare. Collected in the 1920’s, no locality
given.

Ligustrum obtusifolium. Japanese privet, Ibota privet. Native of Asia. Frequent as an escape.
Bluestone Bridge, Rex Avenue, Bells Mill Road, Harper’s Meadow.

Ligustrum vulgare. Common privet. Native of Europe. Frequent as an escape. Walnut Lane,
Cresheim Valley, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

GENTIANACEAE, GENTIAN FAMILY
Bartonia virginica. Bartonia. Wet, acid hollows. Rare. Hartwell Avenue.
Gentiana clausa. Closed gentian, bottle gentian. Wet thickets, stream banks. Infrequent.
Livezey Lane.
Gentiana crinita (Gentianopsis crinata). Fringed Gentian. Meadows, open slopes, etc.
Collected in the valley many years ago. Not seen recently.

APOCYNACEAE, DOGBANE FAMILY
Apocynum androsaemifolium. Spreading dogbane. Dry, open thickets and clearings.
Occasional. Near Harper’s Meadow.
Apocynum cannabinum. Indian Hemp. Thickets and open ground. Infrequent. Walnut Lane.
Vinca minor. Common periwinkle, myrtle. Native of Europe. Widely established in borders
of woods, roadsides, etc. Hermit’s Lane, Henry Avenue, Cresheim Valley, Thomas Mill

WISSAHICKON VALLEY PLANTS a7
Road, Bells Mill Road.

ASCLEPIADACEAE, MILKWEED FAMILY

Asclepias exaltata. Poke milkweed. Occasional in rich woods and clearings. Thorps Lane.

Asclepias incarnata. Swamp milkweed. Swamps and wet thickets. Occasional. Walnut
Lane, Kitchen’s Lane, Thorps Lane.

Asclepias quadrifolia. Whorled milkweed. Woods and thickets. Occasional.

Asclepias syriaca. Common milkweed. Thickets, open banks, clearings, etc. Scattered
throughout. Henry Avenue, Cresheim Valley, Thomas Mill Road.

Cynanchum nigrum (Vincetoxicum nigrum). Black swallow-wort. This twining vine, with
milkweed-like flowers was collected in the early 1900’s in Cresheim Valley near
Mermaid Lane. Not seen in recent years.

Gonolobus obliquus (Matelea obliqua). Angle-pod. Thickets and borders of woods. Rare.

CONVOLVULACEAE, MORNING GLORY FAMILY
Convolvulus sepium (Calystegium silvatica ssp. fraterniflora). je bindweed. Thickets,
open banks, waste ground, etc. Sporadic throughout. Walnut Lan
Ipomoea hederaceae. Morning-glory. Thickets and clearings. Occhnional Bells Mill Road.
Ipomoea pandurata. Wild potato-vine. Dry, open ground. Rare. Above Walnut Lane,
Hartwell Avenue.

(Cuscutaceae)
Cuscuta gronovii. Dodder, love vine. Parasitic on herbaceous vegetation. Occasional
throughout. Henry Avenue, Kitchen’s Lane, Cresheim Valley, Valley Green, Livezey
Lane, Harper’s Meadow.

POLEMONIACEAE, PHLOX FAMILY
Phlox paniculata. Garden phlox. A frequent escape in the valley. Harper’s Meadow.
Polemonium reptans. Jacob’s-ladder. Rich woods and bottomland. Frequent. Walnut Lane,
Kitchen’s Lane, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road.

HYDROPHYLLACEAE, WATER-LEAF FAMILY
Hydrophyllum virginianum. Water-leaf. Rich woods and bottomland. Occasional. Henry
Avenue, Walnut Lane, Bluestone Bridge, Kitchen’s Lane, Mt. Airy Avenue, Livezey Lane,
Valley Green, Thomas Mill Road, above Bells Mill Road, Harper’s Meadow.

BORAGINACEAE, BORAGE FAMIL
Hackelia virginiana (Lappula virginiana). Stickseed, vale s-lice. Woods and thickets.
Occasional. Thomas Mill Road, Thorps Lane.
Lithospermum arvense (Buglossoides arvense). Corn gromwell. An occasional weed of fields
and roadsides
Mertensia virginica. Bluebells, Virginia cowslip. Both native and planted. Occasional.
s Meadow.

Myosotis laxa. Forget-me-not. Moist to dry open ground. Infrequent. Cresheim Creek.

Myosotis scorpioides. True forget-me-not. Stream borders and ditches. Occasional
throughout. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Livezey
Lane, Rex Avenue, Harper’s Meadow.

28 BARTONIA

PHRYMACEAE, LOPSEED FAMILY
(Verbenaceae)
Phryma leptostachya. Lopseed. Rich woods. Fairly frequent throughout. Above Thorps Lane.

VERBENACEAE, VERVAIN FAMILY
Verbena urticifolia. White vervain. Thickets and borders of woods. Frequent. Henry Avenue,
Walnut Lane, Livezey Lane, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

LABIATAE, MINT FAMILY
(Lamiaceae)
Agastache nepetoides. Giant hyssop. Rich thickets and borders of woods.Infrequent. Thorps
L

ane.

Agastache scrophulariifolia. Giant hyssop. Rich woods and thickets. Infrequent. Collected
in 1874, not recently seen.

Collinsonia canadensis. Horse-balm, richweed, stoneroot. Rich woods and _ thickets.
Common. Henry Avenue, Bluestone Bridge, Walnut Lane, Mt. Airy Avenue, Livezey
Lane, Cresheim Creek, Rex Avenue, Thomas Mill Road.

Cunila_ origanoides. American dittany, stone-mint. Dry, open woods and _ clearings.
Infrequent. Thorps Lane.

Hedeoma pulegioides. Mock pennyroyal. Dry woods and thickets. Fairly frequent.

Lamium amplexicaule. Dead-nettle. Occasional as a weed in clearings and waste ground.

Lamium pupureum. Purple dead-nettle. Occasional as a roadside weed. Cresheim Creek,
Harper’s Meadow.

Leonurus cardiaca. Motherwort. An occasional weed near dwellings. Livezey Lane,
Cresheim Valley.

Lycopus americanus. Water-horehound. Low, wet ground. Sporadic. Below Henry Avenue.

Lycopus uniflorus. Water-horehound. Low ground, ditches, etc. Fairly frequent. Bluestone
Bridge, Walnut Lane, Rex Avenue, Bells Mill Road.

Lycopus virginicus. Water-horehound. Rich, moist soil. Occasional. Valley Green.

Mentha arvensis. Common mint. Damp, open situations. Occasional. Walnut Lane, Thorps

ane.

Mentha piperita. Peppermint. Low, wet ground. Infrequent. mae Mill Road.

Mentha spicata. Spearmint. Wet depressions and ditches. R:

Monarda clinopodia, Wild bergamot. Moist thickets and oe banks. Rare. Walnut Lane,

orps Lane.

Monarda didyma, Oswego-tea, bee-balm. Rich woods and bottomland. Rare. Not seen in
recent years.

Monarda fistulosa, Wild bergamot. Dry woods and thickets. Occasional. Leverington Street,
Bells Mill Road.

Nene cataria. Catnip. Occasional as a weed of roadsides and waste places. Bells Mill
Road.

Nepeta hederacea (Glechoma hederacea). Ground ivy. Gill-over-the-ground. Roadsides and
damp, shaded places. Common. Henry Avenue, Bluestone Bridge, Walnut Lane,
Kitchen’s Lane, Livezey Lane, Valley Green, Rex Avenue, Bells Mill Road, Harper’s
Meadow.

Origanum vulgare. Wild marjoram. Roadsides, old fields, and clearings. Rare. Allens Lane.

Perilla frutescens. Perilla. Introduced from Asia. Clearings and alluvial shores. Infrequent.

WISSAHICKON VALLEY PLANTS 29

Henry Avenue.

Prunella vulgaris. Self-heal, heal-all. Open woods, thickets, roadsides, grassy places, etc.
Ours is mostly the variety /anceolata , with leaves about twice as long as broad. Frequent
throughout. Bluestone Bridge, Kitchen’s Lane, Cresheim Valley, Valley Green, Thomas
Mill Road, Harper’s Meadow.

Pycnanthemum verticillatum. Mountain mint. Dry to moist thickets and clearings.
Occasional.

Pycnanthemum virginianum. Mountain mint. Dry to wet thickets, meadows, etc. Occasional.

Salvia lyrata. Lyre-leaved sage. Open woods, clearings, roadsides. Infrequent. Bluestone

ridge.
Satureja vulgaris (Clinopodium vulgare). Basil. Dry, open fields, clearings, disturbed ground,
etc. Occasional around parking and picnic areas.
Scutellaria elliptica, Hairy skullcap. Dry woods and thickets. Rare. Thorps Lane.
Teucrium canadense var. virginicum. Wood-sage. Damp thickets and meadows. Rare.
Trichostema dichotomum. Bluecurls. Dry, open soil. Occasional. Thomas Mill Road.

SOLANACEAE, NIGHTSHADE FAMILY

Lycium halimifolium (L. barbarum). Matrimony vine. Occasional as an escape to recent
clearings and banks. Common between Henry Avenue and Reading R R. bridge, Cresheim
Creek.

Physalis subglabrata. Ground cherry. Fields, roadsides, waste places. Thomas Mill Road,
Harper’s Meadow.

Solanum americanum (S. nigrum). Common nightshade. Roadsides, waste places, etc.
Infrequent. Above Walnut Lane, Mt. Airy Avenue.

Solanum carolinense. Horse-nettle. An obnoxious weed in meadows, clearings, etc.
Occasional. Mouth of creek, Hermit’s Lane, Kitchen’s Lane.

Solanum dulcamara. Climbing nightshade, bittersweet. Thickets and clearings. Occasional.
Henry Avenue, Walnut Lane, Carpenter’s Woods, Livezey Lane, Cresheim Valley, Valley
Green, Rex Avenue.

SCROPHULARIACEAE, FIGWORT FAMILY
Aureolaria pedicularis. Fern-leaved false-foxglove. Rich to dry woods. Rare. Thorps Lane.
Aureolaria virginica. Downy false-foxglove. Dry woods and clearings. Rare. Thorps Lane.
Chelone glabra. Turtle head. Wet thickets, bottomland. Occasional. Bluestone Bridge,
Walnut Lane, Valley Green, Harper’s Meadow.
Cymbalaria muralis. Kenilworth-ivy. Occasional on old walls or the stonework of bridges.
Gerardia tenuifolia (Agalinis tenuifolia). Slender gerardia. Dry, open situations. Rare. Thorps
Lk :

ane.

Gratiola neglecta. Hedge-hyssop. Wet or muddy places. Rare.

Linaria vulgaris. Yellow toadflax, butter-and-eggs. Meadows, roadsides, waste ground.
Occasional throughout.

Lindernia dubia. False pimpernel. Stream shores and damp, muddy places. Rare.

Mazus japonicus (M. pumilus). Japanese mazus. Native of Asia, cultivated in lawns and
gardens, and escaped to grassy fields and wet shores. Livezey Lane.

Mazus_reptans (M. miguelii). Creeping mazus. Sand and gravel shores, escaped from
cultivation, Rare. Henry Avenue.

Melampyrum lineare. Cow wheat. Dry woods and shaded slopes. Rare. Known only from

30 BARTONIA

steep, rocky bank along south side of Thomas Mill Road.

Mimulus alatus. Monkey-flower. Low, wet ground. Rare. Wise’s Mill Road.

Mimulus ringens. Monkey-flower. Stream shores and wet meadows. Occasional. Livezey
Lane, Wise’s Mill Road

Pedicularis canadensis. Wood-betony, lousewort. Moist to dry woods and _ thickets.
Occasional in upper portion of valley. Thorps

Penstemon hirsutus. Beard-tongue. Dry, usually rocky woods and slopes. Occasional.

Scrophularia lanceolata. Figwort. Thickets and borders of woods. Infrequent.

Scrophularia marilandica. Figwort. Carpenter’s Square. Rich woods and _ thickets.
Occasional. Gypsy Lane, Walnut Lane, Kitchen’s Lane, Livezey Lane, Harper’s Meadow.

Verbascum blattaria. Moth mullein. An occasional weed of fields and clearings. Henry
Avenue, Harper’s Meadow.

Verbascum lychnitis. White mullein. clearings, waste places. Rare. Livezey Lane.

Verbascum thapsus. Common mullein. Fields, waste places, roadsides. Occasional. Harper’s
Meadow.

Veronica hederaefolia. \vy-leaved speedwell. Occasional in clearings. Henry Avenue,
Kitchen’s Lane, Valley Green, Rex Avenue.

Veronica officinalis. Common speedwell. Dry woods, thickets, and clearings. Frequent.
Valley Green, Rex Avenue, Harper’s Meadow.

Veronica peregrina, Neckweed. Damp, open places, roadsides. Occasional. Rex Avenue.

Veronica serpyllifolia. Thyme-leaved speedwell. Meadows and picnic areas. Occasional.

BIGNONIACEAE, BIGNONIA FAMILY
Catalpa_ bignonioides. Catalpa, Indian-bean. Cultivated and escaped or naturalized
throughout. Henry Avenue, Bluestone Bridge, Kitchen’s Lane, Leverington Street,
Cresheim Valley, Rex Avenue
Paulownia tomentosa. Princess-tree. Cultivated, widely escaped and naturalized, native of
Asia. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Carpenter’s Woods,
Mt. Airy Avenue, Devil’s Pool, Cresheim Valley, Rex Avenue, Bells Mill Road.

OROBANCHACEAE, BROOM-RAPE FAMILY
Conopholis americana. Squaw-root. Parasitic, mostly on various oaks. Sporadic. Not found
in recent years.
gene ‘virginiana. Beech-drops. Parasitic on the beech tree. Common throughout.
s Lane, Bluestone Bridge, Walnut Lane, Mt. Airy Avenue, Cresheim Valley, Rex
aie Bells Mill Road.

Orobanchae uniflora. One-flowered cancer-root. Parasitic on various herbaceous plants.
Occasional

PLANTAGINACEAE, PLANTAIN FAMILY
Plantago lanceolata. Rib-grass, buckhorn. A common weed of roadsides, fields, clearings,
and di il

Plantago major. Whiteman’s foot. Roadsides, open ground. Occasional.
Plantago rugelii. Rugel’s plantain. Same situations as P.lanceolata and P. major, but far
more abundant. Walnut Lane, Livezey Lane.

WISSAHICKON VALLEY PLANTS 31

RUBIACEAE, MADDER FAMILY

Cephalanthus occidentalis. Buttonbush. Damp thickets. Rare. Henry Avenue, above Walnut
Lane, Livezey Lane

Galium aparine. Cleavers. Woods, thickets, clearings. Fairly abundant. Kitchen’s Lane,
Cresheim Creek.

Galium asprellum. paeieict Low, moist, open or shaded ground. Occasional. Walnut Lane,
Harper’s Mea

Galium circaezans. Wild licorice. Rich to dry woods. Fairly abundant.

Galium triflorum. Bedstraw. Rich woods. Locally abundant. Kitchen’s Lane, Rex Avenue,
Bells Mill Road.

Houstonia caerulea (Hedyotis caerulea). Bluets, Quaker-ladies, innocence. Open woods,
thickets, and grassy clearings. Locally abundant. Hermit’s Lane, Walnut Lane, Carpenter’s
Woods, Bells Mill Road.

Mitchella repens. Partridge-berry. Dry to moist woods. Common throughout. Walnut Lane,
Kitchen’s Lane, Carpenter’s Woods, Mt. Airy Avenue, Cresheim Valley, Valley Green,
Rex Avenue, Thomas Mill Road, Bells Mill Road.

CAPRIFOLIACEAE, HONEYSUCKLE FAMILY

Diervilla lonicera. Bush-honeysuckle. Dry woods and clearings. Rare and local. Not
collected in recent years.

Lonicera japonica. Japanese honeysuckle. The most pernicious weed in the valley; in many
sections completely obliterating the native vegetation by forming solid stands which cover
wooded slopes and thickets.

Lonicera maackii. Maack’s honeysuckle. Occasional escape. Bells Mill Road.

Lonicera sempervirens. Trumpet honeysuckle, coral-honeysuckle.Woods and thickets. Not
seen in recent years and perhaps extinct.

Lonicera tatarica. Tartarian honeysuckle. Native of Eurasia. Widely planted and occasionally
escaping from cultivation. Cresheim Valley, Harper’s Meadow.

Sambucus canadensis. Common elder. Wet thickets and stream borders. Frequent throughout.
Henry Avenue, Walnut Lane, Carpenter’s Woods, Cresheim Valley, Valley Green, Thomas
Mill Road, Bells Mill Road, Harper’s Meadow

Symphoricarpus orbiculatus. Coralberry. Spread from cultivation and escaped along paths
and trails. Cresheim Valley, Thorps Lane.

Viburnum acerifolium. Maple-leaved arrow-wood. Dry or rocky woods. Common throughout.
Walnut Lane, Bluestone Bridge, Carpenter’s Lane, Cresheim Valley, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road, Henry Avenue.

Viburnum dentatum. Southern arrow-wood. Thickets. Hermit’s Lane, Henry Avenue, Walnut
Lane, Cresheim Valley. ee
Viburnum opulus. Guelder-rose, snowball-tree. Native of Europe. Escaped from cultivation.

Carpenter’s Woods, Cresheim Creek, Harper’s Meadow.

Viburnum prunifolium. Black-haw. Occasional in borders of woods.

Viburnum recognitum. Arrow-wood. Wet to dry thickets. Less frequent than V. dentatum,
which it resembles, but from which it differs in its smoother leaves and branchlets.
Carpenter’s Woods, Kitchen’s Lane, Cresheim Creek, Valley Green. =

Viburnum sieboldii. Japanese viburnum. Native of Asia. Escaped from cultivation. Walnut

e, Cresheim Valley.
Viburnum tomentosum (V. plicatum). Snowball. Native of Asia. Escaped from cultivation.

32 BARTONIA
Henry Avenue, Carpenter’s Woods, Cresheim Creek, Cresheim Valley.

VALERIANACEA, VALERIAN FAMILY
Valerianella intermedia (V. umbilicata). Corn-salad, lamb’s-lettuce. Damp to dry woods and
meadows. Rare.

CUCURBITACEAE, GOURD FAMILY
Echinocystis lobata. Wild balsam-apple. Thickets and stream banks. Occasional. Mt. Airy
Avenue, near Harper’s Meadow.
Sicyos angulatus. Bur cucumber. Deep thickets and bottomland. Infrequent. Walnut Lane,
Cresheim Valley.

CAMPANULACEAE, BELLFLOWER FAMILY.
Campanula aparinoides. Marsh bellflower. Low, damp ground, stream banks. Rare, and
local. Allen’s Lane.
Specularia perfoliata (Triodanis perfoliata var. perfoliata). Venus’s-looking-glass.
Occasional in open clearings, waste ground, on roadsides, etc.

LOBELIACEAE, LOBELIA FAMILY

(Campanulaceae)
Lobelia inflata. Indian-tobacco. Fields, roadsides, waste places, sometimes an abundant weed.
Walnut Lane, Carpenter’s Woods.
Lobelia siphilitica. Great lobelia. Rich, low woods and swamps. Occasional. Walnut Lane,
Monastery, Thomas Mill Road, Thorps Lane, Harper’s Meadow.

COMPOSITAE, COMPOSITE FAMILY
(Asteraceae

Achillea millefolium. Milfoil, yarrow. Roadsides, meadows, clearings, etc. Frequent
throughout. Henry Avenue, Thomas Mill Road, Harper’s Meadow

Ambrosia artemisiifolia. Common ragweed. Common throughout on roadsides, banks,
disturbed ground, etc. Walnut Lane.

Ambrosia trifida. Great ragweed. Thickets, bottomland, etc. Common. The variety
integrifolia, with leaves entire instead of three-lobed, is occasional. Walnut Lane,
Cresheim Valley, Rex Avenue, Bells Mill Road, Harper’s Meadow

Antennaria fallax (A. parlinii ssp. fallax). Pussytoes. Open woods, shaded slopes, clearings,
etc. Occasional. Near Harper’s Meadow.

Antennaria neglecta. Pussytoes. Open, wooded slopes, clearings, etc. Fairly common. Above
Bells Mill Road.

Antennaria neodioica. Pussytoes. Dry woods and open slopes. Fairly common. Thomas Mill

Road.

Antennaria plantaginifolia. Pussytoes. Dry, open or shaded slopes, clearings, roadsides, etc.
Common throughout. Below Barren Hill Road, Bells Mill Road.

Anthemis cotula. Mayweed. Roadsides, waste places, etc. Occasional.

Arctium minus. Common burdock. Roadsides, waste ground. Occasional. Bluestone Bridge,
Walnut Lane, Carpenter’s Woods, Devil’s Pool, Cresheim Valley, above Thomas Mill
Road, Harper’s Meadow.

Artemisia vulgaris. Mugwort. Waste places. Near Walnut Lane Bridge, Carpenter’s Woods.

WISSAHICKON VALLEY PLANTS 33

Aster cordifolius. Common blue aster. Open woods and thickets. Fairly common. Henry
Avenue, Bluestone Bridge, Walnut Lane, Livezey Lane, Cresheim Valley, Thomas Mill
Road, Thorps Lane.

Aster divaricatus. White wood aster. Woods and thickets. Common. Henry Avenue,
Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Livezey Lane, Cresheim Valley, Valley
Green, Rex Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Aster ericoides (A. pilosus). Heath aster. Dry slopes and thickets. Occasional.

Aster lateriflorus. Aster. Moist to dry fields and thickets. Fairly frequent. Bluestone Bridge,
Carpenter’s Woods, Bells Mill Road.

Aster linariifolius. Aster. Dry slopes and ledges. Rare. Thomas Mill Road.

Aster novae-angliae. New England Aster. Damp thickets, meadows, etc. Infrequent. Not seen
in recent years, but should be sought for.

Aster patens. Aster. Dry, open woods and thickets. Casual. Thomas Mill Road.

Aster prenanthoides. Aster. Damp thickets, borders of woods, etc. Fairly common. Henry
Avenue, Bluestone Bridge, Walnut Lane, Livezey Lane, Rex Avenue, Thomas Mill Road.

Aster puniceus. Swamp aster. Damp thickets and meadows. Occasional. Cresheim Valley,
Bells Mill Road.

Aster schreberi. Aster. Damp woods and thickets. Locally abundant. Walnut Lane, Rex
Avenue, Bells Mill Road, below Barren Hill Road.

Aster simplex (A. lanceolatus ssp. simplex). Aster. Damp woods and thickets. Occasional.
Henry Avenue, Bluestone Bridge, Walnut Lane, Carpenter’s Woods, Livezey Lane,
Cresheim Valley, Thomas Mill Road, Harper’s Meadow.

Aster undulatus. Aster. Occasional in dry woods and thickets.

Aster vimineus (A. lateriflorus). Aster. Damp to dry woods and clearings. Rare.

Bidens bipinnata. Spanish needles. Roadsides and waste places. Occasional.

Bidens cernua. Stick-tight. Muddy shores, springs, etc. Occasional.

Bidens comosa. Beggar-ticks. Occasional in low, wet ground. Thorps Lan

Bidens frondosa. Stick-tight. A weed along roadsides, or in waste ground. sant Meadow.

Bidens vulgata. Stick-tight. Widely distributed in low ground, ditches, waste places, etc.

Chrysanthemum leucanthemum. Ox-eye daisy. Dry roadsides, clearings, fields, etc. Ours is
the variety pinnatifidum with dissected leaves. Frequent.

Chrysopsis mariana. Golden aster. Dry, sandy soil. Rare. Carpenter’s Woods.

Cichorium intybus. Chicory. Casual throughout. Bells Mill Road.

Cirsium arvense. Canada thistle. Clearings and waste ground. Occasional. Walnut Lane.

Cirsium discolor. Thistle. Thickets, stream banks, etc. Infrequent. Cresheim Valley.

Cirsium lanceolatum (C. vulgare). Bull-thistle. Roadsides and clearings. Occasional. Bells
Mill Road.

Eclipta alba (E. prostata). White eclipta. Muddy shores. Livezey Lane.

Erechtites hieracifolia. Fireweed. Damp thickets and clearings. Occasional. Walnut Lane,
Cresheim Valley, Bells Mill Road, near Harper’s Meadow

Erigeron annuus. Daisy fleabane. Clearings and waste places. Scattered throughou

Erigeron canadensis (Conyza canadensis var. canadensis). Horseweed. An eects weed
of cleared areas and roadsides. Walnut Lane. i

Erigeron pulchellus. Robin’s-plantain. Open woods, thickets, and dry clearings. Sporadic.

tesheim Creek, Rex Avenue.

Erigeron strigosus. White-top. Open slopes, roadsides, etc. Infrequent.

Eupatorium fistulosum. Joe-pye-weed. Damp thickets, borders of woods, meadows. Common

34 BARTONIA

throughout. Bluestone Bridge, Walnut Lane, Cresheim Valley, Thomas Mill Road, Bells
Mill Road, Harper’s Meadow.

Eupatorium perfoliatum. Boneset, thoroughwort. Damp thickets and meadows. Fairly
abundant. Walnut Lane, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Eupatorium purpureum. Sweet joe-pye-weed. Damp woods and thickets, moist shaded
roadsides. Common. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt.
Airy Avenue, Valley Green, Rex Avenue, Thomas Mill Road, Bells Mill Road, Harper’s
Meadow.

Eupatorium rugosum. White-snakeroot. Rich woods, thickets, and clearings. Abundant
throughout. Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Carpenter’s
Woods, Cresheim Valley, Rex Avenue, Thomas Mill Road, Harper’s Meadow

Galinsoga ciliata (G. quadriradiata). Galinsoga. A ubiquitous weed of clearings, roadsides,
etc. Walnut Lane, Livezey Lane, Cresheim Valley, Rex Avenue, Thomas Mill Road,
Harper’s Meadow

Gnaphalium obtusifolium. Cudweed, everlasting. Dry slopes and clearings. Occasional.

Helenium autumnale. Sneezeweed. Rich thickets and fields. Rare. Below Walnut Lane.

Helenium nudiflorum (H. flexuosum). Sneezeweed. Roadsides and clearings. Rare.

Helianthus decapetalus. Thin-leaf sunflower. Woods and thickets. Frequent. Mouth of creek,
Henry Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Valley
Green, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Helianthus divaricatus. Sunflower. Dry, sterile or sandy soil. Rare.

Helianthus tuberosus. Jerusalem artichoke. Thickets, borders of woods, etc. Occasional.

Heliopsis helianthoides. Ox-eye. Open woods and thickets. Rare.

Hieracium paniculatum. Hawkweed. Open woods and shaded slopes. Frequent. Henry
Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim
Valley, Valley Green, Thomas Mill Road, Bells Mill Road.

Hieracium scabrum. Hawkweed. Dry woods and thickets. Occasional. Bells Mill Road.

Hieraceum venosum. Rattlesnake-weed. Open woods and clearings. Occasional. Carpenter’s
Woods, Thomas Mill Road, Bells Mill Road.

Inula helenium. Elecampane. Cleared ground. Rare. Found years ago near Thorps Lane.
Perhaps no longer growing in the valley.

Krigia biflora. Dwarf dandelion. Dry, usually sandy soil. Rare. Carpenter’s Woods.

Lactuca biennis. Tall lettuce. Thickets, borders of woods, and clearings. Common
throughout. Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Mt. Airy Avenue, Cresheim
Valley, Valley Green, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Lactuca canadensis. Wild lettuce. Thickets, borders of woods, and clearings. Infrequent.

Lactuca floridana var. villosa. Blue-flowered lettuce. Dry thickets and roadsides. Rare. Rex
Avenue.

Lactuca scariola (L. serriola). Prickly lettuce. Roadsides and clearings. Occasional.

Prenanthes altissima. Rattlesnake-root. Damp to dry woods and thickets. Common
throughout. Walnut Lane, Carpenter’s Woods, Mt. Airy Avenue, Valley Green, Rex
Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Prenanthes trifoliolata. Gall-of-the-earth. Rich to dry woods. Fairly frequent. Cresheim
Valley, Valley Green, Thomas Mill Road.

Rudbeckia hirta. Black-eyed-Susan. Dry, open places. Occasional.

Rudbeckia laciniata. Tall coneflower. Rich, moist thickets and bottomlands. Common. Henry
Avenue, Bluestone Bridge, Walnut Lane, Kitchen’s Lane, Devil’s Pool, Cresheim Valley,

WISSAHICKON VALLEY PLANTS 35

Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Rudbeckia triloba. Coneflower. Naturalized occasionally in open woods, thickets, and
clearings.

Senecio aureus. Golden ragwort. Occasional in low, swampy ground. Cresheim Creek, Bells
Mill Road, Harper’s Meadow.

Sericocarpus asteroides (Aster paternus). White-topped aster. Dry, open woods and
clearings. Rare. Carpenter’s Woods, along bridle path, south of Bells Mill Road, near
summit of trail.

Silphium perfoliatum. Cup-plant. Rich woods and thickets. Infrequent. Below Henry Avenue,

near Harper’s Meadow

Solidago altissima (S. mesilate Tall goldenrod. Dry to damp thickets and clearings.
Fairly frequent. Hermit’s Lane, Walnut Lane, Cresheim Valley, Thomas Mill Road,
Harper’s Meadow.

Solidago arguta. Goldenrod. Open woods, thickets, and clearings. Infrequent. Not seen in
recent years.

Solidago bicolor. Silver-rod. Frequent in dry woods, thickets, and fields. Frequent. Walnut
Lane, Carpenter’s Woods, Cresheim Valley, Thomas Mill Road, Bells Mill Road

Solidago caesia. Blue-stemmed goldenrod. Damp, rich woods and thickets. Common. Henry
Avenue, Bluestone Bridge, Walnut Lane, Carpenter’s Woods, Mt. Airy Avenue, Cresheim
Valley, Rex Avenue, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Solidago canadensis. Canada goldenrod. Rich woods and thickets. Rare.

Solidago gigantea. Goldenrod. Rich thickets, swamps, etc. Freqent. Walnut Lane, Cresheim
Valley, Thomas Mill Road, Bells Mill Road, Harper’s Meadow.

Solidago graminifolia (Euthamia graminifolia var. graminifolia). Flat-top goldenrod.
Thickets and meadows. Carpenter’s Woods, Cresheim Valley, Thomas Mill Road.

Solidago juncea. Goldenrod. Dry, open places. Rare. Bells Mill Road.

Solidago latifolia (S. flexicaulis). Goldenrod. Rich woods and thickets. Common. Henry
Avenue, Bluestone Bridge, Walnut Lane, Carpenter’s Woods, Mt. Airy Avenue, Livezey
Lane, Thomas Mill Road, Bells Mill Road, Harper’s Meadow

Solidago nemoralis. Goldenrod. Dry, sterile clearings and fields. Row Carpenter’s Woods,
Bells Mill Road, below Barren Hill Road.

Solidago odora. Anise-scented goldenrod. Dry, open woods and meadows. Rare. Thorps
L

ane.

Solidago puberula. Goldenrod. Dry, sterile slopes and clearings. Rare.

Solidago rugosa. Goldenrod. Wet thickets and stream banks. Frequent. Henry Avenue,
Bluestone Bridge, Walnut Lane, Carpenter’s Woods, Thomas Mill Road, Cresheim Valley,
Bells Mill Road.

Solidago squarrosa. Goldenrod. Dry, rocky woods and shaded slopes. Rare. Collected in
1866 but no locality given.

Solidago ulmifolia. Goldenrod. Dry, rocky woods and thickets. Rare. Thorps Lane.

Sonchus asper. Sow-thistle. Roadsides, waste ground, etc. Sporadic. Henry Avenue.

Taraxacum officinale. Common dandelion. An ever-present weed of roadsides, clearings,
hoy ground, etc. Bluestone Bridge, Livezey Lane, Thomas Mill Road, Harper’s
Me

‘trast Vovabobacienie Iron-weed. Wet thickets and low, swampy ground. Occasional.
Cresheim Valley.
Xanthium italicum (X. strumarium var. canadense). Cocklebur. Low ground, stream banks

36 BARTONIA

and waste places, etc. Thorps Lane.
Xanthium pensylvanicum (X. strumarium var. canadense). Cocklebur. Bottomland, waste
places, etc. Occasional. Cresheim Valley, Harper’s Meadow.

Bartonia No. 59: 37-47, 1996
The Identity of Anemone riparia (Ranunculaceae)

CARL S. KEENER
Department of Biology, The Pennsylvania State University, University Park, PA 16802

EDWARD T. DIx
Bureau of Forestry, PO Box 8552, Harrisburg, PA 17105

BRYAN E. DUTTON
Brooklyn Botanic Garden, 1000 Washington Ave., Brooklyn, NY 11225

In recently reviewing the Anemone virginiana L. complex in Pennsylvania, it has become
apparent that the name, Anemone riparia Fernald (as described by Fernald in 1899), has been
applied to a discordant set of elements. In general, systematists have determined specimens
as A. riparia based on either (1) larger sepal size (up to 2 cm long), or (2) presence of
cuneate-based leaf divisions. These disparate treatments have led to a considerable
divergence regarding the identity of A. riparia, as well as different opinions with respect to
its range throughout Canada and the eastern United States. Consequently, the principal aim
of this study is to clarify the identity of Fernald’s A. riparia and to apply these results to a
more discriminating analysis of the A. virginiana complex within North America.

HISTORICAL OVERVIEW

In 1899, M. L. Fernald described a new species of Anemone characterized by relatively
large sepals (1.5-2.0 cm long), subcylindric fruiting heads, suberect styles, and cuneate-based,
thin leaflets and which occurs chiefly on calcareous river banks, rocky shores, and moist
cliffs in eastern Canada and northeastern United States (Fernald 1899; Robinson & Fernald
1908). Because of continued confusion between A. riparia and A. virginiana, Fernald (1917)
later attempted to sharpen the differences by comparing leaf segment bases, anther lengths,
thickness of fruiting heads and style orientation. Fernald’s remarks are pertinent: "Contrasted
with A. virginiana it [A. riparia] has the leaf-segments usually more cuneate at base,
although this character is by no means absolute; anthers 0.7-1.2 mm. long, those of the more
southern A. virginiana running from 1.2-1.6 mm. long; its fruiting head 7-11 mm. thick, as
contrasted with A. virginiana in which the heads are 1.2-1.5 cm. thick; and the subulate pale
styles ascending or subascending in fruit, as contrasted with the firmer, more divergent styles
of A. virginiana" (Fernald 1917). Moreover, Fernald (1917) noted that both taxa have
polymorphic sepals ranging from small, more or less inconspicuous to large, white petaloid
forms, leading him to name several of these variants. Fernald also named one color form
(f. rhodantha) of A. riparia. These observations were repeated in Fernald’s key to Anemone
in the eighth edition of Gray’s Manual of Botany (Fernald 1950, p. 661).

Meanwhile, other taxonomists formed their own opinions concerning the identity of A.
riparia. In both his large-scaled treatments of the flora of the southeastern United States,
Small (1903, 1933) distinguished 4. riparia from A. virginiana on the basis of sepal color,

aq

38 BARTONIA

size of fruiting head, and style orientation, A. riparia having clear white sepals, fruiting
heads less than 1 cm thick, and achenes with appressed or ascending styles. Anemone
virginiana, on the other hand, is distinguished by its greenish sepals, fruiting heads greater
than 1 cm thick, and achenes with spreading styles. Moreover, Small claimed that A. riparia
occurred on "[bJanks, often calcareous stony soil, Blue Ridge and more northern provinces,
N.C. to Tenn., Alb., and Que." (Small 1933, p. 517).

In the second edition of their ///ustrated Flora, Britton and Brown (1913) listed A. riparia
as a synonym of A. virginiana. But in the revision of Britton and Brown’s //lustrated Flora,
Gleason (1952) followed Fernald in recognizing both A. riparia and A. virginiana.
Gleason’s key (1952, Vol. 2, p. 180) distinguished both species on the basis of leaf segment
patterns and style orientation. However, Gleason remarked that "the taxonomic status of the
group of plants described under this name [A. riparia] is still open to question" (1952, p.
180). By the time the one-volume Manual was produced (Gleason & Cronquist 1963),
Gleason and Cronquist reverted to Britton and Brown’s earlier view by lumping A. riparia
with A. virginiana. Gleason and Cronquist observed, however, that "[s]ome plants from the
northern part of the range diverge toward A. cylindrica [distinguished from the A. virginiana
complex by its 5-9-leaved involucre, naked peduncles, and generally narrower, more
cylindric fruiting heads] in their somewhat narrower heads and in having the I[ea]f-segments
more cuneate below. These have been segregated as A. riparia Fern., but they do not appear
to be sharply separable from A. virginiana" (Gleason & Cronquist 1963, p. 316). In the
second edition of the Manual, Gleason and Cronquist (1991, p. 52) continued to include A.
riparia with A. virginiana, and with only the addition "...var. alba A. Wood or...," the same
comment quoted above is repeated in the newer edition.

Over the years these disparate views concerning the taxonomic disposition of A. riparia
have been reflected in the floristic treatments of other authors. For example, Rydberg (1932,
pp. 332-333) generally followed Fernald, but considerably expanded the range of A. riparia
("River banks: Me.-Va.-Alta.") and distinguished A. riparia from A. virginiana chiefly on
the basis of style orientation, and sepal color and length ("Petals [sic] white, 1.5 cm. long”
vs. "Petals greenish white, 1 cm. long or less," respectively). Deam (1940) viewed A.
virginiana as a variable species, especially with respect to the stamen length, and sepal size,
shape, color, and texture, and concluded that "after a careful study of 77 specimens from all
parts of [Indiana] I have decided that the characters are too variable to be of taxonomic
value." Both Scoggan (1957) and Seymour (1969) recognized A. riparia, but except for leaf
pubescence (Seymour), they added no new characters separating these two species.

In a study of the flora of the prairie provinces of Canada, Boivin (1968, 1969) concluded
that, based on the smaller flowers (sepals +1 cm long, greenish) and narrower, more cylindric
fruiting heads, the elements of the A. virginiana complex in that region should belong to 4
taxon he named var. cylindroidea. Boivin observed that his new variety "[rJanges in Canada
from southern Quebec west to northern British Columbia," but "[g]rades eastward into var.
riparia and var. virginiana" (Boivin 1969). Boivin (1969) noted also that "[s]upposed
differences in leaf shape have not proved worth retaining," and that the "name var. riparia
(Fern.) Boivin 1966 for the large flowered eastern plant should be replaced by the earlier and
correct var. alba Wood 1861" (see Boivin 1966, 1967, and Wood 1861, p. 203).

During the past decade, Mitchell and Dean (1982) reviewed the Anemone virginiana
complex for New York and concluded that "[t]his seldom-studied group of plants presents
some interesting problems in variation.". They suggested, moreover, that "var. alba W
[= A. riparia] was derived through ancient hybridization of A. virginiana and A. cylindrica

IDENTITY OF ANEMONE RIPARIA 39

[generally sympatric with A. virginiana northward], and has now backcrossed to A.
virginiana sufficiently to form a morphological bridge as well as an ecological cline.
Variety alba has a much broader northern range than typical var. virginiana, and has
apparently inherited the heterozygosity needed for postglacial invasion of cool, moist
habitats—thus expanding the range of the species." Voss (1985, pp. 229-230) treated A.
virginiana "in a broad sense" and pointed out that large-flowered plants "were named as A.
virginiana var. alba Wood and were included in A. riparia Fern., a name often applied (but
frequently with some misgivings) to certain plants in our area." Voss noted that "[d]istinc-
tions in anther length, sometimes cited, appear useless, and the angle of the style is scarcely
better. Typical var. virginiana tends to have fatter, more ovoid fruiting heads (to 17 mm
thick) and to be larger-leaved, the leaf segments more ovate, than var. alba, which in some
ways approaches A. cylindrica." Finally, in the Flora of the Great Plains, Sutherland (1986)
merely listed A. riparia as a synonym under A. virginiana without comment.

In summary, two points should be stressed: (1) taxonomists are not in agreement
concerning the recognition of the taxonomic group variously named A. riparia Fernald or
A. virginiana L. var. alba A. Wood, and (2) taxonomists have separated A. riparia from A.
virginiana on the basis of a differential set of characters. These are chiefly sepal size and
color or shape of leaflet bases, although anther length, style orientation, thickness and shape
of fruiting heads, and leaf pubescence have been used also as discriminators. Undeniably,
by recognizing different sets of characters, taxonomists have developed disparate taxonomic
treatments of this complex. The aim of this study, therefore, is to clarify the identity and
taxonomic status of Anemone riparia by means of an analysis of the character states
Previously identified as possible discriminators between this taxon and A. virginiana.
secondary aim is to evaluate the status of Boivin’s var. cylindroidea, and thus round out a
preliminary taxonomic treatment of the Anemone virginiana complex.

MATERIALS AND METHODS

A total of 341 specimens from throughout the range of the A. virginiana complex
(specimens were borrowed from A, CM, DAO, F, GH, NEBC, NYS, PAC, PH) were scored
for the shape of the involucral leaf base (truncate-obtuse or cordate-reniform; involucral
leaves were selected because not all specimens had basal leaves which are larger but similar
in overall shape), and the shape of the basal portion of the central leaflet divisions (concave,
Straight, convex). Subsamples with well-preserved floral organs were selected from
herbarium material of each taxon. The lengths of sepals and anthers and the length and
thickness of the fruiting heads were measured to the nearest 0.1 mm. Style orientation and
leaf pubescence are too indeterminate and so were omitted from any further analysis. Flower
and fruiting head measurements were compared by using the Ryan-Einot-Gabriel- Welsch
multiple range test option of the SAS analysis-of-variance procedure. This test minimizes
rejection of the null hypothesis (that two samples are equal) when it is indeed true. e
taxon with the minimum number of preserved flowers (N = 32) or fruiting heads (N = 30)
determined the sample size for the REGW/ANOVA. The analysis was done on the IBM
3090/180 computer at the University Park Campus of The Pennsylvania State University.

Ideally, all characters should be measured from a set of complete specimens. Given that
useful characters included both flowering and fruiting material, this proved impossible in
Practice. This also precluded any attempted correlation analysis involving, e.g., sepal length
and fruiting head length and thickness all obtained from the same specimen. Consequently,

40 BARTONIA

for purposes of this study, we initially classified all specimens into three groups based on
a critical comparison with the type specimens of Anemone riparia and A. virginiana var.
cylindroidea. Measurements were taken from subsamples of each group. It would have
been preferable to have pooled all the data and then derive a principal components analysis,
but the specimens we utilized simply lacked one or more of the critical characters. Our
study, thus, reflects an analysis based on a pooled grouping of initially classified herbarium
specimens and therefore lacks some of the rigor that a thorough population-based study
would have yielded.

RESULTS

The results of the multiple range test indicate that the differences in sepal length among
all three taxa are significantly different (Table 1). Anemone virginiana var. cylindroidea
shows relatively little variation around the shortest mean length. The sepal measurements
of var. alba overlap the range of var. virginiana but average 2 mm longer. The differences
in anther length and fruiting head thickness between var. alba and var. cylindroidea are not
significant but these taxa have significantly shorter anthers and narrower fruiting heads than
var. virginiana. The three taxa are not significantly different in fruiting head length.

TABLE 1. Statistical pi cleues of selected characters of the varieties in the Anemone virginiana complex with
mean + s.d., range, and NV.

Character cylindroidea alba virginiana

Sepal length (mm) P2412", 5210233 15.24+3.3°, 9-21, 33 12.9+3.5°, 6-21, 39
Anther length (mm) 0.940.1"°, 0.7-1.2, 32 1,040.2’, 0.8-1.5, 32 1.3+0.2°, 0.9-1.7, 32
Fruiting head length (mm) 19.8+4.5*, 13-36, 30 19.042.4*, 13-24, 34 19.243.2", 12-28, 30
Fruiting head thickness (mm) 8.7+1.2", 7-11, 30 9.1+1.0°, 8-11, 33 11.041.2°, 9-14, 33

‘Results based on a Ryan-Einot-Gabriel-Welsch multiple range test (alpha = 0.01). Mean values followed by the
same letter are judged not significantly different.

With respect to the involucral leaves, the base of the central leaflet or division in var.
virginiana is straight-sided in about half the specimens, the other half being divided about
equally between those with concave sides and those with convex sides (Table 2). In var.
cylindroidea the basal leaflet edges are again predominantly straight, with about 25% of the
specimens having concave sides and fewer than 5% are convex. However, in var. alba the
predominant shape is concave and although about one-third of the specimens are straight-
sided, no convex edges were observed. Anemone virginiana var. alba also shows a higher
percentage of leaves with truncate-obtuse bases whereas var. virginiana (and vat.
cylindroidea—despite missing data) generally has cordate leaf bases (Table 2).

TABLE 2. Involucral leaflet morphology.

pe ee
Lower margin of central leaflet (percent) Base of involucral leaves (percent)
Taxon concave straight convex truncate cordate
var. cylindroidea (N=57) 25 70 5 _ cee
var. alba (N=64) 65 35 0 =71) 82 18
var. virginiana (N=58) =. 21 53 26 =58) 20 80

IDENTITY OF ANEMONE RIPARIA 41

Clearly, no single character demarks unequivocally any variety from the other two.
Furthermore, the continuous nature of the variables makes the determination of some
specimens, especially from the northern interior of the geographic range rather problematic.
Nevertheless, at the limits of the overall range, the three taxa are sufficiently distinct with
respect to the ensemble of five characters (viz., sepal and anther length, fruiting head
thickness, margins of central involucral leaflets, base of involucral leaves; all five characters
should be employed to avoid the confusion resulting from utilizing differential sets of
characters) that in our opinion they should be recognized as varieties of a wide-ranging
polymorphic species. Critical population studies might well change this conclusion, and, in
fact, it would be desirable to conduct such studies in the northern part of the range of
Anemone virginiana, especially in the area from the Great Lakes eastward to Newfoundland.
In our study, no evidence of hybridization appeared to exist. Hybridization experiments
involving all three taxa would be desirable.

TAXONOMY
Anemone virginiana L.

Tall, caudiciferous or short-rhizomatous, perennial herb; stems 0.5-1.0 m tall, loosely
pubescent. Basal leaves compound or deeply 3-parted, broadly reniform to cordate-ovate in
outline, to 15 cm wide; central segments or leaflets rhombic-ovate in outline, variously
parted, cleft, lobed or incised, margins crenate to serrate distally, basally concave, cuneate,
or convex, thinly pubescent; petioles to 2.8 dm long, variously pubescent. Involucral leaves
similar to basal leaves, usually smaller, lower whorled, upper opposite; base of leaf outline
tcordate or reniform to nearly truncate; petioles 1-8 cm long; central segments or leaflets
to 12 cm long and 6 cm wide, firm, green to dark green above, lighter or pale green below,
variously pubescent on both surfaces, apices acute, margins variously lobed, incised, serrate
or crenate, bases convex, cuneate, or rounded, base to lower lobe notch 2.6-6.4 cm, petiolules
to 1 cm long. Inflorescence terminal, (1-)3-5(-9) flowered, usually with an opposite-leaved
secondary involucre. Sepals typically 5, separate, whitish or greenish-white, ovate to
obovate, 5-21 mm long, leathery to thin, variously pubescent abaxially; petals or staminodia
absent; stamens numerous, anthers cylindrical, 0.7-1.7 mm long, filaments capillary, to7 mm
long, glabrous. Fruiting heads ovoid, oblong-cylindric to cylindric, to 3.6 cm long, 0.7-1.4
cm thick; achenes brownish, compressed, obovoid, ca. 1.5 mm. broad, densely woolly; styles
greenish to purplish apically, spreading to ascending, 1-2 mm long, short-pubescent or
sometimes glabrous apically. Chromosome no.: = 8. Three intergradient varieties.

Key to Varieties of Anemone virginiana

1. Sepals greater than 1 cm long, usually thinly pubescent abaxially; anthers usually greater
than 1 mm long; wide-ranging plants of moist habitats ......-.----+++++-5: 2

1. Sepals less than 1 cm long, densely tomentose abaxially; anthers usually less than | mm
long; mostly Canadian plants of dry woods, sandy ridges and grasslan
ee CART OR SW 14 ws ee var. cylindroidea

2. Ge inveteceal saute " i te r reniform (rarely subtruncate), the margins of the
cin Pon narnia nahiaclee to eve nek generally light green, variously
lobed or serrate, variously pubescent; anthers typically greater than 1.1 mm long; fruit-

42 BARTONIA

ing heads ovoid to ovoid-cylindric, often greater than 11 mm thick; plants of dry rocky
open woods, slopes, moist thickets, etc.; wide-ranging ......... var. virginiana
Base of involucral leaves often truncate to subtruncate (sometimes reniform or
cordate), the margins of the central leaflets basally concave to straight-sided, generally
deep green, apically incised, thinly pubescent; anthers typically less than 1.2 mm long;
fruiting heads tovoid-cylindric, ca. 10 mm thick; plants of riparian areas, — cliffs
and ledges, etc.; chiefly New England ao adjacent Canada ......... alba

a

1. ANCHORS VaNEIBIA® L. var. virginiana (Fig. 1)

Anemone virginiana L., Sp. Pl. 1: 540. 1753. TYPE: Habitat in Virginia (Lectotype: LINN-IDC Microfiche 710.19,
designed by Reveal et al. 1992).

Anemone virginiana f. leucosepala Fernald, Rhodora 19: 140. 1917. TYPE (so designated on the herbarium sheet
ia GH): North Carolina, Biltmore, June 28 and August 9, 1897, Biltmore egal no. 54b [GH!; parse.
of 28 June (in flower and mounted on left side of sheet) herein designated as TYPE inasmuch as ther
are ig ad of two different dates on the same sheet (9 Aug specimen in fruit)].

ae virginiana f. rubrosepala House, New York State met ee 243-244: 26. 1923. TYPE: New York,
Greene Co., Calo, June 18, 1918, L. S. Slater s.n. (Holotype: NYS!).

ibaa virginiana f. plena E. J. Palmer & Steyermark, Brittonia 6 113. 1958. TYPE: Missouri, St. Louis Co.,

ockw oods Reservation, 1956, G. E. Moore s.n. (Holotype: F!).

Se |
fhe oy

\

Fig. 1. Distribution of Anemone virginiana var. virginiana in North America.

Base of involucral leaves +cordate or reniform to rarely subtruncate; central leaflets lobed
and variously toothed, basal margins mostly convex to cuneate, typically light green d
variously pubescent. Sepals 6-21 (ave.: 12.9; N = 39) mm long, thinly pubescent abaxially;
anthers 0.9-1.7 (ave.: 1.3; N = 32) mm long; fruiting heads 12-28 (ave.: 19.2; N = 30) mm

IDENTITY OF ANEMONE RIPARIA 43

long, 9-14 (ave.: 11.0; N = 33) mm thick, ovoid to ovoid-cylindric. Flowering in June to
August; fruiting in July to September. Herbs of dry rocky open woods, slopes, thickets, and
river banks throughout eastern North America from southeastern Georgia to northeastern
Louisiana, eastern Oklahoma, eastern Kansas and eastern and northern Nebraska, north to
western North Dakota, east to central Maine and south-central Quebec (Roberval) (Fig. 1).

Anemone virginiana f. leucosepala Fernald is a large-sepalled (18 mm long) form with the
typical anthers (1.2 mm long) of A. virginiana var. virginiana. Except for the concave
leaflet bases (comparable to A. virginiana var. alba) the specimens (flowering, one fruiting
peduncle) are otherwise similar to var. virginiana.

Anemone virginiana f. rubrosepala House is a reddish-sepalled variant of var. virginiana.
Its general habit, long anthers (1.5 mm long), and convex involucral leaflet bases certainly
place it with var. virginiana.

Anemone virginiana f. plena E. J. Palmer & Steyermark is based on a large plant bearing
flowers with many sepal-like sterile stamens.

2. Anemone virginiana var. alba (Oakes) A. bd (Fig. 2)
Anemone cylindrica var. alba Oakes, M ag. Hort. Bot. vii: 182. 1841. TYPE: veers sag Co., Paha potas a6
obbins s.n. (Lectotype, herein designated: GH!; specimen dis shos rte igh
side of sheet). Anemone virginiana & alba (Oakes) A. Wood, rage ook Bot (ed. 1847): 140. 1847; Anemone
virginiana 8 alba [Oakes] A. Wood, ese book Bot. (ed. 1861): 2
Anemone riparia Fernald, Rhodors 1: 51. 1899. TYPE: Maine 5 Risse n uis eae Dover, June 25, 1894, M. L.
Fernald s.n. (Lectotype, herein ie GH!). Anemone virginiana var. riparia (Fernald) B. Boivin,
Naturaliste Canad. 94: 645. 1967.
Anemone riparia f. rhodantha Fernald, Rhodora 19: 139. 1917. TYPE: Canada, Quebec, Gaspé County, banks of
e Grand River, July, 1902, G. H. Richards s.n. (Holotype: GH!). Anemone virginiana var. riparia forma
rhodantha (Fernald) B. Boivin, Naturaliste Canad. 94: 645. 1967.
Anemone riparia f. inconspicua Fernald, Rhodora 19: 140. 1917. TYPE: Canada, Quebec, Gaspé County, Percé
Mt., Percé, July 25, 1905, Williams, Collins & Fernald s.n. (Holotype: GH!).

Base of involucral leaves truncate to subtruncate, occasionally reniform or cordate; central
leaflets sharply toothed to incised, basal margins usually cuneate to concave, typically dark
green and sparsely pubescent. Sepals 9-21 (ave.: 15.2; N = 33) mm long; anthers 0.8-1.7
ata 1.0; N = 32) mm long; fruiting heads 13-24 (ave.: 19.0; N = 34) mm long, 8-11 (ave.:

9.1; N = 33) mm thick, +ovoid-cylindric. Flowering in mid-May to mid-July; fruiting in late
June to September. Herbs of ledges, bluffs, rocky outcrops, open woods, alluvial river
thickets, lake shores, and roadsides, often in calcareous soil, ranging throughout the New
England states and adjacent Canada from western Newfoundland and Anticosti Island to
Victoria Co., Nova Scotia, south through eastern Quebec and New Brunswick to Connecticut
and southeastern New York, north through southern Ontario to Moose Factory ochrane
Terr. Dist.), southwestward to northern Michigan, Minnesota, and eastern Wisconsin (Fig.
2).

In 1841 when William Oakes named a new variety of Anemone cylindrica his protologue
was: "Anemone cylindrica Gray. Torrey and Gray, Flora, I., 13. var. alba Oakes. Flowers
larger, sepals white. On rocky ledges in Castleton, Burlington, and other places in Vermont,
also in Uxbridge, Mass. Dr. Robbins, 1829" (Oakes 1841). Shortly thereafter, this variety
was taken up by Alphonso Wood in the second (1847) edition of his Class-Book of Botany
under the name of A. virginiana ("8 alba. Oakes. Fis. larger; sep. white—Ledges, Vt. Dr.
Robbins."). However, when M. L. Fernald described A. riparia (Rhodora |: 51. 1899) he
listed two synonyms: "A. cylindrica, vat. alba, Oakes, Hovey’s Mag. vii. 182.

BARTONIA

Fig. 2. Distribution of Anemone virginiana var. alba in North America.

A. virginiana, var. alba, Wood, Class Book, 203." Nevertheless, it appears that Fernald
consulted A. Wood’s edition of 1861 in which var. alba was given as "B. alba. Fis. larger;
sep. white——Ledges, Vt. (Dr. Robbins.)." Consequently, we conclude that in the 1861 edition
Wood simply deleted Oakes as author of the basionym, but Fernald, apparently unaware of
Wood’s earlier (1847) transfer of Oakes’s var. alba to A. virginiana, assumed that Wood was
the only author of A. virginiana var. alba and so listed the name ("A. virginiana, var. alba,
Wood, Class Book, 203") in his description of A. riparia in 1899. This mistake has been
perpetuated recently by Boivin (1969, p. 282): "The name var. riparia (Fern.) Boivin 1966
for the larger flowered eastern plant should be replaced by the earlier and correct var. alba
Wood 1861." Mitchell and Dean (1982), and Voss (1985) followed suit in referring to "var.
alba Wood," but they did not recognize this taxon (i.e., Fernald’s A. riparia) as occurring
in New York and Michigan, respectively.

Lectotypification of Anemone cylindrica A. Gray var. alba Oakes is based on a sheet in
GH bearing a label "Anemone virginiana alba - grandiflora. Castleton, Vt. Dr. Robbins.
Oakes" and appears closest to Oakes’s protologue as given above. The sheet consists of two
specimens, apparently representing two different plants. Both specimens have cuneate-based
central involucral leaflets. However, one specimen has light-green leaves, sepals ca. 17 mm
long, anthers ca. 1.4 mm long, and has the general aspect of A. virginiana var. virginiana.
The second specimen has dark-green leaves, sepals ca. 14 mm long, anthers ca. 1.0 mm long,
and is referable to Fernald’s Anemone riparia. Accordingly, we are designating this
specimen (with the smaller sepals and shorter anthers, and mounted on the right side of the
sheet) as the lectotype for A. cylindrica var. alba Oakes (= A. virginiana var. alba (Oakes)

IDENTITY OF ANEMONE RIPARIA 45

A. Wood).

Anemone riparia f. rhodantha Fernald, based on a very poor specimen (one flower with
purplish sepals 15 mm long and anthers ca. 1 mm long, no basal leaves, but with badly
pressed involucral leaves), probably belongs to var. alba.

Anemone riparia f. inconspicua Fernald is based on a specimen with two flowers having
sepals 9-11 mm long and anthers ca. | mm long. Except for the relatively shorter sepals
(though not as short as those in var. cylindroidea), the type is clearly var. alba.

3. Anemone virginiana var. cylindroidea B. Boivin (Fig. 3)
Anemone virginiana var. cylindroidea B. Boivin, Phytologia 18: 281. 1969. TYPE: Canada, Saskatchewan, Pike
Lake, July 31, 1949, Boivin, Russell & Breitung 6733 (Holotype: DAO)).

¢

Fig. 3. Distribution of Anemone virginiana var. cylindroidea in North America.

Base of involucral leaves cordate to reniform; central leaflets variously lobed and toothed,
basal margins typically cuneate (sides straight), usually light green to ashen and often silky
pubescent. Sepals 5-10 (ave.: 7.2; N = 33) mm long, usually densely tomentose abaxially,
anthers 0.7-1.2 (ave.: 0.9; N = 32) mm long; fruiting heads 13-36 (ave.: 19.8; N = 30) mm
long, 7-11 (ave.: 8.7; N = 30) mm thick, cylindric. Flowering in early June to early August,
fruiting in late July to late August. Herbs of open woods, grasslands, sandy ridges, moist
river banks, etc., ranging from eastern Quebec (Gaspé Pen.) and western New Brunswick to
southern Quebec and adjacent northern New York, southern Ontario, central Minnesota, and
Scattered northwestward through the grassland areas of Manitoba, Saskatchewan, and Alberta
to central British Columbia (Fig. 3).

46 BARTONIA

With respect to the three varieties of A. virginiana, var. cylindroidea is closest
phenetically to A. cylindrica A. Gray. Both A. virginiana var. cylindroidea and A. cylindrica
have short sepals with densely tomentose abaxial surfaces and cylindrical fruiting heads.
Anemone cylindrica usually has >3 involucral leaves, lacks involucels on the secondary
peduncles, and in contrast to the greenish styles of A. virginiana, has crimson styles. These
characters usually —_— A. cylindrica from A. virginiana var. cylindroidea, but
intergradient forms and rare hybrids sometimes occur, which suggests possible local
introgression following ote ee of the last glacial advance.

ACKNOWLEDGEMENTS

We extend our grateful thanks to the following herbaria for loans of specimens and critical
ty , CM, O, F, GH, NEBC, NYS, PH, plus ALU, NCU, UNA, and US. We
statis acknowledge ‘the financial support of the Eberly College of Science of The
Pennsylvania State University in underwriting the use of the IBM 3090/180 Computer at the
University Park Campus. Special thanks are due to Drs. K. N. Gandhi, Marilyn Park,
Ronald A. Pursell, James L. Reveal, Lloyd Stark, and Edward G. Voss for their helpful
comments, but we do not impute any errors of fact or judgment to these esteemed
colleagues.

LITERATURE CITED

BoIvin, B. 1966. Enumération des plantes du Canada III. 217. open ty Naturaliste Canad. 93: 584-591
[Anemone virginiana L. var. riparia (Fernald) B. Boivin, p. 587, i
Bolvin, B. 1967. Enumération des plantes du Canada VII. ee Cana om 625-655 [Anemone virginiana
L. var. riparia (Fernald) B. Boivin, p. 645, is validly published].
BoIviN, B. 1968. Flora of the Prairie Provinces, Pt. Il. Phytologia 16(3); 219-261 [Ranunculaceae, pp. 232-254].
BoIvVIN, B. 1969. Flora of the Prairie Provinces, Pt. Il: Additions and corrections. Phytologia 18(5): 281-282 [173-
174].
BRITTON, N. L. & A. BROWN. 1913. An illustrated flora of the northern United States, Canada and the British
possessions, ed. 2, Vol. II. Charles Scribner’s Sons, New York.
C. 1940. Flora of Indiana. Publ. by the Dept. of Consetvetion, Division of Forestry, Indianapolis.

6 pp.
FERNALD, M. L. 1899. Two plants of the Crowfoot Family. Rhodora 1: 48-52.
FERNALD, M. L. 1917. Some color forms of American anemones. Rhodora 19: 139-141.
FERNALD, M. L. 1950. Gray’s manual of botany, eighth ed. American Book Co., New York. Ixiv + 1632 pp.
GLEASON, H. A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent
Canada. Vol. 2. Lancaster Press, Lancaster.
GLEASON, H. A. & A. CRONQUIST. 1963. Manual of vascular plants of northeastern United States and adjacent
Canada. D. Van Nostrand Co., Princeton. lii + 810 p
GLEASON, H. A. & A. CRONQUIST. 1991. Manual of ie plants of northeastern United States and adjacent
anada, 2nd ed. The New York Botanical Garden, Bronx. Ixxv + 910
pete: R. S. & J. K. DEAN. 1982. Ranunculaceae (Crowfoot family) of New York State. Contributions to
Flora of New York State IV. Bull. No. 446, New York State Museum, Albany, N.Y. vi + 100 pp
Guat W. 1841. Notice of some rare plants of New England, with descriptions of some new species. Mag. Hort.
Bot. 7: 178-186.
REVEAL, J. L., C. E. JARvis, F. R. BARRIE, & B. E. DuTTON. 1992. On the typification of seven names in
Anemone (Ranunculaceae) proposed by Linnaeus. Bartonia 57: 28-31.
ROBINSON, B. L. & M. L. FERNALD. 1908. Gray’s new manual of botany, 7th ed. American Book Co., New York.

926 pp.
RYDBERG, P. A. 1932. Flora of the prairies and plains of central North America. The New York Botanical garden,
New York. viii + 969 pp.

IDENTITY OF ANEMONE RIPARIA 47
SCOGGAN, H. J. 1957. Flora of Manitoba. National Museum of Canada. Bull. No. 140, Biol. Series No. 47. vi

pp.
SEYMOUR, F. 1969. The flora of New England. Charles Tuttle Co., Rutland. xvi + 596 pp.
SMALL, J. K. 1903. Flora of the Southeastern United States. Publ. by the author, New York. xii + 1370 pp.
SMALL, J. K. 1933. Manual of the Southeastern flora. Publ. by the author, New York. xxii + 1554 pp
SUTHERLAND, D. 1986. i rhea Pages 84-107 in Great Plains Flora Association. Flora of the Great Plains,
Univ. of Kansas Press, Lawrence.
Voss, E.G. 1985. Michigan flor pt. II: Dicots (Saururaceae-Cornaceae). Cranbrook Institute of Science, Bull.

Woop, A. 1847. A class-book of botany. Crocker & Brewster, Boston. ii + 645 pp
Woop, A. 1861. Class-book of botany. A. S. Barnes & Burr, New York. viii + 832 pp.

awe ¥
su

es

PRU tae “vie

ey

pr?

Bartonia No. 59: 49-54, 1996

Distribution, Life Forms, Taxonomic Categories, and
Habitats of the Endangered and Threatened Vascular
Plants in Pennsylvania: A Summary

JEFFREY L. WALCK'
Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Pkwy., Philadelphia, PA 19103
and Morris Arboretum of the University of Pennsylvania, 9414 Meadowbrook Ave., Philadelphia, PA 19118

Two-thousand and seventy-six plant taxa are known to be native to Pennsylvania, i.e. part
of the original flora before settlement by Europeans or occurring due to natural means of
dispersal (Rhoads & Klein 1993). This rich flora is the result of a diversity of habitats. For
example, Pennsylvania contains the southern extent of the northern hardwood forest and the
northern extent of the mixed mesophytic forest (Kiichler 1964). Physiographically,
Pennsylvania includes a portion of the Coastal Plain in the southeast and the Central
Lowland in the northwest (Pennsylvania Geological Survey 1962; Fig. 1). Furthermore,
several geological substrates (e.g. limestone, serpentine, shale) support endemic taxa.

The state of Pennsylvania maintains a list of plants of special concern. Of the 2076 native
plants, 15% are officially classified as endangered (E) or threatened (T) and 5% are
extirpated. Another 9% are rare, vulnerable, or tentatively undetermined. Of these, 6 taxa
are listed and 37 are candidates for listing on the Federal Endangered and Threatened
Species List (Pennsylvania Natural Diversity Inventory 1993). Two Pennsylvania plants are
Considered extinct throughout their ranges (Kirkland et al. 1990): Elodea schweinitzii and
Micranthemum micranthemoides. Elodea schweinitzii has only been collected in New York
in 1816 and in Pennsylvania between 1829 and 1832; however, its taxonomic status is
questionable (Cook & Urmi-Kénig 1985). Micranthemum micranthemoides was found in
freshwater tidal shores from New York to Virginia (Fernald 1950).

I evaluated the known occurrences of the E and T taxa in the state with respect to
glaciation and physiographic provinces. Distribution within the state was compared to the
total geographic range of the taxon to determine if its occurrence in Pennsylvania was
endemic, disjunct, or peripheral. The biological aspects examined included life form and
taxonomic relationships. Lastly, I classified the habitat type in which each taxon was found.
Brenner (1986) evaluated the distribution and habitats of the E plants in Pennsylvania;
however, no comparison to this study is made since only 21 taxa were listed as E at that
time and almost all of these are included in this study.

METHODS

The taxa of vascular plants listed as E (227) and T (77) in Pennsylvania constitute the data
Set for this study (Pennsylvania Natural Diversity Inventory 1993; Rhoads ‘& Klein 1993).
Data on distribution in Pennsylvania in relation to Wisconsinan Glaciation (Fig. 1),
Occurrence in physiographic provinces (Fig. 1), life form, and taxonomic category were
tabulated from information in Rhoads & Klein (1993).

Ser nis Sut
‘Present address: School of Biological Sciences, University of Kentucky, Lexington, KY 40506

49

50 BARTONIA

BR

Fig. 1. Physiographic Provinces in Pennsylvania: Central Lowland (CL), Appalachian Plateaus (AP), Valley and
Ridge (VR), Blue Ridge (BR), New England (NE), Piedmont (PD), and Coastal Plain (CP). Shaded area represents
land covered by Wisconsinan Glacier. Modified from Pennsylvania Geological Survey (1962, 1981). Physiographic
boundaries and names are not those currently used by the Pennsylvania Geological Survey.

Data on the total geographic range and habitat were compiled from regional and state
floras, taxonomic monographs, and other research papers. The area of each physiographic
province in Pennsylvania was determined by weight of paper/area relationship (total area =
117,413 km’, United States Geological Survey 1970).

A comparison was made between the distribution of each taxon in Pennsylvania (Rhoads
& Klein 1993) and the total geographic range of the taxon to determine which of the
following categories was most appropriate:

Disjunct. Taxa whose main ranges lie considerably distant from Pennsylvania and do not

occur in an adjacent state.

Peripheral from the north. Taxa that range from Newfoundland to Alaska and south to

ennsylvania (e.g. Carex aurea).

northern Penn |
Peripheral from the east. Taxa that occur from Newfoundland or Long Island south to

Florida along the coast, west to Texas, and north in the interior to Kentucky and
Missouri (e.g. Ptilimnium capillaceum). These taxa have occurrences in eastern
southeastern Pennsylvania. The ranges in the northeast of a few of these taxa extend
inland from the Atlantic Coast to the Great Lakes (e.g. Ammophila breviligulata), and
these taxa have occurrences in northwestern and/or southeastern Pennsylvania

Peripheral from the south. Taxa that occur from southem New Jersey to southern
Illinois to Florida and Texas. Some of these taxa have a restricted range: coastal plain
taxa that reach their northern limit in southeastern Pennsylvania (e.g. Arnica acaulis)
and southern Appalachian taxa that reach their northern limit in southcentral am
southwestern Pennsylvania (e.g. Cymophyllus fraseri).

ENDANGERED AND THREATENED PLANTS IN PENNSYLVANIA 51

Peripheral from the west. Taxa that range from southern Ontario to Montana south to
western Pennsylvania, western Tennessee, and Kansas (e.g. Diarrhena obovata).
Continuous. Taxa whose main ranges include Pennsylvania. Some are northern plants
that occur in the southern Appalachians at high elevations (e.g. Potentilla tridentata).
Habitat types were divided into six categories:
a. Still or flowing water.
b. Wet, open canopy (e.g. tidal to freshwater marshes and shores, boggy swales, bog mats,
fens).
c. Wet, closed canopy (e.g. shrub fens, swamps, alluvial bottomlands).
d. Dry, open canopy (e.g. glades, prairies, dunes and dry shores, fields, savannas).
e. Dry, closed canopy (e.g. shrub thickets, xeric to mesic coniferous and deciduous
forests).
f. Cliffs.
The most frequently mentioned habitat was selected for a taxon for which several habitats
were listed.

RESULTS

GLACIAL RELATIONSHIP. Fifteen percent of the E and T taxa occur only in the glaciated
portions of the state, while 48% are restricted to areas south of the glacial boundary. The
remaining 37% show no relationship to glaciation.

PHYSIOGRAPHIC PROVINCES. Eleven percent of the E and T taxa occur only in the Valley
and Ridge Province, 11% are restricted to the Appalachian Plateaus, and 7% to the Piedmont.
The Coastal Plain and Central Lowland contain 5% and 4%, respectively. None of the E and
T taxa occur exclusively in the Blue Ridge or New England Provinces. On a taxa/area basis,
the Coastal Plain and Central Lowland had the greatest concentration of E and T taxa
restricted to a single province (Table 1).

TABLE 1. Number-area relationships of E and T plant taxa restricted to a single physiographic province in
Pennsylvania.

Physiographic Area* # restricted taxa/area
i (km?) taxa (#/km? x 10°)

Appalachian Plateaus 68985 35 0.51

lue Ridge 803 0 0
Central Lowland 575 11 19.13
C Plai 596 14 23.49
New England 643 0 0
Piedmont 12012 22 1.83
Valley and Ridge 33796 33 0.98

*Determined by weight of paper/area relationship given the area of the state as 117,413 km? (U.S. Geological
Survey 1970).

The remaining 62% of the E and T taxa have occurrences in more than one province, but
none have occurrences in all seven provinces. Most taxa occur in adjacent provinces, with
the Appalachian Plateaus and Valley and Ridge the most common combination. Several
other taxa have occurrences in disjunct provinces; the most extreme being two that occur in
diagonally opposite comers of the state, in the Central Lowland and Coastal Plain.

TOTAL GEOGRAPHIC RANGE RELATIONSHIP. Cerastium arvense var. villosissimum had been

52 BARTONIA

considered endemic to Pennsylvania until its recent discovery in Maryland (Latham 1993).
Three E and T taxa are disjunct: Montia chamissoi (Rhoads et al. 1985), Aster —
(included in A. parviceps—Hart 1990), and Carex geyeri (Keener & Park 1986). An
additional disjunct taxon could be Dodecatheon amethystinum (Klotz & en 1993);
however, most populations in Pennsylvania are too variable to separate reliably from D.
meadia (Klotz & Walck 1993; Rhoads & Klein 1993).

Twenty-eight percent of the E and T taxa are peripheral from the north, 22% from the
south, 22% from the east, and 5% from the west. Twenty percent of the taxa have a
continuous distribution.

LIFE FORM. Seventy-five percent of the E and T taxa are herbaceous polycarpic
perennials, 10% are annuals, and six taxa are biennials/monocarpic perennials. Seven
percent are deciduous shrubs, and three taxa are evergreen shrubs. Seven taxa are deciduous
trees, one taxon is an evergreen tree, five taxa are herbaceous perennial vines, and one taxon
is a woody vine.

TAXONOMIC CATEGORY. Forty-five percent of the E and T taxa are monocotyledons, 51%
are dicotyledons, and 4% are pteridophytes. The E and T taxa belong to 76 families; those
with the greatest number (greater than 7) are Cyperaceae (65), Poaceae (26), Asteraceae (25),
Orchidaceae (14), Potamogetonaceae (10), Ranunculaceae (10), Juncaceae (9), and Rosaceae
(8). Of the 171 genera represented on the E and T list of plants, Carex (32), Eleocharis
(11), Potamogeton (10), and Juncus (9) contain the largest concentrations of E and T taxa.
Thirty-one families and 125 genera are represented by only one taxon.

HABITAT. Thirty-seven percent of the E and T taxa occur in wet, open canopy; 23% in

, Open canopy; 18% in dry, closed canopy; 11% in wet, closed canopy; 8% in still or
flowing water; and 3% on cliffs.

DISCUSSION

Seventy-seven percent of the E and T taxa in Pennsylvania are at the periphery of their
total geographic range. Further, the Wisconsinan Glacial boundary is the limit of the
northern distribution of half of the E and T taxa in Pennsylvania. In North Carolina, U.S.A.
(Bostick 1981); Massachusetts, U.S.A. (Field 1979); Ontario, Canada (Argus 1992); Finland
(Lahti et al. 1991); and Sheffield, England (Hodgson 1986) the majority of rare plants are
at the limits of their range. Argus (1992) has argued from biological, social, and political
views for the preservation of peripheral populations; however, his most compelling reason
is genetic differentiation of peripheral populations.

The Valley and Ridge and Appalachian Plateaus Provinces have the highest concentration
of E and T taxa restricted to a single province. This is probably associated with the larger
area of the provinces (Table 1), greater diversity of habitats, and a greater representation of
distributional patterns. Both provinces have taxa represented in each of the habitat types
outlined above with the majority of taxa in four habitat types (ca. 14 taxa for each except
cliffs and still or flowing water). Each of the other provinces do not have taxa represented
in two or four habitat types, and the majority of taxa only occur in one habitat type (ca. 11
taxa in each province), e.g. wet, open canopy in Coastal Plain and Central Lowland and dry,
open canopy in the Piedmont. Furthermore, the Valley and Ridge and Appalachian Plateaus
Provinces have taxa represented in all distributional categories; whereas, the other provinces
do not have E and T taxa represented in two or three categories. igh Snot found a
similar physiographic pattern for E and T wetland plants in Pennsylvani

ENDANGERED AND THREATENED PLANTS IN PENNSYLVANIA 53

Seventy-seven percent of the E and T taxa are herbaceous polycarpic perennials (including
herbaceous perennial vines), which is not unexpected since the majority of plants in northern
North America (Thieret 1977) and Pennsylvania (Rhoads & Klein 1993) are herbaceous
perennials. Furthermore, 12% are annuals or biennials/monocarpic perennials, 8% are
shrubs, 3% are trees, and one taxon is a woody vine. These percentages about equal the
percentages of the life forms among the native taxa in Pennsylvania (Rhoads & Klein 1993).
A similar pattern has also been found for the rare plants in New England (Field 1979).

In Pennsylvania, 5% of the total native flora are pteridophytes, 1% are gymnosperms,
33% are monocotyledons, and 61% are dicotyledons (Rhoads & Klein 1993). These
percentages are about equal to those for the northeastern United States: 3%, 1%, 28%, and
68%, respectively (Fernald 1950). However, 45% of the E and T taxa are monocotyledons
and 51% are dicotyledons, and none is a gymnosperm. Field (1979) found a similar pattern
for rare plants in New England.

Families with the largest number of E and T taxa are also the largest families in
Pennsylvania and the northeastern United States, namely the Cyperaceae, Poaceae, and
Asteraceae (Fernald 1950; Rhoads & Klein 1993). The Rosaceae, the fourth largest family
in Pennsylvania and the northeastern United States (Fernald 1950; Rhoads & Klein 1993),
is underrepresented in number of E or T taxa. Carex is the largest genus in Pennsylvania
(Rhoads & Klein 1993).

Fifty-seven percent of the E and T taxa in Pennsylvania are associated with water-
dependent habitats and 43% with dry habitats. Further, most of the E and T taxa in
Pennsylvania are associated with open canopy habitats; a result also obtained for New
England (Field 1979). In a recent study, Davis (1993) reported that 57% of the 579 species
listed as "Plants of Special Concern in Pennsylvania" were wetland species. Thus, the
majority of the E and T taxa in Pennsylvania occur in water-dependent habitats, which
Occupy about 2% of the state’s land surface (Tiner 1989).

ACKNOWLEDGMENTS

I thank AE. Schuyler, J. and C. Baskin, and A.F. Rhoads for advice and for reviewing
the manuscript; and K. Fogarasi for computer assistance. I especially thank Virginia D.
Einwechter for her warm hospitality during my stay in Philadelphia.

LITERATURE CITED

ARGUs, G.W. 1992. The Pgs: of rare vascular plants in Ontario and its bearing on plant conservation.
Canad. J. Bot. 70: 469-

BOsTICK, P.E. 1981. hace analysis of the flora of the Carolinas. I. The Carolina spectrum. Castanea 46:
140-153.

BRENNER, FJ. 1986. Habitat preservation and pe for rare and endangered species management. Pages 37-
52 in S.K. Majumdar, F.J. Brenner, and . Rhoads, eds. Endangered and threatened species programs in
Pennsylvania and other states: causes, issues sai management. Pa. Acad. Sci., Easto

Be C.D.K. & K. URMI-KONIG. 1985. A revision of the genus Elodea ce aE Aquat. Bot. 21: 111-

bain A.F. 1993. Rare wetland plants and their habitats in Pennsylvania. Proc. Acad. Nat. Sci. Philadelphia 144:
254-262.

FERNALD, M.L. 1950. Gray’s manual of botany. 8th ed. Amer. Book Co., New York.

FIELD, K.G. 1979. Biological and ecological characteristics of rare ig species in New England: A comparison
with common species. Master’s Thesis, Boston — Bosto:

HART, R. 1990. Aster depauperatus: a midwestern migrant on eastern Fa ela barrens? Bartonia 56: 23-28.

54 BARTONIA

Hopason, J.G. 1986. Commonness and rarity in plants with special reference to the Sheffield flora. Part I. The
identity, distribution and habitat characteristics of the common and rare species. Biol. Conserv. 36: 199-252.
C.S. & M.M. PARK. 1986. An overview of the vascular plant geography of Pennsylvania. Pages 111-144
in S.K. Majumdar, F.J. Brenner, and A.F. Rhoads, eds. Endangered and threatened species programs in
Pennsylvania and other states: causes, issues and management. Pa. Acad. Sci., Easton.
G.L., Jk., AF. RHOADS & K.C. KIM. 1990. Commentary: Perspectives on biodiversity in Pennsylvania
and its maintenance. J. Pa. Acad. Sci. 64: 155-159.
Kotz, L.H. & J.L. WALCK. 1993. Rare vascular plants associated with limestone in southwestern Franklin County,
Pennsylvania. Bartonia, 57 Supplement: 16-41.
KUCHLER, A.W. 1964. Potential natural vegetation of the conterminous United States. Amer. Geogr. Soc. Spec. Publ.
No, 36
E. KEMPPAINEN, A. KURTTO, & P. UOTILA. 1991. Distribution and biological characteristics of threatened
vascular aya in Finland. Biol. Conserv. 55: 299-314.
THAM, R.E. 1993. The serpentine barrens of temperate eastern North America: critical issues in the management
of rare species and communities. Bartonia 57 Supplement: 61-74.
‘AL SURVEY. 1962. Physiographic provinces of Pennsylvania. Pa. Geol. Surv., 4th ser.,
Map 13. heiee ”
. 1981. Glacial deposits of Pennsylvania. Pa. Geol. Surv., 4th ser., Map 59. Harrisburg.
PENNSYLVANIA NATURAL DIVERSITY INVENTORY. 1993. Plants of special concern. The Nature Conservancy,
Western Pennsylvania Conservancy, and Pa. Dept. Environ. Res., Middletown.
RHOADS, A.F. & W.M. KLEIN, JR. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. Amer.
Phil. aig Philadelphia.
_—_—_—, R.H. MELLON, & R.E. meen 1985. Montia chamissoi redi d along the Del River

iJ

in Sa County, Pennsylvania. Bartonia 5
J.W. 1977. Life-forms in the siahioas ee Mich. Bot. 16: 27-3
TINER, R.W., n 1989. Current status and recent ee in thea et s wetlands. Pages 368-378 in S.K.
Majumdar, R.P. Brooks, F.J. Brenner, and R.W. Tiner, Jr., eds. Wetlands ecology and conservation: Emphasis
in Pennsylvania. Pa. Acad. Sci., Easto
UNITED STATES GEOLOGICAL SURVEY. 1970. The national atlas of the United States of America. U.S. Dept. Interior,
Washington.

salah eemmprmeneee=

Bartonia No. 59: 55-70, 1996
The Genus Rhexia in New Jersey

DAVID SNYDER
New Jersey Department of Environmental Protection
Division of Parks and Forestry, Office of Natural Lands Management
New Jersey Natural Heritage Program, CN-404, Trenton, NJ 08625

Rhexia is a small North American genus of 13 recognized taxa (11 species, 2 varieties;
fide Kral & Bostick 1969). The majority of the taxa occur on the Coastal Plain of the
southeastern United States (James 1956; Kral & Bostick 1969), which is the region of
greatest taxonomic complexity. This complexity results from variation within taxa and
hybridization between taxa (James 1956; Kral & Bostick 1969). In New Jersey the genus
is represented by five taxa: Rhexia aristosa, R. x brevibracteata (here named and described),
R. mariana var. mariana, R. mariana var. ventricosa, and R. virginica. The distribution of
the genus within New Jersey mirrors the distribution of the genus as a whole: only one taxon
is widespread in the state, the other taxa are restricted to the Coastal Plain and reach their
greatest abundance and diversity in the southernmost portion of that province. Cape May
is the only county containing all the taxa of specific or subspecific rank.

In New Jersey material, the taxonomical distinctions between the taxa are normally sharp,
and they usually can be distinguished readiliy in both the field and herbarium. When
identification difficulties are encountered, these invariably involve distinguishing R. mariana
var. ventricosa from either R. mariana var. mariana or R. virginica, and R. aristosa from
R. virginica. The difficulty with R. mariana var. ventricosa is largely an herbarium problem,
a result of trying to put a name to incomplete or poorly prepared herbarium material. It can
be rectified by collecting only complete and fully mature specimens which are accompanied
with good field notes. Identification of R. aristosa and R. virginica is apparently
complicated by hybridization of the two taxa, at least there are morphologically intermediate
specimens between these otherwise discrete taxa. Natural hybrids involving various
combinations of R. mariana var. mariana, R. mariana var. ventricosa, and R. virginica, have
been reported (James 1956; Kral & Bostick 1969), but none of these have been confirmed
from New Jersey. Occasionally specimens are encountered that are difficult to place, and
hybridism is a possibility, but this is not easily verified from a study of herbarium
specimens, especially when the collection is represented by a single specimen lacking
detailed field notes. When collecting in this genus, it is important to note if other taxa of
Rhexia are present, and if the specimen collected is representative of the population instead
of an aberrant individual. In cases where morphological variation or intermediacy is
observed, a representative series of specimens should be collected.

The taxonomy and distribution of Rhexia in New Jersey has not been discussed previously
in any significant detail. In the early catalogs and floras of Britton (1889), Stone (1912), and
Taylor (1915), the distribution, abundance, and habitat is summarized for the three taxa then
known to occur in New Jersey (R. aristosa, R. mariana, R. virginica). County distribution
maps and terse comments are presented in Hough (1983) for the same taxa. e purpose
of this paper is to update and clarify the distribution, more fully describe habitat and biology,
and to describe the taxa within the framework of current taxonomic concepts. Specimen

55

56

BARTONIA

citations are primarily based on specimens at CHRB, GH, NY, PH, and my own collection
(DBS). Nomenclature follows Kartesz and Kartesz (1980).

A:

i‘

KEY TO NEW JERSEY RHEXIA

Rootstocks not tubiferous, usually shallowly set, stiff, horizontally spreading, freely
rhizomatous, and wide creeping (if characters of rootstocks are unavailable, go to heading

N WwW
ww

eee ee tee ee ee ee ee Soe ee ee a ee See. oOo CO Re ee Se a Se ee eS Se CE RS 8 8 Le ee Se

. Stem faces unequal, one pair of opposing faces broader and convex, the other, much

narrrower and concave, stems appearing cylindrical at mid stem, flowers pink, to
SAE ED as at AS sigan he 8 Tek g <P 2ceent 1. R. mariana var. mariana
2. Stem faces all approximately equal, stems appearing square at mid stem, flowers
CHOTIIATIN) DORE 10 TARORIA 8 as re oi crsh oa is + cep 2. R. mariana var. ventricosa
Rootstocks tuberiferous (or at least some plants within the population producing tubers),
soft and flexible, rhizomes when produced, soft and flexible ...............-- 3
3. Stem faces unequal, one pair of opposing faces broader and convex, the other
much narrrower and concave, stems appearing cylindrical at mid stem, stem angles
not winged; flowers pink to almost white ....... 1. R. mariana var. mariana

iv)

. Stem faces all approximately equal, stems appearing square at mid stem, angles
4

winged or not winged; flowers (normally) rose to magenta ...........--

4.

os

Inflorescence leafy bracted, the leaves only gradually reduced and not at all
scale-like; sepal lobes linear to narrowly lanceolate, at maturity 4-7 mm long,
tipped with a stout Demon! awn, + 1 mm long; hypanthium rim beset with
coarse, yellowish trichomes 2. 5 mm or more we (often as long as the calyx
lobes); surface of seeds joaginaantty ridged, the concentric ridges relatively
broad, low, smoothish to somewhat verrucosely margined, continuous or a
series of contiguous low domes, the surface appearing merely wrinkled or
pebbled at low magnification

. Inflorescences not leafy bracted, appearing naked, the uppermost bracteal leaves

greatly reduced or scale-like and many times shorter than the mature hypanth-
ium; sepal lobes deltoid to lanceolate, at maturity 1-4.3 mm long, tipped with
a slender, usually gland-tipped hair 0.5-1 mm long; hypanthium rim with or
without coarser yellowish trichomes, when present 2.0 mm or less long and
shorter than the sepal lobes; surface of seeds muricate or papillose, with or
without low, smoothish, domes or ridges, the processes forming nearly
concentric, longitudinal, relatively narrow and high rows or ridges, surface
appearing more granular than wrinkled at low magnificaition .......-- 5
. Surface of seeds finely papillose, the projections forming, neat, low,
closely concentric ridges, without a well developed dorsal crest
Cha. Seek ee ks cu Cees ee. 2. R. mariana var. ventricosa
Surface of seeds muricate or coarsely papillose, the ridges more irregular
and less closely concentric, with or without a well developed weed
CEO eek ay ee ee pede ee Oe ee ee ey
6. Stems conspicuously winged (wings usually more than 0.25 iiss
sepal lobes short, mostly 2.5 mm or under, deltoid or lanceolate;
hypanthium rim without coarse yellowish trichomes 3. R. virginica
Stems wingless or only inconspicuously winged (wings mostly 0.25

an

a

RHEXIA IN NEW JERSEY 57

mm or less); sepal lobes longer, 2.1-4.5 mm; hypanthium rim beset
with coarse, yellowish, trichomes ....... 4. R. x brevibracteata

1. Rhexia mariana L. var. mariana.

This is the only taxon in Rhexia occurring in New Jersey that does not have a square
stem. Although the stem is not truly cylindrical, the combination of concave and convex
opposing faces give it the appearance of being round, and it rolls easily when pushed lightly
between thumb and forefinger. When in flower, it is easily recognized by its pale pink
(rarely darker) to nearly white petals, all other taxa being rose colored, varying to crimson
or magenta. Although the taxon is variable in a number characters (Fernald Ju James
1956; Kral & Bostick 1969) throughout its range, New Jersey material is fairly uniform,
especially in leaf shape and vestiture. Leaves are predominantly elliptic, "rae
narrrowly so, and hirsute above and on the veins below. The stem is also hirsute, giving the
plant its characteristic bristly appearance.

It is common, and occurs on the Coastal Plain from Massachusetts south to southern
Florida, then west to Texas. In the interior it occurs from Georgia to Virginia, and northwest
to Missouri, Illinois, and Indiana (James 1956; Kral & Bostick 1969).

Rhexia mariana var. mariana is frequent in the southern portion of New Jersey, and after
R. virginica, it is the secondmost common Rhexia occurring in state. It has been collected

om all Coastal Plain counties, except for the two northernmost, Mercer and Middlesex.
Although widespread throughout this region, it tends to be somewhat local in its distribution,
reaching its greatest abundance in Atlantic and Cape May counties and in portions of the
counties bordering the Delaware bayshore. Since it spreads by a system of horizontally
5 rhizomatous rootstocks, it is usually abundant where found and forms extensive
colonie

It i. in a variety of mostly successional habitats which are at least seasonally wet and
is commonly found in moist sandy or peaty thickets, moist shallow depressions and
shoulders of roadsides and trails through pitch pine lowland forests, meadows, moist fields,
upper peaty shores of ponds, and occasionally sphagnous wetlands. Its stoloniferous habit
makes it weedy, and therefore a good colonizer, and it can be found abundantly in railroad
and powerline right-of-ways, bottoms of inactive sand pits, banks of ditches, and other
disturbed habitats. It is the least likely taxon in Rhexia to be found in permanently saturated
soils or standing water in New Jersey.

Flowering is from early July (rarely earlier) into mid October. The foliage turns bright
Orange or red-orange in the Autumn, - the leaves persisting late into the season, and
usually not falling until after a hard fros

This is a distinctive species and at identifiable in the field and herbarium, unless
Specimens are incomplete or poorly prepared. In which case, specimens may be confused

with R. mariana var. ventricosa.
VE NEW JERSEY SPECIMENS EXAMINED. Atlantic Co.: Moist open sandy soil near Great Egg

S of Clementon, 3 Sep 1939, L. P. Hynes 1440, PH. Cape May Co.: Swale along Crooked Creek, E of Cape May
Court House, 11 Sep 1938, B. Long 53070, PH. Cumberland Co.: Old cart road through woods SE of Haleyville,
10 Aug 1933, B. Long 41496, PH. Gloucester Co.: Edge of low sandy thicket ca 1 mi NW of Bridgeport, 13 Feb
1949, B. Long 68712, PH, Monmouth Co.: 1.5 mi W of Fort Plains, 4 Aug 1955, V. L. Frazee s. n., PH. Ocean
Co.: Wet meadows very near the salt marshes, Coxs, E of Tuckerton RR and the Shore Road, 11 Oct 1910, B. Long
5480, PH. Salem Co.: Sandy peaty border of pond-hole, 1.5 mi NNE of Olivet, 6 August 1935, B. Long 47092, PH.

58 BARTONIA

2. Rhexia mariana L. var. ventricosa (Fern. & Grisc.) Kral & Bostick

This taxon possesses the unique combination of square stems and stoutish, horizontally
creeping and freely rhizomatous rootstocks that are not tubiferous. Only R. mariana var.
mariana has similar rootstocks but its stem is distinctly subcylindrical. The remaining New
Jersey taxa are all square stemmed with tubiferous rootstocks. While R. mariana var.
ventricosa never produces tubers, it can not be said that all plants within populations of R.
virginica or R. aristosa, will have tubers. However, at least some plants within the
population will be tuberiferous, unless all are very young. When tubers are not present (or
not collected) the rootstocks of R. virginica and R. aristosa usually differ from R. mariana
var. ventricosa by being softer (spongy thickened when growing in standing water), more
clustered, commonly descending, and therefore more deeply set. In contrast, those of R.
mariana var. ventricosa are more woody and therefore stouter and less flexible; they spread
horizontally just below the surface of the soil and are freely rhizomatous. Rhexia virginica
and R. aristosa occasionally produce long creeping roots and even rhizomes, but these are
never woody or stout. Surface sculpturing of mature seeds of R. mariana var. ventricosa is
distinctive and is usually diagnostic (see discussion under R. virginica). The seeds are finely
papillose with concentric rows of closely and evenly spaced papillae, and lack a well
developed dorsal crest. Seeds of all other taxa are either muricate, coarsely and irregularly
papillose, verrucose, or smooth ridged. In absence of rootstocks and mature seeds, it is
sometimes difficult to distinguish R. mariana var. ventricosa from R. virginica. In such
situations, the two taxa can usually (but not infallibly) be separated by the lack of a well
developed wing on the stem of R. mariana var. ventricosa, and by the length of the
hypanthium neck relative to the length of its body: in ventricosa the neck is equal to or
longer then the body; in virginica the neck is predominantly shorter then the body (James
1956; Kral & Bostick 1969). If care is taken in collecting complete specimens with mature
fruit, and if the specimens are then prepared so that stem characteristics are not lost in
pressing (i.e. using foam rubber pads between the blotters and cardboard ventilators), 2.
mariana var. ventricosa can be easily and reliably identified.

Somewhat local in distribution, R. mariana var. ventricosa is restricted to the eastern
United States where it occurs on the Coastal Plain from southern New Jersey to South
Carolina (James 1956; Kral & Bostick 1969).

In New Jersey, R. mariana var. ventricosa is apparently restricted to the Cape May
peninsula where it is documented from five extant localities and two historical sites from
Cape May north to Cape May Court House. Although recently rediscovered in New Jersey
(Snyder 1994), preliminary field investigations suggest that it may not be exceptionally rare,
but perhaps overlooked and undercollected. Rhexia mariana var. ventricosa was first
reported as occurring in New Jersey by James (1956) based on specimens collected from
Cape May by Mackenzie in 1909 (Snyder 1994). In 1991, Andy Windisch and I discovered
a large population just north of Kay’s Pond (Snyder 1994). Subsequently, I located one
additional herbarium specimen (Brown s. n. CHRB) and four additional occurrences in the
field. Whether this taxon is restricted to Cape May County or more widespread remains to
be seen after more searching. Unless specifically searched for, it is easy to pass over R.
mariana var. ventricosa for R. virginica. This may be why the species has gone largely
unrecognized in the state until now. It may be too, that this is example of southern taxon
expanding its range northward.

All documented extant occurrences of this variety are in sandy soils of open, or somewhat
thicket-covered, inactive sand pits located within oak-pine or oak dominated forests. These

RHEXIA IN NEW JERSEY 59

pits all have seasonally fluctuating water levels, and in this respect function much like
Coastal Plain intermittent ponds. Some characteristic associated plant species include
Eleocharis quadrangulata, Spiranthes odorata, S. vernalis, Rotala ramosior, and Hedyotis
uniflora. Like R. mariana var. mariana, var. ventricosa tends to be weedy, and successfully
exploits disturbed habitats. Apparently only Mackenzie’s 1909 specimens (4438 NY) labeled
"moist meadow" were collected in what could be a natural habitat.

Flowering occurs from mid July into mid October. The foliage turns a brilliant yellow-
orange in the Autumn, with the leaves persisting late into the season, and usually not falling
until after a hard frost.

This taxon has been recognized at the specific level by Fernald (1935; 1950) and James
(1956), reduced to subspecific rank by Kral & Bostick (1969), and most recently, subsumed
under R. interior by Cronquist (Gleason and Cronquist 1991). Although a distinctive and
easily recognized taxon, R. mariana var. ventricosa suggests an intermediate between R.
mariana var. mariana and R. virginica. It has the stout, horizontally spreading rhizomatous
rootstocks and the long necked hypanthia of R. mariana var. mariana but the square stems
and the generally more ovate shape of its leaves are characters of R. virginica. This
intermediacy led James (1956) to speculate that R. mariana var. ventricosa may be of hybrid
origin with R. mariana var. mariana and R. virginica the putative parents. However, Kral
& Bostick (1969) noted that the hybrid between these taxa was quite different. Further,
chromagraphic, cytologic, and morphologic studies conducted by these authors showed that
ventricosa bore a much closer relationship to typical mariana then it did to R. virginica, thus
supporting the reduction of ventricosa to varietal level under mariana (Kral & Bostick 1969).
Although artificial hybrids between R. mariana var. ventricosa and both R. mariana var.
mariana and R. virginica have been produced experimentally (Kral & Bostick 1969),
naturally occurring hybrids between these taxa apparently have not been observed (James
1956; Kral & Bostick 1969)

REPRESENTATIVE NEW JERSEY SPECIMENS EXAMINED. Cape May Co.: Moist meadow, Cape May, 18 Sep 1909,
- si Mackenzie 4438, NY; Damp soil, Cold Spring, 25 Aug 1936, O. H. Brown s. n., CHRB; Extensive colony

wing in moist sand of inactive sand pit, Pierces, 0.4 mi NW of Kay Pond, 15 Oct 1991, D. Snyder and A.
Poe CHRB, DBS, PH, VDB; Common in wet sandy soil in thickets and open, sedgy bottoms of inactive sand
Pit, Higbee Beach, 0.4 mi NE of Davey’s Lake, 29 July 1992, D. Snyder, CHRB, DBS; Scattered colonies in moist
to wet sand in thickets and more open areas of excavated portions dredge spoil bank, Petticoat Bridge, just SE of
= ~ May, 29 Jul 1992 and also 5 Oct 1992, D. Snyder, CHRB, DBS; Large, scattered colonies in moist to

andy bottoms of a series of inactive sand and gravel pits, just NNW of Cold Spring, 29 Jul 1992 and also
: Oct oh, D. Snyder, DBS, PH; Extensive colony in moist, sandy edge of thicket in partially regraded and filled
sand pit along eastern edge of Goshen Sports Complex, ca 1 mi N of Cape May Court House, 30 July 1992, D.
Snyder, DBS.

3. Rhexia virginica E

This species is easily distinguished from all other New Jersey taxa by its conspicuously
wing-angled stems. The wings are usually prominent, but they can be as narrow as 0.
(James 1956). Wings near the minimal width are often obscured in pressing, so it best to
observe this character on living specimens. In general aspect, R. virginica closely resembles

- mariana var. ventricosa, however, the rootstocks of the former are not tuberiferous, an
those of R. virginica are commonly tuberously thickened. Since the tubers are

, Many herbarium specimens lack tubers. Additionally, the tuberous hackles 3 are

relatively short-lived and eventually degenerate and may persist for as little as one year
(Posluszny et al. 1984). A number plants may have to be examined before the tuberiferous
rootstock is observed. The seeds of R. virginica are usually sharply distinct from R. mariana

60 BARTONIA

var. ventricosa, the surface being irregularly ridged, the processes being muricate or coarsely
papillose, with those of the dorsal crest being coarser and better developed, in contrast to the
uncrested, fine, and neatly rowed papilliae of R. mariana var. ventricosa. Seed morphology
of R. virginica is quite variable, and while most fit the previous description, specimens vary
from nearly tuberculate to merely papillose. Exceptional specimens that I collected from the
edge of a water filled sand pit near Big Goose Pond, Egg Harbor City, Atlantic County (7
Oct 1993, Snyder DBS), had seeds which were finely papillose with the processes of the
dorsal crest only slightly more coarse, and essentially indistinguishable from seeds of R.
mariana var. ventricosa. However, the roots of these specimens were tuberously thickened
and the necks of the hypanthia were distinctly shorter than their bodies, both characteristics
of R. virginica.

This is the most widespread and most common species of Rhexia. It occurs from southern
Canada south to northern Florida, and west to Missouri, lowa, and eastern Texas (James
1956; Kral & Bostick 1969).

Rhexia virginica is also the most abundant and most widely distributed taxa occurring in
New Jersey. It reaches its greatest abundance in the Coastal Plain, especially within the Pine
Barrens, and occurs sporadically northward, mostly in the Piedmont, to Hunterdon, Morris,
Passaic, and Bergen counties.

It grows in a diversity of open, mostly acidic, wetland habitats including pine barren
savannas, shores and shallow bottoms of Coastal Plain intermittent ponds, swampy woods,
peaty or sandy thickets, seepage areas along small streams, edges of Atlantic white cedar
swamps, and meadows. It is frequently associated with sphagnum mosses. Although this
species is not especially weedy, it occurs abundantly in disturbed or successional habitats
such as abandoned cranberry bogs, ditches, fields, inactive sand and gravel pits, and also
powerline, railroad, and road right-of-ways. In northern New Jersey it has been collected
from seasonally moist, open thickets over trap rock. When growing in permanently saturated
soils or in standing water, the stems and rootstocks become spongy-thickened.

Flowering is from early July into late September, rarely later. The foliage turns deep red
to red-purple in the Autumn. It is the first Rhexia to loose its leaves in the Autumn, which
commonly drop shortly after flowering has ceased, and seeds have matured, which is often
well before the onset of a hard frost. From late September through mid October, when the
foliage of other taxa of Rhexia is green or only beginning to reveal Autumnal colors, plants
of co virginica are conspicuous with their often bare stems and dehiscent capsules (Snyder
1994).

My treatment of R. virginica is sensu James’ (1956) R. virginica var. virginica and
excludes the var. purshii (Spreng.) James. The var. purshii is a southern element, perhaps
ecotypic (R. Kral pers. comm.), that occurs from North Carolina south to Florida, then west
to Mississippi (James 1956). It is distinguished from the typical variety by its nearly
glabrous stems, narrower and typically lanceolate leaves, longer sepal lobes (up to 4 mm),
and smaller, usually glabrous hypanthia. In its extreme, it is distinctive, but intermediates
are frequent, and for this reason the variety is usually subsumed under R. virginica (Kral &
Bostick 1969; Kral pers. comm.). My reason for following James’ circumspection is strictly
one of convenience. As noted by James (1956), some of the morphological characters 0
var. purshii approach those of R. aristosa. Since var. purshii has not been documented for
New Jersey, I feel that it is unnecessary to complicate key construction by including the full
spectrum of rangewide variations of R. virginica.

Experimentally produced, as well as naturally occurring hybrids between both varieties

RHEXIA IN NEW JERSEY 61

of R. mariana and R. virginica have been reported (James 1956; Kral & Bostick 1969), but
these hybrids have not been documented for New Jersey. Fertile hybrids between R. aristosa
and R. virginica have been produced experimentally (Kral & Bostick 1969) and specimens
of what have been determined as naturally occurring hybrids between these taxa have been
reported (James 1956; R. Kral pers. comm.). See comments under R. aristosa for further
discussion.

REPRESENTATIVE NEW JERSEY SPECIMENS EXAMINED. Atlantic Co.: Common on black sphagnous ps of large
clearing in cedar-birch-cranberry, petals lavender-rose, 6 mi NW of Tuckahoe, 10 September 1964, R. Kral 22620,
PH. Bergen Co.: Moonachie, near Losen Slofe Branch of Hackensack River, 22 JUL 1970, W. See 1593, PH.
Burlington Co.: Bog ca 3 mi S of Cookstown, 15 Aug 1950, B. Long 72007, PH. Camden Co.: Open swamp along
Pump Branch of Albertson Brook, Ancora, 28 Aug 1926, J. W. Adams 504, PH. Cape May Co.: Swamp ca | mi
SW of Marshallville, 29 Aug 1946, W. M. Benner 1024], PH. Cumberland Co.: Dam breast of old cranberry bog
on Panther Rd., ca 1 mi SE of Coopers Mill, 18 May 1946, B. Long 64879, PH. Gloucester Co.: Sandy, peaty
margin of Sand. on Little Ease Run, Franklinville Lake, eee 25 Jul 1940, B. Long 54868, PH. Haatiiun
Co.: Lambertville, 1887, J. H. Butterfoss s. ., PH. Mercer Co.: Princeton Junction, 19 Aug 1894, T. Seal s. n., PH.
Middlesex Co.: Along path in moist woods betwee five Avenue and Farrington Lake, S of Church Lane, Patrick’s
Corner, 11 Aug 1983, J. S. mete et al. s. n., CHRB. Monmouth Co.: Swampy woods, | mi N of Fort inge 1
Aug 1955, V. L. Frazee s. n., PH. Morris Co.: Sandy bog, MT. Tabor, 19 Aug 1920, K. K. Mackenzie s
Ocean Co.: i oy bog; Warren Grove, | Aug 1941, R. H. Pohl 3227, PH. Passaic Co.: Moist, open eck
on trap rock hillside, Garrett Mountain Reservation, 13 Aug 1992, D. Snyder, DBS. Salem Co.: Sphagnous border
of pond-hole along intermittent streamlet, ca 1 mi E of Union Grove, 4 Sep 1937, B. Long 51453, PH. Somerset

6.: Local on RR track, W of Middlebush, 8 Jul 1945, M4. A. Chrysler s. n., CHRB. Union Co.: Terminal moraine
behind Jefferson School, Summit, 6 Aug 1936, J. Kezer s. n., NY

4. Rhexia aristosa Britt.

Rhexia aristosa is characterized by its square, wingless, or essentially wingless stems with
glabrous internodes; its nearly glabrous, linear-lanceolate to lanceolate leaves commonly
under 6 mm wide (rarely more than 8 mm) with prominent veins below, strongly ascending
to almost erect, and gradually reduced in the inflorescence; the bracts often as long or longer
than the mature hypanthia; its long (3-7 mm), linear to lanceolate, stiffly awn-tipped sepal
lobes; the long (2.5-4.5 mm), coarse, yellowish, eglandular trichomes congested at the rims
of the hypanthia; and the seeds with low, more-or-less concentrically arranged, smooth to
verrucosely margined ridges or domes, which give them a wrinkled or brain-like appearance.
Kral & Bostick (1969) and some subsequent authors (Godfrey & Wooten 1981; Gleason &
Cronquist 1991) describe the flowers as being "dull lavender" in contrast to the "bright rose-
lavender" color of flowers of R. virginica. 1 have found no significant color difference in
Side-by-side living plants of R. aristosa and R. virginica in New Jersey. 1 would describe
both as rose or magenta. Darker or lighter color forms are occasionally encountered in both
species.

Rhexia aristosa is extremely local to rare throughout its range. It is restricted to the
Coastal Plain in North Carolina, South Carolina, Georgia, and Alabama, with disjunct
occurrences in Delaware and New Jersey' (The Nature Conservancy Biological and
Conservation Database). It reaches its greatest abundance in North Carolina and Georgia (R.
Kral, A. Weakley pers. comm.) The species is currently under federal review as being
Possibly threatened or endangered in the United States (U. S. Fish and Wildlife Service
1990)

'Stalter & & Lamont (1990) report collecting a eee of R. aristosa from Assateague Island, Virginia. However
this specimen has been reidentified as R. v: irginica (Frank Hirst pers. comm.). Beaven & Oosting (1939) report

4S Occurring in the "Burned Area” which is wholly within Delaware (Bill aoe pers. comm.) and is currently
known as the Burnt Swamp. There appears to be no documentation supporting a Maryland occurrence of R.
aristosa.

62 BARTONIA

This is the rarest taxon of Rhexia occurring in New Jersey. It is currently documented
from three extant occurrences: two contiguous ponds in north central Atlantic County and
one site in northern Cape May County. Two additional historical sites are documented from
the same general area that the extant Atlantic County occurrences are found. One of these
historical sites is the type locality where plants were discovered in 1888 by J. C. Gifford and
E. H. Kilmer (Britton 1890). Neither the type (NY) nor the isotypes (GH, NY) contain any
specific information on their labels that pinpoints the type locality, and its exact location
remains uncertain. Type specimens are labeled "Egg Harbor City"? along with subsequent
collections by the original collectors except for one labeled "near Egg Harbor" (Gifford s.
n. PH). Attached to the type specimen is a letter written by E. H. Kilmer to N. L. Britton
dated 11 February 1889, which states: "We were at Egg Harbor City (seven miles from
[Mays Landing]) and the Rhexias attracted our attention as we were walking on the
embankment of the railroad. The plants were growing in a narrow ditch-like excavation
three feet wide." Between 1888 and 1898, numerous collections were made from "Egg
Harbor City," all presumably from the type locality. A second, and apparently new locality,
was discovered in 1898 at Cologne, about 2.5 miles southwest of Egg Harbor City (Saunders
s. n. PH; Stone 1912). Despite diligent field searching (Fables 1962; Hirst & Hirst 1962),
R. aristosa went undetected for nearly sixty years, until a third locality was discovered in
1957 by Frank and Robert Hirst at Little Goose Pond, southwest of Egg Harbor City (Hirst
& Hirst 1962). Little Goose Pond is a natural pond located about a mile southwest of the
railroad, and is clearly not the type locality. However, the outlet stream to the pond is
crossed by the railroad just to the northwest of Egg Harbor City, and it is conceivable that
seeds from Little Goose could have been dispersed into drainage ditches along the railroad.
If this was source of the plants discovered at the type locality, it might provide an
explanation as to why R. aristosa can no longer be found in the drainage ditches along the
railroad: the habitat was suitable to briefly support a population, but conditions were
inadequate to sustain the population. Rhexia aristosa was next collected in 1959 (R. Hirst
s. n. PH) from a shallow, water filled borrow pit located about 1.5 mile southeast of Egg
Harbor City. Although the specimen is labeled "Cologne," it was collected about a mile
northwest of that community (F. Hirst pers. comm.). It can not be determined whether this
is the same site as C. F. Saunder’s 1898 collection, or possibly even the same as the type
locality. Repeated searches of the site has failed to relocate the species. In 1982, I found
a small population in Big Goose Pond. Big Goose, which is located immediately adjacent
to Little Goose, had been thoroughly searched by Frank and Robert Hirst in the late 1950s
for R. aristosa (F. Hirst pers. comm.), as well as by many others in the intervening years.
The species’ seemingly abrupt appearance in recent years at Big Goose may be explained
by either the presence of dormant seeds in the seed bank, or recently spread seeds by
waterfowl, deer, box turtles (present by the dozens buried in the mud one dry summer) oF

*Both James (1956) and Kral & Bostick (1969) cite the type locality as "May’s Landing, Egg Harbor City, Atlantic

" The two place names refer to distinct localities separated by about seven miles. The error of citation is no
doubt a result of a newspaper article from the 16 Feb. 1889 edition of the May’s Landing Record entitled, "May's
Landing Adds a New Flower to the Flora of the State” which is attached to the type of Rhexia aristosa. The article
is written by J. E. Peters who sent the original specimens collected by Kilmer and Gifford to Britton. Peters, who
lived at May’s Landing, was being overly parochial in his use of "May’s Landing” as the name of the collection
locality and he does not mention Egg Harbor City anywhere in the article. Kilmer in his letter to Britton clearly
states that the specimens came from Egg Harbor City.

RHEXIA IN NEW JERSEY 63

even accidentally introduced as seed stuck on the boots of one of the many botanists that
have since visited these ponds. In 1982 this population consisted of a few, scattered plants,
but in 1992 I counted ca. 1400 plants. No new locations were documented until 1991 when
I discovered a small population in an inactive sand pit near Petersburg, Cape May County.}

In New Jersey, Rhexia aristosa is restricted to a very narrow range of habitat types.
Habitats of extant occurrences are similar in two respects: all are in wetlands characterized
by fluctuating water levels and located in either pitch pine lowland or pine-oak forests within
the Pine Barrens. The habitats can be further divided into natural systems and those created
or maintained anthropogenically.

Two of the three extant occurrences are in Coastal Plain intermittent ponds. Big and Little
Goose ponds are the largest natural ponds occurring in the Pine Barrens. The largest of the
two is Big Goose Pond at ca. 22 acres. Water source is either from rainfall or ground water
seepage, and drainage is a combination of external (ephemeral streams) and internal
(percolation and evaporation). Despite their acreage, the ponds are shallow bottomed, and
during years of average precipitation, they largely dry out by mid summer. Surface soils
range from sand or sandy loam with no, or only a shallow layer of organic matter, and are
underlain by gravelly sand. When dry, the bottoms of the ponds are covered by herbaceous
vegetation, with species of grass, sedge, and rush dominating. Characteristic herbaceous
species occurring in one or both of these ponds include Carex striata, Cladium mariscoides,
Rhynchospora fusca, R. inundata, Scleria reticularis, Panicum hemitomon, P. verucosum,
Juncus pelocarpus, Xyris difformis, X. smalliana, Lachnanthes caroliniana, Eriocaulon
aquaticum, Sagittaria teres, Drosera intermedia, Utricularia juncea, U. olivacea, and Lobelia
boykinii.

Rhexia aristosa has been documented from two types of artificially created habitats. As
Previously noted, the type specimens were collected from a drainage ditch along a railroad.
The Hirst specimen from Cologne and the recently discovered Petersburg population are
from borrow pits. In structure and function these pits are much like natural intermittent
ponds. Both of these pits are shallow excavations in sandy soils with fluctuating water
levels. Water levels are determined by rainfall and ground water seepage. Drainage is
internal or through evaporation. These two borrow pits, as well as numerous other similar
ones that occur throughout New Jersey’s Coastal Plain, support floras that are similar to
those found in their natural counterparts.

There is no evidence that R. aristosa is weedy, nor does it seem to readily colonize
disturbed or artificially created habitat, or for that matter, natural ponds. Although seed
Production in R. aristosa is prolific, the species is extremely rare throughout most of its
range. This suggests that either (or more likely both) the methods of seed dispersal or the

‘Previous reports for Cape May County (Stone 1912; Fairbrothers and Hough 1973) prove to be erroneous.
Specimens collected by Gerry Moore (15 Jul 1992 835; 19 Jul 1992 863 PH) from a Coastal Plain intermittent pond
SSE of Hunters Mill, Cape May County, New Jersey were initially identified as R. aristosa by R. Kral (G. Moore
Pers. comm.). Specimens that I collected from this population (26 Oct 1992 s. n. VDB), were identified by R. Kral

R . mariana var. ventricosa. Although these late season specimens were rather poor (all stem leaves had dropped

More thorough study of this population is needed to resolve its identification.

64 BARTONIA

availability of suitable habitat are the critical factors limiting the distribution of R. aristosa.
Seeds could be dispersed into contiguous wetland habitats by water movement, but long
range dispersal into noncontiguous habitats presumably is achieved through the movement
of animals, most likely by waterfowl, which would likely favor natural ponds or other large
water filled depressions such as sand pits.

Rhexia aristosa’s occurrence in disturbed habitats is infrequent throughout its range and
such occurrences may be short lived. Rather than colonizing disturbed habitats, it is possible
that disturbances such as forrest clearing, ditching and sand mining break the dormancy of
seeds contained in the seed bank, where perhaps, they are relics from a time when more
favorable conditions prevailed. In Delaware, a population of R. aristosa was recently
discovered in a area that had been clear cut, ditched, and planted with Pinus taeda seedlings
(B. McAvoy pers. comm.). This suggests that the species was present in the seed bank prior
to the clearing. A study of aerial photography dating back to the 1930's, indicates that a
number of depressional ponds may have existed at this site which were eventually succeeded
by woody vegetation (B. McAvoy pers. comm.).

Rhexia aristosa, \ike all the taxa of Rhexia occurring in New Jersey, is a perennial.
However, under certain conditions or in certain habitats, the species may be very short-lived
or be annual. Reproduction is both sexual and asexual. Asexual reproduction is through
underground tubers and the very occasional rhizome. Like those of Rhexia virginica, the
tubers of R. aristosa are likely short-lived. Under adverse environmental conditions (i.e.
successive years of exceptionally high or low water levels) it is possible that the entire
rootstock may succumb. Like many species adapted to cyclical or dynamic habitats, seed
banking is key to long term survival. Seed production in R. aristosa is copious with the
seeds capable of persisting in the seed bank for an undetermined number of years. Sutter
and Boyer (unpublished) found R. aristosa to be a prolific seed banker: one soil sample
contained 6,872 seeds per square meter. They also found that seeds germinated only when
the substrate remained moist or was subjected to simulated drawdown. Seeds grown in
permanently saturated or inundated substrates did not germinate.

These data support the inferred conclusions of many observers familiar with the yearly
fluctuation of population sizes of R. aristosa. My five year observations of the Little Goose
Pond population provide a dramatic example. 1989 and 1990 were years of exceptionally
high rainfalls in New Jersey. Only a few plants were found in 1989 and none were located
in 1990. By July, 1991, with a return of more typical pattern of rainfall, thousands of
seedling plants were observed in dry, or nearly dry portions of the pond. In late August of
1992, the pond was almost completely dry, and I estimated 38,000 plants. Most were small,
unbranched or with a few branches. Only a small percentage matured sufficiently to produce
flowers. In the summer of 1993 much of New Jersey was experiencing near drought
conditions. At the height of a record breaking week of temperatures near or above 100
degrees, the mortality rate for Rhexia aristosa was high: few seedlings survived and
thousands of the mature plants were blackened or wilted. Few flowering plants were
observed, but the few that were, were restricted to portions of the pond where standing water
remained. Rain showers, although infrequent or brief during much of the summer, became
more regular in September and October. Despite the earlier high mortality, more than 50,000
plants were estimated in October, with a high percentage of these being large, flowering
plants. Countless seedlings, apparently recently recruited, and young plants were also
present.

Flowering is from mid July to late September, occasionally a little earlier, or later,

RHEXIA IN NEW JERSEY 65

depending on water levels. The foliage turns dark purple-brown, appearing almost black,
and frequently tinged with deep red, especially along veins, stems, and hypanthia. The
purple-brown pigmentation often begins to show in late July as blotchy, dark spots, the
coloration remaining uneven throughout the remainder of the season. When growing in deep
water, stem leaves are commonly deciduous, otherwise the leaves persist until frost.

As acknowledged by James (1956) and Kral & Bostick (1969), Rhexia aristosa is one of
the most distinctive taxa in the genus. In morphological characters, R. aristosa is most
similar to R. virginica, and ordinarily the two are easily differentiated. Rhexia virginica is
somewhat variable (more so in the southern part of its range than in New Jersey material)
and some specimens superficially suggest R. aristosa. However, the narrow leaves, leafy
inflorescences, coarse trichomes on the rims of the hypanthia, and very ene; attenuated sepal
lobes tipped with stout, eglandular awns are diagnostic for R. aristosa.* Fertile hybrids
between R. aristosa and R. virginica have been artificially produced by Kral & Bostick
a ea James (1956) noted that among specimens of R. aristosa and R. virginica that he

ined, were a number of specimens collected from several different localities, that were
fines intermediate between the two." I have examined New Jersey material so
annotated by James and others, as well as living material, and discuss them below as Rhexia
x brevibracteata.

REPRESENTATIVE NEW JERSEY SPECIMENS EXAMINED. Atlantic Co.: Wet place, Egg Harbor City, 1 Aug 1888,
J. E. Peters s. n., (type and isotype) NY°; Near Egg Harbor City, 15 September 1896, G. M. Beringer s. n., NY;
Egg Harbor City, 17 Aug 1897, B. Heritage s. n., PH; Egg Harbor City, 17 Aug 1898, C. D. Lippencott s. n., PH;
Little Goose Pond, 20 Jul 1957, Bob Hirst s. n., PH; Shallow water, Little Goose Pond, 6 Aug 1960, A. J. Hoiberg
1013, CHRB; Shallow water, Little Goose Pond, Egg Harbor City, 20 Aug 1979, D. B. Snyder 599-2RU, CHRB;
Abundant (ca. 38,000 plants) on nearly dry bottom of a large intermittent Coastal Plain pond, Little Goose Pond,
Egg Harbor City, 31 Aug 1992, D. Snyder 599-5, DBS; Thousands (ca. 50,000) of plants scattered throughout dry,
to nearly dry bottomed Coastal Plain intermittent pond, Little Goose Pond, Egg Harbor City, 7 Oct 1993, D. Snyder
599-7RU, CHRB, DBS; 1,000 or so plants growing on dry bottom of large intermittent Coastal Plain pond, SW bay
of Big Goose Pond, Egg Harbor City, 31 Aug 1992, D. Snyder 599-15, DBS; Berlin Ave. and RR, Cologne, 29 Jul
1959, B. Hirst s. n., PH. Cape May Co.: Small colony on nearly dry bottom of shallow sand pit, ca. | mi W of
Petersburg, 18 July 1991, D. Snyder 599-18; Small colony on nearly dry bottom of shallow sand pit, ca. | mi W
of Petersburg, 29 Jul 1991, G. Cavileer, K. Seager, and D. Snyder 599-20 DBS; Small colony on dried bottom of
shallow sand pit, ca. 1 mi. W of Petersburg, 10 Sept 1993, D. Snyder 599-25RU, CHRB.

5. Rhexia x brevibracteata Snyder, hybr. nov.

Rhexia aristosa Britt. x Rhexia virginica L.

Hybrida naturalis inter parentes media.

Erect perennial herb to 72 cm from tuberiferous rootstocks, usually without horizontal
stoloniferous rhizomes. Rhizomes when produced, slender, short or long creeping, but soft
and flexible. Roots and stems frequently spongy thickened when growing in water for
Prolonged periods of time. Stems four angled, all sides being nearly equal in width, the
angles wingless or with narrow wings, internodes nearly glabrous except for a very few

"The tich trichomes on the hypanthia and sepal lobes of both R. aristosa and R. virginica become brittle upon drying

dare often broken or missing on specimens. This is also true for specimens collected very late in the season
asin have weathered or dehiscent fruit capsules. The nature of the trichomes, especially the presence or absence
of mo is best observed on living specimens or on dried specimens having buds, flowers, and reasonably mature

Capsul
‘Although these specimens were collected by E. H. Kilmer and J. C. Gifford (Britton 1890), only J. E. Peters’ name
appears on the preprinted label.

66 BARTONIA

spreading, mostly glandular hairs, nodes with conspicuous tufts of coarse hairs, these either
glandular or not, few to bushy branched. Leaves predominately lanceolate, occasionally
broadly so, sometimes ovate on lateral branches, 2.2-4.3 cm long, 0.5-1.9 cm wide (mostly
1 cm or more), sessile, apex obtuse to acute and mostly mucronately tipped, adaxial surfaces
sparsely hirsute to nearly glabrous, abaxial side prominently tripled-nerved, the nerves
yellowish, often with an additional pair of less conspicuous nerves along the margins,
secondary nerves none to present, faint and not well developed when present, the surface
glabrous except for a few appressed coarse hairs along the nerves, margins serrulate, the
teeth mostly divergent and aristate-tipped. Leaves reduced (commonly abruptly) in the
inflorescence, the uppermost becoming scale-like and many times shorter than the hypanthia.
Inflorescence an open or contracted cyme, few to many flowered. Flowers deep rose to
magenta, up to 3 cm across, petals broadly obovate, asymmetrical, tipped with a mostly
glandular hair, abaxial surfaces with numerous glandular hairs. Hypanthia at maturity 5-11
mm long, with necks slightly shorter to barely equalling the bodies, sparsely glandular-hairy
throughout, the rims with more numerous and coarser hairs up to ca. 2 mm long, calyx lobes
lanceolate to triangular-lanceolate, 2.1-3.1 (4.5) mm long, tipped with a slender hair ca. 1
mm long and mostly gland-tipped. Seeds cochleate, 0.5-0.7 mm long, surface irregularly,
but concentrically ridged, the ridges relatively high and comprised of contiguous to isolated
muricate processes varying to more regularly ridged with papillose processes.

Specimens of this presumed hybrid were first collected from New Jersey by K. K.
Mackenzie on 10 August 1913 near Egg Harbor City, Atlantic County (5584 CHRB, GH,
NY, PH). Annotations on his specimens, and notes entered in his collecting journal (unpub-
lished), suggest that Mackenzie struggled with the identification of these specimens. On
some specimens "Rhexia" is written in ink, followed by "aristosa" in pencil; others are
followed by "virginica" which was then crossed out and replaced with "aristosa." Still other
specimens are identified only as "Rhexia." In his journal, specimens of 5584 are entered as
"Rhexia aristosa Britt.," but followed by the comment, "5584 may be only a form of R.
virginica produced under artificial conditions." On one of the labels of his specimens (NY)
he wrote, "This species [my emphasis] is merely a form of R. virginica." Mackenzie
apparently never saw genuine R. aristosa in the field; if he did, he never collected
specimens. Based on his 5584 series, and the few specimens of R. aristosa available to
taxonomists early this century, it is understandable why Mackenzie would question the
taxonomic validity of R. aristosa.

Mackenzie was not alone in having difficulty in identifying these specimens. F. W.
Pennell in 1918 annotated one of the Mackenzie series (NY; named as R. aristosa by
Mackenzie) as R. virginica. This specimen, as well as two others of the series (also NY)
were then annotated to "?Rhexia aristosa x R. virginica" by James in 1955. In 1986, R.
Bounds annotated one of the specimens to R. aristosa (the one annotated to R. virginica by
Pennell) and two as R. virginica. In unpublished notes, Bounds (pers. comm.) says about
the specimen he annotated as R. aristosa, "I somewhat agree [with James’ annotation of
?Rhexia aristosa x R. virginica] but would say perhaps some virginica genes have
introgressed into this." Current opinions are also at odds. Frank Hirst (pers. comm.), who,
with his brother Robert, rediscovered Mackenzie’s locality in the late 1950s, considered these
plants to be possibly hybrids of R. aristosa and R. virginica. A. E. Schuyler collected and
labeled a specimen as R. virginica (23 Jul 1980, 5456, PH). Specimens from this locality
(DBS; VDB), plus identical material that I collected from two other nearby localities, were
identified by Robert Kral as R. aristosa, although he commented (pers. comm.) that these

RHEXIA IN NEW JERSEY 67

were, "tough calls indeed." Over the last 14 years that I have been studying this material,
I have variously identified these specimens as R. aristosa, R. virginica, or hybrids between
these taxa.

James (1956) stated that R. aristosa and R. virginica are "typically quite distinct
morphologically." According to Kral & Bostick (1969), R. aristosa is "one of the most
distinctive in the genus." Further, specimens of R. aristosa collected throughout its range
are consistently similar and show little significant morphological variation. This being so,
then why has there been such a divergence of opinions on the proper disposition of
Mackenzie’s 5584 and subsequent collections of this material? Slight discrepancies in the
keys of some manuals may have contributed somewhat to the uncertainty, but the primary
obstacle has been the lack of sufficient material to study (sheets of Mackenzie’s 5584 are
few, scattered between herbaria, and due to the usual wear and tear that specimens are
Subjected to, are often deficient in critical characters). With the relocation of the two
populations known to Mackenzie, and the discovery of two additional populations, a critical
evaluation is now possible.

In general appearance, the hybrid plants are more suggestive of R. virginica. The leaves
are On average wider, and more broadly lanceolate then is typical for R. aristosa, and the
leaves of the inflorescence are conspicuously reduced, unlike the leafy bracted inflorescence
of R. aristosa. On closer examination, the rims of the hypanthia have coarse yellowish
trichomes (strictly a characteristic of R. aristosa) and the sepal lobes are longer than those
of typical R. virginica.” However, the trichomes, on average, are shorter and less coarse than
those of R. aristosa, and many are gland-tipped (more characteristic of R. virginica). The
sepal lobe is not tipped with a stout, eglandular awn, as is R. aristosa, but is tipped with a
fairly slender, usually glandular hair as is usually RR. virginica. The seeds of R. x
brevibracteata are ridged with processes that are distinctly muricate or papillose (although
a few smoother ridges may also be present), giving the surface a decidedly rough or granular
appearance, especially at lower magnifications, in contrast to the smoother, more wrinkled
look of seeds of R. aristosa. Based on the length and shape of the sepal lobes and the
Presence of coarse trichomes on the hypanthium, R. x brevibracteata appears more closely
related to R. aristosa then to R. virginica, and if it were not for the greatly reduced bracteal
leaves of the inflorescence, it would be difficult to distinguish it from robust plants of R.
aristosa.

Rhexia x brevibracteata has been documented only from Atlantic County, New Jersey.
Four extant populations are currently known from three separate sites located between Egg
Harbor City and Port Republic. The greatest distance between sites is 3.7 miles. All sites
are located very near documented occurrences of R. aristosa. No site is farther than 3.7 miles
from a historical occurrence of R. aristosa or 8.4 miles from an extant occurrence. Although
natural occurring hybrids between R. aristosa and R. virginica have been reported in the
southern range of R. aristosa (R. Kral pers. comm.), there is no evidence that these hybrids
are morphologically coherent, persistent, stable, or taxonomically significant. Clearly more
Study is needed.

“The pur. Purshii phase of R. virginica has sepal lobes which in size (up to 4 mm) and shape (lanceolate) approach those
of R. aristosa or R. x brevibracteata . However, purshii material lacks the coarse hypanthial trichomes (hypanthia
often completely glabrous) of these taxa and the surficial sculpturing of its seeds are more like those of typical R.
virginica.

68 BARTONIA

All known occurrences for R. x brevibracteata are in inactive sand pits, similar to those
described under R. aristosa. Two pits (Egg Harbor City) are shallow, water-filled
excavations, which are normally dry by mid summer and are well vegetated with a diversity
of wetland sedges, grasses, and herbs. Characteristic species include Eleocharis microcarpa,
Rhynchospora knieskernii, Scirpus cyperinus, Muhlenbergia torreyana, Panicum verrucosum,
P. virgatum, Xyris difformis, Ludwigia linearis, Gratiola aurea, and Eupatorium luecolepis.
The other two sites (South Egg Harbor, Port Republic) are much deeper excavations and
have areas in which water levels regularly drawdown or dry out completely, but are also
interspersed with much deeper pond-like excavations that usually hold water throughout the
year. Most of the above listed taxa are present with Xyris smalliana, Ilex glabra, Vaccinium
macrocarpa, and Coreopsis rosea as notable additions. In these latter sites, R. X
brevibracteata grows in areas of fluctuating water levels in shallow depressions and "pond"
margins and bottoms and also in higher topographic positions characterized by nearly barren,
and comparatively dry sand—a habitat decidedly drier than that of R. aristosa or R. virginica.
Populations range in size from several hundred to several thousand plants. There is little
variation within and between populations, except in stature, branching, and leaf dimensions,
which is perhaps more a result of age and fluctuation of water levels. Rhexia virginica is
present at all documented sites, ranging in abundance from rare to infrequent. Only at one
site (near Port Republic) were plants collected that could not be definitely referred to R.
aristosa, virginica, or x brevibracteata. These plants were few, and grew mixed with typical
specimens of R. x brevibracteata and R. virginica. Historically, R. aristosa was
documented at one R. x brevibracteata site (Cologne, 29 Jul 1959, B. Hirst s. n., PH) but
despite repeated searches, it has not been found there since, or has it been discovered at any
of the other R. x brevibracteata sites.

Flowering normally occurs from mid July through late September, but varies somewhat,
depending on seasonal water levels. Fall coloration of the leaves is similar to R. aristosa
with leaves turning a blotchy purple-brown in early summer, the pigmentation becoming
darker and red tinged later in the season. Leaves persist until frost.

Although artificial hybrids between R. aristosa and R. virginica have been produced
experimentally (Kral & Bostick 1969), naturally occurring hybrids between these taxa are
apparently quite rare (R. Kral pers. comm.). Given the rarity of R. aristosa and the usually
different habitats occupied by each species (intermittent ponds vs. herbaceous wetlands), this
is not surprising. Field observations suggest that R. x brevibracteata isa fertile hybrid. Seed
production is copious, with a high percentage of full seeds produced. Numerous seedlings
have been observed at all the known localities. Cyclical fluctuations of population sizes
similar to those reported for R. aristosa have also been observed at one locality.
populations declined during successive years of high water levels and recovered in years of
average or lower than average water levels, with recruitment apparently through the seed
bank. Further, viable seeds provide a more likely explanation as to how R. X brevibracteata
has been dispersed to other sand pits which are separated from each other by a few miles as
well as a myriad of roads, agricultural lands, small towns, and other barriers to dispersal.
Such barriers are easily crossed by waterfowl. Each population could have arisen
independently as a result of chance hybridization, but given the rarity of R. aristosa in New
Jersey and the distance between documented sites of it and the hybrid, it seems the least
likely explanation.

Rhexia x brevibracteata \ikely arose as the result of the chance hybridization of R.
aristosa and R. virginica, and through subsequent backcrossing, possibly with R. aristosa,

RHEXIA IN NEW JERSEY 69

it has become a stable, morphologically distinct, and apparently sexually reproducing taxon.
Evidence supporting this conclusion is sumarized as follows: (1) fertile hybrids between the
putative parents have been produced experimentally; (2) both putative parents are
morphologically distinct, each possessing unique morphological characters which do not
overlap; (3) the putative hybrid posseses no unique morphological characters, but combines
characters from each of the supposed parents in an unique assemblage; (4) the hybrid is
documented from only a very small geographical area where both putative parents occur
together; (5) all documented occurrences of the hybrid occur in artificially created habitat.
It is clear that R. x brevibracteata can not be taxonomically accommodated as a phase of R.
aristosa or R. virginica, to do so, would blur the morphological distinctions of the latter two
sharply defined taxa.

(3 : NEW JERSEY: Atlantic Co.: W side of the Garden State Parkway, between
Indian Cabin Road and Clarks Mill Stream, ca. 1.2 mi W of jct of Mill Street and Main
Street, Port Republic. Thousands of plants scattered throughout a complex of inactive sand
pits. Plants cepa in shallow water along edge of pond-like excavations, 16 Sept 1993,
D. Snyder, PH (ISOTYPES: CHRB, DBS, NY).

ADDITIONAL NEW JERSEY SPECIMENS EXAMINED: Atlantic Co.: W side of the Garden State Parkway,
between Indian Cabin Road and Clarks Mill Stream, ca. 1.2 mi W of jct of Mill Street and Main Street, Port
Republic. Thousands of plants scattered throughout a complex of inactive sand pits. Growing in wet sand of
shallow excavations. Scattered plants of R. virginica present but no R. aristosa present, 17 Aug 1992, D. Snyder,
DBS, PH, VDB; Shallow artificial pond, Egg Harbor City, 10 Aug 1913, K. K. Mackenzie 5584, CHRB, GH, PH,
NY; 1.4 mi SE of Egg Harbor City, W side Berlin Ave, S side of RR. Scattered on now dry, sandy bottom of
inactive borrow pit, 21 Aug 1980, G. Cavileer, F. Hirst, and D. Snyder, DBS; 2.2 km SE of Egg Harbor City, SW
side of small pond west of Berlin Ave and RR. Scattered plants in moist sand, 23 Jul 1980 4. E. Schuyler 5456,
PH; 1.4 mi SE of Egg Harbor City, W side Berlin Ave, S side of RR. Abundant in shallow water and in wet sand
on nearly dry bottom of inactive borrow pit, 7 Oct 1993, D. Snyder, CHRB, DBS, NY, PH; Berlin Ave and RR,
Cologne, 29 Jul 1959, B. Hirst s. n. (mounted with a specimen of R. aristosa), PH; 1.4 mi SE of Egg Harbor City,
E side Berlin Ave, N side of RR. Abundant (3,000+) on nearly dry, sandy bottom of an inactive sacha pit, 25 Sept
1991, D. Snyder, DBS, PH, VDB; 1.4 mi SE of Egg Harbor City, E side Berlin Ave, N side of RR. Scattered
throughout mostly dry bottom of inactive borrow pit, 7 Oct 1993, D. Snyder, DBS; Sand oi on S side of
Belladonna Ave., South Egg Harbor. Ca. 1,700 plants, mostly growing in drier portions dy peu sand pits, some
growing in seemingly dry sand, 5 Aug 1992, D. Snyder, CHRB, DBS, NY, PH, VDB; Sand pits on S side of
Belladonna Ave., South Egg Harbor. poate erg growing in moist to dryish sand on oe bottom of a series of
inactive sand pits, 7 Oct 1993, D. Snyder,

ACKNOWLEDGMENTS

I thank curators at CHRB, GH, NY, PH and Gerry Moore for making specimens available.
Richard Bounds, Frank Hirst, Bill McAvoy, Ernie Schuyler, Rob Sutter, and Alan Weakley
helped with various aspects of this paper. Larry Morse reviewed an early draft. My special
thanks to Robert Kral for reviewing the final manuscript, annotating specimens, and his
Many insightful comments and discussions on the Rhexia taxa of New Jersey.

LITERATURE CITED

BEAVEN, G. F. & H. J. OosTING. 1939. Pocomoke Swamp: a study of a cypress swamp on the Eastern Shore of
Maryland. Bull. Torrey Bot. Club 66: 367-389.
BRITTON, N. L. 1889. Catalogue of plants found in New Jersey. Geol. Surv. N. J., Final Rep. State Geol. 2: 27-642.
~——. 1890. A new Species of Rhexia. Bull. Torrey Bot. Club 17: 14-15
FABLES, D., JR. 1962. 20 "Lost Plants." Bartonia 3: 7-10. :
U.S. Fish AND WILDLIFE SERVICE. 1990. Endangered, and threatened wildlife and plants; review of plant taxa for
29.

listing as endangered, or threatened species. Federal Register 55: 6184

70 BARTONIA

FAIRBROTHERS, D. E. AND M. Y. HOUGH. me resend 1975). Rare or endangered vascular plants of New Jersey.
Science Notes No. 14, N. J. State Mus.,

FERNALD, M. L. 1950. Gray’s manual of seca "he ed. American Book Co., New York.

L. GRISCOM. 1935. Three days of botanizing in southeastern Virginia. Rhodora 37: 167-189.

GLEASON, H. A. & A. CRONQUIST. 1991. Manual of vascular vita ants of northeastern United States and adjacent
Can

GopFREY, R. K. & J. W. WOOTEN, 1981. Aquatic and wetland of southeastern United States, Dicotyledons.
The Univ. of Georgia, Athen

earl F. & B. Hirst. 1963. Additions to the flora of Atlantic and Burlington Counties (including some relocated

cies). Bartonia 32: 11-13.

OUGH, M. Y. 1983. New Jersey wild plants. Harmony Press, New Jers

JAMES, C. W. 1956. A revision of Rhexia (Melastomataceae). Brittonia a 201-2 30.

KARTESZ, J. T. & R. KARTESZ. 1980. A synonymized checklist of the vascular flora of the United States, Canada,
and Greenland. Univ. of North oo Press, Chapel Hill.

KRAL, R. : P. E. BOSTICK. 1969. The genus Rhexia oe ee Sida 3: 387-440.

MACKENZIE, K. K. sto: alee bial of plants collected in New Jersey for years 1910-1918. N.Y. Bot. Gard.,
special aihiiohie

PosLusZzNy, U., M. J. SHARP, & . Keppy. 1984. a eens in Rhexia virginica: some
incenbolcaical ecological leat Canad. J. Bot. 62: 2118-2

SNYDER, D. B. 1994. Additions, range extensions, reinstatements, and aaa in the New Jersey flora. Bartonia

: 79-96,
STALTER, R. & E. LAMONT. 1990. The vascular flora of Assateague Island, Virginia. Bull. Torrey Bot. Club 117:
48-56.

STONE, W. 1912. The plants of southern New Jersey, with especial reference to the flora of the Pine Barrens. Ann.
Rep. N. J. State Mus., 1910, part IT; 21-828.

SuTTER, R. D. & M. Boyer. (unpublished) The seed bank of three rare species in southeastern pond cypress
savannas: Rhexia aristosa, Rebel boykinii, and Oxypolis canbyi . Southern Heritage Task Force, The Nature
Conservancy, Chapel Hill

TAYLOR, N. 1915. Flora in the vicinity of New York. Mem. N. Y. Bot. Gard. vol. 5.

Bartonia No. 59: 71-79, 1996

The Botanical Work of the Ruth Brothers of
Bucks County, Pennsylvania, and its Significance Today

PAMELA WHITE
New York Botanical Garden, Bronx NY 10458

ANN F. RHOADS
Morris Arboretum of the University of Pennsylvania, 9414 Meadowbrook Ave, Philadelphia PA 19118

In December 1881 22-year-old John Ruth and his 16-year-old brother Harvey decided to
"commence the study of botany." They sent away for a copy of "Gray’s Manual and
Lessons in Botany" (Gray 1867) for $2.16 and began to "get some knowledge of the
rudiments of the science." At the first sign of spring they set forth to tramp the woods and
fields in the vicinity of their home in the village of Monroe with the stated intent of
Preparing a "Catalogue of the Flora of Durham." Seven years of intense joint collecting
ended in 1889 when John Ruth moved to Bethlehem to take a job with the Lehigh Valley
Railroad. In succeeding years John added specimens to their joint herbarium from the
Vicinity of West Bethlehem, where he lived until September 1913, and later from Clifton,
NJ. Harvey, who moved to Pequest, NJ in 1902, continued to add to the collection until his
death in 1904.

John A. Ruth (1859-1918) and Harvey F. Ruth (1866-1904) were the sons of a German
Lutheran farm family of northeastern Bucks County. Their father, Charles Ruth, later gave
up farming and went to work as a blacksmith at the Durham Iron Works. John and Harvey
attended public school in Durham Township. At the age of seventeen John was sent to the
West Chester Normal School for six months after which he taught school from 1877 to 1888.
Following the completion of his formal education in the local public schools, Harvey was
employed in the chemical laboratory of the Durham Iron Works.

The observations and collections of this pair of amateur botanists were recorded in
detail by John Ruth in seven volumes of botanical notes totalling more than 750 pages and
nearly 300 pages of correspondence now preserved in the archives of the Academy of
Natural Sciences of Philadelphia (Ruth 1881-1917). The notebooks, written in a neat hand
and an almost poetical prose, contain only botanical observations. Few, if any, clues to other
events in their lives were included. However, a 44 page typewritten autobiography by John
Ruth (Ruth 1897) and a biographical sketch of John Ruth (Fackenthal 1926) provided
additional background. In addition, John and Harvey each contributed papers to the
Buckwampum Literary and Historical Society of which they were founding members. Titles
by John A. Ruth included "The Flora of Buckwampum" (Ruth 1888) read at a meeting held
June 14, 1888 and "Our Local Flora” (Ruth 1889) read June 8, 1889. Harvey read a paper
on the history of the village of Monroe (formerly Lehenburg) on June 13, 1896 (Ruth 1896).
In addition to the botanical notes, the brothers left a "Catalogue of the Flora of Durham and
Nockamixon" (Ruth 1887) containing a chronological record of 676 herbarium specimens
collected from April 1882 to September 1885, and a 109 page unpublished "Flora of
Northeastern Bucks County" (Ruth 1907).

71

72 BARTONIA

The notebooks also included clippings from the local newspapers (mainly the Doylestown
Intelligencer) which dealt with topics of a botanical nature such as notices of visits to Bucks
County by noted botanists, new botanical finds and obituaries of Asa Gray, Thomas C.
Porter and other prominent botanists. "The Ferns of Durham and Vicinity" (Moffat 1889),
a paper presented to the Bucks County Historical Society by Miss Margaret Moffat, and also
carried in the newspaper, provides a clue to the source of the brothers’ early botanical
interests. Miss Moffat was Harvey Ruth’s teacher at the Monroe School.

John Ruth’s notes recorded the brothers’ initial efforts at specimen preparation and the
improved methods they eventually adopted. John spent many winter evenings working on
the collection which he treasured until his death. The Ruth brothers’ herbarium, which
eventually consisted of 4000 specimens, became the property of a cousin, Benjamin F.
Fackenthal, Jr., after John Ruth’s death in 1918. Mr. Fackenthal "loaned" the collection to
the Academy of Natural Sciences where it remains (Fackenthal 1926; Stone 1920). Users
of the local herbarium at the Academy of Natural Sciences of Philadelphia will be familiar
with the Ruth brothers’ label, i 9 from an Easton firm in 1884, with J. A. and H. F.
Ruth printed boldly across the bott

Like many other amateur Beales of the 1800s, John and Harvey Ruth reflected a strong
Victorian era interest in natural history. Before they turned to botany they had amassed a
collection of some 400 specimens of minerals and fossils and 3500 Indian artifacts carefully
excavated at the site of a prehistoric village along the Delaware River near Kintnersville, all
of which are now in the collections of the Mercer Museum in Doylestown (Ruth & Ruth
1878). Both were also very interested in local history and genealogy and made contributions
in this area also.

When John and Harvey Ruth made the decision to apply themselves to botany it was a
total effort. Starting with "Gray’s Manual" mentioned above, they soon built up an extensive
botanical library. Green’s "Botany", "Trees, Plants and Flowers" (Baily 1870), "Familiar
Lectures on Botany" (Phelps 1852), "An Introduction to the Study of Botany" (Comstock
1832), and "Class-book of Botany" (Wood 1881) were purchased in 1883. The following
year they added "Grasses of the United States” (Vasey 1883), "Grasses and Forage Plants"
(Flint 1858), Carpenter’s "Vegetable Physiology and Botany", Cook’s "Manual of Botanic
Terms", "The Sylva Americana" (Browne 1832), "Catalogue of North American Carices"
(Bailey 1884), "The Botanical Collector’s Handbook" (Bailey 1881), "Manual of the Mosses
of North America" (Lesquereux and James 1884), and "The Naturalist’s Directory"
(Anonymous 1865-66). In 1885 more books were purchased including "Synoptical Flora of
North America" (Gray 1878, 1884), "Catalogue of the Plants Growing Without Cultivation
in the State of New Jersey" (Willis 1874) and "Our Native Ferns and Their Allies"
(Underwood 1881). In addition the notebooks contain frequent references to such periodicals
as: American Naturalist, Torrey Botanical Bulletin, Botanical Gazette and Proceedings of
the American Academy of Arts and Sciences.

Their equipment included a dissecting microscope made by Harvey from a pocket
magnifier purchased for $1.50 and homemade plant presses. In 1883 they purchased 4
Universal Household Microscope (70 to 290 diameters) for $12.00, with which to examine
mosses and other small specimens.

CONTACTS WITH THE BOTANICAL COMMUNITY

By 1883 John was in regular contact with Dr. Isaac S. Moyer, of Quakertown, author of

RUTH BROTHERS OF BUCKS COUNTY 73

the "Catalogue of the Flora of Bucks County" (Thomas and Moyer 1876; Moyer 1884) and
Dr. C. D. Fretz of Sellersville, Dr. Moyer’s colleague and successor. The Ruth brothers sent
specimens to Moyer and Fretz for confirmation and were proud to add, by John’s
calculations, over 30 species to the Bucks County list. Fretz’ revised catalogue credited J.
A. and H. F. Ruth with 26 taxa (Fretz 1905).

One of their first new county records was Harvey’s discovery of Ophioglossum vulgatum
on August 2, 1883. They immediately sent a specimen to Dr. Moyer and received the
following letter by return mail:

Quakertown 8-4-83
My dear sir: Your brother has made a very fine discovery. Dr. C. D. Fretz and myself have been scouring the upper
and middle of the county and I. C. Martindale the lower for years, for Ophioglossum, and without success. Be kind
enough to secure me a few good specimens, root and whole plant. Many thanks for your kindness. It has given
me much pleasure. I, S. Moyer

John’s comment, penned following the transcription of the letter in his notebook, was:
"This pays us richly for what we have spent on botany. It has been our good fortune to add
three new species to the Flora of Bucks Co.”

The discovery of Ophioglossum was followed quickly by Harvey’s collection of
Gentianella quinquefolia (formerly Gentiana baer hoi on October 3, 1883. Upon
receiving a specimen, Moyer had the following comm "Your plant is Gentiana
quinquefolia sure enough, and is an excellent find . . . arn to your researches-we can add
it to the flora of good old Bucks." A letter soon arrived from Fretz congratulating the
brothers on their finds, requesting specimens of both and promising to send "specimens of
mine" in return.

John and Harvey also corresponded with Thomas C. Porter, of Lafayette College in Easton
whose "Flora of Pennsylvania" (Porter 1903) was published posthumously. From 1885
through 1889 the Ruth brothers exchanged specimens with Porter and through their collecting
efforts, contributed several new state records including: Panicum boscii (formerly Panicum
latifolium var. molle), Scirpus cyperinus (formerly Scirpus eriophorum var. cyperinus) and
Scirpus sylvaticus, John later discovered Anthoxanthum aristatum (formerly Anthoxanthum
puelii), a European grass not previously collected in the state, growing in his lawn in West
Bethlehem. Porter repeatedly urged the brothers to, "pay me a visit and take a look at my
herbarium and botanical library and appliances." However, there is no evidence that they
ever did.

Other professional botanists with whom the Ruth brothers were in occasional contact
included L. H. Bailey, then of the Michigan Agricultural College, whom they consulted
regarding several Carex species; George Vasey, agrostologist at the Smithsonian Institution
in Washington, DC who identified several grasses for them, and Dr. N. L. Britton, of the
Columbia College Herbarium and later the New York Botanical Garden. They also sent
specimens to A. S. Hitchcock, of the State University of lowa and later the Smithsonian,
and Dr. H. H. Rusby of Columbia College.

By 1885 John and Harvey were actively involved in exchanging herbarium specimens with
other amateur botanists whom they located through the "Naturalists Directory". Correspon-
dence and lists of specimens offered, as well as "dissiderata" were included in the
correspondence (Ruth & Ruth 1882-1918). There were frequent references in the notebooks
to collecting specimens for exchange. Exchanges were recorded with other local PA and NJ
botanists as well as correspondents in NY, MT, ME, IL, CA, NE, FL, and MA.

14 BARTONIA

Although their herbarium and notes were eventually deposited in the Academy of Natural
Sciences of Philadelphia, there is little evidence that John and Harvey had much contact with
that institution during their lives. John Ruth sent a specimen of Rumex to the Academy in
1912 which Stewardson Brown identified as R. crispus L. (Anonymous 1912). John wasn’t
satisfied with that determination and subsequently sent specimens to the National Arboretum
and the New York Botanical Garden where botanists identified the plant as Rumex
triangulivalvis (formerly R. mexicanus).

The Philadelphia Botanical Club was not founded until 1891, after John had left Bucks
County to live in West Bethlehem. An 1896 card from Dr. C. D. Fretz to Harvey contained
an invitation to join a field trip to the Nockamixon Rocks by the Philadelphia Botanical Club
and the Torrey Botanical Club. Whether Harvey did so is not known.

COLLECTING SITES

John and Harvey Ruth botanized the northeastern corner of Bucks County. With few
exceptions, their activity was limited to the area they could reach on foot from their home
in the village of Monroe located along the Delaware River two miles south of Riegelsville.
Quite by chance this corner of the county has a highly varied geology which contributes to
the diversity of plants found in the region. The Monroe Border Fault, which intersects the
Delaware River near the village of Monroe, forms an abrupt boundary between the Triassic
shale, sandstone, quartzite and diabase to the south and a band of Cambrian limestone and
Precambrian gneiss, which are part of the Reading Prong, to the north (Geyer & Bolles
1987). In addition a fairly extensive deposit of Quaternary sand and gravel occurs along the
river in the vicinity of Kintnersville (Greenman 1955).

Collecting sites visited frequently by the Ruths included the Nockamixon Cliffs, Wykers
(formerly Lynn) Island, Buckwampum Mountain, Cooper and Hewitt’s pond, woods near
Durham Church, and the Nockamixon swamps. Although the intensity of collection and
degree of description of these sites varies, the information in the notebooks and manuscript
floras allowed us to make some comparisons between the appearance and species makeup
of these sites in the 1880s and their condition today. Of particular interest are early
collections of species now classified as endangered (PE), threatened (PT), rare (PR),
vulnerable or undetermined (TU) in Pennsylvania (Commonwealth of Pennsylvania 1993).
Overall, the Ruth collections included 17 taxa of concern to the Pennsylvania Natural
Diversity Inventory.

KAMIXON CLIFFS. The most famous of the sites frequented by the Ruths was the
Sheer, northwest facing cliff along the Delaware River between Kintnersville and
Narrowsville. Several streams cascade down the 300 foot cliff face forming cool, moist
ravines which harbor such rarities as the arctic-alpine species, Sedum rosea (PE). Even
before T. C. Porter’s discovery of Sedum rosea at the site in 1867, the Neckudioans Cliffs
drew the attention of botanists. C. S. Rafinesque called the cliffs "the greatest natural
curiosity on the Delaware River . . . | found here the Adlumia and many other rare plants."
(see Benner 1935). Other notable local botanists who have examined the flora at the site
have included John Harshberger; A. P. Garber (a student of Porter’s), S. S. Van Pelt, J. W-
Eckfelt, Thomas Seal, C. S. Williamson, Harold Pretz and Walter Benner (Harshberger 1899,
1909). Specimens of Sedum rosea collected by all of them can be found today in the
Academy of Natural Sciences Herbarium.
Aster ericoides (TU), collected at a rocky promontory at the top of the cliff known 4s

RUTH BROTHERS OF BUCKS COUNTY 75

"Top Rock" by John and Harvey, was found at the exact same spot in September 1991.
However several other species, including Taxus canadensis (TU), described by the Ruths
as abundant on the cliffs along the river from Monroe south, are apparently gone. Over
browsing by Virginia white-tailed deer may well be the cause, Taxus is preferred winter
browse (Allison 1990) and deer trails, pellets and browse lines are abundant on upper slopes
above the cliffs.

A 28 acre section of the Nockamixon Cliffs was recently added to the adjacent Delaware
Canal State Park through the efforts of the Heritage Conservancy’s Significant Natural Areas
Protection Program.

WYKERS ISLAND. Formerly known as Lynn Island, Wykers Island is located in the
Delaware River opposite the Nockamixon Cliffs near the town of Kintnersville. The island,
which is covered with a dense floodplain forest, is approximately 14 acres in area. As is
typical of river islands, the upstream end is characterized by alluvial sand and cobble
deposits with dwarfed and distorted tree and shrub growth which is frequently scoured by
flooding. The downstream end of the island is higher, less frequently flooded and supports
mature forest. Although John Ruth referred to an earlier time when part of the island was
farmed, it does not appear that there has been any disturbance in the past 100 years other
than that caused by the river itself. One Liriodendron tulipifera with a four foot trunk
diameter and a number of trees exceeding three feet dbh were observed in 1992.

In six visits to the island between August 1884 and October 1887 the Ruths compiled a
list of 197 species. In May and August visits in 1992, we noted a total of 151 taxa,
approximately 55 of which appeared on the Ruth brothers’ list. 1992 data included 40
species of trees and shrubs of which one of the most interesting was Diospyros virginiana,
the native persimmon, which formed a grove near the upper end of the island. John an
Harvey noted "a small tree well in fruit" 100 years earlier.

The island proved to be fertile ground for new county records to add to John and Harvey’s
list. Eragrostis capillaris, Spartina pectinata (formerly S. cynosuroides) and Lythrum
salicaria were first collected on September 9, 1884 and Cyperus dentatus was added in 1885.
A specimen of Lythrum salicaria, "a single plant along the eastern shore near the water’s
edge," was sent to Moyer for confirmation, and pronounced by him as "very rare" and "a
splendid find". However the species was not seen the following year leading John to
conclude that it had "probably been destroyed." By a hundred years later Lythrum salicaria
had spread up and down the river and become a major threat to native wetland species
throughout the northeastern states (Thompson et al. 1987).

Prunus pumila and Lupinus perennis (both PR), were collected by the Ruths on the island.
On May 20, 1887 John recorded the following notes regarding these species: "One of the
first plants found was Prunus pumila in flower. It is spread over the stony northern end .

Near by we found large beds of lupine in flower ...." Recent searches indicated that
these species were not present on the island, presumably because there was less open area
at the upstream end. Other differences between John and Harvey’s species list and our 1992
data support the conclusion that formerly open areas on the island’s northern end have grown
up in the past 100 years perhaps due to less frequent scouring by ice. The Ruths listed 65
Species in the families Poaceae and Asteraceae, in contrast to the 22 species which we
observed in 1992. Prunus pumila is still abundant on islands further upstream where the
habitat is more open. '

Another state rarity, Gratiola aurea (TU), collected by John and Harvey in mud at the
north end of the island, could not be relocated in 1992. It has not been found recently at

76 BARTONIA

other former locations along the river in Bucks County either.

Tradescantia ohiensis (TU) was abundant throughout the island’s interior in May 1992.
While John and Harvey did not list this species, they did record the presence of T.
virginiana. Gray did not recognize T. ohiensis (Gray 1867, 1887) in the manuals the Ruths
had access to, and described T. virginica as highly variable, smooth or hairy. Since the two
species are distinguished mainly by the pubescent sepals and pilose pedicels of 7. virginiana
vs. glabrous sepals and pedicels of T. ohiensis (Fernald 1950), it is possible that the species
we saw in 1992 was the same thing that was present 100 years earlier. Unfortunately there
was no Ruth herbarium specimen of Tradescantia from the island so it was impossible to
be certain what they saw. Interestingly, a Ruth specimen labelled as 7. virginiana collected
from the riverbank at Monroe was correctly identified by today’s definition of that species.
T. virginiana had also been collected at other nearby sites.

A further complication was the fact that 7. ohiensis is frequently cultivated and may have
been introduced to the island via flood waters as have many other species. The process,
described by John Ruth in the following excerpt, "seeds from distant localities . . . brought
by the freshets and deposited among the drift heaps where in time they spring up and
become established,” has continued over the years. Today the herbaceous flora of much of
the island’s interior is dominated by exotics such as Alliaria petiolata, Anthriscus sylvestris,
Hesperis matronalis, Microstegium vimineum, Lonicera japonica, Lamium purpureum,
Glechoma hederacea, Allium vineale and Polygonum cuspidatum, none of which were
reported 100 years ago.

BUCKWAMPUM MOUNTAIN. Located between Durham Road and Route 412 in Springfield
Township, Buckwampum Mountain was the childhood home of John and Harvey Ruth an
later a favorite collecting site. John Ruth’s notes record 21 visits to the site between 1882
and 1887. It was there in a "large open bog on the east side of the hill" that they collected
Drosera rotundifolia in 1884. Another local rarity Platanthera psycodes (formerly
Habenaria psycodes), was found in abundance along streams southwest of the "bog" in 1886.
Neither of these species was found during repeated searches of the eastern slopes of the
mountain in recent years. Although much of the area remained forested, and appeared intact,
the canopy at the "bog" was fairly dense and the area was apparently much shadier than it
was at the time of the Ruths’ visits. Abundant deer trails and a generally sparse shrub layer
indicated that browsing by deer may also have been a factor in altering the species
composition. Hydrastis canadensis, \isted as vulnerable on the state list, and not mentioned
at all by the Ruths, formed an extensive stand on the mountain in 1992.

COOPER AND HEWITT’S POND. This vernal pond located just south of Rattlesnake Hill, was
one of two sites at which the Ruths collected Gentiana flavida (formerly G. alba) (TU).
John and Harvey were very intrigued with this find and corresponded with Thomas C. Porter
about it. At the time, Porter struggled with the occurrence of this gentian in eastern PA and
hypothesized that it was transported from the southwest part of the state by Indians.
Subsequent collections from Lehigh and Northampton Counties have confirmed that the
range of this species included eastern PA. John Ruth described the pond as "surrounded by
a growth of oak, maple, dogwood and other trees . . . dry at all seasons except the spring.”
In 1992 the only evidence of its former location was a slight indentation in an expanse of
cultivated fields, an indication of the fragility of this habitat type.

WOODS NEAR DURHAM CHURCH. These woods, which occur on the southern slope of the
Reading Prong on Precambrian gneiss, were visited several times by the Ruths, and were the
subject of John’s reverent admiration. Local knowledge of the botanical curiosities of this

RUTH BROTHERS OF BUCKS COUNTY 4 |
wooded ravine no doubt inspired him when in 1884 he wrote the following in his notebook:

A stream of purest spring water, with its several branches, winds in and out among the majestic trees... . Gray,
mossy, gnarled old oaks that have weathered the storms of a century skirt the banks of the little stream. Tulip
poplars, straight as an arrow, rise to the height of a hundred feet. Fine maples, beech, and chestnuts add their
foliage. A feeling of awe comes over me when in such a grove. No temple reared by human hands is more
imposing.

Although John and Harvey reported such local rarities as Obolaria virginica, Galearis
spectabilis (formerly Orchis spectabilis), Liparis liliifolia, and Asclepias exaltata, none of
these were located in a May 1992 visit. Non-native species such as Rosa multiflora and
Lonicera japonica had invaded the peripheral areas, but the central portion of this stream
valley remained an intact native woodland. The forest of Fraxinus americana and
Liriodendron tulipifera appeared to be the result of at least partial cutting 50 or 60 years ago.
Steep slopes contained a more mature beech-oak forest. Scattered along the stream were
Many seepage areas with moss covered rocks and dense stands of Symplocarpus foetidus.
Early reports indicated that Tro/lius laxus (PE) once grew in one of these boggy ravines
(Thomas & Moyer 1876). Although it was mentioned in a newspaper clipping preserved in
John’s notes, there was no evidence that the Ruth brothers ever came across this species in
their botanizing.

OCKAMIXON SWAMPS. The Nockamixon Cliffs were formed by partly metamorphosed
shale and siltstone hardened by the heating effect of a massive igneous intrusion (Greenman
1955). The diabase sill which resulted forms a poorly drained, rocky upland above the cliffs
of which Coffman Hill is the high point. Much of the area, which is characterized by
Sluggish, boulder-strewn streams, wet meadows and moist, rocky woods, is presently
included in State Game Lands Number 56. The remainder is abandoned farms and low
density rural lots with numerous small, manmade ponds.

John Ruth found the meadows and swamps of this region most interesting. On June 1,
1886 he described them as, "a mass of flowers . . . one of the most beautiful sights you ever
beheld... large patches golden yellow with the flowers of Krigia biflora (formerly Cynthia
virginica) and Senecio aureus .. . acres blue with Jris versicolor. . . and here and there small
patches of scarlet which announce the presence of Castilleja coccinea . . . ditches are filled
with . . . Orontium aquaticum ..." Other rarities found in the area included Carex
buxbaumii, Carex tetanica and Eriophorum gracile.

ile meadows tend to be early successional habitats, and thus not stable, long-term plant
communities, areas such as those described by John Ruth can still be found in the diabase
region of Bucks County. Repeated searches in recent years have failed to locate extant
Populations of Orontium aquaticum (PR) or Eriophorum gracile (PE); but other species such
as Carex buxbaumii (TU), Carex meadii (TU), Castilleja coccinea (TU) and Phlox pilosa
(TU) were still present. In addition species such as Caltha palustris, Carex squarrosa, Phlox
maculata, Lilium canadense, Platanthera lacera, Scirpus pendulus, Liatris spicata,
Andropogon gerardii, Sorghastrum nutans, Helianthus giganteus, Rudbeckia triloba, R
fulgida, Cirsium muticum, Eupatorium fistulosum, Gentiana andrewsii, and Gentianopsis
crinita continue to make these wet meadows a botanical delight as well as a succession of
color throughout the season.

78 BARTONIA
CONCLUSION

Change has come to a number of the collecting sites John and Harvey Ruth frequented
100 years ago. The most severe is altered land use which has totally eliminated a vernal
pond and continues to threaten other areas such as the Nockamixon swamps and the lower
slopes of Buckwampum Hill. The least disturbed sites are Wykers Island and the
Nockamixon Cliffs, which are protected in part by their difficult access. However, even
those sites have lost some of their earlier diversity. Changes in the flora of Wykers Island
appear to be related to reduced scouring by ice and the constant introduction of exotic
species via flood waters. Succession and over browsing by Virginia white-tailed deer are
apparently factors in the loss of species from several sites.

In many ways John and Harvey Ruth fit the description of amateur botanists of the
nineteenth century developed by Keeney (1992). By collecting intensively in a local area
they were able to provide a welcome extension of the work of the professionals, who were
in turn willing to assist the brothers with difficult identifications, exchange specimens with
them, and offer encouragement. They also became active participants in a nationwide
network of amateur and professional botanists involved in the exchange of botanical
specimens.

However, unlike Keeney’s profile of an amateur as one primarily driven by a desire for
self improvement, John and Harvey were clearly interested in adding to the scientific
knowledge of the flora of their area. They kept careful records of their botanical
observations and noted with great pride the additions to the lists of species known from
Bucks County and the Commonwealth of Pennsylvania which resulted from their work.
Their botanical specimens are today an integral part of the local herbarium at the Academy
of Natural Sciences and are included in the Flora of Pennsylvania Database (Rhoads & Klein
1993) where they contribute to documenting the state flora.

Equally significant is the "snapshot in time" which John Ruth’ s botanical notes provided.
They have given us a unique opportunity to better understand the changes in the flora of
northeastern Bucks County over the past 100 years.

ACKNOWLEDGMENTS

We thank librarians at the Academy of Natural Sciences of Philadelphia and the Bucks
County Historical Society for their assistance in locating historical documents and Patricia
Pingel for help in relocating some of the Ruth brothers’ collecting sites.

REFERENCES

ALLISON, T. D. 1990. The influence of deer browsing on the reproductive biology of Canada yew Taxus canadensis
Marsh.) Oecologia 83:523-529.

toe 1865-66. The naturalists directory. Parts I. & II., North America and West Indies. Salem Publishing
Co le

ANONYMOUS. 1912. Additions to the museum. Proc. Academy of Natural Sciences 64:569-577.

BAILEY, L. H. 1884. Catalogue of North American Carices, Cambridge

BAILEY, W. W. 1881. The botanical collector’s handbook. Naturalist’s handy series, No. 3. George A. Bates, Salem.

BAILY, W. L. 1870. Trees, plants and flowers: where and how they grow. J. B. Lippincott and Co., Philadelphia.

BENNER, W. M. 1932. The flora of Bucks County, Pennsylvania. Published by the author, Philadelphia.

Brown, D. J. 1832. The sylva americana. W. Hyde and Son, Boston

COMMONWEALTH OF PENNSYLVANIA. 1993. Pennsylvania Code, Title 25. Chapter 82. Conservation of Pennsylvania
native wild plants. 23 Pa.B. pes (October 30, 1993)

RUTH BROTHERS OF BUCKS COUNTY 719

wae tay J. L. 1832. An introduction to the study of botany. D. F. Robinson and Co., Hartford
ACKENTHAL, B. F., JR. 1926. John A. Ruth biographical notice. Bucks County aces Society Papers Vol.
V:275-27 8.
FLINT, C. L. 1858. Grasses and forage plants. A. O. Moore, Agricultural Book Publisher, New Y
FRETZ, C. D. 1905. Flora of Bucks County, Pennsylvania. Pages 7-54 in W. W. H. Davis. History oe bee County,
2nd ed., vol. Il, appendix. A. E. Lear, Inc. Pipersville.
GEYER, A. R. & W. H. BOLLES. 1987. Outstanding scenic geologic features of Pennsylvania, Part 2. Bureau of
Topographic and Geologic Survey, Department of Environmental Resources, Commonwealth of Pennsylvania,
Harrisburg.
GRAY, A. 1867. Manual of the botany of the northern United States. Ivison, Blakeman, Taylor and Co., New York.
GRAY, A. 1878. Synoptical flora of North America, vol. II, part 1. Ivison, Blakeman, Taylor and Co., New York.
Gray, A. 1884. Synoptical flora of North America, vol. I, part 2. Ivison, Blakeman, Taylor and a New York.
GRAY, A. 1887. Gray’s lessons and manual of botany, revised edition. Ivison, Blakeman and Co., New York.
GREENMAN, D. W. 1955. Ground water resources of Bucks County, Pennsylvania. Bulletin hes ‘eet and
Geologic Survey, Department of Internal Affairs, Commonwealth of eg ta Harrisburg.
HARSHBERGER, J. W. 1899. The botanists of Philadelphia and their work. T. C. D: aoe: Sting Philadelphia.
HARSHBERGER, J. W. 1909. Plant formations of the Nockamixon Rocks. Bull. Torrey Bot. Club 36:651-673.
HITCHCOCK, é S. 1935. Manual of grasses of the United States. United States Government Printing Office,
Washingto
KEENEY, E. 2 1992. The EL gperscots University of North Carolina Press, Chapel Hill
LESQUEREUX, L. & T. P. JAMES. de Manual of the mosses of North America. Bradlee Whidden, Boston.
Morrat, M. J. 1889. The os of Durham and vicinity. Bucks County Historical Society Papers 1:209-212.
Moyer, I. S. 1884. Indigenous a naturalized flowering plants, ferns and fern allies of Bucks County. Bucks
County Historical Society Papers 1:161-172.
PHELPS, A. H. L. 1852. Familiar lectures on botany. Francis J. Huntington, New York.
PorTER, T. C. 1903. Flora of tc Ac Ginn and Company. New York,
RHoaps, A. F. & W. MCK. KLEIN Aa ie vascular flora of Pennsylvania: annotated checklist and atlas.
American Philosophical Society, Ph ilad
RuTH, H. F. 1896. Lehnenburg, sora aech of a small village. B. F. Fackenthal Collection, Fol. 44, Bucks
ey Historical eine tone
UTH, H. F.& J. A . 1878. laine notes and reg of aboriginal implements collected in Bucks
County, 4 vols. rier hele Historical Society, Doylestow
RuTH, J. A. 1881-1917, Botanical notes, 7 vols. Manuscript ‘Cilietind #694, Academy of Natural Sciences,
fone
RuTH, J. A. 1887. Catalogue of the flora of Durham and Nockamixon, Manuscript Collection #694, Academy of
Natural Sciences, Philadelphia
RUTH, J. A. 1888. The flora of Buckwampum: Manuscript Collection #386, Fol. 1, Bucks County Historical Society,
Doylestown.
Ruth, J. A. 1889. Our local flora. Bucks County Historical Society Papers 5:222-231.
RuTH, J. A. 1897. Autobiography. Manuscript Collection #386, Fol. 2, Bucks County Historical Society,
Doylestown.
RuTH, J. A. 1907. Flora of northeastern Bucks County, parts 1 and 2. Manuscript Collection #694, Academy of
Natural Sent Philadelphia.
& H. F. RuTu. 1882- 1918. Correspondence. Manuscript Collection #694, Academy of Natural

Stisiices: days

STONE, W. W. 1920. Letter to B. F. Fackenthal dated July 19. 1920. Manuscript Collection #206, Academy of
Natural Sciences, Pemage nem

Co J. & I. S. Moyer. 1876. Plants, birds and mammals of Bucks County, oo appendix pages 3-27

n: W. W. H. Davis, ce of Bucks County. Published by the author, Doyles

Thies D. Q., R. L. StuCKEY, & E. B. THOMPSON. 1987. Spread, impact, a case of purple nigel
(Lythrum salicaria) in North American wetlands. Fish and Wildlife Research 2. United States Department
Interior, Fish and Wildlife Service, Washington.

UNDERWOOD, L. M. 1881. Our native ferns and their allies. H. Holt and Co.

VASEY, G. 1883. The grasses of the United States. U. S. Government Printing Office, Washington.

WILLs, O. R. spe Catalogue of the plants growing without cultivation in the state of New Jersey. W.

Sc d Co.
Woon, A. 1881. Class-book of botany. A. S. Barnes and Co., New York.

eased 4 Ag
(HIG! eA io

Poptart a acyihoes «

2 EOD LE 3
A Yano
fe

abe.
aay

Bartonia No. 59: 81-85, 1996
Additions to the Flora of Bradford County, Pennsylvania

ROBERT F. C. NACZI AND JOHN W. THIERET
Department of Biological Sciences, Northern Kentucky University
Highland Heights, KY 41099-0400

This account lists 80 species, 29 genera, and 2 families as new to Bradford County,
Pennsylvania, as well as two species, Carex Gilet ep Carex texensis, that appear to
be new state records. This determination was made from information in a recent floristic
atlas (Rhoads & Klein 1993) and an associated database lb 2 Flora Database 1995).
Voucher specimens, collected by Naczi in 1990 and by Naczi and Thieret in 1994, have been
deposited in the herbarium of the Academy of Natural Sciences of Philadelphia (PH), with
duplicates of some collections at Northern Kentucky University (KNK) and the University
of Michigan (MICH). Nomenclature largely follows that used in Rhoads & Klein (1993);
if we use a different name, the one used there is given in parentheses. Species that
apparently have been introduced to Pennsylvania are noted, based on data presented in
Rhoads & Klein (1993). Collection numbers in the 2000s are those of the senior author
alone; those in the 4000s, of both authors. The following list is alphabetical by family and
species.

CEAE. Acer platanoides—Towanda, trees to 12 inches DBH, with Fab smaller ones and saplings,
woods along railroad tracks behind Riverside Cemetery, 4242, 16 Jun 1994, introduc

APIACEAE. Aegopodium podagraria pee form)—Towanda, wooded drainage area - outside of
culvert, behind Riverside Cemetery, 4248, 16 Jun 1994, introduced. Anthriscus sylvestris is—ca. 3.3 mi S of West
Franklin: roadside, 4269, 16 Jun 1994, Asai aS Osmorhiza claytonii—ca. 13 mi SW of ae aaa maple
forest, 4296a, 17 Jun 1994,

ARALIACEAE. Panax quinquefolius—ca. 13 mi SW of Towanda, sugar maple forest, 4297, 17 Jun 1994,
ris—Sayre, disturbed area in railroad yard, 4329, 18 Jun 1994, introduced.

BRASSICACEAE. Mista a thaliana—Sayre, weedy, gravelly area in s einat yard, 435], 18 Jun 1994,
. ; ; anda, along railroad track at un
Station, 4237, 16 Jun sy en edge of Troy, roadside, 4/59, 15 Jun 1994, pean Sa Nasturtium officinale—c
4 mi SW of Troy, in small stream, 4222, 15 Jun 1994, introduced. Sisymbrium loeselii—Sayre, weedy, gravelly area
in railroad yard, 4350, 18 Jun 1994, introduced.

CABOMBACEAE. Brasenia schreberi—ca. 15.5 mi SW of Towanda, many leaves washed up on shore, Sunfish
Pond County Park, 4308, 17 Jun 1994

CAESALPINIACEAE. Gleditsia pinodbithoe—<. 11 mi SW of Towanda, two trees at edge of woods, 43/6,

7 Jun 1994.

CANNABACEAE. Humulus japonicus—Towanda, disturbed area along Susquehanna River at route 6 bridge,
4239, 16 Jun 1994, introduced. i

CAPRIFOLIACEAE. Lonicera maackii—ca. 3.3 mi S of West Franklin, one shrub 12 ft tall, perhaps originally
planted, eng 16 Jun 1994, introduced. Lonicera morrowii—Troy, lightly shaded stream bank, 4/58, 15 Jun 1994,
introduce

chavaniece Cerastium glomeratum—Troy, garden weed, 4/57, 15 Jun 1994, introduced. Sagina

81

82 BARTONIA

procumbe ae in fares at edge of woods, back edge of Riverside Cemetery, 4244, 16 Jun 1994.
Scleranthus annuus—ca. mi S of Sylvania, ene 4167, 15 Jun 1994, introduced. Silene antirrhina—Sayre,
weedy, gravelly area in sete yard, 4348, 18 Jun 1994, introduced.

CYPERACEAE. Carex albicans—ca. 4 mi ‘i of New Albany, mesic deciduous woods, 4/33, 14 Jun _—

ar atlantica ssp. capillacea—ca. 3.4 mi SW of West Franklin, wetland, 4284, 16 Jun 1994. Carex blanda—c
SSE of Towanda, lightly shaded, moist, sandy silt along Towanda Creek, 26/5, 16 Jun 1990; Sayre, meee
pho area at edge of woods at edge of railroad yard, 4335, 18 Jun 1994. Carex bromoides ssp. bromoides—ea. 2.5
NW of mage in alder thicket along stream, 2589, 15 Jun 1990; ca. 4 mi N of New Albany, mesic

pe iduous woo 34, 14 Jun 1994. Carex ee eeepegtig 2 mi SSE of West Franklin, swampy r
preonaeNaae gdb 2622, 16 Jun 1990; ca. 5 m W of Troy, moist to wet red maple woods, 4/87, 15 Jun
1994; ca. 4 mi WSW of Troy, roadside in alae ii 4198, 15 Jun 1994. Carex crawfordii—Sayre, weedy,
moist area at edge of woods at edge of railroad yard, 4336, 18 Jun 1994. Carex glaucodea—ca. 1.3 mi NW of
Frankliadale, lightly shaded slope above creek, 26/0, se ie 1990. Carex O llensne oa 2.5 mi NW of
Franklindale, 2597, 15 Jun 1990; Sayre, weedy, moist area at edge of woods at edge of railroad yard, 4340, 18 Jun
1994. Carex grayi—ca. 2 mi SSE of Towanda, lightly shade moist clay along Towanda Creek, 2620, 16 Jun 1990.
Carex grisea—ca. 2 mi SSE of Towanda, lightly shaded, moist, sandy silt along Towanda Creek, 26/3, 16 Jun 1990;
Sayre, weedy, moist area at edge of woods at edge of railroad yard, 4347, 18 Jun 1994. Carex hirtifolia—ca. 2 mi
NW of Franklindale, mesic deciduous forest near stream, 2596, 15 Jun 1990. Carex leavenworthii—Towanda, edge
of woods, back edge of Riverside Cemetery, 4245, 16 Jun 1994. Carex mesochorea—Towanda, edge of woods,
back edge of Riverside Cemetery, 4246a, 16 Jun 1994. Carex molesta—Sayre, weedy, moist area at edge of woods
at edge of railroad yard, 4337, 18 Jun 1994. Carex novae-angliae—ca. 2.1 mi S of West Franklin, moist red
maple/hemlock woods, 2632, 17 Jun 1990; ca. 5 mi WSW of Troy, moist to wet red maple woods, 4/85, 15 Jun
1994; ca. 4 mi SW of Troy, wet, sunny site near small pond in larch swamp, 4230, 15 Jun 1994, Carex
pensylvanica—ca. 2 mi SE of Towanda, rocky, oak/red maple/hemlock forest, 26/2, 16 Jun 1990; ca. 5 mi WSW
of Troy, moist to wet red maple woods, 4/84, 15 Jun 1994. Carex prasina—ca. 2.5 mi NW of Franklindale, in alder
thicket along stream, 2590, 15 Jun 1990; ca. 4 mi WSW of Troy, 4 in caphos aa. 4202, 15 Jun 1994. Carex
sparganioides—ca. 1.5 mi S of Sylvania, deciduous forest, 4/60, 15 Jun 1994. Carex stricta—ca. 3.4 mi SW of
West Franklin, ath wetland, 4282, 16 Jun 1994. Carex tenera—ca. fF mi N of Evergreen, moist shallow ditch,
4155, 14 Jun 1994; Sayre, weedy, moist area at edge of woods at edge of railroad yard, 4346, 18 Jun 1994. Carex
texensis—Towanda, moist, shaded lawn, back edge of Riverside Cemetery, 4247, 16 Jun 1994, probably introduced.

m

2631, 17 Jun 1990; ca. 1 mi N of Evergreen, dry rocky roadbank, 4/50, 14 Jun 1994. Carex willdenowii—ca. 13
mi NW of Franklindale, ase soil on slope above creek, 2608, 15 Jun 1990; ca. 4 mi N of New Albany, deciduous
forest, 4/30, 14 Jun 1994. Eleocharis acicularis—ca. 15.5 mi SW of Towanda, forming mats in shallow water along
shore, Sunfish Pond County Park, 43/2, 17 Jun 1994. Schoenoplectus subterminalis—ca. 15.5 mi SW of Towanda,
plant fragments washed up on shore, Sunfish Pond County Park, 43//, 17 Jun 1994. Scirpus atrocinctus—ca. 13
mi SW of Towanda, edge of open, marshy pond, 4305, 17 Jun 1994.

DROSERACEAE. Drosera rotundifolia—ca. 16.5 mi SW of Towanda, in peatland, 43/7, 17 Jun 1994; ca. 4
mi WSW of Troy, shrubby, sunny, wet area with mosses, 4/94, 15 Jun 1994; ca. 3.4 mi SW of West Franklin,
wetland, sphagnous, 4280, 16 Jun 1994,

ERICACEAE. Rhododendron maximum—ca. 4 mi § of West Franklin along Schrader Creek, shaded cliffs along
creek, 264/, 18 Jun 1990; ca. 1.7 mi S of West Franklin, edge of depression dominated by shrubs, 4263, 16 Jun
1994.

FABACEAE. Coronilla varia—Towanda, roadside along route 6, 423/, 16 Jun 1994, introduced.
GERANIACEAE. Geranium pusillum—Towanda, weedy lawn along Susquehanna River, 4234, 16 Jun 1994,
introduce

IRIDACEAE. Iris pseudoacorus—ca. 8.5 mi SW of Towanda, sunny, moist roadside, 4292, 17 Jun 1994,
troduced.

JUNCACEAE. Luzula tr ent ae 4 mi WSW of Troy, shrubby, sunny, wet area, 4/95, 15 Jun 1994.
LAURACEAE. Lindera benzoin—ca. 1.7 mi S of West Franklin, swampy area, 425/, 16 Jun 1994.
her degene Smilacina si Ps 1.7 mi S of West Franktin, in shaded sphagnum under shrubs, 4254, 16

1994.

EYCORODIACEAE —— inundata—ca. 3.4 mi SW of West Franklin, sandy spots at pond margin,
4273, 16 Jun 1 m hickeyi—ca. 5 mi WSW of Troy, moist to wet red maple woods, 4/97, 15 Jun
1994; ca. 3.4 mi on of West ‘Presa rocky, dryish forest, 4270, 16 Jun 1994.

LYTHRACEAE. Decodon verticillatus—ca. 3.4 mi SW of West Franklin, in shrubby wetland, 4278, 16 Jun

BRADFORD COUNTY FLORA 83

MENYANTHACEAE. Menyanthes trifoliata—ca. 3.4 mi SW of West Franklin, in shrubby wetland, 4269a, 16
Jun 1994.

NYMPHAEACEAE. Nymphaea odorata—ca. 15.5 mi SW of Towanda, in shallow water, Sunfish Pond County
Park, aes 17 Jun 1994,

PINACEAE. Picea abies—ca. 3 mi S of West Franklin, uneven-aged and -sized trees, reproducing, in disturbed
woods, née 16 Jun 1994, introduced. Picea glauca—ca. 3 mi S of West Franklin, uneven-aged and -sized trees,
reproducing, in disturbed woods, 4265, 16 Jun 1994, introduced. Picea rubens—ca. 3.4 mi SW of West Franklin,
moist wey 4277, 16 Jun 1994,

AE. Avena sativa—Wysox, disturbed area, 4352, 18 Jun 1994, introduced. Lolium perenne—Towanda,
along frst tracks at railroad station, 4236, 16 Jun 1994, introduced.

PO OGETONACEAE. Potamogeton epihydrus—ca. 15.5 mi SW of Towanda, plant fragments washed up
on shore, Sunfish Pond County Park, 43/5, 17 Jun 1994.

PRIMULACEAE. Lysimachia piettdniehe's 9 16.5 mi SW of Towanda, peatland, 4327, 17 Jun 1994.

RANUNCULACEAE. Ranunculus bulbosus—ca. 2 mi § of Sylvania, moist, shady roadside, 4/76, 15 Jun 1994,
introduced. Ranunculus repens—ca. 4 mi WSW of Troy, shady, wet roadside, 4207, 15 Jun 1994, introduced.

ROSACEAE. Geum rivale—ca. 4 mi SW of Troy, larch swamp, 4223, 15 Jun 1994. Sorbaria sorbifolia—ca.
3 mi S of West Franklin, large colony for ca. 30 m along roadside, perhaps from original planting, 4266, 16 Jun
1994, introduced.

SAXIFRAGACEAE. Saxifraga pensylvanica—ca. 4 mi SW of Troy, larch swamp, 4224, 15 Jun 1994.

SCROPHULARIACEAE. Chaenorrhinum minus—ca. 8 mi SW of Towanda, roadside, 4290, 17 Jun 1994,
introduced. Melampyrum lineare var. americanum—ca. 1.5 mi S of Sylvania, rocky roadbank at edge of mixed
woods, 4/64, 15 Jun 1994. Veronica peregrina—Towanda, along railroad track behind Riverside Cemetery, 4243,
16 Jun 1994,

“Sb ar Sa Ailanthus altissima—Towanda, bluff along Susquehanna River, 4232, 16 Jun 1994,
introduc

DISCUSSION

Of the 80 species we report as new to Bradford County, 47 (59%) are vsti to
Pennsylvania and 33 (41%) are introduced to the state. The total number of sp and
infraspecific taxa previously recorded from Bradford County is 899, of which 712 (79%) are
native and 187 (21%) are introduced (Pennsylvania Flora Database 1995). Including our
discoveries, 979 total species and infraspecific taxa are now recorded for the county, with
759 (78%) natives and 220 (22%) exotics. The large proportion of introduced species among
our county records indicates at least two things. First, introduced plants probably have been
neglected in previous botanical exploration of the county. Second, such species probably
have increased in importance in the flora in recent time. Acer platanoides, Anthriscus
sylvestris, Carduus nutans, Coincya monensis (Hutera cheiranthos), and Lonicera morrowii
may be recent invaders of Bradford County. Rhoads and Klein (1993) include these exotics
in a list of species that apparently are spreading within Pennsylvania. A large proportion of
the natives (22 of 47, or 47%) are members of Carex, the most species-rich and taxonomical-
ly complex genus of vascular plants in Pennsylvania. Many field botanists avoid collecting
Carex plants because they believe them to be too difficult to identify. Thus, neglect of these
plants in past collecting in Bradford County probably accounts, at least somewhat, for the
large number of Carex species among our records.

Many of our county records, both natives and exotics, are quite expected. Such species
as Carex bromoides, C. pensylvanica, C. eoogee Drosera rotundifolia, Lindera benzoin,
Osmorhiza claytonii, Potamogeton epihydrus, Rhododendron maximum, and Tussilago
Jarfara are recorded from adjacent counties, often commonly (Rhoads & Klein 1993). shat
records are rather surprising. Two of the county records listed in this paper are
new state records. Carex crawfordii has been reported from Pennsylvania (Rhoads & Been

84 BARTONIA

1993) based on one collection [Pike Co.: Twin Lakes, Bright 19320, 29 Jun 1944, two sheets
at herbarium of Carnegie Museum of Natural History (CM)]. These specimens were recently
re-determined by Anton A. Reznicek as the common Carex scoparia Schk. (A.A. Reznicek
pers. comm.; S. Thompson pers. comm.), eliminating C. crawfordii from the recorded flora
of Pennsylvania. Thus our collection (also determined by Reznicek) appears to be the first
of C. crawfordii from the state. Carex texensis also appears to be a new state record; it is
not included by Rhoads & Klein (1993).

Though C. texensis is native in the eastern United States (Gleason & Cronquist 1991), this
species is probably not native to Pennsylvania. Carex texensis is a weedy species that often
invades lawns (Waller 1929). Its geographic range seems to be expanding northward and
eastward, often in cemetery lawns (A.A. Reznicek pers. comm.), the habitat in which we
found it. Like C. texensis, C. crawfordii is native in the eastern United States, but farther
north (mostly north of Pennsylvania). Though we collected it with several introduced
species in a disturbed area at the edge of a railroad yard, it is likely native to Pennsylvania.
Evidence for this contention includes (1) the frequent occurrence of native populations of C.
crawfordii in disturbed areas (A.A. Reznicek pers. comm.), (2) its occurrence in three
counties of nearby New Jersey, all farther south than our locality (Hough 1983), and (3) its
occurrence in Tioga County, New York, which borders Bradford County on the north (New
York Flora Association 1990).

Carex glaucodea, C. leavenworthii, C. mesochorea, and C. willdenowii are somewhat
disjunct from their other Pennsylvania occurrences. Carex glaucodea and C. willdenowii are
near the northern limits of their ranges, at which their populations are quite scattered (Naczi
pers. observation). Interestingly, C. leavenworthii and C. mesochorea occurred in the same
cemetery lawn as C. texensis. Like C. texensis, these species appear to be expanding their
ranges; they may be adventive in Bradford County.

Our discoveries of Coincya monensis and Sisymbrium loeselii are surprising, too. Rhoads
& Klein (1993) record the introduced C. monensis from only Clearfield and Luzerne
counties, but they report that it is expanding its range and suggest that it is probably more
frequent than herbarium records indicate. Our two collections from Bradford County support
this suggestion. Previously, S. /oeselii was known in Pennsylvania from only Pike County,
apparently from one population (Rhoads & Klein 1993). Gleason & Cronquist (1991)
comment that this native of southeastern Europe and western Asia is only "occasionally
found in our range.”

Most of the species recorded in this paper are infrequent to abundant in Pennsylvania, but
three are plants listed or proposed for listing under the Pennsylvania Wild Plant Conservation
Act: Carex crawfordii, C. mesochorea, and Panax quinquefolius (Rhoads & Klein 1993).

We gathered a wealth of county records in only 7 days in a brief period of late spring.
Apparently, floristic knowledge of Bradford County had been rather incomplete prior to our
collecting. The county’s flora is better known, and further exploration will doubtless yield
more records.

ACKNOWLEDGMENTS

We thank Dr. Ann Fowler Rhoads (Morris Arboretum) for providing a list of the plant
taxa previously recorded from Bradford County and various statistics describing them, all
from the Pennsylvania Flora Database; Dr. Anton A. Reznicek (University of Michigan) for
identifying some of our sedge specimens and sharing his knowledge of sedge distribution,

BRADFORD COUNTY FLORA 85

ecology, and literature; and Dr. Alfred E. Schuyler (Academy of Natural Sciences of
Philadelphia) for determination of a specimen of Scirpus. The late A. B. Cox showed the
senior author many Bradford County habitats that proved fruitful in the search for
noteworthy records. National Science Foundation doctoral Dissertation Improvement Grant
BSR-9001260 provided financial support for the 1990 field work; Northern Kentucky
University’s Department of Biological Sciences and Office of Research, Grants, and
Contracts contributed financially to the 1994 field work.

LITERATURE CITED
GLEASON, H. A. & A. CRONQUIST. 1991. Manual of vascular plants of northeastern United States and adjacent

: nx.
HouGH, M. Y. 1983. New Jersey wild plants. Harmony Press, Harmony.
NEW YORK FLORA ASSOCIATION. 1990. Preliminary Vouchered Atlas of New York State Flora. New York State

PENNSYLVANIA FLORA DATABASE. 1995. Morris Arboretum, University of Pennsylvania, Philadelphia.

Ruoaps, A. F. & W. M. KLEIN, JR. 1993. The Vascular Flora of Pennsylvania. Annotated Checklist and Atlas.
American Philosophical Society, Philadelphia

WALLER, A. E. 1929. Origin of cultivated plants as illustrated by a sedge. Ecology 10: 415-419.

Re

Nea ee

aia:

Sey

ele as \
pS i ae

Pap

PRET

e254

Ha!

Css

2

eae

UNG.

ry

oyveat

Bartonia No. 59: 87-93, 1996

Population Study of Eriophorum gracile Koch
(Cyperaceae) at its Southern Range Limit in Pennsylvania

CAMILLE BARR'
Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018

In 1989 a new population of Eriophorum gracile Koch was found in northeastern
Pennsylvania in Pike County. It has been found at three other sites since 1988; however,
it was reported extirpated at twenty-two historical sites (Pennsylvania Natural Diversity
Inventory). A similar trend is occurring in New Jersey, with one possible extant population
and at least nine historical. The range of E. gracile is circumboreal and extends south in
North America to Pennsylvania, New Jersey, Indiana, Iowa, Colorado, and California
(Gleason & Cronquist 1991). Its southern limit closely follows that of northern dwarf-shrub
bogs and the terminal moraine of the Wisconsin glaciation which retreated approximately
20,000 years ago (Lynn & Karlin 1985). Eriophorum gracile is classified as endangered in
Pennsylvania by the Pennsylvania Department of Environmental Resources (Commonwealth
of Pennsylvania 1993).

Because Pennsylvania represents part of the southern limit of the plant’s geographical
range, the question arises: Are factors associated with the habitats at the southern limit of
this species’ range reducing the fecundity and/or genetic composition of these populations
causing the species to become rare? Because both reduced fecundity and genetic diversity
can limit a population’s ability to rebound from extirpation, these two parameters were
investigated using the newly-found Pennsylvania population of E. gracile. Genetic diversity
was measured using allozyme electrophoresis. Because comparisons between congeneric
species have been shown to be illustrative of the causes and consequences of biological rarity
and commonness (Pleasants & Wendel 1989; Cabin et al. 1991; Standley & Dudley 1991;
Standley et al. 1991) reproductive success was investigated here by comparing seed set
between E. gracile and E. virginicum L., a species which also grows in Smith’s Swamp bog.

There are numerous possible causes behind the loss of E. gracile from its historical sites.
The first, physical disturbance, is a strong possibility considering the extensive development
in the northeastern United States and the evidence of physical disturbance completely
eliminating some bog habitats and their associated plants. Sites that have undergone
disturbance in the form of beaver damming, man-made damming, road building, construction,
peat mining, etc., can experience ecological changes such as increased sedimentation, altered
Water flow and nutrient flux, and actual habitat elimination making the area unsuitable for
E. gracile survival. Stewart & Nilsen (1993), in a vegetative comparison of five bogs in
West Virginia and Tennessee, observed that the one site impacted by agricultural fields, with
resultant higher nutrient levels than the other four sites, was the only one that did not contain
Eriophorum virginicum, another bog sedge, and other species associated with more nutrient-
Poor bogs. A second possibility is that the historic populations were sink populations (sensu

eens
‘Current address: Department of Botany, University of Hawaii, Honolulu, HI 96822

87

88 BARTONIA

Pulliam 1988) dependent upon outside source populations for recruitment. If a change in
propagule dispersal from source to sink occurred, the sink population could die out. A third
possibility is that succession, either on a site-specific or regional scale (such as occurs with
the retreat of glaciers and the associated climatic changes) is affecting the habitat suitability
for E. gracile. Populations at geographical limits would seem to show the effects of climatic
changes more noticeably than those more centrally located. Although the literature on the
ecological consequences of populations at their geographical limits is sparse, those studies
that have been performed implicate both biological and climatic causes of rarity (Pigott &
Huntley 1978, 1981; Weis & Hermanutz 1988; Pigott 1989; Neilson & Wullstein 1983;
Houle & Bouchard 1990; Cabin et al. 1991). Finally, these habitats might be inferior in
quality to other, more centrally-located areas, resulting in reduced fecundity and genetic
variability of the populations, and making them more susceptible to extirpation by stochastic
events. It is this last theory which I chose to investigate.

The study site is a glacial kettlehole bog approximately one hectare in size in Pike County,
Pennsylvania (Table 3). It is surrounded by a large highbush blueberry (Vaccinium
corymbosum)—red maple (Acer rubrum) swamp. The bog shows clear concentric zonation
from larch (Larix laricina) and black spruce (Picea mariana) at the periphery, grading to a
grounded bog mat dominated by leatherleaf (Chamaedaphne calyculata) and Sphagnum spp.,
to a floating mat of sedges (Carex canescens, Carex limosa, Rhynchospora alba, and
Eriophorum gracile) along with Xyris montana and Sphagnum spp. (Sphagnum cuspidatum,
S. fallax, S. magellanicum, and S. recurvum), and finally to an area of open water roughly
30 m across. Shrubs are low, and the few trees that are in the bog are generally not over
2m. Eriophorum gracile occurs as a pioneer species estimated at 5000 individuals at the
edge of the open water in dense clumps and forms a floating mat of vegetation. Individuals
nearer the mat of Sphagnum and Chamaedaphne are partially rooted in the substrate, and are
scattered throughout this mat. The pH of water in the floating mat is about 4.0, and the
whole bog is exposed to full sun.

METHODS

DENSITY COUNTS. Density counts along a microsite gradient were taken from m’ quadrats
along a 6 m long transect during the period of May 25—July 14, 1991. The transect ran from
grounded bog mat containing Chamaedaphne calyculata, and Sphagnum spp., to open water,
to a small grounded island ~2m in diameter containing C. calyculata and two small trees of
Larix laricina and Picea mariana, each approximately 2 m in height. The transect then ran
back to open water. Numbers of vegetative and flowering shoots were recorded for each
quadrat and compared to microsite type which was either grounded bog mat dominated by
shrubs of C. calyculata and Sphagnum spp., open water, or a combination of both. This was
done to determine any differences in plant densities between microsites, and thus
microhabitat preference of E. gracile. Associated species in each quadrat were also recorded.

ENZYME ELECTROPHORESIS. Leaf material was collected from 5 plants each in 5 semi-
distinct clumps of plants around the bog. Well-spaced, selected samples were taken
unsystematically in early September 1992 from each clump in order to maximize the
likelihood of sampling all genotypes in the population. Starch gel electrophoresis was Tun
on fresh and frozen material following the protocol of Soltis et al. (1983). Frozen samples
were quick-frozen in liquid nitrogen then stored at -80°F. Gels were stained for alcohol
dehydrogenase (ADH), malate dehydrogenase (MDH), phosphoglucoisomerase (PGI), >

PENNSYLVANIA POPULATION OF ERIOPHORUM GRACILE 89

phosphogluconate dehydrogenase (6-PGD), fructose-1,6-diphosphatase (F-1,6-DP), shikimate
dehydrogenase (SKDH), adolase (ALD), and glutamate oxaloacetate transaminase (GOT).
The first five ran on a discontinuous citric acid-histidine-HCI buffer system; SKDH and ALD
ran on a continuous tris-citric acid buffer system; and GOT ran on a discontinuous lithium
borate and tris-citrate buffer system. All buffer system recipes came from Soltis et al.
(1983). Staining recipes for PGI, 6-PGD, ALD, F-1,6-DP, MDH, and SKDH were obtained
from Soltis et al. (1983); that for ADH from Vallejos (1983); and that for GOT from Harris
& Hopkinson (1976). All putative loci were consistently scorable across all gels. All
enzymes were interpreted as dimeric except ALD which was tetrameric, and SKDH, which
was monomeric.

FECUNDITY. Twenty-seven inflorescences were collected from the same 5 clumps as the
leaf material on July 14, 1991 after fruit was set and spikelets were just beginning to
disassociate. These were dissected by spikelet. From these spikelets, numbers of spikelets
per reproductive ramet, total numbers of flowers (determined after the flowers had senesced
by counting the persistent scales, each of which originally subtended a single flower), total
numbers of presumed seeds (determined by enlarged ovaries), and the total maximum
number of presumed viable seeds (determined from visual inspection of the seeds—those that
were at a minimum 3/4 swollen with a hard seed and containing white-colored (as opposed
to yellow or dark) endosperm were judged to be filled and potentially viable) were counted.
Potential viability, which was calculated as the number of flowers produced per reproductive
ramet, and actual viability, which was calculated as the maximum number of potentially
filled seeds per reproductive ramet, were determined. From these measures, average
potential viability, average actual viability, and average total percent viability (actual seed
Set/potential seed set) were then calculated.

RESULTS

DENSITY COUNTS. The density counts, habitat types, and list of associated species of each
quadrat are listed in Table 1. In numbers of vegetative shoots, the plant clearly grew better
in the open water microsite, confirming its role as a pioneer species. Numbers of flowering
shoots did not directly reflect the microsite type except that there were no flowering shoots
on the quadrat that was completely on the grounded mat; thus effect of microsite on total
fitness of the plant is not certain, except that little growth—vegetative or reproductive—oc-
curred on the completely grounded mat. ae

ENZYME ELECTROPHORESIS. The eight enzymes that stained showed no variation in
putative loci among the 25 plants. Two enzymes, ADH, and SKDH, were polymorphic and
heterozygous indicating that the population sampled is homogeneous.

FECUNDITY. Of 2517 flowers examined, only 116 actually developed into potentially
viable seeds resulting in an actual fecundity of 4.3 viable seeds/plant (Table 2). The actual
ecundity may be even lower as the assessment of potentially viable seeds was done
conservatively. This is in sharp contrast to Eriophorum virginicum which had an actual
fecundity of 200 viable seeds/plant. While the number of E. virginicum plants with intact
spikelets used was only one compared to 27 of E. gracile, examples of a number of partially
fallen apart E. virginicum spikelets showed that nearly all scales contained a swollen, hard
seed. This was never seen with E. gracile spikelets. Total percent fecundity was 4.6% for
E. gracile and 87.3% for E. virginicum.

90 BARTONIA

Table 1. Density counts per m’ of E. gracile along a 5 m long transect of microsite gradients and associated
species

No. of Shoots?

Quad.' Veg. Flow. Total Habitat Type Associated Species
bh Ae 15 184 Half grounded mat, half © Chamaedaphne calyculata, Sphagnum spp., Carex
open water canescens, cei pens ore limosa
< 2ee 12 598 Open water Utricularia sp., palus:
ns jase Ai 0 101 ‘Island grounded mat Chamaedaphne abet er ae spp., Carex
canescens, Carex lim

4 53 10 63 ‘Partial island grounded cisunaaiaiied ee Carex limosa, Carex
mat, mostly open water canescens, Sphagnum spp., Xyris montana
Rhynchospora alba, Drosera rotundifolia, Drosera
intermedia
pa 7 190 Open water Carex limosa, Carex canescens, Drosera intermedia,
Chamaedaphne calyculata (j

!
ape = “a of vegetative shoots, Flow. = No. of flowering shoots, Total = Total number of shoots.

Caution must be used comparing different species. E. virginicum shoots grow widely
separated from each other and the species does not appear to be clonal at this site. These
factors may cause it to rely more strongly on seed production and/or on longer rhizomes
than E. gracile, therefore the two species may have differing growth and reproductive
strategies, and different morphologies.

Table 2. Quantitative seed data of Eriophorum gracile compared with E. virginicum.

E. gracile E. virginicum

Total no. reproductive ramets 27 1
Total no. spikelets 86 7
Total no. flowers 2517 229
Total no. seeds produced 1456 200
Total max no. filled seeds 116 200
Avg. no. spikelets/reproductive ramet 3.2 7

vg. potential viability (no. flowers/reproductive ramet) 93.2 229
Avg. actual viability (max. no. filled seeds/reproductive ramet) 43 200
Avg. total % viability ({actual viability/potential viability] x 100) 4.6 87.3

hore ios nore a

DISCUSSION

Allozyme results show that all 25 individuals examined were homozygous for 6 loci and
heterozygous with the same alleles for ADH and SKDH, suggesting that the population of
Eriophorum gracile might have been established from a single founder individual. The
homogeneity of allozyme loci and the low rate of seed set compared to its congener E
virginicum also indicate that genetic self-incompatibility mechanisms might be operating i
this species. Self-incompatibility with lack of exogenous pollen flow could lead to the
absence of successful sexual reproduction. Because the Smith’s Swamp population is iso-

PENNSYLVANIA POPULATION OF ERIOPHORUM GRACILE 91

Table 3. Extant and Historical Sites of Eriophorum gracile Koch. in Pennsylvania, recorded by herbarium
specimens.*

PIKE COUNTY. Extant site 2.1 km N of Lords Valley, Smith’s Swamp: bog, very edge of Sphagnum mat with
Carex limosa, 13 May 1991, C. Barr, PH; edge of Sphagnum mat with Carex limosa, 13 May 1991, C. Barr, PH;
edge of bog, with Carex limosa and Sphagnum, 16 May 1991, C. Barr, PH; abundant at edge of mat and in open
water with Carex limosa, 16 May 1991, A. Rhoads, MOAR; in open water adjacent to bog mat, with Carex
canescens and Carex limosa, abundant, 16 May 1991, A. Rhoads, MOAR.

BUCKS C . Argus [Bucks and Montgomery cos.], 3 May 1893, J. Crawford, PH. Nockamixon Rocks,
Narrowsville, in a meadow, 30 May 1893, B. Heritage, PH; 30 May 1893, S. Brown, PH; 30 May 1893, C.D.
Lippincott, PH; bog near Nockamixon Cliffs, 30 May 1893, T. C. Porter, PH; 30 May 1893, S. Brown, PENN. Ca.

mi NW of Pleasant Valley, boggy swale, head of tributary of Cooks Creek, 9 Jun 1923, B. Long, PH.
Quakertown, tunnel [prob. same as Rockhill], a Jun 1866, I.C. Martindale, PH; near Quakertown ‘ied same as
Rockhill], 3 Jun 1894, S. Brown, PH; Quakertown [prob. same as Rockhill], 3 Jun 1894, S. wn, PENN;
Rockhill, Jul 1881, C.D. Fretz, PH; 1 Jun nor J.B. Brinton, PENN; bog, 31 May 1903, A. aa: ht 30 May
1906, S. Brown PENN; 30 May 1906, E.B. Bartram, PENN; small bog rt. of RR above station, 30 May 1906, S.S.
VanPelt, PH. Sellersville, 1866, A.P. Garber, PH; near Sellersville, on trap rock range, N. Penn RR, abundant, Jun
1866, I.C. Martindale, PH. Springfield Township [prob. same as Pleasant Valley], 5 Jun 1899, C.D. Fretz, PH.

MONTGOMERY COUNTY. No locality or date, A.P. Garber, PH. Sassamansville, 31 May 1910, JM. Mumbauer,
PH

DELAWARE COUNTY. Rhodes Swamp, Marple, swamp above Rhodes’ [Tan?] yard, undated, A.H. Smith, PH;
Rhodes Swamp, Marple, undated, A.H. Smith, PH; Rhodes Swamp, Springfield, Jun 1864, C.E. Smith, PH; Rhodes
Swamp, Marple, 18 May 1865, T.C. Porter, PH; Rhodes Swamp, Marple, Springfield Township, Jun 1865, A.H.
Smith, PENN.

BERKS COUNTY. 0.25 mi N of Gouglersville, rare and local in bog, 24 May 1941, W. Wilkens, PH; 0.5 mi N
of Gouglersville, alt. 700 feet, marsh, 23 May 1943, D. Berkheimer, CM. Lobachsville, bog, 27 July 1915, B. Long,
PH. Seiningers Swamp, Spring —— 14 May 1881, TJ. Oberly, PH.

N Hellertown, 1863, T.C. Porter, PH. Johnsonville, wet meadows along trolley road,
30 May 1908, CS. Williaa, at lohenueille: bog south of RR, 0.25 mi E of station, 31 May 1908, S.S.
VanPelt, PH; 0.25 mi E of Johnsonville, along DL&W RR, 5 Jun 1910, H.W. Pretz, PH. Mount Bethel, 31 May
1908, E. B. Bartram, PH. Williams Township, meadow, 4 Jun 1868, T.C. Porter, PH.

LEHIGH COUNTY, Allentown, 3.25 es SSW of Center Square, open, marshy spring partly scrubby place along
south side of stream, 2 Jun 1935, H.W. Pretz, PH. 1.5 mi E of Center Valley, meadows along streamlet, 9 Jun 1912,
H.W. Pretz, PH. 2.5 mi ESE of souskaad? below Swamp Church, swampy place along county line road, 21 May
1911, H.W. Pretz, PH. 2.13 mi SSE of Limeport, marshy meadows heading Licking Creek, 14 Jun 1914, H.W.
Pretz, PH. 0.75 mi NNE of Sigmund, in a meadow beside Indian Creek, 25 Jun 1916, H.W. Pretz, PH. Sigmund,
open marshy meadows north of road just east of Sigmund PO (Hampton Furnace), 17 Jun 1923, H.W. Pretz, P
About | mi SSE of Spring Valley, in the meadows along streamlet on Triassic ridge [could be same as Center
Valley], 13 May 1911, H.W. Pretz is

RREN COUNTY. Extant site near Columbus, Toplovich Bog, small kettle pond (less than .1 acre) surrounded
by Chamaedaphne, 29 May 1990, J.K. Bissell et al., CLM. Extant site about 1.7 km N of Columbus, small stand
in open wet peaty depression fed by seep at base of low slope, 25 Aug 1993, A.E. Schuyler 7837, PH.

ERIE COUNTY. One extant site at Edinboro Lake, N shore, fen meadow shrub thickets along fen spring channel
on embankment, 21 Jul 1988, J.K Bissell et al, CLM; Carex hummocks along W side of channel within center of
fen, 10 Jun 1989, J.K. Bissell et al, CLM. Presque Isle, 30 May 1879, G. Guttenberg, CM; Presque Isle, N Misery

ay, wet meadow, J. Miller, CM.

“Abbreviations of herbaria are those of Holmgren et al. 1990.

lated from other conspecific populations, wind- or insect-mediated exogenous rate flow

is likely to be limited to nonexistent. The presence of E. gracile filled seeds, though low

in number, and of marginal quality, might be explained by apomixis. Alternatively, self-

compatibility with lack of subsequent seedling establishment could also explain the results.
Suboptimal habitat quality may also be affecting reproductive capability. Habitats south

of the Wisconsin terminal moraine do not have the qualities of the glaciated region; even

habitats at the edge of the moraine could experience southern influences. Clonal structure

92 BARTONIA

itself may limit successful fertilization in self-incompatible species (Handel 1983; Aspinwall
& Christian 1992), however other factors such as temperture-limited ovule development,
insufficient time to mature fruits due to short growing seasons, and decreased pollen viability
and/or volume have been implicated as reducing successful sexual reproduction of a northern
clonal plant species (Weis & Hermanutz 1988) and could also be acting at other range limits.

Eriophorum gracile in this habitat is likely to be in an early vegetatively colonizing stage
and its survival depends upon a stable habitat until the population develops adaptive genetic
variability. However, because the E. gracile population studied is farily isolated from other
conspecific populations, the chance for building variability by combination of alleles is
drastically reduced.

Further research is needed to determine the causes of extirpation of E. gracile at its
southern limit. It is first necessary to perform comprehensive site surveys of all historical
sites to verify that E. gracile is surely gone from them. Table 3 lists herbarium specimen
data of E. gracile from Pennsylvania with indications whether it is extant or extirpated at
sites where it was previously collected. Repeating the same experiments performed in this
study on peripheral and more centrally-located populations and comparing the results
obtained there would be useful to determine whether low fecundity and low genetic
variability are correlated with the southern limit. Investigating the breeding and dispersal
systems of E. gracile would be extremely useful for clarifying the genetic and reproductive
dynamics of the populations. Finally, comparative water and substrate chemistry analyses
of the habitats that have lost E. gracile with habitats supporting vigorous populations could
indicate possible environmental changes that are associated with extirpation.

ACKNOWLEDGMENTS

I thank Drs. Ann Rhoads, Brenda B. Casper, and Alfred E. Schuyler for all their help,
resources, and inspiration. I also thank the Pennsylvania Department of Environmental
Resources for access to my study site, the Pennsylvania Chapter of The Nature Conservancy
for invaluable information, and E. Karlin for identifying Sphagnum.

LITERATURE CITED

ASPINWALL, N. & T. CHRISTIAN. 1992. Pollination iba seed alg e population savin in Queen-of-
the-prairie, Filipendula rubra (Rosaceae) at Botkin Fen, Miss Amer. J. Bot. 79: 488-4

ae a . RAMSTETTER, & R.E. ENGEL. 1991. Tvolace par siae ? a locally rare ee Rhodora

COMMONWEALTH OF PENNSYLVANIA. Pennsylvania Code, Title 25, Chapter 82. Conservation of Pennsylvania
native wild plants. 23 Pennsylvania Bulletin (October 30, 1993).

GLEASON, H.A. & A. CRONQUIST. 1991. Manual of vascular plants of northeastern United States and adjacent

anada. 2nd ed. New York Botanical Garden, New York.

Gray, AJ. 1984. Genetic change during succession in plants. In A.J. Gray, M.J. Crawley, & P.J. Edwards (eds.)
Colonization, succession, and stability. Blackwell Scientific, London.

HAMRICK J.L. 1983. The distribution of genetic variation within and among natural plant populations. Pages 335-
ni in ie Schonewald-Cox et al. Genetics and Conservation. Benjamin/Cummings Publishing Co., Inc.,

Howe s. N. 1983. Pollination ste plant population structure, and gene flow. Pages 163-211 in L. Real,
ed., Pollination ecology. York.
RRIS, H. & D. A. HOPKINSON. 1976 Handbook of enzyme electrophoresis in human genetics, 2.7.5.1, 1. North-
Holland Publishing Co., Amsterdam.
HOLMGREN, R.K., N.H. HOLMGREN, & L.C. BARNETT. 1990. Index herbariorum part 1: the herbaria of the world.
th ed. New York Botanical Garden, Bronx

PENNSYLVANIA POPULATION OF ERIOPHORUM GRACILE 93

HOULE, G. & F. BOUCHARD. 1990. Hackberry (Celtis occidentalis) at the northeastern .. = by distribution in
North America: eet structure and radial growth patterns. Canad. J. Bot. 68: 2685-

LYNN, L.M. & E.F. KARL 1985. The vegetation of the low shrub bogs of northern ky i and adjacent
New York: Suitie - their southern limit. Bull. Torrey Bot. Club 112: 436-444.

NEILSON, R.P. & L.H. WULLSTEIN. 1983. Biogeography of two southwest American oaks in relation to atmosphere
dynamics. J. Biogeog. 10: oe 297.

PENNSYLVANIA NATURAL DIVERSITY INVENTORY. Unpublished. PNDI sentaeh The Nature Conservancy, The
Western Bie aoa Dolan and DER Bureau of Forestry, Middlet

PicotT, C.D. & J.P. HUNTLEY. 1978. Factors rar the distribution wy Tia cordata at the northern limits
of its geographical range. New Phytol. 81: 429-441.

PicoTT, C.D. & J.P. HUNTLEY. 1981. Factors controlling the distribution of Tilia cordata at the northern limits
of tis geographical range. III Nature and causes of seed sterility. New Phytol. 87: 817-839.

PicotT, C.D. 1989. Factors controlling the distribution of Tilia cordata at the northern limits of its geographical
range. IV Estimated ages of the trees. New Phytol. 112: 117-121.

PLEASANTS, J. M. & J. F. WENDEL. 1989. Genetic erg in a clonal narrow endemic, Erythronium propullans,
and in its widespread progenitor, Erythronium albi Amer. J. Bot. 76: 1136-1151.

PULLIAM, H.R. 1988. Sources, sinks, and population po aa Amer. Nat. 132: 652-661

SOLTIs, D.E., C.H. HANFLER, D.C. DARROW, & G.J. GASTONY. 1983. Starch gel electrophoresis of yom a
compilation of iflinding buffers, gel and electrode buffers, and staining schedules. Amer. 3

STANDLEY, L.A. & J.L. DUDLEY. 1991. Vegetative and sexual reproduction in the rare sedge, Carex ss

STANDLEY, L.A., J.L. DUDLEY, & L.P. BRUEDERLE. 1991. Electrophoretic variability in the rare sedge, Carex
-450.

polymorpha (Cyperaceae). Bull. Torrey Bot. Club 118: 444
TEWART, C.N. & E.T. NILSEN. 1993. Association of edaphic factors and vegetation in several isolated

Appalachian peat bogs. os Torrey Bot. Club 120: 128-135.
U.S. GEOLOGICAL SuRVEY. Rowland, PA (map). 1:24000 7.5 minute series (topographic). Wahsington, DC.
G

—
VALLEJOS, C. E. 1983. Enzyme ial ee Pages sa in 8. D. Tanksley & T. J. Orton, eds., Isozymes

in plant genetics ne breeding, fe A. Elsevier, Amste
WEIS, I. M. & L. A. HERMANUTZ. ee The population biology nie the arctic dwarf birch, Betula glandulosa: seed

rain and the secndiable seed bank. Canad. J. Bot. 66: 2055-2061.

eohaiics

Sant Pests cer mee s

pris ooh he tale eae Ge

Bartonia No. 59: 95-96, 1996
Taxonomic Status of Panicum hirstii Swallen

ALFRED E. SCHUYLER
Academy of Natural Sciences of Philadelphia
1900 Benjamin Franklin Parkway, Philadelphia PA 19103-1195

Panicum hirstii Swallen was first collected in Sumter County, Georgia, in 1900 and
"doubtfully referred to" Panicum roanokense Ashe a decade later (Hitchcock and Chase
1910). In 1958 it was discovered in Atlantic Count, New Jersey, and described as a new
species soon afterward (Swallen 1961). More recently, P. hirstii was considered to be a
probable "glabrous variant of Panicum neuranthum Griseb." (Kral 1983) or conspecific with
Panicum aciculare Desv. (Gleason and Cronquist 1992). In order to determine if P. hirstii
warrants recognition as a species, I studied plants identifiable as it, P. roanokense, P.
neuranthum, and P. aciculare in the field and herbarium in 1993. Plants of P. hirstii and
P. roanokense were examined in the field in Sussex County, Delaware, P. aciculare in
Wicomico County, Maryland, and P. neuranthum in Baker County, Georgia. Numerous
herbarium specimens of all taxa were examined in collections at the Academy of Natural of
Sciences (PH), New York Botanical Garden (NY), Smithsonian Institution (US), and
Vanderbilt University (VDB).

As do its relatives, P. hirstii produces erect leafy stems with terminal inflorescences early
in the season, then develops axillary shoots with inflorescences later in the season, and
finally persists as low over-wintering leafy rosettes close to the ground. The axillary shoots
are sparingly branched and not much reduced. The slender plants grow to about one meter
tall and have narrow inflorescenes with upwardly appressed branches. The pedicels are less
than twice as long as the glabrous elliptic spikelets. The first glume is short and broadly
obtuse, the second glume and sterile lemma have nine nerves, and a hyaline sterile palea is
present. The plants grow in ponds or on pond bottoms with sandy peaty substrates, which
intermittently have standing water. Considerable water level fluctuation may occur
throughout the growing season.

Panicum hirstii is more similar to P. roanokense than to any other described taxon. The
latter is considered conspecific with Panicum (Dichanthelium) dichotomum by some authors
(Gleason and Cronquist 1992; Gould and Clark 1978). Both P. hirstii and P. roanokense
have glabrous spikelets about the same size, both have about equally prominent nerves in
the leaves, second glumes, and sterile lemmas, and both have similar hyaline sterile paleas.
Panicum roanokense is readily distinguished from P. hirstii, however, by having more
branched and reduced axillary shoots, more open inflorescences with pedicels often twice
or more as long as the spikelets, and more variation the number of nerves (7-9) in the second
glumes and sterile lemmas. Instead of ponds or pond bottoms, P. roanokense is found in
Swales, meadows, swamps, and low woods (Fernald 1950; herbarium specimen data).

Panicum hirstii bears a superficial resemblance to P. neuranthum, which is considered
conspecific with Panicum (Dichanthelium) aciculare by some authors (Gould and Clark
1978). The following characters that distinguish P. hirstii and P. neuranthum, also effectively
distinguish P. hirstii and P. aciculare sensu stricto. The inflorescences of P. neuranthum
may be narrow with upwardly appressed branches and resemble those of P. hirstii to some

95

96 BARTONIA

extent. Plants with open inflorescences, however, may grow in the same populations with
plants having upwardly appressed inflorescenes. In contrast to P. hirstii, P. neuranthum
often has pedicels over twice as long as the spikelets and also has much more prominent
nerves in the leaves, second glumes, and sterile lemmas. It further differs from P. hirstii by
having pubescent spikelets, seven instead of nine nerves in the second glumes and sterile
lemmas, and no sterile paleas. The habitat is drier with P. neuranthum more often found in
savannas (Hitchcock and Chase 1910) and low sandy peaty areas that are intermittently wet
(herbarium specimen data). It is not known from ponds or pond bottoms although it may
occur in low areas adjacent to ponds.

No apparent infraspecific variation was encountered in specimens of P. hirstii throughout
its limited range from New Jersey to Georgia. Its range within the four states from which
it is known is very local: two sites in Atlantic County, New Jersey; one site in Sussex
County, Delaware; two sites in Onslow County, North Carolina; one site in Sumter County,
Georgia; and one site in Calhoun County, Georgia (data from herbarium specimens at NY,
PH, and US). Its limited occurrence may be attributed to its specialized habitat in ponds
characterized by considerable water-level fluctuation.

ACKNOWLEDGMENTS

Dana Peters arranged for this study to be funded by the U.S. Fish and Wildlife Service.
Patricia Holmgren (New York Botanical Garden), Warren Wagner (Smithsonian Institution),
and Robert Kral (Vanderbilt University) provided access to herbarium specimens at their
respective institutions. For help, directions, and/or guidance with field work, I thank Jim
Allison, Frank Hirst, Robert Kral, William McAvoy, Robert Norris, Tom Patrick, Patricia
Schuyler, and Ron Wilson. Terrance Mann prepared the manuscript for publication.

LITERATURE CITED

FERNALD, M. L. 1950. Gray’s manual of botany. 8th ed. American Book Company, New York.

GLEASON, H. A. & A. CRONQUIST. 1992. Manual of ene plants of northeastern United States and adjacent
Canada. 2nd ed. The New York Botanical Garden,

GOULD, F. W. & C. A. CLARK. 1978. Hidliontbelien reed in the United States and Canada. Ann. Missouri
Bot. Gard. 65: 1088-1132.

HITCHCOCK, A. : & A. paper 1910. The North American species of Panicum. Contr. U.S. Natl. Herb. 15.

KRAL, R. 1983. A report on some rare, threatened or —~ forest-related vascular plants of the South. USDA
Forest aig Technical Publication R8-TP2, A 2Vv

SWALLEN, J. R. 1961. A new species of Panicum sats New Jerscy. Rhodora 63: 235-236.

Bartonia No. 59: 97-106, 1996

Soil Chemistry Data for Species of Special Concern in
Pennsylvania

RICHARD MELLON
Mellon Biological Services, P.O. Box 63, Morrisville, PA 19067

While most vascular plants in Pennsylvania were described over 100 years ago, I am
unaware of any single source for detailed habitat requirements for these species. With the
increasing emphasis on habitat creation and restoration, and reintroduction projects, soil
chemistry characteristics for individual species becomes increasingly important for successful
plantings. This may be especially true for rarer species, since generally they either have
limited habitat within the state or their habitat requirements are very narrow. Unfortunately,
definitive soil chemistry data for most species are limited or nonexistent.

Over the last 15 years, I have collected, and have had analyzed, soil samples from 360
sites in eastern Pennsylvania (as far west as Fulton and Tioga counties) and Central and
Southern New Jersey. These samples were collected as parts of various studies for
Bowman’s Hill Wildflower Preserve Association, the Pennsylvania Department of
Environmental Resources (as part of the field team for Morris Arboretum of the University
of Pennsylvania and separately), the National Park Service study of the Upper Delaware
Scenic and Recreational River (Morris Arboretum), and numerous individual consulting
wetlands projects.

METHODS

Soil was collected from the root zone adjacent to the species listed. This zone varied,
depending on the habitat, from barely decayed peat in bogs, to the "Ao" horizon, the "A"
horizon and occasionally the "B" or "C" horizons in disturbed areas. The soils were stored
in plastic bags, then air dried, screened and sent to the Agricultural Analytical Services
Laboratory (formerly known as the Merkle Laboratory) at The Pennsylvania State University,
University Park, Pennsylvania. The laboratory tested for calcium, magnesium and potassium
(exchangeable cations [milli- equivalents/100 grams] and percent base saturation), phosphorus
(pounds/acre), pH, cation exchange capacity and organic matter (ash combustion). Over the
years of collecting data, the method of measuring organic matter changed from a titration
method, which had limited accuracy over about 5 percent, to an ash combustion technique,
which is currently used. Therefore, all pre-ash combustion data were eliminated from
calculations and left blank in Table 1. The nomenclature follows Rhoads & Klein (1993),
€xcept for Dodecatheon amethystinum, which follows Gray’s Manual (Fernald 1950).

RESULTS

A partial list of Pennsylvania endangered, threatened and extirpated plants and the soil

97

98 BARTONIA

chemistry data collected for each are given in Table 1. Since these are very rare species,
multiple sites were not always available. Therefore, the data should be considered as
preliminary. Locality information for the sample sites is given in Table 2.

ACKNOWLEDGMENTS

I thank the Bowman’s Hill Preserve Association, Dr. Ann Rhoads and others at the Morris
Arboretum, the Pennsylvania Department of Environmental Resources, and Ann Newbold
for verifying the identification of the rare plants.

LITERATURE CITED
FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co., New Yo:

rk.
Ruoaps, A. F. & W. M. KLEIN, JR. 1993. The Vascular Flora of Pennsylvania, Annotated Checklist and Atlas.
American Philosophical Society, Philadelphia.

99

SOIL CHEMISTRY DATA FOR RARE PLANTS

69- IP

OPT-€ 17

09°0 €7-61 LI-6 r-£ LI1-901 €r-0'r er

TIFLS TIF67Z = EOF’ LOFS7Z 9 OF8L'0 TFIZ PFE FP SFI Lorl + (4 11suIyjO2 xaaD)
SL-IP SOEGSZ SI-TI L0-+'0 870-070 S-P €-1 I-l 7S-9F 6 ELE €

$1¥09 O7F98T «=O OOFETI 1 0F9'0 ~0'OFET0 OFS 1FZ OF1 €Fos LOF8'€ (9) 4 pipjjng xaivD
i¢

7 VL O€ 01 Le0 tr SI ¢ 8€ 89 q Saplosaisp DIuOIOg
9

Zz 19 WA 80 60°0 Ip rl I sil 8b d Saplojwuapig suapig
—_ 6L-79 LET L1-70 90'0-90°0 Ly-Ll €1-Z I-l TE-8 Ors: 61 ‘St ‘et

_ 9 0FEL 0 1FS7 > 0F9'0 00'0F90°0 €lF9E SF8 OFI OIF6I yOFL’s (¢) 4 sipiqojoads 421sy
8E-87 67ESEL S7-EO ¢0-S°0 07 0-110 8-7 t-Z I-l bE-S SrLe €

SFEE LEPTET SO FH 00FS°0 S0'0F9I'0 €FS IF€ OFI $1F07 POF 'b ML n3jaq-140u 4ajsy
11-8 p6I-LZl 07-07 O'SI-O'1l 07 0-S1'0 91-01 €8-9L I-l 8-9 04-09 SZ “IZ

701 €€Fl9l OOFOT OTTO El €0 0F81'0 eFEl pFO8 OF I IFL ¢ OFS 9 (OL snyosadnvdap saisy
L@-LI SZEBLZ 067-081 ¢ZI-S'8 Ol'I-00'1 89-59 I€-0€ v-Z Zr-0£ SLTL lz

SF7Z yTFTOE S$ SFS EZ 07S 01 $0 0FS0'I TFLO IFI€ FE 9F9E LOFrL (%) 4 suayisaa wniuajdsy
67

er Vil 70 £0 ¢70 I z I 08 Se L iajposq wnjuajdsy
$9

OL 781 ZI 90 400 L v 0 9 Or I psoqjng psnyja4y
Lp

1Z 181 O€ 60 veo LI s z aa sr Is Sisualanossnu siqoay

L6-78 TICS a: Ses €1-9°0 Ol I-ve0 S1-9 St rl €67-7P Crle Zr ‘OF puDIMID4J4Dg

SFE6 OSFI'ST 60FSZ 70760 97 0F09'0 vFOl Fb 1#Z L8FELI 7 O0F0'Y (9) 4 daiyouojauy

310 % og9 Le) aw x 8) 3 x d Hd 07 saioadg
300 [/bout % oB/Sq] a(N) 81S

‘BluBAASUUdd Ul WII0U0D jeIDeds Jo sjuRd Joy BIeP ANSIUIIYS [IOS “| IQRI

IL
OL ‘¥9 “PS

L6O-€ TET OFI-07 rsS-€'0 08°7-01'0 pS-l€ €7-L v1-Z 9Ly-r1 0'9-8°€
€vFE9 PLETyI «= LEFI'9 6 lFr7Z €7 1F bl STEP 9F¢1 SF8 Z9IFOLI 6 OF8'F (9) a =: Saploosiapue wnipy] >)
Is
p9-r1 61E-PLI OSE II OSI-0'F 67 0-110 89-Lb =: OH EZ 1-0 6€-F1 TL09 0S ‘6h ‘BP
61F9p 6VFLST 6TIF6 HT 9 €Fh'6 90 0F61°0 8F6S SFSE OF I 8F67 yo-L9 (Ss) L polupjay Xa)
79-6 61E-Orl OSb-8'6 OSI-O'b 67 0-110 OL-Ly 9-67 7Z-0 6£-+1 84-99 IS ‘0S 6b
6IFLYP 19F797 = € vIFO'8Z 6 FP OL 90 0F7Z°0 6F8S LF8€ IF 6767 vOFOL (9) L SIidays X40
06-8 Pee-sZl «OL 0 9 €-70 0S 1-010 €S-p LI-Z 9-1 771-7 LS-9€ Sb ‘Er ‘bE
87F0P €LFSOZ 7@6Z LIFUT pr 0F6S 0 9IFLI CFS IF€ 9EFP9 9 0FE'P (rl) a pydioudjod xaav>
OL ‘%S
L6°S8 rcesol O8t-ce 68-11 Ob'€-9F'0 SS-Ol 87-P [I-€ $81-89 Srse ‘Ib ‘OP 6E
SF16 TOFLLZ «=LSFO'OI ZT EFE PY €0 IF8L'1 6IFLE —OOLF9I €F9 SPFIEl €OFI bP (9) L pynoaadnod xan
ra)
~ 6 €8 Or! 67 Or'l 6h Ol S Sel Ov q p4oyfionnd xasv)
Z
fe) 19
| ‘09 ‘6S ‘LS
BS L6-0¢ LLES OIL = =SEI-TI S60 IS ZI-Pl 0 9€-L oz I€-1 Peer SSSE ‘SS ‘PS ‘IP
91F68 SSFO7TZ 87489 Vt € €l €Fh6 7 SFO OFS | LFZI 68F7ZL1 CS OFE' YF (OD L putsadsosijo xai0D
19-6 6I1E-Orl OSb86 OSI-0'r 67 0-110 OL-Lbp =: 96 Z-0 6£-07 84°99 IS ‘0S ‘6r
I@FIP S9FPST BS EIFOLZ 0 PF9'6 LO'0F@Z'0 OIF09 = LF9E 1F1 LFIE ¢ OFO'L (bp) L panlf xado)
S£-7Z O8I-ZPl O9I-OTI 07S I $7 0-910 eFis UF1l I-l 87-9 TL89 €€
9FS7Z PIFI9OL SFE PI 7 OF8'I €0°0FIZ0 OFL8 IFZ1 OF I 8F91 L0FO'L (py) pauinga XxadD)
76°L8 697-1 bZ Ob SE 9°S-5'0 09°7-0F'Z 79-95 17-61 11-6 €67-87Z 9S-1'S r9 ‘79
€F06 VIFSST €PFL6E CTFIE 01 0F0S'Z €F6S 1¥0Z 1F01 €£F19Z 7 OFb'S (2) L p4pubip xaiD>
B10 % 99 Ze) 3 x ie) 3W > d Hd 007 saisadg
Sho joan % ov /SqI aN) 22S

100

(p.quoo) | aqe],

101

SOIL CHEMISTRY DATA FOR RARE PLANTS

Le uyjauyorg

rl S€l sl €0 670 II z z sil 8b q wnwayjuDijayy
69

61 691 o's r0 S10 O€ Z l 18 cP q s1jpas0g DLaadp)

ral apyupnovoad!

LI Ll o's 07 170 62 ZI I 091 wy q piqaoydng
p9-SE 6 IES EZ O'SH-O'ST O'SI-S'8 67 0-110 89-Lp = OF IE 1-0 6f-F1 TL99 ~—sIS ‘OS ‘6b UNIDUADIIPLAIA
OFS O€FIBT GTIFOIE 67F9' TI L0 0F77'0 8FSS 9F0P OF I 6FIE € 0F8'°9 (s) L wnsoydoiug

91
IZ Lgl O€l €€ zr'0 8L 0z € p91 69 L pajaiodu visi
PZ snupiuljodp9
II vel €€ Ol IZ°0 vZ L z vz 6% q sndojuvydapq
v-Z LE-BE Z€-0'1 vI-L0 $1'0-L0°0 L797 ~—- O91 7-2 9b7Z-01Z €S-£'S 9
IF€ yTFE9 LIFlZ €OFI'I pO OFIT'O 1FLZ 7F81 OFZ 81F8ZZ O'OFE'S (2) 4 pnaind siapys0ayy
9 laspad “IRA
a 8°01 Ov rl 40:0 Le €l I raul es q DSNIGO StapYy20a/q
Ss

a 8'P ct ZI 010 EL SZ Z €rl 69 q laljom pojysoulyay
$9-6S 67EN6L ET 0'1-9°0 0L'0-7E'0 OC-L 97 €-I Z17-06 Pre 69 ‘89 odaidojAduino
7F19 OSFEEZT BR OFSE 1 OF8'0 Sl OFLS0 SFSI Fb IF€ OSFSLI ¢ OFE'F (p) 4 staajododkq,

97
81 791 Z0l 80 $0 9 8 € €9 6% J = PIppau uoaymwoapog
87 wnursdyaun
~— L97 £tz x4 £50 68 6 z vZ TL L uoaywoapoq
310 % eo): 6) a) 3 x Le) 3 x d Hd 07 saisadg
300] /bow % 9e/Sq[ q(N) lS

(p.1uos) | 21qeL

BARTONIA

102

rr
ol Lvl $0 Z0 Z10 € l l €l le A paopfiun viBsaquayynyy

£9

_ rs st 90 oro or ll € 19 VL a loss}MDYy? DLO)
06-0 LIe-+s8 87-70 LI-10 $7'0-90'0 8I-l Ll 1-0 zs-9 6re€ €

I€FLb 18F90Z LOFO'l € 0FS'0 90 0FE1'0 9FL TZ OF I €IFLI v 0F0'b (3) 1 - pupm3.1a DyousDy
Zp-9Z SEZ-O8I OF€-80 0'I-r'0 Z1'0-40'0 LI-€ $-Z 1-0 Is-91 0S-0'F b‘€

LFvE LaSiZ = O1FS'I € 0F9'0 70 0F01'0 L¥8 IF€ OF I SIFLE SOFE b (OL pjpiadia vyousopy
Lt-v $ 67-0'S ¢ 7-80 O'I-r'0 Or 1-400 O€-F 11-2 9-0 9r-€1 9S-6€ €Z ‘07

8F91 S8FPr Ll LOFT € 0FL0 7S OFOS 0 OIFSI vF9 77 Z1F9Z LOF9'P (4 pupiiput DIuOAT
19-6 61E-OFl O'Sb-0°86 OSI-0'F 67 0-£70 OL-Lp =: 9-6 raat 6£-07 81-99 OS ‘6F

pTFtp LUFL YT «8 vIF6 67 S yF0'01 €0 0FS70 OlFzs =—s_s LF 8E OF I SFO 90FOL (¢) 4 muyoy DYaqoT
lL

€ vr oz £0 1) a) Sb L rd rl 09 L Duupulsop DI/agOT
gS

6 Cll €0l 01 €70 68 6 z tw 9L L 14440} snoune
€s

I o£ 80 z0 £00 4 l I 61 rs q Se
€l

tl yA aa 90 010 0z 8 z 67 oF q snsopfig snounf
6L pSl68 O7-80 L0-€0 910-600 €1-8 t-£ I-I 86-E7 8r-OPr £

178 €€F+7Z1 VD OFFI 7 OFS 0 pO OFE1 0 Fl Fb OF | 8EF19 1 OFL > (04 snmojoysip snouny
Lv-67 €v739 E07 60-10 Z£0-61 0 SS-8 €l-€ €-l L8-SS TOI Pb ZE ‘OE

6F8E S8F9SI 60F6T 1 0F8°0 L0 07970 p7FZE SF8 1F7Z 9IFIL 11F7's WL povdo xajj

310 % 949 8) 3 | 2) aw >| d Hd 07 saisadg

oe

Seoi hon % /sq\ oN) BIS

(p,juo0s) | age]

103

SOIL CHEMISTRY DATA FOR RARE PLANTS

|
91 ssl Or! 97 ero €L rl z is 19 L pppnpofial Baad
of
Ll ssl 0 z0 110 € I 1 tr 6¢ a Dywruapiay DIMA”
uw
IL-1Z EvVEOIZT OLIVE 07-01 06°0-Z£'0 rS-81 8-S rl 9IE-8E TSSb ‘99 ‘9S ‘SE
91F9S 9PF99T 9FF06 OFS | 61 0F79'0 ZIFEE #9 {FZ COIFPLI COTS (3) 4 iunag wenyruEtjog
ZL-81 OZI-+6 OS-0F TUE 170-070 €S-€E€ EZ" t-2 €8-Zr 9S-6F at
7iFPr ViFrol 6S OFer PrOrL | 0F070 SFU SF91 oFz LIFs9 cores (g) 1 smusngos wmuo3ijog
1z
7 er 80 10 S00 fal 0z I 9 6 aq ussiun> vppBjog
98-98 SEECIE OFl-0'7Z 61-91 91 O-LL0 71-69 xs 0-0 gI-Zl Ls-Ls uw aptunig-uDAa
0F98 €lF97ZE O1FOEZ TOFS’ I ZO OFFI 0 TIL 1F9 0 eFs1 oor. s (Oa MIN IMOMIIO
I
14 v'07 s¢ 07 00 lz ol I 601 oF 2. puadipnyed vog
9-6 61E-OFl OSt86 OSI-0'r 67 0-110 OL-Lb =: 9-6 7-0 6E-+1 8299 IS ‘OS ‘6b
6IFLP 19F79Z = € PIFO'8TZ 6 €Fr 01 90° 0F77'0 6F8S LF8E 1F1 667 bp OFOL (g) 4 ponn]3 visspuswd
Le
o€ 191 30 70 L410 ¢ I I €8 tr q wnsdydoymex wnomg
IL
€ tr 07 €0 oro SP L 4 rl 09 1 minypauar wnypydoudy
€6-bL SOEPLZ OB8I-LE 75-90 08 I-rr'0 $$-Zl 0@-Z Ll IS 1-79 Sree sr
OIFPs LIFI6% = LFEOI € 767 89 0FTZI'I C7FPE Os SFI eFb SPFLOl OFT Pp rales 2y03 voy
BIO % O90 te) 3W x Te) aw S| d Hd 07 saisadg
oe
200 1/bour % /SQ| aN) 81S

€L JRA puns “dss
ZI nig ez 60 070 ze ll € 00°LZ LS q xajduas o3ppyos
$1-9 €9l-L pl E€7-70 80-£ 0 Zr 0-710 SI-Z S-Z €-1 E1E-8 6rSh O16 ‘8 ‘€ wnouyunpio
PFI 9OFTSI = 8 OF O'l 7 OFS'O ZL OFETO SFL IF€ 1F7Z pT1F66 TOFL Pb (>) 4 wniyoursdasis
I€-LZ S6I-LLI O'TI-O0I LUSz 6£ 0-8£ 0 6S-1$ OIE ZZ 8S-6P gS-S's fal
TF67 LOFL81 vv O0FIOI 1 0F97Z OF6E'0 €FS¢ IFS OFZ pFTZS OFS'S (¢) 4 pasoa uinpas
19-6 6I€-O'bl O'Sb-8'6 SI-p 67 0-€7:0 OL-Lb 9-67 rad | 6£-0% 81-99 OS “6r
7-7 LL-Lvt 8 vl-6'67 S yF0l €0 0FS70 OIFLS —s LF BE OFI 8FOE 9 0FO'L (2) 4 DIDI[INAIA DLAJIS
€1-8 pol-LZl = E707 SI-II 07'0-F1'0 91-01 = 98-94 I-l Z1-9 04-09 SZ ‘IZ
TI LATLI dL OFIZ L Ft €0 OFLI'0 TZ p08 OF I 7F6 $ 0FS'9 (pb) L paopfionnd v1sajas
9 qyinus
b CL av a rl O€ LI Z SLI ce q snizajdouaoyog
<
z S-b 9OI-E8 TrOE 07-71 bZ0-910 Or-9€ ~— GIS I 7-2 877-871 9S-S'°¢ 6 ‘7
© IFS V1FS6 9 0FO'E tv 0F9'I 0 0F07'0 TF8E TFLI OFZ OSF8L1 L0F9'S (7) 4 DULIA]OI DLADIN3DS
< 19-6 61E-0'rl O'Sb-8°6 OSI-0'r 67 0-€70 OL-Lp =: 9-67 ZI 6£-07 8-99 OS ‘6b paanjpidos
p7FCPp LLAL YT -8'v1F6'6Z S'PF0'01 €0 0FST0 OIFLS = ss LF8E OF I SFE 9 0F0'L (9) 4 vaodsoysudyy
LIL L8t-Ti 2 'ES'0 L0-£0 0£ 0-600 1€-€ 6-7 tl ZrI-EZ ESTP et
PFT OPFTIL 6 0F9'I 7'0F9'0 80 OF P70 ZIFLI €F9 FE €PrF9pr p 0F6'b (s) 4 pupLiput pIxXayYy
vl 67E68 S70 80-€ 0 9b'0-L0'0 €I-l a | Z-0 86°8 Srl L‘b'€
OCF6E 19FL%Z = SOFT lOFS'0 LL OFSI'O PFS 1¥Z 1F1 67FEE € OF (OD 4 soyjayd snosanO
L
LZ Lz cz 80 9r'0 6 € s 19 I'v q pywajof snosan©
310 % Od9 2) 3 x ie) 3 x d Hd 0] sainads
oB/s A 4
300 [/bow % /SQ| aN) .B1S

104

(p.auos) | a1qe1

105

SOIL CHEMISTRY DATA FOR RARE PLANTS

“WIMAIOGIY SIUOJ 9} 18 SI UaUTIDAds JAaYONOA vB aIROIPUI SIaquINU Plog AY] “Z I[Qe, Ul UdAIT UONRIO| JUEId SI JaquINU = 907,
‘(ajdwies suo Ajuo sem aloy} sues Jaquinu ou) sajduies j1os Jo Joquinu si sasoyjuared ul Jequinu = (WV),
‘paredinxa—x ‘pouajwaiyj—], ‘pasasurpus—q :viuralAsuuag ul smeys sjued = BIS,

9Z-1Z Z9T-SIZT «=. TO 80-L'0 Z€0-S1'0 6-1 €-€ I-l €S-7Z Cre € pupiuong “dss
£Fv7 bv T+6 £7 60FT 1 $0 0FSL0 60 0F¢7 0 p+¢ OFE OF1 O1F8E 1 0FST Pb (2) 4 ppifinpad vjol,
p9-ss 097-F SI i Sal Tl 1-70 710-010 Sint et 1-0 €1-8 6 8e £
p09 CS eFS 17 6 0F6 0 £0FS 0 10 0FI1 0 LFS CFE OF 1 TF01 bp OFl b (s)4 wnpnu WnuAnNg!A
oF
Ol +6 £S Et 8b 0 9S 8l S 617 $9 q ponnjs piuouia,
76718 [p2-ret O6E-1'SE €r-so0 09'T-0F I $9-79 6l-6l 11-9 8T7-8L 9S-7S rs
9FL8 £ OF8 €7 SIFT LE lFr 7 09'0+00 7 1¥¢9 OF61 £46 CLFES1 LoFs's (Z) L Plpatwsajul pLivjnoiaiy)
B10 % fe: 6) Ze) 3W x 3, 3 y d Hd 07 saisadg
Sho] bod ¥ oe/Sq| a(N) .B1S

(p.quos) | 919%,

106 BARTONIA

Table 2. Plant locations corresponding with location numbers in Table 1.

PENNSYLVANIA

Berks Co: Union Twp (1).

Bucks Co: Bensalem Twp (2), Bristol Twp (3-8), Bristol Boro (9, 10), Bristol Twp (11), Falls Twp
(12), Lower Makefield Twp (13), Nockamixon Twp (14), Solebury Twp (15), Upper Makefield
Twp (16).

Chester Co: Coatesville Boro (20), Elk Twp (21, 22), West Whiteland Twp (23).

Delaware Co: Haverford Twp (24), nec Twp (25).

Franklin Co: Montgomery Twp (26, 27).

Lancaster Co: E. Hempfield Twp 8), Hemfield Twp (29), Drumore Twp (30), Martic Twp (31, 32).

Lehigh Co: North Whitehall Twp

Luzerne Co: Dennison Twp (34), Sam Twp (35), Fairview Twp (36), Pittston Twp (37).

Monroe Co: Coolbaugh Twp (38, 39), va Sega Twp (40-42), Tunkhannock Twp (43-45).

Montgomery Co: Lower Providence Twp (46), Upper Hanover Twp (47).

Northampton Co: Hellertown Boro (48), Upper Mt Bethel (49-51).

Pike Co: Porter Twp (52, 53).

Sullivan Co: Colley Twp (54, 55), Davidson Twp (56), Laporte Twp (57).

Tioga Co: Charleston Twp (58), Chatham Twp (59), Shippen Twp (60, 61).

Wayne Co: Buckingham Twp (62), Manchester Twp (63), Mt Pleasant Twp (64), Preston Twp (65-
70), Scott Twp (71, 72), Sterling Twp (72).

York Co: Lower Chanceford (73).

NEW JERSEY
Burlington Co: Woodland Twp (17, 18), Southampton Twp (19).

Bartonia No. 59: 107-111, 1996

Soil Chemical and Physical Properties Associated
with Two Pennsylvania Threatened Plant Species
(Juncus biflorus and Aster dumosus) at Crow’s Nest Farm,
Chester County

JASON T. LAROCCO, ANNE K. WEBSTER, AND R. KELMAN WIEDER’
Department of Biology, Villanova University, Villanova, PA 19085

The grass rush (Juncus biflorus Ell.) and the bushy aster (Aster dumosus L.) have been
proposed for listing under Title 25, Chapter 82 of the Pennsylvania Wild Plant Conservation
Act (Rhoads & Klein 1993). The Vascular Plant Technical Committee of the Pennsylvania
Biological Survey has recommended threatened status for J. biflorus, while the status of A.
dumosus remains undetermined (E. Dix, Pennsylvania Natural Diversity Inventory, pers.
comm.). Juncus biflorus, a facultative wetland species (sensu Reed 1988), commonly is
found in moist or wet areas within meadows and shores; Aster dumosus, a facultative upland
species (sensu Reed 1988), generally is found in dry or moist, and often sandy areas, near
or on the coastal plain (Gleason & Cronquist 1991). Both species occur in a small meadow
(about 0.5 ha) surrounded by successional deciduous forest at the Crow’s Nest Farm site,
Chester County, PA. Red maple (Acer rubrum) and tulip poplar (Liriodendron tulipifera)
may be encroaching upon the meadow. Invasion of the meadow by woody vegetation would
likely alter local environmental conditions, which in turn may lead to decline or local
elimination of J. biflorus and A. dumosus. As a first step toward developing a management
plan for maintaining these two threatened species, this study was undertaken to characterize
Surface soil chemical and physical properties in the meadow and in the surrounding forest.
Results have relevance to the broader question: are the environmental conditions at the
Crow’s Nest Farm site suitable for continued survival of J. biflorus and Aster dumosus?

STUDY SITE

The section of Crow’s Nest Farm that was studied consists of a meadow, 0.5 ha,
Surrounded by mixed deciduous forest, containing red maple (Acer rubrum) and tulip poplar
(Liriodendron tulipifera). The soil was classified as a Croton silt loam, characterized by
poor drainage on uplands, with 3 to 8% slopes (USDA 1963). However, in northern Chester
County, where our study site is located, soils tend to be shallower over bedrock, allowing
for better surface drainage than soils characteristic of the Croton series. Surface soils for this
Series generally consist of a dark-brown silt loam layer, with a subsoil layer of yellowish-
brown and reddish-brown silty clay loam. A claypan 38-56 cm below the surface tends to
Slow internal drainage. Croton silt loam soils promote the growth of black oak, red oak,
cottonwood, maple, ash, hickory, and elm comprising hardwood forests.

—— eee
‘Corresponding author

107

108 BARTONIA

2 0 10 20 30 40 50
80

Distance (m)

Distance (m)

Figs. 1 & 2. Fig. 1, Sampling grid showing meadow (unshaded area) surrounded by forest (shaded area). Fig. 2,
Contour plot of pH, one of the three variables that separated the forest from the meadow in the multivariate stepwise
discriminant analysis. The plot was generated using negative exponential smoothing, using the 53 grid point
measurements of soil pH; contour interval is 0.5 pH units (SYSTAT 1993).

METHODS

A sampling grid was established in the meadow and adjacent forested area (Fig. 1). At
each of the 53 grid nodes, we collected two soil samples (10 cm deep, 5 cm diameter) using
an AMS soil core sampler with hammer attachment. For one soil sample from each location,
soil wet weight was recorded before oven-drying the sample for one week at 60° C; bulk
density and water content were then calculated. The dried soil samples were sieved through
a 2 mm mesh screen prior to determining percentages of sand, silt and clay using the
hydrometer method (Day 1965). Sand was defined as particles ranging from 50 pm to 2mm
in diameter, silt as particles ranging from 2 to 50 um, and clay as particles smaller than 2
uum. For the second soil sample from each location, a subsample was analyzed immediately
for pH (1:1 soil/water slurry). Within 24 hr of soil collection, separate soil subsamples were
extracted with dilute acid-fluoride (Bray’s P; Olsen & Sommers 1982) and 10 mM CaCl,
(Adams 1974). The dilute acid-fluoride extracts were analyzed for PO}--P colorimetrically
(stannous chloride method). The CaCl, extracts were analyzed for SO}--S and NOj-N by
ion chromatography and for NH;-N using the colorimetric phenate method. Soil water
contents and bulk densities were used to convert chemical concentrations determined for
fresh soils to a per gram dry mass or cm”® basis.

JUNCUS BIFLORUS AND ASTER DUMOSUS 109

& q
3 $
8 Sg
2 §
: a
i ibe Beceem T T ras i
0 10 20 30 40 50
Distance (m) Distance (m)

Figs. 3 & 4. Fig. 3, Contour plot of available inorganic N (ug/cm’), one of the three variables that separated the
forest from the meadow in the multivariate s tepwise discriminant analysis. The plot was generated using negative
€xponential smoothing, using the 53 grid point measurements of available soil inorganic N; contour interval is
ug N/cm’ (SYSTAT 1993). Fig. 4, Contour plot of Bray’s P (ug/cm’), one of the three variables that — ti
forest from the meadow in the multivariate stepwise discriminant analysis. The plot was generated using n
eae smoothing, using the 53 grid point measurements of soil P; contour interval is 5 ug P/cm’ (SYSTAT
).

RESULTS

The meadow soils and forest soils did not differ with regard to any of the physical
variables and differed with regard to only one of the chemical variables, soil pH, which was
higher in the meadow than in the forest (Table 1). A multivariate stepwise discriminant
analysis (SAS 1990), using all soil chemical and soil physical variables, distinguished the
forest and meadow areas of the study site on the basis of soil pH (R* = Ag p = 0.0104),
total CaCl,-extractable inorganic N (NO3-N + NHN) concentrations (ug cm’; R? = 0.0981;
P = 0.0238), and Bray’s P concentrations (ug cm™; R? = 0.0430; p = 0.1442). Contour plots
(Figs. 2-4) reveal spatial variability in these three soil variables within the study site. These
Plots further substantiate a general pattern of higher soil pH, lower available inorganic N,
and lower Bray’s P in the meadow than in the forest (cf. Fig. 1).

DISCUSSION

Juncus biflorus and Aster dumosus, although broadly distributed throughout the eastern
United States, have been proposed for listing under Title 25, Chapter 82 of the Pennsylvania

110 BARTONIA

Table 1. Summary information for forest and meadow at Crow’s Nest Farm (values represent mean + std. error;
N= 34 for forest, N = 19 for meadow).

Variable Forest Meadow Variable Forest Meadow
Soil pH 4940.1" °°. 5.F2 0:1 Bray’s P (ug/g dry soil) 16.3+1.7 12.9+0.9
H,O (% w/w) 21544) 32952038 Bray’s P (ug/cm’) WAH. $33 250
H,O (g/cm’*) 0.39 + 0.01 0.43 + 0.01 SO?--S (ug/g dry soil) 8.9413 £1.72 2.1
NH;}-N (g/g dry soil) TOL09- “964212 SO;-S (ug/em’*) 9.5+1.4 12.9 + 2.6
NH}-N (yg/cm’) 716+ O08. 100-5 13 Bulk density (g/cem*) 1.05 +0.03 1.05 + 0.04
NO;-N (ug/g 2 soil): 0.94 0.2 ° ~12 403 Sand (%) §6424176° 353412
NOj-N (ug/em 09+02. 1304 Silt (%) 385213 398 + 10
Clay (%) 5.2 +06 49+0.8

*Statistically significant at the 5% level; student’s t-test.

Wild Plant Conservation Act. At the Crow’s Nest Farm study site, the J. biflorus population
consists of at least 50 individuals, mostly in the northern half of the meadow. Though not
positively identified in 1994, 4. dumosus was confirmed by the Pennsylvania Natural
Diversity Inventory a few years ago to be located at the southern end of the meadow
(Stevens Heckscher, pers. comm.). Neither species occurs in the surrounding forest. As a
first step toward developing a management plan for J. biflorus and A. dumosus at Crow’s
Nest Farm, this study compared soil chemical and soil physical properties in the meadow and
the forest.

Reconnaissance of the field site suggested that the meadow area may represent an
abandoned farm road connecting the public road just to the south of the meadow with a field
just to the north. Therefore, we expected to see a greater soil bulk density in the meadow
than in the forest resulting from compaction of the surface soils along the farm road. If the
meadow is indeed an abandoned farm meadow, this past disturbance is not reflected in any
of the soil physical properties at the site today.

Both meadow and forest soils contained about 56% sand, 39% silt, and 5% clay,
classifying the soils as sandy loams. Although we found no information on edaphic
tolerances for A. dumosus, the preference for sandy soils appears to be indicative of other
related species of aster. Abrahamson & Solbrig (1970) reported that A. cordifolius, A
undulatus, and A. lowreianus preferred soils that had a high sand percentage with a favored
range in soil pH of 5.5 to 6.8.

Meadow soils differed from the forest soils in terms of chemical properties. The lower
soil pH in the forest than in the meadow is likely the result of the decomposition of the
forest leaf litter. The lower available inorganic N and Bray’s P in the meadow than in the
forest may indicate less favorable conditions for tree growth in the meadow. However, both
red maple and tulip poplar, abundant in the surrounding forest, are notoriously good
colonizers of open sites (Harlow et al. 1979). Their encroachment upon the meadow should
be monitored. Tree invasion of the meadow is a possible threat to the persistence of J.
biflorus and A. dumosus. However, a potentially greater threat to J. biflorus and A. dumosus
is the presence of Japanese stilt-grass (Microstegium vimineum (Trin.) A. Camus). This non-
native aggressive, invasive species has been expanding its range in Pennsylvania in recent
years (Rhoads & Klein 1993). Microstegium is widely dispersed throughout the meadow and
has the potential, if left unmonitored, to outcompete and exclude J. biflorus and A. dumosus.

e persistence of J. biflorus and A. dumosus at this site will depend on the complex
interactions of a wide variety of abiotic and biotic factors that influence plant success. The

JUNCUS BIFLORUS AND ASTER DUMOSUS 111

descriptive nature of this study precludes definitive conclusions about the role of soil
chemical properties in affecting these two plant species. However, it appears that Aster
dumosus might be living at the lower end of its sand and pH tolerance. Therefore,
encroachment by the surrounding forest should be closely monitored to assess its possible
impact on soil physical properties and pH. In addition, the population of Japanese stilt grass
should be monitored because of its potential to exclude both J. biflorus and A. dumosus.
Therefore, competition with Japanese stilt-grass and tree invasion of the meadow appear to
represent potentially serious short-term and — threats, respectively, to the persistence
of J. biflorus and A. dumosus at Crow’s Nest Farm

ACKNOWLEDGMENTS

We thank Dr. Stevens Heckscher, Taylor Memorial Arboretum, Natural Lands Trust, for
introducing us to the Crow’s Nest Farm site and for providing useful information throughout
the study. Mr. Scott Starr contributed valuable assistance in the field and laboratory.

LITERATURE CITED

ABRAHAMSON, W. A. & O. ‘3 SOLBRIG. 1970. Soil preference and variation in flavanoid pigments in species of
ster. i a i TZ2al

ADAMS, F. 1974. Soil anid Pages 441-448 in E. W. Carson, ed. The plant root and its environment.
“yo-anga = ee ress, Charlottsville
Day, P. R . Particle fractionation and particle-size analysis. Pages 545-565 in C. A. Black, ed. Methods

of soil — Part 1, physical and Valea ta properties, including statistics of measurement and sampling.
American Society of Agronomy, Madiso

GLEASON, H. A. & A. CRONQUIST. 1991. Mana of vascular plants of northeastern United States and adjacent
Canada. The New York Botanical Garden, Bro

HARLOW, W. M., E. S. HARRAR, & F. M. WHITE. 1979, Textbook of dendrology, 6th ed. McGraw-Hill Book Co.,
New York.

OLSEN, S. R. & L. E. SOMMERS. 1982. Phosphorus. Pages 403-430 in A. L. Page, R. H. Miller, & D. R. Kenney,
eds. Methods of 285 analysis, Part 2, Chemical and microbiological properties, 2nd ed. American Society of
Agronomy, Madiso

REED, P. B., JR. 1988 National list of plant species that occur in wetlands: 1988 national summary. U.S.
Department of the Interior, — ses Wildlife Service, St. Petersbur

RHoaDs, A. F. & W. M. KLE 1993. The vascular flora of Pennsylvania—annotated checklist and atlas.
American Philosophical roto Philadelphia

SAS. 1990. SAS/STAT user’s guide, version 6, 4th ed. ane Institute, Inc., Cary.

SYSTAT. 1993. SAS/STAT 5.02 for Windies SYSTAT, Inc., Evanston.

USDA. 1963. Soil — of Chester County, Pennsylvania. v, S. Department of Agriculture, Soil Conservation
Service, Washingto

ca bre

be ‘
pe Bid Be a | i
af % i Hh z
hd he > 4

wie!

my ee
‘ethay 4 a

Poors et

shane

sit rate!
tre? i

hbase $e S27

Bartonia No. 59: 113-117, 1996

Polypodium appalachianum, P. virginianum, and Their
Hybrid in New Jersey and Pennsylvania

JAMES D. MONTGOMERY
Ecology III, R.R. 1, Berwick, PA 18603

In the early 1950s, Irene Manton and her co-workers discovered that the common eastern
Polypodium virginianum L., or rock-cap fern, included both diploid (n=37) and tetraploid
(n=74) races (Manton 1950, Manton & Shivas 1953). Evans (1970) reported that triploid
hybrids also existed which had aborted spores and morphology intermediate between the
diploid and tetraploid. The morphological differences between the diploid and tetraploid
were summarized by Kott & Britton (1982). Haufler and co-workers (Haufler & Windham
1991, Haufler et al. 1995) determined that the tetraploid originated as a hybrid between the
eastern diploid and a far northern species P. sibericum Siplivinskij, established that P.
virginianum L. was the correct name for the tetraploid, and named the eastern diploid P.
appalachianum Haufler and Windham.

New Jersey Ferns and Fern Allies (Montgomery & Fairbrothers 1992) and The Vascular
Flora of Pennsylvania, Annotated Checklist and Atlas (Rhoads & Klein 1993) were
published while work on this group was in progress. In both works the authors indicated
that two species and a hybrid were included, but did not attempt to describe or map the
geographic distribution of the species and hybrid.

The purpose of this paper is to present maps of the geographic distribution and discuss
information on the habitats of Polypodium appalachianum, P. virginianum, and their hybrid
(which does not have a binomial) in New Jersey and Pennsylvania.

METHODS

Herbarium specimens were examined for New Jersey and Pennsylvania from the following
herbaria: BUPL, CHRB, CM, NY, PH (including PENN), US, YUO, and the author’s
Personal herbarium. The specimens were identified using the characteristics listed by Kott
& Britton (1982), Haufler & Windham (1991), and Haufler et al. 1993). As indicated by
these authors, the most reliable characteristic to distinguish the diploid P. appalachianum
from the tetraploid P. virginianum is the number of sporangiasters (modified sporangia) in
the sorus (>40/sorus in P. appalachianum; < 40/sorus in P. virginianum). This characteristic
is useful in the field and in most herbarium specimens collected in the late summer (when
the spores mature) through early winter. Material collected in the spring and early summer
usually has the sorus too disintegrated or too immature, and the relative number of
Sporangiasters cannot be determined. Blade shape (deltate and broadest near the base in P.
ppalachianum, narrowly elliptic and broadest near the middle in P. virginianum), and shape
of the tips of the blade segments (acute or pointed in P. appalachianum, rounded in P.
virginianum) are useful characteristics in herbarium specimens and in the field. Spore size
Was used in the laboratory to confirm identification of herbarium material. The hybrid can
only be distinguished with certainty by its aborted spores. Many hybrid specimens can be

113

114 BARTONIA

identified tentatively by their intermediate morphology: the blade is rather straight-sided

elow the tapering apex with many segments nearly equal in length, and some of the
segment tips are pointed. As Kott & Britton (1982) pointed out, removal of these hybrid
specimens makes distinguishing the species much less difficult.

Maps of the distribution were prepared using base maps from Montgomery & Fairbrothers
(1992) for New Jersey, and Rhoads & Klein (1993) for Pennsylvania. It was anticipated that
some difference might be found in the habitat preference for the two species; therefore,
habitat data were tabulated from those specimens that contained this type of information.

RESULTS AND DISCUSSION

Somewhat less than 60% of the specimens from the two states represent P. appalachi-
anum. In New Jersey, 54% are P. appalachianum, 35% P. virginianum, and 10% P.
appalachianum x virginianum. In Pennsylvania, 59% are P. appalachianum, 31% P.
virginianum, and 10% the hybrid. Assuming the species and hybrid have been collected in
proportion to their occurrance, which seems reasonable given their morphological similarity,
these numbers represent their relative abundance in New Jersey and Pennsylvania.

P. appalachianum has been recorded from 18 of New Jersey’s 21 counties (Fig. 1). It is
common in the northern part of the state and on the Inner Coastal Plain, but nearly absent
from the Outer Coastal Plain. There is a single record from the Pinelands, from an old well.

P. virginianum has been recorded from 14 counties (Fig. 2). It is unrecorded from the
urban counties of the northeast and much less common than P. appalachianum on the Inner
Coastal Plain. There are four records from the Pinelands. Most of the records are from the
Ridge and Valley and Highlands in the northwestern part of the state.

Figs. 1-3. Distribution of Polypodium in New Jersey. Fig. 1, P. appalachianum. Fig, 2, P. virginianum. Fig. 3,
P. appalachianum x virginianum. Base map from Montgomery & Fairborthers (1992).

POLYPODIUM IN NEW JERSEY AND PENNSYLVANIA 115

Erie * 7 e r
. & e Susquet® :
ott Tioga Bradford usquehanna
® e
2-2 - ei <r St
e
® ee su liyan gp 7 geo
. COMI e.6hU6°
: ra ei oe
e?
* } ) North inp
2 rent £ fe
fe
¥ Bracks
ere
°
n

ee . Re
Hs . ® ry PPG!
e Bedigg /o Frankl e Pa ngsier eee
e/e e
Gieéde Faygiee Someffet ” Fyton dee * Adgms Yor s &,
: od Po a ° » © *%e

ces es he
Warren McKean © Potter Tioga Bragjord Susque@ana
°

e bd . »
- % e ‘i °, wg ° J
os - Bedior Fran —. ‘4
wets var bd ais Figen bed eal % e . . 7 %s
es e e eo

Figs. 4 & 5. Distribution of Polypodium in Pennsylvania. Fig. 4, P. appalachianum. Fig. 5, P. virginianum. Base
map from Rhoads & Klein (1993).

P. appalachianum x virginianum has been recorded from 9 of New Jersey’s 21 counties
(Fig. 3). There is only one collection from southern New Jersey and none from the
Pinelands. Records are scattered in northern New Jersey. The lack of records from Sussex
and Morris counties is surprising, since both parents are common there.

Polypodium appalachianum has been collected in all of Pennsylvania’s 67 counties (Fig.

4), and is well represented from all geographic provinces. There are multiple records for

116 BARTONIA

Ce
ae
Erie
Warren McKean Potter Tioga Bradi@rd Susquehanna
Crawtord
2
e126 ee
Venapgo
Mercer
a
Clearfield CofMre
. C
e
- Bedford Franklin -
Fayette ets Fulton Adams York Sf
Greene e os
é 2 | e s

Fig. 6. Distribution of Polypodium appalachianum x virginianum in Pennsylvania.

most counties (except Crawford). The higher number of collections from eastern and
western Pennsylvania probably reflects more active collectors in these parts of the state,
rather than species distribution.

P. virginianum has been collected from 62 of the 67 counties in Pennsylvania (Fig. 5).
This species also occurs in all parts of the state, but is apparently uncommon in the north
central counties on the Allegheny Plateau. Some of this may be due to collecting bias, but
P. appalachianum has been collected more frequently in this area. More collecting is needed
in Warren, McKean, Potter, Tioga, and Bradford counties.

P. appalachianum x virginianum is recorded from 45 of the counties in Pennsylvania (Fig.
6). It has been collected from most parts of the state, but appears to be rare or absent from
the Poconos. The hybrid is much less frequently collected than the parents, as indicated by
the single records from many counties. The concentrations in Columbia, Montour, and
Northumberland counties represent searching by the author. The concentrations in
northwestern Pennsylvania represent intensive collecting by C. F. Chuey (YUO) and
personnel at the Carnegie Museum of Pittsburgh (CM). The hybrid is probably relatively
more common in Pennsylvania than indicated.

Most of the specimens of P. appalachianum and P. virginianum have been collected on
rocks, boulders, ledges, cliffs, or rocky woods. In New Jersey, of the specimens with habitat
information on the labels, 45% of the P. appalachianum and 37% of the P. virginianum were
collected in rocky woods or wooded slopes (or some similar combination). Boulders oF
rocks account for 17% of the P. appalachianum and 15% of the P. virginianum specimens,
and cliffs and ledges 19% of the P. appalachianum and 11% of the P. virginianum. A few
specimens were collected on tree trunks or bases of trees. In southern New Jersey most
specimens of both species were collected from sandy banks of streams or roads. A few were
collected from old mortar or outcrops of ironstone.

POLYPODIUM IN NEW JERSEY AND PENNSYLVANIA By

In Pennsylvania 19% of both P. appalachianum and P. virginianum were collected on
rocky woods or wooded slopes, 31% of the P. appalachianum and 30% of the P.
virginianum were collected on boulders or rocks, and 22% of the P. appalachianum and 23%
of the P. virginianum were collected on cliffs or ledges. Smaller percentages were collected
on banks, tree trunks or tree bases in Pennsylvania.

The similarity of these figures indicates that there is no difference in habitat between the
two species. The differences between the two states are probably a reflection of different
habitat availability. New Jersey has a more extensive Coastal Plain and less mountain
topography than Pennsylvania.

Of the P. appalachianum specimens where a specific type of rock is indicated, 18% were
collected on limestone, 59% on sandstone, 16% on shale, and 7% on other rock types
(granite, schist, gneiss, conglomerate, or trap rock). Of the specimens of P. virginianum,
14% were collected on limestone, 39% on sandstone, 37% on shale, and 10% on other rock
types. Neither species shows any affinity for calcareous or acidic rock from these data.

There is less information on which to base conclusions about the hybrid. The same
general habitat types as the parents are listed on herbarium labels from both states. Most
specimens of the hybrid were collected from sandstone or shale; relatively few were noted
as collected on limestone. There is very little data on whether this hybrid usually grows
with its parents, but the field experience of the author indicates that at least one of the
parents is usually present.

Polypodium appalachianum and P. virginianum are both common and widespread species
in New Jersey and Pennsylvania. Their habitats and geographical ranges overlap throughout
both states. The hybrid is also fairly common. Field botanists in both states should become
familiar with these plants.

LITERATURE CITED

EVANS, A. M. 1970. A review of systematic studies of the pteridophytes of the southern Appalachians. In The
distributional history of the biota of the southern — Part II. Flora. Research Division Monograph
2, Virginia Polytech. Inst. and State Univ., Blacksbur

HAUFLER, C. H. & M, D. WINDHAM, 1991. New species of North American Cystopteris and Polypodium, with
comments on their gio erga Amer. Fern J. 81:7-23.

5. Reticulate evolution in the Polypodium vulgare complex. Syst. Bot. 20:89-109.

5-323 in Flora of North America north

FA: LANG, a A. WHITMORE. 1993. Polypodium. Pages 31
of Mexico. — of N. A. Editorial Comm. Oxford Univ. Press, New York.
tie L. S. & D. M. BRITTON. 1982. A comparative study of sporophyte morphology of the three cytotypes of
Polypodium virginianum in Ontario. Canad. J. Bot. 60:1360-1370.
MANTON, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge Univ. Press, Cambridge.
“ a SHIVAS. 1953. Two cytological forms of Polypodium mune in eastern North America. Nature

172:4

cba J. D. & D. E. FAIRBROTHERS.
Brunswick.

RuHoaps, A. F. & W. M. KLEIN. 1993. The vascular flora of Pennsylvania, annotated checklist and atlas. Amer.

E,
Philosoph. Soc., Philadelphia.

1992. New Jersey ferns and fern allies. Rutgers Univ. Press, New

¢ 4
‘ *
° . 23 “ye Z
. , ; ; OY dee gs 4 , .
A * epee : ee

a if pascie aR

yp

ae

: hoe ae,

i jp SNPa eS
Pee

née
“yn
3

5 ae ‘4 : Sone cj Pe ered

ER Ghar

xg Pia -
eer is =

p>

‘>

Bartonia No. 59: 119-121, 1996
Observation of Polyembryony in Trillium and Smilacina

JOHN F. GYER
P.O. Box 185, 243 Jessup Mill Rd., Clarksboro, NJ 08020

TRILLIUM OBSERVATIONS. Three observations of a pair of germinated embryos
emerging from a single seed were made in 1993 during the course of experiments designed
to study seed dormany in Trillium. Two double-embryo seeds were found in 80 seeds
collected from two 7. reliquum Freeman berries. One appeared in a sample of over 1,000
seeds bulked together from many 7. grandiflorum (Michx.) Salisb. berries. After root growth
reached about 2 cm and frost danger was past, the double-embryo seeds were planted into
garden soil, but did not produce cotyledons the next spring. Since these seeds were not
dissected, it is not certain that the two embryos were associated with twin endosperms as
would be expected if they arose from separate embryo sacs that in turn resulted from the
development of both dyad cells. This, however, is a more plausible origin of the observed
double embryos than is apomixis from the structures described by Swamy (1948) or Jeffery
& Haertel (1947).

The seeds for this work came from plants in the garden at Winterthur Museum, Garden,
and Library at Winterthur, Delaware. The Winterthur collection was developed by H. F. du
Pont during the early 1900’s with significant additions at mid-century. Since some of the
species in the collection are now considered rare or endangered, Winterthur’s Garden

partment wanted to explore the option of propagation from seed of their own specimens.
The immediate objective of the author’s volunteer research was to find ways to reduce seed
germination time from the expected two or more years. The first step was a series of careful
observations of the germination of hand-pollinated seeds from the collection. Hand
pollination resulted in large numbers of 7. grandiflorum seeds from the Azalea Woods
section of the garden and lesser quantities from the individual specimens of other species.

The germination procedure generally followed the technique proposed by Deno (1993).
Freshly harvested seeds were placed in moist paper towels in polyethylene bags at room
temperature (about 75°F) until the arils had disintegrated (about a week). The seeds were
removed from the towel, rubbed gently in a sieve under running tap water to remove
degraded tissue, and returned to a fresh moist towel and bag. In the experiments where
double embryo seeds were found, the seeds were held at room temperature for 3 months,
stored 3 months at 40°F, and returned to room temperature. The bags were inspected
Periodically for germination, the removal of rotted seed, and the proper moisture level. The
double embryo seeds germinated during the second storage at room temperature. One of the
T. reliquum double-embryo seeds is shown in Fig. 1. ? ;

The T. reliqguum plants that produced the double-embryo seeds were particularly vigorous.
One had two blooming stems, the other had one. Their rhizomes grew at the soil line
beneath two to three inches of oak leaf litter that protected the leaves from damage due to
rain-splashed soil. The environment had been stable for at least three years because a clump
of seedlings, some with monofoliate and some with trifoliate leaves (two and three year
plants) grew where a berry had fallen and shed seeds that were not carried off by ants.

119

120 BARTONIA

Fig. 1. Two germinating embryos emerging from a single seed of Trillium reliquum Freeman. 1 = root, 2 =
rhizome, 3 = cotyledon petiole, 4 = collar of starchy endosperm.

The two adult plants were cross-pollinated twice in May and the berries were harvested in
August. The strong growth of the plants and the high density of pollen on the stigmas
undoubtedly contributed to the development of double-embryo seeds.

The double-embryo 7. grandiflorum could not be traced to an individual plant, but the
Azalea Woods population at Winterthur contains numerous very strong plants and is actively
reproducing. No photograph was taken of this double-embryo seed, but recently Solt (1995)
photographed a double-embryo T. grandiflorum in the course of her research.

The origin of double-embryo seeds in Trillium has not been previously reported, but
Jeffery & Haertl (1939a & 1939b) claim that apomixis exists in Trillium while Swamy
(1948) reports the development of multiple embryo-like structures in the developing
endosperm of 7. undulatum Willd. Although either process could produce multiple embryo
seeds, neither author reports observing multiple mature embryos or the germination of
multiple embryo seeds. The Jeffery & Haertl report of apomixis is not supported by Howe
(1940), the detailed work of Blain (1945), or that of Berg (1958 & 1962). In fact Berg
(1962) states, "Apparently a statement by Jeffery and Haertl should be assumed wrong rather
than correct, unless sustained by observations made by other investigators." Swamy reports
multiple embryo-like structures in 19% of the fertilized 7. undulatum Willd. ovules he
examined. If Swamy’s embryo-like structures occur in other Trillium species and commonly
result in viable embryos, the large number of seeds germinated in the Winterthur experiments
should show numerous instances of multiple embryo germinations. They do not. Only
double-embryo germinations were observed and these were very rare.

The simplest explanation of the origin of twin embryos is to hypothesize that both dyad

POLYEMBRYONY IN TRILLIUM AND SMILACINA 121

cells in the developing megaspore continue to grow and are fertilized. This would result in
two embryos each with its own endosperm encased in a single integument. Since one dyad
cell disintegrates under normal conditions, the development of both may be favored by
particularly strong plant growth while the fertilization of both would be favored by a heavy
load of pollen on the stigma. Both of these coop let existed for the 7. reliquum plants
when ad produced the observed double-embryo S.

S INA RACEMOSA OBSERVATION. va pair of germinated embryos emerged
from a she seed of Smilacina racemosa (L.) Desf. during experiments based on Deno’s
(1993) germination protocol as summarized above. The seed population was from a single
panicle collected in September. The berry pulp was removed by gentle abrasion in a sieve
under running tap water. The twin embryos produced roots about five cm long before they
were planted with normal, single-embryo seeds near an existing population in a deciduous
woods. No photographs were taken of the twin embryo see

Gorham (1953) described normal early megaspore developencnt in S. racemosa, stating
that the upper dyad cell develops into an embryo sac while the lower disintegrates. If both
dyads survived, produced embryo sacs, and were fertilized, then twin embryos could form
as is hypothesized for Trillium.

Gorham’s (1953) claim of apomictic seed production is accompanied by these observations
that suggest the possibility of unobserved fertilization: (1) viable pollen, (2) pollen tubes
growing in stylar tissue, (3) pollination required for seed development, (4) polar nuclei that
"—fuse suddenly, some time after pollination," (5) degeneration of ovules that do not produce
endosperm.

A fully satisfactory explanation of the development of the twin embryos in the double-
embryo seed germinated in my experiments awaits the application of currently available
techniques in a reinvestigation of reproduction in S. racemosa.

LITERATURE CITED
BERG, R. Y. 1958. Seed a morphology, and phylogeny of Trillium. Mat.-Naturv. Klasse 1
BERG, R. Y. 1962. Contribution to the comparative embryology of the Liliaceae: Soliopus, Trillium, Paris, and

Cs :
BLAIN, A. 1945. The Sexual Sari ae of Trillium. T. erectum L., T. grandiflorum (Michx) Salisb.) Ph.D.
thesis, McGill University, unpubl
DENO, N.C. 1993. Seed a ieee nl ory, and practice, 2nd ed. Published by the author, State College.
GORHAM, A. L. 1953. The question of fertilization in Smilacina racemosa (L.) Desf. Phytomorphology 3(1,2):44-

50.
Howe, T. D. 1940. Development of the embryo sac in Trillium grandi iflorum. Amer. J. Bot. 27:11s
JEFFREY, E. C. & E. J. HAERTL. 1939a. Apomixis in Trillium. LaCellule 48:78-88.
JEFFREY, E. C. & E. J. HAERTL. 1939b. The production of unfertilized seeds in Trillium. Science 90:81-82.
SOLT, S. & J. F. GyeR. 1995. Trillium ee hres and growth, unpublished seminar, October 25, 1995, New
England Wild Flower Society, Garden in the Woods, Framingham.
Swamy, B. G. L. 1948. On the peepee oth development of Trillium undulatum. LaCellule 52:5-1.

HSTOver

ae

ogttis 20 ¢ Kia
one, aa &

ia
ee a

od” --
fest blues

“ap Spc tarts

Soa

Bartonia No. 59: 123-124, 1996

The Gold-Cone Tamarack (Larix laricina forma lutea,
Pinaceae) in Pennsylvania

ROBERT F. C. NACZI AND JOHN W. THIERET
Department of Biological Sciences, Northern Kentucky University
Highland Heights, KY 41099

During field work in Bradford County, Pennsylvania, in June 1994 we explored a
tamarack (Larix laricina) swamp near Troy. There, among the many purple-coned tamracks
we noticed a single tree that stood out quite obviously from the rest: its cones were clear
yellow without a trace of purple. Cone color was the only difference we noted between the
yellow-coned tree and the other tamaracks. We had never seen a yellow-coned individual
even though we have, between us, observed tamaracks in nine Canadian provinces from
Newfoundland to Alberta, in Canada’s Northwest Territories, and in the U.S. from Maine
to Minnesota.

Later investigation in the literature revealed that a yellow-coned "variety" of L. /aricina
was described in a paper on North American Larix by Jaurés and de Ferré (1949) as var.
lutea Jaurés: "A laricina typica differt strobili colre luteo." The type specimen (at Harvard
University Herbaria [A!]), said to be from a cultivated tree, was collected by J. B. Moyle
(no. 172) on 5 Jul 1930 ina "Forest Nursery" in Clearwater County, Minnesota. Our inquiry
among several foresters in that county revealed that such a nursery no longer exists; none
of the foresters to whom we spoke recalled seeing a yellow-coned variant of tamarack.

Further investigation revealed that most of the descriptions of L. Jaricina we checked
make no mention of such a variant. Even Flora of North America describes the seed cones
as being "at first dark red to violet, later turning yellow-brown" (Parker 1993). The cones
On our tree were certainly not "at first dark red to violet."

Den Ouden & Bloom (1965), treating the variant as a form (a rank we consider
appropriate), published the new combination L. Jaricina “forma lutea (Jaurés & de Ferré) den
Ouden & Boom." Of the plant they wrote "Flowers yellow; found in Clearwater County,
U.S.A."

Farjon (1990) included "var. /utea Jaurés et Y. de Ferré" in synonymy under L. Jaricina.
However, in his description of the cones of that species, "yellow" does not appear. He wrote
of their color as ibe vee red or greenish" when immature and "orange-brown with
purplish tinge" at matu

Welch & Haddow p's listed "forma /utea Den Ouden/Boom." They commented that
the variant was "named for a tree bearing yellow flowers found in Clearwater County, Pa.
[sic]”

Only three infraspecific epithets—none of them /utea—appear under L. laricina in Cope
(1986). And, finally, in all of Larix the only infraspecific taxon based on cone color appears
to be "var. lutea" of L. laricina (Kriissmann 1970).

Jaurés & de Ferré stated that intermediates between their var. /utea and the “forme

typique” are frequent. These authors worked only with herbarium specimens; we suspect

123

124 BARTONIA

that sharp distinctions between the color variants become blurred as the specimens fade with
age. Such fading may be evident on the holotype: the scales of its cones, after some 66
years in a herbarium, are now light brown (tan) with no trace of purple. The small bracts
subtending the cone scales are somewhat darker brown.

Authorship of the epithet /utea is given by several authors (Farjon 1990; den Ouden &
Boom 1965; Kriissmann 1970) as "Jaurés et de Ferré." In the original publication of the
epithet no authorship is indicated at the place (page 6) where var. /utea (one of two varieties
proposed under L. /aricina) is described. Elsewhere in the paper, though, the statement is
made (page 2) that "Ces deux variétés s’appelleront: L. /utea et L. parvistrobus Jaurés.'
Clearly de Ferré has been excluded.

Tamarack appears to be less common in cultivation than some of its Old World congeners.
The yellow-coned variant, being u pommel a well be desirable as a specimen plant. We
sugest the name "gold-cone tamarack" for

Further observations on the occurrence 2 this variant would be welcome.

Our voucher specimen is deposited in the herbarium of Northern Kentucky University
(KNK): Pennsylvania, Bradford County, in Larix swamp, along S side of Fallbrook Road,
0.3 mi W of jct with Swamp Road, ca. 4 mi SW of Troy, 15 Jun 1994, Naczi & Thieret
4225. We collected a sheet of the typical tamarack, with its purple cones, at the same time
(Naczi & Thieret 4221, KNK).

We thank Dr. David E. Boufford and Dr. David M. Brandenburg for aid.

LITERATURE CITED

Cope, E. A. 1986. Native and cultivated conifers of northeastern North America. Comstock Publishing Associates,
a Division of Cornell University Press, Ithac

DEN QOUDEN, P. & B. K. BOOM. 1965. Manual of same conifers. Martinus Nijhoff, The Hague.

FARIJON, A. 1990. Pinaceae. Koeltz ileaitie Books, Koenigstein

JAURES, R. & Y. DE FERRE. 1949. A propos des Larix d’ pened du Nord. Bull. Soc. Hist. Nat. Toulouse 84:1-
16.

KRUSSMANN, G. 1970. Handbuch der Nadelgehdlze. 4. Leiferung, Bogen 10-12:169-176. Paul Parey, Berlin.

PARKER, W.H. 1993. Larix. Pages 366-368 in Flora of North America north of Mexico, volume 2. Oxford Univ.
Press, New York.

WELCH, H. & G. HADDOW. 1993. World checklist of conifers. Landsman’s Bookshop, Buchenhill.

Bartonia No. 59: 125-128, 1996

Paintings of Fungi by Lewis David von Schweinitz
in the Archives of the
Academy of Natural Sciences of Philadelphia

DANA M. LYNCH
ademy of Natural Sciences of Philadelphia
1900 sp eialin Franklin Parkway, Philadelphia PA 19103-1195

The Academy of Natural Sciences of Philadelphia has a rich and remarkable collection

of historical botanical items including the herbaria of many famous American botanists,

ooks, manuscripts, and works of art. In its collection, the Academy is fortunate to have
Volumes I, II, III, and V of Fungorum Nieskiesium Icones [Icones] (1804) by Lewis David
von Schweinitz (1780-1834). These four volumes include over 1,000 original paintings of
fungi species. They are expertly and artistically rendered in watercolor by Schweinitz, and
after nearly 200 years still retain their vivid color and beauty. A review of the species
illustrated in Icones indicates that 71 are those that were originally described by Schweinitz
and his mentor Johannes Baptista von Albertini in their published work Conspectus
Fungorum in Lusatiae superioris agro Niskiensi crescentium e methodo Persooniana
[Conspectus] (1805). Although the number and quality of the original paintings in the four
volumes is exceptional, they are archival materials that have had very limited recognition or
appreciation. In order to generate further interest and draw the attention of the mycological
community to these historical paintings, a listing of the 71 species that were newly described
in Conspectus and occur in Icones is provided in Table 1.

Lewis David von Schweinitz was born in Bethlehem, Northampton County, Pennsylvania,
on February 13, 1780. His great-grandfather Nicholas Lewis Count Zinzendorf, was the
founder of the Unitas Fratrum or Moravian Brethren church, and his father was in charge of
the fiscal and secular concerns of the Moravian Brethren in America (Pennell 1934). From
the age of seven to seventeen, Schweinitz was sent to study at the all boys school, Nazareth
Hall, in the Moravian community near Bethlehem. At Nazareth Hall, Schweinitz came under
the tutelage of Samuel Kramsch, a minister for the Moravian church and an ardent botanist
(Barnhart 1934). Under Kramsch and in the company of Christian Frederick Denke, a fellow
Student and botanist, Schweinitz’s interest in botany began.

In 1798, the von Schweinitz family moved to Germany, and Schweinitz entered the
Moravian Theological Seminary at Niesky in Silesia. There he met Professor Albertini, who
was also intensely interested in botany. Together Schweinitz and Albertini collected and
Studied fungi throughout the Niesky region and in 1805 published their Conspectus, which
contained descriptions of 1,130 fungi species including 127 newly described species. It was
during this same time period, probably in conjunction with the development of Conspectis,
that Schweinitz painted Icones. The Conspectus established Schweinitz’s and Albertini s
reputation among European mycologists leading to an honorary degree for Schweinitz from
Kiel. This work and two subsequent works by Schweinitz alone on fungi, Synopsis
Fungorum Caroliae superioris secundum obervationes in 1822 and Synopsis Fungorum in

125

126 BARTONIA

America Boreali media degentium in 1932, and his large herbarium are responsible for
designating Schweinitz as the founder of mycology in America (Benedict 1934). Although
interested in all the fields of botany throughout his life, von Schweinitz’s chief interest was
fungi (Benedict 1934). For more complete information on Schweinitz, see Bartonia No. 16,
1934

In 1834, when Schweinitz died, his large herbarium was bequeathed to the Academy of
Natural Sciences of Philadelphia. si extensive was his herbarium (23,000 species) that the
Academy’s previous herbarium was simply added to the Schweinitz series (Pennell 1934).
In 1903, George E. de Schweinitz presented Jcones volumes 2, 3, and 5 to the Academy, and
in 1913, Mrs. W. A. Lemly (Emily de Schweinitz Lemly) donated volume 1 to the Academy.
A letter from the librarian at the University of North Carolina at Chapel Hill states that
volume 4 is at this institution (Phillips and Phillips 1963). Years ago Eugene A. Rau stated
that he had seen seven volumes of /cones (Phillips and Phillips 1963). Assuming this to
be the case, the whereabouts of volumes 6 and 7 is unknown. We are proceeding on with
a project to list all the fungi in Schweinitz’s /cones with their associated names and
descriptions from the Conspectus.

Table 1. Species newly described in Conspectus and also illustrated in Jcones Volumes |, II, Il], and V

Species in Conspectus Volume Plate Figure
Sphaeria lutea I 9 2
Sphaeria rimosa I 1] 2
Sclerotium vaporarium I 37 y
Sclerotium scutellatum I 38 2
Lycogala I 49 3
Licea strobilina 1 60 s
Aecidium Violae I 65 2
Amanita porphyria I 82 8
Agaricus bulbiger (under Lepiota in /cones) Vv 354 245
Agaricus porphyropus (under Cortinaria in cones) Il 112 75
Agaricus lepidomyces (under Cortinaria in Icones) Vv 356 251
aricus luteo-virens (under Gymnopus in Icones) Il 117 91
Agaricus bosus (under Gymnopus in Icones) il 114 83
Agaricus camarophyillus (under Gymnopus > Icones) Vv 359 259
Agaricus vitellum (under Cortinaria in Jcone Il 102 50
Agaricus purpurascens (under Gymnopus in ae Vv 361 263
Agaricus tricholoma (under Gymnopus in Icones) Vv 360 261
Agaricus scabellus (under Gymnopus in Icones) Il 161 208
Agaricus tricolor (under Omphalia in Jcones) Vv 374 303
Daedalea rubescens Vv 381 3
Boletus ae (sistotrematoides in /cones) Vv 384 D2
Boletus con Ill 181 11
Boletus r Vv 389 59
Boletus sanguinolentus Vv 391 64
Sistotrema pendulum Vv 393 10
Sistotrema viride (aeriginosum in Icones) Ill 199 5
Hydnum vi ns Vv 395 19
Hydnum fasciculare Vv 399 26
m castaneum Vv 398 25
Hydnum bicolor Vv 398 23
Hydnum calvum Ill 206 13
calvum Vv 399 27

SCHWEINITZ PAINTINGS OF FUNGI 127

Table 1 (cont'd)

Thelephora a gaa Ill ei 9
Thelophora puni Vv 402 36
Thelophora concentrica Vv 401 35
Thelephora marginata Ill 219 31
Leotia truncorum 411 5
Helvella fistulosa Vv 414 6
Tremella saligna iI 235 8
Peziza clavus 423 120
Peziza betuli Il 240 9
Peziza fusco-can Il 241 10
Peziza carbonaria il 242 15
eziza fascicularis Hl 243 16
Peziza digitalis Vv 418 105
Peziza membranacea Vv 419 109
Peziza brunnea Ill 250 44
Peziza ink ea ll 251 45
Peziza Ill 250 43
Pez. ‘0- Bee Vv 422 117
Peziza theleboloides 423 121
Peziza albo-violascens ll 252 49
Peziza leucotricha Vv 422 116
Peziza sericea Ill 247 31
Peziza porioides Ill 251 46
Peziza lonicerae lll 251 47
Peziza laciniata Hl ao) 65
Peziza sanguinolenta Il 261 80
Peziza virens lll 261 81
Peziza betulina Ill 261 719
Peziza verrucaria (verucaria in Icones) V 422 118
Peziza pulveracea Il 265 93
Peziza led V 424 123
Peziza c lea Vv 422 119
Ascobolus lignatilis I 267
Stilbum hyalinum Vv 426 1]
Periconia flavovire Vv 430 4
eratium m pores (neeclik in Icones) Vv 429 2
fsaria monilioi Vv 432 4
Erineum toons Vv 445 8
Mesenterica grisea Vv 452 2
Mesenterica sanguinolenta hs 453 3

ACKNOWLEDGMENTS

I thank A. E. Schuyler at the Academy of Natural Sciences of Philadelphia for suggesting
this project and directing me throughout its pai and Carol Spawn for access to the
archived artwork of Schweinitz at the Academy.

REFERENCES

ALBERTINI, J. B. v. & L. D. v. SCHWEINITZ. 1805. Conspectus fungorum in Lusatiae superioris agro Niskiensi
arescentium e methodo Persooniana. Lipsiae Sumtibus Kummerianis

BARNHART, J. H. 1934. The botanical correspondents of Schweinitz. Bartonia 16: 19-36.

BENEDICT, D, M. 1934. Lewis David von Schweinitz, the mycologist. Bartonia 16: 12-14.

128 BARTONIA

JOHNSON, W. R. 1835. A memoir of the late Lewis David von Schweinitz, P.D. with a sketch of his scientific
labou urs. ANE by order of the Academy of Natural Sciences of Philadelphia. William B. Gibbons,

phia.
PENNELL F. W. 1934. The botanist Schweinitz and his herbarium. Bartonia 16: 1-8.
PHILLIPS, V. T. & M. E. PHILLIPS. 1963. Guide to the manuscript collections in the Academy of Natural Sciences
of Philadelphia. Special Publ. No. 5, Academy of Natural Sciences of Philadelphia
SCHWEINITZ, L. D. v. 1804. Fungorum Nieskiensium icones, Pars. I, II, III, and V. Uripublishea:

Bartonia No. 59: 129-133, 1996

OBITUARIES

Gilbert Raymond Cavileer (1911-1992)

On I January 1992, Gilbert Raymond Cavileer, Pine
Barrens native and natural history counselor, died in
Philadelphia’s Hahnemann Hospital. His was the passing
of yet another extraordinary field botanist, approximately 3
years after the death of Louis Edwin Hand (Bartonia No.
55:1-3, 1989).

I met Gil through Joseph M. Welch, then Assistant
Manager of Brigantine National Wildlife Refuge, when |
was working in Washington, D.C. It was the beginning of
an enduring friendship of over 3 decades. At first when we
got together, were mainly explored the Pine Barrens; in
recent years we were more likely to visit such places as
Brigantine National Wildlife Refuge, South Cape May
Meadows and Higbee’s Beach, Corson’s Inlet, the Seven
Bridges area near Tuckerton, or Brigantine Beach. My
memories of Gil are thus intertwined with pitch pine stands,
cedar bogs, sand and surf, the tangy smell of the salt marsh,
monarchs migrating, snow geese yipping—in summary, a host of priceless hours, shared with
a cherished companion.

Gilbert was of English, Dutch, and Irish background. His ancestors included lineages
which trace their history in America to the 1600s, for example Cavileer, Veale and Claypool,
and the late 1700s and early 1800s, for example Endicott and Maloney.

Gil was the oldest of three boys (a 4th child, James C., died in infancy in 1924). William
M. was born in 1917 and Watson S. in 1922. Bill was mainly a boat builder; he is now a
part-time security guard at William B. Kessler Memorial Hospital in Hammonton, New
Jersey. Watson became the Supervisor of Manufacturing for Beckman Instruments in
Fullerton, California; he died several months after Gil.

Their father, Raymond S. (1879-1933), had a variety of occupations, largely oriented to
the sea. He was a sea captain—with 2 brothers he owned a vessel, and with a cousin he
Owned a shipyard in Camden, New Jersey; he also was a boat builder, clam digger, clam and
Oyster buyer, carpenter, and farmer.

Their mother, May Maloney Cavileer (1887-1977), was a nurse and teacher. She was a
warm and cheerful person, who would sing while doing house work. Her "fling for the
year," as Bill Cavileer expressed it, was an annual trip to the Atlantic City racetrack. In her
declining years she moved to California, where Watson and his family were living, to be in
@ warmer climate.

Gil’s mother probably was more influential in his life than was his father; certainly she
was home nearly all the time, whereas Gil’s father often was away from home. Also Gil’s
father favored Bill over the other boys. However, Raymond Cavileer’s admonition, "If you
can’t improve on silence, keep quiet,” definitely was part of Gil’s philosophy.

129

ce

130 BARTONIA

The family home was on the edge of the Pine Barrens in Port Republic, New Jersey. The
Cavileers grew their own vegetables, had chickens and turkeys, raised pigs which were
bedded in marsh hay harvested by the boys, and obtained a variety of fish and shellfish from
nearby creeks, inlets, and bays. Young Gilbert was curious about animals; he brought home
snakes and turtles, and at age 12 was given a subscription to Nature Magazine. The house
was always full of books, and the boys were also exposed to classical music and opera from
a crank-up Victrola.

After graduating from Egg Harbor City High School, Gil dug clams and trapped for a
while before heading west. These were depression years, when a popular song was "Brother,
can you spare a dime?" During these years, Gil worked at Civilian Conservation Corps
camps in Idaho and California, and ranches in California. He was building boats in Atlantic
City when he was drafted into the U.S. Signal Corps in World War II. Sent to Germany,
he stayed there for about a year and a half after the war ended, replacing communication
lines. He then returned to New Jersey and lived with his mother. When jobs were scarce
after the war, Gil and Bill pulled sphagnum moss and sold it to nurseries, but Gil made his
living mainly as a boat builder and clam digger.

He never married. A special friend of recent years was Betty Hann of West Wildwood,
New Jersey.

Gil’s residence from 1955 until his death was 106 Main Street in Port Republic. The left
part of the house was built between 1790-1800, and the right half in about 1870.
Housekeeping and record keeping were relatively low on Gil’s list of priorities. I would
return after a year’s absence, for example, to find that the open space on the kitchen table
had become even smaller because of the proliferation of more magazine, letters (some
unopened), announcements of meetings (past, present, and future), subscription blanks,
financial appeals, utility bills, Social Security forms, bank slips, discarded (?) silverware, salt
and pepper shakers, pill containers, pens and pencils, erasers, plant remains (vascular and
non-vascular), etc. I often thought of the table top as a potential archaeological site! A
similar fate overtook the house in general, and also his truck. It was not at all unusual to
find a stray item of interest, for example a vintage letter, in an odd and unexpected place.

Gil essentially was a self-taught naturalist, although he did take a few courses, for
example at the Wetlands Institute in Stone Harbor, New Jersey. For the identification of
vascular plants, he used both Gray’s Manual and Britton and Brown, but preferred the keys
in Gray’s. In the field he carried an 8x hand lens, and at home he had a binocular
microscope. His knowledge of natural and local history was legendary, and always freely
shared.

He led wildflower walks and gave programs for a number of organizations, for example,
the Philadelphia Botanical Club, the Wetlands Institute, the Cape May Geographic Society,
the Audubon Society, and the Jersey Cape Shell Club. He was a member of the Philadelphia
Botanical Club and the New Jersey Chapter of The Nature Conservancy. With John
Gallegos, Gil conducted rare plant surveys in the late 1970s and early 1980s in four acid
meadows: Big and Little Goose Ponds, Barkwoods Pond, and Hirst Pond. The results are
detailed in two unpublished reports, and in a paper, "The influence of fluctuating water
levels upon the incidence of rare plant species of southern New Jersey," which was read at
a meeting of Stockton College in Pomona, New Jersey. Gil also assembled vascular plant
lists of other areas, for example the Holgate Unit of Brigantine National Wildlife Refuge,
The Nature Conservancy’s South Cape May Meadows, and the U.S. Coast Guard Station at
Cape May. In his latter years he worked part-time for Inman Environmental of Pleasantville,

OBITUARIES 131

New Jersey, where among other duties he assisted in compiling a vascular plant list of
Atlantic County. He collaborated on Snyder and Vivian’s Rare and Endangered Vascular
Plant Species in New Jersey, which was published in 1981, and looked for endangered
species at Brigantine National Wildlife Refuge in more recent years. Gil also banded birds
for a number of years.

He also traveled whenever his finances and health allowed it. In the latter part of his life
he journeyed to the northeastern United States and far eastern Canada, Florida, Wisconsin,
the northwest, and the southwestern United States. On one western trip he was joined by
Ernest A. Choate and on another by the author of this obituary, but usually he traveled alone,
staying with friends at his destination. His field companions over the years included Hollis
Koster of Green Bank, Frank Hirst of Stockton, Maryland, Keith Seager of Cape May, and
David B. Snyder of New Jersey’s National Heritage Program.

Gil continued his botanical explorations up to the time of his fatal illness. On one of our
last outings, we were joined by Seager and Snyder on a hot, muggy day near Cold Spring
in Cape May County; we visited two areas, one for the spike-rush Eleocharis halophila, and
the other for the rush, Juncus coriaceus. The heat index was 100 degrees F, and we were
all feeling its effects, especially, or so it seemed, Gilbert, who suddenly blacked out and
collapsed after our last hike. But I wondered if it was entirely from the heat, because he had
told me he had been having dizzy spells recently. A few days later, when we said goodbye
at the Pomona Airport, I wondered if we would ever get together again. On the phone a few
weeks later, Gil told me that he had Lyme Disease, but this diagnosis proved to be incorrect.
He had leukemia. His condition worsened, and he died in the hospital. In his younger years
Gil sometimes walked on a beach in a storm; there was a strong Northeaster on the day of
the funeral, and, as Budd Wilson, Jr., a long-time friend of Gil’s, wrote in a letter to me,
"Somehow going out in a Northeaster seemed appropriate for Gil." He was buried in the
family plot by St. Paul’s Methodist Church in Port Republic.

When asked about his health, Gil was apt to say, "Struggling along." His attitude toward
life was simple and to the point: enjoy each day, yet realize that death is natural and
inevitable.

Snyder and Seager took responsibility for Gil’s herbarium. Earlier collections are at the
Academy of Natural Sciences and Rutgers University.

Peter Kalm, a Swedish naturalist and student of Linnaeus, met John Bartram in 1748; his
comments about the Quaker botanist remind me of Gil, and might serve as his epitaph: "He
has acquired a great knowledge of natural philosphy and history, and seems to be born with
a peculiar genius for these sciences .... We owe to him the knowledge of many rare plants
which he first found .... He has shown great judgment and an attention which lets nothing
escape unnoticed. Yet with all these qualities . . . he did not care to write down his
numerous and useful observations .... I have often been at a loss to think of the sources
whence he obtained many things which came to his knowledge. I, also, owe him much, for
he possessed that great quality of communicating everything he knew" (Peter Kalm’s Travels
in North America. The English version of 1770. Revised from the original Swedish and
edited by Adolph B. Benson. Volume I. Wilson-Erickson Inc., New York, 380 pages, pp.
61-62).

132 BARTONIA

I thank Bill Cavileer for information on Gil and the Cavileer family, and Keith Seager for
biographical details. The photo for this obituary was taken on 17 September 1981 near Cold
Spring, Cape May County, New Jersey.

KENNETH I. LANGE
Department of Natural Resources, Devil’s Lake State Park, Baraboo, WI 53913

Charles T. Brightbill (1935-1994)

On July 26, 1994, Pennsylvania lost one of its most valued
teachers, naturalists, photographers, historians, and friends,
Chuck Brightbill, who died after struggling more than a
year with cancer.

Born in Chambersberg, Pennsylvania, on November 25,
1935, Chuck lived most of his adult life in nearby Mercers-
burg. He was a 1958 graduate of Lebanon Valley College
and received a master’s degree in music education in 1968
from Trenton State College, New Jersey. He taught music,
mostly elementary and vocal and beginning band instru-
ments, for 10 years in New Jersey and retired in 1989 after
21 years in Pennsylvania.

I first met Chuck in the summer of 1979 while he was ecology director at Sinoquipe Scout
amp. That summer I retook Bird Study merit badge so that I could learn more about birds;
I had already heard about Chuck’s reputation as a keen birder. I remember vividly the first
contact I had with him as he and I stood on the porch of the Dining Hall and he pointed out
a chipping sparrow singing in a Virginia pine. For 15 years Chuck was a close friend and
mentor—he encouraged my interests in biology and taught me to cherish knowledge and share
it with others through walks and slide presentations. Chuck, his mother, my parents, and I
took many trips together, sometimes solely for birding while other times solely for plants;
for several years, we spent New Years birding at Chincoteague, Virginia.

Chuck was best known for his nature walks and slide presentations. His personality and
teaching methods aided children and adults in their enjoyment of his talks. He developed
presentations based on his travels (e.g., "Wildlife of Alaska"—the photo in this obituary was
taken while he was birding in Alaska on 16 July 1978), the local flora and fauna, and
historical events. He was a talented actor; thus, in his historical presentations he was well-
known for giving talks on special topics and dressing as an Indian, frontiersman, Civil War
soldier, or Scot Highlander. In fact, he took his role so seriously as to get a Mohawk haircut
instead of wearing a wig.

Chuck was active in various community organizations and served several leadership
capacities: Tuscarora Education Association, Conococheague Audubon Society, Boy and Cub
Scouts, Towne Singers in Chambersburg, Fort Loudon Ruritan Club, three local historical
societies, and the Maryland Ornithological Society in Washington County. He volunteered
for 17 years with the Pennsylvania Bureau of State Parks and worked for 6 summers as a
nature director at Cowans Gap State Park. He helped to organize the Tuscarora Wildlife

OBITUARIES 133

Education Project and its Natural History Museum, and was involved in rebuilding the
historic Fort Loudon on the original site.

Chuck, as a self-taught botanist, gave leads to rare plants and special habitats in Franklin
County, Pennsylvania. For example, Chuck for several years led a field trip to southwestern
Franklin County for the Conococheage Audubon Society (in which | participated), and
stopped at a mesic upland woods to show the luxurient herbaceous layer. Chuck recognized
that the flora as unusual but never thoroughly investigated it. Starting in 1986, Dr. Larry
Klotz (Shippensburg University, Pennsylvania) and I studied the flora in detail and recorded
five state rarities including one species new to the state (see Bartonia 56:29-33; 1990). Larry
and I led a field trip for the Philadelphia Botanical Club to this area on 5 May 1991 (see
Bartonia 58:141-142; 1994). Chuck also gave us leads on several additional rare species in
the nearby area as well as the bluffs along the West Branch of the Conococheague Creek
(see Bartonia, Supplement to 57:16-41; 1993).

Chuck will be immensely missed, but will be remembered for his work. The nature center
at Cowan’s Gap was named "Brightbill Interpretive Center" in 1994, and the museums at the
James Buchanan High School for the Tuscarora Wildlife Education Project and at the Fort
Loudon Historical Site will be named after him. In 1994, he received the Meritorious
Service Award from the National Audubon Society.

JEFFREY L. WALCK
School of Biological Sciences, University of Kentucky, Lexington, KY 40506-0225

Bartonia No. 59: 134, 1996

REVIEW

Pennsylvania Higher Plants Volume I. Field Manual, by James A. Schmid. Schmid and
ompany, Inc., Media. 1994. 182 pp. $35.

New Jersey and Pennsylvania Higher Plants Volume II. Desk Manual, by James A.
Schmid and John T. Kartesz. Schmid and Company, Inc., Media. 1994. 443 pp. $55.

These two volumes are an attempt to make available to others the working files of the
Schmid and Co. regarding the wetland indicator status and state and federal rare plant
ces of vascular plants. While the compact size of Volume I (5 x 8 in.) may make

t a handy field reference, the term manual is misleading. In fact this book is a list and the
infansalen therein only accessible to one who has already identified the plants in question.

Volume I actually consists of two lists, one alphabetical by genus and species and a
second alphabetical by common name; family name and wetland and rarity codes are
repeated in each section. It has taken some ingenuity to come up with unique common
names for each species in the list, an effort that has involved inventing names for some
sedges and other plants that do not have generally accepted common names. Schmid and
Co. have also augmented the U.S. Fish and Wildlife Service wetland indicator codes, based
on their own field experience.

Volume II repeats the information in Volume I with the addition of rather cryptic coded
notes on historic wetland indicator status, according to several other sources, and a growth
habit code. While Volume I includes some synonymy in the alphabetical listings, Volume
II contains a more extensive listing of "Obsolete Latin Names" referenced to "Current Latin
Names." The larger format of Volume II (8.5 x 11 in.) makes it less useful in the field,
perhaps one characteristic in its favor, however, is the inclusion of lists for both Pennsylva-
nia and New Jersey. For a user interested in only Pennsylvania, there is little to be gained
by investing in Volume II, the Rhoads and Klein 1993 Vascular Flora of Pennsylvania:
Annotated Checklist and Atlas (American Philosophical Society, Philadelphia) is a much
better buy at $50. It also contains wetland and rarity status as well as synonymy, growth
habit, habitat and distribution maps for each species within the state.

According to the introductions in both volumes, the 3373 Pennsylvania plants inlcuded
in these references were derived from a list compiled by John Kartesz and represent species
"known to grow wild or to persist without continued, deliberate cultivation . . . plus a few
weedy exotics that we have encountered thriving in the field but not found in previous
reports." This compares with 3318 taxa included in the Rhoads and Klein book. Diffrences
in taxonomic interpretations account for some of the discrepancy. However, the scientific
validity of Schmid and Kartesz’ work would be enhanced if the source of their records were
more clearly defined and new state records of occurrence were properly vouchered with
herbarium specimens.

ANN F. RHOADS

134

Bartonia No. 59: 135-137, 1996

NEWS AND NOTES

FIRST RECORD OF DRYOPTERIS CLINTONIANA (D. C. EATON) DOWELL IN
LANCASTER COUNTY, PENNSYLVANIA. While botanizing in a familiar spot near Safe
Harbor, in Lancaster Co., PA, on September 26, 1994, James C. Parks discovered a
population of about 30 plants of the fern genus Dryopteris. The plants were of various sizes,
indicating a stable, reproducing population. They were growing on an elevated flood plain
terrace above a small stream in a generally steep-sided, forested ravine. The overstory trees
were Platanus occidentalis and Acer negundo. The shrub layer contained Lindera benzoin.
Herbs present included Hemerocallis, Impatiens spp., and Polystichum acrostichoides. The
largest Dryopteris ferns grew at the edge of the terrace at the base of a steep slope, in rich,
organic soil, but several small ones were near the streamside bank. The largest fonds
exceeded 0.8 m in length and were initially thought to be D. celsa or a hybrid of it. Both
are reported from further south in Lancaster Co. Careful examination of specimens led Parks
to determine that they were D. clintoniana. Subsequently, this determination was confirmed
by James T. Montgomery. He noted that the basal pinnae were triangular, the scales a
uniform, dull brown, and very few sporangia had not shed, all indicative of this fertile
hexaploid species. Specimen records, provided by Ann Rhoads from the Flora of
Pennsylvania Database at the Morris Arboretum, indicate that D. clintoniana is recorded
from York and Dauphin Counties but that most of the PA records are from the glaciated
northeastern region. As Montgomery observed, D. clintoniana has a strange distribution in
Pennsylvania and is uncommon even where it is found. This thriving population is the
southernmost known in Pennsylvania and adds to this perplexing distribution. Voucher
specimens have been deposited at The Academy of Natural Sciences of Philadelphia (PH)
and Millersville University (MVSC). J. C. PARKS AND J. T. MONTGOMERY.

1993 NORTHERN DELAWARE RARE PLANT SURVEY HIGHLIGHTS. The 1993
botanical field season in New Castle County and northern Kent County proved fruitful in the
discovery of new state records, relocations of historical records, and of new rare plants sites.
In the Brandywine drainage of the Piedmont physiographic province, discoveries included
the first record of Cardamine angustata ($2) for that drainage (previously only known from
the Red Clay Valley); Delaware’s second site for Veronicastrum virginicum (S1); new
Stations for Andropogon gerardii (S1) and Bromus pubescens (S2); and a modern record for
the more northern Callitriche palustris (former SH). The Red Clay drainage yielded a
second site for Carex granularis (S1) and the first site for Carex gracilescens (former SH);
Euonymus atropurpurea (former SH) growing near new populations of Smilax hispida (S1)
and Andropogon gerardii; Juncus subcaudatus (S1) growing in a low wet meadow; and
finally Leptoloma cognatum (former S1) growing in abundance on a dry roadbank and an
adjacent mown pasture. The White Clay drainage produced the new state record Agrimonia
striata; a third Delaware station for Heliopsis helianthoides; and Zannichellia palustris
(former SH). Other Piedmont drainages also provided Calystegia spithamaea (former
in bloom in a young dry open woods; Delaware’s second extant station for Chamaelirium
luteum (S1); a third population of Geum vernum (S1); two stations of the new state recor
Osmorhiza claytonii; and Stellaria longifolia (former SH). An additional find was a small

135

136 BARTONIA

but healthy blooming population of Trillium erectum, not seen in the state for over a century,
and in an apparently native setting.

On the Coastal Plain of central and southern New Castle County, finds included a second
population of Zannichellia palustris in a tidal habitat; two populations of Silene virginica
(former SH); a second population of Matalea caroliniensis (S1); Cyperus refractus, a new
state record; and finally, the wetland shrub Cornus stricta (unofficially a new state record
apparently overlooked by Tatnall, although previously recorded for the state by W. S. Taber
in Trees of Delaware (1930). Old spoilage and dredge pits along the C & D Canal were
found to contain unusual pond, bank, and ridge habitats where we discovered Juncus
diffusissimus and Aristida curtisii, both new state records; the latter was found growing with
a small population of Hudsonia tomentosa (S4), not previously recorded for New Castle
County. A freshwater impoundment provided the first modern collection of Cicuta bulbifera
(former SX), not seen in more than a century. Limnobium spongia (a former SH), and
Utricularia gibba (S2) were also found at this site.

On the Coastal Plain into Kent County, finds in the western part of the county included
Sporobolus clandestinus, Lespedeza stuevei (both former SH species), Commelina erecta v.
angustifolia (S1), Paronychia fastigiata (S1), and another population of Cyperus refractus.
Several late-season forays yielded more populations of Leptoloma cognatum and small
colonies of Quercus prinoides (S1) and Pycnanthemum torrei (S1). Finally, explorations
along the New Castle County coastline turned up several populations of Leptochloa
fascicularis v. maritima within disturbed wetlands habitats; this indicates that it may be
found more frequently in similar situations in the future. These rare plant surveys were
funded by the Delaware Natural Heritage Program. JANET EBERT AND JACK HOLT

PENNSYLVANIA SCIENCE OFFICE OF THE NATURE CONSERVANCY 1993
VEGETATION SURVEY HIGHLIGHTS. During the 1993 field season biologists from The

ature Conservancy’s Pennsylvania Science Office (PSO) conducted county natural areas
inventories (CNAI) in Sullivan and Wyoming counties. Surveys of sites identified from
aerial photographs and historical data helped us locate several new sites for state-listed plant
species (Dept. Environmental Resources 1987, 1993) in the two counties. Nomenclature is
that used by Rhoads & Klein (1993). State Satus is from Title 25, Chapter 82: Conservation
of Pennsylvania Native Wild Plants (Dept. of Environmental Resources 1987, 1993).

Jacob’s Ladder (Polemonium van-bruntiae Britt.), Pennsylvania Endangered, was located
in one forested swamp in eastern Sullivan County. There are several vague historical records
from this vicinity but recent searches had been unsuccessful (A. E. Schuyler, pers. comm.);
the species was last reported in 1934. The only other known extant site in Pennsylvania is
in Wayne County. The Wayne County population has been declining for several years and
was reduced to one plant as of 1993. This site in Sullivan County may represent the only
viable population of this species in Pennsylvania.

Floating bladderwort (Utricularia radiata Small) had been seen last in 1927 in southern
Bucks County and was believed extirpated (Rhoads & Klein 1993). Populations were found
in two lakes in eastern Sullivan County while conducting searches for Potamogeton
confervoides. Its discovery makes it a Pennsylvania Endangered species (Dept. of
Environmental Resources 1987).

Slender cliff-brake (Cryptogramma stelleri (Gmel.) Prantl), a Pennsylvania Endangered
fern, was found at one new location in western Sullivan County. The population is only the
fifth extant in the state and appears to be the second largest.

NEWS AND NOTES 137

Tuckerman’s pondweed (Potamogeton confervoides Reichenb.), Pennsylvania Threatened
and a candidate for listing under the Endangered Species Act, was located at two new sites
in western Wyoming County during the CNAI. One of the two new sites is a small glacial
kettlehole lake with several homes around it and the other is a beaver pond located on state
game lands. We also attempted to find it at several other known sites in the two-county area
as part of a separate wetland plant contract. Only one of the species’ known sites could be
reconfirmed. At the time, October 1993, the population appeared on the verge of being
crowded out by another aquatic plant.

Also noteworthy were new populations of several other listed species discovered during
the CNAI. These include five locations for creeping snowberry (Gaultheria hispidula (L.)
Muhl. ex Bigel.), four locations for Labrador tea (Ledum groenlandicum Oeder) and one for
bog sedge (Carex paupercula Michx.), a county record.

These County Natural Areas Inventories are supported by the individual counties and by
the PA Department of Community Affairs through Recreational Improvement and
Rehabilitation Act grants. The wetland plants survey was sponsored through a grant to the
Dept. of Environmental Resources from the United States Environmental Protection Agency
under the Clean Water Act’s 1993 State Wetlands Protection Development Grant Program.

The information on rare species and exemplary natural communities from the CNAIs is
contributed to the Pennsylvania Natural Diversity Inventory data base. The PSO works in
cooperation with the Western Pennsylvania Conservancy and the Bureau of Forestry, PA
Department of Environmental Resources, to maintain the PNDI data base of the state’s rarest
species, exemplary natural communities, geologic features and managed areas. As of this
writing, there are over 11,000 extant and historical records of plants, animals and
communities.

LITERATURE CITED

DEPARTMENT OF ENVIRONMENTAL RESOURCES. 1987. Title 25, Chapter 82: Conservation of Pennsylvania native
wild plants. PA Bull. 17 (49) of December 5. ; :
DEPARTMENT OF ENVIRONMENTAL RESOURCES. 1993. Title 25, Chapter 82: Conservation of Pennsylvania native

wild plants. PA Bull. 23 (25) of June 19.
RHOADS, A. F. & W. M. KLEIN, JR. 1993. The vascular flora of Pennsylvania: Annotated checklist and atlas.
American Philosophical Society, Philadelphia.

ANTHONY F. DAVIS, JULIE A. LUNDGREN AND JOHN R. KUNSMAN

Bartonia No. 59: 138-142, 1996

1993 FIELD TRIPS

16 May: Furnace Hills, Lancaster Co., PA. We hiked through the gamelands to see
numerous plants of Galearis spectabilis, Trillium cernuum, Cypripedium pubescens (nearly
past bloom), Adlumia fungosa, Hydrastis canadensis, Triosteum aurantiacum, and
Rhododendron periclymenoides. At Camp Shan area we saw Aplectrum hyemale and
Waldsteinia fragarioides. At Middle coke Wildlife Management Area we saw a stand of
about 200 plants of Jsotria verticillata in full bloom within an area of 20 square feet. A
small group ventured to Shenk’s Ferry to see Corallorhiza wisteriana in peak bloom.
Leaders: Tim Draude and Mark Larocque.

22 May: Woodlands Cemetery, 40th St and Woodland Avenue, Philadelphia, PA.
Formerly the country estate of William Hamilton, the Woodlands was an impressive
landscape garden in the 18th and early 19th centuries. John Lyon and Frederick Pursh
worked as gardeners and plants from the Lewis and Clark expedition were cultivated here.
Ginkgo biloba, Populus nigra "Italica", Ailanthus altissima, and Acer platanoides were
introduced into cultivation in North America at the Woodlands. To get some idea of what
is there now, we recorded the following woody plants: Acer pseudoplatanus, A. rubrum,
Aesculus hippocastanum, Broussonetia papyrifera, Buxus sempervirens, Castanea mollissima,
Celtis occidentalis, Chamaecyparis pisifera, Fraxinus americana var. biltmoreana, Ginkgo
biloba, Gleditsia triacanthos, Gymnocladus dioica, Juglans nigra, Liquidambar styraciflua,
Morus rubra, Picea pungens, Prunus serotina, Quercus palustris, Q. rubra, Sassafras
albidum, Sophora japonica, Styrax japonica, Taxus baccata, T. cuspidata, Thuja orientalis,
Tilia americana, T. cordata, Ulmus sp., Zelkova serrata. Leader: Alfred E. Schuyler.

5 June: Muskee Creek, Manumuskin River, Tuchahoe River watersheds, Pine Barrens of
Atlantic and Cumberland cos., NJ. We stopped at an open cedar bog in the Lawrens Branch
of the Manumuskin to see excellent stands of Eriocaulon compressum and Orontium
aquaticum in flower. Arethusa bulbosa in bloom and Pogonia ophioglossoides in bud were
also noted, as well as fine stands of Kalmia latifolia and K. angustifolia in the adjacent pine-
oak uplands. At Aetna Furnace along the Tuckahoe we saw patches of Liparis liliifolia on
mounds of slag with Ophioglossum vulgatum and Pyrola americana. In an old timber cut
in the Muskee Creek headwaters we noted a state record Chamaecyparis thyoides with a cbh
of 9 ft 7.5 in. The sedge genus Carex was studied throughout the day and the following
species were noted: C. abscondita, C. albicans v. emmonsii, C. albolutescens, C. annectens,
C. atlantica, C. barrattii, C. bullata, C. canescens, C. collinsii, C. crinita, C. debilis, C.
exilis, C. folliculata, C. howei, C. lurida, C. intumescens, C. cpa C. pensylvan-
ica, C. swanii, C. striata, C. stricta, and C. trisperma. Leader: Gerry Moo

12 June: Nockamixon State Park and Haycock Mountain, Bucks Co., “eg We stopped
along PA route 563 to see one of the five extant locations in the state for Phlox pilosa.
Other plants seen along the road include Krigia biflora, Linum intercursum, Lobelia spicata,
Pedicularis canadensis, and Scutellaria integrifolia. At Thatcher Road meadow we noted
Phlox maculata, Castilleja coccinea, Liatris spicata, and Lilium canadense. At the Parkway
Diner a Wood Turtle, Clemmys insculpta, was observed eating the ripened fruits of Fragaria
virginiana. Along the 0.5 mile trail to the top of Haycock Mountain, >50 species were

138

1993 FIELD TRIPS 139

encountered, including Adlumia fungosa, Obolaria virginica, Goodyera pubescens, Rubus
odoratus, and a single plant of Liparis liliifolia. Botrychium virginianum was abundant and
very ripe, ining large clouds of spores when touched. Leader: Greg Edinger.

13 June: Tyler Arboretum, Lima, Delaware Co., PA. This trip focused on the Pink Hill
serpentine barren and nearby woods. Noteworthy species included Pryrola virens, Ilex
verticillata, Chimaphila maculata, and Phlox subulata. Leader: John Ballas.

19 June: Pennypacker Mills County Park, Schwenksville, Montgomery Co., PA. Several
areas of the park have been allowed to revert to "natural" conditions that may have prevailed
during Governor Pennypacker’s ownership. In the first entrance field, which is moist, and
in a nearby wooded area, we encountered 26 species of carices, including Carex squarrosa,
C. bushii, C. hirtifolia, C. conoidea, C. glaucodea, C. laxiculmis, and C. lanuginosa. There
were many attractive wild flowers blooming such as Lobelia spicata, Salvia lyrata, and
Mentha spp. At the nearby property of club members Goeffrey Kaiser and Bruce Grimes,
we enjoyed a bountiful habitat of woodlands, wetlands, stream, and field. Leaders: Ann
Newbold and Heinrich Zoller.

20-24 June: Lancaster, PA. The 1993 Joint Field Meeting of the Philadelphia Botanical
Club, the Torrey Botanical Club, and the Northeastern Section of the Botanical Society of
America was housed at Millersville University near Lancaster. There were three full days
of field trips. Localities visited included the Furnace Hills area in northern Lancaster
County; serpentine barrens in southern Lancaster County; limestone cliffs in central
Lancaster County; and areas along the Susquehanna River in Lancaster and York counties.
Trip leaders were Timothy Draude, William Olson, Roger Latham, Ann Newbold, and Fred
Habegger. Evening speakers included Stephen Sylvester, who discussed the geological
resources of the Lancaster area; Roger Latham, who spoke about Pennsylvania’s serpentine
and glacial till barrens; Guy Steucek, who presented a program about Lancaster County
woodlands; and Mike Batcher, who program dealt with the management of threatened flora
and fauna. As a result of aggressive development, degradation of water resources, or
invasion by exotic plant species, little remains of the native woodlands of Lancaster County.
There are bright spots, however: small patches of native plants are being protected by the
Nature Conservancy and other local organizations, and there are efforts to link remaining
core habitats with riparian corridors. The chairperson was Cyane Gresham, and the treasurer,
Karl Anderson. :

10 July: Tyler Arboretum, Lima, Delaware Co., PA. We visited the Pink Hill serpentine
barren particularly to see the beautiful display of Lilium philadelphicum. Noteworthy
associated species were Oenothera fruticosa, Lobelia spicata, and Ceonothus americanus.
Leader: John Ballas. ae

11 July: Green Acres Natural Area, Medford Leas, Burlington Co., NJ. We were invited
by the Arboretum Oversight Committee of Medford Leas to conduct a field trip of their
Natural Area for the express purpose of updating their species list, developed by the leader
in 1992. The Natural Area is comprised of four major inner coastal plain communities: 1)
palustrine nontidal emergent wetland, 2) palustrine forested wetland, 3) mesic mixed oak
forest, and 4) successional forests (one of mature Pinus virginiana; the other of young
Liquidambar styraciflua). Focus was on the palustrine forested wetland along Sharp’s Run
and Southwest Branch of Rancocas Creek (= Haynes Creek). Situated at the foot of a steep
Slope, this rich floodplain, underlain by sandy loam with some glauconite content, is
characterized by a mixed forest community of fast-growing hardwoods. Important trees,
several of them huge specimens, include Acer rubrum, Platanus occidentalis, Magnolia

140 BARTONIA

virginiana, Salix nigra, Quercus palustris, Q. falcata, Fraxinus americana, Liriodendron
tulipifera, Liquidambar styraciflua, Ulmus americana, Carya spp., Prunus serotina, Carpinus
caroliniana, Ilex opaca, Betula nigra, and some Acer saccharinum, Cornus florida, and
Sassafras albidum. Common shrubs and vines scattered throughout are Lindera benzoin,
Viburnum dentatum, V. prunifolium, Cornus amomum, Toxicodendron radicans, Euonymus
americanus, Vaccinium corymbosum, Clethra alnifolia, Ilex verticillata, Sambucus
canadensis, Parthenocissus quinquefolia, and Mitchella repens. A typical diverse herb layer
also prevails, including such species as /mpatiens capensis, Lobelia cardinalis, Chelone
glabra, Caltha palustris, Anemone quinquefolia, Podophyllum_ peltatum, Aquilegia
canadensis, Symplocarpus foetidus, and an array of ferns. Although infused with various
plantings (including exotic species) by the residents, this palustrine forest has retained its
native character and anticipated species composition. We added 13 taxa not previously
recorded: Chenopodium album, Polystichum acrostichoides, Deparia acrostichoides (=
Athyrium thelypteroides), Dryopteris carthusiana, Potamogeton epihydrus, Ceratophyllum
demersum, Acer platanoides, Ulmus rubra, Corylus americana, Poa pratensis, Solanum
dulcamara, Physalis subglabrata, and Rudbeckia laciniata. The communities we examined
were well known to Witmer Stone, who explored the Medford flora at the turn of the
century, documenting some 750 species while based at a cabin almost on site. In comparing
some of our discoveries to those reported by Stone (1911), we were surprised, for example,
by his omission of Acer saccharinum and Fraxinus americana. Of the former Stone says
"sparingly southward within our limits"; of the latter (of which we saw huge specimens) he
states "rare southward." Finally, all were given an opportunity to see the largest reported
hickory in the state: a Bitternut, Carya cordiformis, with a cbh of 13 ft 3 in. Leader: Ted
Gordon.

14 August: Smithville Mansion and other parks in western Burlington Co., NJ. At Mill
Creek Park in Willingboro, high water levels kept us from fully exploring the intertidal zone,
but we did find Lobelia chinensis, a rapidly-spreading adventive, in bloom at the high tide
line on a sandy shore. In the same area were Eleocharis erythropoda, Lycopus rubellus, and
Helenium autumnale. Upland woods nearby had Physocarpus opulifolius and an unusually
large stand of Lycopodium obscurum, while the shores of a small, stagnant pond produced
Eclipta prostrata, Hypericum mutilum, and Veronica anagallis-aquatica. At the Rancocas
Nature Center in Westhampton, the shores of a nearby pond produced Zizania aquatica,
Triadenum virginicum, Rhexia mariana, Juncus acuminatus, Rhynchospora capitellata,
Galium tinctorium, Utricularia gibba, and many other species. Wood edges had Humulus
lupulus and Juglans cinera growing near J. nigra. Finally, the shores of Smithville Lake in
Easthampton Township had Lobelia cardinalis in good bloom, with Verbena hastata and
Ascelpias incarnata, Mikania scandens, Apios americana, Lysimachia_ terrestris, Carex
crinita, and Lindernia dubia also present. Of particular interest here were numerous plants
of Azolla caroliniana, growing with Lemna in warm, shallow water; this floating fern,
seldom seen in New Jersey, was first noticed here in 1992. The liverwort Riccia natans was
also observed. Leader: Karl Anderson.

15 August: Savannahs and Cedar Swamps, Batsto River, Burlington Co., NJ. Extending
along ancient oxbows and coves, the cedar swamps, savannahs, and quaking bogs of the
Batsto River above the village of Batsto attracted botanists as early as 1805 and, perhaps,
even earlier. On a savannah well above the village, we had little difficulty finding our target
species: Rhynchospora oligantha, Xyris fimbriata, Schizaea pusilla, and flowering specimens
of about a dozen Platanthera integra and two P. cristata. The remarkably diverse flora

1993 FIELD TRIPS 141

encountered here included many other rare as well as common species: Nathecium
americanum, Juncus caesariensis, Muhlenbergia torreyana, Carex livida, Pogonia
ophioglossoides, Platanthera clavellata, Calopogon tuberosus, Hypericum denticulatum,
Sabatia difformis, Lophiola aurea, Lobelia canbyi, L. nuttallii, Rhynchospora cephalantha,

. alba, R. gracilenta, R. fusca, Xyris torta, Iris prismatica, Polygala cruciata, Smilax
laurifolia, Scleria reticularis v. pubescens (= S. muhlenbergii), Cladium mariscoides,
Orontium aquaticum, Calamovilfa_ brevipilis, Eleocharis robbinsii, E. tuberculosa,
Proserpinaca pectinata, the typical carnivorous plants, Sphagnum cyclophyllum, and S.
portoricense (the latter, a new record). In shallow water along the margins of the stream,
we observed Utricularia purpurea, Potamogeton confervoides, Nymphoides cordata,
Sclerolepis uniflora, Peltandra virginica, Pontederia cordata, Sagittaria latifolia, and
Eriocaulon decangulare. At the request of the N.J. Forest Service, we conducted a survey
for successional herbs at a 1989-1991 cedar clearcut of 3.4 hectares at Penn Swamp Branch
west of the Tuckerton Trail. Prior to the harvest, a mature Chamaecyparis stand with a
sparse herb layer occupied this site, which now serves as a cedar regeneration experimental
plot. The site produced a substantial sum of 40 species of herbs, vines, and ferns. Although
no rare species were encountered, we were surprised by the occurrence of Bidens polylepis,
an interloper. Leader: Ted Gordon.

21 August: Sandy Branch, New Castle Co., DE. Along the western border of Delaware
but just across the state line in Maryland, we observed a small population of Luzula
acuminata in a vegetative state. (This taxon has only been recorded once in Delaware.) On
the edge of a somewhat drawn-down beaver impoundment, we saw extensive stands of
Deschampsia flexuosa on the slopes of an open chestnut oak stand. Along an adjacent
powerline cut we observed the following species, a number of which are normally found
only in the Piedmont: Cacalia atriplicifolia (mostly finished flowering), yards away from a
pocket of Lycopodiella appressa, Helianthemum propinquum, Lechea villosa, Castanea
Pumila, and Euphorbia ipecacahuanae. In a dry meadow above and near the cut, a single
stem of Cyperus refractus, at only its second Delaware station (it was only found int he state
this year), grew with Euphorbia corollata, Croton glandulosus, and Cenchrus sp. the
Moister trail down from the meadow Elephantopus carolinianus was blooming, and further
along more typically Piedmont species as Lactuca floridana, Cimicifuga racemosa, and
Polystichum acrostichoides. Toward the end of the uncultivated section of the powerline cut,
we observed Paronychia canadensis along a wood edge and on a side trail, Ceonothus
americanus, a large but local population of Helianthus divaricatus, and a small colony (long
past flowering) of Silene virginica at one of only two known Delaware stations. Finally, the
leaders had planned to take the group a few miles north to see a population of Matalea
Caroliniensis on the Delaware/Maryland state line, but conveniently we found a small but
very large-leaved colony in flower along a rich wooded trail near the powerline cut.
Leaders: Jack Holt and Janet Ebert.

11 September: Batsto River, Burlington Co. and Route 30, Atlantic Co., NJ. We ventured
into Long Savannah along the eastern side of the Batsto River hoping to see Spiranthes
laciniata. We were unsuccessful, although our search of the previous year had produced
approximately 15 scattered plants here. Among the species seen wee Aster nemoralis, A.
novi-belgii, Eriocaulon decangulare, Schizaea pusilla, and Hypericum canadensis. A few
miles east of Hammonton along Route 30 we examined a fine station of >100 Gentiana
autumnalis with about 50% of the plants in bloom. Associated species in full bloom were
Heterotheca (Chrysopsis) mariana, Liatris graminifolia, Aster gracilis, and A. spectabilis.

142 BARTONIA

Leader: Mark Larocque.

18 September: Washington Park, Washington Township, Gloucester Co., NJ. It appears
that this central Gloucester County park had not been botanized previously by the club.
Trails, edges of fields, a series of young woodlands, and a few patches of mature forest
composed of plants common to the inner coastal plain were examined during intense
downpours. A significant feature was the presence of an Atlantic White Cedar swamp
supporting a number of common species associated with this community: Carex striata,
Chamaedaphne calyculata, Drosera rotundifolia, Juncus canadensis, and many others.
Observed elsewhere on the site were Chelone glabra, Juncus validus (a new station for the
county), Myrica heterophylla, Toxicodendron vernix (dense in most red maple swamps),
Aster novi-belgii, Agalinis purpurea, and Cyperus rivularis. Leader: Joe Arsenault.

Bartonia No. 59: 143-146, 1996

1994 FIELD TRIPS

I May: Fern Hill Farm, Gloucester Co., NJ. Club members toured the woodland garden
of this landscaped farm owned by the leaders. Many spring woodland ephemerals were in
bloom, including 18 species of trillium. Photographers in the group had a fine opportunity
for closeup photography at a leisurely pace. Leaders: John and Janet Gyer.

7 May: Williamson Park and Shenk’s Ferry, Lancaster Co., PA. We ventured onto a
limestone outcrop overlooking the Conestoga River to see Dodecatheon amethystinum (Pink
Shooting Star), Arabis lyrata (Lyre-leaf Rockcress), Aquilegia canadensis (Columbine), and
Pellaea atroupurpurea (Purple-stem Cliffbreak). At Shenk’s Ferry Wildflower Preserve in
the southern part of the county, we saw many spring flowers, including Mertensia virginica
(Virginia Bluebells), Polymnia canadensis (White-flower Leafcup), Hybanthus concolor
(Green Violet), Triosteum aurantiacum (Wild Coffee), Phlox maculata (Wild Sweetwilliam)
and Corallorhiza wisteriana (Spring Coralroot). Leader: Mark Larocque.

11 June: Atsion and Penn Swamp Branch, Wharton State Forest, Burlington Co., NJ.
Within the confines of the old "village" of Atsion were fine clusters of Opuntia humifusa,
thickets of fruiting Prunus maritima, and extensive cushions of flowering Minuartia
caroliniana. Along the abandoned railroad tracks nearby, we briefly explored a pitch pine
lowland forest (including moist depressions) with a history of frequent wildfire. Here we
noted in anthesis Melampyrum lineare, Iris prismatica, Panicum mattamuskeetense (=
Dichanthelium dichotomum var. dichotomum), and Fimbristylis puberula. Not yet in flower
were Muhlenbergia torreyana and Crotonopsis elliptica (Rushfoil), a tiny, wiry, branched
plant with a silvery aspect, growing in exposed sand. At our next stop, the primary mission
was to search the wetland along Penn Swamp Branch (= Goodwater Run), a small Batsto
tributary southeast of Quaker Bridge, for a small population of Eriophorum tenellum (Few-
nerved Cottongrass). However, an intensive search for this state-endangered sedge, last
observed here in the late 1960s, was unsuccessful. Within expansive Penn Swamp Pond,
ringed by Chamaedaphne calyculata, we focused on the shallow, upper half, comprised of
Carex exilis and C. atlantica hummocks associated with Chamaecyparis thyoides, Lophiola
aurea, and Pogonia ophioglossoides, the latter two in bud. Flowering species included
Utricularia fibrosa, U. subulata, Rhynchospora fusca, Eriocaulon compressum, Vaccinium
macrocarpon, and Gaylussacia dumosa. Areas of somewhat deeper water were dominated
by Cladium mariscoides, while less saturated zones contained patches of Muhlenbergia
forreyana, It appears that beaver activity has resulted in raising more permanently the water
level of this complex. With difficulty we also penetrated a mature cedar forest northeast of
the pond. A number of the larger trees were well over 100 years old. Windthrown es
lay helter-skelter. Beneath a typical ericaceous shrub layer, shaded carpets of sphagn
mosses harbored only a few species: Carex collinsii, Mitchella repens, Thelypteris ouastris,
and fine patches of 7. simulata. Future searches for Eriophorum tenellum should focus on
the 0.7 mile long wetland corridor extending west of the pond to a small bridge downstream,
nearer the Batsto River. Leader: Ted Gordon

26-30 June: Frostburg, MD. The 1994 Joint Field Meeting of the Philadelphia Botanical
Club, the Torrey Botanical Club, and the Northeastern Section of the Botanical Society of

143

144 BARTONIA

America was housed at Frostburg State University in western Maryland. There were three
full days of field trips. Localities visited included Green Ridge State Forest, Finzel Swamp,
the Frostburg City Watershed, and the C & O Canal in Maryland; and Larenim County Park
and Dolly Sods in West Virginia. Habitats seen included shale barrens, flood plains, shrub
swamps, deciduous woodlands, heath barrens, and grass balds. Evening programs included
an overview of Natural Communities of Western Maryland, by Edward Thompson of the
Maryland Department of Natural Resources; a look at the Geology of Western Maryland, by
Dr. Tom Small of the Frostburg State University Geology Department; a Survey of the
Herbaceous Flora in two Western Maryland Counties, by Dr. James Howell of Allegany
Community College; a discussion of the Trilliaceae, by Dr. Victor Soukup; and a program
about the Vegetation of the Gauley and Bluestone Rivers, by Dr. Bill Grafton of West
Virginia University. The Chairperson was Simon Dadydeen, of the Biology Department of
Frostburg State University; the Treasurer was Karl Anderson.

10 July: Taylor’s Preserve, Cinnaminson, Burlington Co., NJ. This trip investigated wood
edges, the margins of cultivated fields, shallow freshwater wetlands, and the gravelly
intertidal zone of the Delaware River. A list of 207 plants was compiled, of which 80 were
non-native species, as might be expected in a long-farmed, disturbed site. A copy of the list
is available on request. Some less-common species included Cyperus microiria, Carex
frankii, Galinsoga parviflora, and some very large, old, planted specimens of Quercus
macrocarpa. A single plant of Polygonum perfoliatum was found; this is the second
reported location for this invasive introduced plant in New Jersey, and the first record for
Burlington County. Another introduction, Lobelia chinense (in bloom), dominated the
intertidal zone, growing with Rumex verticillatus and scattered plants of Amaranthus
cannabinus, Myosotis laxa, and a non-flowering Polygonum sp. In addition to plants,
numerous species of butterflies and birds enlivened the trip. Leader: Karl Anderson.

18-28 July: Our club experienced a first—an out-of-bounds trip to Europe. Joined by a
number of American Botanical Society members, we traveled to Switzerland, primarily to
see its bountiful flora under the botanical supervision of one of the foremost taxonomists of
Switzerland, Dr. Heinrich Zoller, Professor Emeritus of the University of Basel. From
Zurich we looped through the Alps using a boat on Lake Lucerne, rackrailways on the
subalpine Central Alps, our own bus through mountain passes, a bus-on-a-train through
tunnels much too long to hold your breath, and cable cars for the upper alpine stages,
botanizing all the way (whenever we weren’t catching our breath from the wonder of the
scenery). At Zermatt we viewed the Matterhorn from Hotel Riffelberg very high up on its
magnificent slope, then walked to the celebrated Gorner Glacier where we picnicked on one
of our delicious Swiss lunches (hard bread, cheeses, yogurt, fruit and Swiss chocolate). The
plants we saw and named were legion, including gentians, tiny and blue, tall and yellow,
orchids by the dozens, species of Campanula, Silene, and Saxifraga, and of course the not-
to-be-forgotten Edelweiss growing alongside our trail. Leaders: Henrich Zoller, Max Seiler
of the Basel Botany Club, and Ann Newbold.

23 July: Millersburg, Dauphin Co., PA. We Ventured into a wet meadow along the
Susquehanna to see 75-100 Platanthera peramoena in mid-bloom. Other flowering plants
here were Ascelpias incarnata, Vernonia noveboracensis, and Rudbeckia laciniata. Invasive
grasses seem to have had a detrimental effect on the orchid’s habitat. The past two years
there have been >200 robust specimens (4-5 ft tall) of the Purple Fringeless Orchid. We
traveled to Route 324 to see a roadside cut with a dozen plants of Platanthera ciliaris,
Goodyera pubescens, Epipactis helleborine, Sabatia angularis, and a puzzling Eupatorium

1994 FIELD TRIPS 145

sp. Leader: Mark Larocque.

6 August: Malaga Lake and Vicinity, Franklin Township, Gloucester Co. and Upper
Pittsgrove, Salem Co., NJ. We explored both the west shore of Malaga Lake and walked
the broad, shallow bottom of the upper reaches of the lake. The disturbed shoreline at the
end of Malaga Drive supported Rhynchospora capitellata, R. macrostachya, R. chalaroceph-
ala, Eleocharis olivacea, E. tuberculosa, E. robbinsii, Cyperus rivularis, Dulichium
arundinaceum, Rhexia virginica, Lycopus uniflorus, Juncus canadensis, J. pelocarpus,
Panicum longifolium, Vaccinium macrocarpon, and Apios americana. In the lake submerged
species such as Utricularia purpurea and U. fibrosa were intertwined with floating leaved
plants of Nuphar lutea, Nymphaea odorata, and Nymphoides cordata, forming dense beds.
Half of the group explored a dense Chamaecyparis swamp along the Scotland Run stream
corridor above the main body of the lake. The shallow water surrounding the cedar stand
supported the same submerged and floating-leaved species listed for the lake, in addition to
Decodon verticillatus, Rhynchospora fusca, R. alba, Carex canescens, Eleocharis tenuis,
Triadenum virginicum, Xyris difformis, Drosera filiformis, D. rotundifolia, and D.
intermedia. The other half of the group concentrated on the oak-pine forest west of the lake.
This forest supported all of the typical oaks and pines as well as Carya pallida, Cornus
florida, Dichanthelium villosissimum, and D. acuminatum. On Nature Conservancy property
in Brotmanville, we investigated a small portion of a large tract of land west of the Maurice
River, north of Garden Road. Here a sand mine and surrounding disturbed land contained
Hudsonia ericoides and Monarda punctata, and Carya pallida was a significant component
of the oak-pine forest. A series of paths leading into red maple swamp bordering the
Maurice River led to pockets of Toxicodendron vernix. Leader: Joe Arsenault.

6 August: Montauk Point, Long Island, NY. The focus of this joint trip with the
American Association of Field Botanists was orchids. We visited a roadside cut in
Riverhead to see Malaxis bayardii, which differs from the similar M. unifolia in flower
structure and overall habitat range. At Napeague Harbor we saw the newly described
Platanthera pallens, which has been separated from P. cristata on the basis of flower
structure. Field examination of the two revealed significant differences in the lip, petals, and
coloration. The lip of P. pallens is significantly recurved, and the flower is a pale yellow.
We also saw a large population of Sabatia stellaris in a salt marsh adjacent to the P. pallens
Population. Finally, we visited several coastal pond shores to see Coreopsis rosea and
Gratiola aurea. Leaders: Eric Lamont and Mark Larocque. é

13 August: Martha Furnace and Oswego River, Burlington Co., NJ. We immediately
began exploring the diverse flora of the savannahs, quaking bogs, and cedar swamps that
border the Oswego above Martha. The presence of several flowering species of bladderwort
in close proximity allowed for convenient comparison. We examined Ufricularia fibrosa,
U. cornuta, U. juncea (including forma virgatula), the tiny pinheads of U. subulata forma
cleistogama, and the delicate purple flowers of a few lingering specimens of U. resupinata.
Widely distributed species included Rhynchospora fusca, R. alba, Eleocharis robbinsii, E.
tuberculosa, Muhlenbergia uniflora, Cyperus dentatus, Xyris torta, Juncus canadensis, and
J. Pelocarpus. Occupying congenial pockets were Sabatia difformis, A galinis virgata, Rhexia
virginica, Polygala cruciata, and P. brevifolia, each contributing a speck of color. Jutting
from the stream bed were robust specimens of Xyris smalliana, with their prominent beet-red

S. Conspicuous on extensive carpets of sphagnum mosses were the large goblets of
Sarracenia purpurea, associated with dense ranks of Narthecium americanum and Lophiola
aurea, both gone to seed. Only a few plants of Tofieldia racemosa were seen. Despite

146 BARTONIA

diligent search of this savannah, we did not relocate Rhynchospora oligantha, a rare sedge
first seen here by the leader in 1975. However, we did rediscover two small patches of rare
peat mosses, Sphagnum angustifolium and S. portoricense. On a gentle slope nearby were
Schizaea pusilla, Lycopodiella caroliniana, and the foliage of Platanthera blephariglottis.
In a spung below the site of Martha Furnace, we encountered Rhynchospora gracilenta,
Juncus caesariensis, Scleria reticularis v. pubescens, several flowering stems of Ludwigia
hirtella, and Platanthera clavellata in fruit. A short search in this vicinity failed to turn up
a recently extant population of Ophioglossom vulgatum. Leader: Ted Gordon.

10 September: Chesapeake and Delaware Canal, New Castle Co., DE. After assembling
in Glasgow, we journeyed to an assortment of shallow ponds in dredge spoil on the north
side of the canal. There we observed such species as Elocharis quadrangulata, E. olivacea,
and Schoenoplectus purshianus in well defined wetland zonal communities, along with
Juncus diffusissimus and, on nearby sandy soil, Cyperus grayii, usually found on beach
dunes. We visited a tidal ditch paralleling the south side of the canal. Here we found
Sagittaria calycina and Potamogeton foliosus. Our last stop was a sandy spoil deposit at the
southeast end of the canal, where we observed a small population of Hudsonia tomentosa,
but found that we were too early for Aristida dichotoma var. curtissii. Leaders: Jack Holt
and Janet Ebert.

Bartonia No. 59: 147-151, 1996
1995 Membership

ALDHAM, ALBERT—1660 Hemlock Farms, Hawley, PA 18428, 717-775-6773
AMOos, SANDRA—41 Laurel Rd, Clementon, NJ 08021, 609-346-2242
ANDERSON, KARL H.—Rancocas Nature Center, 794 Rancocas Rd., Mt Holly, NJ 08060, 609-261-2495
TTARDI, VINCENT—PO Box 162, Norwood, NJ 07648, 201-767-2425
~gea Jusa—160 Kendal Dr., Kennett Square, PA 19348, 215-388-8656
UX & RON BEAUCHAMP—634 W. Ellet St., Philadelphia, PA 19119, 215-438-5183
SALAS, Siesta 4265, Elwyn, PA 19063, 610-566-7392
BARR, CAMILLE—Botany Dept., University of Hawaii, 3190 Maile Way, Honolulu, HI 96822
BATH, PRISCILLA—157 Coleman Rd, Trenton, NJ 08690, 609-587-4849
BAUCHSPIES, JAMES T.—1205 Washington St., Easton, PA 18042, 215-253-8498
BEATTY, GEORGE—PO Box 412, Lemont, PA 16851
BESITKA, SISTER MARY ANN—7310 Torresdale Ave., Philadelphia, PA 19136, 215-332-8299
BIEN, WALTER F.—144 Summit Ave., Langhorne, PA 19047, 215-752-3762
BIER, CHARLES W.—372 Kepple Rd., Sarver, PA 16055, 412-281-2777
BIRD, JOSEPH P.—145 Garfield Ave, Trenton, NJ 08609, 609-392-3640
BOWMAN’S HILL WILDFLOWER PRESERVE—PO Box 103, Washington Crossing, PA 18917, 215-862-2924
Boyb, HOWARD—232 Oak Shade St., Tabernacle, NJ 08088, 609-268-1734
BRANDYWINE CONSERVANCY, DAVID HARPER—PO Box 141, Chadds Ford, PA 19317, 610-388-2700
RESLER, CARL—264 W. Wolfert Station Rd., Mickleton, NJ 08056, 619-467-3642
fe RINTON, EDWARD & JOAN—896 Roundelay, West Chester, PA 19382, 215-793-1582
BROADDUS, LYNN—106 Alapocas Dr., Wilmington, DE 19803, 302-651-9598
BROTHERSON, ROBERT—Box 179, Revere, PA 18953, 215-847-5074
BRUEDERLE, LEO P.—Biology, Univ. of Colorado, PO Box 173364, Denver, CO 80217, 303-556-3419
BUCK, WILLIAM R.—New York Botanical Garden, Bronx, NY 10458, 718-817-8624
BURGESON, DENNIS K.—236 Adams Ave., Barrington, NJ 08077
CHEETHAM, ROBERT M.—237 § 2Ist St. #3, Philadelphia, PA 19103
Cortiz, CYNTHIA—32 pagel ne Yardley, NJ 08610
ae JOHN E. & MARILYN—439 Gladstone Ave., aap eine NJ 08033, 609-429-4987
CRICHTON, OLIVER W.—94 clea Ave., Wilmington, DE 19083, 3(2-764-5588
Gusiexe ALLISON W.—Division of Natural Areas ODNR, ae tain Square, Columbus, OH 43224, 614-265-6471
DABYDEEN, SIM MON—Frostberg State University, Frostberg, MD 21532, 301-689-4213
DALGLEISH, DAvE—1900 Sycamore St., Haddon Heights, NJ 08035, 619-547-4823
DANIEL, MARY—HC 1 Box 1117, Blakeslee, PA 18619, 717-646-3326
DARKE, RICK—Longwood Gardens, PO Box 501, Kennett Square, PA 19340, 215-388-6741
DAvis, CHARLES A—1510 Bellona Ave., Lutherville, MD 21093, 410-252-4154
sipbie LINK—14 Cemetery Ln., Schwenksville, PA 19473, 215-287-6635

DICKER, NAOMI D.—309 W. 93rd St. 6A, New York, NY 10025, 212-222-5179
DIEDRICH, ARMON W, JR.—502 Highland Terrace, Pitman, NJ 08071, 609-589-8455
Diorio, JOHN E.—P.O. Box 228, 28 Main St., Heislerville, NJ 08324, 609-785-1643
DOLAN, THOMAS IV—721 Glengarry Rd., Philadelphia, PA 19118, 215-242-6136
DRAUDE, TIMOTHY—415 Poplar St., Lancaster, PA 19103, 717-393-7233

DUNMORE, RALPH—21 Briarwood Drive, Elverson, PA 19520

EBERT, JANET—394 Smith Bridge Rd., Chadds Ford, PA 19317, 215-459-0585

EDINGER, GREG—RR 1, Box 365, Warner Hill Rd., Schoharie, NY 12157, 518-295-7570
ELWELL, CATHARINE R.—36 S 9th St., Allentown, PA 18102

ESHERICK, HELEN K.—2346 Dublin Rd., Orefield, PA 18069, 610-398-0521

EVANDER, EDITH E.—1401A Cuba Rd., Hunt Valley, MD 21032, 410-771-0534

EVANS, JANET—Library, Pa. Horticultural Society, 325 Walnut St. Philadelphia, PA 19106
EVERETT, ALAN C.—304 Runnymede Ave., Jenkintown, PA 19046, 215-887-0347
FARLEY, ELIZABETH B.—319 Bala Ave., Bala Cynwyd, PA 19004, 215-667-0625

147

148 BARTONIA

FELTON, ANNA & PAUL—37 Crawford Rd., Audubon, PA 19403, 215-666-5922

FIELD, STEPHEN R. & THERESA—S Evelyn Ave., Vineland, NJ 08360, Sa 5868

FINE, NORMAN—16 Overhill Rd., East Brunswick, NJ 08816, 908-257-244

FINGERUT, JOYCE—2106 Pennsylvania Ave., Ft. Washington, PA 19034, ne 542-0153

FLANIGAN, TONI ANNE—1238 S. Sheridan St., Philadelphia, PA 19147, 215-755-7375

FOGARASI, KASIA—317 Roxborough Ave., Philadelphia, PA 19128, 215-482-3835

FRANK, SUSAN—2508 Pine St., Philadelphia, PA 19103, 215-732-9280

FREET, FRANCES E.—15 Appalachian Dr., Carlisle, PA 17013, 713-243-8666

FREYBURGER, HELEN R.—5258 34th Ave. North, St. Petersburg, FL 33710, 813-526-1579

FRICK, JULIA W.—Blair 213, 1400 Waverly Rd., Gladwyne, PA 19035, 215-645-8863
ARBACK, MARY E.—3839 Janice St., Philadelphia, PA 19114, 215-332-7105

GEHRIS, CLARENCE W.—Shell Point Village, 1133 Cameo Ct., Fort Myers, FL, 33908, 813-466-6407

GLASS, AMELIA—135 Washington Ave., Pitman, NJ 08071, 609-589-6435

GOFF, ELINOR—791 College Ave., Apt. 1, Haverford, PA 19041, 215-649-2933

GoopD, NORM

periecate THEODORE & PATRICIA—31 Burr’s Bill Rd., Southampton, NJ iri 609-859-3566
GRESHAM, CYANE—498 Siegfriedale Rd., Kutztown, PA 19530, 610-683-145
GYER, JOHN—Box 185, 243 Jessup Mill Rd., Clarksboro, NJ 08020, pvt
HALLIWELL, THOMAS B.—19 Kings Rd., Netcong, NJ 07857, 201-347-6071
HA TY, GAIL B—488 Big Oak Rd., Morrisville, PA 19067, 215-295-4734
HARRIS, JESSIE M.—4401 W St. NW, Washington, DC 20007, 202-338-9083
HART, ROBIN—Natural Res. Dept., PO Box 8, Sarasota, FL 34230, 941-378-6113
HASSELL, LLOYD V.—45 Danbury Rd., Lancaster, PA 17601, 717-569-2368
HAWK, L. JEFFERY—16 Andrea Lane, aay NJ 08619
HECKSCHER, STEVENS—10 Ridley Dr., hen ig PA Age
HEILMAN, LYNN—1004 Cherry Circle, Lansdale, PA 1944
HILL, Roy L.—180 W. Drexel Ave., Lansdowne, PA ioak 215-626-7743
HIRST, FRANKLIN S.—500 Little Mill Rd., Stockton, MD 21864, 410-632-1362
HOLT, ROBERT J.—3032 Taft Rd., Norristown, PA 19401, 610-584-5578
HUNT, LYNN F.—PO Box 553, New Gretna, NJ 08224, 609-296-4922
HUTCHEON, DAviD J.—25 Caledonia Dr., Warminster, PA 18974, 215-957-0976
cr RICHARD D.—403 Georges Rd., Dayton, NJ sep eave}
IRETON, MARY LOU—213 4th Ave., Haddon Heights, NJ
pane CAROL A.—2325 Oakdale Ave., Glenside, PA ieee 215-885-8912
Jess, ROBIN A.—S55 Lahiere i Edison, NJ 08817, 908-572-5928
JOHNSON, ELIZABETH—The Nature Conservancy, 200 Pottersville Rd., Chester, NJ 07979, 908-879-7262
JOHNSON foment K.—3039 ee Dr., Export, PA 15
JOHNSSON, ROBERT G. & FANNY M.—7422 Ridge Rd., Frederick, MD 21701, 301-371-5215
JOHNSTON, KAREN M.—RD 1, Box 488F, Liverpool, PA 17045, AES ie
KAISER, oe Box 222, Sumneytown, PA 18084, 215-234-8
KAPLAN, PAULA WEST—1085 Huntingdon Rd., Abington, PA 19001, Nei 386
KELLER, Buzaser—14 Wyomissing Blvd., Wyomissing, PA 19610, 215-374-3458
K , JA W.—35 Arlington Dr., Pittsford, NY 14534, 716-381-3906
Paces DONALD—100 Three Bcidae Rd., Shamonk, NJ 08088, 609-268-2226
KLOTZ, LARRY H.—Biology Department, Shippensburg University, ‘ghia PA 17257, 717-532-1402
KOERBER, WALTER A. JR—1380 Valley Green Rd., Etters, PA 17319, 717-938-9618
KOLAGA, VALERIE—186 Dilworthtown Rd., West Chase PA 19382, 610-399-3136
KRAFFT, CAIRN—57 Willow Rd., Churchville, PA 18966, 215-322-8553

KRAIMAN, CLAIRE—Flying Hills, 7 Chip Lane, Reading, PA 19607, 215-775-9737
LACE, JANICE—PO Box 1793, Frisco, CO 80443
LADEN, MILTON—William Penn House #1407, 1919 Chestnut St., Philadelphia, PA 19103, 215-568-6599
LAIDIG, KIM—712 Wood Lane, Haddonfield, NJ 08033

ONT, ERIC—717 Sound Shore Rd., Riverhead, NY 11901, 516-722-5542
pie, ee 20 State Rd., Box 537, Paoli, PA 19301, 610-644-1890

THAM, ER—Box 57, Wallingford, PA 19086, 610-565-8979

npr, pguee H.—107 Blackthorn Rd., Wallingford, PA 19086, 610-566-2823
LAUER, DAVID M.—49 Cornell Ave., Churchville, PA 18966

1995 MEMBERSHIP LIST 149

LEAPMAN, HERSH R.—114 Wellington Rd., Lancaster, PA 17603, 717-397-6080

LECK, MARY—Biology Dept., Rider College, 2083 Lawrenceville Rd., Lawrenceville, NJ 08648, 609-896-5092

LEVIN, MICHAEL—414 Mill Rd., Havertown, PA 19083

LIGHTY, RICHARD W.—Mt. Cu he Center, PO Box 3570, Greenville, DE 19807, 302-239-4244

LIVERSIDGE, DAVID—46A Brainerd St., Mt. Holly, NJ 0

LLOYD, LAWRENCE—RD 3084A, Mohnton, PA 19540, 610-777-7702

LOFURNO, MICHAEL J.—2028 Fitzwater St., Philadelphia, PA 19146, 215-732-0849

LOEFFLER, CAROLE C.—Department of Biology, Dickinson College, Carlisle, PA 17013, 717-245-1360

LOGAN, MICHAEL—1027 Morris St., Philadelphia, PA 19148, 215-334-6151

LONKER, MILTON L.—8704 Patton Rd., Wyndmoor, PA 19118, 215-233-4818

LYNCH, DANA M.—1006 Robin Dr., West Chester, PA 19382

MACIARELLO, MICHAEL J.—Dept. of Agriculture and Natural Resources, Delaware State University, Dover, DE
19901, 302-739-5120

MANCYwopbA, GEORGE—609 Hunters Lane, Waterford, NJ 08089, 609-768-0114

MANTELL, SONIA—PO Box 1002, Berwyn, PA 19312, 610-644-5474

MaPEs, CAROL C.—Biology Department, Kutztown University, Kutztown, PA 19530, 215-683-4312

MARTIN, HARRIS W.—229A Presidential Dr., Greenville, DE 19807, 302-424-8444

MAURICE, KEITH R.—186 Main St., Linfield, - sci 215-495-7951

McAvoy, WILLIAM—35 Windflower Dr., Newark, DE 19711, 302-734-3219

McCaABE, MARIANA F.—803 N. Franklin: St, iain PA 19464, 215-326-3082
— ean W.—20 Sylvan Lake Ave., Haddonfield, NJ 08033, 609-429-4628
MCCONVILLE, LYNN—1835 Wynnewood Rd., Philadelphia, PA 19151, 215-877-4311

Scan, see IV—56 W Main St., Suite 303, Christiana, DE 19702, 302-737-9335
MCLAUGHLIN, WILLIAM—PO Box 645, Medford, NJ 08055, 609-268-8010
MCLEAN, WILLIAM & ELIZABETH—139 Cherry Lane, Wynnewood, PA 19096, 610-642-4196
MEADE, JOHN A.—16 Tall Oaks, East Brunswick, NJ 08816, 908-257-8743
MEAGHER, WALTER L.—2 Parkside Way, Robbinsville, NJ 08961, 609-448-9189
MEEHAN, THERESA—Woodcrest Gardens, PO Box 528, Ardmore, PA 19003, 610-649-2849
MEYER, gee C. JR.—5 Railroad Lane, Whitehouse Station, NJ 08889, 908-534-9587
MICKLE, ANN M.—Department of Biology, LaSalle University, Philadelphia, PA 19141, 215-951-1254
MILNER, sna, M.—1131 Harbour Dr., Palmyra, NJ 08065, 609-829-3142
MONTGOMERY, JAMES D.—Ecology III, RD 1, Berwick, PA 18603, 717-542-2191
MOOBERRY, F. M.—106 Spottswood Lane, Kennett Square, PA 19348, 215-444-5495
Moore, GERRY—Department of Biology, Vanderbilt University, Nashville, TN 37235, 615-385-4577
Moore, JULIA E.—Molyneaux Rd., Camden, ME, 04843, 207-231-8338
MORGANSTERN, a River Rd., Asbury, NJ 08802
Morse, LARRY E.—The Nature Conservancy, 1815 N. Lynn St., Arlington, VA 22209, 703-841-5361
Moss, MIRIAM hoor Brookside Rd., Elkins Park, PA 19117, 215-635-0176
NACZI, ROBERT F. C.—Dept. of Biological Sciences, Northern Kentucky University, Highland Heights, KY 41099,
606-572-6929
NEWSTEAD, EDWIN & CHARLOTTE—270 Roseland Ave., Essex Fells, NJ 07021, 201-226-7651
NICHOLS, HORATIO R—412 Federal City Rd., Pennington, NJ 08543, 609-737-7442
NOBLE, WILFRED R—PO Box 4331, Philadelphia, PA ~ 215-233-0733
NYSTEDT, JOHN—PO Box 1153, Haddonfield, NJ 080.
O’BRIEN, MICHAEL—22 Richards Ave., Pine Hill, NJ ado 609-783-1103
O’DELL, GERALD—892 E. Schuylkill Rd., Pottstown, PA 19464, 215-323-3971
OLSON, WILLIAM F.—2 Main St., Apt. B, Farmingdale, NJ 07727, 908-938-3187
PARKS, JAMES C.—865 Letort Rd., Washington Boro, PA 17582, 717-872-5206
N, W. R.—20 State Road, Box 537, Paoli, PA 19301, 610-644-1890

PINGEL, PATRICIA A.—650 Freemansville Rd., Shillington, PA 19607

YLER, DOROTHY D.—18 Bridle hee Chadds Ford, PA 19317, 215-459-3969
RADIS, RICHARD P.—69 Ogden Ave., Rockaway, NJ 07866, 201-586-0
RAYSER, JEANNE—2 Sawmill Rd., "Modio NJ 08055, 609-983-1617
tay ANN F.—3 Blythewood Rd., Doylestown, PA 18901, 215-348-8139

IDES, CHARLES A. JR.—107 Stony Creek Ave., Lansdale, PA 19446, 215-368-9591

RICKENBACK, PENNY—PO Box 181, Oley, PA 19547, 610-987-6585
RIGG, ELIZABETH—655 Caley Rd., King of Prussia, PA 19046, 610-265-1184

150 BARTONIA

RISKA, MICHAEL E.—Delaware Nature Society, Box 700, Hockession, DE 19707, 302-239-2334
ROBACK, HELEN M.—188 Redwood Rd., King of Prussia, PA 19406, 215-265-1158
ROWAN, JANE O.—120 Governors Dr., Wallingford, PA 19086, 610-490-1080
RupyJ, Eric §S.—915 Olive Branch Ct., Egdgewood, MD 21040, 410-676-5782
RUSSELL, EMILY W. B.—Box 430, Mt. Tabor, NJ 07878, 201-625-3382
SALGANICOFF, MATILDE—556 N 23rd St., Philadelphia, PA 19130, 215-751-0396
SCHAEFFER, ROBERT L. JR—1940 Turner St., Apt. 105, Deon PA 18104
ScHoss, JAY—545 N. Edgemere Dr., West Allenhurst, NJ 077
SCHNEIDER, GEROGE—345 Nursery Rd., a we PA tae, 717-292-4035
a WILLIAM L.—228 Canterbury Dr., West Chester, PA 19380, 610-431-2449
Sc , W. JUERGEN—1 Stream Valley Ct., Laytonsville, MD 20882
seaceuilk ALFRED E.—Academy of Natural Sciences, 1900 Benjamin Franklin Pkwy., Philadelphia, PA 19103,
215-299-1193
SCHWEITZER, DALE F.—66E Main St., Box 30B, Port Norris, NJ 08349
ScoTT, CONNIE—139 Myrtle Ave., Havertown, PA 19083, 610-446-6259
Scott, JOHN D.—RD 1, Box 249D, Hertzog School Rd., Mertztown, PA 19539, 215-682-2809
SEAGER, KEITH A.—278 Fishing a Rd., Cape May, NJ 08204, 609-884-8778
SEIPLER, MARY JANE—4200 Tamarack Dr., Murrysville, PA 15668
SETTLEM KENNETH T.—219 Maple i, Jersey Shore, PA 17740, 717-398-2546
—218 Cummings Ave., Elberon, NJ 07740, 908-728-1989
ite WILLIAM—512 Red Bluff Ct., Millersville, MD 21108
G S.—728 Seymour Rd., Bear, DE 19701, 302-324-9468
eri DAviD—12 Chesterfield-Georgetown Rd., Trenton, NJ 08620, 609-298-1555
STAILEY, HELEN M.—8701 Macon St., Philadelphia, PA 19152, 215-673-8163
STALTER, RICHARD—St. John’s University, Grand Central & Utopia Pkwy., Jamaica, NY 11439, 718-990-5237
ae Sdne Si Maltbie Ave, Apt. 19B, Midland Park, NJ 07432, 201-612-9069
TEVENS, RLES E.—615 Preston Place, ae VA 22903, 804-293-8658
micnloade quis Ravenwood Rd., Exton, PA 19341, 610-524-2373
STUCKEY, RONALD L.—1315 Kinnear Rd., Calais OH 43212, 614-292-6095
SWEETMAN, HAROLD—Jenkins Arboretum, 631 Berwyn Baptist Rd., Devon, PA 19333, 215-647-8870
TUCKER, ARTHUR O.—Delaware State College, Dover, DE 19901, 302- 739-5120
UDELL, VAL—2746 Yost Rd., Perkiomenville, PA 18074, 610-754-7163
VERLENDEN, DONALD—PO Bx 116, Landsdowne, PA 19050, 610-622-0227
VIRKLER, ROBERT J.—429 Elliger Ave., Ft. Washington, PA 19034
VISION, TODD—EEB, Princeton University, Princeton, NJ 08544, 609-258-1656
VOLLMER, JOHN—42 Burrs Mill Rd., Southampton, NJ 08088, 609-859-2805
WALKOVIC, JOANNE H—539 Woodland Ave., Media, PA 19063, 610-566-0861
WALLS, JERRY G.—657 Second St., Trenton, NJ 08611
WEINER, JACOB—Department of Biology , Swarthmore College, Swarthmore, PA 19081, 215-328-8038
WENGER, prctohentey L.—1091 Street Rd, Oxford, PA 19363, 215-932-5361
WHEELER, STEPHEN A.—312 W. 6th St, Boyertown, PA 19512, 215-369-3768
WIEBOLDT, shies F._—2488 Crab Creek Rd., Christiansburg, VA 24073, 540-382-9492
WILDMAN, HOWARD—PO Box 381, New Lisbon, NJ 08064, 609-894-4870
WILEN, RALPH & ELLEN—143 Ridge Rd., Southampton, NJ 08088
WILLIAMS, CARL—165 W. Ridge Pike, Ulnveritk: PA 19468, 610-489-6331
WILLIAMS, DAviD L.—641 Coppermine Rd., Princeton, NJ 08540, 201-297-0642
WILLIG, SARAH ANDERSEN—26 Sycamore Lane, Phoenixville, PA 19460, 215-933-3539
ILSON, RONALD—3740 Ridge Rd., Snow Hill, MD 21863, 410-632-3892
WINDISCH, ANDREW & MARTHA—PO Box 312, Chatsworth, NJ 08019, 609-726-9054
WITMAN, DEANNE M.—PO Box 294, Lyons Station, PA 19536, 610-682-7967
WoLrFF, EMILY T.—295 E. Rose Tree Rd., Media, PA 19063, 215-566-4907
WOLFF, JOHN—2640 Breezewood Dr., Lancaster, PA 17601, 717-569-6955
Woop, HOWARD P.—3300 Darby Rd. C-802, Haverford, PA 19041, 215-642-9963
ZIMMERMAN, L. WILBUR—922 Montgomery Ave. K2, Bryn Mawr, PA 19010, 610-527-3340
ZOLLER, HEINRICH, 50 Renninger Rd., Bechtelsville, PA 19505

1995 MEMBERSHIP LIST 151
Honorary Members

BOYLE, E. MARIE—Unitarian-Universalist Home, 224 W. Tulpehocken St., Philadelphia, PA 19144
EWAN, JOSEPH & NESTA—Missouri Botanical Garden, PO Box 299, St. Louis, MO 63166, 314-577-9505
McGraATH, JAMES K.—304 Derwyn Rd., Lansdowne, PA 19050, 215-284-2594

NEWBOLD, ANN—S0 Renninger Rd., Bechtelsville, PA 19505, 215-754-7573

SHAEFER, CHARLOTTE—Bellingham, Apt. B10, 1615 E. Boot Rd., West Chester, PA 19380, 610-918-2246
WOODFORD, ELIZABETH—6 Sawmill Rd., Medford, NJ 08055

Life Members

ARSENAULT, JOSEPH—961 Clark Ave., Franklinville, NJ 08322, 609-697-2459

GREENLAND, CHRISTINE M.—790 E. Street Rd., Warminster, PA 18974, 215-322-4105

PATRICK, RUTH—PO Box 4095, Philadelphia, PA 19118

ROBERTS, WILLIAM H.—1922 Rittenhouse Sq., Philadelphia, PA 19146, 215-569-5632

RYAN, NANCY PETERS—419 S. Carlisle St., Philadelphia, PA 19146, 215-735-6189

THOMPSON, SUE—Carnegie Museum of Natural History, 4400 Forbes Ave., Pittsburgh, PA 15213, 412-622-3295

EDITOR’S NOTE

Biographical Accounts of Hans Wilkens and Grace M. Tees will be published in Bartonia
No. 60. Friends and colleagues of Hans and Grace are encouraged to provide information
to the editor concerning things they remember about them. As many know, both made
Substantial contributions to support botany at the Academy of Natural Sciences and were
longtime members of the Philadelphia Botanical Club.

% oj enonwdit

isinaniee

Contents (cont'd)

NIN i ois 9 ing See A gr Ae se 2 ee ee 129
OO ca ise es ka Tp Re eas ween nt Coe eee ee ee ee 134
Pe MOUND oe ye lig ea ee ee ee 135
SR OWN AVM oe Sc ee ee ee ee 138
vefithashiny Od, , Ct re eR a Na eR ce ce isin 143
premmersnes Liat os oh oe os ee 147
RO ON i ee ee ee 151

Program of Meetings 1993 and 1994

Date Subject Speaker
1993
28 Jan Toluene Removal from Indoor Air by Dieffenbachia .................+4-. John R. Porter
28 Feb Changes in the Flora of Northeastern Bucks County, 1890-1990 ............ n F. Rhoads
25 Mar Plowers of the Swit Alps. 25.5 ee Heinrich Zoller
Ann Newbold

22 Apr Recent Rare Plant Surveys in Pennsylvania .............-....+-eeeeees John Kunsman
27 May The Foliage is the Fruit Hypothesis and the Sexual Specialization of the Female

InGlosescence of Dulinlograts 3 ks ea iw ye James A. Quinn
23 Sep Slide Ilustrated Report of Summer Botanizing .................2-00-5- Club Members
28 Oct Netonad History of Water Plans 2. io a a Alfred E. Schuyler
18 Noy Native Azaleas and Rhododendrons of the Eastern United States................ Jim Plyler
16 Dec Woodland Gardens Close Up: Pollination Mechanisms and Seed Production

John and Janet Gyer

1994
24 Feb A Visit to Lake Baikal in Southern _ a ee Ann Rho
24 Mar Alpine Wildflowers of Colorado and Wyoming ..........----5.++++005- Sioux Baldwin
28 Apr Botanical Diversity in the Moist —_ a ple Hite, Be oct. Wayt Thomas
26 May Walnford: An Historic Landscape Restoration ........-...0.-02-00005 Elizabeth McLean
22 Sep Slide Illustrated Report of Summer heeding ee ee aks Club Members
27 Oct Wetlands and Polllies Charles A. Rhodes
17 Nov Academy’s Treasury of Botanical Literature .......-..-- 220s essere Elizabeth McLean
Alfred E. Schuyler
Howard P. Wood
15 Dec Botanical Exploration of the Mekong and Yangtzi River Divide in Northwest

Viens: China oo ea es ee ee Harold Sweetman