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PHILADELPHIA BOTANICAL CLUB

he Philadelphia Botanical Club

Bartonia No. 61: 1-13, 2002

Structure and Composition of a
Northern Hardwood Forest Exhibiting Regeneration Failure

iology Department, University of Scranton, Scranton, PA 1851
townsendd 1@UofS.edu

DANIEL S. TOWNSEND, JENNIFER S. SEVA', CAROL HEE-SEAGLE? AND GLEN MAYERS?
)

ABSTRACT. We surveyed trees (dbh >10.0 cm), saplings (>140 cm tall to dbh <10.0 cm),
seedlings/sprouts (11 to 140 cm tall) and shrubs of a hemlock-mixed northern hardwoods forest at
Lacawac Sanctuary in northeastern Pennsylvania. The canopy (24 species, 490.5 stems ha”) was
dominated by eastern hemlock (Tsuga canadensis) with red maple (Acer rubrum) and red oak (Quercus
rubra) as subdominants. Other canopy trees, in descending order of importance, were chestnut oak
(Quercus prinus), American beech (Fagus grandifolia), sugar maple (Acer saccharum), white pine (Pinus
strobus) and sweet birch (Betula lenta). The sapling stratum (22 species, 239.5 stems ha") was dominated
by T. canadensis with F. grandifolia, A. rubrum and A. saccharum as subdominants. Oak saplings were
rare (Q. rubra) or absent (Q. prinus). Sapling density was only 49% that of canopy density. Small
saplings (0.1 to 2.5 cm dbh) were very sparse (14.0 stems ha’) and 82% were T. canadensis or F.
grandifolia. The seedling/sprout layer (23 species, 444 stems ha’) was dominated by F. grandifolia (236
stems ha"), which occurred solely as root suckers. Only 6% of seedlings/sprouts (24.7 stems ha’) were
>30 cm tall, the majority of which were F. grandifolia (18.6 stems ha') and T. canadensis (4.8 stems
ha"). Hence, the Lacawac Sanctuary forest was almost devoid of woody stems from 31 cm tall to 2.5
cm dbh, exhibiting much lower densities than most other eastern U.S. forests, either second-growth
or old-growth, that we surveyed. Only forests with very high densities of white-tailed deer (Odocoileus
virginianus) had comparably low seedling and sapling densities.

INTRODUCTION

Lacawac Sanctuary is a 202-ha forested nature preserve located on the Pocono Plateau
(Allegheny Plateaus Province) in northeastern Pennsylvania. Although it lies within Braun’s
(1950) hemlock-white pine-northern hardwood forest region, Lacawac’s location is quite close
(ca. 15 km east) to the border between the northern hardwood and Appalachian oak forest
regions (Kuchler 1964; Rhoads and Klein 1993). Russell et al. (1993) viewed Lacawac as
occupying the transition zone between the two forest biomes, but assigned it to the
Appalachian oak forest based on the fairly high percentage of oak (Quercus) pollen in a
sediment core from Lake Lacawac. We conducted this study, in part, to address the question
of Lacawac Sanctuary’s forest type affiliation by quantitatively documenting its current
structure and composition.

Dramatic changes occurred in eastern North American forests subsequent to European
settlement, principally as a consequence of human activities (W hitney 1994). Timbering and
clearing of land for agriculture resulted in widespread destruction of eastern forests during

, rey a 15.1 Denver Road, Denver, PA 17517; "Department of Marine Science, University of North
Carolina at Chapel Hill, Chapel Hill, NC 27599; 374 Indian Rocks, Lake Ariel, PA 18436

Manuscript submitted 8 January 2001, revised 20 April 2001. MISSOURI BOTANICAL
JUL 0 8 2002
GARDEN LIBRARY

2 BARTONIA

the eighteenth and nineteenth centuries (Whitney 1994). Land use changes during the
twentieth century have allowed forests to recover in many areas, including Pennsylvania,
although forest composition and structure have changed considerably from pre-settlement
forests (Whitney 1994). Two changes in eastern North American forests that have occurred
during the twentieth century include an apparent decline in oak dominance (Abrams 1992)
and the increased importance of red maple (Acer rubrum; Abrams 1998). Oak decline is
attributed to widespread ion failure arising from a combination of factors, including
understory competition from late successional, shade-tolerant species, selective logging, and
long-term fire suppression (Abrams 1992). The increasing importance of red maple seems to
arise from its adaptability to a range of light and edaphic conditions, combined with
widespread fire suppression (Lorimer 1984; Abrams 1998).

Browsing by overabundant white-tailed deer (Odocoileus virginianus Zimmerman) is
another factor that has dramatically altered eastern forest structure and composition in the
twentieth century (Whitney 1994). Deer at densities greater than 8 to 10 deer km” can
significantly affect forest community structure (Whitney 1984, 1990; Frelich and Lorimer
1985; Tilghman 1989; Anderson and Katz 1993; Rooney and Dress 1997), plant population
ecology (Anderson 1994; Balgooyen and Waller 1995; Rooney 1997) and other wildlife
(DeCalesta 1994). Excessive browsing can significantly reduce or eliminate regeneration of
woody species (Anderson and Loucks 1979; Alverson and Waller 1997; Healy 1997; Rooney
and Waller 1998), reduce density and reproductive success of shrubs and herbs (Allison
1990a, 1990b, 1992; Anderson 1994; Rooney 1997) and lower the diversity of forest herbs
(Rooney and Dress 1997).

At Lacawac Sanctuary, anecdotal observations of high deer abundance and browsing
damage to woody and herbaceous vegetation date from the early 1970s (Arthur Watres, pers.
comm.). Low levels of deer hunting occurred on the sanctuary until 1984, when hunting was
banned. When we began this study in 1991, an obvious browse line at 1.5 to 1.6 mand a
dearth of understory plants were evident throughout the sanctuary. Annual deer hunts,
reinitiated at Lacawac in 1994, have harvested an average of 22 deer yr“. In a 2000 census
at the sanctuary, we estimated deer density at 19 to 29 deer km~ (Townsend, unpubl.). It
is therefore likely that deer densities prior to the study on which we report here were at
least as high. Our second objective in this study was to quantitatively document the
condition of the woody understory in the sanctuary forest.

FIELD SITE DESCRIPTION

Lacawac Sanctuary (41°23’ N, 75°17’ W) is a 202-ha nature preserve located at the
northern edge of the Pocono Plateau, in Paupack Township, Wayne County, Pennsylvania.
The Pocono Plateau is a moderately rolling elevated tableland that lies in the eastern
glaciated region of the Allegheny Plateaus Province. The Pocono Plateau forest represents
the southeastern-most extent of the hemlock-white pine-northern hardwoods region, and is
bounded to the west, south and east by Appalachian oak forest (Braun 1950; Kuchler 1964;
Rhoads and Klein 1993).

The climate of Wayne County is typified by cold winters and moderately warm summers,
with temperatures ranging from a mean daily minimum of -11.4° C in January-February to
mean daily maxima of 26.7° C and 25.9° C in July and August, respectively. Average daily
temperatures range from -5.1° C in January to 19.9° C in July (Martin 1985). Mean annual
Precipitation is 98.2 cm, about 55% of which falls during the growing season of April

REGENERATION FAILURE IN A HARDWOOD FOREST 3

through September. Mean annual snowfall is 145.3 cm; on average, 52 days a year have at
least 2.5 cm of snow on the ground (Martin 1985).

The sanctuary, situated on the north shore of Lake Wallenpaupack (a man-made
hydropower reservoir), includes a 20.2-ha glacial lake, Lake Lacawac, and virtually its entire
watershed. Sanctuary elevations range from 366 m to 460 m. Most of the sanctuary has
moderately rolling topography, with steeper slopes and bedrock outcrops occurring as ledges
that drop more than 60 m to the shore of Lake Wallenpaupack in the southern part of the
sanctuary. The bedrock of the sanctuary area is red Catskill sandstone (Mowbray 1971).
Upland soils are derived mainly from glacial till, and are comprised of extremely stony loams
of the Lordstown, Mardin, and Oquaga Series, and moderately well drained and somewhat
poorly drained extremely stony loams of the Wellsboro Series (Martin 1985). Soil depths
range from very shallow (in the ledges area) to greater than 1.5 m in some portions of the
sanctuary (Martin 1985). Lacawac Sanctuary is more than 95% forested.

METHODS

We surveyed four areas of upland forest on the sanctuary, covering approximately 100 ha,
during June-August 1991-92. Based on sanctuary historical records, the time since last major
timbering in the four stands ranged from 65 to 110 years.

We used a point-centered quarter method to sample canopy trees (Cottam and Curtis
1956; Brower et al. 1990), which we defined as those with dbh (diameter at breast height =
diameter at 1.37 m) >10 cm. Sampling points were spaced at random distances between 25
and 50 m along parallel transects spaced at random intervals of 25 to 50 m. Any canopy tree
with multiple trunks at breast height was counted as a single individual and represented by
the sum of the basal areas of all living trunks.

Saplings, seedlings, sprouts, shrubs and lianas were sampled in one 10-m x 10-m plot at
each point. Plots were placed at random in one of the four quarters at each point, with a
plot corner at the point and two sides extending out along the cardinal compass points that
defined the quarter. Saplings were defined as stems > 1.4 m tall and O.1cm < dbh < 10.0
cm. Seedlings and sprouts were defined as stems 11 to 140 cm tall; we did not systematically
record seedlings or sprouts $10 cm tall. We examined each seedling-sized plant at ground
level to determine whether it was a true seedling or a sprout. The vast majority of sprouts
that we recorded (98.3%) were root suckers of Fagus grandifolia; a few others were basal
sprouts of other species. In each plot, we identified and counted the number of saplings,
seedlings, sprouts, shrubs and lianas; and we measured the dbh of each sapling, and the
height of each seedling, sprout or shrub. si

We indexed browsing damage to the root suckers of F. grandifolia under four categories
(unbrowsed, light, moderate, heavy) based on evidence of deer bites that removed live
terminals or branches (we did not include shoots that had been browsed in previous years
or that had not produced new growth). “Unbrowsed” suckers were those that lacked any
evidence of browsing on the terminal or any branch. “Light” browsing was recorded for any
sucker with at least one, but no more than 50%, of its terminal and branches removed.
“Moderate” browsing was recorded for suckers with more than 50% but not all of their
shoots removed. “Heavy” browsing was recorded when all live shoots on a root sucker had
been bitten off.

For each canopy tree and sa
frequency, relative frequency (RF), basal area,

pling species, we calculated density, relative density (RD),
relative basal area (RA), and importance value

+ BARTONIA

(IV = (RD + RF + RA)/3) (rower et al. 1990). For seedlings and sprouts, shrubs, and
lianas, we computed density, RD and RF, and IV = (RD + RF)/2) (Brower et al. 1990). We
computed summary statistics for all variables using the values from each of the four stands
as independent data points.

For each forest stratum in each of the four stands, we computed species richness,
Simpson’s diversity index, D,, and Simpson’s coefficient of evenness, E, (Simpson 1949,
Brower et al. 1990). We then compared means for each variable among strata using one-way
ANOVA (Zar 1999). We computed Morisita’s index of community similarity ([,,) to make
pairwise comparisons of the composition of canopy, sapling, and seedling/sprout strata
(Horn 1966, Brower et al. 1990). Morisita’s index ranges from zero when communities have
no overlap to one when they are identical. All nomenclature follows Gleason and Cronquist
(1991).

We compared Lacawac’s forest structure and composition to several published studies in
eastern U.S. forests. We developed an initial list of studies in which measures of composition
and structure were given for all species in the canopy, sapling and seedling strata. We
selected studies from the list so as to include both old-growth and second-growth forests that
were representative of several eastern deciduous forest regions (Braun 1950), including mixed
mesophytic, oak-hickory, oak-chestnut, beech-maple, and northern hardwood forests. We
also included studies that covered more than one site, especially when sites differed in deer
density and consequent browsing pressure.

RESULTS

We sampled 392 points and associated plots (total plot area sampled = 3.92 ha), and
recorded 43 taxa of canopy trees, saplings, seedlings/sprouts, shrubs, and lianas. We recorded
1,568 canopy trees (dbh >10 cm) comprising 24 species, yielding a density of 490.5 stems
ha” and a total basal area of 34.7 m? ha" (Table 1). Canopy trees had a mean dbh of 30.0 cm
and a mean basal area of 708 cm? tree",

Tsuga canadensis was the dominant canopy tree, with an IV almost twice that of either
Acer rubrum or Quercus rubra, the two subdominant trees (Table 1). These three species
accounted for 65.6% of total IV. Ace? rubrum had higher density and frequency but lower
basal area than Q. rubra (Table 1), a result of Q. rubra stems being less common but much
larger (mean basal area = 1,147 cm? tree) than A. rubrum (mean basal area = 509 cm? tree’).
Quercus prinus, Fagus grandifolia and Acer saccharum constituted a third group of important
canopy trees (Table 1). The three oak species we observed, Q. rubra, Q. prinus, and Q. alba
(white oak), together comprised 22.2% of canopy density and 26.4% of canopy IV.

One hundred ninety single-trunked canopy trees had dbh >40 cm, including 7. canadensis
(36% of total), Q rubra (30%), Q. prinus (11%), A. saccharum (8%), A. rubrum (4%) and P.
strobus (3%). The largest individual trees that we recorded included P. strobus (dbh = 81cm),
Q. rubra (77.5 cm) and Liriodendron tulipifera (tuliptree; 72.1 cm).

We recorded 947 saplings comprising 22 species, yielding a mean density among the four
stands of 239.5 stems ha! and a mean basal area of 3.5 m? ha! (Table 2). No saplings
occurred in 94 (24%) of the plots we sampled. The sapling layer was dominated by T.
ca is, with a mean IV (37.6%) greater than that of the three subdominant species (F.
grandifolia, A. rubrum and A. saccharum) combined (Table 2). In contrast to T. canadensis and
Acer Spp., the vast majority (>99%) of F grandifolia saplings were root sprouts. Quercus was
effectively absent from the sapling stratum; we recorded only 10 Q. rubra saplings (2.8 stems

REGENERATION FAILURE IN A HARDWOOD FOREST

5

TABLE 1. Canopy tree density (stems ha’), basal area (m* ha’), relative density (RD), relative frequency
anctuary in 1991-1992; IV is the

yond relative basal area (RA), and i importance value (IV) at Lacawac S$

of RD, RF, and RA. Data are given as means (SEM).

Species Density Basal Area RD RF RA IV
Tsuga canadensis (L.) Carriere 157.2 (30.2) 10.79 (2.05) 32.8 (6.4) 26.5 (4.1) 31.1 (5.9) 30.1(5.4)
Acer rubrum L. 0(22.7) 4.69 (1.23) 19.7(54) 19.9 (4.5) 13.8(3.9) 17.8(4.6)
Quercus rubra L. 74.4 (31.4) 8.26 (2.27) 14.3. (5.1) 14.8 (3.5) 24.067) 17,7(6.1)
Quercus prinus L. 39.5 (12.6) 3.19 (1.02) 7.7(2.4) 85(2.6) 9.0(2.8) 8.4(26)
Fagus grandifolia Ehrhart 31.8 (5.1) 1.65(0.34) 64(06) 7.7(08) 4.7(1.0) 63(08)
Acer saccharum Marshall 29.2 (10.6) —:1.75 (0.82) 5.8 (2.2) 5.9 (2.0) 4.9 (2.3) 5.6(2.1)
Pinus strobus L. 16.6 (6.6) 1.11 0.45) -3.4(1.3) 3.8 (1.5) 3.3 (1.4) 3.5,(1.3)
Betula lenta L 16.1 (6.0) 0.85(0.37) 3.6 (1.6) 43(18) 2.5(1-1) 3.5(1.5)
Fraxinus americana L. 9.8 (4.4) 0.59 (0.31) 2.0 (0.9) 2.7 (1.2) 1.6 (0.9) — 2.1(1.0)
Carya spp 4.8 (1.7) 0.42 (0.19) 0.9 (0.3) 1.3 (0.4) 1.2 (0.5) 1.10.4)
Tilia americana L. 3.7 (3.3) 0.36 (0.30) 0.8(0.7) 1.0(09) 1.0(0.8) 0.9/0.8)
Betula alleghaniensis Britton 8 (1.2 0.19 (0.12) 0.6 (0.3) 0.8 (0.4) 0.6 (0.4) 0.70.3)
Ostrya virginiana (Miller) K. Koch 2.3 (1. 9 0.05 (0.04) 0.5 (0.3) 0.7 (0.4) 0.10.1) 0.4(0.2)
Prunus serotina Ehrhart 1.8 (0.8 0.08 (0.04) 0.4 (0.2) 0.5 (0.2) 0.2 (0.1) 0.40.2)
Liriodendron tulipfera L. 17 (11) 0.28016) 03(0.2) 0.4(0.3) 0.8(0.5) 0.5(03)
Other species* 4.8 2 1.2 1.0
Totals 490.5 34.7
*Includes Carya glabra (Miller) Sweet, C. ovata (Miller) K. Koch and C. tomentosa (Poiret) N
*Other species (IV) include Quercus alba L. (0.3), Nyssa sylvatica Marshall (0.2), Populus randidentate Mi chx. (0.2),

Pinus rigida Mill. (0.1), Carpinus caroliniana Walter (0.1), Picea glauca (Moench) Voss (<0.1), Pinus resinosa Aiton
(<0.1).

TABLE 2. Sapling* density (stems ha’), relative density (RD), relative frequency (RF), relative basal area
(RA), and i importance value (IV) at Lacawac Sanctuary in 1991-1992; IV is the mean of RD, RF, a
RA. Data are given as means (SEM).

Species Density RD RF RA IV
Tsuga canadensis (L.) Carriere 86.5 (14.8) 41.8 (10.6) 33.1(7.9) 45.6 (11.4) 40.2 (10.0)
Fagus grandifolia Ehrhart 40.4 (8.3) 16.8 (1.8) 15.7 (1.2) 13.2 (0.5) 15.2 (1.0)
Acer rubrum L 36.4 (25.1) 11.7 (6.3) 13.9(4.9) 14.9(6.7) 13.5 (5.9)
Acer saccharum M. 22.6 (8.7) 8.6 (2.9) 10.1 G4 8.1 ey) 8.9 e7
a virginiana (Miller) K. Koch 11.7 (4.0) 47(1.5) 5.9(23) 4.9(15) 5.2 (1.

Pk strobus L. pete 10.6 (9.2) 3.2 (2.4) 3.5(2.1) 2.9(25) 3.2 (2.3)

spp.** 6.1 (1.3) 3.1(0.9) 4.8 (1.3) 3.2 (1.2) 3.6 (1.1)
Hamamelis virginiana L 6.1 (5.1) 18 (14) 2.2(16) 07(0.4) 15 (1.1)
Amelanchier spp.*** 5.2 (2.5) 2.1(1.0) 24(09) 1406) 2.008)
Ilex verticillata (L.) A. Gray 3.1 (3.1) 1.9 (1.9) 1.4 (1.4) 0.6 (0.6) 1.3 (1.3)
Nyssa sylvatica Marshall 2.9 (1.7) 1207) 16(1.3) 14(09) 1.4 (1.0)

Valine Ti 2.8 (1.8) 1.0(0.5) 18(0.9) 13(08) 1.3(0.7)
Fraxinus americana L. 1.9 (0.7) 0.8(0.3) 1.305) 07(03) 09(04)
Other species* 32 3 1.1 1.8
Total Density 239.5

*Saplings were defined as stems >1.4 m tall (with 2 minimum dbh = 0.1 cm) and a dbh <10.0 cm.
**Includes Betula lenta L. and B. alleghaniensis Britton.
***Includes Amelanchier arborea (Michx. f.) Fern. and A. laevis Weig.
*Other species (IV) include Carya ovata (Miller) K. Koch (0.6), Carpinus caroliniana Walter (0.3), Abies balsamea
(L.) Miller (0.2), Tilia americana L. (0.2), Acer saccharinum L. (0.1), Crataegus sp. (0.1), and Pinus resinosa Aiton (0.1).

BARTONIA

[__] American beech
GI other species Figure 1. Density of seedlings and sprouts at
Lacawac Sanctuary in 1991-92. Open bars
represent the root sprouts of American beech
(Fagus grandifolia Ehrhart); hatched bars repre-
sent the seedlings and sprouts of all other
species.

Density (stems ha™')
8

11-20 21-30 31-40 41-140
Seedling/sprout height intervals (cm)

ha, 1.3% of sapling IV) and no saplings of Q. prinus or Q. alba (Table 2).

The density of saplings was only 49% of that of canopy trees (Tables 1, 2). In fact, the
ratio of sapling density to canopy tree density was <1.0 for every species except F.
grandifolia (1.27), including T. canadensis (0.55), A. rubrum (0.39), Q. rubra (0.04), Q. prinus
(0), A. saccharum (0.77), Betula spp. (0.32), P. strobus (0.64), and Fraxinus americana (white
ash; 0.19). The dearth of saplings was especially acute for the smallest size interval (dbh =
0.1 to 2.5 cm), in which we recorded only 57 stems (14.0 stems ha"), 82% of which were T.
canadensis or F. grandifolia (Table 3).

We recorded 1,600 seedlings and sprouts (11 to 140 cm tall), comprising 23 species with
a mean seedling/sprout density of 444 stems ha" (Table 4). Fagus grandifolia dominated this
stratum, with 42% of seedling/sprout IV; however, it occurred only as root sprouts, never
as true seedlings or basal stem sprouts. Among the seedlings/sprouts of other species (” =
743; 190 stems ha"), Q. rubra and T. canadensis were most important, while A. rubrum,
Amelanchier spp. (serviceberry) and P. strobus comprised a third tier (Table 4). In contrast
to F. grandifolia, most seedling-sized plants of other species were true seedlings; basal stem
sprouts were rare (n = 21) and there were no root sprouts.

Most F. grandifolia root sprouts (91.5%) were <30 cm tall (Figure 1) and showed evidence
of extensive and repeated browsing. Of 857 beech root sprouts that we recorded, none were
unbrowsed, and only a few (2%) showed light browsing. The vast majority were moderately
(34%) or heavily (64%) browsed. Moderately to heavily browsed sprouts typically occurred
as low, shrubby plants with thickened stems bearing numerous short branches (usually less
than 10 cm) and a few small leaves.

For canopy species other than beech, the great majority of seedlings and sprouts (96.8%)
were <30 cm tall (Figure 1); we recorded only 24 true seedlings that were >30 cm tall and
79% of those were T. canadensis. Although Q. rubra seedlings were fairly abundant (18.3%
of seedling/sprout density), only 5 seedlings were 21 to 30 cm and none were >30 cm. We
recorded only 10 Q. prinus seedlings/sprouts (four were basal stem sprouts); all were 11 to
20 cm. Seedlings and sprouts of understory tree and tall shrub species, including Hamamelis
virginiana (witch-hazel), Ilex verticillata (winterberry), Rhododendron maximum (white-
laurel), Vitis labrusca (fox grape), Kalmia latifolia (mountain-laurel), and Vaccinium
corymbosum (highbush blueberry) were rare or absent (Table 4), and always <20 cm tall
when present.

Because the heavy shade of T. canadensis may often result in low regeneration, we tested
the hypothesis that seedling density varied with hemlock density among the four stands
using the Pearson product-moment correlation coefficient (Zar 1999). The density of true
seedlings (excluding sprouts) ranged from 79 to 301 seedlings ha? among the four forest
stands, while density of T: canadensis (canopy trees and saplings combined) ranged from 154
to 348 trees ha’. The correlation (7, = -0.686, 2 = 4, P >0.10) was not significant, although

REGENERATION FAILURE IN A HARDWOOD FOREST 7

TABLE 3. Sapling size distribution at Lacawac Sanctuary i in 1991-1992. n = number of stems recorded
in 392 100m? plots. Dashes indicate that a species did not occur in a size class.

Dbh Intervals (cm)

Species n 0.1-2.5 2.6-5.0 5:1-7:5 7.6-10.0
Tsuga canadensis (L.) Carriere 347 21 73 93 160
Acer rubrum 137 — 28 50 59
Fagus grandifolia Ehrhart 165 26 39 61 39
Acer saccharum Marshall 97 4 36 25 32
Ostrya el oe (Miller) K. Koch 49 : 14 15 18
Pinus strobus L 38 = 13 11 14
Betula spp.* 22 1 7 5 9
Hamamelis os de. 23 1 20 - 2
Amelanchier spp.** 19 - i 4 4
Quercus rubra L. 10 _ 1 2 7
Nyssa sylvatica Marshall 10 — 2 5 3
Fraxinus americana L 8 1 3 2 2
Other species* 23 1 13 6 3
Density (stems ha’) 239.5 14.0 65.8 70.6 89.1

*Includes Betula alleghaniensis Britton and B. lenta

**Includes Amelanchier arborea (Michx. f.) Fern. ‘ed A. laevis Wieg.

*Includes Abies balsamea (L.) Miller, Acer saccharinum L., Carpinus caroliniana Walter, — ovata (Miller) K
Koch, Crataegus sp., Ilex verticillata (L.) A. Gray, Pinus resinosa Aiton, and Tilia americana L

TABLE 4. Seedling and sprout* density (stems ha”), relative density (RD), relative frequency (RF), and

importance value (IV) at Lacawac Sanctuary in 1991-1992; IV is the mean of RD an . Data are
given as means (SEM).
Species Density RD RF IV
Fagus grandifolia Ehrhart 235.9 (86.8) 49.9 (6.7) 34.0 (2.6) 41.9 (4.0)
Quercus rubra 85.6 (25.0) 18.3 (4.2) 30.1 (7.6) 24.2 (5.8)
Tsuga cantalensis (L.) Carriere 54.6 (12.1) 15.9 (5.3) 15.6 (2.9) 15.7 (3.9)
cer rubrum L. 22.7 (9.7) 4.4 (1.4) 5.4 (1.2) 4.9 (1.2)
melanchier spp.** 15.5 (5.9) 3.8 (1.3) 4.2 (1.6) 4.1 (1.5)
Pinus strobus L 11.3 (6.8) 2.0 (1.4) 2.9 (1.7) 2.5 (1.5)
Fraxinus americana 3.5 (1.4) 1.6 (1.0) 2.3:{12) 2.0 (1.1)
Quercus prinus L 2.3 (0.8) 1.0 (0.6) 1.7 (0.8) 1.3 (0.7)
Acer pensylvanicum L. 2.2 (2.2) 0.5 (0.5) 0.5 (0.5) 0.5 (0.5)
hododendron maximum L 1.9 (1.9) 0.4 (0.4) 0.2 (0.2) 0.3 (0.3)
Prunus serotina Ehrh. 1.5 (0.9) 0.2 (0.1) 0.4 (0.2) 0.3 (0.2)
i/ icillata (L.) A. Gray 1.5 (1.5) 0.3 (0.3) 0.2 (0.2) 0.3 (0.3)
Carya spp.*** 1.2 (0.9) 0.7 (0.7) 1.1 (0.9) 0.9 (0.8)
Other species* . i
Total Density 443.6

*Seedlings and sprouts were defined as stems 11 to 140 cm tall.
**Includes Amelanchier arborea (Michx. f.) Fern. and A. laevis Wieg.
***Includes Carya glabra (Miller) Sweet, C. ovsta (Miller) K. Koch, and C. tomentosa (Poiret) Nut
*Other species (IV) include Betula spp. (0.3), Tilia americana L. (0.3), Vitis labrusca L. (0.2), Acer melee Marshall
(0.1), Hamamelis virginiana L. (0.1), and Nyssa sylvatica Marshall (0.1)

8 BARTONIA

TABLE 5. Shrub and liana density (stems ha"), relative density (RD), relative frequency (RF), and
importance value (IV) at Lacawac Sanctuary in 1991-1992; IV is the mean of RD and RF. Data are
given as means (SEM).

Species Density RD RF IV
Vaccinium angustifolium Aiton* 251.3 (108.5) 50.5 (21.0) 41.9 (12.1) 46.2 (16.3)
Rhododendron maximum L. 182.4 (133.7) 36.2 (18.2) 26.5 (12.0) 31.4 (15.0)
Vitis labrusca L. 20.4 (8.9) 9.1 (6.3) 17.0 (6.5) 13.1 (6.2)
Kalmia latifolia L. 6.3 (3.2) 1.0 (0.4) 3.6 (1.5) 2.3 (0.9)
Vaccinium corymbosum L. 4.2 (2.7) 0.6 (0.4) 2.6 (1.5) 1.6 (0.9)
Rubus hispidus L. 2.5 (0.8) 0.7 (0.3) 3.3 (1.3) 2.0 (0.8)
Berberis thunbergii DC. 1,7 (1.7) 0.5 (0.5) 1.4 (1.4) 1.0 (1.0)
Parthenocissus quinquefolia (L.) Planchon 2.3 (13) 1.0 (0.7) 1.6 (0.9) 1.3 (0.8)
Viburnum acerifolium L. 0.4 (0.4) 0.1 (0.1) 0.4 (0.4) 0.2 (0.2)
0.6 (0.6) 0.1 (0.1) 0.4 (0.4) 0.2 (0.2)
Sambucus racemosa L. 0.3 (0.3) 0.2 (0.2) 0.9 (0.9) 0.5 (0.5)

*may include some plants of V. pallidum Aiton.

the negative sign and size suggest a potential inverse relationship between the two variables
(statistical power was low due to small sample size). However, even in one forest stand
where T. canadensis was third in importance (IV = 13.7) behind A. rubrum (IV = 25.2) and
Q. rubra (IV = 18.8), seedling density was only 288 seedlings ha’, 98.8% of which were 11
to 30 cm tall.

The shrub layer was sparse, with 196 (50%) of the plots we surveyed lacking any shrubs
or lianas. Vaccinium angustifolium (lowbush blueberry) was the most common shrub,
occurring in 27.8% of all plots at a mean density of 251 stems ha‘ (Table 5). The majority
(92.5%) of V. angustifolium shrubs were <30 cm tall. Rhododendron maximum, the only
other common shrub, occurred in 15.3% of all plots at a mean density of 182 stems ha"
(Table 5). Kalmia latifolia and Vaccinium corymbosum were minor constituents of the shrub
layer. Vitis labrusca was the most common liana (Table 5).

There were significant differences among the four strata (trees, saplings, seedlings/sprouts
and shrubs) in species richness (F;,, = 9.80, P = 0.002), species diversity (F,,. = 11.00, P
<0.001), and species evenness (F;,, = 8.60, P = 0.003). Multiple comparisons tests for all
three variables indicated that significant ANOVAs were solely a consequence of significantly
lower richness, diversity and evenness of shrubs (all P <0.05) compared to the other three
strata. Although means of all three variables declined from trees to saplings to seedlings, the
declines were not statistically different.

Morisita’s index of community similarity indicated fairly strong overlap between canopy
trees and saplings (Mean I,, = 0.82, SEM = 0.05), a much lower overlap between saplings
and seedlings/sprouts (Mean I, = 0.45, SEM = 0.04), and the lowest overlap between
canopy trees and seedlings/sprouts (Mean I, = 0.38, SEM = 0.05). The differences in
similarity values were significant (Foi. = 21.62, P <0.001).

DISCUSSION

Forest Type Affiliation
The Lacawac forest is a hemlock-red maple-red oak mixed hardwoods forest, at least in
its canopy composition. We agree with Russell et al. (1993) that it could be viewed as a

REGENERATION FAILURE IN A HARDWOOD FOREST 9

transition between the Appalachian oak and northern hardwoods biomes, and our canopy
data confirm their findings of a stronger oak presence than would be expected of a typical
northern hardwoods forest (Table 1). However, for several reasons, we suggest that Lacawac
should be regarded as more closely affiliated with the northern hardwoods than the
Appalachian oak forest (Braun 1950, Kuchler 1964). First, only two Quercus species were
common, an . rubra is not an unusual constituent of northern hardwoods forests,
especially in eastern areas of the Allegheny Section (Braun 1950). Second, besides T.
canadensis and A. rubrum being the two most common species, F. grandifolia, A. saccharum,
P. strobus and B. lenta were all fairly well represented in the canopy. Third, sapling
composition was strongly representative of a typical northern hardwoods forest (Table 2),
with a virtual absence of Quercus.

Our results add Lacawac to a list of eastern U.S. forests in which Quercus is declining in
importance (Crow 1988, Abrams and Downs 1990, Abrams 1992). Although still important
in the canopy, the striking dearth of Q. rubra or Q. prinus in the sapling and seedling strata
portends a steady oak decline at Lacawac over the next several decades as canopy trees die.
Oak regeneration failure at Lacawac is likely the result of a combination of factors, including
heavy shade (especially in hemlock-dominated stands), the absence of fire for at least the last
50 years, and deer browsing (Crow 1992; Abrams 1992). There have been no fires in the
sanctuary forest since 1948, and probably for several decades before that (Arthur Watres,
pers. commun.).

Regeneration Failure

The forest understory at Lacawac Sanctuary was largely devoid of sizeable seedlings or
sprouts and small saplings. Densities of seedlings/sprouts and saplings were among the lowest
values reported in the literature for any of several eastern U.S. forests that we surveyed,
either second-growth or old-growth (Table 6). In second growth forest stands in Wisconsin,
Anderson and Katz (1993) found seedling densities (stems 15 cm tall to 1.3 cm dbh by their
definition) ranging from 7,079 stems ha’ in stands “subjected to intense deer browsing
pressure” to 16,055 stems ha" in stands where deer had been excluded for 27 years. Using
more liberal size limits (11 cm tall to 2.5 cm dbh), we recorded densities of only 458 stems
ha? (Figure 1, Table 3). In a Pocono Plateau hardwood forest located about 50 km south of
Lacawac, Held and Giuliani (1989) reported sapling densities (stems 1.0 m tall to 10.0 cm dbh
by their definition) of 1,658 stems ha’. By contrast, stems in the same size range at Lacawac
occurred at a density of 241 stems ha’. Even discounting Held and Giuliani’s (1989) inclusion
of Kalmia latifolia among saplings (we report them as shrubs), sapling density at their site
was still about six times greater than at Lacawac.

In a western Pennsylvania old-growth forest, where deer browsing had severely impacted
the understory, Whitney (1984) found densities of small saplings (30 cm tall to 2.5 cm dbh
by his definition) that ranged from 28 to 110 stems ha. Applying the same size criteria, we
found densities of 38.7 stems ha! at Lacawac. Whitney’s (1984) was the only eastern forest
site that we found in the literature with seedling and sapling densities as low as those at
Lacawac.

We propose the use of the ratio of sapling density to canopy density as a useful measure
of understory condition. In most forests, sapling to canopy density ratios are >1.0 (Table
6). The ratio at Lacawac (0.49) was much lower than in any study we examined, except for
a second-growth forest in northern Wisconsin (sapling/canopy density = 0.11) noted to have
been subjected to “intense deer browsing pressure” (Anderson and Katz 1993).

BARTONIA

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REGENERATION FAILURE IN A HARDWOOD FOREST 11

We hypothesize that the striking failure of regeneration in the forest at Lacawac is, at least
in part, a consequence of overbrowsing by white-tailed deer. The very low density of
saplings, especially those in the 0.1 to 2.5 cm dbh size class (Table 3), suggests that
regeneration failure extends many years into the past. Observations made subsequent to this
study (Townsend, unpubl. data) of seed production and the density of seedlings <10 cm tall
suggest that neither seed rain nor germination rates have limited recruitment into the size
class that was so badly depleted of woody stems. Furthermore, there are no other vertebrate
browsers in the Lacawac forest that would constitute alternative causes of regeneration
failure (David Byman, pers. comm.).

Shading by T. canadensis likely accounts, in part, for the very low densities of seedlings
and small saplings that we observed in the Lacawac forest. However, we did not find a
significant correlation between hemlock density and seedling density among the four stands
we studied, and regeneration failure occurred in all four stands that we studied, including one
that was dominated by A. rubrum and Q. rubra, with lower T. canadensis importance.
Furthermore, even in stands dominated by T. canadensis, sprouting (by root suckers or basal
sprouts) should represent an important alternative regeneration mechanism for many woody
species. But we found a virtual absence of sprouts of canopy trees, understory trees or tall
shrubs (except for F. grandifolia) everywhere in the sanctuary. And even F. grandifolia,
reportedly a less preferred food for deer (Bramble and Goddard 1953; Richards and
Farnsworth 1971), exhibited significant evidence of browsing.

ACKNOWLEDGMENTS

This study was funded by University of Scranton faculty research grants (to D.S.T.), and
undergraduate student research awards (to J.S.S. and C.H.) under a Howard Hughes Medical
Institute Grant to the University of Scranton. We are grateful to Don Waller, Tom Rooney,
Ned Fetcher, Roger Latham and two anonymous reviewers for providing constructive
comments on various versions of the manuscript.

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REGENERATION FAILURE IN A HARDWOOD FOREST 13

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Bartonia No. 61: 15—26, 2002

Interrelationships Among Achene Weight, Orientation,
and Germination in the Asters
Doellingeria umbellata var. umbellata,
Symphyotrichum novae-angliae and S. puniceum
(Asteraceae)

JERRY G. CHMIELEWSKI AND SONAM RUIT
Department of Biology, Slippery Rock University of Pennsylvania, Slippery Rock, PA 16057
jerry.chmielewski@sru.edu

umbellata, Symphyotrichum novae-angliae, and S. puniceam. Achenes of S. novae-angliae and D.
umbellata that germinated were on average heavier than were those that did not germinate. All achenes
of S. puniceum germinated irrespective of treatment or size. Achenes of all species germinated three
to four times more quickly in agar medium trials, irrespective of orientation, than in filter-paper trials.
Species varied in response to germination media: a greater proportion of D. umbellata var. umbellata
achenes germinated on agar than on filter paper; more S. novae-angliae germinated on filter paper; and
germination of S. puniceum was similar in both. Total germination was not affected by orientation for
any species. Achene weight was not correlated with the time taken to germinate for either S. puniceum
or D. umbellata, irrespective of orientation. In contrast, for S. novae-angliae the time to germinate was
negatively correlated with achene weight regardless of orientation. However, achenes of S. novae-
angliae oriented in the up position took significantly longer to germinate than did those in the
horizontal or down positions. The results of this study indicate that differential allocation of resources
occurs in the three commonly co-occurring, weedy aster species and that germination success in them
may or may not be related to these allocations, genetic factors, or environmental conditions, but not
orientation.

INTRODUCTION

Certain physical and chemical requirements, such as those related to light quality,
moisture availability, oxygen concentration, pH, and temperature regime, must be satisfied
for seed germination to occur (Evanari 1956; Toole et al. 1956; Bradbeer 1988). The
relationship between and among these requirements, microsite characteristics and seed
morphology will define both total germination and time taken for germination to occur
(e.g., Harper 1977; Jones 1978; Bewley and Black 1982; Baskin and Baskin 1979, 1985;
Prinzie and Chmielewski 1994). Although interspecific differences occur in a variety of seed
characteristics (e.g., seed size or seed coat characteristics), intraspecific differences, that
is, the production of morphologically different seeds within a species (e.g., Weiss 1980;
Forsyth and Brown 1982; Maun and Payne 1989; McGinley 1989; Zhang 1993; Venable et
al. 1995), appear to be restricted, with exceptions, to relatively short-lived, fugitive species,

Manuscript submitted 30 August 2000, revised 19 December 2000.

15

16 BARTONIA

particularly weeds, in the families Asteraceae, Brassicaceae, Chenopodiaceae, and Poaceae
(Harper 1977).

Species whose seed types have different constraints on their evolution, such as ray and
disk achenes in the Asteraceae (specifically in subfamily Tubuliflorae), provide a unique view
of the adaptive consequences of seed heteromorphism. In these species, seed functions tend
to diverge (Venable 1985). Reduced dispersability and delayed germination are characteristic
of the outer achenes, whereas distance dispersal and quick germination are more typical of
the central achenes (Forsyth and Brown 1982; McEvoy 1984; Venable and Levin 1985;
Tanowitz et al. 1987).

Studies that previously reported total germination in aster species (Nichols 1934; Delisle
1938; Wetmore and Delisle 1939; Avers 1953; Havercamp and Whitney 1983; Jones 1978;
Kothman and Kallah 1978; Peterson and Bazzaz 1978; Baskin and Baskin 1979, 1985; Allen
et al. 1983; Pitelka et al. 1983; Clampitt 1987; Chmielewski 1991; Prinzie and Chmielewski
1994; Bosy and Aarssen 1995) collectively did not differentiate between the essentially
monomorphic achenes produced by ray and disk florets. Both Chmielewski (1991) and
Prinzie and Chmielewski (1994) indicated their uneasiness in excluding such a potential
source of variation in germination trials and recommended a procedural guide for future
studies dealing with the evolutionary significance of achene germination strategies in
heterogamous composites (Chmielewski and Huff 1995). Although achenes derived from
either ray or disk florets were indistinguishable in the aster Eurybia divaricata (L.) Nesom
(Aster divaricatus L.), ray achenes were on average lighter than disk achenes, and this
difference was due to increased allocation to the embryo rather than the pericarp. Further,
pericarp thickness decreased with increasing achene weight in the disk achenes. Neither total
germination nor the time taken to germinate by ray or disk achenes was correlated with
achene weight or embryo weight, but instead varied inversely with pericarp weight for ray
achenes only. The authors recommended that, unless the consequences of differential
allocation to the embryo and pericarp of achenes are documented, future germination studies
should differentiate between ray and disk achenes. A second study, using Doellingeria.
umbellata (Miller) Nees var. umbellata (Aster umbellatus Mill. var. umbellatus), indicated that
the number of days to germination was not correlated with any measured aspect of achene
weight for either ray or disk achenes whether the Pappus was removed or intact
(Chmielewski 1999). The production of morphologically and ecologically indistinguishable
ray and disk achenes would suggest that the significance of the heterogamous capitula
(specifically the ray florets) rests in their attractiveness to pollinators (Chmielewski 1999).
Based on these studies we conclude that the information gained by separating disk and ray
achenes for use in germination studies does not warrant the additional effort necessary to
distinguish the two.

Although the effects of physical and chemical requirements on germination have been
studied for a variety of species (e.g., Bewley and Black 1982), as have general seed
morphology (e.g., Williams and Harper 1965; McEvoy 1984; Venable and Burquez 1989) and
dimorphism (see Chmielewski 1999), the relationship between patterns of within-achene
allocation and seed orientation have not (Rowlee 1893; Sheldon 1974; Peart 1984; Bosy and
Aarssen 1995).

Prior to 1994 the literature treats the aster species included in this study as members of
Aster, but chloroplast DNA evidence (Xiang and Semple 1996) and gene sequence data
(Noyes and Rieseberg 1999) demonstrate that Doellingeria, Symphyotrichum, and other genera
are distant from Aster. Reference to the aster literature in general follows recently defined

EFFECTS ON GERMINATION IN THREE SPECIES OF ASTER 17

infraspecific classifications (Semple and Chmielewski 1987; Semple et al. 1991) and revised
generic combinations (Nesom 1994). For the sake of clarity the Aster epithet referenced will
appear in parentheses with each first use of the new aster epithet.

In this study we examine the relationship between within-achene allocation (total weight,
embryo weight, and pericarp weight) and orientation on germination in three clonal species,
Doellingeria umbellata var. umbellata, Symphyotrichum novae-angliae (L.) Nesom (Aster novae-
angliae L.) and S. puniceum (L.) Léve and Love (Aster puniceus L.). These species were
selected for this study because each is considered weedy through at least part of their
respective ranges (Alex et al. 1980; Anonymous 1990; Mulligan 1992), they often co-occur
in open, abandoned fields, they are easily distinguishable, mean achene weight occurs in the
upper, middle, and lower weight classes found in the asters, and clonal structure differs
among the species. Doellingeria umbellata forms clones that are neither freely spreading nor
compact, but rather continuous in the sense that the inflorescences of adjacent stems touch
or overlap, but are not crowded. Symphyotrichum puniceum forms clones that are freely
spreading and open; adjacent stems may be quite distant. Symphyotrichum novae-angliae
forms clones that are extremely compact, the stems being tightly clustered (Semple et al.
1996

METHODS

Mature capitula of Doellingeria umbellata, Symphyotrichum novae-angliae, and S. puniceum
were harvested from groupings of stems (likely clones) growing in an abandoned field on
the Slippery Rock University campus (Slippery Rock, Pennsylvania) during the fall of 1998.
The capitula were placed in manila envelopes and returned to the laboratory where they
were allowed to air dry for 1 week. Following air drying, the capitula were stored (-4 + 2°
C) in the dark for 20 weeks. The following spring >400 achenes of each species that
appeared full and resisted a gentle pinch with forceps were randomly selected for
germination trials. Achenes were not separated on the basis of whether they were of the disk
or ray type.

For some germination trials, the pappus was teased from the achene. Teasing entailed
lightly brushing the pappus bristles with fine forceps until the majority had detached; those
few that remained were actively grasped and plucked. Individual achenes were then weighed
on a Cahn C-33 microbalance (+2 pg). Just prior to sowing, achenes were surface sterilized
with a 30% bleach solution to minimize or eliminate fungal growth.

The first germination trial entailed the horizontal placement of achenes on moistened
filter paper in 150 x 15 mm petri dishes. These were placed in a Biotronette Mark III
environmental chamber with a temperature of 24 + 2° C, 12 h photoperiod, and light
intensity of 218 pmol s! m?. The petri dishes were randomly re-positioned on a daily basis.
Achenes were examined daily for 135 days (coincident with the last day of the agar study
trial); germination was indicated by either the emergence of the radicle or cotyledons,
whichever appeared first. Petri dishes were moistened regularly as needed. This germination
trial allowed for the post-germination recovery of pericarps. Pericarps were harvested for
each germinated achene as soon as they could be removed from the seedling. Air-dried
pericarps were subsequently individually weighed. Embryo weight was determined to be the
difference between total achene weight and pericarp weight. The results of this germination
trial were used to: (1) establish the relationships among total achene weight, pericarp weight,
and embryo weight for each species; (2) determine the relationship between within-achene

18 BARTONIA

allocation to both total germination and time taken to germinate; and (3) provide baseline
data for the achene orientation germination trial.

The second germination trial was designed to determine the relationship between achene
weight and orientation on germination. Following surface sterilization with a 30% bleach
solution, individually weighed achenes were sown into a 0.8% agar medium prepared with
de-ionized water (following the methodology of Bosy and Aarssen 1995) and poured to a
depth of approximately 1 cm into 150 x 15 mm petri dishes. Individual achenes (100 per
position for each species) were placed below the agar surface and positioned vertically
upward (pappus end up), horizontally, or vertically downward (pappus end down). Each
petri dish contained a maximum of 30 achenes oriented in only one position. The viscosity
of the agar was such that sown achenes were not only suspended in the agar in a specific
orientation, but also completely in contact with it throughout the germination trial.
Conditions for germination were similar to the previous trial. The petri dishes were
randomly repositioned within the environmental chamber on a daily basis. Achenes were
examined daily for 38 days during which the agar remained moist and the majority of
germination had occurred. Germination was indicated by either the emergence of the radicle
or cotyledons, whichever came first.

SAS/STAT (SAS Institute Inc. 1989) procedures and options were used for data analysis.
The UNIVARIATE procedure was used to summarize descriptive statistics (achene weight,
pericarp weight, embryo weight, and time taken to germinate) for the respective species. The
Shapiro-Wilk W statistic was used to test for normality. Because much of the data were not
normally distributed, the non-parametric Mann-Whitney or Kruskal-Wallis tests (using the
x? approximation of the NPARIWAY procedure) were used for between- and among-group
comparisons respectively. The FREQ procedure and TESTP TABLE option were used to
compare total germination in respective trials. Relationships between combinations of
characters were determined for each species by calculating correlation coefficients. The
CORR procedure, SPEARMAN option was used for this purpose.

RESULTS

Doellingeria umbellata var. umbellata (Table 1)

The mean achene weights of randomly selected samples assigned to each germination trial
were not significantly different. Thus, differences between or among trials with respect to
total germination or time taken to germinate cannot be attributed to differences in achene
weight. Significantly fewer achenes germinated in each of the agar trials compared to the
filter-paper trial. However, much of the germination in the filter-paper trial occurred beyond
the 38-day limit of the agar trial, with only 9.7% of the filter-paper trial achenes germinating
within the agar trial time frame. Total germination was not affected by orientation in the
three agar trials. Germination was weakly correlated with achene weight for each of the
germination trials. Achenes that germinated in each trial were heavier than achenes that did
not, but the weight of germinated achenes did not differ across trials. For the filter-paper
trial, embryo weight was positively correlated with both achene weight and the number of
days taken by an achene to germinate. Also for this trial, pericarp weight was negatively
correlated with the number of days taken to germinate. No other correlations among achene
weight, pericarp weight, embryo weight, and days taken to germinate were significant for
any of the trials. The time taken by an achene to germinate was significantly greater for the
filter-paper trial compared to any other. The time taken by an achene to germinate was not
affected by orientation in the three agar trials.

EFFECTS ON GERMINATION IN THREE SPECIES OF ASTER 19

TABLE 1. Summary of descriptive statistics (+ standard deviation) for achenes of Doellingeria umbellata
var. umbellata germinated in petri dishes on filter paper or agar medium in one of three orientations.
Weight in mg.

Agar Orientation

Filter Paper

Pappus
End Up

Pappus
Horizontal

Pappus
End Down

103
0.758 + 0.115
re

100
0.761 + 0.167

100
0.744 + 0.158

100
0.727 + 0.154
re

2
0.694 + 0.152

ean ungerminated achene weight 0.712 + 0.126 0.734 + 0.178 0.702 + 0.160
Number of germinated achenes 66 29 35
Mean germinated achene weight 0.784 + 0.100 0.825 + 0.116 0.821 + 0.122 0.814 + 0.127
Comparison of mean achene weight x? = 7.6742 xX° = 4.2695 x? = 12.412 x? = 11.961

(germinated versus ungerminated;

df = 1 P = 0.0056 P = 0.0388 P = 0.0004 P = 0.0005
Mean pericarp weight 0.221 + 0.043
Mean embryo weight 0.563 + 0.102
Mean number of days to germinate 739 £337 2.5 £75 25.3 + 6.5 22.0 + 7.2
Median number of days to germinate 83 23 25 22
Symphyotrichum novae-angliae (Table 2)

Mean achene weights of randomly selected samp] igned to eac h trial were

not significantly different. Thus, differences betw

o °o
een or among trials in total germination

or time taken to germinate cannot be attributed to differences in achene weight. Significantly
more achenes germinated in each of the agar trials compared to the filter-paper trial.
Further, only 35% of the filter-paper trial achenes germinated within 38 days, the duration
of the agar trials. Total germination was not affected by orientation in the three agar trials.
Germination was not correlated with achene weight for each of the germination trials.
Achenes that germinated were heavier than achenes that did not germinate, but the weight

TABLE 2. Summary of descriptive statistics (+ standard deviation) for achenes of Symphyotrichum

novae-angliae germinated on filter paper or agar medium in one of three orientations. Weight in mg.
Agar Orientation
Pappus Pappus Pappus
Filter Paper End Up Horizontal End Down
Number of achenes sown 102 100 100 100
Mean achene weight 0.374 + 0.074 0.362 + 0.095 0.369 + 0.096 0.379 + 0.090
Number of ungerminated ach 55 15
ean ungerminated achene weight 0.378 + 0.070 0.313 + 0.109 0.264 + 0.113 0.341 + 0.093
Number of germinated achen 77 79
Mean germinated achene weight 0.371 + 0.079 0.377 + 0.086 0.388 + 0.079 0.389 + 0.089
Comparison of mean achene weight x? = 0.3857 x2 = 6.3853 x? = 14.139 x? = 4.8604
(germinated versus ungerminated;
df = 1) P = 0.6250 P = 0.0115 P = 0.0115 P = 0.0115
Mean pericarp weight 0.076 + 0.025
Mean embryo wi 0.294 + 0.081
Mean number of days to germinate 22.6 + 24.3 8.5 + 7.5 5.1 + 4.1 6.9 + 5.1
6 4 6

Median number of days to germinate 9

20 BARTONIA

of germinated achenes did not differ across trials. For the filter-paper trial embryo weight
was positively correlated with achene weight. For the agar trials achene weight was
negatively correlated with the number of days taken by an achene to germinate. No other
correlations (among achene weight, pericarp weight, embryo weight, and days taken by an
achene to germinate) were significant. The time taken by an achene to germinate was not
only significantly different among the four trials, but also among the agar orientation trials.

Symphyotrichum puniceum (Table 3)

ean achene weights of the randomly selected samples assigned to each germination trial
differed; those used for the agar trial and sown horizontally were, by chance, significantly
lighter than others. Although 100% of the achenes germinated irrespective of treatment, only
82% of the filter-paper trial achenes germinated within 38 days, the duration of the agar
trials. Germination was not correlated with achene weight. For the filter-paper trial embryo
weight was positively correlated with achene weight. Pericarp weight was positively
correlated with achene weight, but negatively correlated with the number of days taken by
an achene to germinate. No other correlations among achene weight, pericarp weight,
embryo weight, and days taken to germinate were significant for any of the trials. The time
taken by an achene to germinate was significantly greater for the filter-paper trial compared
to any other. The time taken by an achene to germinate was not affected by orientation in
the three agar trials.

TABLE > Summary of descriptive statistics (+ standard deviation) for achenes of Symphyotrichum
puniceum germinated on filter paper or agar medium in one of three orientations. Weight in mg.

Agar Orientation

Pappus Pappus Pappus

Filter Paper End Up Horizontal End Down
Number of achenes sown 101 100 100 100
Mean achene weight 0.293 + 0.053 0.294 + 0.048 0.267 + 0.054 0.284 + 0.051
Number of germinated achenes 101 1 100
Mean germinated achene weight 0.293 + 0.053 0.294 + 0.048 0.267 + 0.054 = 0.284 + 0.051
Mean pericarp weight 0.100 + 0.028
Mean embryo weight 0.193 + 0.046
Mean number of days to germinate 18.2 + 19.3 4.0 + 2.1 4.5 + 2.6 4.8 + 3.0
Median number of days to germinate 8 3 3 3

DISCUSSION

Achene weight is quite variable among the asters, with at least a 10-fold difference
evident (Delisle 1938; Wetmore and Delisle 1939; Peterson and Bazzaz 1978; Havercamp an
Whitney 1983; Pitelka et al. 1983; Chmielewski 1991, 1999; Prinzie and Chmielewski 1994)
among the weedy species (Alex et al. 1980; Anonymous 1990; Mulligan 1992). Eurybia
divaricata, the single non-weedy aster for which achene weight is available, produces
comparatively light achenes (Chmielewski and Huff 1995).

Achenes of Doellingeria umbellata, Symphyotrichum novae-angliae, and S. puniceum, all
wetland or facultative wetland species (U.S. Department of the Army 1987), occur in the
upper, middle, and lower weight classes, respectively, found in the asters. Achenes of D.
umbellata are, on average, nearly twice the weight of those produced by many aster species.

EFFECTS ON GERMINATION IN THREE SPECIES OF ASTER 21

In this study, achenes of D. umbellata were significantly heavier, averaging 0.8 mg, with
some > 1.1 mg, than reported values (0.6 mg in Chmielewski 1999). Although achene weight
in S. novae-angliae was first reported as 0.41 to 0.45 mg (Delisle 1938; Wetmore and Delisle
1939), a significantly lower mean value of 0.20 mg (pappus removed) was subsequently
reported (Havercamp and Whitney 1983). In considering that the pappus of aster achenes
typically weighs between 0.06 and 0.08 mg (Chmielewski 1999) the values reported in the
present study (0.36-0.38 mg) are comparable to those in the former two reports (weight
includes adjoining pappus), but not the latter. These values are also comparable to that
reported for a white-flowered hybrid (0.41 mg) of the species (Delisle 1938). Achene weight
in S. puniceum (0.26-0.30 mg), the only weedy, obligate wetland aster species for which
values are available, tends toward the lowest achene-weight class in the asters. Variation in
achene weight both within and among aster species may be attributable to cytotypic
differences, clonal variation, yearly variation, or a combination of these or other factors.
Past studies on asters suggested that because achene weight in some species may be correlated
with total germination, the evolutionary significance of achene weight is associated with pre-
germination phenomena such as dispersal and post-germination phenomena such as seedling
survival or vigor (Chmielewski 1991, 1999; Prinzie and Chmielewski 1994). None of these
phenomena were part of this investigation.

Following late autumn dispersal, aster achenes imbibe on the soil surface and are exposed
to stratification temperatures during the winter months, germinating in early spring (Baskin
and Baskin 1979). Germination is controlled by the phytochrome system, with red and
white light promoting germination and darkness and far-red light inhibiting germination
(Peterson and Bazzaz 1978; Baskin and Baskin 1979). Despite this controlling system,
germination immediately following harvest is quite variable. Some of the weedier species,
for example Symphyotrichum ericoides (L.) Nesom (Aster ericoides L.), S. falcatum (Lindl.)
Nesom (A. falcatus Lindl. in Hook.), S. lanceolatum (Willd.) Nesom (A. lanceolatus Willd.),
S. pilosum (Willd.) Nesom (A. pilosus Willd.), S. sericeum (Vent.) Nesom (A sericeus Vent.),
and S. tradescantii (L.) Nesom (A. tradescantii L.) germinated readily (60-80%) immediately
following harvest (Keever 1950; Jones 1978; Baskin and Baskin 1979, 1985). These values are
contrasted by comparatively low immediate post-harvest germination in S. lateriflorum (L.)
Live and Léve (A. Jateriflorus [L.] Britton), S. praealtum (Poir.) Nesom (A. praealtus Poir.),
S. novae-angliae (L.) Nesom, S. oolentangiense (Riddell) Nesom (A. azureus Lindl. in Hook.),
S. drummondii (Lindl.) Nesom (A. drummondii Lindl. in Hook.) and S. sagittifolium
(Wedem. ex Willd.) Nesom (A. sagittifolius Wedem. ex Willd.) (Keever 1950; Jones 1978;
Baskin and Baskin 1979, 1985).

The relationship between cold-temperature exposure and germination in Symphyotrichum
novae-angliae and other asters is poorly defined. Germination in the red form of S. novae-
angliae (32%) exceeded that of the typical form (8%), but the conditions under which
germination occurred were not specified (Rowlee 1893). No difference in total germination
was reported between refrigerated (exposed to an outdoor cold frame; 64.5%) or unrefrigerat-
ed (61.5%) achenes sown directly in the greenhouse (Nichols 1934). Germination
i diately following | as rep ly low (6%), but increased considerably following
two months of refrigeration in a moist atmosphere (Jones 1978). In our study, germination
of refrigerated achenes ranged between 15 and 54%, depending on trial. No comparable cold
treatment data are available for Doellingeria umbellata or S. puniceum.

Based on germination, viability is maintained for at least 135 days by D. umbellata, 110
days by S. novae-angliae, and 67 days (all achenes of the species had germinated by this day)

22 BARTONIA

by S. puniceum. Germination of surface sown achenes of D. umbellata on fine-grained
vermiculite occurred until the 90th day of a 150-day germination trial (Chmielewski 1999).
Uninjured, soaked achenes of S. novae-angliae, which did not germinate spontaneously,
remained viable and germinable for over eight months provided they were maintained in a
moist atmosphere (Jones 1978). Nichols (1934) had previously reported that refrigerated
achenes of S. novae-angliae remained viable for 28 days and germinated in a minimum of 14
days, whereas those that were not refrigerated remained viable for at least 78 days and
germinated in a minimum of 43 days. These results suggest that viability, as applied in each
of the respective studies, varies not only among species, but also within a species. Clonal
differences, yearly variation, or a combination of these or other factors may account for the
latter. Although the longevity of buried aster achenes is not well documented (Livingston
and Allessio 1968; Rabinowitz 1981; Johnson and Anderson 1986; Leck and Leck 1998),
viability of at least three months is necessary to allow for spring germination in these and
other aster species.

Achene weight had a relatively insignificant role in affecting total germination in
Doellingeria umbellata and no effect in either Symphyotrichum novae-angliae or S. puniceum.
Previous studies on other species also reported that achene weight may or may not affect
germination. For example, heavy achenes of Oclemena acuminata (Michx.) E.L. Greene

ster acuminatus Michx.) exhibited significantly higher germination than did the lighter
achenes (Pitelka et al. 1983). Both tetraploid and hexaploid achenes of S. lanceolatum did not
begin to germinate with any regularity unless above minimum weights (Chmielewski 1991).
Germination was positively correlated with achene weight for S. pilosum (Prinzie and
Chmielewski 1994), but was not related to achene weight for either ray or disk achenes of
Eurybia divaricata (Chmielewski and Huff 1995). Achene weight did not affect total
germination in ray or disk achenes of D. umbellata for which the pappus was removed or
remained intact (Chmielewski 1999). Our results in conjunction with previous reports
suggest that the relationship between achene weight and total germination in asters is
species-specific.

Variability in achene weight presumably reflects a balance between architectural
constraints associated with the capitulum and the evolutionary pressures associated with
dispersal, persistence, predator avoidance, germinability, or seedling competition (Harper
1977; Jolls and Werner 1989). The three species included in this study exhibited different
strategies in the allocation of resources within achenes and between allocation patterns and
the time taken to germinate. As achene weight increased in Doellingeria umbellata, embryo
weight also increased, but pericarp weight was static. Differential allocation between heavy
and light achenes was not previously reported for the species (Chmielewski 1999). The time
taken by these achenes to germinate was positively correlated with achene weight, but
negatively correlated with pericarp weight. As achene weight increased in Symphyotrichum
novae-angliae so did embryo weight, but the time to germination was not affected. Both
pericarp weight and embryo weight increased with increasing achene weight in S. puniceus
but the time to germination was not affected. A significant inverse relationship between
pericarp weight fraction and achene weight, with heavier achenes having proportionately
thinner pericarp, was predicted by the crude model presented in Prinzie and Chmielewski
(1994). Because it is the embryo, which by imbibition and cell expansion provides the
physical force required to rupture the pericarp (Cook 1980), heavier achenes should
germinate with greater regularity and should do so more quickly than lighter achenes. Ray
achenes of Eurybia divaricata were significantly lighter than disk achenes, the difference

EFFECTS ON GERMINATION IN THREE SPECIES OF ASTER Pe

being due to increased allocation to the embryo as opposed to pericarp, but total germina-
tion did not differ between the two (Chmielewski and Huff 1995).

The embryo within an aster achene is arranged so that if the achene lands pappus end
up, the cotyledons are up and the hypocotyl is down. With this arrangement the embryonic
parts occur in the most advantageous position for germination (Rowlee 1893), the

lationship between germination and seed orientation reflecting differential effects of gravity
on the embryo (Bosy and Aarssen 1995).

The pappus of aster achenes assists in dispersal (Rowlee 1893; Carlquist 1967; Sheldon and
Burrows 1973) and it also facilitates at least initial optimization of orientation for
germination to occur (Rowlee 1893; Sheldon 1974; Bosy and Aarssen 1995). Following
several days of soaking, the pappus of these and other aster species irreversibly collapses and
the achenes rest more or less horizontally on the substrate. The pappus in some aster species
may be less important in its effects on germination than anticipated, as germination in disk
and ray achenes of Doellingeria umbellata was 2.5 to 3 times greater among achenes from
which the pappus was removed compared with intact achenes (Chmielewski 1999). Similar
results were reported for disk and ray achenes of Eurybia divaricata (Chmielewski and Huff
1995). In contrast, the status of the pappus, whether removed or intact, had little or no
effect on germination in Oclemena acuminata (Pitelka et al. 1983).

Germination in Doellingeria umbellata was not affected by orientation in this study.
However, the species did exhibit a range of germination percentages for horizontally
oriented achenes (pappus removed) germinated on vermiculite (19.6% in Chmielewski 1999),
filter paper (64%), and agar (35%). Because mean achene weights among these trials were
similar, the observed differences must be attributable to some factor other than achene
weight. Although the duration of the vermiculite (150 days) and filter paper (135 days)
germination trials was comparable, the mean number of days to germinate on vermiculite
(16), was considerably less than on the filter paper (76). Because germination of aster achenes
was previously shown to be protracted (Chmielewski and Huff 1995), the authors suggested
that it would be prudent to examine median, rather than mean, time to germination. Median
germination time and mean time to germination for achenes of D. umbellatum were
comparable within each of the respective germination trials. Also, if germination on filter
paper was monitored for the same number of days as was germination on agar, only 9.7%
of the achenes germinated compared to 64%, and the mean number of days to germinate
decreased from approximately 76 to 18.

Total germination in Symphyotrichum novae-angliae for achenes sown with the pappus end
down versus up was variable, but typically low (4 and 12% respectively; Rowlee 1893). Total
germination in the red-colored form was similarly not affected by orientation, but the values
were considerably higher at 28 and 32% respectively (Rowlee 1893). Contrasting total
germination values of 67, 10, and 77% were reported for achenes sown with the pappus
down, pappus up, or pappus oriented horizontally, respectively (Bosy and Aarssen 1995).
In our study, total germination was not affected by orientation and was similar to that
reported by Bosy and Aarssen (1995) for achenes sown horizontally. Further, inasmuch as
orientation did not affect the number of days to germination in the latter study, results from
our study indicate that horizontally sown achenes germinate more quickly than those with
the pappus oriented down, and that these germinate more quickly than do those with the
pappus oriented up. Median germination time and mean time to germination for achenes
of S. novae-angliae were comparable for the three achene positions in agar, but approximate-
ly 2.5 times longer in the filter-paper trial. The disparate values for germination in the four

24 BARTONIA

studies cited, in concert with the germination values from this study for achenes sown
horizontally on filter paper, suggest that germination in the species is more related to the
source of the achene (ramet, clone, population, etc.) as opposed to orientation or
germination medium. Subsequent comparative germination studies should consider the
source of the achene in defining intraspecific variation.

Complete germination in Symphyotrichum puniceum, irrespective of achene weight,
orientation, or medium suggests that behaviorally the species is the most general in terms
of germination requirements. Median germination time and mean time to germination were
comparable within each of the agar trials, but like S. novae-angliae, approximately 2.5 times
shorter than the filter-paper trial.

Assuming that an achene has “found” a suitable site, many internal or local factors could
affect germination. The results of this and previous studies indicate that within a species
achene weight is variable, that immediate post-harvest germination and the consequences of
refrigeration are variable, and that the duration of germination is variable among species.
The significance of differential allocation of resources within achenes of the three species 1s
witnessed in different times to germination (though the same is not necessarily true for other
aster species). Total germination is typically not affected by orientation per se for these three
species. However, total germination for at least the horizontally oriented achenes is affected,
at least in part, by the medium upon which the achenes are sown. Germination may also
be correlated with moisture availability on the various media, but in all cases saturation of
the achenes was attempted. We conclude that for these three aster species, orientation affects
germination in a specific site to the extent that it affects water availability or imbibition.

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Bartonia No. 61: 27-47, 2002

A Review of the Nomenclature in Witmer Stone’s
The Plants of Southern New Jersey

GERRY MOORE
Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 11225
gerrymoore@bbg.org

ABSTRACT. In 1912, Witmer Stone’s landmark work on the plants of southern New Jersey was
published. This work is the only floristic treatment for southern New Jersey and it continues to be
used today. However, its use is made difficult by the nomenclature, which included elements of the
1905 International Code of Zoological Nomenclature and the defunct American (Brittonian) code of
botanical nomenclature. As a result, many of the names used by Stone are no longer in current use.
Stone’s authorship citations were often incomplete and confusing because he cited only basionym
authors even for combinations and inconsistently used parentheses to indicate basionym authors in
combinations. Also Stone did not clearly indicate the rank of the 60 trinomials he used. For this
paper, all names employed by Stone in The Plants of Southern New Jersey have been reviewed, and it
is concluded that Stone’s trinomials should be regarded as validly published, unranked names. The
19 nomenclatural innovations published by Stone in The Plants of Southern New Jersey are discussed.
A table comparing all of the accepted names in Stone’s manuals with those in two currently used
manuals is available from the author upon request.

INTRODUCTION

Witmer Stone had a nearly lifelong association with the Academy of Natural Sciences of
Philadelphia (Anonymous 1940). He assumed charge of the Academy’s bird collections at
the age of 22 and published on diverse topics of zoology including ornithology, mammalogy,
and zoological nomenclature (see Anonymous 1940; Pennell 1940). He is best known for
his contributions in ornithology, serving as editor of The Auk for over twenty years and
publishing his very well-received Bird Studies at Old Cape May (1937). He was also a
member of the International Commission on Zoological Nomenclature from 1927 until his
death in 1939 (Melville 1995). As a result of his contributions to natural history, he received
an honorary doctorate from the University of Pennsylvania in 1913 (Ledwell 2000).

Besides his interests in zoology, Stone also had research interests in botany, particularly
the flora of southern New Jersey. He was an original member of the Philadelphia Botanical
Club and published numerous papers on the local flora (1902, 1903a, 1903b, 1906, 1907a,
1907b, 1908a, 1908b, 1909, 1910a, 1910b, 1911, 1912b), culminating in 1912 with The Plants
of Southern New Jersey with Especial Reference to the Flora of the Pine Barrens and the
Geographic Distribution of the Species.’ This is the only comprehensive floristic treatment for

Manuscript submitted 15 December 2000, revised 15 April 2001.
'The publication date of Stone’s work has been variously cited as 1910 (the date given on the cover page of the

Annual Report of the New Jersey State Museum in which Stone’s work appeared), 1911 (the date given on the title
page of Stone’s work), and 1912. Pennell (1940) clearly indicated that the work was not effectively published until

27

28 BARTONIA

southern New Jersey, and it continues to be used today. The keys, based on material from
southern New Jersey, and the precise phenological information provided by Bayard Long
make it an excellent field manual. Stone’s detailed discussions on plant biogeography and
his eloquent narratives of his experiences make for informative and pleasant reading when
not in the field.

Harley H. Bartlett (1912), who was harshly critical of Stone’s method of nomenclature,
lauded his flora stating it was “safe to predict that none of the local floras now current will
be longer held in esteem by botanists generally than this one of Mr. Stone’s.” Roland
Harper (1912) regarded it as “one of the best local floras ever published.” Merritt Fernald
(1934a) clearly approved of Stone’s work since he referred to it as “scholarly” (see Jones 1945
for a discussion of Fernald’s use of the word “scholarly”). Fernald (Gentiana stoneana in
Rhodora 41: 555; 1939) and Homer D. House (Viola stoneana in Bulletin of the Torrey
Botanical Club 32: 253; 1903) have named taxa in Stone’s honor (neither name is in current
use). In naming the Pine Barrens gentian (now known as G. autumnalis L.) in honor of
Stone, Fernald noted: “no plant could more appropriately commemorate Witmer Stone than
this retiring and distinguished gentian, with its flowers of true blue.”

In the preface of his work, Stone commented on the confusion surrounding botanical
nomenclature during the early 1900s and noted, “the botanist in America, at least, is on the
horns of a dilemma.” He went on to note that two very different botanical codes were in
use — the American (Brittonian) Code? (Arthur et al. 1907) and the Vienna (International)
Code (Briquet 1906) — and stated that zoologists “will probably find it quite impossible to
accept certain of the features of these codes which are at variance with the International
Zoological Code [International Commission on Zoological Nomenclature 1905].”

Stone commented that the general problems of zoological and botanical nomenclature
were identical and noted that the zoologists “have been ‘playing the game’ seriously longer
than the botanists.” He predicted that “men of the same intellectual ability will eventually
adopt the same methods.” Stone’s prediction of a uniform code or at least unified methods
has yet to come true and a current effort to gain acceptance of a harmonized biological code
of nomenclature (Greuter 1996; Greuter and Nicolson 1996; Greuter et al. 1996; Orchard
et al. 1996) has failed (Barrie and Greuter 1999).

Stone indicated that he would make no attempt to revise botanical nomenclature and
would follow the Brittonian Code. This allowed Stone to use tautonyms (e.g., Sassafras
sassafras), which are not permitted in current botanical nomenclature (Greuter et al. 2000)
but are accepted in zoological nomenclature (I.C.Z.N. 1905; Ride et al. 1999).

However, an important exception Stone made in following the Brittonian Code was “in
the treatment of species and sub-species in separate categories, a most pernicious rule which
botanists will in all probability ultimately reject” (1912a, p. 35). This assignment of
coordinate status to specific and infraspecific ranks is why Stone permitted older varietal
epithets to displace later specific epithets at the rank of species, and it is also why Stone
assigned specific and infraspecific names coordinate status in the index. This practice was
harshly criticized

early in 1912, the first copies being distributed on 14 February. I have Lee Sowden’s (an early member oF ie

Philadelphia Botanical Club) copy of Stone’s flora and in it is a note from Stone indicating that he gave the book

her pre on 14 February 1912. This work was reprinted by Quarterman Publications in 1973 (reviewed by
tafleu 1973).

"Nicolson (1991) argued that the American Code should be called the Brittonian Code since not all American
botanists embraced this code over the Vienna Code

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 23

by botanists, most notably Harley H. Bartlett (1912) who found Stone’s nomenclature
“thoroughly objectionable.”

Specific and infraspecific names having coordinate nomenclatural status has been a long-
standing point of disagreement in biological nomenclature. This practice persists in
zoological nomenclature (Ride et al. 1999), where specific and infraspecific ranks are part of
the species group and the “[p]riority of the name of a nominal taxon is not affected by
elevation or reduction in rank of the taxon within the family group, genus group, or species
group” (Article 23.3.1). Zoological nomenclature (Ride et al. 1999) only recognizes one
formal infraspecific rank, the subspecies (Article 45.6). Coordinate status for specific and
infraspecific ranks was permitted in the original Brittonian Code (Arthur et al. 1904) but
was abandoned in the 1907 version (Arthur et al. 1907). Contrary to Stone’s prediction, the
current botanical code (Greuter et al. 2000) still does not assign coordinate status to specific
and infraspecific names.

Further complicating the use of Stone’s work is the way in which Stone cited authorship.
He followed the zoological tradition of only citing the basionym author, even when the
name used is different in rank or position from that of the basionym. Bartlett (1912)
regarded this a “useless and uncalled for violation of botanical tradition.”

When Stone used a combination he usually cited the basionym author parenthetically, but
L have noted at least 26 instances where Stone used a transfer name and cited the basionym
author without parentheses. Included on this list are five Stone innovations (Panicum
commonsianum addisonii, Blephariglottis x canbyi, Polygonum hydropiperoides opelousanum,
Ammodenia peploides maritima, Rubus villosus enslenit).

It could be argued that in the cases of transfers from specific to infraspecific rank, Stone
did not regard these changes as nomenclatural innovations, since he gave the specific and
subspecific rank coordinate nomenclatural status. However, there are numerous cases of
Stone citing the basionym author parenthetically when a change from specific to
infraspecific rank (or the inverse) was made, including nine Stone innovations (Paspalum
laeve circulare, Chaetochloa imberbis versicolor, Scleria reticularis torreyana, Dioscorea villosa
glabrifolia, Polygonum eciliatum, Lespedeza oblongifolia, Acer rubrum carolinianum, Euthamia
graminifolia nuttallii, Doellingeria umbellata humilis). Thus, Stone’s occasional failure to
parenthetically cite the basionym author for these combinations must be regarded as simple
errors.

Stone’s inconsistent usage of parentheses in author citations combined with other errors
of citation, such as citing only the transfer author in combinations (e.g., Mublenbergia foliosa,
Eragrostis pectinacea spectabilis, Veronica americana), using a basionym but citing its author
parenthetically (e.g., Carex interior capillacea, Oenothera biennis, Galinm lanceolatum), citing
two authors in two separate sets of parentheses (e.g., Sorghastrum nutans), not citing an
author at all (e.g., Carex varia emmonsit), and citing the wrong author (e.g., Quercus palustris,
Prunus maritima, Epilobum coloratum, Asclepias amplexicaulis, Utricularia cleistogama) make
his author citations difficult to follow. Also, the practice of citing only the basionym
author makes it difficult to know when Stone effected a transfer. Indeed, in the 19 cases
where Stone effected a nomenclatural innovation, there is no indication from his authorship
citation that he did so. Bartlett (1912) viewed this as “a practice which cannot be too
strongly condemned.” put)

Perhaps the most difficult nomenclatural issue in Stone’s work is his employment of
trinomials with no clear indication of rank (e.g., subspecies, variety, form). Most of the
nomenclatural innovations in The Plants of Southern New Jersey are trinomials. However,

30 BARTONIA

Stone when using trinomials used the terms “variety,” “subspecies,” and “form,” almost
interchangeably, thus making it difficult to ascertain at what rank the trinomials were
published. For example, under Osmunda cinnamomea Stone discussed “var. O. cinnamomea
glandulosa” and under Panicum commonsianum addisonii, he stated “it seems more reasonable
to regard . . . [it] as a sub-species rather than as full species.” He regarded Rhynchospora
glomerata leptocarpa as a “more slender form” and when discussing Viola palmata, V. palmata
triloba, and V. palmata sororia, he noted that “these three forms show all sorts of
intergrades.” In a paper on Viola, Stone (1903b) described but did not name an infraspecific
taxon of Viola papilionacea Pursh and used the designation “subsp.” after the specific epithet.

In Stone and Cram’s (1904) popular guide to the mammals of North America, the
trinomials employed were referred to as “varieties.”

Stone’s general views on infraspecific rank can best be gleaned in the introductory
comments in his work on Viola (1903b) where he stated:

- .. the point that I would particularly emphasize is that we have in these forms, which grow

often side by side, just as clearly differentiated races as the geographical sub-species of vertebrates,

and that they can be designated by trinomials to better advantage than by the binomial method
now generally in vogue, even though the grouping be admittedly arbitrary in some instances.

The careless use of the trinomial or varietal name in botany for all sorts of variation, purely

individual, albinistic, etc., of course acts as a prejudice against applying it to well-established

racial forms of this kind, but with the tremendous increase in species that we have currently

Witnessed in some genera the benefits of the trinomial system should be apparent. Unless we

are thoroughly familiar with the genus, it is impossible to tell in a strictly binomial system

which forms are clearly defined species and which are slightly differentiated races of a well-

known type, while the use of the trinomials indicates this at a glance.
Stone (1903c) took a similar position on the use of trinomials in an article in Condor.
Stone’s views on this subject are in close agreement with those put forth in the American
Ornithologists’ Union Code (Coues et al. 1886), the last American zoological code of
nomenclature before the establishment of the International Commission on Zoological
Nomenclature (see Melville 1995), which criticized the binomial system and advocated the
use of “trinomina” for organisms “known now to intergrade in physical characters” but not
for “those slightly distinct and scarcely stable forms. . .”

In The Plants of Southern New Jersey, Stone commented that “one authority, the authority
of the specific or subspecific name has been given.” However, most of the trinomial authors
cited by Stone published their names at the rank of variety. The fact that Stone in The
Plants of Southern New Jersey never consistently used one term to represent his infraspecific
rank and used the terms subspecies, variety, and form seemingly interchangeably when he
discussed trinomials makes it impossible to assign an infraspecific rank to them.

This has resulted in subsequent works inconsistently citing Stone’s trinomial innovations
as subspecies, varieties, or unranked infraspecific taxa. For example, in the International
Plant Names Index (The Plant Names Project 1999), the trinomial Paspalum laeve circulare

ash) W. Stone was simultaneously listed as being both a varietal and subspecific name.
U.S.D.A. (2000) regarded Paspalum laeve circulare (Nash) W. Stone, Polygonum hydropiper-
oides opelousanum (Riddell ex Riddell) W. Stone, and Euthamia graminifolia nuttallii (Greene)
W. Stone as having been published as varieties but treated Carex leptalea harperi (Fernald)
W. Stone as a subspecies. Hitchcock and Chase (1951) treated Stone’s three grass
innovations — Chaetochloa imberbis versicolor, Panicum commonsianum addisonii, Paspalum
laeve circulare — as having been published as an unranked trinomial, a subspecies, and a
variety, respectively.

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 31

I have concluded that under the current botanical code (Greuter et al. 2000), Stone’s
nomenclatural innovations involving trinomials should be regarded as validly published
unranked names (Article 35.3). As such, these names are inoperative in questions of priority
except homonymy (Article 53.4). The lack of a clear indication of rank does not invalidate
Stone’s trinomials since they were published prior to 1953 (Article 35.1).

It could be argued (see Voss 1966) that the trinomials published in Stone’s work should
be regarded as subspecies, since this is the term that he used in the introductory comments
of his work (Voss 1966, p. 35). However, this would, in effect, force one to regard all
previously published varietal names recognized in Stone’s work as trinomials as being
transfers (from variety to subspecies). This would greatly increase the number of
nomenclatural innovations in Stone’s work, since all 60 trinomials would have to be
regarded as innovations. Current nomenclatural indices (e.g., the International Plant Names
Index) do not treat all of Stone’s trinomials as innovations, but rather regard as innovations
only those in which Stone was the first to publish the name as a trinomial.

Regarding Stone’s trinomials as subspecific names would also contradict his statement in
the preface, “the names given in Britton’s Manual have been adopted except where changes
have been suggested in subsequent publications.” If Stone had intended for his trinomials
to function as transfers from the rank of variety to subspecies, then in those cases where he
used a trinomial to represent a varietal name that was also a basionym, he should have cited
the basionym author parenthetically; however, he did not (e.g., Panicum virgatum var.
cubense, Panicum columbianum var. thinium, Cyperus filiculmis var. macilentus, Carex
umbellata var. tonsa). This seems to have been the practice of botanists working under the
American Code, such as Pennell (1935) who commented: “In now adopting ‘subspecies’ for
precisely the concept that has often been called ‘variety,’ I wish it clearly understood that
only a superficial change of name is involved and that I do not consider the trinomials
appearing in this study to be new combinations.” However, unlike Pennell (1935), Stone
did not consistently use the term “subspecies” when referring to his trinomials.

Also, recognizing the trinomials as subspecies in some cases (see later discussions under
Carex umbellata abdita and Ammodenia peploides maritima) will result in a variety including
the type of the correct name of a subspecies, but not including the type of the correct name
of the species, having a different epithet than the subspecies. This is contrary to
Recommendation 26A.1 of the current botanical code (Greuter et al. 2000).

It should be noted that the International Plant Names Index (hereafter abbreviated as
LP.N.I) indicated that Stone published the combination “Utricularia inflata var. radiata” in
his Plants of Southern New Jersey. However, while Stone printed that combination (see p.
693) he did not accept it as a distinct taxon and thus under the current botanical code
(Greuter et al. 2000) the name is not validly published (Article 34.1).

NOMENCLATURAL INNOVATIONS

For each of Stone’s 19 nomenclatural innovations a review of all pertinent original
literature was made and a discussion on each innovation is provided below. A complete list
of nomenclatural (i.e., homotypic or objective) synonyms is also provided for each
innovation. With regards to trinomials, only those for which Stone was the first to publish
the name as a trinomial are considered as innovations. Abbreviations of books, periodicals,
and authors of botanical names follow Stafleu and Cowan (1976-1988), Lawrence et al. (1968)
and Brummitt and Powell (1992), respectively.

32 BARTONIA

01. Paspalum laeve [unranked] circulare (Nash) W. Stone, Pl. So. New Jersey 187. 1912a.

Paspalum circulare Nash in Man. FI. N. States (ed. 1): 73. 1901

P. laeve var. circulare (Nash) Fernald in Rhodora 36: 22. 1934.

Stone was the first author to transfer the name to the infraspecific level and Fernald
(1934b) was the first to recognize this taxon at varietal rank. In 1934 (and again in 1935),
Fernald did not indicate that he was effecting a transfer and attributed the name at the
varietal level to Stone. However, Fernald (1950) later cited himself as the transfer author.

Hitchcock and Chase (1951), U.S.D.A. (1982), and Kartesz (1994) recognized this taxon
as a variety of Paspalum laeve. Gleason and Cronquist (1991) and U.S.D.A. (2000) regarded
P. circulare as a synonym of P. laeve Michx. Hitchcock and Chase (1951), Kartesz (1994),
and U.S.D.A. (2000) incorrectly regarded Stone as the transfer author for the name P. /aeve
var. circulare, while U.S.D.A. (1982) regarded Fernald as the transfer author of this name.

02. Panicum commonsianum [unranked] addisonii (Nash) W. Stone loc. cit. 205.
Panicum addisonii Nash in Bull. Torrey Bot. Club 25(2): 83. Feb. 1898.
P. commonsianum var. addisonii (Nash) Fernald in Rhodora 37: 391. 1935; Pohl in Amer. Midl.
Naturalist 38: 582. 1947 (isonym).

P. ovale var. addisonti (Nash) C.F.Reed in Phytologia 67(6): 452. 1989.

Dichanthelium ovale var. addisonii (Nash) Gould and C.A.Clark in Ann. Missouri Bot. Gard. 65:
1115. 1978.

Panicum addisonii Nash and P. commonsianum Ashe are two Dichanthelium-type panic
grasses that occur in the eastern United States primarily in sandy soils on the Coastal Plain.
In Ashe’s (1898) treatment, P. addisonii and P. commonsianum were not close, being the 24th
and 60th taxa, respectively. Despite their wide separation, Ashe’s original description of P.
commonsianum closely matched his description and Nash’s (1898) original description of P.
addisonii. In Ashe’s key, the two taxa were separated by “leaves scattered on the stem, all
except the lowest narrowed to the base” for P. addisonii vs. “leaves crowded near the base
of the stem, upper not reduced” for P. commonsianum. Both names were typified by
material from Cape May County, New Jersey (P. addisonii: E.P. Bicknell s.n., 30-31 May
1897, Wildwood [NY]; P. commonsianum: A. Commons 34, June 1898, Cape May. [NCU).

Subsequent works by Hitchcock and Chase (1910, 1951) and Hitchcock (1935) placed
Panicum addisonii and P. commonsianum in the species-group Columbiana. The taxa were
distinguished by spikelet size (2.0 mm.-2.1 mm. in P. addisonii vs. 2.2-2.4 mm. in Fi
commonsianum) and panicle characters (dense with ascending branches in P. addisonit VS.
open with spreading branches in P. commonsianum). Unlike Ashe’s treatment, the two taxa
were treated as sister species and it was noted that P. addisonii closely resembled P.
commonsianum. Other works during this time (Robinson and Fernald 1908; Britton and
Brown 1913a; Deam 1929; Small 1933) also recognized the two taxa as distinct species.

Witmer Stone (1912a) was the first to regard Panicum addisonii and P. commonsianum as
conspecific (but distinct at the infraspecific level) under P. commonsianum. Stone’s use of
P. commonsianum in this sense is incorrect because P. addisonii Nash (published Feb. 1898)
has priority over P. commonsianum Ashe (in J. Elisha Mitchell Sci. Soc. 15: 55. Nov. 1898).
Fernald (1935) perpetuated Stone’s error by effecting the incorrect combination p
commonsianum var. addisonii (Nash) Fernald. The combination that Fernald should have
published is “P. addisonii var. commonsianum”; this combination has yet to be effected.

In 1974, Gould elevated Panicum subgenus Dichanthelium Hitchcock and Chase to generic
rank. Gould and Clark (1978) later recognized P. addisonii and P. commonsianum 2%
convarietal under Dichanthelium ovale (Elliott) Gould & C.A.Clark. Gould and Clark (1978)

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 33

recognized that P. commonsianum var. addisonii (Nash) Fernald was the earliest published
varietal name but they did not indicate that Fernald’s combination was incorrect. And
working under the Leningrad Code’s (Stafleu et al. 1978) Article 26 (autonyms — in this case
P. commonsianum var. commonsianum — “are not to be taken into consideration for
purposes of priority”), the combination D. ovale var. addisonti (Nash) Gould & C.A.Clark
was effected. D. ovale var. addisonii was used by Kartesz and Kartesz (1980), Clewell (1985),
Kartesz (1994), and U.S.D.A. (2000). Reed (1989) adopted Gould and Clark’s species and
varietal concepts but did not recognize the genus Dichanthelium and effected the transfer P.
ovale var. addisonii (Nash) C.F.Reed.

Interestingly, the name Dichanthelium ovale var. addisonii (and Panicum ovale var.
addisonii) is correct under the current botanical code (Greuter et al. 2000). The incorrect
assumption that P. commonsianum has priority over P. addisonii and the change in how
autonyms are treated with regards to priority (under the current code they are to be taken
into consideration) effectively cancel each other out.

However, all is not well regarding current usage since some currently used works do not
recognize Dichanthelium as a distinct genus and did not follow Gould and Clark’s (1978)
species and varietal concepts. These works regard Panicum addisonii and P. commonsianum
to be conspecific under P. commonsianum (Radford et al. 1968; Gleason and Cronquist 1991)
or distinct varietally under the names P. commonsianum var. commonsianum and P.
commonsianum var. addisonii (Fernald 1950; Gleason 1952a; Gleason and Cronquist 1963;
U.S.D.A. 1982; Ownbey and Morley 1991). Barring conservation of P. commonsianum over
P. addisonii, P. commonsianum would have to be replaced by P. addisonii and a new
combination (“P. addisonii var. commonsianum”) would have to be effected if the two taxa
were to be considered distinct varietally under P. addisonit.*

However, conserving Panicum commonsianum over P. addisonii would force a disadvanta-

eous nomenclatural change for those who currently follow the taxonomy of Gould and
Clark (1978) since, under the current autonym rule (Article 11.6), the name Dichanthelium
ovale var. addisonii would be incorrect and a new combination would be needed. Thus, as
long as there remain differences of opinion regarding the taxonomic rank and position of
P. addisonii and P. commonsianum a name change will need to be made for solely nomencla-
tural reasons, regardless of whether or not P. commonsianum is conserved over P. addisonit.
This argues against formal conservation of P. commonsianum.

In the LP.NJ. under the entry of “Panicum commonsianum subsp. addisonit (Nash) Stone,”
it was noted that “[e]Jven though Stone did not state the rank of his infraspecific taxon, his
discussion on the status of P. commonsianum and P. addisonii (p. 206, paragraph 1) indicates
the rank to be subspecific.” While Stone did use the term “sub-species” in his discussion of
P. commonsianum var. addisonii, he (as discussed earlier) also used the terms “variety” and

>Of course not updating innovations effected prior to the Sydney Code (Meikle et al. 1983) has also resulted in
names in current use being incorrect. An example is Dichanthelium sabulorum var. patulum (Scribner & Merrill)
Gould & C.A. Clark in Ann. Missouri Bot. Gard. 65: 113. 1978. This name was circumscribed by Gould and Clark
to include Panicum nashianum Scribner var. nashianum and P. nashianum vat. patulum Scribner & Merrill and
hence is incorrect. However, Gould and Clark’s taxonomy and nomenclature have been adopted by Clewell (1985),
Kartesz (1994) and U.S.D.A. (2000).

‘The nomenclatural problem addressed in this paper is not relevant if the taxonomy of Lelong (1984) or Hansen
and Wunderlin (1988) is followed; Lelong recognized Panicum addisonii and P. commonsianum to be synonyms of
P. ovale var. pseudopubescens (Nash) Lelong and Hansen and Wunderlin recognized them as synonyms of
Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. acuminatum.

34 BARTONIA

“form” when discussing some of his other taxa with trinominal names. Thus, I regard his
use of the term “sub-species” in this case to be incidental. Other currently used works treat
P. commonsianum addisonii (Nash) W. Stone as having been published at the rank of
subspecies (Hitchcock and Chase 1951), variety (Kartesz 1994; U.S.D.A. 2000) or as an
unranked trinomial (Hithcock 1935)

03. Chaetochloa imberbis [unranked] versicolor (E.P. Bicknell) W. Stone loc. cit. 213.

Chaetochloa versicolor E.P. Bicknell in Bull. Torrey Bot. Club 25: 105. 1898.

Panicum versicolor (E.P. Bicknell) Nieuw]. in Amer. Midl. Naturalist 2: 64. 1911, non Doell 1877.

nom. illeg.

Chaetochloa Scribn. (in U.S.D.A. Div. Agrostol. Bull. 4: 38. 1897) is now recognized under
Setaria P.Beauv. (in Ess. Agrostogr.: 51. 1812), which is conserved over Setaria Ach. ex
Michx. (in Fl. Bor.-Amer. 2: 331. 1803). Bicknell’s basionym as well as the two combinations
based on it were not cited by Fernald (1950), U.S.D.A. (1982, 2000), Gleason and Cronquist
(1991) or Kartesz (1994). C. versicolor has never been transferred into Setaria.

Bicknell’s description most closely resembles that of Setaria geniculata (Lam.) P.Beauv.
recognized in Fernald (1950), U.S.D.A. (1982, 2000), Gleason and Cronquist (1991), and
Kartesz (1994). Hitchcock and Chase (1951) also regarded Chaetochloa versicolor and Stone’s
transfer based on it as synonyms for S. geniculata. Unlike Pasaplum laeve circulare and
Panicum commonsianum addisonii, Hitchcock and Chase (1951) did not regard Stone’s
innovation as having been published at the varietal or subspecific rank but cited C. imberbis
versicolor (E.P. Bicknell) W. Stone as an unranked trinomial in synonymy under 5S.
geniculata. No currently used work (e.g., Fernald 1950; U.S.D.A. 1982, 2000; Gleason and
Cronquist 1991; Kartesz 1994) cited Stone’s trinomial in synonymy.

04. Scleria minor W. Stone loc. cit. 283.
Scleria triglomerata var. gracilis Britton in Ann. New York Acad. Sci. 3: 230. 1885, non S. gracilis
Elliott 1824 nec Rich. 1792.

S. triglomerata var. minor Britton, Illus. Fl. (ed. 1) 1: 282. 1896. nom. illeg.

This taxon was originally named as Scleria triglomerata var. gracilis. However, upon
learning that S. gracilis had already been published by Elliott in 1824 (which itself is a later
homonym of S. gracilis Rich.), Britton published S. triglomerata var. minor. Britton’s name
is illegitimate, since it was nomenclaturally superfluous (the name S. triglomerata var. gracilis
was available) when published (Article oS

Britton (1913a) discussed nomenclature in the introduction of vol. 1 of the I//ustrated Flora
and provided 10 basic rules of nomenclature which were said to be derived from “the Co
of Nomenclature devised by the Paris Botanical Congress in 1867, as modified by the rules

Sciences at the meetings held in Rochester, New York, in August 1892 and at Madison,
Wisconsin in August, 1893.” Rule 4 stated, “the original name is to be maintained, whether
published as a species, subspecies, or variety.” However, the Brittonian Code (Arthur et al.,
1907) dropped this Provision and names no longer had priority outside of their own rank.

When Stone recognized this taxon at specific rank, he used the name Scleria minor and
he indicated that he was publishing a transfer of Britton’s name by citing it as “Scleria minor
(Britton).” However, Stone’s innovation was not a new combination but rather a nomen
novum. S. minor is a replaced synonym for, and is typified by, S. triglomerata var. gracilis.
Therefore, the proper citation is S. minor W. Stone and not S$. minor (Britton) W. Stone as
in Fernald (1950) and U.S.D.A. (1982). Gleason and Cronquist (1991) also recognized this

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 35

taxon as a species and correctly cited Stone as the author, but they incorrectly indicated that
this taxon would be “perhaps better treated as S. triglomerata var. minor Britton.” The
correct name for this taxon when treated as a variety of S. triglomerata Michx. is S.
triglomerata var. gracilis Britton.

05. Scleria reticularis [unranked] torreyana W. Stone loc. cit. 283.

Scleria laxa Torr. in Ann. Lyceum Nat. Hist. New York 3: 443. 1836, non R.Br. 1810. nom. illeg.

S. mublenbergii’ Steud., Nomencl. Bot. (ed. 2): 543. 1841 (nom. nov. for S. laxa Torr.).

S. torreyana Walp. in Ann. Bot. Syst. 3: 696. 1852-53 (nom. nov. for S. laxa Torr.). nom. illeg.

Torrey (1836) was the first to describe this taxon but his name is illegitimate because it
is a later homonym of Scleria laxa R.Br. Steudel (1841) was the first to rename this taxon
when he published S. muhlenbergii. Walpers also renamed Torrey’s S. axa but his name is
illegitimate because Steudel had already published an avowed substitute for S. /axa Torr.

Stone adopted Walpers’ name at the infraspecific level, legitimizing the name at this level
in a fashion similar to the situation with Scleria minor W. Stone. (Article 58.1). However,
no one has ever transferred this name to a distinct infraspecific rank.

The taxonomy of the Scleria reticularis complex is controversial. When Torrey first
described his S. laxa he distinguished it from S. reticularis on the basis of S. laxa’s possessing
pitted achenes “marked in a somewhat spiral manner with transverse hairy rugae.” Britton
added another hairy-fruited Scleria with his description of S. reticularis var. pubescens Britton
(in Ann. New York Acad. Sci. 3: 232. 1884) and commented that it frequently grew with S.
laxa Torr. from which it could be distinguished “by the absence of transverse or spiral
ridges.” Thus there are three morphological types of concern: 1) typical S. reticularis with
glabrous achenes; 2) S. mublenbergii (=S. laxa Torr.) with pitted achenes marked in a spiral
to transverse manner with hairy rugae; and 3) S. reticularis var. pubescens, which like S.
muhlenbergii has pubescent achenes but lacks the spiral or transverse ridges.

Stone did not regard the two hairy-fruited types to be taxonomically distinct, citing Scleria
reticularis var. pubescens as a synonym of S. reticularis torreyana. Rather than use Britton’s
varietal name, he transferred Walpers’ name because (as discussed earlier) he believed that
earlier specific epithets should displace later varietal epithets.

Current usage is divided on whether to recognize the hairy-fruited material as distinct
from the glabrous-fruited material. Like Stone, no later work followed Britton in
recognizing the two hairy-fruited types (i.e., Scleria mublenbergii, S. reticularis var. pubescens)
as taxonomically distinct. Fernald (1950), Kartesz (1994), and U.S.D.A. (2000) recognized
S. mublenbergii and cited S. reticularis var. pubescens as a synonym. U.S.D.A. (1982) and
Fairey (1967) treated S. muhlenbergii as a synonym of S. reticularis var. pubescens. Gleason
and Cronquist (1991) did not regard the hairy-fruited and glabrous-fruited types as
taxonomically distinct and listed S. mubhlenbergii and S. reticularis var. pubescens as taxonomic
synonyms of S. reticularis. None of these works cited S. reticularis torreyana W. Stone in
synonymy nor is it listed in the LP.NL.

*Steudel’s original spelling was Scleria mueblenbergii. However, Henry Muhlenberg’s father adopted the anglicized
form (Muhlenberg) of the original family name (Miihlenberg). All of Henry Muhlenberg’s letters were signed and
all of his articles (except those edited by Willdenow) appear J und he Henry M Pat Bs
Merrill and Hu 1949). Therefore, I have treated Steudel’s original spelling as an error to be corrected (Article 60.1).

36 BARTONIA

06. Carex varia [unranked] emmonsii (Dewey ex Torr. & A. Gray) W. Stone loc. cit.
304.

Carex davisii Dewey in Amer. J. Sci. 10: 279. 1826, non Schwein. & Torr. 1825. nom. illeg.

C. emmonsii — ex Torr. & A. Gray in Ann. Lyceum Nat. Hist. New York 3: 411. 1836 (nom.

nov. for C. davis

C. novae-angliae var. saan (Dewey ex Torr. & A. Gray) J.Carey ex A. Gray, Manual (ed. 1):

556. 1848.

C. lucorum var. emmonsii (Dewey ex Torr. & A. Gray) Chapm., Fl. South. U.S. (ed. 1): 539. 1860.

C. albicans var. emmonsii (Dewey ex Torr. & A. Gray) J.Rettig in Sida 14: 133. 1990.

This taxon was originally described as Carex davisii by Dewey but this name is
illegitimate because it is a later homonym of C. davisii Schwein & Torr. It was later
renamed C. emmonsii in Torrey’s (1836) Monograph of North American Cyperaceae. In
Torrey’s paper the name was attributed to Dewey but there was no accompanying statement
that he contributed in any way to the paper other than providing specimens for
examination. However, Torrey did acknowledge “most valuable aid” provided by Asa Gray
and stated “the Synopsis of North American Carices, I wish to have considered as our joint
performance.” Hence the proper citation should be Carex emmonsii Dewey ex Torr. & A.
Gray. More recent authors either cited the basionym authorship as “Dewey” (Fernald 1950,
U.S.D.A. 1982) or “Dewey ex Torr.” (Kartesz 1994, U.S.D.A. 2000).

Stone, like other early authors (e.g., Robinson and Fernald 1908), misapplied the name
Carex varia Muhl. ex Schkuhr to this taxon. Also, Stone did not cite any author after his
C. varia emmonsii trinomial and below this entry he incorrectly cited the basionym as
“Carex varia var. emmonsii ‘Dewey,” Torr.”

Rettig (1989) originally recognized Carex emmonsii, even though it was considered to be
conspecific with the earlier C. albicans Willd. ex Spreng. (Syst. Veg. 3: 818. 1826). C. albicans
was not used because it had been “ ‘widely and persistently’ used for the taxon now known
as C. emmonsii var. emmonsii,” even though the type of C. albicans “belongs to C. emmonsi
var. mublenbergii.” Rettig never proposed to formally reject this name and later (1990)
adopted it and recognized C. emmonsii as a variety of C. albicans. Currently used works
either recognized this taxon as a species (Fernald 1950, U.S.D.A. 1982) or as a variety of C.
albicans (Gleason and Cronquist 1991, Kartesz 1994, U.S.D.A. 2000). None of these works
nor the LP.NJ. cited C. varia emmonsii (Dewey ex Torr. & A. Gray) W. Stone in
synonymy.

07. Carex umbellata [unranked] abdita (E.P. Bicknell) W. Stone loc. cit. 305.
Carex umbellata var. brevirostris Boott, Ilustr. Carex 2: 99. 1860, non C. brevirostris Cedestr. 1857
nec Blytt 1857.
C. umbellata var. brachyrhina Piper in Contr. U.S. Nat. Herb. 11: 175. 1906 (nom. nov. for ©.
umbellata var. brevirostris). nom. illeg.
C. abdita E.P. Bicknell in Bull. Torrey Bot. Club 25: 492. 1908 (nom. et stat. nov. for C. umbellata
var. brevirostris).

Bicknell in his discussion of Carex umbellata Schkuhr var. umbellata for Nantucket stated:
This plant agrees so well with Schkuhr’s illustration of Carex umbellata that there can be little
doubt that it is definitely typical, although a form with longer-beaked more pubescent perigynia

‘In The Plants of Southern New Jersey, Stone used quotation marks around an author’s name to indicate that this
author had a plant name ascribed to him by another author.

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 37

would seem to have been commonly understood in that sense. The plant here in view,
notwithstanding its comparatively short beak is not the var. brevirostris Boott. The latter plant
or one which must be referred to it on the basis of descriptions, is frequent on Long Island,
N.Y., where the typical plant seems to be rare, the prevailing form being the var. tonsa Fernald.
The var. brevirostris often occurs with the latter and differs from it constantly in definite
characters. It commonly forms larger, closer tufts and has much longer, narrower, more erect
and less rigid leaves and more slender culms, some of which are capillary and elongated and bear
a slender-pedicelled, bracteate fertile spike near the base of the staminate one; the perigynia are
mostly pubescent and only 2.5-3 mm long, with the short beak only 0.5-1 mm., the achene pale
brown, rather shining and about 1.5 mm. long by 1 mm. thick. I have long regarded this plant
as a distinct species and have deposited specimens from Richmond Hill, Long Island in the
herbarium of the N.Y. Botanical Garden labeled Carex abdita.

Based on the passage above, I have concluded that Carex abdita was published as a nomen
et status novon for C. umbellata var. brevirostris Boott (non C. brevirostris Cedestr.). This
conclusion is based on Bicknell’s statement that, based on descriptions, the material in
question from Long Island must be referred to C. umbellata var. brevirostris and the fourth
sentence in the above passage which links the name C. umbellata var. brevirostris to the
description. However, since Bicknell stated that he had regarded this plant to be a distinct
species, it could be argued that C. abdita was published as a species novum (typified by the
Richmond Hill, Long Island material).

Stone incorrectly indicated that Bicknell had published the name as Carex umbellata var.
abdita. Most recent works (U.S.D.A. 1982, 2000; Gleason and Cronquist 1991; Kartesz 1994)
regard Carex abdita as a taxonomic synonym of C. umbellata. Fernald (1950) recognized C.
abdita as a distinct species. None of these works cited C. umbellata abdita (E.P. Bicknell)
W. Stone in synonymy. The LP.N.I listed Stone’s trinomial as having been published at the
rank of subspecies.

This case represents an example of how recognizing Stone’s trinomials as subspecies could
have the disadvantageous effect of a varietal name including the type of the correct name of
a subspecies, but not including the type of the correct name of the species, having a different
epithet than the subspecies name. If Stone’s trinomials were regarded as subspecies then
the correct name for this taxon at the rank of subspecies would be Carex umbellata subsp.
abdita (E.P. Bicknell) W. Stone. However, the correct name for the same taxon at the rank
of variety would be C. umbellata var. brevirostris Boott. Such a situation (i.e., the name of
the variety is different from the name of the subspecies even though the name of the variety
is typified by the type of the subspecies but not the type of the correct name of the species)
is possible under the current botanical code (Greuter et al. 2000) but is contrary to
Recommendation 26A.1.

08. Carex leptalea [unranked] harperi (Fernald) W. Stone loc. cit. 305.

Carex harperi Fernald in Rhodora 8: 181. 1

C. leptalea var. harperi (Fernald) Weath. & Griscom in Rhodora 36: 39. 1934.

C. leptalea subsp. harperi (Fernald) Cald. & Tayl. in Canad. J. Bot. 43: 1391. 1965.

This taxon was originally described by Fernald (1906), who commented that the plant was
similar to Carex leptalea, but could be distinguished on the basis of its perigynia (longer and
more strongly overlapping), scales (whitish and acuminate), and achenes (puncticulate,
sharply angled). Currently used works treated Carex harperi as a taxonomic synonym of C.
leptalea (U.S.D.A. 1982; Gleason and Cronquist 1991) or as a variety (Fernald 1950) or
subspecies of C. leptalea (Kartesz 1994; U.S.D.A. 2000). Recognizing this taxon as a variety

38 BARTONIA

of C. leptalea, Fernald (1950) attributed the transfer to Stone. Kartesz (1994) and U.S.D.A.
(2000) recognized this taxon at subspecific rank and also attributed the transfer at this rank
to Stone. U.S.D.A. (1982) treated this name as a synonym of Carex leptalea and cited “Carex
leptalea var. harperi (Fernald) Fernald ex W. Stone” as a synonym. This exemplifies the
undesirable result that can occur when the basionym author is cited but not parenthetically.
In this case Stone published “C. leptalea harperi Fernald.” This was later interpreted by
U.S.D.A. (1982) to indicate that Stone was ascribing the transfer to Fernald, when all he was
actually doing was citing the basionym author. Bartlett (1912) regarded this practice as
“falsely branding” the basionym author as the author of a new combination. This issue
indicates a potential problem with adopting the zoological practice of author citation when
parentheses are inadvertently left out. It also highlights how this practice would allow
transfers to be effected in an ambiguous fashion.

Since Stone cannot be regarded as having published the name at the varietal level the
question of course is who was the first person to recognize this taxon as a variety and hence
effect the transfer. This appears to be Weatherby and Griscom (1934; see Voss 1966), who
incorrectly ascribed the transfer to Stone. Thus, the correct citation should be Carex leptalea
var. harperi (Fernald) Weatherby & Griscom. The 1 P.N.J. treated Stone’s trinomial as having
been published at the rank of subspecies.

09. Dioscorea villosa [unranked] glabrifolia (Bartlett) W. Stone loc. cit. 358.

Dioscorea paniculata var. glabrifolia Bartlett in U.S.D.A. Bur. Pl. Industr. Bull. 189: 1915. 1910.

D. villosa var. glabrifolia (Bartlett) Blake in Rhodora 20: 49. 1918.

D. villosa forma glabrifolia (Bartlett) Fernald in Rhodora 39: 401. 1937.

Bartlett (1910) displaced the name Dioscorea villosa L. with D. paniculata Michx. on the
grounds that D. villosa L. was being misapplied and that the earliest name available for this
taxon was D. paniculata Michx. However, Stone did not accept Bartlett’s interpretation and
transferred Bartlett’s name as D. villosa glabrifolia. Blake (1918) also did not accept Bartlett's
views on this matter and recognized Bartlett’s name as a variety of D. villosa. Fernald (1937)
later recognized this taxon as a form of D. villosa.

Two later works (U.S.D.A. 1982; Gleason and Cronquist 1991) did not recognize
Bartlett’s name at any level and did not even cite his name in synonymy. Bartlett’s plant
would key to Dtoscorea villosa in Gleason and Cronquist. Fernald (1950) continued to
recognize this taxon as a form of D. villosa. Kartesz (1994) and U.S.D.A. (2000) recognized
this name as a synonym of D. quaternata J.F.Gmel. None of these works cited D. villosa
glabrifolia (Bartlett) W. Stone in synonymy. The LP.N.I. listed Stone’s trinomial as having
been published at the rank of subspecies.

10. Blephariglottis x canbyi (Ames) W. Stone loc. cit. 368.

Habenaria canbyi Ames in Rhodora 10: 70. 1908.

Platanthera xcanbyi (Ames) Luer, Native Orchids Florida: 151. 1972.

Stone made this combination in the discussion under Blephariglottis blephariglottis. This
nothotaxon is the result of crossing between Habenaria blephariglottis (Willd.) Hook. and H.
cristata (Michx.) R.Br. In all recent works the fringed orchids were placed in either
Habenaria Willd. or Platanthera Rich.

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 39

11. Gyrostachys beckii (Lind.) W. Stone loc. cit. 375.

Spiranthes beckii Lindl., Gen. Sp. Orchid. PL: 472. 1840.

Ibidium beckii (Lindl.) House in Muhlenbergia 1: 128. 1906. nom. invalid.

Triorchis beckii (Lindl.) House in Amer. Midl. Naturalist 6: 206. 1920. nom. invalid.

The transfers listed above were not due to disagreements over generic level taxonomy but
as to what the earliest published name was for this genus of plants. The name Jbidium was
first published as a nomen nudum by Salisbury (in Trans. Hort. Soc. London 1: 291. 1812).
This name was later validated with a description by Small (Fl. Southeast U.S. (ed. 2): 318.
1913). Before Small published his validating description, House made “transfers” into
Ibidium, including Lindley’s name. However, these “combinations” are not valid since they
were published before Small’s validating description. Also, House’s (1920) publication of
the name “Triorchis beckii” is not valid because Triorchis Petiver is pre-Linnaean and hence
invalid.

The genus name Gyrostachys was first published (as Gyrostachis) by Persoon (Syn. Pl. 2:
511. 1809) but it was invalid at the time of publication because it was not accepted by
Persoon. Blume (Coll. Orchid.: 127. 1859; Fl. Javae Nov. Ser.: 107. 1859) later validated this
name by accepting it as a genus. However, by this time the name Spiranthes had been
validly published by L.C. Richard (De Orchid Eur.: 20, 28, 36. 1837).

Spiranthes is conserved over the nomenclatural synonym Orchiastrum Ség. (Pl. Veron. 3:
252. 1754). It is not conserved over Aristotelea, Ibidium, Gyrostachys, and Helictonia Ehrh.
Aristotelea Lour. (Fl. Cochinch.: 522. 1790) is an earlier name and is homotypic with
Spiranthes. However, Aristotelea Lour. should be regarded as a parahomonym (Article 53.3)
of Aristotelia L’Her. (Stirp. Nov. 31: 16. 1785), which is conserved over Aristotela Adans.
(Fam. 2: 125. 1763). The valid publication date of Spiranthes predates the valid publication
dates of Ibidium and Gyrostachys; Helictonia Ehrh. is invalid.

The taxonomic limits of this taxon have been controversial as have been the application
of names in Spiranthes. The taxon in question has been recognized under three different
basionyms and combinations based on them: S. simplex A. Gray (1867), non Grisebach 1864
(=S. grayi Ames), S. beckii Lind. and S. tuberosa Raf. The recognition of this taxon under
S. beckii (and combinations based on it) was due to the misapplication of this name to
material now considered to be conspecific with S. gracilis (Bigelow) Beck (Correll 1950, Luer
1975). Gray described S. simplex (Manual (ed. 5): 506. 1867) for this taxon believing that S.
tuberosa Raf. also applied to S. gracilis.

Ames (1947) regarded Spiranthes tuberosa to be comprised of more than one species and,
in view of the lack of a type specimen, chose to abandon it and use S. gray: and S. gracilis
var. floridana (Wherry) Correll for the two elements involved. Fernald (1946) disagreed
stating, “as to the identity of Spiranthes tuberosa Raf. . . . there is certainly no doubt” and
applied the name to the only small-flowered taxon in the eastern United States that possesses
atuber. Correll (1950) followed Ames’ views on application and recognized this taxon as
Spiranthes grayi. All other currently used works followed Fernald’s views (Fernald 1950;
Luer 1975; U.S.D.A. 1982, 2000; Gleason and Cronquist 1991; Kartesz 1994).

12. Polygonum hydropiperoides [unranked] opelousanum (Riddell) W. Stone loc. cit. 422.
Polygonum opelousanum Riddell, New Orleans Med. Surg. J. 9: 611. 1853 (as nom. nud. by Riddell
in New Orleans Med. Surg. J. 8: 760. 1852).
Polygonum hydropiperoides var. opelousanum (Riddell) Gleason, Illus. Fl. vol. 2: 82. 1952.
Persicaria opelousana (Riddell) Small, Fl. Southeast. U.S. (ed. 1): 378. 1903.

40 BARTONIA

This epithet was originally published by Riddell as a nomen nudum in 1852. Riddell
validated his name with a description in 1853. This second publication is not cited by the
major nomenclatural indices such as the /.P.N.J. Thus, some Riddell names in these indices
are given incorrect author citations. For example in LP.N.J. the citation for this Riddell
name is Polygonum opelousanum Riddell ex Small. In light of this overlooked publication,
Riddell entries in these indices need to be reviewed and updated.

Stone was the first to recognize this taxon at the infraspecific level under Polygonum
hydropiperoides. Gleason (1952b) was the first to recognize this taxon as a variety and cited
Stone as the transfer author at this rank.

Fernald (1950) recognized Polygonum opelousanum. Gleason and Cronquist (1991)
recognized this taxon as a variety of P. hydropiperoides and cited Stone as the transfer author.
U.S.D.A. (1982, 2000) and Kartesz (1994) treated P. opelousanum as a taxonomic synonym
of P. hydropiperoides and in synonymy cited “P. hydropiperoides var. opelousanum (Riddell ex
Small) Riddell ex W. Stone.” Besides incorrectly citing Stone as the transfer author at the
rank of variety, these works incorrectly cited Small as the validating author of the basionym
due to the overlooked status of Riddell’s 1853 publication (cited above). They also
incorrectly cited Riddell as an ascribed author of the transfer, because Stone cited Riddell
without parentheses after P. hydropiperoides opelousanum. This is another instance of the
“false branding” problem that Bartlett (1912) discussed. The I.P.N.I. lists Stone’s trinomial
as having been published at the rank of subspecies.

13. Polygonum eciliatum (Small) W. Stone loc. cit. 423.

Polygonum punctatum var. eciliatum Small in Bull. Torrey Bot. Club 20: 214. 1893.

Small distinguished this from typical Polygonum punctatum by its “glossy appearance
... perfectly smooth and eciliate ochreae, and the slightly narrower and more reticulated
achene.” In Stone’s key to the Polygonaceae, P. punctatum and P. eciliatum are separated
based on the ochreae (fringed with cilia vs. n :

In Stone’s discussion of this plant he stated, “This handsome Polygonum was first
discovered by Mr. Alexander McElwee 11 September 1892 at Wildwood. He proposed to
describe it as new, but for some reason did not do so and his manuscript description is sull
attached to the specimen at the Philadelphia Academy herbarium.” Stone went on to stale
that the “pink flowers are quite showy.” This is similar to Small’s statement in his original
description that “the color of the flowers and racemes .. . is very striking.” Further
evidence that Small recognized the material examined by Stone as belonging to his P.
punctatum var. eciliatum (holotype: Pringle 4213, Valley of Toluca, Mexico State, Mexico
[NY] is that he mentioned New Jersey in the range of this plant in his Flora of the
Southeastern United States (1933).

No works in current use recognize Small’s taxon, nor do they cite P. punctatum vat.
eciliatum Small or P. eciliatum (Small) W. Stone anywhere in synonymy. Small’s plant
closely matches the descriptions of Polygonum punctatum in Fernald (1950) and Gleason and
Cronquist (1991).

14. Ammodenia peploides [unranked] maritima (Raf.) W. Stone loc. cit. 441.
um maritimum Raf., New FI 1: 62. 1836.
Ammodenia maritima (Raf.) E.P. Bicknell in Bull. Torrey Bot. Club 37: 58. 1910. ;
Arenaria peploides var. robusta Fernald in Rhodora 11: 114. 1909 (nom. et stat. nov. for Adenarium
maritimum Raf.).

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 41

Honckenya peploides subsp. robusta (Fernald) MHultén, Fl. Aleut. Isl: 173. 1937.
Honkenya peploides var. robusta (Fernald) House in New York State Mus. Bull. 243/244: 56. 1923.
Minuartia peploides subsp. robusta (Fernald) Mattf. in Bot. Jahrb. Syst. 57: 27. 1921.

The name Ammodenia was not accompanied by a description when it was published by
J.G. Gmelin (ascribed to Patrin, Fl. Sibir. 4: 160. 1769) so it was an invalid name at the time
of Ehrhart’s publication of Honkenya (in Beitr. Naturk. 2: 180. 1788, non Willd. ex Cothen.
1790), which also predated Adenarium Raf. (in Amer. Monthly Mag. & Crit. Review 2: 266.
1818).

Honkenya Ehrh. (1788) was named in honor of Gerhard August Honckeny. This led
Bartling to change the spelling to Honckenya (in Ord. 305. 1830). Most recent works also
spelled the name Honckenya (Gleason and Cronquist 1991; Kartesz 1994; U.S.D.A. 2000).

However, before Bartling’s correction of Honkenya Ehrh to Honckenya, Willdenow had
published Honckenya (in Usteri, Del. Opusc. Bot. 2: 200. 1793) to replace Honkenya Willd.
ex Cothen. (Disp. 19. 1790; =Clappertonia Meisn. 1837) in the Tiliacae. In my opinion,
Honkenya Willd. ex Cothen. and all orthographic variants based on it (Honckenya;
Honckenia Pers., Syn. 1: 416. 1805; Honckeneya Steud., Nomencl. Bot. (ed. 2) 1: 774. 1840)
should be treated as later homonyms of Honkenya Ehrh. (Article 53.3).

Rafinesque was the first to distinguish this taxon from Arenaria peploides on the basis of
its entire leaves (vs. subserrate in A. peploides) and referred to this taxon as Adenarium
maritimum. This was later transferred into Ammodenia by E.P. Bicknell (1910).

Fernald (1909) chose not to regard these maritime plants as generically distinct from
Arenaria and regarded Adenarium maritimum as a variety of Arenaria peploides. However,
“since the name maritima has since been twice used for species in the genus Arenaria and
as a varietal designation would be quite meaningless for a local variation of a uniformly
maritime species,” Fernald introduced the name Arenaria peploides var. robusta.

Stone was aware of Fernald’s varietal name but used Rafinesque’s name in his transfer
because of his belief that specific and infraspecific names have coordinate nomenclatural
status. Fernald’s variety was transferred to Honkenya peploides by House (1923). It was also
later transferred to subspecific rank under H. peploides by Hulten (1937) and under Minuartia
peploides by Mattfeld (1921). More recent works either treated this taxon as H. peploides ssp.
robusta (U.S.D.A. 1982, 2000; Kartesz, 1994), H. peploides var. robusta (Gleason and
Cronquist 1991), or Avenaria peploides var. robusta (Fernald 1950). None of these works
cited Ammodenia peploides maritima (Raf.) W. Stone in synonymy. The LP.N.L. listed
Stone’s trinomial as having been published at the rank of subspecies. te ©

This is an example of the disadvantageous effect of recognizing Stone’s trinomials as
subspecies. If Ammodenia peploides maritima (Raf.) W. Stone is recognized as a subspecies
then the recently used names Honckenya_ peploides subsp. robusta (Fernald) Hulten and
Minuartia peploides subsp. robusta (Fernald) Mattf. are incorrect since Stone’s name would
have priority at the subspecific rank and new combinations in Honckenya and Minuartia
would have to be effected. The earliest name at the rank of variety would be Arenaria
peploides var. robusta. Thus, like the Carex umbellata abdita example discussed earlier, this
is a case where recognizing Stone’s trinomials as subspecies would have the disadvantageous
effect of a varietal name including the type of the correct name of a subspecies, but not
including the type of the correct name of the species, having a different epithet than the

subspecies name.

42 BARTONIA

15. Rubus villosus [unranked] enslenii (Tratt.) W. Stone loc. cit. 480.

Rubus enslenii Tratt., Rosac. Monog. 3: 63. 1823

Stone misapplied the name Rubus villosus Aiton for a taxon that is properly known as
R. flagellaris Willd. (Bailey 1943) and he recognized R. enslenii at the infraspecific level under
this taxon. No other author has recognized R. enslenii at a distinct infraspecific rank under
either R. villosus or R. flagellaris. All current works either recognize R. enslenii as a distinct
species (Fernald 1950; U.S.D.A. 1982; Gleason and Cronquist 1991) or as a taxonomic
synonym of R. flagellaris (Kartesz 1994; U.S.D.A. 2000). None of these works cited Stone’s
trinomial in synonymy. Bailey (1943) recognized R. enslenii and cited Stone’s trinomial as
a synonym and recognized it as having been published at the rank of variety. The LP.NI
recognized Stone’s trinomial as having been published at the rank of subspecies.

16. Lespedeza oblongifolia (Britton) W. Stone loc. cit. 509.

Lespedeza hirta var. oblongifolia Britton in Trans. New York Acad. Sci. 12: 66. 1893.

When Britton described this taxon he regarded it as “very well-marked” and placed it
under Lespedeza hirta “on account of its long peduncles.” Stone regarded this taxon as a
species and distinguished it from L. hirta on the basis of its oblong leaflets. Stone stated that
this plant was “not common.” Neither author discussed any fruit characters.

Britton and Brown (1913b), under Lespedeza hirta stated “[a] plant known as Lespedeza
hirta oblongifolia may be a hybrid with L. angustifolia (Pursh) Ell. We do not know its
fruit.” Fernald (1941, 1950) regarded L. hirta var. longifolia (DC.) Torr. & A. Gray (Fl. N.
Am. 1:368. 1840.) and L. hirta var. oblongifolia to be convarietal under L. hirta var. longifolia.
All other currently used works (Gleason and Cronquist 1991; Clewell 1966; Kartesz 1994;
U.S.D.A. 1982, 2000) recognized L. longifolia DC. and L. oblongifolia to be distinct hybrids,
L. x longifolia (L. capitata x L. hirta) and L. x oblongifolia (L. angustifolia x L. hirta). In
the L.P.N.I. this name was listed as L. oblongifolia Stone; however, there seems to be no basis
for not listing Britton as the author of the basionym.

17. Acer rubrum [unranked] carolinianum (Walter) W. Stone loc. cit. 544.

Acer carolinianum Walter, Fl. Carol.: 250. 1788.

Rufacer carolinianum (Walter) Small, Fl. S.E. U.S.: 826. 1933.

Under Acer rubrum, Stone stated the following: “The Red Maples of our region are
puzzling in their variability. We have a form with very small three-lobed leaves, usually

rarely tomentose beneath which may be referable to true rubrum. ... The division of
specimens . . . is arbitrary, but the differences between the extremes is striking.” Stone’s A.
rubrum carolinianum with its small three-lobed leaves is undoubtedly equivalent to A.
rubrum var. tridens A.W.Wood and A. rubrum var. trilobum Torr. & A. Gray ex K.Koch.
Most currently used works (Fernald 1950; U.S.D.A. 1982, 2000; Gelderen et al. 1994; Kartesz
1994) recognized this taxon under the name A. rubrum var. trilobum. Gleason and
Cronquist (1991) regarded the A. rubrum complex as “. . . variable, but indivisible.” None
of these works cited Stone’s trinomial in synonymy. The /.P.N.I. listed Stone’s trinomial
as having been published at the rank of subspecies.

NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 43

18. Euthamia graminifolia [unranked] nuttallii (Greene) W. Stone loc. cit. 752.
Euthamia nuttallii Greene in Pittonia 5: 73. 1902.
E. graminifolia (Greene) var. nuttallii Sieren in Rhodora 83: 564. 1981.
Solidago graminifolia var. nuttallii (Greene) Fernald in Rhodora 10: 92. 1908.
S. nuttallii (Greene) Bush in Amer. Midl. Naturalist 5: 168. 1918. pro syn., nom. invalid.
TAXONOMIC SYNONYMS
Euthamia floribunda Greene in Pittonia 5: 74. 1902, non Solidago floribunda Phil. 1894.
E. graminifolia var. polycephala (Fernald) Moldenke in Revista Sudamer. Bot. 6: 29. 1939.
Solidago polycephala Fernald in Rhodora 10: 93. 1908 (nom. nov. for E. floribunda Greene).
S. graminifolia var. polycephala (Fernald) Fernald in Rhodora 17: 12. 1915.
S. floribunda (Greene) Bush in Am. Midl. Naturalist 5: 167. 1918, non Phil. 1894) pro syn., nom.
invalid.

Greene (1902) described two closely related taxa of Euthamia from the E. graminifolia
complex, E. floribunda and E. nuttallii. They were distinguished from typical E. graminifolia
on the basis of their pubescent leaves and inflorescence. E. floribunda was distinguished
from E. nuttallii on the basis of its tiny involucre (3-3.5 mm. long) and its appressed deltoid
green tips of the bracts.

Currently used works either treat these plants as Solidago L. (Fernald 1950) or retain them
in the segregate genus Euthamia Nutt. as interpreted by Strother 2000 (U.S.D.A. 1982, 2000;
Kartesz 1994; Gleason and Cronquist 1991). Fernald (1950) recognized E. floribunda and E.
nuttallii as varieties of S. graminifolia. U.S.D.A. (1982) regarded Greene’s two names as
synonyms of E. graminifolia. Gleason and Cronquist (1991) regarded “var. nuttallii” as a
hirtellous phase of E. graminifolia. Kartesz (1994) and U.S.D.A. (2000) recognized E.
floribunda and E. nuttallii as convarietal under E. graminifolia var. nuttallit. Gleason and
Cronquist (1991), Kartesz (1994), and U.S.D.A. (1982, 2000) incorrectly cited Stone’s
trinomial as having been published at the rank of variety. However, Stone did not effect
a transfer at the varietal rank and it appears that this combination (E. graminifolia var.
nuttallii) was first published by Sieren (1981; see also Voss 1996). The LP.N.L index listed
Stone’s trinomial as having been published at the rank of subspecies, although it incorrectly
ascribed the combination to Stone.

19. Doellingeria umbellata [unranked] humilis (Willd.) W. Stone loc. cit. 763.
Aster humilis Willd. in Sp. Pl. 3: 2038. 1804.
Aster umbellatus var. humilis (Willd.) Britton et al., Prelim. Catalogue New York, 27. 1888.
Doellingeria humilis (Willd.) Britton in Ilus. Fl. 3: 392. 1898. ,
Stone cited this plant as occurring in “open swamps 1n the Pine Barrens; not common.
In their review of Aster in the Willdenow Herbarium, Jones and Hiepko (198 1) stated “at
the rank of variety, the name A. humilis is to be placed in synonymy under A. umbellatus
var. latifolius A. Gray.” In currently used works, Stone's Doellingeria umbellata humilis is
referable to either A. umbellatus (Fernald 1950; Gleason and Cronquist 1991), A. umbellatus
var. latifolia (U.S.D.A. 1982; Kartesz 1994) or Doellingeria seriocarpoides Small (U.S.D.A.
2000). However, none of these work cited Stone’s trinomial in synonymy. The LP.N.L
listed Stone’s trinomial as having been published at the rank of subspecies.

A table has been prepared that provides full authorship citations for the names used in
The Plants of Southern New Jersey and compares Stone’s nomenclature with that in Fernald
(1950) and Gleason and Cronquist (1991), two commonly used field manuals for the
northeastern United States. Copies of it can be obtained by writing to the author.

44 BARTONIA
ACKNOWLEDGMENTS

I would like to thank Ted Gordon, Dan Nicolson, and David Snyder for reviewing earlier
drafts of this manuscript. Ernie Schuyler and Ed Voss are thanked for reviewing the
submitted manuscript. Tracy Delius is thanked for assistance in preparing the manuscript.
Financial support is acknowledged from the Brooklyn Botanic Garden.

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of Botanical Nomenclature Adopted by the Thirteenth International Botanical Congress, Sydney
August 1981. Regnum Vegetabile 111. 472 pp.

MELVILLE, R. V. 1995. Towards Stability in the Names of Animals; a History of the International
Commission on Zoological Nomenclature. International Trust for Zoological Nomenclature,
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MERRILL, E. D. AND S. HU. 1949. Work and publications of Henry Muhlenberg, with special
attention to unrecorded or incorrectly recorded binomials. Bartonia 25: 1-66.

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89.

NICOLSON, D. 1991. A history of botanical nomenclature. Annals of the Missouri Botanical Garden
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ORCHARD, A. E., W. R. ANDERSON, M. G. GILBERT, SEBSEBE D., W. T. STEARN, AND E. G. VOSS.
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Ownsry, G. B. AND T. MORLEY. 1991. Vascular Plants of Minnesota. University of Minnesota Press,
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PENNELL, F. W. 1940. The botanical works of Witmer Stone. Bartonia 20: 33-37.

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RETTIG, J. H. 1989. Nomenclatural changes in the Carex pensylvanica group (section Acrocystis,
Cyperaceae) of North America. Sida 13: 449-452.

RETTIG, J. H. 1990. Correct names for the varieties of Carex albicans/Carex emmonsii. Sida 14: 132-133.

RIDE, W. D. L., H. G. CoGGER, C. Duputs, O. KRAUS, A. MINELLI, F. C. THOMPSON, P. TUBBS,
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Nomenclature, London. 306 pp.

ROBINSON B. L. AND M. L. FERNALD. 1908. Gray’s New Manual of Botany (7th ed.). American Book
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994 pp.

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NOMENCLATURE OF STONE’S PLANTS OF SOUTHERN NEW JERSEY 47

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Agaricus subsect. Gymnopus fertilis
(sensu Schweinitz) [=Entoloma
olivacebrunneum (L.) Kumm.]

Boletus frondosus (sensu
Schweinitz) [= Grifola frondosa
(Dickson:Fr.) $.F. Gray]

Boletus fomentarius (sensu
Schweinitz) [= Fomes fomentarius
(L.:Fr.) Kickx.]

Bartonia No. 61: 48-53, 2002

Lewis David von Schweinitz’s Mycological Illustrations

DAVID HEWITT*
Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway,
Philadelphia PA 19103-1195
dhewitt@oeb.harvard.edu

ABSTRACT. The early nineteenth-century mycologist, Lewis David von Schweinitz, is known to
have produced seven bound volumes of paintings of fungi; four of these are at the Academy of Natural
Sciences of Philadelphia and one each are at the University of North Carolina at Chapel Hill,
University of Michigan, and the Farlow Herbarium at Harvard University. The volumes in
Philadelphia and Chapel Hill together form a single set; the volume in Michigan and the volume at
Harvard are not part of that set.

INTRODUCTION

Lewis David von Schweinitz (1780-1834), “the mycological father of us all” (Rogers 1977),
was the first American to focus his botanical efforts on fungi (Johnson 1835; Benedict 1935;
Pennell 1935; Rogers 1977). Born and educated in Bethlehem, P lvania,
lived and studied in Silesia, where, in 1805, he and Johannes Baptista von Albertini published
their seminal work, Conspectus Fungorum in Lusatiae superioris agro Nieskiensi Crescentium
e methodo Persooniana [Conspectus]. This 412-page volume contains descriptions and
illustrations of fungi from Lusatia, a region along the Neisse River in what is now eastern

rmany and western Poland, and includes the descriptions and illustrations of 93 new
species (Benedict 1935). Included in the Conspectus are 12 color plates, and on each of these,
except for plate XI, is written either “v. S.” or “L. v. S.,” indicating that Schweinitz prepared
the illustrations. Schweinitz returned to the United States in 1812, and for the rest. of his
life collected cryptogams and phanerogams, painting them and writing about them and
corresponding with the pantheon of botanists who lived and worked in early nineteenth-
century Europe and the United States. Upon Schweinitz’s death, his herbarium was
deposited at the Academy of Natural Sciences of Philadelphia (PH), forming the basis of its
nascent Botany Department, and much of his correspondence and manuscript materials have
since been deposited in the archives at the Academy. ee cs ee

Schweinitz was a prodigious illustrator, and bound volumes of his paintings of fungi, his
Icones Fungorum, have been deposited at a number of institutions. Seven of these volumes,
each different from the others and none ever published, are known to exist. There are four
volumes at the Academy and one volume each at the University of North Carolina at
Chapel Hill, University of Michigan, and the Farlow Herbarium at Harvard University.
All told, in the seven volumes over 2,000 fungi are illustrated. In addition, there are 81
unbound plates of botanical illustrations in the Academy archives; 18 of these depict fungi
(Table 1) and the rest are of vascular plants.

Schweinitz also

——————
Manuscript submitted 6 November 2000, revised 30 March 2001. :
*Current address: Farlow Herbarium, Harvard University, Cambridge, MA 02138

49

50 BARTONIA

TABLE 1. Index to the Academy of Natural Sciences of Philadelphia’s unbound mycological
illustrations by Lewis David von Schweinitz. All are line drawings plus watercolor, except
for plate no. 48, which is just in pencil. Plates 18 through 47 are of vascular plants.

Plate Fungus Illustrated Plate Fungus Illustrated
1 Phallus sp. 13. Sphaeria Persicaria
2 Phallus coccineus. Salem Sphaeria capitata
3 Amanita caesarea Salem Sphaeria Geoglossum
Boletus calopus. Salem Sphaeria subterranea
Boletus floccosus Salem 14 § ia friabili
4 Agaricus Lact. Indigo. Salem Sphaeria Diaspyri [')
Agaricus cinabarinus [!] Sphaeria smilacicola
[unnamed (Agaricus?)] Sphaeria discincola
5 Agaricus Myc. nigripes S[r?] Salem Sphaeria Viburni
Arachnion candidum S[r?] Sal. Sphaeria olivaceo-nigro ad undulat
Agaricus P[?] lamellosus S[r?] Salem Sphaeria grisea
Agaricus Gym. Rhodoxanthus S[r?] Salem Sphaeria spissa
Merulius olivaceus S[r?] Salem 15 Sphaeria Sacculus
Boletus merulioides S[r?] Salem Sphaeria molliuscula
6 Agaricus niger S[r?] Salem Sphaeria enteroleuca
Leoti ida Salem Sphaeria annulata
{unnamed (Agaricus?)] Sphaeria (melograma) turbinulute [!]
{unnamed (Sphaeria?)] 16 Sphaeria protracta P.
Agaricus P[?] pectinatus. Salem Sphaeria Scirpi
7 Dacryomyces pellucidus. Salem Sphaeria Yuccae glorios.
Agaricus Gym. illudens Salem Sphaeria Aristidae
8 Amanita ampla Salem Sphaeria Sarraceniae
9 Agaricus. Micromph. venosus Sal Sphaeria Typhae.
Boletus Betula s{r?] Salem Sphaeria Tetragonathecae
Merisma fastidiosum 17 Sphaeria acervulata
Agaricus Myc. siccus s[r?] Salem. Sphaeria gregalis
10 [unnamed (Sphaeria?)] Sphaeria cohortina
11 Sphaeria nigripes Sphaeria hydnicola
Sphaeria teres Sphaeria caulina
Sphaeria capreolatae Sphaeria horrida
Sphaeria radicalis 48 Gilled mushroom
12 Sphaeria lenta
Sphaeria lobata
Sphaeria ochracea
Sphaeria poc
Sphaeria atropurpurea
Sp
Sphaeria ceratosperma

ea ee

The bound volumes at the Academy, titled the Fungorum Nieskiensium Icones, are volumes
1, 2, 3, and 5 of apparently a single series. They are designated, respectively, on their title
pages: “Pars I,” “Pars II,” “Pars III,” and “Pars V” and contain plates numbered 1-273, 350-
453 (excluding numbers 33, 40, 44, 45, 46, 47, 50, 51). The Chapel Hill cones is volume 4
of this series (Phillips and Phillips 1963). Eugene A. Rau, who deposited the bulk of the

LEWIS DAVID VON SCHWEINITZ’S MYCOLOGICAL ILLUSTRATIONS 51

Schweinitz materials in the Academy archives, said that he had seen six or seven volumes
of Icones (Krieger 1940; Phillips and Phillips 1963) and so it was believed for decades that
there were volumes 6 and 7 still unaccounted for.

RESULTS AND DISCUSSION

I recently came across evidence that refutes the existence of volumes 6 and 7 of the
Academy/Chapel Hill Jcones. In the manuscript collection of the American Philosophical
Society, there is microfilm of letters from Schweinitz to John E. LeConte. In a letter dated
25 June 1820, Schweinitz writes: “The following is the Contents of my V volumes of fungi
- Vol. I From Sphaeria to Phallus exclus. of the latter II Agarici only III Boleti to the
Byssocladium or Naematelia excl. vol IV Supplement to I, V Suppl to II and Il...”
Schweinitz is surely referring to the Academy/Chapel Hill Jcones: (1) The contents that he
lists correspond to those in the Academy Jcones. (2) If volume 5 is a supplement to volumes
2 and 3, it would be expected that there would be an overlap in genera treated in the
volumes. This is the case: 14 of the 16 genera treated in volume 3 are treated in volume 5;
3 of the 3 genera treated in volume 2 are treated in volume 5 (Table 2). (3) There is no five-
volume mycological work in the Schweinitz bibliography appended to Walter R. Johnson’s
1835 obituary of Schweinitz, nor have I seen any reference elsewhere for such a work; thus
there is no other known work that Schweinitz could have been referring to.

And so, there are most probably only five volumes of the Academy/Chapel Hill cones.
The possibility remains that Schweinitz painted additional supplements and they are the
volume(s) to which Rau referred. Could the Michigan or Farlow Icones be such volumes?

A 1938 letter from Dr. E. B. Mains of the University of Michigan, tucked into volume
1 of the Academy Icones, states that the volume at University of Michigan “is entitled Icones
Fungorum on the back . . . There is no title page.” This is unlike the Academy Icones, all
of which have title pages upon which are written Fungorum Nieskiensium Icones. “Clearly
not a part of the Fungorum Nieskensium Icones series of ANSP” is handwritten at the
bottom of Mains’ letter. The letter is addressed to Willman Spawn, but according to his
wife, Carol Spawn, former Academy archivist, the note is not in Willman Spawn’s hand-
writing (C. Spawn, pers. comm.). It is unknown who wrote this note, and its author did
not explain why the Michigan cones is “clearly not a part” of the Academy Icones. Krieger
(1940) describes the Michigan Icones as “about 8 cm thick, with boards measuring 20 by 30
cm... Stamped in gold on the back are the words ‘Icones Fungorum.’ ” The Academy
Icones are not as thick, their other dimensions are greater (23.5 cm x 37.5 cm (vols 1, 2, 3;
vol. 5 is 23.2 cm wide)), and they are not stamped in such a manner. Thus the Michigan
Icones is most probably not a part of the Academy/ Chapel Hill Icones.

A recent trip to the Farlow Library yielded the following information about the Farlow
Icones. The cover measures 18.5 cm x 25.5 cm, while the interior pages measure 18 cm x
25 cm. The interior pages were apparently cut to their present size after the illustrations
were done, as some of the illustrations and captions are cropped. The cover is red cloth
with Icones Fungorum Nieskiensium printed in gold leaf. The Farlow volume contains 50
plates, and according to a listing prepared by Dr. Donald Pfister, curator of the Farlow
Herbarium, it contains 248 illustrations. Mesenterica grisea, a species illustrated in the Farlow
Icones, is in Dana Lynch’s list of species ‘Illustrated in the Academy Icones that were also
newly described in Albertini and Schweinitz’s Conspectus (Lynch 1996). The known
Provenance of the Farlow Icones is incomplete. According to Dr. Pfister, this volume came

52 BARTONIA

TABLE 2. Index to the genera of fungi illustrated in Lewis David von Schweinitz’s Fungorum
Nieskiensium Icones at the Academy of Natural Sciences of Philadelphia.

PAGE(S) GENUS NAME PAGE(S) GENUS NAME
Pars I 231-232 Helvella
64-67 Aecidium 201-207 nu
57 Arcyria Leotia
42 Bovista Merisma
75 Conoplea 238-266 P
59 Cribraria 197-200 Sistotrema
76-77 "yathus 270-272 Stilbum
52 208-219 The a
41 Geastrum 233-237 Tremella
26-28 Hysterium
ae Pars V oa
49 L a
62-63 ft a, <oee
4 ‘ortinaria
54 preeie 359-369 Gymnopus
6 Pilabodes 352-355 Lepiota
72-73 inia i poche
8 Selevaderene 374-375 phalia
37-39 Sclerotium 376-378 Pleuropus
* 372 Pratella
58 Stemonitas 373 Russula
36 Sphaerobo 351 mani
1-25 Sphaeria 425 Ascobolus
55-56 Trichia 428 Ascophora
Trin 382-392 Boletus
35 Tubercularia sec Botrytis
61 Tubioling 429 Ceratium
43 Tulostoma 405-409 Clavaria
68-71 ale 381 Daedalea
30-32 Xyloma 438-442 Dematiu
Epichysium
Pars II 443-445 ineum
Agaricus 410 Geoglossum
127-131 Coprinus 419, 425, 109b Helotium
90-102 Cortinaria 414 Helvella
103-120, 161 mopus 449-450 Himantia
138-143 Lactifluus 394-399 Hydnum
83-89 Lepiota 431-433 Isaria
121-126 Mycena 411 Leotia
152-161 Omphalia 379-380 Merulius
162-168 Pleuropus 404 Merisma
132-137 Pratella 452-453 Mesenterica
144151 Russula 435-437 Monilia
78-82 Amanita 1 Morchella
169-174 Merulius 430 Periconia
416-424 eziza
ee , 350 Phallus
B 446-448 Racodium
267 Ascobolus 451 Rhizomorpha
poh oe 393 Sistotrema
175 412 Spathularia
298 Geisleiieds 426 : Stilbum ¢

268-269 Helotium 415 Tremella

LEWIS DAVID VON SCHWEINITZ’S MYCOLOGICAL ILLUSTRATIONS 53

to the Farlow via Maselio Schaechter, a member of the Boston Mycological Club, who
bought it at a bookseller’s in Bungay, England, and then donated it to the Farlow. Maselio
Schaechter’s story of this volume can be found on pages 102 and 103 of his book, Jn the
Company of Mushrooms, Harvard University Press, 1997 (Pfister, pers. comm.). On the
inside front cover of the Farlow Icones, there is a tag in the upper left corner on which is
printed: “W. Webster/ Bookseller & Stationer/ Late G. Fell/ 60 Piccadilly”. Also on the
inside front cover is a bookplate on which is printed “ORA E SEMPRE/ Sir Francis Denys
Bart.” Written in ink, on an interior page, is “Catherine Eliza Perceval/ Philadelphia/ March
8th 1826.”

The Icones at the Farlow is not part of the series at the Academy. The Farlow volume
differs from the series at the Academy as follows: (1) it is much smaller in format than the
volumes at the Academy (18.5 cm x 25.5 cm vs. 23.5 cm x 37.5 cm (Academy vols 1, 2, 3;
vol. 5 is 23.2 cm wide)); (2) it does not have a title page; (3) its plates are numbered 1-50,
these plate numbers are duplicated in volume 1 of the Icones at the Academy, and the
“missing” volumes (6 or 7) would contain plates with higher numbers; (4) many of its plates,
in addition to painted illustrations with names and numbers written in ink, also have
numbers and names and sketches which are written and drawn in pencil. The Academy
Icones contain nearly exclusively painted illustrations with names and numbers written in
ink.

CONCLUSION

The volumes of Schweinitz’s paintings at the Academy and Chapel Hill form a single set,
and the volumes at Michigan and the Farlow are not part of that set. The Michigan and
Farlow Icones are different in format from each other and from the Academy/Chapel Hill
Icones and the letter from Schweinitz to LeConte strongly suggests that the Fungorum
Nieskiensium Icones was only intended to be a five-volume set.

ACKNOWLEDGMENTS

This paper is dedicated to Donald P. Rogers (1908-2002), biographer of Lewis David von Schweinitz.
Dr. Rogers reviewed an earlier manuscript of this paper, and I am grateful for his many helpful
comments. ;

I thank Dr. A. E. Schuyler of the Academy of Natural Sciences of Philadelphia and Dr. Donald
Pfister of the Farlow Herbarium for their generous contributions of information, advice, and
encouragement. I thank the staff of the Archives of the Ewell Sale Stewart Library for allowing me
to examine the Academy’s set of Schweinitz’s [cones (Collection 437). I thank the staff of the Farlow
Reference Library of Cryptogamic Botany fo
Schweinitz’s Icones.

LITERATURE CITED

BENEDICT, D. M. 1935. Lewis David von Schweinitz, the mycologist. Bartonia 32: 12-14 eee
JOHNSON, W.R. 1835. A memoir of the late Lewis David von Schweinitz, P. D. with a sketch of his scientific labors.
Read before the Academy of Natural Sciences of Philadelphia, 12 May 1835. William P. Gibbons, Philadelphia.
KRIEGER, L. C. C. 1940. Unpublished color plates of fungi by Lewis David von Schweinitz. Bartonia 20: 28-29.
LYNCH, D.M. 1996. Paintings of fungi by Lewis David von Schweinitz in the archives of the Academy of Natural
Sciences of Philadelphia. Bartonia 59: 125-128.
PENNELL, F. W. 1935. The botanist Schweinitz and his herbarium. Bartonia 16:
PHILLIPS, V. T. AND M. E. PHILLIPS. 1963. Guide to the manuscript collections in
of Philadelphia. Special Publ. No. 5, Academy of Natural Sciences of Philadelphia. im
ROGERS, D. P. 1977. L. D. de Schweinitz and early American mycology. Mycologia 69: 223-245.

1-8
the Academy of Natural Sciences

Original figure of ANSP 3854 taken from Lesquereux (1879:

pl. 55, fig. 3). (Note that Lesquereux obliterated Mansfield’s

catalogue number and replaced it with his own.)

= ;
(i

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GN

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Gaytw.

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*~, ..

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Lesquereux, Lectotype,

ANSP 3854 (Lesquereux #90, PAGS #11163).

Sphenopteris goniopteroides

Bartonia No. 61: 54-80, 2002

A Catalogue of the Type Specimens of the Taxa Erected by
Leo Lesquereux in the Coal Flora (1879, 1880, 1884)

JAMES LENDEMER
Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway,
Philadelphia PA 19103-1195

ABSTRACT. Leo Lesquereux’s Coal Flora has long been considered the basis for American
Carboniferous paleobotany and is a monument to his idominable will to succeed under trying
conditions. Throughout the four volumes that make up the Coal Flora, 247 new species are introduced.
The type specimens upon which these species are based are known to be scattered throughout the
collections of numerous institutions. A catalogue of these specimens has never been compiled although
one was begun by Watt in 1974. This paper provides, for the first time, a catalogue of the type
specimens of all of the Coal Flora taxa.

INTRODUCTION

Darrah (1969: 1) noted of Lesquereux’s Coal Flora “It is a paradox that Leo Lesquereux,
who performed prodigious work under trying conditions, on one hand provided a
magnificent panorama of the ‘Coal Flora’ and on the other, left a trail of unsettled details
that remain to this day extremely difficult to interpret.” Lesquereux’s Coal Flora (1879, 1880,
1884) can be considered the “foundation of American Carboniferous paleobotany” (Darrah
1969: 2). Many of the newly erected species throughout its pages were based on specimens
from the collections of R.D. Lacoe of Pittston, Pennsylvania and LF. Mansfield of
Cannelton, Pennsylvania. However, the collections of J.H. Britts, Professor Eugene A.
Smith, S.S Strong, William Lorenz, and many others, also were utilized by Lesquereux. The
importance of the Coal Flora lies partially in its broad coverage. The taxa described within
its pages were not limited to Pennsylvania but also included those from as far west as
Arkansas, Missouri, Iowa and Illinois. This broad coverage also allowed Lesquereux to draw
valuable conclusions on the relationships of various Carboniferous floras of the United
States, as well as relationships between the coal floras of America and those of Europe.

since the transfer of the collection to the Pennsylvania Geological Survey in 1889.

All of the specimens originally held for the “Geological Museum” were kept in dead
storage at the Academy of Natural Sciences in Philadelphia (= ANSP). The collection was
later transferred to the Pennsylvania State Museum (W illiam Penn Memorial Museum) in
Harrisburg. However, a small portion of Lesquereux’s collection was apparently retained at

Manuscript submitted 13 November 2000, revised 2 April 2001.

fs

56 BARTONIA

ANSP. These specimens were rediscovered by Darrah who simply identified them as
“Lesquereux’s Coal Flora types.” The collection as a whole has been overlooked by
apparently every author with the exception of Darrah who identified one holotype (=
lectotype) from the material. The small portion of the collection held at ANSP has recently
been extensively studied and the types reported by Spamer and Lendemer (2000; a revision
of Spamer, 1988).

REPOSITORIES OF LESQUEREUX’S COAL FLORA TYPES

Lesquereux’s Coal Flora types are deposited mainly at two institutions, Harvard and the
United States National Museum. This is because many of Lesquereux’s species were based
either upon specimens from the collection of R.D. Lacoe or were later bought by him. The
majority of Lesquereux’s early specimens from Lacoe’s collection were given to Harvard.
However, by far the majority of Lesquereux’s Coal Flora types were still in Lacoe’s
possession when his collection was donated to the National Museum.

Smaller collections identified and described by Lesquereux in the Coal Flora can be found
in a number of other museums. These are listed below with the institutional acronym that
is used here: ANSP — Academy of Natural Sciences of Philadelphia; AGS SM — Alabama
Museum of Natural History (Tuscaloosa); CP — Pratt Museum of Natural History (Amherst,
Massachusetts); HU — The Botanical Museum, Harvard University (Cambridge, Massachu-
setts); ISM — Illinois State Museum (Springfield); LC- Lafayette College (Easton, Pennsylva-
nia); USNM — National Museum of Natural History (Washington, D.C.); SMP —
Pennsylvania State Museum (William Penn Memorial Museum; Harrisburg); UP — Chicago
Field Museum.

Of the 247 taxa erected in the Coal Flora, 37 currently are not represented by any known
type specimens. Some of these may be at Harvard. However, the author has been unable to
examine the collections there. A list of these taxa is provided below (Appendix A) to
facilitate later workers in locating or verifying specimens which may be suspected as type,
or for designating neotypes. However, this is not intended as the final word on the matter
as some or all of the types may be found at a later time.

ARRANGEMENT OF THE CATALOGUE

The species in this catalogue are listed alphabetically by their specific epithet (and indexed
by genus in Appendix B). Each of the binomials are the original nomenclatural combinations
given by Lesquereux and only primary types (holotypes, isotypes, lectotypes, and syntypes)
are listed here. In the case of some species, it could not be determined whether Lesquereux
had only a single specimen (a holotype) or a suite of specimens (syntypes) in his possession
when the species was described. Thus, a single specimen is treated as a holotype only when
it is known to have been the only specimen upon which the species was described.
Otherwise, a single specimen is treated as a syntype. Catalogue numbers that are linked by
an addition sign (+) are either contiguous pieces of one specimen or part and counterpart
(in cases of the latter it is indicated which is the part and which is the counterpart). Finally,
no systematic changes or revisions are made here. The names used are in their original state
as given by the author at the time of description (this includes the original orthography):
This is meant to place them in their proper historical context and avoid the ambiguity that

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX ay

often arises from the changes made by later authors. For the majority of the species listed
here notes on synonomy and current usage are not included.

DESIGNATION OF LECTOTYPES

Relatively few lectotypes have been designated for Lesquereux’s Coal Flora taxa. Merrill
(1907), Carluccio et al. (1966), Darrah (1969), and Goubet et al. (2000) are, in fact the only
previous authors to have done such, and Merrill (1907) did so inadvertently. Under the
advice of H.W. Pfefferkorn and E.E. Spamer (pers. comm.), the author has kept with this
trend. Only one lectotype is designated in this paper, for the taxon Sphenopteris goniopter-
vides Lesquereux. S. goniopteroides is not well understood and the only known type
specimens were recovered by the author during the preparation of the catalogue of type
specimens at ANSP. This paper does, however provide a review of the few lectotypes that
have been designated previously.

CITATIONS AND PUBLICATION DATES OF THE COAL FLORA

The manner in which the descriptions of the taxa erected in the Coal Flora are cited has
changed almost as often as the names themselves. Darrah (1969) treated all of the taxa
described in the Text volumes (1880) with indication to the Atlas (1879) as having been
erected by name and figure(s) in the Atlas. More recently however, these same taxa are
considered to have been erected in the Text with indication to the figures in the Atlas, this
is how these taxa are cited in this paper. Volume three of the Coal Flora (1884) is considered
separately from the first two volumes.

CATALOGUE

abbreviata, Odontopteris Lesq. ' ca
Lesquereux (1879: pl. 21, fig. 7; 1880: 138-139). Holotype, USNM 14827. Carbon Hill shaft, “C. vein,
ittston, Pennsylvania.

abnormalis, Rhabdocarpus Lesq.
Lesquereux (1884: 818-819, pl. 110, figs. 48, 49). Syntypes, USNM 26370. Cannelton, Beaver County,
Pennsylvania; collected by I.F. Mansfield.

Both of the illustrated specimens are preserved on the same rock.

adamsii, Trigonocarpus Lesq. .
Lesquereux (1884: 820, pl. 110, figs. 51-57). Syntypes, USNM 26727-26733 (figs. 51-57 respectively; all
of the specimens carry Lacoe #957). Stark County, Illinois.

agassizi, Neuropteris Lesq.

Lesquereux (1879. pl. 17, Tes. 1-4; 1880: 117-118). Syntypes, HU 5822+5824 +5826 +5827 +USNM

10050 (fig. 1), HU 5833 (fig. 2), HU 5843 (fig. 3), HU 5853 (fig. 4). Mount Hope coal mines, Portsmith,

Rhode Island. |
The specimens in figs. 1-3 were “communicated” to Lesquereux by Mr. J.H. Clark of Newport. Fig.

4 was copied from Teschmacher (Boston Journal of Natural History 5: 34).

58 BARTONIA

aldrichi, Callipteridium Les
Lesquereux (1879: pl. 39, oy + 1880: 171-172). Syntypes, USNM 11238 (fig. 1), USNM 12087 (fig.
2), USNM 11242 (fig. 3). Black Creek Seam, Jefferson mines, Alabama; collected by T.H. Aldrich.

ambigua, Alethopteris Lesq.
Lesquereux (1879: pl. 31, figs. 1-4; 1880: 182-183). Syntypes, USNM 13066 (fig. 1), USNM 41056 (fig.
2), USNM 41057 (fig. 3), HU 7465 (fig. 4). Henry County, Missouri.

ampullaeformis, Trigonocarpus Lesq.
Lesquereux (1884: 823, pl. 109, figs. 18-20, 212). Syntypes, USNM 26536, inert County,
Arkansas; USNM 26538, USNM 26537, Tracy City, Tennessee; communicated by F.L. Harvey;
donated by R.D. Lacoe.

All of the specimens carry Lacoe #975 The specimens are figured as follows: fig. 18 is USNM 26538,
fig. 19 is USNM 26536, and fig. 21 is USNM 26537. The specimen in fig. 20 is currently unlocated.

anceps, Stemmatopter
Lesquereux (1884: 838) cana USNM 17139 ee 4 + 17140 (counterpart; Lacoe #611). Cannelton,
Beaver County, Pennsylvania; donated by R.D. Lac

Lesquereux erected this taxon as a questionable new james

andrewsi, Lepidodendron Lesq.
Lesquereux (1879: pl. 64, fig. 6; 1880: 389-390). Holotype, USNM 15430+ 15431. Mazon Creek, Grundy
County, Illinois.

gularis, Lepidocystis Le
pares (1879: pl. 69, fies 16, 17; 1880: 456). Syntype, ees 25255425256 (fig. 17). Campbells
Ledge, Port Griffith?, Pennsylvania; donated by R.D. Lac
The specimen in fig. 16 is currently unlocated.

stata, Stemmatopteris Lesq.
Lesquereux (1879: pl. 59, fig. 5; 1880: 339). Holotype, USNM 17078. Cannelton, Beaver County,
Pennsylvania; collected by LF. Mansfield.

anomala, Neuropteris Lesq

Lesquereux (1879: pl. 7, fig, 1; 1880: ie slg Syntype, HU 5860 (MCZC 430). Gray roof shale of
Morris, Illinois; collected by Joseph E

apiculatus, Rhabdocarpus

Lesquereux (1884: 819, pl. 110, fig. 50). Holonri USNM 26756 (Lacoe #979). Stanton Mine, Wilkes:
Barre, Pennsylvania; donated by R.D. Lacoe

apiculatus, Cordaicarpus Les sq.
Lesquereux (1880: 551). Syntype, USNM 25322. Cannelton, Beaver County, Pennsylvania; collected
by LF. Mansfield.

This taxon is not erected in the Atlas as Lesquereux noted in the errata of volumes one and two.
Instead the taxon is erected in volume two with indication to the figures in the Atlas esque 197?
pl. 83, figs. 6, 6a). The explanation of the plates in the Atlas identifies the figures as “ Co
gutbieri.” —— later also emended the reidentification of the figures as “Cordaicarpus ap icalave
sp. nov.” to “ C. ovalis sp. nov.” (Lesquereux 1880: errata).

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 59

arborescens, Lycopodites Lesq.
Lesquereux (1884: 778-779, pl. 106, fig. 1). Syntype (Holotype?), USNM 15559 (Lacoe #257). veo s
coal bank, Washington County, Arkansas; communicat ted by F.L. Harvey; donated by R.D. L

aspera, Neuropteris Les
Lesquereux (1879: pl. 13, fies: 10-12; 1880: 121), Syntype, HU 5862. Cannelton, Beaver County,
Pennsylvania; collected by LF. Mansfield.

One of the two missing syntypes may be SMP 11101 (Lesquereux #28).

biformis, Neuropteris Les
Lesquereux (1879: pl. 13, fig 7; 1880: 121). eve i AGS-SM 34. Coal Measures, Tuscaloosa,
Alabama; collected by Professor Eugene A. Sm

Goubet et al. (2000) moved this taxon into = genus Neuralethopteris Cremer ex Laveine.

brevifolia, Calamostachys
Lesquereux (1884: 718, pl. ps fs 5, 5a). Holotype, USNM 18358. Cannelton, Beaver County,
Pennsylvania; collected by I.F. Mansfield.

brevifolium, Taeniophyllum Lesq
Lesquereux (1884: 788-789, pl. 108, fs 3, 3a). Syntype, USNM 19276 + 19277 (Lacoe #774). Cannelton,
Beaver County, Pennsylvania; donated by R.D. Lacoe.

brittsii, Lepidodendron Lesq. '
a ape (1879: pl. 63, fig. 1, 2). Syntype, USNM 15482 (fig. 1). Clinton Coal, Missouri; collected
Dr. J.H. Britts.
” The specimen in fig. 2 is currently unlocated.

brittsii, Sphenopteris Lesq ; : : .
Lesquereux (1879: pl. 55, C 2; 1880: 277-278). Syntype, USNM 14953 (fig. 2). Clinton, Missouri;
collected by Dr. J.H. Britts

broadheadi, Conostrichus Lesq. . :
Lesquereux (1879: pl. B, figs. 1, 25 1880: 15-16). Holotype, USNM 10250. Vernon County, Missouri
Pottsville Group, Pennsylvanian.

butleri, Lepidostrobus Lesq ; ,
Lesquereux (1884: 240-841). Syntype, USNM 16175 (Lacoe #681). Butler Mine, near Pittston,
Pennsylvania; donated by R.D. Lacoe.

butlerianus, Carpolithes
Lesquereux (1884: 824-825, wi 111, fig. 19). Holotype, USNM 26355 (Lacoe #990). Butler Mine,
Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe

Lesquereux (1879: pl. ny fies. 1-8; 1880: 115). Syntypes; HU 5796 (fig. 1), New Philadelphia,
canine HU a (fig. , HU 5800-5802 (figs. 3-5). Pomeroy, Ohio; HU 5797 (fig. 6), HU 5799
(fig. 7), Wilkes-Barre, Pennsylvania; HU 5803 +5804 (fig. 8), Pomeroy, Obio. d ules, figs. 6 and
Darrah (1969: 97) noted, “I would exclude from this = the two cyclopteri eA a i? “ai
7, and include questionably the cyclopterid pinnule, fig. 8.” Stockmans poi P Igs. hme
refigured the specimens in Lesquereux’s figs. 2, 3, 4, tie 8 Fuld e specimen in fig.

refigured by Darrah (1969: pl. 72, fig. 1)-

60 BARTONIA

campbellianum, Lepidophyllum Lesq.
Lesquereux (1884: 786, pl. 107, figs. 6, 7). Syntypes, USNM 16377 (fig. 7), USNM 16376 (fig. 6), USNM
16378-16380. Campbells Ledge, Pittston, Luzerne County, Pennsylvania.

cheathami, Eremopteris Lesq.

Lesquereux (1884: 770-771, pl. 104, figs. 2-4). Syntypes, USNM 15009 (fig. 2), Washington County,

Arkansas; USNM 15025 (fig. 3), USNM 15026 (fig. 4), Tracy, Tennessee; donated by R.D. Lacoe.
All of the specimens carry Lacoe #1038.

circularis, Cardiocarpus Lesq.
Lesquereux (1884: 812, pl. 110, fig. 10). Holotype, USNM 25592 (Lacoe #963). Vermilion County,
Indiana.

clarkii, Pecopteris Lesq.
Lesquereux (1879: pl. 41, fig. 10; 1880: 261-262). Holotype, HU 7390. Mount Hope Coal mines, Rhode
Island; collected by Mr. J.H. Clark

clavatum, Trochophyllum Lesq.

Lesquereux (1879: pl. 3, figs. 21-23; 1880: 65). Holotype, USNM 16643. In a bed of friable black shale
in the barren measures, about one hundred feet lower than the Pittsburgh Coal, between Irwin Station
and Pittsburgh, Pennsylvania; collected by W.D. Moore.

clintoni, Pecopteris Lesq.
Lesquereux (1879: pl. 42, figs. 1-5; 1880: 251-252). Syntypes, USNM 13172 (fig. 3), USNM 30857 (fig.
1), USNM 13173 (fig. 5).

The specimens in figs. 2 and 4 have not been located.

communis, Macrostachya Lesq. ‘
Lesquereux (1879: pl. 3, figs. 17, 18; 1884: 828-829). Syntypes, USNM 17740 (fig. 17), USNM 17751 (fig.
18), USNM 10974, Cannelton, Beaver County, Pennsylvania; 17742, Olyphant, Lackawanna County,
Pennsylvania.

communis, Sphenopteris Lesq.

Lesquereux (1884: 762-763, pl. 104, figs. 1, 1a). Syntypes, USNM 14340 (fig. 1), USNM 14345, USNM
14346 (all of the specimens carry Lacoe #442). Arkansas; Dade, Georgia; Tracy, Tennessee; donated by
R.D. Lacoe.

compacta, Knorria j
Lesquereux (1884: 839-840). Syntypes, USNM 15800-15802 (all of the specimens carry Lacoe #816).
Thayer, Kansas; donated by R.D. Lacoe.

conglobatus, Cardiocarpus Lesq.
Lesquereux (1884: 810-811, pl. 109, figs. 10, 11). Syntype, USNM 25565 (fig. 11). Washington County,

Arkansas.

conicus, Carpolithes Lesq

Lesquereux (1884: 824, pl. 109, fig. 17). Holotype, USNM 26483 (Lacoe #959). Shamokin, Pennsylvania
donated by R.D. Lacoe.

corrallum, Rhacophyllum Lesq.
Lesquereux (1879: pl. 57, fig. 4, 4a; 1880: 317). Syntype, USNM 19469 (fig. 4). Mazon Creek, Grun#Y
Co inois.

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 61

riaceum, Lepidophyllum Lesq.
ens (1884: 787, pl. 107, fig. 10). Lectotype, USNM 15908 (fig. 10; Lacoe #672). Cannelton,
Beaver County, Pennsylvania; donated by R.D. Lacoe.
USNM 15908 was designated as the lectotype of Lepidophyllum coriaceum Lesquereux, by Merrill
(1907: 210).

cornuta, Odontopteris Lesq.
Lesquereux (1879: pl. 22, figs. 7-9; 1880: 128-129). Syntypes, USNM 10348 (fig. 9), USNM 10345 (fig.
7), USNM 10346 (fig. 8). Cannelton, Beaver County, Pennsylvania; donated by IF. Mansfield.

cornutum, Rhacophyllum Le
Lesquereux (1879: pl. 57, fig. 1880: 317-318). Holotype, HU 7708. Cannelton, Beaver County,
Pennsylvania.

cornutus, Rhabdocarpus
Lesquereux (1879: pl. 85, = 5 1880: 583). Syntype, HU 8019 (fig. 15; Lesquereux #T. 29). Mazon
Creek, Grundy County, Illinoi

crassa, Volkmannia
Lesquereux re a pl. 90, fig. 1). Syntypes, USNM 18091 (Lacoe #851), USNM 18091. Dade
County, Geo

The Gipueedh nan is USNM 18091. The second syntype is currently unfigured.

crassus, Cardiocarpus Les
Lesquereux (1884: 812, pl. “09, fig. 12; pl. 110, figs. 6-9). Syntypes, USNM 25598-25602 (all of re
specimens carry Lacoe #963). Campbells Ledge, Pittston, Luzerne County, Pennsylvania; donated b
R.D. Lacoe

The e specimens are "pene as follows: fig. 6 is USNM 25599, fig. 7 is USNM 25600, fig. 8 is USNM
25601, fig. 9 is USNM 2560

cuspidata, Annularia L
Lesquereux (1884: 725, 92, figs. 7, 7a). Holotype, USNM 17851 (Lacoe #788). Rushville, Ohio;
Allegheny Group, Pennsylvanian; donated by R.D. Lacoe.

cuspidatum, Lepidodendron Lesq. nee
Lesquereux (1879: pl. 64, fig. 7). “Hole, { USNM 15437. “Plymouth E vein, Pittston, Luzerne
County, Pennsylvania; donated by R.D. Lac

cyclostigma, Lepidodendron Lesq. : ‘ : .
Lesquereux (1879: pl. 62, fig. 5; 1880: 394-395). Syntype, USNM 15501 a
donated by Dr. J.H. Britts.

cyclostigma, Stemmatopteris Les hant, P l
Lesquereux (1880: 341). Syntypes, USNM 11050, USNM 17188, USNM 17089. Olyphant, Pennsylvania.

decipiens, Neuropteris Lesq. ne
Lesquereux (1880: 93). Syatypes, U USNM 10621, USNM 10623. Mazon Creek, — Illinois.

Both specimens consist of a part and a counterpart. Lesquereux noted that he ad specimens
from the Centralia coal shaft in Illinois. These specimens have yet to be loca

62 BARTONIA

dentata, Megalopteris Lesq
Lesquereux (1884: 833-834), — hail 11448-11450 (all of the specimens carry Lacoe #783).
Rushville, Ohio; donated by R.D. Lac

denticulata, Archaeopteris Lesq.
Lesquereux (1884: 774). Syntype, USNM 17008. Rushville, Ohio.

dichotomum, Dicranophyllum Le
Lesquereux (1880: 553-554, pl. 87, fig: 9). Syntype, USNM 25108 (fig. 9). Cannelton, Beaver County,
Pennsylvania; collected by IF. Mansfield.

dichotomus, Cordaianthus Lesq.
Lesquereux (1879: pl. 76, fig. 7 1880: 546-547). Holotype, USNM 19212. Henry County, Missouri.

dilatatus, Cardiocarpus Le
Lesquereux (1884: 806-807, ok 110, fig. 2). Syntype, USNM 25384 (fig. 2; Lacoe #961). Washington
County, Arkansas; donated by R.D. Lacoe.

dilatatus, Lepidophloios Lesq.
Lesquereux (1879: pl. 68, figs. 6, 7; 1884: 781-782, pl. 105, figs. 1-4). Syntypes, USNM 15947 (fig. 6),
Clinton, Henry County, Missouri; USNM 15941+15942 (fig. 7), Cannelton, Beaver County,
Pennsylvania; USNM 15944 (fig. 1), Clinton, Henry County, Missouri; USNM 15945 (fig. 2), USNM
15956 (fig. 3), USNM 15943 (fig. 4); spiced ANSP 3721 (Lesquereux 3 [11076] (Lesquereux, 1887:
99)]), SMP 11107 (Lesquereux 34 [Lesquereux 1889: 101]), Cannelton, Beaver County, Pennsylvania.
The specimen in fig. 3 is currently unlocated. USNM 15941 and 15942 are part and counterpart.
ANSP 3721 and USNM 15943 are currently unfigured. L. dilatatus was erected in the third volume of
the Coal Flora (1884: 781-783). In the description, Lesquereux placed the two figures from the Atlas
(1879: pl. 86, figs. 6, 7) under this name. The specimens from Cannelton were sent to Lesquereux by
LF. Mansfield. Spamer and Lendemer (2000) reported that ANSP 3721 was the counterpart of the
specimen in fig. 2. This however, is in error. The specimen figured was from Missouri while the ANSP
specimen was from Cannelton, Pennsylvania. It is, nonetheless an unfigured, questionable syntype.

diminutivus, Cardiocarpus
Lesquereux (1879: pl. 85, fig. 48; 1880: 570). cena USNM 26418. Sub-conglomerate of Pittston,
Luzerne County, Pennsylvania; donated by R.D. Lac

dimorpha, Pseudopecopteris Lesq.
Lesquereux (1879: pl. 35, figs. 1-6; 1880: ae rich CP 481 (fig. 6). Coal of Mount Hope, ea
Newport Rhode Island; donated by J.H. C

diplotesta, Cardiocarpus Le

Lesquereux (1884: 812-813, pl. 110, fig. 13). Holotype, USNM 25672 (Lacoe #272). Campbells Ledge
Pittston, Luzerne County, Pennsylvania.

divergens, Cardiocarpus

Lesquereux (1884: 811, pl. re S 4). Syntype, USNM 2551 (fig. 4; Lacoe #964). “E vein,” Pittsto®
Luzerne County, Pennsylvania.

ebracteatus, Cordaianth

Lesquereux (1884: 844). =a USNM 19216. “Coal A,” Boston Mine, Pittston, Luzerne County:
Pennsylvania.

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 63

elegans, Lepidophyllum Lesq.
Lesquereux (1884: 787-788, pl. 107, fig. 11). Holotype, USNM 16230+16231 (Lacoe #737). Mazon
Creek, Grundy County, Illinois; donated by R.D. Lacoe

USNM 16230+ 16231 are part and counterpart.

elrodi, Neuropteris Lesq.

Lesquereux (1879: pl. 13, fig. 4; 1880: 107-108). Syntype, USNM 11839 (fig. 4), Black Warrior Basin,
Alabama; syntype(?), SMP 11587 (Lesquereux #514 [11587] (Lesquereux 1889: 121)]), Whetstone
Quarries, Indiana.

Lesquereux found USNM 11839 in the collection of the Alabama State Geological Survey without
labels. Gastaldo (pers. comm.) has observed the specimen and noted that the lithology and preservation
corresponds to that of the Black Warrior Basin of Alabama. Lesquereux noted that he also had
specimens from the Montevallo Seam, Alabama (via T.H. Aldrich) and the Whetstone Grit of Orange
County, Indiana (via E.T. Cox). Goubet et al. (2000: 15) placed USNM 11839 in synonomy with
Nestidethontinis schlehanii (Stur). Goubet et al. (2000: 27) also placed all of the specimens later figured
by Lesquereux as Neuropteris elrodi into synonomy with Neuralethopteris biformis (Lesquereux).

emarginata, Stemmatopteris Lesq.
Lesquereux (1880: 337-338). caeaies USNM 17142. Cannelton, Beaver County, Pennsylvania;
donated by IF. Mansfield.

USNM 17142 was designated as the lectotype by Merrill (1907: 324). Lesquereux did not give an
enumeration of the specimens upon which he based this species. The description is not clear as to
whether one specimen was used or a suite of specimens was used. Thus, we treat the specimen as a
syntype that was designated as the lectotype by Merrill.

emersoni, Annularia
Lesquereux (1880: nee Spabgie USNM 17841. St. Clairsville, Ohio; Monongahela Formation,

ennsylvanian.

emersoni, Danaeites Lesq.
Lesquereux (1879: pl. 28, figs. 1-3; 1880: 157-158). Syntypes, USNM 12711 (fig. 1), USNM 12710 (fig.
2). Shale above the Coal, St. Clairsville, Ohio.

The specimens were sent to Lesquereux by Mr. P.W. Emerson of St. Clairsville. The specimen in
fig. 3 is currently unlocated.

evansii, Alethopteris Lesq.
Lesquereux (1884: 834). Syntypes, USNM 11886-11890 (all of the specimens carry Lacoe #1013). Tracy
City and Rockwood, Tennessee; donated by R.D. Lacoe.

€xpansum, Rhacophyllum Le:
Lesquereux (1879: pe 57, 7, fig Pi "4880: 323-324). Holotype, USNM 19435. Olyphant, Pennsylvania;
donated by R.D. Lac

fallax, Lepidophyllum Le

Lesquereux (1884. 786, pl 1 107, figs. 4, 5). Syntypes, USNM 15897 (fig. 5; Lacoe #892), Cannelton,

Beaver County, Pennsylvania; 158% (fig. 4; Lacoe #892a), Rhode Island; donated by R.D. Lacoe.
— (1907: 210) included USNM 15898 as one of the type specimens. However, the description

ssiacant = upon only two specimens, those that were figured. Thus, USNM 15898 is not part of the

64 BARTONIA

fasciculata, Megalopteris Lesq.
Lesquereux (1879: pl. 24, fig. 2; 1880: 150-151). Holotype, USNM 11704. Lower beds of the coal
measures of Illinois; collected by Mr. J.H. Southwell.

fasciculatus, Asterophyllites Lesq.
Lesquereux (1879: pl. 3, figs. 1-4; 1880: 41-42). Syntypes, USNM 18292+HU 8075 (fig. 2), USNM 18291
(fig. 1), USNM 18297 (fig. 3), Clinton, Henry County, Missouri.

The specimen in fig. 4 is currently unlocated.

fasciculatus, Cardiocarpus Lesq.
Lesquereux (1879: pl. 85, figs. 30, 3a; 1880: 570). Syntype, USNM 25775. Campbells Ledge, Pittston,
Luzerne County, Pennsylvania.

fertilis, Volkmannia Lesq.
Lesquereux (1884: 720-721, pl. 90, fig. 4). Syntypes, USNM 17772 (fig. 4), Roof of “coal B,” Archbald,
Pennsylvania; USNM 17773-17775, Stark County, Illinois.

flexuosus, Cordaianthus Lesq.
Lesquereux (1884: 802, pl. 109, fig. 2). Syntypes, USNM 19217 (fig. 2), USNM 19218 (both specimens
carry Lacoe #846). Campbells Ledge, Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

fucoideum, Rhacophyllum Lesq.
Lesquereux (1879: pl. 58, fig. 6, 7; 1880: 325). Syntypes, USNM 19443 (fig. 7), USNM 19442 (fig. 6).
Mazon Creek, Grundy County, Illinois; collected by S.S. Strong.

georgiana, Pecopteris Lesq.
Lesquereux (1884: 759-760, pl. 98, figs. 6, 6a). Syntypes, USNM 12420+12421 (fig. 6; Lacoe #855),
USNM 12422 (Lacoe #855a). Dade County, Georgia; donated by R.D. Lacoe.

gibsoni, Alethopteris Lesq. ; age?
Lesquereux (1879: pl. 28, figs. 4-6; 1880: 183-184). Syntype, USNM 11964 (fig. 4). Danville, Illinois.
The specimens in figs. 5 and 6 are currently unlocated.

giffordi, Caulopteris Lesq. .
Lesquereux (1879: pl. 60, figs. 1, 2; 1880: 343-344). Holotype, UP 422. Coal measures near Alta, Peoria
County, Illinois; donated by William Gifford.

giffordi, Trigonocarpus Lesq.
Lesquereux (1879: pl. 85, figs. 5, 6; 1880: 592-593). Holotype, USNM 26755. Lower coal measures, neat
Alta, Peoria county, Illinois; donated by William Gifford.

goniopteroides, Sphenopteris Lesq.

Lesquereux (1879: pl. 55, figs. 3, 4). Lectotype [designated here!], ANSP 3854 (Lesquereux #90 [11169

(Lesquereux 1889: 104)]). Cannelton, Beaver County, Pennsylvania; collected by LF. Mansfield.
The specimen in fig. 3 (ANSP 3854) is now partially broken, with a small triangular fragment _

missing. Also, at one time in the past, the specimen has been broken in two and glued back together

The identification which Lesquereux (1889: 104) gave for the specimen was Pseudopecopteris new

Lesquereux. This is in error. Both Lesquereux’s and Lesley’s catalogue numbers match. Furthermor®

a comparison of the specimen with the protologue has found that the specimen matches st 2 *

respect. This proves that either Lesquereux was in error or that he reidentifed the specimen before

it was sent to the Pennsylvania Geological Survey.

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 65

gracile, Lepidophyllum Les
Lesquereux (1884: 786-787, at 107, fig. 8). Holotype, USNM 16245 (Lacoe #894). Campbells Ledge,
Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

gracilis, Calamites Lesq
Lesquereux (1879: pl. 5, fig. 17; 1880: 29-30). Holotype, HU 4479. Upper coal strata, Western
Kentucky.

gracilis, Equisetites Lesq.
Lesquereux (1884: 729-730, pl. 90, fig. 5). Holotype, USNM 18236 (Lacoe #938). Dade County,
Georgia; donated by R.D. Lacoe.

grand’euryi, Sigillaria Lesq.
Lesquereux (1884: 795-797). pd USNM 16399, USNM 16400. Cannelton, Beaver County,
Pennsylvania; donated by R.D

Both specimens carry Lsce: ey

grandis, Lesleya Les
Lesquereux (1879: pl. os, figs. 1-3; 1880: 143). Syntype, ISM 8654 (fig. 2). Soft shaly sandstone, base of
the Chester Limestone.

The two remaining specimens were cited by Watt (1974) as being at Harvard. However, no
catalogue numbers were given. All of the specimens were communicated to Lesquereux by Prof. A.H.
Worthen.

grandis, Trigonocarpus Le
Lesquereux (1884: 821, pl. 111 cx 1-3). Syntypes, USNM 26020 (fig. 3), USNM 26018 (fig. 1), USNM
26019 (fig. 2; all of the specimens carry Lacoe #973). Stark County, Illinois; donated by R.D. Lacoe.

griffithii, Neuropteris Lesq
Lesquereux (1884: 737-738, “1 95, figs. 3-8). Syntypes, USNM 10806+ 10807 (fig. 3), USNM 10808-
10812 (figs. 4-8 ennai ia “E vein,” Port Griffith, near Pittston, Luzerne County, Pennsylvania;
donated by R.D oe.

All of the sachsen carry Lacoe #417.

hamulosum, Rhacophyllum Lesq.
ep oe pl. 58, fig. 3; 1880: 321). Holotype, USNM 19445. Clinton, Missours; collected by
Dr. J.H. B

harveyi, Car diocarpus Lesq

Lesquereux (1884: 808, pl. 109, figs. 22, 23). Syntypes, USNM 26159 (fig. 23), USNM 26158 (ig, 22

L specimens carry Lacoe #299). Privatt’s Bank, Washington County, Arkansas; donated by R
acoe

harveyi, Sphenopteris Lesq

Lesquereux (1884: 766, pl. 103, figs. 7, 7b). Syntypes, USNM 14386 (fig. 7), USNM 2664, USNM 41229

2 of the specimens carry Lacoe #370). Sub-conglomerate, Arkansas; collected by F.L. Harvey; donated
y R.D. Lacoe.

hirsuta, Stemmatopteris Lesq
Lesquereux (1879: pl. 59, fig. 1; 1880: 337). Holotype, HU 7717.

66 BARTONIA

hispida, Pseudopecopteris Le
Lesquereux (1884: 755, pl. 98, fig. 3). Holotype, USNM 13737. Cannelton, Beaver County,

Pennsylvania.

ichthyoderma, Lepidophloios Lesq.
Lesquereux (1879: pl. 68, fig. 10; 1880: 426-427). Holotype, USNM 15974. Roof shale of Morris Coal,
Illinois; donated by S.S. Strong.

eaquale, Callipteridium Lesq.
Saeed (1879: pl. 33, se 2-5). Syntypes, USNM 11916-11919 (figs. 2-5 respectively). Cannelton,
Beaver County, Pennsylvan
It should be noted that vont are a number of specimens at SMP and ANSP which are from the type
locality and were identified by Lesquereux.

inaequilateralis, Sphenopteris Lesq
Lesquereux (1884: 765-766, pl. 103, , fis 4-5a). Syntype, USNM 41164 (fig. 4). Washington County,
Arkansas; collected by F.L. Harvey

incertus, Lepidostrobus Le
eral (1879: pl. 69, fig 16 1880: 442-443). Holotype, HU 8086. Shale above the coal, Morris,
Illinois

inflatus, Rhabdocarpus Le.
Lesquereux (1884: 815, pl. 110, fie 36). Syntypes, USNM 26116 6 Be 26), USNM 26117 (both specimens
carry Lacoe #991). Cannelton, Beaver County, Pennsylvania

insignis, Rhabd
Lesquereux (1879: pl. 85, i 26; 1880: 575). Holotype, USNM 41068. Seneca Mine, “coal seam F,”
Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

integrifolia, Whittleseya Le
Lesquereux (1879: pl. 4, fig. 2 "1880: 524). Holotype, AGS-SM 30. (Tr uscaloosa, Alabama)?

This specimen was sent to Lesquereux by Prof. Eugene A. Smith in the same lot as the holotype
of W. undulata Lesquereux (Harvard 7731).

intermedium, Spirangium

Lesq.
Lesquereux (1880: 521). Syntypes, USNM 19626-19630. Campbells Ledge, Pittston, Luzerne County;
Pennsylvania.

kansaseanus, Trigonocarpus Lesq
Lesquereux (1884: 822). Sync | USNM 26999-27002 (all of the specimens carry Lacoe #791). Osages
Kansas; donated by R.D. La

lacoei, Caulopteris Les

q-
Lesquereux (1880: 344). Lectotype, USNM 17106, Syntype, USNM 11051. “Coal #1,” Olyphant
Pennsylvania

USNM 17106 was designated by Merrill (1907) as the lectotype.

lacoei, Cordaites Lesa.
Lesquereux (1880: 535, pl. 87, figs. 2-4). Syntypes, USNM 18917 (fig. 2), USNM 18918 (fig. 3); a
18920 (fig. 4). Shale above “coal E,” Pittston, Luzern e County, Pennsylvania; donated by R. D.L

A fourth (unfigured) syntype is currently unlocated.

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 67

lacoei, Lycopodites Lesq.
Lesquereux (1884: 780-781, pl. 107, fig. 1). Holotype, USNM 10219 (Lacoe #524). “No. 1 vein,”
Olyphant, Pennsylvania; donated by R.D. Lacoe.

lacoei, Sigillaria Lesq.
Lesquereux (1879: pl. 72, fig. 12; 1880: 499-500). Syntype, USNM 16806 (fig. 12). “F vein,” Plymouth,
Pennsylvania; donated by R.D. Lacoe.

lanceolata, Calamostachys Lesq.

Lesquereux (1884: 715-717, pl. 91, figs. 1, 2; pl. 93, fig. 1). Syntypes, USNM 18063 (pl. 93, fig. 1; Lacoe
#558b), USNM 18276 (pl. 911, fig. 2), Dade County, Georgia; USNM 18062 (pl. 91, fig. 1; Lacoe
#558a), Arkansas; donated by R.D. Lacoe.

late-alatus, Cardiocarpus Lesq.
Lesquereux (1879: pl. 85, figs. 46, 47; 1880: 568-569). Syntypes, USNM 25684, USNM 25685 (fig. 46).
Campbells Ledge, Pittston, Luzerne County, Pennsylvania.

late-costatus, Rhabdocarpus Lesq.

Lesquereux (1884: 815, pl. 110, figs. 34, 35). Syatypes, USNM 25830 (fig. 34), USNM 25831 (fig. 35;
both specimens carry Lacoe #969). Campbells Ledge, Pittston, Luzerne County, Pennsylvania; donated
by R.D. Lacoe.

latifolium, Lepidodendron Lesq.
Lesquereux (1879: pl. 63, figs. 7, 8; 1880: 370). Holotype, HU 8200. Coal mines of Olyphant,
Pennsylvania.

The specimen was presented to Lesquereux by Mr. Edward Jones in 1860(?). The specimen also
carries Lesquereux’s personal catalogue number L. 20.

latior, Cardiocarpus Lesq.
Lesquereux (1884: 811, pl. 110, figs. 5, 11, 12). Syntypes, USNM 25585 (fig. 11), USNM 25586 (fig. 12;
both specimens carry Lacoe #977). Rockwood, Tennessee; donated by R.D. Lacoe

latior, Carpolithes Lesq.

Lesquereux (1884: 826, pl. 110, figs. 69, 70). Syntypes, USNM 15964 (fig. 69), USNM 15965, USNM
15966 (all of the specimens carry Lacoe #286). Campbells Ledge, Pittston, Luzerne County,
Pennsylvania; donated by R.D. Lacoe.

leptoderma, Sigillaria Le

sq. : a
Lesquereux (1879: pl. 72, fig. 10; 1880: 489-490). Syntype, USNM 16779 (fig. 10). “F vein,” Plymouth,
Pennsylvania; donated by R.D. Lacoe.

leveretti, Sigillaria Lesq. :
Lesquereux (1884: 800-801, 108, figs. 4, 5). Syatypes, USNM 16445 (fig. 4), USNM 16446 (fig. 5; both
specimens carry Lacoe #480). Spring Run, Augusta Township, Des Moines County, Iowa; donated by
R.D. Lacoe.

linearifolium, Lepidophyllum Le
, Lepidophyllum Lesq. ; |
Lesquereux (1879: pl. 65, fig. 39). Holotype, USNM 16067. Wilkes-Barre lower coal bed, Pennsylvania.

carpus Lesq.
Lesquereux (1884: 805, pl. 109, fig. 16). Holotype, USNM 25345 (Lacoe #983). Cannelton, Beaver
County, Pennsylvania.

68 BARTONIA

lineatus, Lepidocystis Les
Lesquereux (1879: pl. 69, fe 4; 1880: 454-455). — USNM 25254. South Salem vein of Port
Carbon, Pennsylvania; collected by William Lore

longicollis, Cardiocarpus Le
Lesquereux (1884: 808-809, <i “109, figs. 24, 25). Syntypes, USNM 26165 (figs. 24, = USNM 26166
(both specimens carry Lacoe #798). Dade County, Georgia; donated by R.D. Lac

macilenta, Archaeopteris Lesq.
Lesquereux (1884: 773, 775). Lectotype, USNM 16965 (on the same rock as 16964); isotypes, 16962-
16964, USNM 16968-16972, USNM 16978; syntypes, USNM 11320, USNM 16967b, USNM 16973,
USNM 16974, USNM 16976 (unlocated), USNM 16977, USNM 16979, USNM 196966, Meshoppen,
Pennsylvania; donated by R.D. Lacoe.

e suite carries Lacoe # 830. The lectotype was designated by Carluccio (1966). USNM 16%65,
16971, and 16974 each consists of a part and a counterpart. USNM 16977 is the counterpart only; the
part is currently missing.

macrocystis, Lepidostrobus Lesq.
Lesquereux (1879: pl. 69, figs. 1, 2). — USNM 42298 (fig. 1), Mercer County, Illinois; USNM
16130 (fig. 2), Coal Creek, West Vir

The specimen in fig. 1 was mnie ae to Lesquereux by Dr. Salisbury. The species was later raised
to a subgeneric rank by Lesquereux (1880: 443) to accommodate three new species; L. salisburyi Les.
(erected upon the same figures as L. macrocystis), L. quadratus Lesq., and L. mansfieldi Lesq. Chaloner
& Boureaux (1967: 675-676) noted that this taxon merely represents a preservational form of Stigmaria
ficoides (Sternberg), in which the root structure has collapse d and the surrounding tissue has folded into
sections which, coupled with the compressed xylem, gives the appearance of a cross-sectioned cone.

mamillatus, Rhabdocarpus Le
Lesquereux (1884: 816-817, pl. 110, figs. 39-42). Se USNM 26762-5 (figs. 39-42 respectively; al
of the specimens carry Lacoe #665). Stark County, Lllinois

mansfieldi, Callipteridium Lesa.

Lesquereux (1879: pl. 27, figs. 1, 2; 1880: 166). Syntypes, USNM 11930 (fig. 1), USNM 11931+11932
(fig. 2); syntype(?), SMP 11331 (Lesquereux #258 [11331 (Lesquereux 1889: 111)]), Cannelton, Beaver
County, Pennsylvania; collected by I.F. Mansfield.

mansfieldi, Caulopteris Le

Lesquereux (1879: pl. 60, fig, 3; 1880: 346-347). Syntype, USNM 17118 (fig. 3). Cannelton, Beaver
County, Pennsylvania; collected and donated by LF. Mansfield.

mansfieldi, Halonia Lesq.

Lesquereux (1879: we 69, fig. 2; 1880: 414-416). Syntype, USNM 15808 (fig. 2). Cannelton, Beaver
County, Pennsylvania

mansfieldi, Lepidophyllum

Lesquereux (1879: pl. 69, fig. zr 1880: 449-450). Syntype, USNM 15918 (fig. 34). Cannelton, Beaver
County, Pennsylvania; collected by LF. Mansfield.

mansfieldi, Rhabdocarpum Lesq.

Lesquereux (1879: pl. 84, > 21). Holotype, USNM 15977. Cannelton, Beaver County, Penasy Ivan;
collected by I.F. Mansfiel

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 69

This species was erected by name and figure only in the Atlas and was not included in the text of

the Coal Flora

arginata, Megalopteris? Les
oe (1879: pl. 24, fig. 4; 1880: 152-153). Holotype, USNM, 11700. Port Byron, Illinois; collected
by Mr. Southwell.

marginata, Sigillaria Lesq
Lesquereux (1879: pl. 1, fe 5; 1880: 498-499). Holotype, USNM 16798 (Lacoe 460). Plymouth,
Pennsylvania; donated by R.D. Lacoe.

membranaceum, Callipteridium Lesq.
Lesquereux (1879: pl. 27, figs. 4-8; 1880: 172-173). oe USNM 13181-13183 (figs. 4-6 respectively),
USNM 13187. Clinton, Henry County, Missour

The specimen in fig. 8 is currently unlocated an White (1899: 120, pl. 38, figs. 4, 5) placed USNM
13181, 13183 into synonomy with Pecopteris clintoni Lesq. The other specimens, however were
retained in this species.

membranaceum, Rhacophyllum Lesq. : '
Lesquereux (1879: pl. 58, figs. 1, 2). Syntypes, USNM 19465 (fig. 1), USNM 19466 (fig. 2). Clinton,
Henry County, Missouri.

microphylla, Eremopteris Lesq.
Lesquereux (1879: pl. 52, figs. 6, 73 1880: 296). Syntypes, USNM 40110 (fig. 6), HU 7704 (fig. 7).
Helena coal mines, Alabama; collected by Eugene A. Smith.

microphylla, Lesleya Le

Lesquereux (1884: 831- $39), Lectotype, USNM 11744 (Lacoe #314). Osage, Kansas; donated by R.D.
Lacoe.

USNM 11744 was designated as the lectotype by Merrill (1907: 213).

microphylla, Whittleseya Lesq.
Lesquereux (1884: 843). Syntypes, USNM 18880-18882. Washington County, Arkansas; communicated
ey.

microstigma, Stemmatopteris Lesq. i
Lesquereux (1884: 838). Syntypes, USNM 17134-17137 (all of the specimens carry Lacoe #612).
Cannelton, Beaver County, Pennsylvania; donated by R.D. Lacoe.

mimica, Stemmatopteris
orien ‘eth pl. 59, he ‘A 1880: 341-342). Holotype, USNM 17141. Cannelton, Beaver County,
ennsylv.

minor, Macrostachya Le
Lesquereux (1884: 829, ol 3, figs. 19, 19a). raat i USNM 18301 (fig. 19; Lacoe #212). Campbells

, Pittston, Luzerne County, Pennsylv
The lectotype was designated by Merrill (1907. 218).

minutum, Lepidophyllum ,
Lesquereux (1884: 787, pl. = % 9). Holotype, USNM 16243 (Lacoe #673). Archbald, Lackawanna
County, Pennsylvania; donated by R.D. Lacoe.

70 BARTONIA

issouriensis, Eremopteris Le
Lesquereux (1879: pl. 53, fig. 8, “ 1880: 295). Holotype, USNM 30853. Clinton, Missouri; collected
by Dr. J.H Britts.

missouriensis, Neuropteris Lesq.
Lesquereux (1879: pl. 7, figs. 5, 6; 1880: 104). Syntype, USNM 30861 (fig. 6). Clinton, Henry County,
Missouri; communicated by J.H. Britts.

The specimen fig. 5 is currently unlocated.

monstruosa, Odontopteris Lesq
Lesquereux (1884: 741-742, pl. 7, figs. 1-3). Syntypes, USNM 14831 (fig. 2), USNM 19358 (fig. 3).
annelton, Beaver County, Pennsylvania.
The specimen in fig. 1 is currently unlocated.

morrisianum, Lepidophyllum Lesq.
Lesquereux (1879: pl. 69, figs. 40, 41; 1880: 448-449). Syntypes, ne — (fig. 40), USNM 15916
(fig. 41). Shale over the coal of Morris, Illinois; collected by S.S. $

multiplicatum, Spirangium Lesq.
Lesquereux (1879: pl. 75, fig. 11; 1880: _en Syntype, USNM 19675 (fig. 11). Mazon Creek,
Grundy County, Illinois; collected by S.S. Strong

neuropteroides, Callipteridium Le
Lesquereux (1879: pl. 27, fig. 3; 1880: 166-167). Holotype, USNM 11878 + 11879. Mazon Creek, Grundy
County, Illinois

oblongifolia, Neuropteris Lesq.

Lesquereux (1884: 732-733, pl. 94; fig. 3, pl. 95, fig. 2). Syntypes, USNM 10070 (pl. 94, fig. 3) USNM
10071 (pl. 95, fig. 2), USNM 10072 (all of the specimens carry Lacoe #951). Cannelton, Beaver County,
Pennsylvania; donated by R.D. Lacoe.

ophioglossoides, Sorocladus Les
Lesquereux (1879: pl. 48, fig. 11). Hologmpe, USNM 14270. Clinton County, Missouri; collected by Dr.
J.H Britts.

ornata, Pecopteris Les rs
Lesquereux (1884: 760, pl. 111, figs. 30-30b). Syntypes, USNM 13387+13388, USNM 13385 (all of the
specimens carry Lacoe #1008). Mazon Creek, Grundy County, Illinois; donated by R.D. Lacoe.

ovalis, Cardiocarpus
Lesquereux an 810, a 109, figs. 8, 9). Syntypes, USNM 25517, USNM 25516 (fig. 9). Washingto
County, Arkansas

ovalis, Cordaicarpus Lesq. nom. nov. pro. C. apiculatus Lesq.
[See Cordaicarpus apiculatus Lesq.]

ovalis, Sigillaria Lesq. th,
Lesquereux (1879: pl. 71, figs. 7, ig 495). Syntype, USNM 16553 (fig. 7). “F vein,” Plymou
Pennsylvania; donated by R.D

The specimen in fig. 8 is More canlocaiie

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 71

ovatus, Cordaianthus Lesq.
Lesquereux (1880: 545-546, pl. 76, figs. 5, 5a). Lectotype, SMP 11340. Cannelton, Beaver County,
Pennsylvania.

The lectotype was designated by Darrah (1969: 84) by assumption of a holotype. Darrah (1969: 84)
provided a detailed comparison between the lectotype and the original figure (drawn by A.M. Rickey)
which is reproduced below. (The words within the brackets were added by the author.)

“The sketch indicates the ber, g d symmetry of the strobili. The overall

size is reduced approximately 20 percent although the precise limits of the specimen, having the

same monochrome black color of the matrix, are difficult to determine. Minor details have been
idealized or omitted from [the] drawing. The reliability of the illustration without the description

and original specimen is very poor.

pachytesta, Cardiocarpus Lesq.
Lesquereux (1880: 565). Syntypes, USNM 35482-25486. Campbells Ledge, Pittston, Luzerne County,
Pennsylvania; donated by R.D. Lacoe.

pachytesta, Rhabdocarpus Lesq. ;
Lesquereux (1884: 816, pl. 110, figs. 37, 38). Syntypes, USNM 26796 + 26797 (fig. 37), USNM 26798 (fig.
38; both specimens carry Lacoe #989). Mazon Creek, Grundy County, Llinois; donated by R.D. Lacoe.

pardei, Callipteridium Lesq.
Lesquereux (1879: pl. 26, figs. 2, 3). Syntypes, LC uncatalogued (destroyed?).

Both specimens may have been destroyed in a fire at Lafayette College. However, this has not been
confirmed and specimens from the collections of Lafayette College are known to exist in other
institutions. Additionally, at least part of the LC herbarium ended up in ANSP. The Paleobotany
collection at ANSP was searched for the specimens; however they were not found.

patens, Cardiocarpus Lesa.
Lesquereux (1884: 807, pl. 110, fig. 3). Holotype, USNM 25504. Washington County, Arkansas;
Communicated by F.L. Harvey.

Patens, Odontopteris Lesa.
Lesquereux (1884: 740-741, pl. 97, fig. 7). Syntypes, USNM 11274 (fig 7), USNM 11275, USNM 11276
{al of the specimens carry Lacoe #890). Rhode Island

Pectinatus, Lepidocystis Lesq.
Lesquereux (1879: pl. 69, fig. 3; 1880: 454). Holotype, USNM 25252 (Lacoe #432). Subconglomerate
of Campbells Ledge, Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

Pendulus, Lycopodites Lesq.
Lesquereux (1879: pl. 62, fig. 2; 1880: 357). Holotype, HU 7722. Shale of Morris co
€ specimen also carries Lesquereux’s personal catalogue number Ly. 11.

al, Illinois.

b] q-
“esquereux (1884: 825, pl. 111, figs. 22-24). Syntypes, USNM 25950, USNM so po oagetonaar
epg “a carry Lacoe #980). Campbells Ledge, Pittston, Luzerne County, Pennsy ;

Perpusillus, Trigonocarpus Lesq.
“stereux (1884: 820.871, pl. 110, figs. 58-61). Symtypes, USNM 26985, USNM 26982-26984 (all -

© specimens carry Lacoe #971). Stark County, Illinois; donated by R.D. Lacoe.

72 BARTONIA

pittstoniana, Sigillari
Lesquereux (1879: pl. 71, 4 4; 1880: 493). Syntype, USNM 16709 (fig. 4). Plymouth, Pennsylvania;
donated by R.D. Lacoe.

planus, Sporocystis Lesq.
Lesquereux (1879: pl. 69, fig. 15; 1880: 458-459). Syntypes, ANSP 980 icone #420 [11493]
(Lesquereux 1889: 117)]), USNM 25196 - 25199; Syntypes(?), SMP 11492 (Lesquereux #419c [Lesquereux
1889: 117]), SMP 11463 oo 390a [Lesquereux 1889: 116)). ‘Tnter per ee Pittston,
Luzerne County, Pennsylvan
While Watt (1974) gave USNM 25196 as the figured specimen, this must be an error. The figure is
d that no one specimen can be discerned as the type. In fact, Spamer and Lendemer (2000)
penser of ANSP 980 that “the illustration by which this species was erected by name and figure
only is so Beneralized that the identity of this specimen as one of those used to figure the species is
uncertain.” Thus, here I include the specimens at USNM as well as those at ANSP and SMP in the
type series simply because all are known to have been in the possession of Lesquereux at the time of
publication.

polita, Stemmatopteris Lesq.
Lesquereux (1879: pl. 59, fig. 6; 1880: 342). Holotype, USNM 17096. Cannelton, Beaver County,
Pennsylvania; collected by I.F. Mansfield.

praelongus, Lepidostrobus Lesq.
Lesquereux (1880: 433). Holotype, ANSP 3727 (Lacoe #559, Lesquereux #62 [11135] (Lesquereux 1887:
102)})+(?) USNM 41140. “C vein,” Everhart’s colliery, near Pittston, Luzerne County, Pennsylv.
Lesquereux’s citation (1889: 102) for this specimen lists the locality as Cannelton, Beaver Godan
Pennsylvania. Despite this difference in data the specimen seems to be that upon which the miei
was based. The lithology of the specimen also corresponds more with the Pittston locality than t hat
of Cannelton. There is also a specimen in the National Museum (USNM 41140) that is listed as being
the type of this species. The specimen has not been examined and was not listed by Merrill (1907).

pusillus, Cardiocarpus Lesq

Lesquereux (1884: 815, pl. 110, figs. 31-33). Syntypes, USNM 25862 (fig. a) USNM 25923 (ig. 32)
USNM 25924 (fig. 33; all of the specimens carry Lacoe #967), Coal “E and F,” Wilkes Barre,
Pennsylvania; donated by R.D. Lacoe.

quadratifolia, Pecopteris Lesq,
Lesquereux (1880: 234). Lectotype, USNM 12702; isotypes(?), SMP 11175 (Lesquereux #125 [11195]
(Lesquereux 1889: 106)]), SMP 11175 (Lesquereux #102b [11175)] (Lesquereux 1889: 105)]), SMP 11185
(Lesquereux #112a [11185] (Lesquereux 1889: 105)]), Cannelton, Beaver County, Pennsylvania.

The lectotype was designated by Merrill (1907).

radiatus, Cordaites Le
Lesquereux (1880: 540, al 87, figs. 5-7). Syntypes, USNM 18907 (fig. 5), USNM 18908 (fig. 6);
syntype(?) SMP 11501. Cannelton, Beaver County, Pennsylvan

The specimens were collected and sent to Lesquereux by LF. Mansfield. The third, unlocated figured
syntype (fig. 7) may be SMP 11501 (Lesquereux #428) which is currently in the William Pena gnu
in Harrisburg, Pennsylvania. A fourth specimen (unfigured) also remains unlocated.

amosum, Dictyoph Lesq.
pas (1884: 827). omer SMP 11317 (Lesquereux #244). Head of Hill Creek, Cha
township, Tioga County, Pennsylvania, Upper Devonian; collected by Mr. Sherw ood, July <

{
;

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 73

SMP 11317 consists of twelve specimens. Lesquereux noted fifteen specimens however, upon
examination of the collection in 1889 only twelve were found (Lesquereux 1889: 110).
. 7
rigidum, Callipteridium Lesq. .
Lesquereux (1884: 746-747, pl. 99, figs. 1-2a). Syntypes, USNM 13602 (fig. 1), USNM 13601 (fig. 2; both
specimens carry Lacoe #716). Olyphant, Pennsylvania.

rigidum, Lepidodendron Lesq.
Lesquereux (1884: 839). Syntypes, USNM 15515-15517 (all of the specimens carry Lacoe #918). Dade
Mine, Georgia; donated by R.D. Lacoe.

robusta, Alethopteris Lesq.
Lesquereux (1884: 835). Syntypes, USNM 12174-12177 (all of the specimens carry Lacoe #743).
Cannelton, Beaver County, Pennsylvania; donated by R.D. Lacoe.

robusta, Pecopteris Lesq. ;
Lesquereux (1879: pl. 39, figs. 7, 8; 1880: 229-230). Syatypes, USNM 12524 (fig. 8), USNM 12523 (fig.
7). Subconglomerate ledge, Pittston, Luzerne County, Pennsylvania.

rotundifolium, Idiophyllum Lesq. -
Lesquereux (1879: pl. 23, fig. 11). Holotype, USNM 10258. Mazon Creek, Grundy County, Illinois.

royi, Sphenopteris Lesq. ;
Lesquereux (1884: 768, pl. 104, figs. 7-10). Syntypes, USNM 19853 (fig. 7), USNM 19852 (fig. 8),
USNM 19851 (fig. 9), Rockwood, Tennessee; USNM 19854 (fig. 10), USNM 19855, Washington
County, Arkansas; donated by R.D. Lacoe.

of the specimens carry Lacoe #1037.

Tugosus, Cordaianthus Lesq. a ‘
Lesquereux (1884: 803-804). Syntypes, USNM 19214 (Lacoe #929), “Coal C,” Shamokin, Pennsylvania;
USNM 19211 (Lacoe #943), Mazon Creek, Grundy County, Illinois; donated by R.D. Lacoe.

salisburyi, Lepidostrobus Lesq. nom. nov. pro. L. macrocystis Lesq.

Lnele-chieVineo 443-444). pe seen USNM 42298, Mercer County, Illinois; USNM 16130, Coal
Creek, West Virginia. , :

_ The specimen in fig. 1 was presented to Lesquereux by Dr. Salisbury. This species was ere :
indication to the same figures upon which Lesquereux had based Lepidostrobus macrocystis. When he
taised the species to subgeneric status he proposed the name L. salisburyt to replace L. macrocystis.
Thus, L. salisburyi Lesquereux, is treated here as a nomen novum for L. macrocystis. See the entry for
L. macrocystis for notes on the synonomy provided by Chaloner & Boureaux (1967: 675-676).

cted with

‘cutatum, Lepidodendron Les ; :

5) q- i |
Lesquereux (1879: pl. 63, fig. 6; 1880: 369-370). Syntype, USNM 15448 (fig. 6). Clinton, Missouri,
collected by Dr. J.H Britts.

‘erpillifolia, Pecopteris Le |
sq- :
2, USNME (1879: pl. 46, figs. 1-3; 1880: 237-238). Syarypes, USNM 13929 (fig. 1), USNM 13190 (fig.
13931 (fig. 3). Mazon Creek, Grundy County, Ulinols. | ‘
mr worth noting that Lesquereux sent a series of specimens of this ben : arr
Prongniart However, Brongniart never gave Lesquereux his views on the species. 1 have no
M locate these specimens (though if they still exist they are most
Useum of Paris).

74 BARTONIA

serrata, Megalopteris Lesq.
Lesquereux (1884: 834). Syntypes, USNM 11463, USNM 11464 (both specimens carry Lacoe #798).
Rushville, Ohio; donated by R.D. Lacoe.

sigillarioides, Lepidophloios Lesq.

Lesquereux (1879: pl. 68, fig. 8; 1880: 13-14). Holotype, USNM 6173. Clinton, Missouri.
Lesquereux cited Mr. I.H. Britts as the donator of the specimen. This is, however in error. The

specimen were sent to Lesquereux by Mr. J.H. Britts.

simplex, Asterophycus Lesq.
— (1879: pl. B, figs. 7, 8; 1880: 13-14). Syntypes, USNM 10255. Beaver County, Pennsylvania
of the illustrated specimens are preserved on the same r

simplex, Cardiocarpus Lesq
Lesquereux (1879: pl. 85, re 49, 50; 1880: 569). Syntypes, USNM 15630 (fig. 49), USNM 15631 (fig
50). Campbells Ledge, Pittston, Luzerne County, Pennsylvania.

simplex, Lycopodites Lesq.
Lesquereux (1884: 779-780, pl. 106, fig. 2). Holotype, USNM 15552 (Lacoe 258). Campbells Ledge,
Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

smithsii, Neuropteris Lesq
Lesquereux (1879: pl. 13, nia 1-3; 1880: 106). ee AGS-SM 25c. Black Creek vein, New Castle
Coal, Alabama, collected by Eugene A. Smi

Of the three specimens upon which “nani based this species, only this single representative
has been found (it is illustrated in Lesquereux’s fig. 2). The lectotype was designated by Goubet et al.
(2000: 19). Lesquereux (1880: 106) cited page 76 of the 1876 Geological Report of Alabama as as the
location of the original description for this species; however, it is a nomen nudum there. Thus, by
default, the place of publication shifts to the next available date which is the Avlas to the Coal Flora.
Goubet et al. (2000: 19-23) also provide a review of the systematic placement of this taxon in which
it is placed into the genus Neuralethopteris Cremer ex Laveine.

smithsii, Taeniopteris Lesq. ran
Lesquereux (1879: pl. 25, fig. 7; 1880: 153-154). Holotype, AGS-SM 78, Alabama Coal Fields, ol
Coal Mines, Alabama; from Professor Eugene A. Smith (locality inferred by Lesquereux (1880: 154)

solida, Sphenopteris Lesq
Lesquereux (1884: 769, pl. 01, fig 3. Halorgen USNM 14213414214. Mazon Creek, Grundy Count)
ois; communicated by F.T.
The specimens are part and Aine

southwelli, Megalopteris Lesq. Coal
Lesquereux (1879: pl. 24, fig. 1; 1880: 158-149). Syntype, USNM 41171 (fig. 1). Lower bed of me
Measures, sub-conglomerate, near Port Byron, Illinois; collected by Mr. J.H. Southwell.

speciosa, Pseudopecopteris Lesq.
Lesquereux (1879: pl. 51, fig. 1; 1880: 216-217). Holotype, USNM 41170. Helena coal mines,
collected by Professor Eugene A. Smith.

Alabama:

specious, Cardiocarpus Lesq County,
Lesquereux (1884: 807-808, a 110, fig. 0. Holotype, USNM 26205. Cannelton, Beaver
Pennsylvania; collected by LF. Mansfield

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 75

spectabilis, Lepidostrobus Lesq.
Lesquereux (1880: 435). Syntypes, USNM 16059, USNM 16060. Cannelton, Beaver County,
Pennsylvania.

sphenophyllifolia, Archaeopteris Lesq.
Lesquereux (1884: 773, 775). Syntypes, USNM 16980-16982 (all of the specimens carry Lacoe #829).
Meshoppen, Pennsylvania; donated by R.D.

sphenopterioides, Odontopteris Lesq.
Lesquereux (1879: pl. 21, figs. 3, 4; 1880: 139-140). Syntypes, USNM 13531 (fig. 3), Mazon Creek,
Grundy County, Illinois; USNM 13532 (fig. 4), Clinton County, Missouri.

The specimen in fig. 3 was communicated to Lesquereux by Dr. J.H. Brits.

spicatus, Cordaianthus
Lesquereux (1884: 802- 2 oat 109, fig. 3). Syntype, USNM 19226 (fig. 3; Lacoe #940). Campbells
Ledge, Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

spinosum, Rhacophyllum Le
Lesquereux (1879: pl. 58, fig. Pi 5 1880: 320-321). Syntype, USNM 19436 (fig. 4). Clinton, Missouri;
collected by Dr. J.H. Britts.

The specimen in fig. 5 is currently unlocated.

squamiferum, Lepidodendron Lesq.
Lesquereux (1879. pl. 62, fig. 3; 1880: 376-377). Holotype, AGS-SM 18. Helena coal mines, Alabama;
donated by Professor Eugene A. Smith.

squamosa, Rachiopteris Le
Lesquereux (1884: 838-839). ‘Saas USNM 19608-19610 (all of the specimens carry Lacoe #456).
Cannelton, Beaver County, Pennsylvania; donated by R.D. Lacoe

Squamosa, Stemmatopteris Lesq.
Lesquereux (1879: pl. 59, fig. 2; 1880: 339). Holotype(?), USNM 17073 (fig. 2). Cannelton, Beaver
County, Pennsylvania, donated by LF. Mansfield.

stantoni, Lepidophyllum Le
Lesqureux (1884: 841). PE USNM 16221-16223 (all of the specimens carry Lacoe #657). Stanton
€, Wilkes-Barre, Pennsylvania; donated by R.D. Lacoe.

‘tarkianus, Trigonocarpus
Lesquereux (1884: 822, pl. vial gs 7.13). Syntypes, USNM 26983, USNM 26982, USNM 26504
USNM 26985 (all of the specimens carry Lacoe #972). Stark County, Illinois; donated by acoe

striata, Dechenia Lesq.
Lesquereux (a7s. pl. e7, fig. 3). Holotype, USNM 10259. Alta, Illinois; donated by R.D. Lacoe(?).

‘ubcrenulata, Pseudo opecopteris

ir ole pl. 37, figs. 7, 3). pce HU 6236 (fig, 7), HU 6237 (fig. 8)- Mazon Creek, Grundy

“ berenulata, Odontopteris Le
teux (1879: pl. He figs. 5, @. Syntypes, USNM 35317 (fig. 5), USNM 10378 (fig. 6). Mazon

Creek K, Grundy C County, Illinois; donated by R.D. Lacoe(’).

76 BARTONIA

subcylindricus, Trigonocarpus Les
Lesquereux (1879:pl. 84, figs. 9, 10). aie, HU 8035. Locality unknown.

subfalcata, Neuropteris Les
Lesquereux (1879: pl. 13, figs, 5, 6; 1880: 102-103). Syntypes, AGS-SM 32 (lot consists of two
specimens). Gray shale, Helena coal mine?, Alabama; communicated by Professor Eugene A. Smith.

sub-globosus, Rhabd
Lesquereux (1884: 817, pl. 110, figs. 43-45, 62). Syntypes, USNM 26331, USNM 26330 (fig. 43), USNM
26332 (fig. 44; all of the specimens carry Lacoe #978), Butler Dam, near Pittston, Luzerne County,
Pennsylvania.

The specimen in fig. 45 is currently unlocated.

tenax, Rhabdocarpus Le
Lesquereux (1884: 818, pl. NO, figs. 25, 26). Syntypes, USNM 26458, USNM 26459. Cannelton, Beaver
ounty, Pennsylvania.

timidum, Lepidophyllum Le
Lesquereux = 448). ate USNM 16218 (Lacoe #565). Wilkes-Barre, Pennsylvania; donated by

tracyana, Sphenopteris Lesq.
Lesquereux (1884: 766-76, pl. “to, fig. 2). Holotype, USNM 14132 (Lacoe #930). Tracy City, Tennessee.

transsectus, Carpolithes L
Lesquereux (1884: 826, pl. iL figs. 27-27b). Syntypes, USNM 25810, USNM 25811 (both specimens
carry Lacoe #279). Washington County, Arkansas; communicated by F.L. Harvey.

trichoideum, Rhacophyllum Lesq.
Lesquereux (1880: 322). Syntypes, USNM 19453, USNM 19454. Oakwood colliery, Wilkes-Barre,
Pennsylvania.

truncata, Taeniopteris Lesq.
Lesquereux (1884: 743-744, iA 94, fig. 8). Holotype, USNM 11699 (Lacoe #860). Campbells: Ledge,
Pittston, Luzerne County, Pennsylvania; donated by R.D. Lacoe.

truncatum, Rhacophyllum
Lesquereux (1879: pl. 50, fig. e “1880: 311-312). Holotype, USNM 19414. Red shale of the Vespertis
on the bluffs of the Susquehanna River above Pittston, Coxten Narrows, Luzerne e Cousty;
Pennsylvania.

undulata, Whittleseya Les

Lesquereux (1879: pl. 4, fig. > 1880: 525). Holotype, HU 7731. (Tuscaloosa, Alabama)(?) if
The specimen was sent to Lesquereux by Prof. Eugene A. Smith in the same lot as the holotype

W. integrifolia Lesquereux.

vestita, Pecopteris Lesq. fg
Lesquereux (1879: pl. 43, figs. 17; Pu 252-253). Syntypes, USNM 13127 (fig. 1), USNM 13160 (8
3). Clinton, Henry County,
All specimens with the sees of those in figs. 1 and 3 are currently unlocated.

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 77

williamsi, Sigillaria Lesq.
Lesquereux (1880: 488-489). Syntypes, USNM 16745 (Lacoe #719), USNM 16746 (Lacoe #536).
Olyphant, Pennsylvania.

zonulatus, Cardiocarpus Lesq.
Lesquereux (1879: pl. 85, figs. 44, 45; 1880: 568). Syntypes. USNM 25678 (fig. 44), USNM 25679 +27680
(fig. 45). Subconglomerate, Pittston; donated by R.D. Lacoe.

ACKNOWLEDGMENTS

Firstly, I wish to thank Hermann Pfefferkorn (PENN), Ethel McCardell, and Lisa Mikula
for reviewing the early drafts of the manuscript, as well as C.J. Cleal, Robert A. Gastaldo,
athird anonymous reviewer, and again Hermann Pfefferkorn for their helpful comments and
copious notes on the later drafts of the manuscript. Thanks are also extended to William
DiMichele (USNM), Robert Sullivan (SMP), Linda Klise (YPM), and Susan Rossi-Wilcox
(HU) for their help in locating and verifying the specimens in their institutions. Lastly, I
wish to draw attention to the small army of previous workers without whom many of these
specimens would have been lost, destroyed, or left unrecognized in the vast collections of
our nation’s museums.

LITERATURE CITED

CARLUCCIO, L. M., HUEBER, F. M., AND BANKS, H. P. 1966. Archaeopteris macilenta, anatomy and
morphology of its frond. American Journal of Botany 54: 719-730.

CHALONER, W.G. & BOUREAUX, E. 1967. Lycophyta, in (E. Boureaux, Editor) Traite de Paleobotan-
ique 2: 435-782.
DARRAH, W. C. 1969. A Critical Review of the Upper Pennsylvanian Floras of [the] Eastern United

States with Notes on the Mazon Creek Flora of Illinois. W.C. Darrah, Gettysburg. 220 pp.
UBET, P., PFEFFERKORN, H., GILLESPIE, W. 2000. Neuralethopterids (T rigonocarpalean
Pteridosperms) from the Early Pennsylvanian of Eastern North America. PaleoBios 20: 11-37.
LESQUEREUX, L. 1879. Atlas to the Coal Flora of Pennsylvania, and the Carboniferous Formation
Throughout the United States. Board of Commissioners, for Second Geological Survey, Harr isburg.
Second Geological Survey of Pennsylvania, Report of Progress. 85 pp. P1-P18, pls. A, B. ;
LESQUEREUX, L. 1880. Description of the Coal Flora of the Carboniferous Formation in Pennsylvania
and Throughout the United States. Board of Commissioners, for Second Geological Survey,
Harrisburg. Second Geological Survey of Pennsylvania, Report of Progress (2 vols.). 694 pp.. pls.
86,87. Vol. I:1. Cellular cryptogamous plants, fungi, thalassophytes. 2. Vascular cryptogamous
Plants, Calamariae, Filicaceae (ferns). Vol. I: 1. Lycopodiaceae. 2. Sigillariae. 3. Gymmosper ms,
LESQUEREUX, L. 1884. Description of the Coal Flora of the Carboniferous Formation in Pennsylvania
and Throughout the United States. Board of Commissioners, for Second Geological Survey,
ea isburg. Second Geological Survey of Pennsylvania, Report of Progress. Pp. P695-P977, pls. 88-

LESQUEREUX, L. 1889. Fossil plant collection. Pp. 99-122 in Geological Survey of Penn haan
rtalogue of the Geological Museum, Part 3, Board of Commissioners, for he Seton ST
RRILL, G. P. 1907. Catalogue of the T d Figured Specimens of Fossils, Minerals, and Ores.
2, . ype and rigured spec ;
Government Printing Office, Washington. Bulletin of the United States National Museum, Bulletin

: z . Or ,
52, part 2: Fossil Vertebrates. 2. Fossil Plants. 3. Minerals, Rocks, and Ores app “ite oo

78 BARTONIA

SPAMER, E. E., LENDEMER, J. C. 2000. Type specimens of fossil plants in the Paleobotany Collection,
of the Academy of Natural Sciences of Philadelphia — revised catalogue. Proceedings of the
Academy of Natural Sciences of Philadelphia 150: 315-328.

STOCKMANS, F. 1933. Les Neuropteridees des Bassins houillers Belges (1° pt.), Memoirs of the Royal
Museum of Natural History of Belgium 57: 1-61.

WATT, A. D. 1974. Catalog of Specimens Illustrated in the Lesquereux Coal Flora. Paleobotanical
Section of the Botanical Society of America. University of Indiana, Bloomington, Indiana. 42 pp.

WHITE, D. 1899. Fossil flora of the Lower Coal Measures of Missouri. United States Geological Survey
Monograph, No. 37.

APPENDIX A. List of taxa for which no types have been located. The taxa are listed
alphabetically by genus in their original combinations as given by Lesquereaux.

Taxon Year Described Taxon Year Described
Calamostachys praelongus 1880 Megalopteris rectinervis 1884
Carpolithes minimus 1884 Megaphitum grand’euryt 1880
Conostrichus prolifer 1880 Neuropteris blissii 1884
Cordaianthus scabar 1884 Neuropteris carrit 1884
Cordaicarpus cinctus 1884 Neuropteris obscura 1880
Cordaicarpus stabilis 1884 Odontopteris affinis 1884
Dendrophycus desorii 1884 Odontopteris dilatata 1884
Halonia 1880 Pecopteris carvit 1884
Lepidocystis quadrangularis 1880 Pecopteris obsoleta 1884
Lepidocystis obtusus 1880 Pecopteris venulosa 1880
Beart heme lanceolatum 1880 Pseudopecopteris denudata 1880
Lepidophyllum cultriforme 1884 Rhabdocarpus emarginatus 1884
Lepidostrobus aldrichii 1880 Rhabdocarpus howardi 1880
Lepidostrobus lacoei 1880 Sigillaria lorenzi 1880
Lepidostrobus latus 1884 Sphenopteris microcarpa 1880
Lepidostrobus mansfieldi 1880 Sphenopteris pseudo-murrayana 1880
Lepidostrobus quadratus 1880 Stemmatopteris schimperi 1880
Lycopodites flexifolius 1884 Stigmaria amoena 1880
Lycopodites ortoni 1880 Trigonocarpus multistriatus 1884
Lycopodites strictus 1880 Trigonocarpus saffordi 1880
Megalopteris abbreviata 1880

APPENDIX B. List of taxa erected in the Coal Flora arranged alphabetically by genus 6 -
original combinations as given by Lesquereaux. <p for which no type specimens “
been located are listed separately in Appendix A

ee
Althopteris amibgua Asterophycus simplex

Alethopteris evansi Asterophyllites fasciculatus

Alethopteris gibsoni Calamites gracilis

Alethopteris robusta Calamostachys brevifolia

Annularia emersoni Calamostachys lanceolata

Archaeopteris denticulata Callipteridium aldricht

Archaeopteris macilenta Callipteridium ineaquale

Archaeopteris sphenophyllifolia Callipteridium mansfieldi

COAL FLORA TYPE SPECIMENS OF LEO LESQUEREUX 79

Callipteridium membranaceum
Callipteridium neuropteroides
Callipteridium pardet

Cardiocarpus conglobatus
Cardiocarpus crassus
Cardiocarpus dilatatus
Cardiocarpus diminutivus
Cardiocarpus diplotesta
Cardiocarpus divergens
Cardiocarpus fasciciulatus
Cardiocarpus harveyi
Cardiocarpus late-alatus
Cardiocarpus latior
Cardiocarpus longicollis
Cardiocarpus ovalis
Cardiocarpus pachytesta
Cardiocarpus patens
Cardiocarpus pusillus
Cardiocarpus simplex
Cardiocarpus specious

Carpolithes perpusillus

Caultoperis mansfieldi
Conostrichus broadheadi
Cordaicarpus lineatus
Cordaicarpus ovalis
Cordaianthus dichotomus
Cordaianthus ebracteatus
Cordaianthys flexuosus
Cordaianthus ovatus

lanthus rugosus
Cordaianthus Spicatus

ites lacoei
Cordaites radiatus
Danaeites emersoni
Dechenia striata
Dicranophyllum dichotomum
E ophytum ramosum
E teris cheathami

"emopteris microphylla

Eremopteris missouriensis

‘ites gracilis

_ a mansfieldi
Idiopbyllum rotundifolium

Knorria compacta

Lepidodendron andrewsi
Lepidodendron brittsii
Lepidodendron cuspidatum
Lepidodendron cyclostigma
Lepidodendron latifolium
Lepidodendron rigidum

Lepidophloios sigillarioides
Lepidophyllum campbellianum
Lepidophyllum coriaceum

Lepidophyllum linearifolium
Lepidophyllum mansfieldi
Lepidophyllum minutum
Lepidophyllum morrisianum
Lepidophyllum stantoni
Lepidophyllum timidum
Lepidostrobus butleri
Lepidostrobus incertus
Lepidostrobus macrocystis
Lepidostrobus praelongus
Lepidostrobus salisburyt

] 15

Lycopodites arborescens
Lycopodites lacoet
Lycopodites pendulus
Lycopodites simplex
Macrostachya communis
Macrostachya minor
Megaloptert:

Neuropterts aspera —
Neuropteris biformis

Neuropteris decipiens
Neuropteris elrodi :
Neuropteris griffithit

80

Neuropteris missouriensis
Neuropteris oblongifolia
europteris smithsit
Neuropteris subfalcata
Odontopteris abbreviata
Odontopteris cornuta
Odontopteris monstruosa

Pecopteris georgiana
Pecopteris ornata
Pecopteris quadratifolia
Pecopteris ro
Pecopteris ee
Pecopteris vestz
Pseudopecopteris seine
Pseudopecopteris hispida
Pseudopecopteris speciosa
Pseudopecopteris subcrenulata
Rachiopteris squamosa
Rhabdocarpum mansfieldi
Rhabdocarpus abnormalis
Rhabdocarpus apiculatus
Rhabdocarpus cornutus
Rhabdocarpus inflatus
Rhabdocarpus insignis
Rhabdocarpus late-costatus
Rhabdocarpus mamillatus
Rhabdocarpus pachytesta
Rhabdocarpus sub-globosus
Rhabdocarpus tenax
Rhacophyllum corallinum
Rhacophyllum cornutum
Rhacophyllum expansum
Rhacophyllum fucoideum
Rhacophyllum hamulosum

Rhacophyllum membranaceum

Rhacophyllum spinosum
Rhacophyllum trichoideum
Rhacophyllum truncatum
Sigillaria grand’ euyri
Sigillaria lacoei

Sigillaria leptoderma
Sigillaria leveretti

Sigillaria pittstoniana
Sigillaria williamsi
Sorocladus ophioglossoides

BARTONIA

Sphenopteris brittsit
Sphenopteris communis
Sphenopteris —
Sphenopteris harvey
Sphenopteris inaequilaterals

Sphenopteris tracyana
Spirangium intermedium
Spirangium multiplicatum
Sporocystis planus
Stemmatopteris anceps
Stemmatopteris angustata
Stemmatopteris cyclostigma

Stemmatopteris squamosa
Taeniophyllum brevifolium

Taeniopteris smithsiu

Trigonocarpus adamsi
Trigonocarpus ampullaeformis
Trigonocarpus giffordi
Trigonocarpus grandis
Trigonocarpus kansaseanus
Trigonocarpus perpusillus
Trigonocarpus starkianus
Trigonocarpus subcylindricus
Trochophyllum clavatum
Volkmannia crassa
Volkamnnia fertilis
Whittleseya integrifolia
Whittleseya microphylla

Bartonia No. 61: 81-91, 2002

The Flora of Coastal Plain Pond Herbaceous Communities
on the Delmarva Peninsula

WILLIAM A. MCAVOY AND PETER BOWMAN
Delaware Natural Heritage Program, Delaware Dept. of Natural Resources,
Division of Fish & Wildlife, 4876 Hay Point Landing Rd, Smyrna DE 19977
wmcavoy@state.de.us pbowman@state.de.us

ABSTRACT. Coastal plain ponds on the Delmarva Peninsula are ecologically unique wetlands that
support a diverse assemblage of plant species and are a prominent part of the peninsula’s natural
heritage. As such, conservation of these wetlands is critical in helping to maintain biodiversity on
Delmarva. Very little has been published regarding the vegetation associated with Delmarva coastal
plain ponds, both in terms of the natural community types and the individual plant species that are
closely associated with them. This paper provides a general description of the natural community
types, or vegetation zones, found in Delmarva coastal plain ponds and focuses on the flora of the most
diverse vegetation zone, the herbaceous community. Particular attention is given to the rare and
uncommon flora of these wetlands. The summary of the flora of herbaceous communities in Delmarva
coastal plain ponds provides strong evidence regarding their importance for biodiversity conservation
on the peninsula.

INTRODUCTION

Akin to the well known Carolina Bays of the southeastern U.S., coastal plain ponds on
the Delmarva Peninsula (also known as Delmarva Bays or “whale wallows”) are a unique
type of shallow, seasonally flooded, freshwater depressional wetland. Generally elliptical in
shape and often with a pronounced sandy rim along the southeast side, these wetlands are
typically inundated in the winter and spring and draw down in late summer or early fall
Phillips and Shedlock 1993). They are most commonly found as open, sunny depressions
within areas of forested uplands. The majority of these ponds are small (most are less than
one acre in size), but they are numerous—it is estimated that there are between 1,500 and
2,500 of these ponds on the Delmarva Peninsula (Stolt and Rabenhorst 1987b; McAvoy and
Clancy 1994; Zankel and Olivero 1999).

Coastal plain ponds are recognized as a highly significant part of the Delmarva Peninsula S
Natural heritage (Sipple and Klockner 1984; Tyndall et al.. 1990; Sipple 1999). Their
"Ncommon ecological context and features provide irreplaceable habitat for an abundance
ot rare and uncommon species and unique natural communities. On Delmarva, coastal plain
Ponds are known to harbor 45 rare and uncommon plant species, including eight globally
‘are species and one federally listed species. Coastal plain ponds are also among the oe
‘hreatened natural systems on the peninsula, with hundreds of ponds having been degra %
or destroyed by anthropogenic activities (e.g-, agricultural ditching and clear-cut umber

harvesting methods [McAvoy and Clancy 1994). Despite the estimate of thousands a pee

lima oi
“script submitted 15 November 2000, revised 4 April 2001.

81

82 BARTONIA

occurring on Delmarva, it is often difficult to find ponds of good to high quality (i.e., with
intact and functioning water regimes, open canopy, high species diversity, and rare and
uncommon species).

Coastal plain ponds on Delmarva typically consist of concentric zones of herbaceous,
shrub, and forest natural community types. The most significant and floristically diverse
zone is the herbaceous community. It is within this community that a distinct assemblage
of plant species is found, and where rare and uncommon species are encountered. The focus
of this paper is on the flora of the herbaceous communities of coastal plain ponds on the
Delmarva Peninsula.

DISTRIBUTION OF COASTAL PLAIN PONDS

Coastal plain ponds can be found from Florida to Massachusetts. They are most
conspicuous in North and South Carolina (the “true” Carolina Bays), where they are
typically at their largest and most uniform in shape and orientation (Savage 1982), although
smaller, irregular ponds occur there as well (Sipple 1999). From Virginia northward, they
are typically smaller (usually less than 1 acre), and more variable in shape (Savage 1982).
Although most numerous in the Carolinas, they are still quite abundant on the Delmarva
Peninsula. Rangewide, it has been estimated that there are as many as 500,000 coastal plain
ponds between Florida and Massachusetts (Savage 1982).

On the Delmarva Peninsula, the greatest concentrations of coastal plain ponds can be
found in southwestern New Castle and northwestern Kent Counties, Delaware, the wester
portions of Queen Annes and Kent Counties, Maryland, and Caroline County, Maryland
(Stolt and Rabenhorst 1987b). Outside this region, coastal plain ponds on Delmarva are
scattered and infrequent. They are also locally abundant in portions of Sussex County,
Delaware and Dorchester County, Maryland.

The Delmarva Peninsula (Figure 1) is an area lying entirely within the Atlantic Coastal
Plain physiographic province of the eastern United States. The Peninsula lies south of the
Fall Line (a term applied to the boundary between the Appalachian Piedmont province an
the Atlantic Coastal Plain) of New Castle County, Delaware and Cecil County, Maryland,
and is bordered on the east by the Delaware River, Delaware Bay and the Atlantic Ocean,
and on the west by the Elk River and Chesapeake Bay. It includes the Coastal Plain
province of the state of Delaware (three counties), the Eastern Shore of Maryland (nine
counties), and the Eastern Shore of Virginia (two counties).

ORIGIN OF COASTAL PLAIN PONDS

Despite decades of research and debate, the geologic origins of coastal plain ponds are i
unresolved. The debate actually began in 1933 (Savage 1982), when Melton and Schriever
proposed that the presence of Carolina Bays were actually the result of meteorite strikes
They argued that the incredible uniformity in shape and orientation, along with the preseat”
of a raised sandy rim around each bay, indicated a single widespread phenomenon, with ; 4
impacts of meteorites being the most plausible explanation. Over the years, this theory
been highly controversial, with many strong arguments both for and against, and it st ; :
many adherents to this day (Savage [1982] provides an excellent overview of the develoP
ment of the theories on the origin of Carolina Bays).

DELMARVA PENINSULA COASTAL PLAIN PONDS 83

Whether or not the meteorite theory is true in
the Carolinas, it is unlikely to be so on the Del-
marva Peninsula (Stolt and Rabenhorst 1987b).
The pond features that strengthen the meteorite
theory—uniformity of shape and orientation,
raised sandy rims, overlapping rims—are much
less prevalent in coastal plain ponds on
Delmarva. Currently, the most plausible theory
is that Delmarva coastal plain ponds began
forming between 15,000 and 20,000 years ago as
depressions in coastal areas in a climate that was
much colder and somewhat drier (Stolt and
Rabenhorst 1987b). Stolt and Rabenhorst’s
theory suggests that strong winds created these
depressions in unvegetated sandy dune areas and
deposited the sand as the elevated rim found
around many of the wetlands today.

GEOMORPHOLOGY, HYDROLOGY AND SOILS
OF DELMARVA COASTAL PLAIN PONDS

Stolt and Rabenhorst (1987a) identified two
basic types of coastal plain ponds on the
Delmarva Peninsula: the basin-fill and the sandy
bottom. Of the two types, the basin-fill ponds
are generally deeper, with steeper slopes extend-
ing down to distinct low areas. These low areas

ene
NE are usually filled with loess deposits—silty loam
ae. material that was originally deposited by the
he a wind when the ponds were forming. These
basin-fill deposits are typically 1 to 3 m deep, but

Figure 1. may be as deep as 5 m. The second pond type,
ey o paces terial it sine tonal bottom, usually has some basin-fill
New Castle County Debiware ast ae sediments as well, but in much smaller amounts.
Part of the peninsula. The sandy bottom ponds typically have a rela-
tively flat bottom, with a thin layer of mucky
loam over deep sands. sate
_ The two pond types also appear to differ in distribution. The steep-sided, basin-fill type
's found in the northern part of the peninsula, especially in the southwestern region of New
Castle County, Delaware and western Kent County, Maryland (pers. obs. by the first
author; Frank Hirst and Ron Wilson, pers. comm.). As one moves south into Queen Annes
and Caroline Counties, the sandy, flat-bottom ponds start appearing, and are the prevalent
< type in southern Delmarva (pers. obs. by the first author; Frank Hirst and Ron
Son, pers. comm.). nt
Ina study of hydrology in coastal plain ponds in Delaware, Phillips and Shedlock (1993)
nis that Delmarva coastal plain ponds function as seasonally inundated wetlands.
Ypically, they are full in the winter, often to a depth of more than 1 m. In the summer,

84 BARTONIA

as temperature and evapotranspiration increase, the water level drops. The amount and the
timing of drawdown depend on annual weather patterns, with the substrate being
completely exposed in hotter, drier years. In wetter, cooler years, many ponds remain
inundated (though not full) throughout the summer.

Studies of water chemistry in Delmarva coastal plain ponds have typically found the
waters to be very soft and acidic (Stolt and Rabenhorst 1987a; Newman and Schalles 1990),
Rasmussen (1958) reported near neutral (6.2 to 7.1) pH values for a site in Sussex County,
Delaware; Phillips and Shedlock (1993) found the pH to be below 5.0 at sites in southwest
ern New Castle County, Delaware.

DELMARVA COASTAL PLAIN POND NATURAL COMMUNITIES

Historically, the vegetation dynamics of Delmarva coastal plain ponds have been poorly
documented, although a few studies have been conducted in recent years (Tyndall et al 1990;
Berdine and Gould 1999; Tyndall 2000). The natural vegetation communities of Delmarva
coastal plain ponds cover a wide range of variation, from closed-canopy forests to dense
shrublands to communities dominated by herbaceous plants (Berdine and Gould 1999).
Most commonly, these different community types occur as a complex of concentric
vegetation zones within an individual pond (Tyndall et al 1990; Berdine and Gould 1999;
Tyndall 2000). The center of the pond is usually open and dominated by herbaceous
species. Moving away from the center and toward the upland perimeter of the pond, there
is typically a transition to a shrub-dominated zone and finally to a forested community.
Each of these zones is defined by the Delaware and Maryland Natural Heritage Programs
as an individual community type (i.e., comprised of a characteristic suite of dominant plant
species occurring in a consistent, repeating pattern on the landscape). Although zones are
defined as individual community types, they occur as a complex, which exhibit repeating
patterns from pond to pond. However, it is possible to find Delmarva coastal plain pon
that have succeeded to purely shrub or forest communities, although they are less common
than the pattern described above (pers. obs. by the first author).

FOREST COMMUNITIES

Forested communities can occupy the entire depression, but more frequently are found
at the perimeters of pond openings. Whether they dominate the depression or are at me
periphery, these communities typically have a rather dense shrub layer of Clethra alnifolia
Vaccinium corymbosum, Rhododendron viscosum and Leucothoe racemosa, and are usually very
low in herbaceous species diversity. The canopy is typically composed of Acer rubrum,
Liquidambar styraciflua, Nyssa sylvatica, Pinus taeda (in southern Delmarva), Quercus p lust
and Quercus phellos (Berdine and Gould 1999; Bowman 2000).

SHRUB COMMUNITIES

There are several shrub communities that occupy Delmarva coastal plain ponds, but they
all feature the same shrub species: buttonbush (Cephalanthus occidentalis). This extreme!)
wide-r anging species can be found in a variety of wetland habitat types throughout ibe
peninsula, but is characteristic for Delmarva coastal plain ponds (pers. obs. by the firs
author). In many cases, buttonbush is the sole dominant in the pond, and essentially forms
a monoculture with few other species present (Bowman 2000).

DELMARVA PENINSULA COASTAL PLAIN PONDS 85

HERBACEOUS COMMUNITIES

Delmarva coastal plain pond herbaceous communities (DCPPHC) are unique wetland
community types on the peninsula. Classifying these communities is often a challenge, as
species composition and abundance can vary significantly from year to year in individual
ponds (pers. obs. by the first author; Ron Wilson, pers. comm.). These fluctuations in
species composition and abundance are almost certainly attributable to variation in
precipitation and temperature. However, no studies to date have looked at the correlation
between species composition and weather data.

Despite these challenges, numerous attempts have been made to describe DCPPHCs, with
various studies identifying five to ten different types (Tyndall et al. 1990; McAvoy and
Clancy 1994; Berdine and Gould 1999). While it is beyond the scope of this paper to
describe these various communities, it is worth noting that these studies have identified

iffe b iti pying basin-fill ponds and sandy-bottom ponds. There
seems to be a greater variety of sandy-bottom pond communities (Bowman 2000), and these
communities often have species with a southern affinity as dominants or co-dominants
(Table 1). These species are much less common in the basin-fill type ponds (pers. obs. by the
first author; Frank Hirst and Ron Wilson, pers. comm.).

THE VASCULAR FLORA OF
DELMARVA COASTAL PLAIN POND HERBACEOUS COMMUNITIES

The vascular flora of DCPPHCs (Table 1) is based primarily on observations made
through the years by the first author while visiting an abundance of coastal plain ponds
throughout the peninsula. A site visit usually consisted of traversing through the DCPPHC
while recording each species observed. Certain sites that were high in species diversity or
contained rare and uncommon taxa were visited annually to record observations on species
assemblages and the presence/absence of rare and uncommon taxa. Voucher specimens were
frequently collected and deposited in the herbarium of the Delaware Natural Heritage

rogram. In the future, these specimens will be transferred to the Claude E. Phillips
Herbarium, Delaware State University (DOV; herbarium acronyms follow Holmgren et al.
1990). The vascular flora of DCPPHCs is also based on consultations with knowledgeable
individuals (primarily Frank Hirst and Ron Wilson), natural community data from the
Delaware and Maryland Natural Heritage Programs (McAvoy and Clancy 1994; Berdine and
Gould 1999; Bowman 2000), herbarium specimens (primarily PH and DOV), and the
literature (Tatnall 1946; Sipple and Klockner 1984; Tyndall et al. 1990; Tyndall 2000).
he flora of DCPPHCs includes 78 native species and varieties of vascular plants,
“Presented by 22 families and 41 genera (Table 1). The largest families are Cyperaceae with
species and Poaceae with 15 species. The largest genera represented are Carex 9 species)
and Rhynchospora (8 species). Graminoids (grasses, sedges, and rushes) comprise the majority
of her baceous plants with 47 species, or 62% of the typical flora of DCPPHCs. Eighty-five
i of the typical flora of DCPPHCs are perennials (66 species) and 16% (12 species) are
Nuals,

jtveive species appear to be regionally restricted to DCPPH
£0 collected from any other habitat type on the peninsula
— of the Delaware and Maryland Natural Heritage Programs;
or; Frank Hirst and Ron Wilson, pers. comm.).
The DCPPHC flora has strong affinities to the southeastern U.S., as 43 species (55%) have

Cs (Table 1), and have not
(Tatnall 1946; rare species
pers. obs. by the first

86 BARTONIA

a more southern distribution, (i.e., are at or near the northern limits of their natural
geographic distribution [Table 1}. In contrast, only ten species of the 78 (13%) have a more
northern distribution (i.e., are at or near the southern limits of their natural geographic
distribution [Table 1]). Species range-wide distribution data are based on Gleason and
Cronquist (1991).

There appear to be differences in species composition between the sandy, flat-bottom type
and the steep-sided, basin-fill type of coastal plain pond on the Delmarva Peninsula. Forty
species occur primarily within the DCPPHC of the sandy, flat-bottom type and are only
rarely found in the steep-sided, basin-fill type (Table 1; pers. obs. by the first author; Frank
Hirst and Ron Wilson, pers. comm.). Of these, 33 species have southern affinities and only
3 species have northern affinities. In contrast, there are 16 species that occur primarily in
the steep-sided, basin-fill type and are only rarely found in the sandy, flat-bottom type
(Table 1; pers. obs. by the first author; Frank Hirst and Ron Wilson, pers. comm.). Of these
16, five have northern affinities and only three species have southern affinities. Twenty-two
species are common to both pond types.

RARE AND UNCOMMON PLANTS OF
DELMARVA COASTAL PLAIN POND HERBACEOUS COMMUNITIES

Delmarva coastal plain pond herbaceous communities contain a high number of rare and
uncommon plant taxa. The data presented below provide strong evidence that conservation
of this unique wetland type is critical in helping to maintain biodiversity on the Delmarva
Peninsula.

In addition, at least four species now thought to be extirpated on the Delmarva xi
may have occurred in coastal plain ponds historically: Juncus elliotii Chapman, -

boykinii Torrey & Gray ex A. DC., Polygala cymosa Walt., and Polygala ram “a me
e ar

. . . * . aes na
extensive ditching and land conversion over time has degraded any remaiming ® ail

“ditches,” which do not specifically describe coastal plain pond habitat. However, -

: ‘ Z aes terri bution
species are often found in coastal plain ponds in other regions within their distrib

DELMARVA PENINSULA COASTAL PLAIN PONDS 87
TABLE 1. Flora of Delmarva Coastal Plain Pond Herbaceous Communities.

The list is arranged alphabetically by family. With some exceptions, nomenclature follows Kartesz
(1994). Synonyms are listed in brackets where necessary for clarification. Taxa considered to be rare
and uncommon on the Delmarva Peninsula by the Delaware and Maryland Natural Heritage Programs
are highlighted in bold. The following indicators follow the scientific name: G = considered to be
globally rare by The Nature Conservancy, a private international conservation organization (globally
rare species are known from only 1 to 100 populations worldwide); 1 = known from only a single
occurrence or population on Delmarva; R = restricted to DCPPHCs on Delmarva; S-B = occurs
primarily in sandy, flat-bottom type ponds on Delmarva (when no indicator is given, the species is
common to both pond types); B-F = occurs primarily in steep-sided, basin-fill type ponds on
Delmarva; § = southern affinities; N = northern affinities.

ALISMATACEAE
Sagittaria graminea Michx. var. graminea (S-B)
TACEAE

Oxypolis canbyi (Coult. & Rose) Fern. (G, 1, R, S-B, S)
ASTERACEAE

Bidens frondosa L.

Boltonia asteroides (L.) L’Her. (S-B, S)

Coreopsis rosea Nutt. (G, R, S-B, N)

Sclerolepsis uniflora (Walt.) B.S.P. (S-B, S)
BLECHNACEAE

Woodwardia virginica (L.) Sm.
CAMPANULACEAE

Lobelia canbyi Gray (S-B, S)
LUSIACEAE

Hypericum adpressum Raf. ex W. Bart. (G, S-B, S)
Hypericum denticulatum Walt. (S-B, S)
Hypericum mutilum L.

CYPERACEAE

haris robbinsii Oakes (S-B)
: ocharis tricostata Torr. (S-B, S)
imbri istylis autumnalis (L.) Roemer & J.A. Schultes
imbristylis perpusilla Harper ex Small & Britt. (G, R, B-F, S)

88 BARTONIA
Table 1 (cont'd)

Rhynchospora chalarocephala Fern. & Gale (S-B, S)
Rhynchospora corniculata (Lam.) Gray (S)
Rhynchospora filifolia Gray (1, R, S-B, S)
Rhynchospora harperi Small (G, S-B, S
Rhynchospora inundata (Oakes) Fern. (S-B, N)
Rhynchospora macrostachya Torr. ex Gray var. macrostachya (S)
Rhynchospora nitens (Vahl) Gray (S-B, S) [Psilocarya nitens (Vahl) Wood]
Rhynchospora scirpoides (Torr.) Gray (S-B, N) [Psilocarya scirpoides Torr.]
Scirpus cyperinus (L.) Kunth
Scleria reticularis Michx. (S-B, S)
GENTIANACEAE
Sabatia difformis (L.) Druce (S-B, S)
HAEMODORACEAE
Lachnanthes caroliana (Lam.) Dandy (S-B, S)
HALORAGACEAE
Proserpinaca pectinata Lam.
JUNCACEAE
Juncus canadensis J. Gay ex Laharpe (N)
Juncus debilis Gray (S)
Juncus repens Michx. (S)
LAURACEAE
Litsea aestivalis (L.) Fernald (G, 1, R, S-B, S)
LE

Utricularia fibrosa Britt. (S-B, S)
Utricularia geminiscapa Benj. (B-F, N)
Utricularia gibba L.
Utricularia inflata Walt. (S-B, S)
Utricularia purpurea Walt. (S-B)
Utricularia radiata Small (S-B, S)
MELASTOMATACEAE
Rhexia aristosa Britt. (S-B, S)
Rhexia virginica L.
NYMPHAEACEAE
Nymphaea odorata Ait. ssp. odorata
ONAGRACEAE
Ludwigia linearis Walt. (S-B, S)
Ludwigia sphaerocarpa Ell.
POACEAE
Coelorachis rugosa (Nutt.) Nash (R, S-B, $) [Manisuris rugosa (Nutt.) Kuntle] -; Cwallea]
Dichanthelium hirstii (Swallen) Kartesz, comb. nov. (G, 1, R, $-B, $) [Panicum hirstil .
Dichanthelium spretum (J.A. Schultes) Freckmann (S-B) [Panicum spretum J.A. Schultes]
Dichanthelium wrightianum (Scribn.) Freckmann (S-B, $) [Panicum wrightianum Scriba.
Eragrostis hypnoides (Lam.) B.S.P. (R, B-F)
Glyceria acutiflora Torr. (R, B-F, N)
Glyceria septentrionalis A.S. Hitchc. (B-F)
Leersia hexandra Sw. (1, R, S-B, S)
Muhlenbergia torreyana (j.A. Schultes) A.S. Hitchc. (G, 1, R, S-B, S)
Panicum dichotomiflorum Michx. (B-
Panicum hemitomon J.A. Schultes (S-B, S)

DELMARVA PENINSULA COASTAL PLAIN PONDS 89

Table 1 (cont'd)

Panicum verrucosum Muhl.

Paspalum dissectum (L.) L. (S)

Saccharum giganteum (Walt.) Pers. (S-B, S) [Erianthus giganteus (Walt.) P. Beauv.]
POLYGONACEAE

Polygonum amphibium L. var. emersum Michx. (B-F) [Polygonum coccineum Muhl. ex Willd.]
PRIMULACEAE

Hottonia inflata Ell. (B-F, S)
RANUNCULACEAE

Panicum rigidulum Bosc ex Nees var. pubescens (Vasey) Lelong (S-B, S) [Panicum longifolium Torr.]
S)

Ranunculus flabellaris Raf. (R, B-F, N)
RUBIACEAE
Oldenlandia uniflora L. (S)
ACEAE

Xyris smalliana Nash (S-B, S)

Table 2. Additional rare and uncommon plant species documented from DCPPHCs that are
not typical for the community type on Delmarva.

Amphicarpum purshii Kunth

Asclepias lanceolata Walt.

Carex typhina Michx.

Centella erecta (L. f.) Fern.

Eriocaulon compressum Lam.

Euthamia tenuifolia (Pursh) Nutt. var. microcephala Greene
Fuirena squarrosa Michx.

Iris prismatica Pursh ex Ker-Gawl.

Juncus pelocarpus E. Mey.

Nymphoides cordata (Ell.) Fern.

Polygala cruciata L. var. aquilonia Fern. & Schub.
Rhynchospora cephalantha Gray var. cephalantha
Rhynchospora cephalantha Gray var. microcephala (Britt.) Kukenth.
Sagittaria engelmanniana J.G. Sm.

Trachelospermum difforme (Walt.) Gray

(Godfrey and Wooten 1981; Weakley 1999). Therefore, it is highly likely that they did occur
‘n coastal plain ponds on Delmarva prior to human disturbance. Another species, also
thought to be extirpated on Delmarva, Echinodorus parvulus Engelm., may also have been
collected from a coastal plain pond. The habitat description given on the specimen label is,
“border of pond, Canterbury” (W. Canby, 1874, Kent County, Delaware, PH). At present,
4small complex of degraded coastal plain ponds still exists in the Canterbury area, but field
“urveys by the first author have failed to rediscover E. parvulus at this site. Lacking clear
documentation that the these five species actually occurred in coastal plain ponds on
Yelmarva, and with no evidence that they still occur anywhere on Delmarva, they were not
included on the overall species list (Table 1).

An additional 15 species of rare and uncommon plants have been documented from
DCPPHCs (Table 2), but their occurrence in this community type is anomalous, for they

90 BARTONIA

are usually found in other wetland types on the peninsula (T atnall 1946; pers. obs. by the
first author; Frank Hirst and Ron Wilson, pers. comm.).

ACKNOWLEDGMENTS

We are extremely grateful to Frank Hirst and Ron Wilson, who have probably visited
more coastal plain ponds than anyone on Delmarva, for their review and valuable comments
on this manuscript. Without their input a paper such as this would not be complete. Many
thanks to Karen Bennett for her time spent reviewing this manuscript and for making
insightful and helpful comments, and for creating the locational map of the Delmarva
Peninsula.

LITERATURE CITED

BERDINE, M. A., AND A. M. A. GOULD. 1999. Identification and Protection of Reference Wetland
Natural Communities in Maryland: Delmarva Bay Wetlands. Maryland Department of Natural
Resources, Annapolis. 87 pp.

BOWMAN, P. 2000. The Natural Communities of Delaware: DRAFT. Unpublished report by the
Delaware Natural Heritage Program, Dover. 70 pp.

GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States
and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx, NY.

GODFREY, R. K., AND J. W. WOOTEN. 1981. Aquatic and Wetland Plants of the Southeastern United
States, Dicotyledons and Monocotyledons. Universtiy of Georgia Press, Athens, GA.

HOLMGREN, P. K., N. H. HOLMGREN AND L. C. BARNETT. 1990. Index Herbariorum Part I: he
Herbaria of the World, 8th ed. New York Botanical Garden, Bronx, NY.

KARTESZ, J. 1994. A Synonymized Checklist of the Vascular Flora of the United States, Canada and
Greenland, 2nd ed. Vol. I. Timber Press, Portland, OR. 622 pp. ;
McAvoy, W. A., AND K. E. CLANCY. 1994. Community classification and mapping criteria for
category I interdunal swales and coastal plain pond wetlands in Delaware. Unpublished report by

the Delaware Natural Heritage Program, Dover. ;

NEWMAN, M. C., AND J. F. SCHALLES. 1990. The water chemistry of Carolina bays: a regional
survey. Archiv fiir Hydrobiologie 118: 146-168.

PHILLIPS, P. J., AND R. J. SHEDLOCK. 1993. Hydrology and chemistry of groundwater and seasontl
ponds in the Atlantic Coastal Plain in Delaware, USA. Journal of Hydrology 141: 157-178.

RASMUSSEN, W. C. 1958. Geology and hydrology of the “bays” and basins of Delaware. Docto

issertation, Bryn Mawr College, Bryn Mawr, Pennsylvania. 206 pp. 1

SAVAGE, H. 1982. The Mysterious Carolina Bays. University of South Carolina Press, Columbia. 1

PP-

SIPPLE, W.S. 1999. Days Afield: Exploring Wetlands in the Chesapeake Bay Region. Gateway Press
Inc., Baltimore.

SIPPLE, W.S., AND W. A. KLOCKNER. 1984. Uncommon wetlands in the coastal plain of Mary at
Pp. 111-138 in Norden, A. W., D. C. Forester, and G. H. Fenwick (eds). Threatened
Endangered Plants and Animals of Maryland. Maryland Natural Heritage Program,
of Natural Resources, Annapolis. 4

STOLT, M. H., AND M. C. RABENHORST. 1987a. Carolina bays on the Eastern Shore of Mary ns
I. Soil characterization and classification. Soil Science Society of America Journal 51: 3943

STOLT, M. H., AND M. C. RABENHORST. 1987b. Carolina bays on the Eastern Shore of Mary
I. Distribution and origin. Soil Science Society of America Journal 51: 399-405. ad

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

DELMARVA PENINSULA COASTAL PLAIN PONDS 91

Delaware, Wilmington

TYNDALL, R. W. 2000. Vegearion change in a Carolina bay on the Delmarva Peninsula of Maryland
ae. an eleven-year period (1987-1997). Castanea 65: 155-164.

TYNDALL, R. W., K. A. MCCARTHY, J. C. LUDWIG, AND A. ROME. 1990. Vegetation of six Carolina
bays in Maced. Castanea 55: 1-21.

WEAKLEY, A. S. 1999. Flora of the Carolinas and Virginia, working draft of 19 January 1999. The
Nature Conservancy, Southeast Regional Office, Southern Conservation Science Department,
Chapel Hill, NC. 833 pp.

ZANKEL, M., AND A. OLIVERO. 1999. Mapping and assessing the conservation status of Delmarva bay
wetlands in Delaware. Unpublished report by the Eastern Conservation Office of The Nature
Conservancy, Boston. 28 pp.

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Bartonia No. 61: 93-116, 2002

Carex (Cyperaceae) in the Potomac River Gorge of
Maryland, Virginia, and the District of Columbia

CHRISTOPHER LEA
National Park Service, 7206 National Seashore Lane, Berlin, MD 21811
chris_lea@nps.gov

CHRISTOPHER T. FRYE
Maryland Department of Natural Resources, Wildlife and Heritage Service,
Tawes State Office Building, E-1, 580 Taylor Avenue, Annapolis, MD 21401

A field survey and herbarium search for taxa of the genus Carex (Cyperaceae) were conducted for
the Potomac River Gorge from 1995 through 2000. In the 120 years prior to this study (1875-1994),
7 taxa had been documented from the gorge. Eighty-two taxa were found during the current
fieldwork; 13 of these are new to this historically well-collected area. The total Carex flora for the
gorge now consists of 90 taxa representing 81 species. Carex corrugata and Carex laxiculmis var. copulata
are reported for the first time from Maryland, and 6 taxa are newly reported from the District of
Columbia. Additional new records for Montgomery County, Maryland and Fairfax and Arlington
Counties, Virginia are noted. Phytogeographical and ecological affinities of taxa groups are discussed.
An annotated checklist of taxa, with notes on local and regional distribution, habitat, and taxonomic
issues, is presented.

INTRODUCTION

_ The Fall Line gorge of the Potomac River is an area of geological, scenic, and biological
interest. This steep-sided, high-gradient reach begins at the intersection of the Coastal Plain
and Piedmont physiographic provinces in Washington, D.C. and Arlington County,
Virginia. It extends upstream some 24 km as the boundary between Montgomery County,

The Potomac Gorge is also ecologically and floristically diverse, as authors of the late
tineteenth and early twentieth century noted (Ward, 1881; Hitchcock and Standley, 1919).

logists’ Field Club with Plummers a
‘nsured that some level of botanical exploration continued throughout the middle part o
nee atighe hial oe RR IEE hat this was reduced after 1930.

raven of Natural Heritage Programs in Maryland (1979), Virginia (1986), - the cage
ot Columbi iti f the gorge as a significant site for
Olumbia (1994) brought new recognition © gorg ni Be Ri area

represented in

Although the botanical elements of this diverse area were relatively well 1919) pub-

‘arly floras of the Washington, D.C. area (Ward, 1881; Hitchcock and Standley,

“uscript submitted 13 November 2000; revised 3 April 2001.

93

mn BARTONIA

lished accounts or even formal listings of vascular plants of the gorge itself have been
surprisingly few. Terrell (1970) gave an account of spring-blooming flora of the Maryland:
District of Columbia side of the gorge, while Grimshaw and Bradley (1973) documented the
vascular flora of Great Falls Park on the Virginia side. In the 1980s and 1990s, Natural
Heritage Program surveys eld a number of rare plant occurrences, but generally
in internal or interagency re

The genus Carex el is by far the most species-rich vascular plant genus in most
areas of temperate eastern North America (e.g., Harvill, 1973; Reznicek and Ball, 1974;
Wheeler and Ownbey, 1984; Reznicek, 1989). Brown and Brown (1984) reported 145 species
(154 total taxa) for Maryland. For Virginia, Harvill et al. (1992) list 137 species, a total that
is conservative in comparison because of the relatively broad concepts of a number of species
employed by the authors. Despite the considerable representation of the genus in the floras
of many areas, the ecologies, distributions, abundances, and, in some cases, systematics of
the various taxa are often poorly known. The large number of taxa and overall complexity
of the genus, the poorly understood systematics of some groups, the somewhat inconspicu-
ous aspect of the plants themselves, and the relatively short seasonal period during which
reliable identifications can be made are all problems that mandate a level of taxonomic study
and effort often impractical for many general field surveys of vascular flora. Despite these
problems, carices are of floristic and conservation interest and are often well represented | in
lists of rare, threatened, and endangered species (Reznicek, 1989). Seventy-seven species of
Carex have been listed as being of some level of conservation concern in Maryland
(Maryland Natural Heritage Program, 1994), although ongoing research (Frye and Lea, 2001)
suggests that a number of these are not rare in the state.

MONTGOMERY

Island DISTRICT OF
Island COLUMBIA
se |
Fletchers
Floodplain
A é t Roosevelt Island
N FAIRFAX Chain Bridge’
COUNTY, VA — - Ry ?
, na nv
é Ss R
0 = ARLINGTON ¢
ARE. ret COUNTY, VA

Figure 1. Potomac River Gorge, showing place names appearing in text.

POTOMAC GORGE CAREX 95

Documentations of Carex in the Potomac Gorge prior to 1995 suggest that there were
knowledge gaps about the floristics of the genus that were similar to those described for
areas of New England by Reznicek (1989). Terrell (1970) found 15 species in the lower
Chesapeake and Ohio (C&O) Canal National Historical Park, 11 of which occurred in the
Potomac Gorge, while Grimshaw and Bradley (1973) compiled 10 species for Great Falls
Park, on the Virginia side of the gorge. In contrast, Hitchcock and Standley (1919)
previously had listed 95 species of Carex for the vicinity of Washington, D.C., an area that
included the valley of the Potomac below Great Falls. It was evident that organized floristic
and ecological information on Carex in the gorge was substantially incomplete.

Description of the Potomac River Gorge

The area of this study encompasses lands along the 24-km reach of the Potomac River
from Theodore Roosevelt Island, which lies at the head of the estuarine section of the
Potomac on the Fall Line, upstream to Great Falls (Figure 1). The study area extends
generally 1 km from the river on both sides to the rim of the gorge, approximately to an
elevation of 100 m above sea level, and to equivalent elevations in tributary stream valleys.
Most fieldwork was conducted on lands managed by the National Park Service (C&O Canal
National Historical Park on the Maryland-District of Columbia side and Great Falls Park,
Turkey Run Park, the George Washington Memorial Parkway, and Theodore Roosevelt
Island on the Virginia side). Small amounts of fieldwork occurred in county parks and on
other private lands.

The gorge is a Fall Line feature formed by accelerated rates of erosion of the river’s bed
as it responded to fluctuations in sea level during the Pleistocene. The resistant nature of the
underlying crystalline metamorphic rock in this section of the Piedmont has produced
bedrock-controlled features such as falls, rapids, exposed rock outcrops, and steep
topography. Modern (Holocene) floodplai d other depositional features are ”"
than on the lower gradient reaches of the Potomac upstream and downstream from the
gorge. Terraces (former floodplains that are rarely to never flooded) of various elevations
and of Pleistocene age or older are prominent. More complete descriptions of the geology
and geomorphology of the gorge are given by Reed at al. (1970) and Tormey (1980).

Recent research (Lea, 2000) has classified and described the diverse, often unusual vascular
piant communities of the gorge and described the strong influence of fluvial processes on
Soils and vegetation. Occasionally, the influence of the underlying bedrock is expressed in
a site’s flora, such as in the presence of bosophiles on soils over amphibolite, a mafic rock.
Although much of the riparian zone in the gorge supports forests similar to those of other
large river floodplains in the mid-Atlantic region, higher gradient sections have unusual,
“our-influenced plant communities that include riverside prairies, open, grassy woodlands,
and xeric, rocky forests. These are concentrated in the 4-km reach immediately downstream
from Great Falls (the Mather Gorge) and the 4 km centered on Chain Bridge (Chain Bridge
Flats). The C&O Canal, restored as a shallow waterway parallel to the Potomac River
throughout the Maryland-District of Columbia side of the study area, provides wet, open
habitat similar to pond edges, while its associated towpath, with its mowed edges, provides
* weedy, Open corridor, as may occur along roadsides.

% BARTONIA
METHODS

The objectives of the study were to record as many Carex taxa as possible in the Potomac
Gorge, to describe the distribution, habitat, and relative abundance of each, and to provide
information for more extensive research on Carex in Maryland and the District of Columbia
(Frye and Lea, 2001). To locate and examine vouchers for the gorge and to plan fieldwork,
the authors searched the collections of GMUF, MARY, NA, PH, and US (acronyms follow
Holmgren et al., 1990) and (with acronyms as assigned here and in lower case letters) those
of the Maryland Department of Natural Resources in Annapolis, Maryland (Tawes), the
Cylburn Arboretum in Baltimore, Maryland (Cylb), the National Park Service at Great Falls
Park, Great Falls, Virginia (Grfa), and the U.S. Geological Survey Water Resource Division
in Reston, Virginia (Usgs). The first author observed and collected some Carex between 1994
and 1999, pursuant to other research (Lea, 2000). Field surveys, primarily by the first author,
were conducted on foot, throughout the study area, with emphasis of the Maryland-District
of Columbia side of the gorge. A canoe was occasionally employed to reach river islands.
Fieldwork was conducted between April 8 and July 20 each year and totaled 12 days in
1999, 14 days in 2000, and a somewhat fewer number of days each year from 1995 to 1998.

Individual Carex plants were identified in the field, and notes were made of the location,
habitat, and relative abundance of each taxon. Specimens were collected as needed to
facilitate identification or as vouchers if a taxon were not previously recorded from the
study area or the applicable political jurisdiction. A 30-power dissecting microscope with an
ocular micrometer often assisted laboratory examination, and voucher specimens from
collections were used for comparison. Both authors examined all voucher specimens
collected during fieldwork, and specialists, particularly A. A. Reznicek (MICH), provided
assistance with some determinations.

RESULTS AND DISCUSSION
Summary of Floristic Findings, Including Significant Taxa

Ninety taxa representing 81 species were confirmed from both field and collections
research. Of these, 13 taxa are newly reported for the Potomac Gorge (Table 1). Nine tax?
that had been historically collected there (Carex annectens var. xanthocarpa, C. davis, a
decomposita, C. glaucodea, C. hystericina, Carex mublenbergii var. mublenbergit, C. pallescens,
C. pellita, and C. seorsa) were not found by the current fieldwork. New records were
determined for Montgomery County and for the District of Columbia based on existing
herbarium vouchers. New records for Fairfax and Arlington Counties were based largely on
Harvill et al. (1992). New jurisdictional records are summarized in Table 1. Carex corruga’4
and Carex laxiculmis var. copulata are newly reported for Maryland. ed

Nine species previously documented from the gorge were considered historical (presum
extirpated) from Maryland at the beginning of this study (Maryland Natural ne
Program, 1994). Of these, five (C. aggregata, C. careyana, C. conjuncta, C. leavenworthit, a
C. striatula) were relocated in the gorge by this study (and in other Maryland location®
during the same period), and two (C. davisii and C. decomposita) were not (C. davis ™*
been rediscovered elsewhere in Maryland). We found that previous reports of the remaining
two historical taxa (C. tenera and C. tetanica) had been based on misidentified pee?
although C. tenera was subsequently found in the gorge, for the first time, during this ste)
Additionally, a tenth species considered historical for the state, C. woodii, was also gine

POTOMAC GORGE CAREX

97

Table 1. Additions to Carex flora of three counties, District of Columbia, and Potomac

Gorge, this study.

Jurisdiction (and no. of
new taxonomic records)

New Taxonomic Records

District of Columbia 6

Montgomery County 8

Fairfax County 3
Arlington County 3
Potomac Gorge 13

Carex caroliniana

Carex cristatella

Carex granularis var. haleana
Carex brevior

Carex corruguata™

Carex digitalis var. macropoda
Carex emoryi

Carex conjuncta
Carex emoryt

Carex emoryt
Carex grayi

Carex albicans var. emmonsit

Carex debilis var. macroc.

arpa
Carex laxiculmis var. copulata*

Carex lupuliformis

Carex grayt
Carex stipata var. maxima
Carex typhina

laxiculmis var. copulata*
Carex lupuliformis

Carex tonsa var. rugosperma
Carex stipata var. maxima

Carex oligocarpa

Carex nigromarginata
Carex stipata var. maxima
Carex stricta

Carex tonsa var. rugosperma

Carex typhina
Carex woodii

“Addition to Maryland flora

Maryland).

Phytogeographical Aspects

for the first time in the gorge during this study (and has been relocated elsewhere in

Based on other research on Carex in Maryland and the District of Columbia (Frye and
Lea, 2001), we note several phytogeographical patterns of the Potomac Gorge Carex flora.

Definitions of physiographic provinces follow Schmidt (1993).
All taxa found by this study are native to eastern North
Maryland. Patterns of collections suggest that Carex aggregata may

its range in, our area due to human activities.
are widely distributed in Maryland. These include both

Of the 90 taxa found, 37 (41%)
taxa found in most or all major regions of the st
tribuloides, C. vulpinoidea) and those widespread
largely absent from the Appalachian Mountains (e.g.,

Forty-two taxa (47%) show a preference for,
Appalachian Mountains. These include taxa that also occ

Chesapeake Bay (e.g., Carex grayi, C. prasina, C. squarrosa,

widespr ead west of the Fall Line, but mo
© gracillima, C. normalis), taxa known exclusively from the

America and, probably, to

have entered, or expande

ate (e.g., Carex cephalophora, C. lurida, C.
in the Piedmont and Coastal Plain, but
Carex umbellata).

or a restriction to, the Piedmont or
ur on the Coastal Plain west of the
C. willdenowit), taxa that are
re strictly limited to that area (e.g., Carex conjuncta,
Piedmont or nearly so (¢.g,,

98 BARTONIA

Carex bushii, C. bystericina, C. planispicata), taxa that are more characteristically Appalachian
(e.g., Carex leptalea ssp. leptalea, C. tonsa var. rugosperma, C. torta, C. woodit), and several
primarily restricted to the valleys of the Potomac and Susquehanna Rivers (e.g., C. cristatella,
C. davisii, C. emoryi, C. jamesii, C. shortiana).

Eleven taxa (12%) are characteristic of the Coastal Plain, with most of these reaching their
western limit along the Fall Line areas of Piedmont rivers. Most seem to be more abruptly
restricted by the Fall Line than do many of their Piedmont counterparts. These taxa include
Carex albolutescens, C. complanata, C. crinita var. brevicrinis, C. seorsa, and C. styloflexa.

Habitat Descriptions

We describe seven typical habitats in the gorge and their Carex flora. Nomenclature of
taxa in genera other than Carex follows Gleason and Cronquist (1991). More detailed
descriptions of vascular plant communities in for the gorge are given by Lea (2000).

Dry-mesic upland forests: Never-flooded upland forests, generally dominated by oaks
(including Quercus rubra, Q. alba, and Q. velutina) and other tree species of well-drained,
mildly acidic soils (Fagus grandifolia, Acer rubrum, Nyssa sylvatica, Liriodendron tulipifera,
Cornus florida). Typical Carex of this habitat include C. albicans var. albicans, C
cephalophora, C. digitalis var. digitalis, C. laxiculmis var. laxiculmis, C. rosea, and C. virescens.

Xeric forests and woodlands: Never-flooded upland forests on shallow, apparently acidic
soils and dominated by Quercus prinus. Also, flood-scoured, rocky woodlands, dominated
by Pinus virginiana and oaks (Quercus rubra, Q. prinus, and Q. stellata), and scoured riverside
“prairies,” dominated by bunchgrasses such as Andropogon gerardii, Schizachyrium scoparium,
Panicum virgatum and by shrubby dry-land tree species. Both of the latter situations occur
on bedrock (strath) terraces and are maintained by infrequent floods with high current
velocities. For all xeric forests and woodlands, the Carex flora is relatively depauperate, but
distinctive, and includes C. albicans var. emmonsit, C. nigromarginata, C. tonsa var. tonsa, an
C. umbellata

Bedrock terrace forests: Dry, rich forests over shallow soils of bedrock terraces influenced
by rare, scouring floods. These occur primarily in the vicinity of Bear Island. Dominant tree
species include several oaks (Quercus rubra, Q. prinus, and Q. alba), Carya glabra, Fraxinus
americana, Juniperus virginiana, and Ostrya virginiana. The Carex flora is diverse and
includes C. albicans var. albicans, C. cephalophora, C. hirsutella, C. laxiculmis vat. Laxiculmis,
C. nigromarginata, C. pensylvanica, C. planispicata, C. retroflexa, C. swanit, C. willdenowt,
and C. woodii.

Floodplain and alluvial terrace forests: Regularly flooded alluvial forests 07 true
floodplains in lower gradient reaches of the gorge, dominated by flood-tolerant tree Spee
including Platanus occidentalis, Acer saccharinum, Acer negundo, Fraxinus pennsylvanica, 20
Ulmus americana. Also, mesophytic forests with Fraxinus americana, Liriodendron eulipifers
Acer saccharum, Celtis occidentalis, Tilia americana, Juglans nigra, Carya cordiforms, is
Ulmus spp. on more elevated, less frequently flooded terraces with sandy, well-drained er
The Carex flora includes C. amphibola, C. blanda, C. grayi, C. grisea, C. jamesi, and C
a with C. careyana, C. oligocarpa, and C. sparganioides present only in the mesophyt¢
orests.

Scoured woodlands: Rocky riparian woodlands scoured by frequent floods and gate
on the active channel shelf, a gently sloping riparian feature that is developed et ail
below that of floodplain (Osterkamp and Hupp, 1984) and is unusually extensive at ¢
Chain Bridge Flats. These woodlands are dominated by stunted flood-tolerant trees

POTOMAC GORGE CAREX |

bicolor, Fraxinus pennsylvanica, Acer negundo, Platanus occidentalis, Diospyros virginiana,
Ulmus americana, Juglans nigra, and Betula nigra) and meadow-forming grasses (Chasman-
thium latifolium, Elymus spp., Panicum clandestinum). Members of Carex are well represented
in this habitat, being generally concentrated in moist, scoured depressions, with 23 species
counted in one 10-m? swale. Typical taxa include C. amphibola, C. blanda, C. conjuncta, C.
crinita, C. frankii, C. granularis, C. grisea, C. leavenworthii, C. lupulina, C. lurida, C.
normalis, C. radiata, C. scoparia, C. shortiana, C. squarrosa, C. stipata var. stipata, C.
tribuloides, and C. vulpinoidea.

Forested vernal pools: Poorly drained depressions, primarily in the upper section of the
gorge on bedrock terraces, holding water in winter and early spring, drying by summer, and
occupied by hydric vegetation, including Acer rubrum and Nyssa sylvatica. The Carex flora
is fairly distinctive, with a number of taxa of primarily Coastal Plain affinity, including C.
albolutescens, C. annectens var. annectens, C. caroliniana, C. debilis var. debilis, C. intumescens,
and C. typhina.

Seepages and spring runs: Moist to mucky soils along drainages in upland forests. The
Carex flora includes C. laevivaginata, C. prasina, C. radiata, and C. styloflexa.

With over one-half of Maryland’s and Virginia’s total Carex flora represented, the
Potomac Gorge is clearly a significant and diverse site for this group, as it has been so
recognized for vascular plants in general. Within this restricted area, the floristic and habitat
diversity allows the opportunity to gain insight into the ecological and taxonomic patterns
of a large portion of each state’s complement of this large and relatively understudied genus.

ANNOTATED LIST OF TAXA

Our treatment of taxa generally follows Rhoads and Block (2000). Treatment of taxa not
listed for Pennsylvania by those authors follows Fernald (1950), with the following
exceptions: the Carex albicans complex (C. artitecta Mackenzie, C. emmonsu Dewey in
Torrey, and C. physorhyncha Liebman of Fernald, 1950) follows Rettig (1990), and C.
tribuloides var. sangemonensis follows Mackenzie (1931).

Our assessment of the relative abundance of a taxon is:

Common: easily found in appropriate habitat and usually will be noted by general botanical
Surveys.

Frequent: more than 10 occurrences seen; if fewer than 10, occurrences have large numbers
of plants.

Infrequent: 4 to 10 occurrences seen; habitat often specialized or limited.

Rare: 1 to 3 occurrences seen, usually with small numbers of individuals.

Historical: At least one collection from earlier than 1995 seen, but not found in the current

fieldwork.

_ Ahabitat description (as described above or in a more generalized manner) for each taxon
‘Ss provided. A discussion of distribution (if restricted within the gorge), regional conserva-
ton Status, and taxonomic concerns follows, as applicable. The local seasonal period for
finding phenologies optimal for identification (1.e., when infructescences of most plants =
Mature, but not senescing) is listed; in some cases, these periods may be Ne gee Y
narrow because of the small number of specimens available for evaluation. A list : ‘tonsoerd
Specimen collection numbers and repositories, comprehensive for the fieldwork of this st y
and also including some representative pre-1995 collections (for infrequent, rare, an

100 BARTONIA

historical taxa) appears at the end of each taxon entry. All collection numbers without an
associated collector name are those of the first author and represent material from the
current fieldwork. Repositories are designated as in the Methods section and also include the
Frostburg State University herbarium, Frostburg, Maryland (Frost), the Anne Arundel
Community College herbarium, Arnold, Maryland (Aacc), and the Delaware Natural
Heritage Program herbarium in Smyrna, Delaware (Dnhp). Vouchers at US will be found
in the Washington, D.C. and vicinity collection (including the Plummers Island collection).

Carex aggregata MacKenzie

Frequent. In open areas in dry woods and less commonly on floodplains. Also in
disturbed soil, such as along C&O Canal towpath. It has been listed as historical for
Maryland (Maryland Natural Heritage Program, 1994). It has been found recently to bea
relatively common species in the state and has been often misidentified or overlooked. Mid-
May to mid-June. 484, 1479 (US); 1488 (MARY); 1561 (GMUE, Grfa); 870, 1020, sn.
(Tawes); s.n. (Aacc).

Carex albicans Willdenow ex Sprengel var. albicans

Common. In dry to dry-mesic upland forests and rich bedrock terrace forests. One of our
most abundant taxa. Also treated as C. artitecta Mackenzie, it occurs in more rich or mesic
(but still dry) forests than var. emmonsii. Evidently included under C. emmonsii Dewey by
Hitchcock and Standley (1919). Mid-April to mid-May. 429, 1340 (US); 1448 (MARY); 779,
790 (Tawes); 952, 1345 (Frost); 773 (Aacc).

Carex albicans Willdenow ex Sprengel var. emmonsii (Dewey) Rettig

Infrequent. In dry, acidic forests, usually under Quercus prinus. Vaso and Hermit Islands,
Donaldson Run ravine, Great Falls Park (Virginia). Often treated as Carex emmonsi Dewey
in Torrey. This is the first report of the taxon for the Potomac Gorge. Mid-April to mid-
May. 436 (US); 442 (MARY); 950 (Tawes); 1333 (Grfa).

Carex albolutescens Schweinitz

Frequent. Edges of forested vernal pools, where it is one of the characteristic tax
Hitchcock and Standley (1919) have misapplied the name C. straminea to this species; while
C. albolutescens of Hitchcock and Standley (1919) evidently represents C. longii Mackenzie,
a Coastal Plain species. Mid-May to late June. 1261, 1485, 1924 (US); 1925 (MARY); iif,
1246 (Tawes); 1709 (Grfa).

Carex albursina Sheldon

Rare. In mesophytic alluvial terrace forests. Found in small numbers at Bear, Vaso; and
High Islands; more numerous at Turkey Run Park. C. albursina is a calciphilic spectes ”
is rare in the Piedmont, where it is apparently restricted to the Potomac and Susqué “—
valleys. Early May to late May. 1455 (US); 452 (MARY); 880 (Tawes).

Carex amphibola Steudel

Common. In floodplain and rich dry-mesic upland forests. This taxon was ——
Carex amphibola Steudel var. rigida (Bailey) Fernald by Fernald (1950), but 1s
considered the type of C. amphibola (Naczi, 1999). Early May to mid-June. 1028, 1048, 11!
1263, 1482 (US); 1204 (MARY); 1178 (Tawes); 1087, 1491 (Frost); 1012 (Aacc)-

POTOMAC GORGE CAREX 101

Carex annectens (Bicknell) Bicknell var. annectens

Frequent. Primarily along edges of forested vernal pools, where it is a characteristic taxon.
Also in other open or disturbed wet areas, in apparently more acidic situations than C.
vulpinoidea. Mid-May to mid-June. 453, 1695, 1704 (US); 1694, 1931 (MARY); 883, 1251
(Tawes); 2090 (Grfa).

Carex annectens (Bicknell) Bicknell var. xanthocarpa (Bicknell) Wiegand

Historical. Single collection in 1932 (det. by F. J. Hermann) from Plummers Island, with
habitat given as “barrens” (most likely, flood-scoured bedrock). Sometimes treated as Carex
brachyglossa Mackenzie, it is said to occur in more basic soils than the typical variety
(Weakley, 1999). Late May. Killip 32611 (US).

Carex atlantica Bailey. ssp. atlantica

Rare. Along spring run. A single station found in the Gold Mine Tract, found. There are
two historical collections (from the same date) from 1899, both possibly from the present
station, and a third from 1915 from the edge of a pool on the mainland opposite Plummers
Island. Early May to early June. 1599, Morris s.n., Pollard s.n., Standley 11663 (US); 1597
(MARY).

Carex blanda Dewey

Common. In a variety of forests and open, often disturbed, habitats, but most common
on floodplains. One of our most frequently encountered Carex species. Late April to early
June. 894, 1037, 1947 (US); 757 (MARY); 460, 466, 857, 998 (Tawes); 875 (Frost); 886 (Aacc);
1469 (Grfa).

Carex brevior (Dewey) Mackenzie ex Lunnell —— :
Rare. Seen at a single location in bedrock terrace forest, Bear Island. This is the first

a, of C. brevior for the Potomac Gorge and for Montgomery County. Mid-May. 1520
awes).

Carex bushii Mackenzie

Rare. Found at a single station on Olmsted Island (Maryland side of Great Falls), in dry
riverside “prairie.” Also known historically from the “District line, Dalecarlia (vicinity of
Little Falls Branch) in 1897 and from “open flat ground, Ft. Ethan Allen” (near rim of gorge
near Fairfax-Arlington County line) in 1929. Late May to early June. Steele s.n. (US); 1932
(MARY); Hasselbring s.n. (NA).

Carex Careyana Torrey ex Dewe age

Infrequent. In rich mesophytic forests on alluvial terraces. Considered historical for
Maryland (Maryland Natural Heritage Program, 1994), it was rediscovered in the state
during this survey. About 12 stations in the gorge, mostly upstream from the Capital
Beltway, are now known, though it is not numerous at any site. This distinctive sedge can
be identified in winter by its wide, purple-based leaves. Late April to mid-May. G. Fleming
8885 (GMUF); 797 (Tawes); 1330 (Grfa); Sigafoos 612 (Usgs).

Carex caroliniana Schweinitz eee
_ Infrequent. Edges of patie vernal pools, Bear Island and Great Falls Park (V irginia) and
in floodplain forest dominated by Quercus phellos at Fletchers Floodplain. This is the first

102 BARTONIA

District of Columbia record for C. caroliniana. Mid-May to early June. 1588, 1692 (US);
1581, 1691 (MARY); 1693 (Tawes); 1711 (Grfa); Baltars 3909 (Cylb).

Carex cephalophora Muhlenberg ex Willdenow

Common in dry-mesic upland forests and in rich bedrock terrace forests, less so in
floodplain forests. Mid-May to late June. 1066, 1162, 1232, 1483, 1593, 1700 (US); 1161, 1698,
1919 (MARY); 1031, 1202 (Tawes); 1160, 1218, 1699, 1701, 1919 (Frost); 1030, 1252 (Aac«)
1715 (Grfa).

Carex communis Bailey var. communis

Infrequent. In dry-mesic forest, usually on steep slopes with bare soil or exposed rock.
Turkey Run Park, High Island, and in ravines of Donaldson Run, Bullneck Run, and Cabin
John Creek. Late April to mid-May. 968, 1445, 1454 (US); 992 (MICH, Tawes).

Carex comosa F. Boott

Infrequent. In wet open areas, including edges of C&O Canal and in an open fresh tidal
swamp. Seen along canal at Bear Island and Cropley and at Roosevelt Island. Early June to
early July. 1224, 1254 (US); 1223 (MARY); 1093 (MICH); 1091 (Tawes); 536 (Frost).

Carex complanata Torrey & Hooker

Infrequent. In bedrock terrace forests and along edges of vernal pools, Bear Island,
Olmsted Island (Maryland side of Great Falls), and Great Falls Park (Virginia) and in
floodplain forest dominated by Quercus phellos at Fletchers Floodplain. Difficult to
distinguish from similar taxa (C. bushii, C. caroliniana, C. hirsutella), and specimens from this
group are frequently misidentified in collections. Mid-May to early June. 869, 872 (US); 1585
(MARY); 1710 (Grfa).

Carex conjuncta F. Boott

Frequent. In moist swales in channel shelf woodlands, less frequently in shaded floodplain
forests. Formerly regarded as historical in Maryland (Maryland Natural Heritage Progra™
1994). Now known to be fairly widely distributed in the Piedmont and Ridge and Valley
provinces in the state. Chain Bridge Flats may be an important population center for this
species, which is evidently uncommon in the region. This is the first report of C. conjuncis
for Fairfax County. Mid-May to late June. 1098 (US); 1089 (MICH); 1702 (GMUF); 7
1094, 1159 (Tawes); 533 (Frost); 2091 (Grfa).

Carex crinita Lamarck var. brevicrinis Fernald of
Frequent. In moist swales in floodplain forests, in forested swamps, and along en" is
ponds and the C&O Canal. Mostly restricted to the Coastal Plain in Maryland, nore

POTOMAC GORGE CAREX 103

the more frequently encountered variety of C. crinita. Mid-May to mid-June. 1574 (US);
1573 (MARY); 550, 1198 (Tawes); 2083 (Grfa).

Carex crinita Lamarck var. crinita

Common. In similar habitat as var. brevicrinis. The dominant variety of C. crinita in the
Piedmont and the Appalachian Mountains of Maryland. Mid-May to mid-June. 1477, 1572
(Us); 1081, 1478 (MARY); 1568 (GMUBF); 1567 (Grfa).

Carex cristatella Britton

Rare. In moist swales in open channel-shelf woodlands at Chain Bridge Flats, in open
fresh tidal swamp at Roosevelt Island, and in marsh at edge of canal near Cropley.
Historically found along Difficult Run and at Plummers Island. This is the first report of
this species for the District of Columbia. Late May to early July. 1072, 2094 (US); 1197,
1212 (MARY); 1214 (MICH); 1220 (Tawes); 2095 (Frost).

Carex davisii Schweinitz & Torre

Historical. Known from a single collection in 1905 from “among rocks at Great Falls of
the Potomac, Maryland.” This species was known from only one other collection in
Maryland and was considered historical for the state (Maryland Natural Heritage Program,
1994) until 2000, when it was found in two locations, one along the Potomac about 40 km
upstream from the Potomac Gorge. It has not been recorded from Virginia. It is rare in the
mid-Atlantic states, where it occurs in rich alluvial woods and meadows. Late May to early

June. Holm s.n. (US).

Carex debilis Michaux var. debilis.

Infrequent. Edges of forested vernal pools and along spring runs. Bear and Vaso Islands,
Great Falls Park (Virginia), and near Plummers Island. This is the first record for C. debilis
in the Potomac Gorge. 451, 1466, 1484 (US); 1580 (MARY); 888 (Tawes); 1466 (Gria).

Carex decomposita Muhlenberg lands

Historical, Apparently occurred along edges of forested vernal pools or other wetlands.
There are several collections at US and NA (dating from 1878 to 1920), from the Widewater
area (section of C&O Canal along Bear Island). These are the only Maryland records for C.

omposita. Specific search efforts were made for this species in the locations and general
habitat described in historical collections. Since this large, distinctive sedge has also not been
seen in recent rare plant surveys in the area, it may well be locally extirpated. C. spe ae
is considered historical for Maryland (Frye and Lea, 2001) and rare in Virginia (Ki . er,
Pres May to early July. Holm s.n., Killip 7108, Maxon 6334, Pollard s.n. (US); Chase

A).

Carex digitalis Willdenow var. digitalis nie

©mmon in dry-mesic to mesic upland forests, less so in rich

8 €ristic upland forest taxon in this area. Early May to mid-June.
67, 1165 (Tawes).

bedrock terrace forests. A

c 1032, 1118, 1441 (US);

Carex digitalis Willdenow var. macropoda Fernald
-In rich, dry forests. Found on Bear and Vaso Is
The dominant form of C. digitalis on the Coast

lands, and in ravine of Cabin John

€
Creek, al Plain to the south of Maryland,

104 BARTONIA

it reaches its apparent northern limit in Maryland, where, contrary to Brown and Brown
(1984), it is much less common than var. digitalis. Distinguished from var. digitalis primarily
by its long-peduncled spikes; we have not noted the coralline-knotty rhizomes described by
Fernald (1950). In Maryland and in the gorge, apparently prefers more basic soils than
typical C. digitalis. This is the first report of the taxon from Montgomery County and from
the Potomac Gorge. Early May to mid-June. 439, 1065 (Tawes).

Carex emoryi Dewey

Frequent. On rocky, scoured river bars and frequently flooded channel-shelf woodlands,
where it can form extensive stoloniferous colonies, as at Chain Bridge Flats and High Island.
Also in an open fresh tidal swamp at Roosevelt Island. Often, relatively few plants in a
colony will flower. We report here the first records for C. emoryi for Montgomery, Fairfax,
and Arlington Counties. Mid-May to late May. 474, 1557 (US); 1586 (MARY); 1053, 1168
(MICH); 1109, 1558 (GMUF); 1059, 1083, 1109, 1167 (Tawes); 1136, 1686 (Frost); 1052
(Aacc).

Carex festucacea Schkuhr ex Willdenow

Rare. In floodplain forest, Fletchers Floodplain and in rich bedrock terrace forests near
Cropley; historically recorded from Great Falls, Maryland. There had been only one
previous collection of C. festucacea from the gorge (in 1907, originally misidentified as C
tenera). Mid-May to mid-June. 1596, 1696, Shreve 1464 (US); 1920 (MARY).

Carex frankii Kunth

Common. In moist, open areas, including scoured channel shelf woodlands, open
floodplain forests, wet meadows, bedrock pools along the river, ditches, and other disturbed
wetlands. Early June to late July. 482, 1101, 1943 (US); 967, 1199 (Tawes).

Carex glaucodea Tuckerman ss

Historical. Known from a single collection, in 1893 near the Aqueduct Bridge, Virginl#
(near present-day Key Bridge in Arlington County) in “shaded places in woods.” T
species, sometimes treated as Carex flaccosperma Dewey var. glaucodea (Tuckerm )
Kiikenthal, is found in dry calcareous or mafic upland soils in Maryland. In the gorge,
may occur over amphibolite. Late May to early June. Holm s.n. (US).

Carex gracilescens Steudel

Rare. In early successional upland forest dominated by Liriodendron tulipifera. Seen only
in Difficult Run stream valley. Historically collected from Bear Island and Little Falls
Branch. This species is locally common along smaller stream valleys in the Marylan
Piedmont and may be more frequent along tributary streams than along the Potome
Hitchcock and Standley (1919) treat it as C. laxiflora Lam. (misapplied to this species). 1?
April to early June. 1384 (US, Grfa); Holm s.n., Maxon & Standley 325, Ward s.n. (US):

Carex gracillima Schweinitz "
Infrequent. In rich, dry-mesic forests along tributary streams to the Potomac and !
mesophytic alluvial terrace forests along the Potomac’s main stem. Found at Bear Isl
Great Falls Park (Virginia), and in the ravines of Cabin John Creek, Bullneck Rus, #
Pimmit Run. Early May to early June. 1374 (MARY); 956 (GMUF); 1451 (Frost):

POTOMAC GORGE CAREX 105

Carex granularis Muhlenberg ex Willdenow var. granularis

Frequent. In moist, scoured woodlands and open floodplain forests. Primarily at Chain
Bridge Flats. Early May to late-June. 813 (US); 1057, 1577 (MARY); 962, 1124 (Tawes); 1936
(Frost); 730 (Aacc).

Carex granularis Muhlenberg ex Willdenow var. haleana (Olney) Porter

Infrequent. Chain Bridge Flats and Roosevelt Island, in habitat similar to that of, and
usually with, the typical variety. Well-marked specimens occur in the gorge and are clearly
differentiated from typical C. granularis (and have been confused with other taxa). As is
reported from elsewhere, intermediate forms (e.g., 1937 [US, MARY, Tawes]) often are
found growing with this and the typical variety and cast some doubt upon the validity of
this taxon. This is the first record of var. haleana for the District of Columbia. Mid-May to
late June. 1127, 1938, Ward s.n. (US); 1938 (MARY); 1938 (GMUF); 1102 (MICH); 961, 1100
(Tawes); 1936 (Frost); 1056 (Aacc).

Carex grayi Carey

Frequent. In floodplain forests, most frequently on fine-textured (silt loam) soils and in
shade. Most plants seen would fit the more southern var. hispidula Gray (see Fernald, 1950),
a dubious taxon (Mohlenbrock, 1999). We report the species, for the first time, from the
District of Columbia and Arlington County. Late May to mid-July. 1076; 1121, 1703 (US);
1149, sn. (MARY); 1122 (Frost); 1077 (Aacc).

Carex grisea Wahlenberg ;

Common. In floodplain forests, open, moist woodlands, and disturbed areas. This species,
sometimes treated as Carex amphibola Steudel var. turgida Fernald, is distinguished from c
amphibola by its more inflated perigynia, which are rounded (not trigonous) in cross-section.
It is more restricted to floodplains than is C. amphibola. Early May to mid-June. 993, 1018,
1021, 1047 (US); 1050, 1155 (MARY); 1051, 1131 (MICH); 1103, 1458 (GMUF); 472, 1017,
1046, 1078, 1092, 1132 (Tawes); 995, 997, 1200, 1450 (Frost); 1019, 1049 (Aacc).

Carex hirsutella Mackenzie Q
Infrequent. In bedrock terrace forests. Found only at Bear Island, where somewhat
common. Mid-May to early June. 1582 (US); 1584 (MARY); 887 (T awes).

Carex hirtifolia Mackenzie . key R
Infrequent. In rich bedrock terrace and mesophytic alluvial terrace forests. Turkey Run

Park, Great Falls Park (Virginia), B d High Islands, and ravine of Cabin John Creek.

‘ ginia), Bear an 5

Ae. May to mid-June. 1447, 1459 (US); 1462 (GMUF); 731, 1027 (Tawes); 1559 (Grfa); 884
c).

Carex hitchcockiana Dewe
a ; mee
Rare. Mesophytic alluvial terrace forests and dry-mesic upland forests over ——
Found only at Turkey Run Park and at Turkey Island. A species of calcareous or malic
soils. Early May to early June. Vasey s.n. (US); 1941 (GMUF); 759 (T awes).

Carex hystericina Muhlenberg ex Willd
; g ex Willdenow :
Historical. Found below Chain Bridge in 1881 and at Lock 5 in 1899. A species of open,

Autrient-rich wetlands. Late May to early June. Steele s.n., Ward s.n. (US).

106 BARTONIA

Carex intumescens Rudge
Frequent. Edges of forested vernal pools, where it is a characteristic species. Early May
to late July. 450, 1236, 2084 (US); 1245 (MARY); 1714 (Grfa).

Carex jamesii Schweinitz

Common. In floodplain forests and open channel-shelf woodlands, most frequently on
finer-textured (silt loam) soils. A species of calcareous soils that is narrowly restricted to the
Potomac River corridor in this area. Late April to mid-June. 464, 1337 (US); 465, 1137
(Tawes); 1560 (Grfa); 1044B (Frost).

Carex laevivaginata (Kiikenthal) MacKenzie

Frequent. In wet, often mucky soils in swamps, floodplain forests, and seepages, and along
spring runs. Usually in more shaded situations than the similar C. stipata. Mid-May to late
June. 960, 1206, 1219 (US); 972 (MARY); 1564 (Grfa).

Carex laxiculmis Schweinitz var. copulata (Bailey) Fernald

Rare. In rich mesophytic forests on alluvial terraces. Found at Turkey Run Park and Bear
Island. This somewhat obscure taxon appears morphologically intermediate between typical
C. laxiculmis and C. digitalis, with pistillate spikes like those of the former and leaf width
and color like those of the latter. Fernald (1950) considered it to be a hybrid between those
two species, a view not shared by Mackenzie (1931), nor supported by limited systematic
research (Manhart, 1986). Rhoads and Klein (1993) consider it a calciphile; its habitat here
and that given for specimens at GMUF from other Virginia locations support this
evaluation. This is the first record of C. laxiculmis var. copulata for the Potomac Gorge and
for Maryland. Late April to early June. 1942, 1945 (US); 1463 (GMUF); 1946 (MICH); 1376,
1921 (Tawes).

Carex laxiculmis Schweinitz var laxiculmis
Common. In dry-mesic, often rich, upland forests and in mesophytic alluvial terrace
forests. Late April to early June. 885, 1142 (US); 891 (Tawes); 1470 (Grfa).

Carex laxiflora Lamarck

Infrequent. In rich dry-mesic forests, often in ravines or on steep slopes with bare soil
exposed. Found at High Island, in the Gold Mine Tract, and in several tributary stream
valleys on the Virginia side, usually in small numbers. Both wide and narrow-leaved -
occur. Treated as C. anceps Muhlenberg by Hitchcock and Standley (1919). C. laxiflora ©
Hitchcock and Standley (1919) evidently refers to C. gracilescens. Late April to early June
1383, 1453 (US); 1379 (MARY); 1024 (Tawes).

Carex leavenworthii Dewe sas

Infrequent. In low, disturbed areas, including open, channel-shelf woodlands, dry ener
prairies, and in openings in floodplain forests. Found at Bear, High, and Roosevelt i
Great Falls Park (Virginia), and Fletchers Floodplain. Has been listed as historical
Maryland (Maryland Natural Heritage Program, 1994), but probably has been enctioee
or misidentified. Mid-May to late June. 1084, 1140, 1153, 2097 (US); 1579 (MARY); i
1085, 1249 (Tawes); 1563 (Grfa); 1144 (Frost).

POTOMAC GORGE CAREX 107

Carex leptalea Wahlenberg ssp. leptalea

Rare. Spring runs and seepages. One station, at Great Falls Park (Virginia). Also, two
historical collections—one from apparently the same location as the current station and one
from Difficult Run. Mid-May to mid-June. 2087 (US, Grfa); Leonard 2176, Standley 11668
(US).

Carex lupuliformis Sartwell ex Dewey

Rare. Edge of forested vernal pool, near Widewater. Very similar to, and often confused
with, the more common C. /upulina. This is the first report of the species for the Potomac
Gorge, Montgomery County, and the Maryland Piedmont. Mid-June to early July. 2092
(US); 1259 (MARY); 1258 (GMUF); 1256 (MICH); 1260 (Tawes); 1257 (Dnhp).

Carex lupulina Muhlenberg ex Willdenow

Common. Moist, partially shaded to open areas, including swales in floodplain forests,
scoured channel shelf woodlands and edges of ponds, the C&O Canal, and forested vernal
pools. Early June to late July. 1096, 1238, 1265, 1266 (US); 552, 1128 (MARY); 1097 (Frost).

Carex lurida Wahlenberg

Common. In a variety of open wet areas, including scoured channel shelf woodlands,
open floodplain forests, wet meadows, edges of ponds and the C&O Canal, and edges of
forested vernal pools. Late May to mid-July. 1095, 1239, 2096 (US); 553 (Frost).

Carex mublenbergii Schkuhr ex Willdenow var. enervis W. Boott

Infrequent. Habitat similar to that of C. aggregata. Locations include near Plummers
land and Cropley. Var. enervis generally has the perigynium nerves less evident than in the
typical variety, but, in some specimens, this difference is not distinct. Mid-May to late June.
1486, 1487 (US); 1918 (Frost).

Carex mublenbergii Schkuhr ex Willdenow var. muhlenbergu ie er
Historical. A single collection in 1895 from High Island. The habitat is most likely dry
woods. Dates unknown; probably as for the var. enervis. Holm s.n. ’

Caret nigromarginata Schweinitz na

Frequent. In dry-mesic to xeric upland forests and rich bedrock terrace forests. This 1s a
- report of the species for Arlington County. Mid-April to early May. 977, 1382 (US);
84, 796 (MARY); 776 (Tawes); 791, 949 (Frost); 976 (Aacc).

Carex normalis M i
ackenzie a iri

Frequent. In moist swales in open channel shelf woodlands, wet areas in sored ae

nd edges, and edges of forested vernal pools. Mid-May to mid-June. 15 7 0, 171 (U ’ ’

Po:
1147, 1589 (MARY); 1157 (Tawes); 1569 (Grfa); 1158 (Frost); 1061 (Aacc).

Carex oligoca |
accom pace alluvial terraces. Found at Bear and High
kk. We report C. oligocarpa for the
1375, 1481 (US); 1026, 1449

); 1014, 1025 (Aacc).

108 BARTONIA

Carex pallescens Linnaeus

Historical. Known from a single collection in 1893 from “dry fields on the Potomac shore
near Aqueduct Bridge, Virginia” (near present-day Key Bridge in Arlington County). This
species, known from several late nineteenth century collections in the District of Columbia,
has not yet been found in Maryland and is considered highly rare in Virginia (Killeffer,
1999). It is apparently a sedge of fields and meadows, and, because of forest succession, may
be today locally less numerous or extirpated. Late May. Holm s.n. (US).

Carex pellita Muhlenberg ex Willdenow

Historical. Two collections at US (as C. lanuginosa Michaux, a misapplied name) from
two nearby locations (Lock 5, High Island) on the same date in 1898. A species of nutrient:
rich fens and other open wetlands. Mid-May. Steele s.n. (US).

Carex pensylvanica Lamarck -

Frequent. Primarily in rich bedrock terrace forests, also dry-mesic upland forests. Some
plants appear as forma gracilifolia (Peck) Kiikenthal. Late April to mid-May. 438, 443, 78,
991, 1344 (US); 430, 846, 979, 1341 (MARY); 980, s.n. (Tawes); 446, 1339 (Frost); 792, 982
Aacc).

Carex planispicata Naczi

Frequent. Rich, dry to mesic forests. Locally common at Bear Island and Great Falls Park
(Virginia). Widespread in the eastern United States, but obscured by nomenclatural and
typification problems. The species was recently described by Naczi (1999), who first
reported it from the gorge, based on an old collection at US from Plummers Island. The
very similar C. oligocarpa, which occurs in similar habitat, has generally fewer perigymia pet
spike, with the perigynia having distinct beaks. Mid-May to early June. 1043, 1067 (US);
1011, 1082, 1139 MARY); 1707 (GMUB); 1045 (MICH); 879, 1042, 1068, 1141 (Tawes); 1708
(Grfa); 1044A (Frost); 1069 (Aacc).

Carex platyphylla Carey

are. On steep slopes, often on bare soil, in rich, dry-mesic forests, evidently over
amphibolite. Found at Turkey Run Park, in the ravine of Cabin John Creek, and neat
Donaldson Run. In our area, an indicator of neutral to basic soils. Late April to late May.
1114, 1446, 1461 (US); 1115 (GMUF).

Carex prasina Wahlenberg ;

Frequent. In mucky soil along spring runs and in seepage swamps, also in wet swales .
floodplain forests. Early May to late June. 847, 957, 1205 (US); 874, 1264 (Tawes); /
(Grfa).

Carex radiata (Wahlenberg) Small f
Common. Primarily in low floodplain forests, open-channel shelf woodlands, ir 4
forested vernal pools, and edges of spring runs. Also occurs in disturbed situations in UP ~
habitats, such as young forests, along trails, and in mowed areas. One of our most ewe
species. Plants treated as C. rosea Schkuhr by Hitchcock and Standley (1919) and Fe pe
(1950) and many at US so identified follow the treatment of Mackenzie (193 1).0
referable to this species (Webber and Ball, 1979, 1984). Early May to late June. 483, 0.
1133, 1442, 1471 (US); 1490 (MARY); 1464 (GMUF); 1016 (Tawes); 1468 (Grfa); 1041 (Aa¢

POTOMAC GORGE CAREX 109

Carex retroflexa Muhlenberg ex Schkuhr
Infrequent. In rich bedrock terrace forest, Bear Island, where locally common. Early to
late May. 871, 890 (US); 1578 (MARY); Strong 87-042 (GMUF); 887, 1591 (Tawes).

Carex rosea Schkuhr ex Willdenow

Frequent. In dry-mesic upland forests, generally in drier and less disturbed habitats than
C radiata. C. convoluta MacKenzie of Hitchcock and Standley (1919) and Fernald (1950) is
referable to this species (Webber and Ball, 1979, 1984). Mid-May to mid-June. 1117, 1172,
1592 (US); 1173 (GMUF); 1203 (Tawes).

Carex scoparia Schkuhr ex Willdenow var. scoparia

Frequent. In moist swales in open channel-shelf woodlands, temporary pools in scoured
bedrock, edges of ponds and the C&O Canal, and along spring runs. Several specimens
appear as forma subturbinata (Fernald and Wiegand) Fernald. Early June to early July. 1143,
1231, 1930 (US); 1230, 2093 (MARY); 1706 (Grfa); 1225 (Frost).

Carex seorsa Howe

Historical. Two collections at US made from two nearby locations — Lock 5 (historically
called First Lock) and the C&O Canal Feeder Dam (near High Island) — on the same date
in 1898. Restricted in Maryland primarily to the Coastal Plain, where it is common on
floodplain forests. Mid-May. Steele s.n. (US).

Carex shortiana Dewe

Infrequent. In moist swales in channel shelf woodlands at Chain Bridge Flats and
Fletchers Floodplain. Also at edge of pond, Great Falls Park (Virginia). In Maryland and
northern Virginia, primarily restricted to rich soils along the Potomac. Mid-May to early
June. 1566 (US); Tanaka s.n. (NA); 1190, 1689 (Tawes); Grimshaw 842a (Grfa).

Carex sparganioides Muhlenberg ex Willdenow

Infrequent. In rich, dry-mesic forests and mesophytic alluvial terrace forests. Plummers
Island area, Bear Island, Turkey Run Park, Great Falls Park (Virginia), and, historically,
from the Virginia side of Chain Bridge. Occurs in less disturbed habitats than the similar C.
aggregata. Early May to mid-June. 1565 (GMUF, Grfa); Tanaka s.n. (NA); 1480, 1583

awes).

Carex squarrosa Linnaeus
Frequent. In moist swales in open floodplain forests or channel-shelf woodlands; also

edges of forested vernal pools. Mid-May to late July. 552 (US); 1129 (Tawes).

Carex stipata Muhlenberg ex Willdenow var. maxima Chapman
ool Tidal and oa ae swamps. Found at Roosevelt Island, Chain Bridge Flats,
and near C&O Canal Lock 5. Although specimens intermediate with the typical variety can
found, very wide-leaved plants clearly represent var. maxima. These are the first Ne
for the taxon for Montgomery County, the District of Columbia, and the Potomac ree.
Mid-May to late June. 1080 (US); 1079, 1576 (MARY); 1130, 1575 (MICH); 1073 (Tawes);
1074 (Frost).

110 BARTONIA

Carex stipata Muhlenberg ex Willdenow var. stipata

Frequent. In wet swales in open floodplain forests and channel shelf woodlands. Also
along edge of C8&O Canal and in other disturbed wetlands, in more generalized habitats
than var. maxima. Mid-May to mid-June. 458, 963, 1476 (US).

Carex straminea Willdenow ex Schkuhr

Rare. Found at a single station, along rocky pools in scoured riverside prairies, Great Falls
Park (Virginia). Historically collected in 1886 from the Maryland side of Great Falls
(specimen was originally identified as C. tenera and later determined by K. K. Mackenzie to
be C. richii Mackenzie, a synonym*for C. straminea). Also collected in 1983 from Bear
Island, in habitat similar to that of the current occurrence. C. hormathodes Fernald of
Hitchcock and Standley (1919) is referable to this species (= C. hormathodes Fernald var.
richii Fernald). C. straminea has been misapplied to C. albolutescens by some older treatments
(e.g., Hitchcock and Standley, 1919). Considered highly rare in Maryland (Frye and Lea,
2001) and in Virginia (Killeffer, 1999). Mid-May to early July. Knowlton s.n. (US); 1705
(GMUF, Grfa); Boone s.n. (Tawes).

Carex striatula Michaux

Rare. In xeric oak-pine bedrock terrace forest. Seen only at Bear Island, with several
historical collections elsewhere in the gorge. Within the gorge, specimens seem fairly distinct
from C. laxiflora, although difficult intermediates between the two occur elsewhere in
Maryland. C. striatula has been considered historical in Maryland (Maryland Natural
Heritage Program, 1994). This assessment may have been because of misidentifications or
from the species being overlooked. Late April to early June. 1922, Shreve 1467, Ward 3.
(US); 1378 (MARY).

Carex stricta Lamarck

Rare. Forested seepage swamp. A single location, at Great Falls Park (Virginia). All
previous material collected from the gorge as C. stricta at US and at the Great Falls Park
herbarium (single specimen, Grimshaw 842) has been determined to be C. emoryi. on
is more characteristic of swamps and riparian wetlands with low current velocities. Locally,
at least, it tends to grow in discrete, dense clumps that are not extensively stoloniferous.
This represents the first confirmed report of C. stricta for the Potomac Gorge. Early May
1472 (US, Grfa).

Carex styloflexa Buckley ;
Rare. Along seepages, spring runs, and edges of forested vernal pools. In the Gold po
Tract and at Great Falls Park (Virginia). Early May to mid-June. 1489, 1595, Shreve 1
(US); 1489, 1594 (MARY); 2086 (GMUF); 2085 (Grfa).

Carex swanii (Fernald) Mackenzie of

Frequent. In dry-mesic upland forests, rich bedrock terrace forests, and along ne
ar pools. Mid-May to late June. 876, 964, 1170 (US); 1233 (MARY); 1163 (Tawes)
Grfa).

Carex tenera Dew d
Rare. Floodplain forest, Fletchers Floodplain. This represents the first confir med byt <
from the Potomac Gorge for this regionally rare species, although collections of ©

POTOMAC GORGE CAREX 111

species have been attributed to it, including one from Great Falls reported by Shreve et al.
(1910). See also entries for C. festucacea and C. straminea. This is the only known extant
occurrence in the District of Columbia, and it is presently considered historical for
Maryland (Maryland Natural Heritage Program, 1994) and highly rare in Virginia (Killeffer,
1999). Late May. 1697 (US, Tawes).

Carex tonsa (Fernald) Bicknell var. rugosperma (Mackenzie) Crins

Rare. Seen only at Bear Island among rock outcrops in xeric oak-pine bedrock terrace
forest. Very similar to C. umbellata, but has a longer perigynium beak. Largely confined to
the Appalachian Mountains in Maryland. We report the taxon here for the first time for the
gorge and for Montgomery County. Early May. Frye and Lea s.n. (Tawes).

Carex tonsa (Fernald) Bicknell var. tonsa

Infrequent, but perhaps overlooked. In rocky riverside prairies, xeric Oak-pine terrace
forest, and on rock outcrops in upland forests. Seen in Gold Mine Tract and at Bear and
High Islands and is likely elsewhere. Mid-April to early May. 973 (US); 1001 (Tawes).

Carex torta F. Boott ex Tuckerman ca

Rare. One station found during this study, along rocky banks and depositional bars of
Cabin John Creek — apparently the same location as the specimen at GMUF. Terrell (1970)
reported a second station along the main stem Potomac. Observations suggest that it may
be declining in the area because of impacts of watershed development on stream flows. Late
April to early May. 1443, Terrell 4222 (US); 1444 (MARY); Miller 152 (GMUF).

Carex tribuloides Wahlenberg var. tribuloides

Common. In moist swales in open channel-shelf woodlands, open fresh tidal swamps,
open floodplain forests, and edges of ponds and the C&O Canal. Some atypical specimens
have been found (see C. projecta and C. tribuloides var. sangemonensis under excluded and
typothetical taxa, respectively). Late May to early July. 1126, 1210, 1234, 1242, 1929, 1939,
1944 (US); 508, 1193, 1196, 1221, 1241 (MARY); 966, 1222, 1248 (Tawes); 2252 (Grfa); 1228
Frost); 1156, 1209 (acc).

Carex typhina Michaux

P Rare. Edges of forested vernal pools, Bear Is

a tecords for the species for the Potomac Gorge an
4y to mid-July. 549, 1690 (US).

Jand and Fletchers Floodplain. These are the
d for the District of Columbia. Late

Carex: umbellata Schk illdenow

Frequent. In pemee wile xeric oak-pine terrace forest, rich renee ete
ress, and dry upland forests. Evidently included under C. tonsa by mare
tandley (1919). Mid-April to early May. 974 (US); 1336 (MARY); 777, 787,
(Grfa); 975 (Frost).

Carex vj

rescens Muhlenberg ex Willdenow
f Common, In dry-mesic cotana and rich bedrock terrace forests, a sn:
Tha, One of our most frequently encountered upland forest carices. Mia-May :

I
"12, 1164 (US); 435 (MARY); 434, 882 (Tawes); 1034 (Frost)

112 BARTONIA

Carex vulpinoidea Michaux

Common. In open, wet, often disturbed, areas, including open channel-shelf woodlands,
open fresh tidal swamp, banks of C&O Canal and ponds, and ditches. Mid-May to early
July. 534, 1099, 1123 (US); 1152 (MARY); 1148, 1154 (Tawes); 1562 (Grfa); 1134 (Frost); 740
(Aacc).

Carex willdenowii Schkuhr ex Willdenow
Frequent. In upland forests and rich bedrock terrace forests. Late April to mid-June. 1380,
2089 (US); 768 (Tawes); 1465 (Grfa); 1335 (Frost).

Carex woodii Dewey

Frequent. In rich, dry forests on bedrock terraces from Bear Island (where locally
common) to Vaso Island; not found on Virginia side. This early flowering species occurs in
several Maryland locations in large stoloniferous colonies reminiscent of, and often along
with, C. pensylvanica, and it has likely been overlooked as that species. Sterile C. woodii
plants have longer leaves that appear more blue-green than do those of C. pensylvanica.
Recently considered historical for Maryland (Maryland Natural Heritage Program, 1994). We
report C. woodii for the first time for the gorge. Late April to mid-May. 431, 1377 (US); 91,
1381 (MARY); 756, 762 (Tawes); 1342 (Frost); 758 (Aacc).

Excluded Taxa

Carex crebriflora Wiegand

A specimen at GMUF labeled collected from the area of Dead Run (a Potomac tributary
in the gorge) in 1987 and labeled C. crebriflora (Strong 87-044) was determined to be C
laxiflora. Inspections of specimens collected as C. crebriflora at GMUF suggest that many
reports of the species from northern Virginia are likely to be based on collections of similar
members of section Laxiflorae. C. crebriflora is a southern species that has not been reported
for Maryland and which collections suggest extends north only into southeastern Virginia,
a range consistent with that given by Fernald (1950).

Carex flaccosperma Dewey
Grimshaw and Bradley (1973) attributed C. flaccosperma to the flora of Great Falls “rs
(Virginia). The voucher specimen so labeled (Grimshaw 930; Grfa) is actually C. grisea.
accosperma (sensu stricto) is a southern taxon known to range north only to southeaster
Virginia (Fernald, 1950), while C. glaucodea (= C. flaccosperma Dewey var. 8
(Tuckerman ex Olney) Kiikenthal) historically has been recorded from the gorge:

Carex interior Baile hased
Hitchcock and Standley (1919) attributed C. interior to the Plummers Island area D A
on a collection at US made in 1915 (Standley 11663). This and a similarly labeled speci™®
from Great Falls, Maryland at US (Morris s.n.) were both determined to be C- atlantion
atlantica. C. interior is presently known from only one Maryland site (Frye and Lea, 200")

Carex nigra (Linnaeus) Reichard based
Hitchcock and Standley (1919) reported C. goodenovii Gay for the Washington arc? Both

on Ward’s (1881) citing of a report of C. vulgaris Fries by Vasey from Chain a to

names are apparently synonyms for C. nigra, a coastal species ranging from Green's

POTOMAC GORGE CAREX 113

Rhode Island (Fernald, 1950). No specimens were seen at US by Hitchcock and Standley
(1919) or by us. The most likely explanation for the Vasey report is a misidentification of
C. emoryi, a species in the same section as C. nigra that is frequent in the vicinity of Chain
Bridge, and is apparently not otherwise accounted for by Ward (188 1).

Carex projecta Mackenzie

Two collections of plants appearing to be C. tribuloides, but somewhat small, with the
lower spikes fairly well separated from one another, and with tips of the perigynia
somewhat spreading (thus, fitting some descriptions of this taxon) were made from Chain
Bridge Flats and Roosevelt Island during this study. A similar specimen was collected in 1888
by Holm (s.7.; NA) in the gorge from “shady woods near the Aqueduct Bridge, Virginia”
(near present-day Key Bridge in Arlington County) and identified as C. tribuloides
Wahlenberg var. reducta Bailey, a synonym of C. projecta. A. A. Reznicek has determined
the current collections to be C. tribuloides, and the 1888 specimen appears to be the same.
The Holm specimen seems to be at least part of the basis for the listing of C. projecta, which
is otherwise restricted to the Appalachian Plateau in Maryland, in the vicinity of
Washington, D.C., by Hitchcock and Standley (1919). We conclude that all material from
this area purported to be C. projecta represents depauperate or atypical C. tribuloides. See
entry for C. tribuloides var. sangemonensis in Hypothetical Taxa.

Carex tetanica Schkuhr .

C. tetanica has been attributed to the Potomac Gorge on the basis of two duplicate
specimens collected by Ward s.n. (US) in 1879 at High Island. One of these had been
annotated as C. granularis var. haleana by F. J. Hermann, and the second was so determined
by this study. No Washington, D.C. area specimens of C. tetanica have been seen, and
reports of its occurrence there (Hitchcock and Standley, 1919) appear to be based on these
oetaggd misidentified specimens or C. woodit (= C tetanica Schkuhr var. woodii [Dewey]

iley).

Hypothetical Occurrences

Carex albicans Willdenow ex Sprengel var. australis (Bailey) Retug
The possible occurrence of this taxon, frequently treated as Carex physorlyncha Liebman,
iN our area is problematic. It is a southern Coastal Plain taxon, said to range north to
southeastern Virginia (Fernald, 1950). Plants appearing much as C. albicans var. albicans, but
More robust, and with short rhizomes, a loosely, rather than densely, cespitose habit, and
wider leaves have been found in the gorge. A specimen collected by Ward in 1880 from
Great Falls, Maryland had been annotated by K. K. Mackenzie as C. physorhyncha, and =
apparently) subsequently annotated by Mackenzie as C. varia Muhlenberg (a paar -
albicans var. albicans or, possibly C. albicans, sensu lato). We have since onhaeeage ar
specimens in the gorge that compare well with syntypes of C. varia Muhlien ae ns
australis Bailey (= C. albicans var. australis) at US and at NY (as a epee é
ASSIA web site; Tracy s.7.). These are fairly restricted to rather xeric 0 a8 os
Consistent with that published for C. physorlryncha (Fernald, 1950, oo » 1 );
though they are sympatric with var. albicans, the typical variety tends to be allotopic .
adjacent, more mesic forests. A. A. Reznicek (pers. comm.) has examined these ak ak “a
Specimens and indicated that similar-appearing plants occur in the Midwest, north of t :
Published range of var. australis. He has concluded that, using a narrow circumscription 0

114 BARTONIA

C. albicans var. australis (perhaps necessary to give the taxon validity), such plants would
most likely be considered var. albicans. He acknowledges that the Potomac Gorge plants are
clearly atypical for var. albicans and that the situation may merit taxonomic review.
Specimens in question include Ward s.n. (US), Lea 981 (Tawes), Lea 1331 (GMUF, US), and
Lea 1332 (MICH).

Carex louisianica Bailey

A single collection of what is possibly this species (Lea 1201, Tawes) was made at Chain
Bridge Flats during this study; the specimen may also be a depauperate C. luputlina. C.
louisianica is primarily restricted to the Coastal Plain in Maryland. It is known historically
from the nearby Anacostia River (Steele s.n., US) and has been recently reported from the
Piedmont of northern Virginia (G. Fleming, pers. comm.).

Carex mesochorea Mackenzie

Hitchcock and Standley (1919) reported C. mesochorea from High Island. We have seen
no specimens from the gorge. It has been found in a number of Piedmont and Coastal Plain
counties in Maryland, usually in dry and open, often disturbed habitats, and it may well
occur in the gorge. It is easily confused with several similar species.

Carex molesta Mackenzie ex Bright

A single collection was made by Terrell s.n. in 1969 (US, det. F. J. Hermann) from along
the Potomac about 8 km upstream from the study area. The species may well occur in the
Potomac Gorge.

Carex tribuloides Wahlenberg var. sangemonensis Clokey

Mackenzie’s (1931) description of this taxon, purported to be a small, primarily southern
form of C. tribuloides, may apply to plants from the lower part of the gorge which key
toward C. projecta, including Lea 1194 (MICH), Lea 1211 (MARY), and the 1893 collection
by Holm s.n. (NA; see discussion under Excluded Taxa). The identity and validity of hcl
sangemonensis, however, is presently too unclear to us for making such a determination. C
tribuloides var. sangemonensis is reported to be highly rare in Virginia (Killiffer, 1999).

ACKNOWLEDGMENTS

We thank Mark Strong, Chris Tuccinardi, and other staff (US), Kevin Conrad (NA); Ted
Bradley (GMUF), Charles Delwiche (MARY), Ernie Schuyler (PH), and staff of the Cylbur
Arboretum for providing access to collections or other support. Anton A. Reznicek f
made determinations of several difficult specimens and generously shared his invalua ;
advice. Maryland Department of Natural Resources volunteer Dan Siehl processed tie
mounted at the Tawes herbarium a number of specimens collected during this study:
National Park Service supported fieldwork, including permitting collection of specimeé
acknowledge Dianne Ingram, Diane Pavek, Dan Sealy, and volunteers Jane an
Gibbins of that agency. Cris Fleming shared her knowledge from her pr
studies. Carol DiSalvo, Joe Kish, Sarah Anderson, Olin Allen, Jil Swearingen,
Steiner, and Dick Wiegand are thanked for various aspects of logistical support "ae. This
Bartgis and an anonymous reviewer provided helpful comments to improve this paper: =
project was funded by The Nature Conservancy’s Virginia E. Crouch Memor ial Eo was
by the Washington Biologists’ Field Club Research Fund. Additional in-kind support

ns; We

POTOMAC GORGE CAREX 115

provided by the Maryland Department of Natural Resources Wildlife and Heritage Service
and by the National Park Service, Assateague Island National Seashore.

LITERATURE CITED

BARTGIS, R., D. FELLER, C. SIMOES, E. THOMPSON, AND R. WIEGAND, 1993. State and nationally
significant natural areas of the Chesapeake and Ohio Canal National Historical Park. Unpublished
report to National Park Service. Maryland Department of Natural Resources, Natural Heritage
Program, Annapolis.

BROWN, M. L. AND R. G. BROWN. 1984. Herbaceous Plants of Maryland. Port City Press, Baltimore.

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. 1987 reprint. Dioscorides Press, Portland,
OR

District of Columbia. In press, Maryland Naturalist.

GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States
and Adjacent Canada, 2nd ed. New York Botanical Garden, the Bronx, 4

GRIMSHAW, S. AND T. R. BRADLEY. 1973. The vascular flora of Great Falls National Park, Fairfax
County, Virginia. Castanea 38: 229-261.

HARVILL, JR., A. M. 1973. Phytogeography of the carices of Virginia. Rhodora 75: 248-257.

HARVILL, JR., A. M., T. R. BRADLEY, C. E. STEVENS, T. P. WIEBOLDT, D. M. E. WaRE, D. W.
OGLE, G. W. RAMSEY, AND G. P. FLEMING. 1992. Atlas of the Virginia Flora, 3rd ed. Virginia
Botanical Associates, Burkeville, VA.

HITCHCOCK, A. S. AND P. C. STANDLEY. 1919. Flora of the District of Columbia and vicinity.
Contributions of the U.S. National Herbarium 21: 1-329.

HOLMGREN, P. K., N. H. HOLMGREN AND L. C. BARRETT. 1990. Index Herbariorum. Part 1,
edition 8. Regnum Veg. 120: x + 1-693.

KILLEFFER, S. E. 1999. Natural Heritage resources of Virginia: rare vascular plants. Natural Heritage
Technical Report 99-11. Virginia Department of Conservation and Recreation, Division of Natural
Heritage, Richmond, VA. Unpublished report.

LEA, C. 2000. Plant communities of the Potomac Gorge and their relationship to fluvial factors.
George Mason University, Fairfax, VA. M.S. thesis.
MACKENZIE, K. K. 1931. North American Flora. 18. Cyperaceae, tribe 2, Caricae. New York Botanical

Garden, New York. ;
MANHART, J. R. 1986. Foliar flavenoids of the North American members of Carex section Laxiflorae
Cyperaceae). Bi i tics and Ecology 14: 85-90.
ee oe. Rare, Threatened, and Endangered Plants of
Maryland. Maryland Department of Natural Resources, Annapolis. '
MCATEE, W. L. 1918. A sketch of the natural history of the District of Columbia, together with an
indexed edition of the United States Geological Survey’s 1917 map of Washington and vicinity.
Bulletin i i iety of Washington 1: 1-142. eis
ea aig gage gs Carex ‘The illustrated flora of Illinois). Southern Illinois
Universi d Edwardsville, IL. .
Nacz1, R. E. sige Digi erar erate a widespread and frequent new species of Carex viene
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of Sci : 37- ;
OSTERKAMP, ae agent C. R. Hupp. 1984. Geomorphic and vegetative characteristics along three
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t

albicans / C. emmonsit. Sida 14: 132-133.

116 BARTONIA

REZNICEK, A. A. 1989. New England Carex (Cyperaceae): T ic probl d phytogeographical
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Maryland. Maryland Weather Service Special Publication Vol. I. The Johns Hopkins Press
Baltimore.

TERRELL, E. E. 1970. Spring flora of the C&O Canal area, Washington, D.C. Castanea 35: 1-25.

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Museum No. 22. U.S. Government Printing Office, Washington,

WEAKLEY, A. S. 1999. Flora of Virginia and the Carolinas, working nee of 19 January 1999. The
Nature Conservancy, Chapel Hill, NC. Unpublished manuscript.

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WEBBER J. M. AND P. W. BALL. 1984. The taxonomy of the Carex rosea (section Phaestoglochin) is
Canada. Canadian Journal of Botany 62: 2058-2073.

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graphical and ecological notes. Rhodora 86: 151-231.

Bartonia No. 61: 117-121, 2002

Rare Plants in the Middle Branch of the Forked River
Watershed, Lacey Township, Ocean County, New Jersey

ALFRED E. SCHUYLER
Academy of Natural Sciences of Philadelphia
1900 Benjamin Franklin Parkway, Philadelphia PA 19103
schuyler@acnatsci.org

TED GORDON
Pine Barrens Inventories, 31 Burrs Mill Road, Southampton NJ 08088

During the spring and summer of 1998, we conducted a field survey to obtain
information about rare plant species and their distributions in the Middle Branch Forked
River watershed. To get information about rare plants previously known from the
watershed, we recorded data from herbarium specimens collected there that are in the
herbarium of The Academy of Natural Sciences. Gordon also supplied data from notes
made on previous visits to the area. We made visits to the area on 27 May, 21 July, 13
August, and 17 September 1998.

Plant nomenclature in this paper follows that in the Manual of Vascular Plants of
Northeastern United States and Adjacent Canada (Gleason and Cronquist 1991). We
determined rarity from listings provided by the New Jersey Department of Environmental
Protection and the Association for Biodiversity Information (www.natureserve.org),
although the rank and status for some of the species have changed from what they were in
the past.

DESCRIPTION OF THE STUDY AREA

The Middle Branch of the Forked River is one of three branches that join to form the
Forked River about 2 km west of where it enters Barnegat Bay. It drains a small watershed
that extends inland to the west for about 7 km with an average north/south width of about
2km. Our survey extends from the headwaters of the Middle Branch, near the abandoned
Tuckerton Railroad grade, downstream to the bridge for Route 9 (Fig. 1), which is about
2 km upstream from the junction of the Middle and North Branches. The Garden State
Parkway and a powerline along its west side bisect the area.

The New Jersey Conservation Foundation owns the land west of the Parkway. The land
between the Parkway and Route 9 is privately owned except for a strip along the stream
Corridor, which is owned by the township. The northern portion of this area 1s residential.
At the southeast end there is an industrial park.

For convenience of reference, we have divided the watershed into three zones from the
headwaters to Route 9. The headwaters portion west of the railroad grade is zone 1, the
rea between the railroad grade and the Parkway is zone 2, and the area between the
Parkway and Route 9 is zone 3. In zones 2 and 3, we recognize north and south portions

Manuscript submitted 3 November 2000, revised 12 April 2001.

117

118 BARTONIA

Cs eel te Fae
residential
= ea SS F orked
2N borrow ~*~ River
pit 3N \
ogler ee Ve
. Pond Ms
2, P At; \
2 LL
ie We Branch f the Forked AN
Dy, 2 “” anes O, Hox
S = c knoll )
Pe (3)
aE fk
alps FS
38 industrial
park 5
ee al = x n ine e Pa
1 km

Sketch map of watershed, Middle Branch of the Forked River. Characters in boldface
the Parkway, and the

Figure 1.
indicate the zones employed in the survey, which are delineated by the river,

railroad grade.
separated by the main stem of the Middle Branch. We have listed the rare plants found
throughout the area in Table 1 and those found in each zone in Table 2.

SEARCH RESULTS

Zone 1: Headwaters of the Middle Branch west of the railroad grade.

The Middle Branch is flooded on the west side of the embanked railroad grade re
functions as a dam. The many dead trees in this extensive pond-like habitat indicate that
conditions must have been drier in the past. We searched carefully for Helonias bullata a
Scirpus longii, two globally rare species collected here in 1915 by Academy Curator Bay
Long. Our searches for them and other rare species known from the watershed we

unsuccessful.

Zone 2N: North of the Middle Branch between the railroad grade and the Garden State
Parkway. ail

We confined our searches in this zone to the powerline clearing along the aati
the Parkway and a large shallow borrow pit. In or along the edges of the powerline oad
north of the borrow pit there were scattered stands of Rhbynchospora knieskernt oa
Calamovilfa brevipilis. Extensive stands of Sagittaria teres, Nymphoides pas
Utricularia resupinata occurred in shallow water of the borrow pit. In damp sa” ee ae
the edge of the pit, there were scattered stands of Rhynchospora knieskernit, and ‘es
of Schizaea pusilla. In drier sand, there were scattered small stands of Gentiana autum

RARE PLANTS OF MIDDLE BRANCH FORKED RIVER WATERSHED 119
TABLE 1. Rare Plants of the Middle Branch of the Forked River Watershed

Calamovilfa brevipilis (Torr.) Scribn.—pine barren reedgrass
Carex barrattii Schwein & Torr.—Barratt’s sedge

Carex livida (Wahlenb.) Willd.—livid sedge

Gentiana autumnalis L.—pine barren gentian

Helonias bullata L.swamp-pink*

Juncus caesariensis Coville—New Jersey rush

Leiophyllum buxifolium (Berg.) Ell.—sand-myrtle

Lobelia canbyi Gray—Canby’s lobelia

Muhlenbergia torreyana (Schultes) A. Hitchc.—pine barren smoke grass
Narthecium americanum Ker-Gawler—bog asph

Nymphoides cordata (Ell.) Fern.—floating-heart

Rhynchospora knieskernii Carey—Knieskern’s beaked-rush
Rhynchospora nitens (Vahl) Gray—short-beaked baldrush
Rhynchospora pallida M. A. Curtis—pale beaked-rush
Sagittaria teres Wats.—slender arrowhead

Schizaea pusilla Pursh—curly-grass fern

Scirpus longii Fern.—Long’s bulrush*

Scleria minor W. Stone—slender nutrush

Utricularia purpurea Walt.—purple bladderwort

Utricularia resupinata B. D. Greene—resupinate bladderwort
Uvularia puberula var. nitida (Britt.) Fern.—pine barren bellwort
Xerophyllum asphodeloides (L.) Nutt.—turkey-beard

*Our 1998 searches for Helonias bullata and Scirpus longii were unsuccessful. They were last collected in 1915.

Carex barrattii and Muhlenbergia torreyana were collected or observed nearby in 1985 by
Gordon. We did not search for them in 1998. We presume they are still present because
they are in an area where habitat conditions have not changed.

Park

We searched the main stem of the Middle Branch and adjacent wetland and terrestrial
habitats in the western portion of this zone. There were occasional stands of Juncus
Caesariensis in cedar swamp openings. Xerophyllum asphodeloides was frequent in open pine
forest bordering swamps. A few small stands of Calamovilfa brevipilis occurred in open
damp sandy/ peaty depressions. We also searched along a southern tributary in the eastern
Portion of this zone where Narthecium americanum occurred in two large stands of
approximately 200 x 30 meters and 100 x 20 meters. Schizaea pusilla was abundant on

ummocks in the larger stand.

Zone 2S: South of the Middle Branch between the railroad grade and the Garden State
Way.

Zone 3N: North of the Middle Branch between the Garden State Parkway and Route
9,

We searched the main stem of the Middle Branch for a distance of about 1 a in the
eastern portion of this zone. Over 10,000 stems of Narthecium americanum occurre in wet
areas along the stream. Carex livida, Juncus caesariensis, and Schizaea pusilla were oe
here too. In the low, open, pine forest at the eastern end of the zone, there were a lew
small stands of Uvularia puberula var. nitida along with scattered stands of ee

ipilis and Leiophyllum buxifolium. Xerophyllum asphodeloides was scattered t rough

120 BARTONIA

TABLE 2. General Locations of Rare Plants in the Middle Branch of the Forked River
Watershed.

Zone 1: Headwaters West of Railroad Grade

Helonias bullata' Scirpus longii'
Zone 2N: North Portion between Railroad Grade and Parkway
Calamovilfa brevipilis Rhynchospora knieskernii
Carex barrattii’ Sagittaria teres
Gentiana autumnalis Schizaea pusi
Mublenbergia torreyana’ Utricularia resupinata
Nymphoides cordata
Zone 2S: South Portion between Railroad Grade and Parkway
movilfa brevipilis chizaea pusilla
Juncus caesariensis Xerophyllum asphodeloides

Narthecium americanum

Zone 3N: North Portion between Parkway and Route 9

Calamovilfa brevipilis Rhynchospora knieskernii
Carex livida Rhynchospora nitens*

Juncus caesariensis Schizaea pusi

Leiophyllum buxifolium Uvularia puberula var. nitida
Lobelia canbyi Xerophyllum asphodeloides

Narthecium americanum

Zone 3S: South Portion between Parkway and Route 9
6 Schi

movilfa brevipilis chizaea pusilla

Juncus caesariensis Scleria minor

Leiophyllum buxifolium Utricularia pu

Narthecium americanum Xerophyllum asphodeloides

Rhynchospora pallida os a ee

‘Collected in 1915. Not relocated in 1998 after careful searching.

*Observed in 1985. Not searched for in 1998, presumed still extant.

*Collected in 1985. Not searched for in 1998, presumed still extant.
‘Collected in 1985. Not searched for in 1998, status uncertain. Site is in heavily disturbed residential area.

portions of open pine forest bordering the extensive swamp along the Middle Branch. We
also searched along the northernmost tributary of the Middle Branch in this zone. se
found Juncus caesariensis and Schizaea pusilla in swamp openings adjacent to the stream. ‘
former also occurred abundantly with numerous plants of Lobelia canbyt at the upper ea
of Dogleg Pond. In a sandy dug depression near the western end of Dogleg Pond, pon
were stands of Rhynchospora knieskernii. A small stand of Calamovilfa brevipilis pose
along the edge of a sand road south of Dogleg Pond. In 1985, Gordon wer
Rhynchospora nitens in a pond heavily disturbed by residential development in the nort
portion of this zone. We did not visit this pond in 1998.

9,
Zone 3S: South of the Middle Branch between the Garden State Parkway ant
South of the extensive cedar swamp that borders Middle Branch is a maze of porns sie
sand roads, dug ditches, dug ponds, and extensive deposits of dredged sand. be

RARE PLANTS OF MIDDLE BRANCH FORKED RIVER WATERSHED 121

scattered stands of dead small trees of Chamaecyparis thyoides that were killed by fire.
Despite this disturbance, the area has a diverse terrestrial/wetland flora. Scattered stands of
Juncus caesariensis, Narthecium americanum, and Schizaea pusilla were in or along the margins
of dug ditches and ponds. Utricularia purpurea was abundant in a long canal-like broad ditch
south of the knoll. A small stand of Scleria minor was on a moist sandy-peaty slope
bordering a ditch northwest of the knoll. Rhynchospora pallida was abundant in moist open
sandy depressions in the southeastern portion of the zone. Calamovilfa brevipilis primarily
occurred along the edges of roads in damp sand throughout lower parts of the zone.
Lewophyllum buxifolium and Xerophyll: phodeloid d th lower parts
of the area in dry sand.

CONCLUSIONS

The Middle Branch of the Forked River harbors a diverse assemblage of globally and
state-rare plant species. It is an area that clearly merits some form of protection and
management to maintain proper conditions for the continued existence of these species.

In some parts of the area, human disturbance has been beneficial for the rare plants. The
large borrow pit, powerline clearing, numerous sand roads (some heavily used by
recreational vehicles), dug ditches and ponds, and extensive deposits of dredged sand all
provide early-successional habitats for rare plants. Periodic logging and fires also create such
habitat conditions.

Some forms of human disturbance, however, are more severe and detrimental to rare
plants. Extensive landscaping associated with residential and industrial development is in
this category. The area between the Parkway and Route 9 is zoned for such development.
We think steps should be taken to prevent this from happening.

Because so many rare species grow in early successional habitats created by human
disturbance, it is not enough to merely protect land by allowing it to stay in a wild state.
Without disturbance, the stronger competitors that grow in late successional habitats would
teplace rarer species. Therefore, a management plan should be designed for the entire
watershed that allows for a balance between human disturbance and the survival of rare
Species,

ACKNOWLEDGMENTS
This work is part of a long-range study supported by The Academy of Natural Sciences
to evaluate and protect environmentally important areas on the west side of Barnegat Bay

in Ocean County, New Jersey. We thank Britton Chance and Ruth Patrick for their
Support and interest in our work.

LITERATURE CITED

GLEASON, H. A. AND A, CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States
and Adjacent Canada. 2nd ed. The New York Botanical Garden, Bronx.

a.)
2 OFEFY Re <4] dagen tres obs wysne «sv i :

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3h pay He SVUGFTET SS 4
—_— 2 oe i tn 7
ee Si im Mees road}
f ites bY- CRED a aa 372 : as 2
: | PERS a) = : se « ; si
u ‘ay © i ey ~~ a9 ' &
: id iment f° sz ne 7 i . -
i+ i ian As a Toa ae q
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; $ Rt dea” at Pen ee cas ss 3
6 Be wr ; le re J
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*
7 — ¥
s <e ree P eT a,
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ot 5 ak y 46 Shake re. ae : M
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ma Ga 5 Sag) ae giteia® a ee te eee ee

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Aner ate OEE, o> Mitel Ay QA og cere mgs cpp regye, > hoe eee
5 J - id 5 ‘ * hs » m eaak —" tie
iat) Gey ET ogee SERRE Gon rnc eis paren e Rares a ae
: : 7 : ba » ie
eee. 405 2 gh Seige ee aS vi ret a ree # Pay aes
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- =i a a ee jee d- ‘ . pts? hese a o+

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Palen E

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7 : sc id 4 A + eA nets a Etipase igas or rg 3 i; = Bee ee he

Bartonia No. 61: 123-130, 2002

The Vascular Flora of Statue of Liberty National Monument,
New York Harbor

RICHARD STALTER AND NELSON TANG
Department of Biological Science, St. John’s University, Jamaica NY 11439

ABSTRACT. The vascular flora of Statue of Liberty National Monument, a 4.86 hectare island in
New York Harbor, consists of 97 species in 82 genera and 40 familes. The largest families in the flora
are the Asteraceae (18 species) and Poaceae (16 species). Together these families comprise 34% of the
flora. Non-native species, 65% of the flora, are a major component of the natural vegetation. A list
of the 97 species of vascular plants is included in an appendix.

INTRODUCTION

The Statue of Liberty National Monument, New York County, New York, is located on
Bedloes Island, an island in the upper bay of New York Harbor (Anonymous 1999). Bedloes
Island is named after its first owner, Isaac Bedloe, who acquired the island as a grant from
the governor of New York prior to 1670. The island’s ownership passed through several
families before it was purchased by New York City to be used as a quarantine station.
During the 1790s, the federal government expresed an interest in the site as a port, and in
1800 the title to the island was transferred from New York State to the federal government.
By 1808, construction began on an 11-point star fort, which was completed in 1811. The
fort was named Fort Wood, in 1814, in honor of Colonel Eleazer D. Wood, an officer killed
in the War of 1812. Until the Civil War began in 1861, the fort was garrisoned with
infantry and artillery; in 1861 it became a recruiting station and ordnance depot. After the
end of the Civil War, from 1865 to 1877, the fort was garrisoned by a small number of
soldiers. In 1877 the fort was selected as the site for the new statue, “Liberty Enlightening
the World” (Anonymous 1999). vat

With the exception of an 1886 photograph showing what probably is Ailanthus, there are
no photos of sufficient quality and clarity to identify the vegetation at the island. There are
references to “Horse Chestnut” in old reports, but no vegetation lists exist for the island.
The designer of the statue, Auguste Bartholdi, envisioned the statue protruding from a tree-
covered island. But trees and people were not compatible and the vegation at the site today
consists of a well maintained lawn, hedges of Taxus cuspidata and Euonymus atropurpurea and
arborescent plantings of Acer platanoides, Platanus acerifolia, Prunus serrulata and Tilia
cordata.

A literature search and consultation with park personnel indicate that there is no
published or “in house” list of the vascular flora of Statue of Liberty National Monument.
The objective of this study is to identify the vascular flora at the monument and compare
the non-native and native vascular plants at this site.

Waa
Manuscript submitted 3 October 2000, revised 16 Feburary 2001.

123

124 BARTONIA
METHODS

Six collecting trips were made to the study area during the growing season from 9
September 1999 to 15 August 2000. Objectives for each trip included collecting voucher
specimens and accumulating information on abundance and apparent habitat preference for
each species. Gleason and Cronquist (1991) was consulted to determine the native status of
each taxon.

More than 250 specimens form the basis for this study. Taxonomically problematic
specimens were sent to various experts for annotation; experts consulted are listed in the
acknowledgments section. A complete set of voucher specimens has been deposited at the
National Park Service (NPS) herbarium at Ellis Island, New York, and partial duplicate sets
have been deposited in the herbaria of Brooklyn Botanic Garden (BKL), Missouri Botanical
Garden (MO), and the University of South Florida (USF). Accession numbers will be
assigned by the National Park Service to the primary set of specimens at the herbarium at
Ellis Island and will be available upon request from the National Park Service.

Nomenclature follows Gleason and Cronquist (1991). Ornamental shrubs and trees were
classified according to Rehder (1990).

RESULTS AND DISCUSSION

The vascular flora of Statue of Liberty National Monument consists of 97 species in 82
genera and 40 families (Table 1). The major families of the total flora are the Poaceae (16
species) and Asteraceae (18 species); 34% of the species comprising the total flora are
contained in these families. Sixty-three species, or 65% of the flora, are not native to New
York State (Gleason and Cronquist 1991). Seventeen families represented in this survey *
cmprised exclusively of non-native species. Families with large numbers of non-native plants
are the Poaceae (11 species) and Asteraceae (10 species). Most species, native and non-nativé
occur principally in disturbed soil near the maintenance building, lawns, the drain
surrounding the base of the statue and along the edge of the sea wall.

By comparison, New York State’s vascular flora consists of 3,603 species, including yf
pteridophytes, 31 gymnosperms and 3,457 angiosperms. The most recent checklist ‘
persisting vascular plants, compiled by Mitchell and Tucker (1997), includes 3,195 species:
of these 2,078 (65%) are native and 1,117 (35%) are non-native. nid

Information in the Cultural Landscape Report (Anonymous 1999) lists several woo?)
species that have been planted by the National Park Service: oriental cherry att
serrulata), Norway maple (Acer platanoides), London plane tree (Tilia cordata), Japanese b wl
pine (Pinus nigra). Horse-chestnut (Aesculus hippocastanum), oaks (Quercus spp.) and om a
(Acer spp.) were present at the island when it was used as a military post. A photograP
the pedestal in 1886 includes Ailanthus altissima; this taxon is represented by a few -
saplings today. Of the planted trees, only Tilia cordata and Gleditsta triacanthos
reproducing at the site.

Notable planted shrubs include a hedge of Japanese yew (Taxus cuspidata) and nee
bush (Euonymus atropurpurea). Additional shrubs planted in the 1960s were Amur Pls)
(Ligustrum amurense), cockspur hawthorn (Crataegus crus-galli), English ivy (H poi ‘a the
and periwinkle (Vinca sp.). Ligustrum amurense and Crataegus hedges were aah : duals
mid-1980s. Ligustrum amurense is represented by a few naturally reproducing 19 "
near the sea wall.

VASCULAR FLORA OF STATUE OF LIBERTY NATIONAL MONUMENT 125

TABLE 1. Statistical summary of the vascular flora of Liberty Island, New York.

Ferns Gymnosperms Dicots Monocots Total
Families 1 1 33 5 40
Genera 1 1 66 14 82
Species 1 1 75 20 97
Native Species 1 0 26 ‘a 34
Introduced Species 0 1 49 13 63

ANNOTATED CHECKLIST OF SPECIES

The vascular plant taxa have been arranged according to the following categories:
pteridophytes, gymnosperms, dicots, and monocots. Within each category, families and
lower taxa are arranged alphabetically. The concept of families follows Cronquist (1988).
Nomenclature primarily follows Gleason and Cronquist (1991), but taxonomic revisions in
Kartesz (1994) were consulted to update the nomenclature. Each entry includes notes on
habitat and frequency in the study area using the categories rare (less than 5 occurrences),
infrequent (5 to 20 occurrences), frequent (more than 20 occurrences).

POLYPODIOPHYTA
Onocleaceae
Onoclea sensibilis L. One individual at southeast base of statue; rare.

PINOPHYTA
axac
*Taxus cuspidata Sieb. & Zucc. Occasional seedlings, living quarters area; rare.

MAGNOLIOPHYTA-MAGNOLIOPSIDA

an
* i, ‘ : : > ;
Amaranthus crispus (Lesp. & Thev.) N. Terrac. Disturbed soil, maintenance area; rare
Amaranthus hybridus L. Disturbed soil near maintenance area, rare.

Anacardiaceae
Rhus copallina L. Disturbed sites near snack bar, one plant; rare. -
Toxicodendron radicans (L.) Kuntze [Rhus radicans L.]. Japanese black pine grove; intreq.

Apiaceae
“Daucus carota L. Lawns and disturbed sites; infreq.

Apocynaceae
Apocynum cannabinum L. Japanese black pine grove; rare.

Pe
Introduced

126 BARTONIA

Asteraceae

* Achillea millefolium L. subsp. lanulosa (Nutt.) Piper. Lawn and disturbed sites; freq.

Ambrosia artemisiifolia L. Disturbed soils, maintenance area; rare.

* Arctium minus Schk. Disturbed sites, near Japanese black pine grove; rare.

*Artemisia vulgaris L. Disturbed sites; freq.

Aster dumosus L. Disturbed soils, maintenance area; rare.

Baccharis halimifolia L. Near base of statue; rare.

Cirsium arvense (L.) Scop. Disturbed soil, maintenance area; infreq.

Conyza canadensis L. (Cronq.) var. canadensis [Erigeron canadensis L. var. canadensis}
Gardens and disturbed sites; freq.

Erigeron strigosus Muhl. Disturbed soil; infreq.

Eupatorium rugosum Houtt. Disturbed soil near park headquarters; rare.

Galinsoga quadriradiata Ruiz & Pavon. Disturbed sites; freq.

*Hieracium caespitosum Dumort. Disturbed sites and lawns; freq.

*Lactuca serriola L. [L. scariola L.]. Disturbed soils, Japanese black pine grove; infreq.

*Matricaria discoidea DC. [Matricaria matricarioides auct. non (Less.) Porter]. Disturbed sites
and lawn near living quarters; infreq.

*Senecio vulgaris L. Gardens, lawns, disturbed sites; freq.

*Sonchus asper (L.) Hill. Disturbed sites; infreq.

*Sonchus oleraceus L. Disturbed sites; infreq.

* Taraxacum officinale Weber. Lawns and disturbed sites; freq.

Brassicaceae

*Alliaria petiolata (Bieb.) Cavara & Grande. Japanese black pine grove; infreq.
*Barbarea vulgaris R. Br. Near sea wall; rare.

*Capsella bursa-pastoris (L.) Med. Disturbed sites; infreq.

Lepidium virginicum L. Disturbed soils; infreq.

Caprifoliaceae
*Lonicera japonica Thunb. Under Japanese black pine; rare.

Caryophyllaceae

*Arvenaria serpyllifolia L. Disturbed soils and pathsides; freq. ides

*Cerastium fontanum Baumg. subsp. vulgare (Hartman) Greuter & Burdet [C. asa
Fries var. vulgare (Hartman) Hylander, C. vulgatum L.]. Lawns and disturbed sites; intreq:

*Cerastium glomeratum Thuillier [C. viscosum L.]. Disturbed sites; infreq.

Caesalpiniaceae
*Gleditsia triacanthos L. Seedlings from cultivated plants; infreq.

Chenopodiaceae

*Atriplex patula L. Near sea wall, maintenance area; rare.
*Chenopodium album L. Gardens and waste areas; freq.
*Chenopodium ambrosioides L. Disturbed soils; infreq.

*Introduced

VASCULAR FLORA OF STATUE OF LIBERTY NATIONAL MONUMENT

Clusiaceae
Hypericum punctatum Lam. Lawn near living quarters; infreq.

Convolvulaceae
*Convoluulus arvensis L. Disturbed sites and on lawns; freq.

Euphorbiaceae

127

Acalypha virginica var. rhomboidea (Raf.) Cooperider [Acalypha rhomboidea Raf.]. Disturbed

soils; infreq.

maesyce maculata (L.) Small [Euphorbia maculata L., E. supina Raf.]. Gardens and

disturbed soil; freq.

Fabaceae

*Medicago lupulina L. Disturbed soil near maintenance area; rare.
"Trifolium aureum Pollich. Lawns; infreq.

*Tnfolium bybridum L. Roadsides and lawns; rare.

“Trifolium repens L. Roadsides, lawns and waste places; freq.

eae
‘Lamium amplexicaule L. Disturbed soils and lawns; infreq.

Malvaceae
*Abutilon theophrasti Medik. Disturbed soils, maintenance area; rare.
ep:

s syriacus L. Seedlings from planted specimens, living quarters; rare.

‘Hibiscus trionum L. Disturbed soil, work area near snack bar; rare.
Malva neglecta Wallr. Gardens and disturbed sites; infreq.

Molluginaceae
“Mollugo verticillata L. Gardens and disturbed sites; infreq.
Moraceae
“Morus alba L. Near sea wall and Japanese black pine grove; rare.
Oleaceae
‘Ligustrum amurense Carr. Near sea wall, maintenance area; rare.
Oxalidaceae
Onalis dillenii Jacq. Gardens and disturbed sites; infreq.
Phytolaccaceae

lacca americana L. Disturbed sites; rare.
Plantaginaceae

o
lantago lanceolata L. Lawns and disturbed sites; freq.
tago major L. Lawns and disturbed sites; infreq.
oe
Toduced

128 BARTONIA

Platanaceae
*Platanus acerifolia (Ait.) Willd. Occasional seedlings from planted trees; rare.

Polygonaceae

Polygonum aviculare L. Disturbed sites; infreq.

Polygonum pensylvanicum L. Disturbed soil, maintenance area; rare.
Polygonum ramosissimum Michx. var. ramosissimum. Near sea wall; rare.
*Rumex acetosella L. Disturbed soils and lawns, living quarters; infreq.
*Rumex crispus L. Disturbed soil, Japanese black pine grove; infreq.

Portulacaceae
* Portulaca oleracea L. Disturbed sites, and lawns; freq.

Ranunculaceae
*Delphinium ajacis L. Disturbed soil, maintenance area; rare.
Ranunculus abortivus L. Japanese black pine grove, 5 individuals; rare.

Rosaceae
*Prunus avium L. Gardens, living quarters, one tree; rare.
Prunus serotina Ehrh. Near sea wall, maintenance area; rare.

Salicaceae
* Populus deltoides Marshall. One plant, base of statue; rare.

Simaroubaceae
* Ailanthus altissima (P. Mill.) Swingle. Several individuals, Japanese black pine grove; rare:

Solanaceae

*Lycopersicon esculentum L. Flower garden, maintenance area; rare.

*Solanum dulcamara L. Disturbed sites; freq.

Solanum nigrum L. var. virginicum L. [S. americanum Miller]. Disturbed sites; rare-
Tiliaceae

*Tilia cordata L. Disturbed sites; rare.

Ulmaceae

Celtis occidentalis L. Japanese black pine grove; rare.
*Ulmus pumila L. Near sea wall, maintenance area; rare.

Violaceae
Viola sororia Willd. [V. papilionacea Pursh]. Lawns, disturbed soils; freq.

Vitaceae
Parthenocissus quinquefolia (L.) Planchon. Japanese black pine grove; rare.

*Introduced

VASCULAR FLORA OF STATUE OF LIBERTY NATIONAL MONUMENT 129

MAGNOLIOPHYTA-LILIOPSIDA
Commelinaceae
*Commelina communis L. Disturbed sites, lawns; infreq.

C

yperaceae
Cyperus esculentus L. Disturbed soil, maintenance area; infreq.

lridaceae

Sisyrinchium angustifolium Miller. Lawn near path around statue; rare.

Liliaceae
“Allium vineale L. Lawns; infreq.
*Omithogalum umbellatum L. Lawn near living quarters; infreq.

Poaceae

Agrostis perennans (Walt.) Tuckerman. Disturbed sites; infreq.

*Aira caryophyllea L. Disturbed soils; infreq.

*Bromus tectorum L. Disturbed soils; infreq.

"Bromus racemosus L. Lawn near living quarters; rare.

“Dactylus glomerata L. Lawn near living quarters; rare.

"Digitaria sanguinalis (L.) Scop. Lawns and disturbed sites; freq.

*Eleusine indica (L.) Geartn. Disturbed sites; infreq.

Festuca rubra L. Lawns and disturbed soils; freq.

“Lolium perenne L. Disturbed soils; infreq.

Panicum dichotomiflorum Michx. Disturbed soil, maintenance area; rare.
° hragmites australis (Cav.) Trin. [P. communis Trin.]. Growing on base of statue; rare.
hing annua L. Lawns, disturbed soil; freq.

zs 04 pratensis L. Lawns; infreq.

Setaria glauca (L.) Beauv. Disturbed soils, maintenance area; infreq.
Setaria viridis (L.) Beauv. Disturbed soils; infreq.

Sorghum halepense (L.) Pers. Disturbed soil, maintenance area; infreq.

———
Introduced

ACKNOWLEDGMENTS

The authors gratefully acknowledge the assistance of the following individuals who are
ee wedged as experts in identification of the plants in the following families: Ihsan Al-
ehbaz (Brassicaceae), Steven Clemants (Amaranthaceae and Chenopodiaceae), Bruce
Mi; n (several families), Eric Lamont (Asteraceae), Robert Meyer (Poaceae), Richard

tchell (Polygonaceae). Appreciation is expressed to Steve Dial who reviewed the
Hanuscript, and to Divina Lopez who assisted in the preparation of this paper. Special
to Dr. Al Farrugio, Ellis Island National Historic Site, who provided collecting
and access to the island. The collection of plants will eventually be housed in the
um at Ellis Island.

Permits
herbari

130 BARTONIA
LITERATURE CITED

ANONYMOUS. 1999. Cultural Landscape Report for Liberty Island, Statue of Liberty National
Monument. History, Analysis and ag: eee Olmstead Center for Landscape
Preservation, National Park Service, Boston.

CRONQUIST, A. 1988. The Evolution and “a of Flowering Plants. 2nd ed. The New
York Botanical Garden, Bronx. 555 pp.

GLEASON, H.A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of the Northeastern United
States, Canada, and Greenland. 2nd ed. The New York Botanical Garden, Bronx. 910 pp.
KARTESZ, J.T. 1994. A Synonymized Checklist of the Vascular Flora of the United sg Canada,

and Greenland. 2nd ed. Volume 1. Timber Press, Inc., Portland, Oregon.

MITCHELL, R.S. AND G.C. TUCKER. 1997. Revised Checklist of New York > Pans Bulletin
No. 490. New York State Museum, State Education Department, Alban

REHDER, A. 1990. Manual of Cultivated Trees and Shrubs. Dioscorides Press, ues Oregon. 99%
Pp.

Bartonia No. 61: 131-136, 2002

Pteridophyte Distribution by Township in
Venango County, Pennsylvania

CARL F. CHUEY
Herbarium, Biological Sciences, Youngstown State University, Youngstown, OH 44555

The distribution of the pteridophyte flora in Venango County, Pennsylvania was
investigated as part of an ongoing study of pteridophyte distribution by township or district
in western Pennsylvania, northern West Virginia and eastern Ohio (Chuey 1976, 1980, 1983;
Chuey and Sturm 1969; Isaac and Chuey 1992a, 1992b).

Venango County, formed in 1800, contains 1,740 km? (679 sq. mi.) and is located in
northwestern Pennsylvania. The 1990 population was 59,381. The density, 34.1 persons
km? (87.9 persons per sq. mi.), is the tenth largest in western Pennsylvania. Major
population centers include Franklin, the county seat, Oil City and Polk. Venango County
increased in population with every census except two through 1940 (Riesenman 1943); this
trend continued with only two declines in the succeeding censuses (Andriot 1980).

All but the western edge is unglaciated (Wherry et al. 1979). Thus, it is in both the
Glaciated and the Unglaciated Allegheny Plateau physiographic provinces. Located in the

egheny River basin, the major tributaries in the county include French Creek, Oil Creek,
Sandy Creek and East Sandy Creek. According to Braun (1950), it is at the interface of the
Hemlock-White Pine-Northern Hardwoods Region and Mixed Mesophytic Forest Region;
Kuchler (1964) put it at the interface of Northern Hardwoods and Appalachian Oak Forest.

METHODS

_ At the Herbarium of Youngstown State University (YUO), the fourth largest herbarium
bie Ohio, Venango County is represented by the largest holding from one county, with 1,403
ore out of the 13,344 specimens from Pennsylvania. Carnegie Museum (CM), the
argest herbarium in western Pennsylvania with 567,000 specimens (Holmgren et al. 1990),
- 2,350 specimens from Venango County. Information on species distribution presented
in this paper (Table 1) is based solely on these specimens. The classification hierarchy is
i Crabbe et al. (1975) and the nomenclature follows that of the Flora of North
menca (Flora of North America Editorial Committee 1993).

RESULTS AND DISCUSSION
Despite the county’s relatively high population density, some taxa such as Equisetum
Dryopteris intermedia, Polystichum acrostichoides and Onoclea sensibilis have been

4rvense
found i ras
und in all townships (Table 1). With additional fieldwork I would expect Diphastastrum

Lo
“uscript submitted 1 November 1995; revised 2 September 1999.

131

BARTONIA

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BARTONIA

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VENANGO COUNTY PTERIDOPHYTE DISTRIBUTIONS 135

digitatum, Lycopodium obscurum, Osmunda cinnamomea, O. claytoniana, Athyrium filix-
femina, Dennstaedtia punctilobula, Pteridium aquilinum and Thelypteris noveboracensis also to
be found in all townships.

Based on the herbarium label data, some of the more unusual taxa such as Asplenium
montanum, A. trichomanes, Dryopteris goldiana, Gymnocarpium dryopteris, Phegopteris
hexagonoptera, P. connectilis, Ophioglossum vulgatum and Botrychium matricariifolium have
not been found since the mid-1960s or occur only in isolated locations. Some may no
longer be present in the flora. Rhoads and Klein (1993) reported Botrychium lanceolatum,
B. simplex, and Ophioglossum pusillum for the county and Parks (1989) reported that Vittaria
and Trichomanes gametophytes are found in Venango County; however, there are no
specimens at either CM or YUO. Lycopodium hickeyi, Equisetum fluviatile, E. sylvaticum,
Lygodium palmatum, Pelleae atropurpurea, Cystopteris bulbifera, C. fragilis, Dryopteris
dintoniana, Woodsia obtusa, Woodwardia areolata and W. virginica have been found in
adjacent counties (Rhoads and Klein 1993) and could reasonably be expected in Venango
County, although there are no specimens from the county at CM or YUO. None of the
taxa represented by specimens at CM or YUO is listed as endangered or threatened (Wild
Resource Conservation Fund 1995).

With increased population pressures requiring more land for homes, shopping malls and
toads, the fern and fern allies can be expected to be reduced in numbers. But Venango
County has an abundance of wild land, including State Game Lands No. 39, No. 45, No.
47, No. 96, and No. 253 (collectively containing 9,123 ha [22,542 acres]), the Allegheny
River area of Clearfield State Forest (1,281 ha [3,165 acres]), Oil Creek State Park (2,843 ha
(7,026 acres]), Two Mile Run County Park (1,091 ha [2,695 acres]), and conservation
casements of the Western Pennsylvania Conservancy in President Township (4,570 ha
(11,300 acres]). We should expect most, if not all, of the taxa to remain a part of the flora
of Venango County.

LITERATURE CITED

ANDRIOT, J. L. 1980. Population Abstract of the United States. Andriot Associates, McLean, VA.
BRAUN, E.L. 1950. Deciduous Forests of Eastern North America, Hafner Publishing Co., New York.
Y,C. F. 1976, Tricounty pteridophytes: a catalog of ferns and their allies occurring in
Trumbull, Mahoning, and Columbiana Counties, Ohio. Laitsch Herbarium Publications of
Youngstown State University 1: 1-26.
HUEY, C. F. 1980, Pteridophyte distribution: study of the ferns and their allies in the upper Ohio
C River basin. Laitsch Herbarium Publications of Youngstown State University 2: 1-57.
HUEY, C.F. 1983. Pteridophyte flora of Ashtabula County, Ohio by township. Laztsch Herbarium
Publications of Youngstown State University 3: 1-48.
CHUEY, C. F, AND N. STURM. 1969. Pteridophyte checklist of Ashtabula, Trumbull and Mahoning
cq cuuaties, Ohio. American Fern Journal 59: 41-44.
ABBE, J.A., A.C. JERMY, AND J. T. MICKEL. 1975. A new generic sequence for the pteridophyte
Ron wtium. Fern Gazette 11: 141-162. ;
RA OF NORTH AMERICA EDITORIAL COMMITTEE. 1993. Flora of North America North of Mexico,

C

“ouucren, P. K., N. H. HOLMGREN, AND L. C. BARNETT. 1990. Index Herbariorum, Part 1, The
erbaria of the World (8th ed.). New York Botanical Garden, Bronx, NY.
= B L. AND C. F. CHUEY. 1992a. Distribution of the pteridophyte flora of Brooke County,
est Virginia, by district. Laitsch Herbarium Publications of Youngstown State University 5: 3-6.

136 BARTONIA

ISAAC, B. L. AND C. F. CHUEY. 1992b. Pteridophyte flora of Forest County, Pennsylvania by
township. Laitsch Herbarium Publications of Youngstown State University 5: 7-11.

KUCHLER, A. W. 1964. Potential Natural Vegetation of the Conterminous United States. American
Gecpraplgeal Society, New York.

PARKS, J. C. 1989. Distribution of gameotophytic populations of Vittaria and Trichomanes in
Pennsylvania. Rhodora 91: 201-06.

RIESENMAN, J., JR. 1943. History of Northwestern Pennsylvania. Lewis Historical Publishing Co.,
New York.

RHOADS, A. F. AND W. MCK. KLEIN, JR. 1993. The Vascular Flora of Pennsylvania: Annotated
Checklist and Atlas. American Philosophical Society, Philadelphia.

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.

WILD RESOURCE CONSERVATION FUND. 1995. Endangered and Threatened Species of Pennsylvania.
Wild Resource Conservation Fund, Harrisburg, P

Bartonia No. 61, 2002

Joseph Andorfer Ewan (1909-1999)

With the death of Joe Ewan, the world lost a scientist
with enormously broad interests and experience. His
expertise ranged across plant systematics and other
fields of botany, the history of natural history and of
the European exploration of the non-European world,
bibliography, biography, etymology, and the history
of the world’s museums, libraries, and botanical
gardens. Joe had a rare ability to carry out rigorous
research with joy and enthusiasm. Colleagues found
him a charming and stimulating collaborator and
mentor, and he touched countless other careers in his
long life. When Joe’s career began, histories of
science and biographies of scientists were almost
always the leisure-time productions of scientists, and
few contributed as much to the professionalization of
these fields as he did.

Joe was born in Philadelphia on 24 October 1909. At the age of three, he moved to Los
Angeles, where he lived until he was 24. It was here, in the sunny mountains and valleys
of southern California that Joe’s interest in the natural world developed. The Ewan
Collection at the Missouri Botanical Garden includes a series of notebooks from his high
school years, full of detailed observations on the birds, trees, and general natural history of
the area. As a senior at Belmont High School, he was president of the Belsci Science Club,
and contributed to the Belmont Sentinel. It was in high school, too, that Joe discovered the

uman side of natural history. The used bookstores of Los Angeles provided him with his
first rare books, and authors’ dedications and previous owners’ names piqued his curiosity.
He tried to trace some of these names, and was quickly led to view books, not just as passive
‘ontainers of data, but as living links between the reader and a chain of previous owners,
publishers, authors and their teachers, colleagues, libraries, and correspondents. This view
9 books as connecting threads running through the fabric of scholarship was one of the
defining themes of Joe’s career.

Joe’s interests found greater focus when he entered UCLA in 1928. His first professional
publications appeared in southwestern birders’ journals in his freshman year, but, influenced

y plant taxonomist Carl Epling, his primary interest soon turned forever to botany. It was
* UCLA also that Joe met fellow student Nesta Dunn, whom he married in 1935. Nesta
as (and is) a talented biologist who shared Joe’s interests in botany and history. She would
collaborator on much of his life’s work, as well as a stimulating and supportive partner
all the major endeavors of his life.
be the advice of his UCLA mentors, Joe transferred to the University of California at
. keley in 1933, where he finished his A.B. in 1934 and began graduate work. Here, he
te four hours a day, 8 to 12 a.m., on the Flora of California project under Berkeley s

ant but emotionally unstable taxonomist, Willis Lynn Jepson, and began his own

in

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138 BARTONIA

research on the taxonomy of the large, predominantly western genus Delphinium. Like
many others before and after him, Joe found Jepson a rewarding but very difficult person
to work with. An immensely talented botanist, Jepson had broad interests and a splendid
way with words (he also published poetry), and he could be a wonderful teacher and
companion, but he was irrationally suspicious and regularly feuded with most of the people
around him, often working himself up into frightening bouts of vehemence. Joe learned
much about the biology of plants and the practice of taxonomy from Jepson and other
Berkeley botanists (notably H. L. Mason), but he also suffered much from Jepson’s
shortcomings, especially the suspicion Jepson felt when Joe became friendly with other
faculty members and other botanists whom Jepson distrusted. Joe’s work in Berkeley taught
him a great deal, but it ended unhappily, as Joe left in 1937 without completing a graduate
degree.

After leaving Berkeley, Joe took up a position as Instructor at the University of Colorado,
Boulder, where he remained until 1944. Joe continued to publish on the botany of western
North America and Hawaii and the taxonomy of Delphinium. The Colorado years were
marked by field trips and collaborative work with other Rocky Mountain biologists, notably
Frederick Clements and Aven Nelson, and by the births of Joe and Nesta’s three daughters.
Joe also wrote a series of articles on early botanical explorers of Colorado for the journal
Trail and Timberline, work that opened the line of research that Joe is best remembered for
— tracing the history of biological exploration of the New World and the development of
an independent American tradition of academic biology.

Like many botanists of the time, Joe joined the wartime botanical inventories of economic
plants, spending 13 months in 1944 and 1945 inventorying wild stands of Cinchona in the
Andes of southern Colombia, in and around the department of Narifio, for the Foreign
Economic Administration. Conditions were difficult and the long separation from his family
was frustrating, but this work introduced Joe to the biota and culture of tropical America,
which remained a lifelong interest.

The end of the war brought a difficult and uncertain time for the Ewans. The job at
Colorado was no longer available. Through the help of Ray Fosberg, a friend from student
days at UCLA, Joe held temporary positions with the Smithsonian Institution and with the
Bureau of Plant Industry, where he served as acting curator of the herbarium of the U. S.
National Arboretum. In 1947 he accepted a permanent position at Tulane University, where
he remained for the rest of his career.

At Tulane, Joe continued his research in plant systematics, working primarily on South
American ferns, Gesneriaceae and Clusiaceae, and keeping in touch with the growing
involvement of U.S. botanists in tropical America. The experience many scientists gained
in the wartime botanical inventories and the greater availability of grant money stimulated
a great increase in field research in Central and South America. New Orleans was the main
transfer point for travel to the south in those days, and Joe kept in close touch with
scientists working all over the Neotropics. More and more, however, Joe’s interests turned
to the process of scientific exploration itself, its history and the way it is shaped by social
and political trends. f

Academic bibliography was transformed in the 1950s and 1960s by the appre
affordable photo-offset reproduction and high quality color printing, and many gee {
and sets of unpublished drawings, previously rare and in some cases almost eure
became available and affordable for libraries and scientists. Joe was at the center of this
transformation. He edited, and wrote substantial introductions for, many importa!

OBITUARY 139

botanical and zoological reprints for several publishers. Most important among these were
eight titles in Hafner’s photo-offset series Classica Botanica Americana, which made many
major early works on North American botany available to the modern scientist, and William
Bartram: Botanical and Zoological Drawings, 1756 - 1788, published by the American
Philosophical Society in 1968. Joe published his own books as well as other people’s. Chief
among these are Rocky Mountain Naturalists (1950), John Banister and his Natural History of
Virginia (with Nesta Ewan, 1970), Biographical Dictionary of Rocky Mountain Naturalists
(with Nesta Ewan, 1981), and a still-unpublished biography of Benjamin Smith Barton, also
a collaboration with Nesta. In the end, his bibliography grew to include more than 400
books, articles, and reviews (a complete bibliography is provided by Dorr and Holland in
an issue of The Archives of Natural History dedicated to Joe).

Throughout the Tulane years, Joe and Nesta traveled widely, attending meetings, visiting
libraries and archives, and meeting new people everywhere. A Guggenheim Fellowship
allowed them to spend a year at the British Museum (Natural History) in 1954-1955, and
summers teaching at Swarthmore and the University of Hawaii further broadened their
experience and contacts. Joe valued a "sticky mind” — one that retained facts and recalled
them at appropriate moments — and his was one of the stickiest. Interesting observations,
significant associations, and odd details that shed light on scientists and their work were a
joy to Joe, and he displayed them, admired them, and traded them with the zeal of a
diamond merchant in a melodrama. He met and corresponded with numerous biologists,
archivists, librarians, historians, and book dealers, and many whose interests and work
included the history of biology found that Joe had become a unique and important part of
their lives. His charm and enthusiasm drew many people into the field; his encouragement
stimulated many casual enthusiasts to begin working systematically; and his willingness to
share his incredible knowledge of the field aided the work of many a professional. Joe
received awards from a suitably diverse array of academic organizations, the most notable
being the Eloise Payne Luquer Medal of the Garden Club of America, the Founder’s Medal
of the Society for the History of Natural History, and the Henry Shaw Medal of the
Missouri Botanical Garden. He was named Ida Richardson Professor of Botany at Tulane
in 1972 and Fellow of the Linnaean Society (London) in 1976, and received honorary Sc. D.
degrees from the College of William and Mary in 1972 and Tulane in 1980.

Joe continued to collect books, and his library grew into a unique collection of great
Significance. Most important is the huge collection of early travel narratives and reports
tom all parts of the non-European world but especially tropical America, invaluable alike
for tracing the provenance of early collections and recording the impressions that early
vestern visitors formed of non-European’ cultures and environments. The collection of
biographies, textbooks, and floras, both historical and modern, are also notable. The library
's given particular significance and charm by numerous association volumes and annotated
working copies of past experts. A stroll through the collection turns up a seemingly endless
Series of unique books and papers: Henry Hurd Rusby’s copy of Bates’ The Naturalist in the
ver Amazons, battered and water-damaged from crossing Brazil and Bolivia with Rusby in
Pe 886, which Joe bought for $1.00 in 1928; Richard Spruce’s copies of Boussingault’s

‘ae Cientificos a los Andes Ecuatoriales and Markham’s Expeditions into the Valley of the
Amazons, 1539, 1540, 1639, both copiously annotated by Spruce with comments and
“rréctions based on his nineteenth-century travels in the same areas; the copy of Edgar

derson’s Plants, Man, and Life that Anderson marked up when planning a revised edition

"was never written; a set of student notes from John Lindley’s Systematical (sic) Botany

140 BARTONIA

class at University College, London, written by Lindley’s nephew, who took the class in
1845-1846; the manuscript of Carl Epling’s massive Flora of Northern Idaho, which Joe and
Nesta tried, unsuccessfully, to find a publisher for in the depths of the Depression; and many
others. For Joe, each volume carried some of the personality of its author and former
owners. He delighted in sharing these treasures with his colleagues — and anyone who was
interested was a colleague. Casual conversations over lunch often turned into explorations
of the stacks, as Joe seldom hesitated to down his lunch in mid-sandwich to answer a
question, check a fact, or acquaint his fortunate companions with a useful or interesting
reference that they had not seen before. “Book openeth book,” Joe was fond of saying, and
by one o’clock the table was often stacked with volumes, every one of them a treasured old
friend to Joe and an enticing new acquaintance to his companions.

Almost as important as the Ewan library are the Ewan papers. Joe maintained — and
preserved — a correspondence with numerous botanists and historians, and he preserved
masses of ephemeral material — articles in newspapers and newsletters, old road maps
(valuable for finding field sites obscured by the renaming or relocation of roads and towns),
photographs, handwriting samples, book jackets, and pamphlets and flyers of all kinds. Here
also are notes and drafts of unfinished projects, especially the massive “Andes and Amazon,’
a huge file that Nesta and Joe assembled on scientific explorers of tropical South America
that was intended to become a big brother to Rocky Mountain Naturalists, but is available
only as a massive and well-organized file of notes.

Joe retired from Tulane in 1977, but he continued to be active in research and teaching.
He served as visiting professor at the University of Oregon in 1978 and 1981 and Ohio State
University in the spring of 1982, and was Regent’s Fellow at the Smithsonian Institution in
1984-85. In 1986, Joe and Nesta sold their library and papers to the Missouri Botanical
Garden. They followed the collection to St. Louis, where they continued to work with and
add to it. Here at a large research institution (the Garden had a permanent staff of more
than 40 research botanists throughout this period), the Ewans were surrounded by people
who could appreciate their unique knowledge and expertise, and the constant stream ©
visiting botanists coming to use the herbarium and library, work with the staff botanists, and
attend the annual Systematics Symposium, brought a volume of personal contact with
colleagues that they had not known since New Orleans ceased to be a major transfer point
for travel to the south. Joe and Nesta quickly became valued members of the Garden
community, and lunch with the Ewans and their collection (then housed in the ornate 1859
Museum Building) was a delight that brought together a diverse group of botanists,
historians, and bibliophiles who still meet regularly to share their views on plants and the
scientists who study them.

Joe suffered a cerebellar stroke in December of 1996. His intellect and memory wer

unaffected by the stroke, but his motor skills were seriously impaired, and he tired -
oved to

d Dick

effort made to save him in case of illness, and told people, with perfect good humor and i
perfect earnestness, that he hoped he wouldn’t live much longer. He came down witht
flu on Thanksgiving Day, 1999. When his overall condition deteriorated about a week later,

OBITUARY 141

he refused hospitalization very forcefully; when asked what should be done if his heart
stopped, he said, tell everybody goodbye. When he died, Nesta was with him, and Kathleen
and Dick were nearby.

Joe was a charming companion and valued resource to numerous friends and correspon-
dents. He loved history as a vista of the myriad ideas and viewpoints on the natural world
that have been held at different times and places, and of the virtues and foibles of the people
who have held them. He loved books as windows into other times and other minds. and
most of all he loved sharing his discoveries with anyone who would truly appreciate shtick
With joy and enthusiasm he brought together biologists, historians, librarians, archivists, and
bibliophiles, starting productive collaborations and enduring friendships among people who
would never have become acquainted with one another — and with one another’s viewpoints
and fields of study — without Joe. The Ewan library and archive at the Missouri Botanical
Garden are unique treasures, and we who knew Joe can never see the library and papers
without seeing him, flitting from shelf to shelf, his face alight with pleasure. “Book openeth
book,” but scholar openeth scholar, too, and Joe’s greatest legacy is the ever-expanding array
of people whose minds have been broadened by contact with him, who have gone on to pass
abit of his understanding and enthusiasm to other colleagues.

Ada Nesta Ewan (1908-2000)

Since these words were written,
Nesta Ewan has also died. She was
_ born Ada Nesta Dunn in northern
Saskatchewan on 8 November 1908.
She grew up in Saskatoon and Calga-
ry, then the family moved to Los
Angeles, where she attended UCLA.
Like Joe, she was strongly influ-
enced by Carl Epling (she not only
studied with him, but frequently
babysat for his children). Nesta met
Joe in botany classes at UCLA and
they were married when she joined
m him in Berkeley after finishing her
masters’ degree. In the early decades
of their marriage Nesta worked as a
librarian, as well as raising their
ghters and collaborating with Joe on research projects. Later, Nesta left her library
devoted her full time to research and bibliography, collaborating with Joe on many
- Nesta lacked Joe’s flamboyant exterior but she shared his broad interests, high
Or academic work, and willingness to offer her time and expertise to help other
Life alone in the nursing home held few rewards for Nesta. When complications
i... biotics Prevented effective treatment of a serious infection, she requested that the
“ation be discontinued and said goodbye to her family. Joe and Nesta were a united

three dau
J0b and
Projects

Wor kers.
ftom anti

142

BARTONIA

couple and an effective team who will be remembered with respect and affection by

hundreds of colleagues.

ALAN T. WHITTEMORE
Missouri Botanical Garden

Current address:

U.S. National Arboretum
3501 New York Avenue NE
Washington, DC 20002-1958

Bartonia No. 61, 2002

Elizabeth Miner Woodford
(22 May 1916-26 November 1999)

The death of Elizabeth “Betty” Woodford of Medford
Township, New Jersey, has left a great void in my
heart and in the hearts of many others whom she also
inspired and whose lives she enriched. Her death at
age 83 at Virtua Health-West Jersey Hospital in Berlin
resulted from complications related to Alzheimer’s
disease. She is survived by her two children, Richard
Wurster of Topanga, California, and Jeanne Woodford
of Medford Township; a brother, Robert Miner of
Rancocas; a sister, Julia Grayham of Edgewater Park;
four grandchildren; and two great-grandchildren.

Born on a Burlington County farm on Bustleton
Road in Florence Township, New Jersey, Betty
moved to Delanco at age four and in 1933 was a
graduate of nearby Palmyra High School. Deeply
involved in the outdoors, she nurtured her botanical
interests by completing a three-year course at the
Arboretum School of the Barnes Foundation in Merion, Pennsylvania, in 1956. She also
studied at the Morris Arboretum in Philadelphia. In the late 1950s, Betty became a member
of the Philadelphia Botanical Club and, much to her delight, was selected to honorary
membership in March 1994. It was in 1957 when she and her husband Jim built their home
ona lake in a 184-acre parcel of pinelands in Medford Township. On a portion of the land,
they developed Cedar Run Wildlife Refuge to nurse back to health injured and orphaned
animals with the intention of releasing them back to the wilds. Gradually the refuge added
40 environmental education component where thousands of local children as well as adults
received their introduction to the beauty concealed in an ecosystem of scrub pines. Betty
steadily developed an extensive library on birds, insects, mammals, herptiles, and natural

story that became the envy of local naturalists, who were graciously encouraged to use it.
Both Woodfords became licensed bird banders, enabling them not only to attract more
‘upporters to the refuge but also to make substantial contributions to our knowledge of
‘gional bird life. At present, the highly successful work of the refuge and education center
. being continued under the direction of their daughter, Jeanne.

In the 1960s and 1970s, many residents statewide who did not know Betty Woodford by
= knew her simply as “the Pine Barrens Lady.” Her reputation as a champion of
ata protection and preservation and as a premier naturalist was hard earned, well

wand widely known. World renowned wildlife photographer Leonard Lee Rue Il,
* Close friend who influenced her photography and at times accompanied her in the field,
Praised Betty for her broad botanical knowledge. Much to the delight of pine barren scholars
= enthusiasts, she lobbied for the republication of two classics long out of print and hard
° obtain. She persuaded Dover Publications to reissue in 1970 John W. Harshberger’s The

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144 BARTONIA

Vegetation of the New Jersey Pine-Barrens (1916) and Quarterman Publications to reissue in
1973 Witmer Stone’s The Plants of Southern New Jersey (1911). She wrote the foreword to the
Stone reprint. In a book called Medford Pioneering Township, published in 1975, she
authored a chapter titled “In Fields and Woodlands.” Also the same year, the Medford
Township Environmental Commission published her illustrated pamphlet, A Home in the
Pines, focusing on landscaping with native plants to protect this fragile ecosystem. For years,
Betty wrote a weekly nature column that appeared as “Wild and Free” in the Burlington
County Times and later as “Ways of the Wild” in The Central Record of Medford. These
articles reveal her broad knowledge of the regional flora and fauna and her understanding
of conservation and environmental issues.

I met Betty in 1968 on a field trip she led into the heart of Wharton State Forest. The
following year I was a participant in her spring evening course on the Pine Barrens of New
Jersey, offered at Lenape Regional High School. It was the beginning of a long friendship,
forged while engaged in mutual exploration and photographic documentation of the flora
of the barrens. On excursions in forest and field, no nuance escaped Betty’s detection. From
her I learned to see afresh, and her influence on my photographic technique, particularly in
composition, was great.

Betty Woodford’s ethic emanated from an innate desire to protect all things wild. In
cherishing nature and admonishing all whose lives she touched to preserve it, she lived
Thoreau’s dictum, “in wildness is the preservation of the world.”

TED GORDON
31 Burrs Mill Road
Southampton, NJ 08088

Bartonia No. 61, 2002

The Schuyler Herbarium Internship

In the spring of 2000, the Philadelphia Botanical Club initiated a campaign to raise
$100,000 for an internship in the herbarium of The Academy of Natural Sciences. The
internship was named for Academy botanist Alfred E. (Ernie) Schuyler, and announced at
his retirement party given by the club at the Morris Arboretum on 27 April 2000. Dr.
Schuyler, who served as a curator in the Academy’s Botany Department from 1 September
1962 to 31 July 2000, is now Curator Emeritus of Botany at the Academy. He edited the
club’s journal Bartonia from 1970 to 1999.

The intern will be expected to perform work related to the care and maintenance of the
Academy’s Local Herbarium/Herbarium of the Philadelphia Botanical Club, which covers
counties in Pennsylvania, New Jersey, Delaware, and Maryland that are within 50 miles of
Philadelphia. The work includes: (1) routine herbarium chores such as mounting specimens,
filing specimens, and identifying specimens, (2) arranging loans for botanical researchers at
other institutions, (3) working with Academy staff to computerize selected portions of the
herbarium, (4) field exploration to update our holdings of plants growing in the local area,
(5) conducting research on problematic plant groups, and (6) participation in botanical club
meetings, field trips, and projects.

Initially this is intended to be a one-year part-time position for college graduates with
backgrounds in botany, and who want to pursue careers dependent on knowledge of botany.
We envision this will ultimately become a full-time position. The work will provide useful
experience for those who want to go on to graduate school, continue with museum work,
or do environmental work for government agencies or private consulting firms.

The Philadelphia Botanical Club has had a close relationship with the Local Herbari-
um/Herbarium of the Philadelphia Botanical Club since its founding in 1891. Goals of the
club and the Academy’s Botany Department are the same in many respects. The Schuyler
Herbarium Internship will enable both organizations to work toward their mutual goals
more effectively.

Thanks to the donors in the gift categories listed below, over $30,000 was contributed to
the Schuyler Herbarium Internship Fund between its inception in April 2000 and 15
December 2000 (over 30% of our goal of $100,000). Many contributions have been received
since then. To make donations, checks should be payable to the Philadelphia Botanical
Club and mailed to: Schuyler Herbarium Internship Fund, The Academy of Natural
Sciences, 1900 Benjamin Franklin Parkway, Philadelphia PA 19103-1195.

TED GORDON

Chairman
Schuyler Herbarium Internship Fund Committee

145

146

BARTONIA

Contributors to the Schuyler Herbarium Internship Fund
Prior to 15 December 2000

Gentiana Contributor ($10,000 or more)
Jeffersonia Contributor ($5,000-$9,999)
Helonias Contributor ($2,500-$4,999)
Kalmia Contributor ($1,000-$2,499)
Mublenbergia Contributor ($500-$999)
Bartonia Contributor ($100-$499)
Schizaea Contributor ($25-$99)

Gentiana Contributors
Robert J. Holt
Elizabeth & William McLean

Kalmia Contributors
Ann F. Rhoads
Alfred E. Schuyler

Muhlenbergia Contributors
Julia W. Frick
William H. Roberts

Bartonia Contributors
Joseph Arsenault

Michael Bowell

Elizabeth B. Farley

Patricia & Ted Gordon

Val Kolaga & Nick Horvath
David Lauer

Cecily Littleton

Norma Milner

Ruth Patrick

Elise J. Payne

Patricia Quigley

Clare E. Rodgers

William & Yvonne Schuyler
Ronald L. Stuckey

Susan P. Treadway

Howard P. Wood

Schizaea Contributors
Dorrell Biddle
Walter F. Bien

Joan Brinton

Robert Brotherson
Bud Cook

Steven Clemants
Linda DeCastro

Jill Dodds

Janet Evans

Jessie Farrell

Kasia Fogarasi
Catharine P. Fussell
Elinor I. Goff
Norma F. Good
Janet & John Gyer
Thomas B. Halliwell
Gay Kimelman
Milton Laden

Mary Leck

Valencia Libby/Jay Danzenbaker
Richard Lighty

Keith Maurice/Judy Harnly
Robert McCombe
Richard McCourt
Andrea McFadden
Robert C. Meyer
John & Carol Mickel
Ann M. Mickle

John C. O’Herron, II
William Olson
Margaret A. Rue
Helen Stailey
Richard Stalter

Sue Thompson

John Vollmer

Bartonia No. 61, 2002—News and Notes

Amaranthus pumilus Raf. (Seabeach Amaranth,
Amaranthaceae) Rediscovered in Sussex County, Delaware

In August of 2000, Amaranthus pumilus was rediscovered in Sussex Co., Delaware after
125 years without a sighting. It was first collected in Delaware in 1875 by Albert Commons
(10 September 1875, A. Commons, s.n., “seabeach, Baltimore Hundred, Delaware,” PH).

Amaranthus pumilus was federally listed as threatened by the U.S. Fish and Wildlife
Service in 1993. Historically, this species was known from Massachusetts south to South
Carolina (Weakley et al. 1996). Amaranthus pumilus was reported as rediscovered at
Assateague Island National Seashore, Worcester County, Maryland in 1998 (Ramsey 2000).
Prior to rediscovery on Assateague Island and in Sussex County, A. pumilus was extant on
Long Island, New York, and in North Carolina and South Carolina.

Lisa Marie Kendall of the Delaware Natural Heritage Program, Division of Fish and
Wildlife, Delaware Department of Natural Resources discovered the first plants on 7 August
2000. Subsequent surveys revealed a total of 41 individuals scattered over 22 kilometers of
Atlantic shoreline. All plants found are within the boundaries of Delaware Seashore and
Fenwick Island State Parks. The largest number of plants (28) was found within a 1.5-km
stretch of shoreline near the swimming beach at Delaware Seashore State Park. This section
of beach is the only area where A. pumilus was found that is off-limits to vehicular traffic.
This area provides the best habitat for the long-term survival of A. pumilus.

Individual plants were found growing on relatively open sand near the base of the
primary foredune. Though overall plant cover was quite sparse, common associates include
Ammophila breviligulata, Cakile edentula (most prevalent), Cenchrus tribuloides, Chamaesyce
polygonifolia, Salsola kali, and Triplasis purpurea. Plants of A. pumilus were often associated
with a wrack line, which appeared to be composed primarily of broken and decomposing
canes of Phragmites australis. Chris Lea, ecologist at Assateague Island National Seashore,
speculates that wrack lines may be acting as a seed trap for A. pumilus (pers. comm.).

n after the rediscovery of A. pumilus, seeds were collected from August to September
and sent to the Mt. Cuba Center for the Study of Piedmont Flora in Greenville, Delaware.
Rick Lewandowski, Director of the center and his staff will attempt to germinate and rear
plants of this annual species for planting next season in the areas where seeds were collected.

In addition to seeds, fresh leaf material was also collected from several plants and sent to
Dr. Kim Hunter at Salisbury State University where comparative DNA studies will be done.
The rediscovery of A. pumilus on the Delmarva Peninsula (Worcester County, Maryland and
Sussex County, Delaware) could be the result of propagules being deposited from southern
Populations (currents run south to north), or from the unearthing of dormant soil seed

- DNA studies may help to answer these questions. ;

In addition to the rediscovery of A. pumilus in Sussex County, another exciting find was
made by Frank Hudson, ecologist at Assateague Island National Seashore. Frank, along with
— from Assateague Island were visiting the Sussex County population and found

Ygonum glaucum growing near individuals of A. pumilus. P. glaucum is considered to be

Blobally rare by The Nature Conservancy and was last collected in Delaware in 1936 (31

147

148 BARTONIA

August 1936, E. Larson, 1216, “sandy beach, 3 miles north of Broadkill Beach,” Sussex
County, Delaware, PH)

The Delaware Natural Heritage Program performs annual surveys for listed and candidate
plant species in Delaware using funds from the U.S. Fish and Wildlife Service.

WILLIAM A. MCAVOY

Delaware Natural Heritage Program
Delaware Department of Natural Resources
Division of Fish and Wildlife

4876 Haypoint Landing Road

Smyrna, DE 19977

LITERATURE CITED

WEAKLEY, A., M. BUCHER AND N. MURDOCK. 1996. Recovery Plan for Seabeach Amaranth
(Amaranthus pumilus Raf.). U.S. Fish and Wildlife Service, Atlanta, GA.

RAMSEY, S., W. TYNDALL AND C. LEA. 2000. Scientific note: the federally threatened Amaranthus
pumilus Raf. (seabeach amaranth, Amaranthaceae) rediscovered on Assateague Island after 31
years. Castanea 65: 165-167.

Bartonia No. 61, 2002—News and Notes

Pine-Barren Golden-Heather (Hudsonia ericoides L.)
Reported for the First Time in Maryland

On 24 May 2000, while searching for rare plants in Anne Arundel County, Maryland, I
made a significant discovery along a Baltimore Gas & Electric Company’s transmission line
right-of-way off Marley Station Boulevard. I chanced upon a new woody vascular plant for
Maryland, the pine-barren golden-heather (Hudsonia ericoides L.). I returned to the site on
3 June to conduct a more extensive search for the heather. I found additional clumps of 11,
2, and 15 plants as I progressed farther along a 230-foot stretch of the right-of-way.

Vascular plants growing at or near these clumps of golden-heather included blackberry
(Rubus sp.), sheep-sorrel (Rumex acetosella), matting rosette grass (Dichanthelium meridionale),
broom panic grass (D. scoparium), glaucous greenbrier (Smilax glauca), frostwort (Helianthe-
mum sp.), broomsedge (Andropogon virginicus), Virginia pine (Pinus virginiana), creeping
bush-clover (Lespedeza repens), a tall hairy bush-clover (Lespedeza sp.), sassafras (Sassafras
albidum), old-field toadflax (Linaria canadensis), southern red oak (Quercus falcata), sweet
vernal grass (Anthoxanthum odoratum), white thoroughwort (Eupatorium album), dwarf
dandelion (Krigia virginica), horseweed (Erigeron canadensis), poverty wild oat grass
(Danthonia spicata), forked rush (Juncus dichotomus), and wavy hair grass (Deschampsia
flexuosa). The tree species in both plots were merely scrub-like due to periodic maintenance
cutting. The soil in the area is Evesboro loamy sand, a very sandy upland soil (Kirby and
Matthews 1973). The forest on both sides of the right-of-way is dominated by Virginia pine
and Virginia pine-mixed oak.

Voucher specimens (Sipple 2032, 2033, and 2034) of the pine-barrens golden-heather have
been submitted to herbaria at Delaware State University, the Philadelphia Academy of
Natural Sciences, and the Anne Arundel County Community College.

According to the Maryland Natural Heritage Program (Lynn Davidson, pers. comm.,
2000) and the Delaware Natural Heritage Program (Bill McAvoy, pers. comm., 2000), this
species has never been reported from Maryland and is considered extremely rare in
Delaware. It was not listed by Steiber (1967, 1971) in his Anne Arundel County flora, and
Brown and Brown (1972) state that it “has never been reported in Maryland, but has been
found in Delaware and adjacent Virginia.” Tatnall (1946) cites only Delaware collections.
Fernald (1950) mentions its presence in Delaware but suggests that reports from Virginia
need verification. Although McAtee (1918) reported it from the Norfolk-Virginia Beach area,
ts not listed for Virginia by Harvill et al. (1992). Gleason & Cronquist (1991) cite only
Delaware for the Delmarva Peninsula. The Natural Resources Conservation Service’s

S database (http://plants.usda.gov/ plants/) lists this species from all of the New
England States, as well as New York, New Jersey, Delaware, and South Carolina, but not
from Maryland or Virginia.

On my two earlier visits and a 10 June 2000 visit to the site, I searched the Virginia pine
and Virginia pine-mixed oak forests on both sides of the right-of-way hoping to find some
°penings in the pines supporting the heather, but found none. The nature of the forest in
nt areas astride the right-of-way suggests that they were more open in the past, and xeric
oe knolls do exist. Given the topography in the area and the sandy soils, I suspect that

€ heather once occurred here independent of the right-of-way.

149

150 BARTONIA

A more detailed account of this discovery and commentary on the significance of rights-
of-way, particularly if they are maintained in natural vegetation by cutting or burning, can
be found in The Maryland Naturalist (Sipple, in press).

William S. Sipple
512 Red Bluff Court
Millersville, MD 21108

ACKNOWLEDGMENTS

The author appreciates the cooperation of Bill McAvoy (Delaware Natural Heritage
Program), Kathy McCarthy, and Lynn Davidson (Maryland Natural Heritage Program)
in providing information on the known distribution of the pine-barren golden-heather in
Delaware and Maryland. Bill McAvoy, Charlie Davis, and two unidentified reviewers
also commented on the larger manuscript cited above from which this note was derived.
I appreciate their input. I likewise appreciate cooperation of Joel Snodgrass, the editor of
The Maryland Naturalist, in allowing this shorter note on the pine-barren golden-heather
to appear in Bartonia.

LITERATURE CITED
BROWN, R.G. AND MLL. BROWN. 1972. Woody Plants of Maryland. Port City Press, Inc., Balti

more.

FERNALD, M.L. 1950. Gray’s Manual of Botany. Van Nostrand Reinhold Company, New York.

GLEASON, H.A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States
and Adjacent Canada. New York Botanical Garden, New York.

HARVILL, A.M. JR., T.R. BRADLEY, C.E. STEVENS, T.F. WIEBOLDT, D.M.E. WARE, D.W. OGLE,
G.W. RAMSEY, AND G.P. FLEMING. 1992. Atlas of the Virginia Flora. Virginia Botanical
Associates, Farmville, VA.

Kirby, R.M. AND E.D. MATTHEWS. 1973. Soil Survey of Anne Arundel County, Maryland. U.S.
Department of Agriculture, Soil Conservation Service, Washington, DC.

MCATEE, W.L. 1918. A sketch of the natural history of the District of Columbia. Bulletin of the
Biological Society of Washington 1: 1-142.

SIPPLE, W. S. (In press.) Pine-barren golden-heather (Cistaceae: Hudsonia ericoides L.): range
extension and a new woody plant for Maryland. Maryland Naturalist.

STEIBER, M.T. 1967. An annotated checklist of the vascular flora of Anne Arundel County,
Maryland. Master’s Thesis, Catholic University of America, Washington, DC. 70 pp-

STEIBER, M.T. 1971. The vascular flora of Anne Arundel County, Maryland: an annotated
checklist. Castanea 36: 263-312.

TATNALL, R.R. 1946. Flora of Delaware and the Eastern Shore: An Annotated List of the Ferns and
Flowering Plants of the Peninsula of Delaware, Maryland and Virginia. The Society of Natural
History of Delaware, Wilmington.

Bartonia No. 61, 2002—News and Notes

What’s New at the Department of Botany,
Academy of Natural Sciences

Many changes are underway in the Academy’s Department of Botany (PH), involving
both personnel and physical plant. Bartonia editor Dr. Roger Latham has invited us to
provide you with a brief overview of developments. We are delighted to share our
enthusiasm for plants and the plant collection here with you and we look forward to
interacting with the readers of the journal over the coming years.

I will begin by telling you a bit about myself, Lucinda
McDade. As the newest staff member in the Department of
Botany, I am still finding my way around and have a great deal
to learn. I arrived here at the beginning of the year from the
University of Arizona where I was curator of the herbarium and
associate professor in the Department of Ecology and Evolution-
ary Biology and the Department of Plant Sciences. My research
interests have focused on three areas: systematics of the family
Acanthaceae, plant reproductive biology, and hybridization as

a mode of evolution and a problem (or not) for phylogenetics.
Lucinda McDade Most recently, I have been involved in some rewarding collabo-
rative efforts to straighten out our understanding of relationships
among the major lineages of Acanthaceae (reprints available on request!). Since my graduate
student days, I’ve done a great deal of field work in the New World tropics. More recently,
. begun to work in Africa as well; I spent three very productive months in South Africa
ast year,

My focus on the tropics perhaps is a natural result of my having grown up in South
Florida. I did my undergraduate work at Tulane University and then went to Duke for my
Ph.D. After completing my doctorate, I had a productive year as a postdoctoral associate
at the Smithsonian Tropical Research Institute in Panama. Before I moved to the University
of Ar izona, I worked for the Organization for Tropical Studies. O.T.S. is a consortium of
universities that has field stations in Costa Rica (some readers may have heard of La Selva
Biological Station) and offers field courses in Costa Rica, Brazil and Peru. I was in charge
of all of O.T.S.’s educational programs and, although my main location was at the North
American office of O.T.S. at Duke in North Carolina, I spent a great deal of time in Costa
Rica. Accepting a faculty position at the University of Arizona represented a move to more
research and teaching, both of which I missed in my position at O.T.S. Moving here
‘epresents another change in professional focus although I intend to continue to be active
in training graduate students; an appointment at U. Penn will make this feasible. Accepting
the position here of course brings many challenges and I look forward to working with
many of you toward meeting these challenges.

I share curatorial responsibility here with Dr. Richard McCourt who writes:

“With the recent arrival of Dr. Lucinda McDade (Associate Curator and Chair) and Dr. James

— (Collection Manager) I have become an old-timer in the department, although I’ve been

che only four years. I moved here from DePaul University in Chicago where I was an

Associate Professor of Biological Sciences. My background and research are in phycology, the

151

152 BARTONIA

study of algae. Although the collections of algae here are small
in comparison to those of land plants, the Academy has a long
record of research in algae. The Academy has been the home of
Dr. Ruth Patrick and Dr. Charles Reimer, experts on diatom
systematics and ecology, and Dr. Francis Drouet, a botany
curator who was a world authority on blue-green algae. |
worked on brown seaweeds in the Gulf of California while
studying for my Ph.D. at the University of Arizona. Later I
moved into the systematics of conjugating green algae (Spirogyra
and relatives) and Characeae (Chara and Nitella), which occur
in the Philadelphia region and worldwide. These green algae
are among the closest living algal relatives to primitive land
plants, such as liverworts and mosses. My laboratory has many
cultures of these algae, which I observe and use in my molecular

> aT systematic research. I extract DNA and sequence genes from
Rick McCourt green algae to decipher the evolutionary tree of green plants.
Ultimately, my research relates to the origin of land plants and

the genetic novelties that accompanied their transition from a wet to a dry habitat some 400 to 500

million years ago.

“Working at the Academy also satisfies my avid interest in historical botany, specifically the
early exploration of the western United States by Lewis and Clark and others. I’m eager to
develop public outreach programs for the upcoming bicentennial celebration (2003-2006) of the
Lewis and Clark expedition. This pioneering trip began in Philadelphia, where Meriwether
Lewis trained with the preeminent botanists of his day. In a sense it also ended here, when the
scientific specimens came back to the city with the explorers. Lewis and Clark's specimens are
but the tip of the iceberg in terms of historically important collections here, and we hope to use
the bicentennial as a springboard to increase awareness of the contributions of botany to our
national history. I look forward to working with the Philadelphia Botanical Club on various
projects, not least among them the development of the Schuyler Internship program for the
enhanced care of our collections.”

Dr. James Macklin is the new collection manager in Botany. He comes to us from
Canada after receiving his Ph.D. in plant systematics from the University of Western
Ontario in London, Ontario.
Currently, James’s research focus
es on the patterns of variation in
Crataegus (Maloideae, Rosaceae).
He is interested in the interaction
of native and invasive species 19

re

become involved in teaching and
outreach projects. He is also a
strong proponent of bioin-
formatics and looks forward to
creating a database that will vastly
James Macklin, James Lendemer, and Rachel Wilson improve access to information 10
the botany collections.

Dr. Rachel Wilson has been a research collaborator in Rick McCourt’s lab since January;
when she began an eight-month sabbatical leave from teaching to investigate cellular,
biochemical, and reproductive features of the alga Spirogyra and a closely related species

WHAT’S NEW AT PH 153

Sirogonium. Her work in the Botany Department began several summers ago, when she
spent a day or two each week in Rick’s lab learning DNA purification and amplification
techniques. She plans to continue her current project at the Academy part-time after
returning to teaching in September at Philadelphia University, where she is an associate
professor of biology.

Sarah Corbett has been the Flora of Pennsylvania Intern since June 2000, after graduating
magna cum laude with a B.S. in biology from Valdosta State University in Georgia. Half
of the internship is herbarium work at the Academy and the other half
is a research project on the Pennsylvania flora supervised by Dr. Ann
Rhoads and Dr. Tim Block at the Morris Arboretum. Sarah’s research
is a floristic inventory of a recent addition to French Creek State Park
near the Pine Swamp Natural Area. She has collected about 250 species
and mapped the plant associations of the tract, which harbors at least
six rare plant species. Sarah writes:
we ‘an “I have come to know the Academy herbarium as few do. My duties bere
Sarah Corbett have included preparing specimen labels, mounting, accessioning specimens,

processing loans, filling information requests, and many other routine herb-
artum activities. In addition, I’ve been an extra set of hands for moving cabinets, specimens, etc.,
during recent drastic changes in the herbarium. The internship is a year-long appointment, and

I will be heading to graduate school in the fall to study paleobotany at the University of

Florida.”

We are very fortunate to have the faithful assistance of one of the Academy’s most active
volunteers, James Lendemer. James is a junior in high school but he has already been
working here for more than three years. James writes:

“When I first began my work in the Department of Botany in early 1998 under the guidance of

Earle Spamer I set out on a total reorganization of the paleobotany collection. The reorganiza-

tion resulted in the recovery of numerous type specimens of fossil plants as well as the recovery

of an interesting collection of fossil plants identified by Leo Lesquereux and included in his work

The Coal Flora {see article in this issue on page 55]. Since that time I have taken up the task of

electronically cataloging the type collection of extant lichens, bryophytes, and fungi. This work

has also led me to actively search and retrieve the many unrecognized type specimens in the

general herbarium. Currently, I am preparing a catalog of the type lichens at PH for eventual

publication,”
_ We are happy to say that Curator Emeritus A. E. (Ernie) Schuyler continues to be active
in the department and in teaching. He is pursuing his research interests in the systematics
and ecology of rare plants, relationships between plant diversity and environmental quality,
and the history of botanical exploration in North America. He periodically teaches
“versity, college, and adult education courses in plant systematics, ecology, and
“AVironmental issues.

Finally, we would like to bring you up-to-date on physical changes that are underway at

the herbarium. The new Lewis and Clark Types Room, across the hall from the previous

room, is essentially complete. It has been outfitted with its own HVAC system, a
“ate-of-the-art fire suppression system, and special dust-free ceiling tiles. This room will
Suse all of our type specimens and other specimens of special historical significance.
Cabinets designed specifically to house the Lewis and Clark collection are on order. We
© acquired a large chest freezer and are hard at work freezing portions of the herbarium
where bugs have been active. Five days at -20°C has been shown to kill these pests. Once
‘Pecimens are freed of bugs we will hold as many of them as we can on the fifth floor in
herbarium cases with tightly sealing, gasketed doors. We have begun a program of baiting

154 BARTONIA

and trapping the beetles that are the culprits. This should enable us to diagnose the problem
and make short-term plans to reduce further damage as much as possible. The longer-term,
and only real, solution is to replace all of the cabinetry with herbarium cases with sealed
doors. We anticipate submitting a proposal to the National Science Foundation in
September for this project.

We have made some changes to the fifth floor with the goal of providing better space for
working with plant specimens. There is a new extensive work area in the alcove, with
windows and lots of space for spreading out and studying specimens. John Kunsman from
The Nature Conservancy has already used it and pronounced it most excellent. We have
also reclaimed one of the areas hidden away behind the compactors for use as a reading

room.

The fifth floor will shortly have two labs up and functioning — one dry lab for
microscope and specimen work and one wet lab for molecular and other work. These will
complement other facilities in the Department of Botany and in the other biodiversity
departments and give us the capacity to accommodate research by ourselves, postdocs, and
students. We anticipate an active and vibrant community of researchers sharing the botany
floors with us. We look forward to showing many of you the changes that are underway!

LUCINDA MCDADE

Associate Curator and Chair

Department of Botany

Academy of Natural Sciences of Philadelphia
June, 2001

Bartonia No. 61, 2002

1997-1999 FIELD TRIPS

1997 Field Trips

2 April: Valley Forge National Historic Park, PA. This was a joint trip with the
Perkiomen Valley Watershed Association on a cold rainy day. Growing on a bank, an
unusually large patch of Epigaea repens, with only a single plant in bloom, had been severely
browsed by deer. This patch was associated with many plants of Hieracium venosum.
Following a path along the Schuykill River from Betzwood Bridge west, we observed the
usual common spring flora including Viola, Glechoma, Oxalis, and Sanicula. Trip leaders: Val
Udell and Ann Newbold.

17 May: Bowman’s Hill Wildflower Preserve, Bucks County, PA. The wildflower preserve
offers a unique opportunity to see a large number of species in one location in a natural
setting. We explored the many trails of the preserve, beginning at the headquarters garden
where we observed Antennaria solitaria, Epigaea repens, Dodecatheon medea, Trillium
cemuum, T. grandiflorum, and T. sessile. On the Parry Trail, we saw Jeffersonia dipbylla,
Hydrophyllum virginianum, Claytonia virginica, Cardamine concatenata, Caulophyllum
thalictroides, Mertensia virginica, Polemonium reptans, Mitella diphylla, and Viola striata. We
also saw the two varieties of Phlox divaricata, var. laphamii with entire corolla lobes and var.
divaricata with notched corolla lobes. On the Azalea Trail, we saw Gaylussacia brachycera,
Viola hirsutula, Anemone quinquefolia, Phlox stolonifera, Saxifraga virginiensis, Packera aurea
(Senecio aureus), Rhododendron periclymenoides, and Cercis canadensis. We also saw very poor
specimens of Galax aphylla. On the Bluebird Trail, Hydrophyllum canadense was seen with

virginianum and Allium tricoccum. The weedy, introduced Ranunculus ficaria was also
observed. The violets on the Violet Path included Viola striata, V. sororia, and V. pubescens.
On the Gentian Trail, we observed Itea virginica, Saxifraga pensylvanica, Ranunculus hispidus
Var. nitidus, Carex prasina, Chelone glabra, Caltha palustris, Iris versicolor, Hypericum pyra-
midatum, Acorus calamus, Carex trichocarpa, Saururus cernuus, Euphorbia purpurea, Stellaria
Pubera, Chrysogonum virginianum, Sedum ternatum, and Meehania cordata. Along the
Harshberger Trail was Exonymus americana, Meehania cordata, Jeffersonia diphylla, Trillium
sessile, T. cuneatum, Dicentra eximia, Cypripedium calceolus, and Uvularia grandiflora. Along
the Medicinal Trail we saw Deparia acrostichoides, Hydrastis canadensis, Moehringta lateriflora,
Cystopteris protusa, Erigenia bulbosa, and Zanthoxylum americanum. The Pocono Laurel Trail
had Pyrularia pubera. The Fern Trail was explored more for spring wildflowers than ferns,
although, a few fern species were observed. These included Cystopteris fragilis and C.
bulbifera, Flowering species included Packera obovata (Senecio obovatus), Brachyelytrum
"rectum, Viola conspersa, V. canadensis, V. palmata, Thalictrum thalictroides, Geranium
robertianum, Dentaria diphylla, and Streptopus roseus. In the bog, we observed Helonias
ta and Chamaecyparis thyoides. The Marsh-marigold Trail had Stylophorum diphyllum,
Collinsia verna, Cypripedium calceolus, Veratrum viride, Symplocarpus foetidus and Mitella
diphylla, Clethra acuminata was observed on the Cornus Bend, and Lupinus perennis was seen
on the Aster Walk. Attendance: 13. Leader: Bill Olson.

155

156 BARTONIA

31 May: Funks Pond Recreational Area north of the Conowingo Dam, Cecil County, MD.
From the Route 1 parking lot, we proceeded down the gravel and woodland trails leading
into the young thickets and forested ravines that dissect this corner of Cecil County. Along
the trails we encountered unusual species such as Carex jamesii (abundant), Veratrum viride,
Triosteum perfoliatum (in a pipeline cut), Hydrastis canadensis, Hydrophyllum virginianum,
H. canadensis, Scutellaria nervosa, and a colony of Lithospermum latifolium in bloom. An
adult and hatchling box turtle, a caerulean warbler, and a fawn were among the fauna seen.
After lunch the party circled Funks Pond, encountering such species as Orobanche uniflora,
Phlox divaricata, Caulophyllum thalictroides, and several escaped goats that were chewing up
the vegetation. We concluded the trip with a long rugged hike up a steep ridge. At the top
we were rewarded by the sight of a large healthy population of Hybanthus concolor in bloom
along with other rarities and unusual species such as Piptochaetum avenaceum, Tradescantia
virginiana, and Cheilanthes lanosa on the dry edge and rocks of the bluff. Attendance: 15.
Leaders: Janet Ebert and Jack Holt.

14 June: Sadsbury Woods, Chester County, PA. This newly established preserve of the
Natural Lands Trust includes one of the largest tracts of continuous forest cover remaining
in western Chester County. Geologically, the site consists of a ridge of Chickies Quartzite
surrounded by gneiss. Small lenses of serpentinite and pegmatite dikes are also present in the
vicinity. Buck Run borders the woods on the south. Sadsbury Woods includes mature red
oak-mixed hardwood forest, palustrine forest along Buck Run, areas of successional forest,
woodland seeps, and a former quarry with successional old-field species. Traces of earlier
habitation including stone walls and an early hand-dug well are present in parts of the
woods. We added at least 59 species to lists compiled during previous visits, with the results
that combined lists included 62 woody taxa (trees, shrubs, and vines), 21 ferns and fern allies,
and 166 herbaceous taxa for a total of 249. A sampling of species on the site includes
Dryopteris carthusiana, Huperzia lucidula, Thelypteris hexagonoptera, Corylis americana, Fagus
grandifolia, Angelica venenosa, Carex glaucodea, C. laxiculmis, Chamaelirium luteum,
Cimicifuga racemosa, Isotrea verticillata, Prenanthes altissima, Rhexia virginica, Trillium
cernuum var. cernuum, and Viola hirsutula. Leader: Ann Rhoads.

22-26 June: Shenandoah National Park, VA. The joint field meeting of the Philadelphia
Botanical Club, the Torrey Botanical Society, and the Northeast Section of the Botanic

Society of America took place in the central district of Shenandoah National Par k. Most
of the participants were housed at historic Skyland Lodge at milepost 41.7 on the Skyline
Drive. Field trips to Hawksbill Mountain, Big Meadows, Millers Head, and Hoover Camp
yielded panoramic vistas, billion-year old granite rocks, a hybrid moss Huperzia appalach we
x lucidula, nine species of orchids including Habenaria (Platanthera) orbiculata, H. viridis,
and Liparis lilitfolia in bloom, and the only known station for Arctostaphylos wva-urst
Virginia. Larry Klotz of Shippensburg University, with assistance from other botanists,
maintained a plant list for all sites, and this was sent to the National Park Service for the
record, Several uncommon plants were found including Aconitum reclinatum, Streptop™
amplexifolius, and Botrychium matricariifolium. Botanist members led groups of participants
on the trails and provided expertise with identification. Chris Ludwig of the Virgin@
Heritage Program led the foray into the 150-acre Big Meadows site, where grasses, sedges,
and ericaceous shrubs dominated the complex of managed, open habitat. Here the -
found Botrychium multifidum, including one specimen with a precocious fertile frond. Eac

1997 FIELD TRIPS 157

evening the group heard the perspectives of guest speakers. Robert Johnsson spoke on the
flora of the upper Potomac Basin and Shenandoah, emphasizing plants that have unusual or
disjunct distribution in the region. Chris Ludwig discussed the rare plants and significant
natural communities of Shenandoah. Tom Blount of the National Park Service spoke about
Shenandoah ecology and gave insight into the historical value and current problems facing
the park. Joan Gottlieb presented “A Portrait of Shenandoah National Park,” a retrospec-
tive focusing on the flora of the site. Attendance: 69. Chairperson for the field

meeting: Joan Gottlieb.

5 July: Bulls Island and Prallsville Mills, Hunterton County, NJ. This was a joint trip with
the Torrey Botanical Society to visit floodplain forest. Among the typical tree species were
box-elder (Acer negundo), silver maple (A. saccharinum), river birch (Betula nigra), green ash
(Fraxinus pennsylvanica), black walnut (Juglans nigra), sweetgum (Liguidambar styraciflua),
sycamore (Platanus occidentalis), and silk-tree (Albizia julibrissin), an invasive. Shrubs included
wild hydrangea (Hydrangea arborescens) and elderberry (Sambucus canadensis). Some of the
more interesting vines were hogpeanut (Amphicarpaea bracteata), Japanese hop (Humulus
japonicus), hop (H. lupulus), everlasting pea (Lathyrus latifolius), moonseed (Menispermum
canadense), climbing false-buckwheat (Polygonum scandens), and bur-cucumber (Sicyos
angulatus). Herbs in bloom included scarlet pimpernel (Anagallis arvensis), dog-fennel
(Anthemis cotula), Chinese mustard (Brassica juncea), black mustard (B. nigra), hemp-nettle
(Galeopsis tetrahit), celandine (Chelidonium majus), Indian strawberry (Duchesnia indica),
wormseed mustard (Erysimum cheiranthoides), common quickweed (Galinsoga quadriradiat. )s
dame’s-rocket (Hesperis matronalis), dwarf St.-John’s-wort (Hypericum mutilum), American
water-willow (Justicia americana), nipplewort (Lapsana communis), motherwort (Leonorus
cardiaca), false-pimpernel (Lindernia dubia), fringed loosestrife (Lysimachia ciliata), monkey-
flower (Mimulus ringens), umbrellawort (Mirabilis nyctaginea), carpetweed (Mollugo
verticillata), forget-me-not (Myosotis scorpioides), catnip (Nepeta cataria), ditch stonecrop
(Penthorum sedoides), plantain (Plantago rugelit), figwort (Scrophularia marilandica), giant
chickweed (Stellaria aquatica), tall meadow-rue (Thalictrum pubescens), round-leaved triodanis
(Triodanis perfoliata), moth mullein (Verbascum blattaria), blue vervain (Verbena hastata),
white vervain (V. urticifolia), and European field pansy (Viola arvensis). Other herbs were
wild chervil (Anthriscus sylvestris), annual wormwood (Artemisia annua), waterweed (Elodea
nuttallir), Virginia stickseed (Hackeli virginiana), bugleweed (Lycopus virginicus), moneywort
(Lysimachia nummularia), long-styled sweet cicely (Osmorhiza longistylis), perilla mint (Perilla
irutescens), plantain (Plantago virginica), curly pondweed (Potamogeton crispus), yellow cress
Rorippa palustris var. hispida), sleepy catchfly (Silene antirrhina), American germander
(Teucrium canadense), rabbit’s-foot clover (Trifolium arvense), wingstem (Verbesina
alternifolia), American speedwell (Veronica americana), and water speedwell (V. anagallis-
“quatica), An interesting sedge was Carex davisii. Among the ferns and fern allies were
“couring-rush (Equisetum hyemale var. affine), Mackay’s brittle fern (Cystopteris fragilis var.
mackayi), purple cliffbrake (Pellaea atropurpurea), and blunt-lobed cliff fern (Woodsia obtusa).
Thanks to William Standaert for compiling the plant list. Attendance: 12. Leader: Patrick
ney,

: July: Bear Swamp West, Downe Township, Cumberland County, NJ. We hiked into
‘he heart of Bear Swamp. There is convincing evidence that what remains of its ancient
rest today has never been cut, probably due to its remoteness and its situation amid

158 BARTONIA

extensive swamps and marshes, although nearby large sections of old-growth were destroyed
by sand mining activities and commercial extraction of lumber earlier in the twentieth
century. Approximately 100 acres of apparently primary forest remain in Bear Swamp West,
along with possibly 200 acres of similar antiquity, composition, and structure in nearby Bear
Swamp East. Red maple (Acer rubrum) is the most abundant tree. In better drained sections
sweetgum (Liguidambar styraciflua) co-dominates. At the lowest elevations, where standing
water is usually present, the co-dominant is blackgum (Nyssa sylvatica). Both these trees
referred to as “gums” attain heights of as much as 120 feet. Sweetbay magnolia (Magnolia
virginiana) occurs as a sizeable tree, with diameters at breast height sometimes exceeding 1.5
feet, according to measurements taken by Club members in previous years. American holly
(Ilex opaca) is conspicuous, and specimens of greater than 60 feet in height are not unusual.
Sweet pepperbush (Clethra alnifolia) is the most abundant shrub, while sphagnum mosses
and cinnamon fern (Osmunda cinnamomea) are conspicuous on the forest floor in the
wettest areas. Mistletoe (Phoradendron leucarpum) is occasionally present, parasitizing only
Nyssa sylvatica in this locality. Attendance: 5. Leader: Stevens Heckscher.

27 July: Whitesbog, Lebanon State Forest, Burlington and Ocean counties, NJ. Near the
village of Whitesbog we examined New Jersey’s only surviving population of Schwalbea
americana (chaffseed). At this northernmost limit of the species’ range, a few fruiting plants
showed damage by herbivory. For more than 20 years the late Louis Hand and the leader
helped preserve this population by select-cutting competing plants and by arranging periodic
mowing. Since 1993, management (including prescribed burning) and monitoring of the site
has been assumed by the Office of Natural Lands Management (N.J. D.E-P.). Growing
nearby was Scleria triglomerata (tall nutrush), with smooth and shining white nutlets, and
the stout culms of Juncus biflorus, with fully-grown reddish-brown capsules. Along the
shrubby border of a reservoir were a few white plumes of Platanthera blephariglottis (white
fringed orchid) associated with the showy orange drumheads of Polygala lutea (orange
milkwort), the pale rose-purple racemes of P. brevifolia (short-leaved milkwort), and the
delicate blue petals of Lobelia nuttallii. The shrubs in flower were Clethra alnifolia (sweet
pepperbush), Rhododendron viscosum (swamp azalea), and Hypericum densiflorum (shrubby
St.-John’s-wort). Still in bud, a lone specimen of Franklinia alatamaha in front of the old
company store in Whitesbog attracted attention. Other non-native species planted by the
late Elizabeth White near her home were noted: Oxydendrum arboreum (sourwood),
Xanthoriza simplicissima (shrub yellowroot), Galax aphylla (wandflower), and Shortia
galacifolia (Oconee-bells). Flowering at the edge of an abandoned blueberry field were two
naturalized heaths, Calluna vulgaris (ling) and Erica tetralix (cross-leaved heather), blending
with the native Rhexia virginica (Virginia meadow-beauty), R. mariana (Mary and meadow:
beauty), Triadenum virginicum (marsh St.-John’s-wort), and Lilium superbum (Turk nd
lily). The surface of nearby Union Pond was dotted with fragrant white water lily including
pink forms, Nymphaea odorata forma rosea. The dull-white, wooly heads of Lachnanthes
caroliniana (redroot) and the bright white flowers of Sagittaria latifolia (broad-leaved
arrowhead) occurred as weeds in cultivated cranberry bogs. Planted to stabilize the dams
surrounding these bogs, Eragrostis curvula (African love-grass) is outcompeting native species.
To the south of Big Tank Reservoir, the leader revealed a previously unreported station “
the endangered Stylisma pickeringii var. pickeringit (Pickering’s morning-glory) in bloom.
Only a single robust specimen, growing in exposed mineral soil devoid of competing plants,
had escaped the onslaught of O.R.V.s. An effort to search difficult swampy tertai® 44

1997 FIELD TRIPS 159

rediscover a giant Chamaecyparis thyoides (Atlantic white-cedar) and several Betula lenta
(cherry birch) was suspended. Thanks to Patrick Cooney for compiling a comprehensive list
of species. Attendance: 18. Leader: Ted Gordon.

2 August: Atsion, Dutchtown, Parkdale, Hampton Furnace in Wharton State Forest,
Burlington and Atlantic counties, NJ. In the vicinity of the abandoned railroad tracks south
of the dam at Atsion Lake, we searched unsuccessfully for a known population of Ludwigia
birtella. Invasion of the site by a dense stand of young pitch pine and shrubs appears to have
crowded out these plants. Nearby, long-standing populations of Fimbristylis puberula,
Rhynchospora torreyana, Leiophyllum buxifolium, Gentiana autumnalis, Crotonopsis elliptica,
and Scutellaria integrifolia continue to prevail. A few flowering plants of Platanthera
blephariglottis were seen. Of note was a single flowering spike of Spiranthes vernalis along
the shoulder of Route 206. More at home along the coast, the spring ladies’-tresses
periodically shows up here and in a few other interior pine barrens spots. Southeast of
Dutchtown in a fire-prone lowland pitch pine forest, the coppery-yellow flowers of
icum denticulatum stood out. In and around bog-ore swales, we observed a fine
assemblage of sedges: Carex barrattii, C. bullata, C. debilis, C. livida, C. striata, Eleocharis
tenuis, E. microcarpa, E. tuberculosa, Scleria triglomerata, Rhynchospora cephalantha, R. pallida,
Scirpus cyperinus, and S. longii. Distinguishing characters of Long’s woolgrass were pointed
out, particularly the sticky black band at the base of the involucre and the saucer-shaped
tussocks of long-established plants. Among the grasses were Glyceria obtusa, Calamovilfa
brevipilis, Amphicarpum purshii, Mublenbergia torreyana, and Erianthus giganteus. On wet sand
asmall patch of an endangerd peatmoss, Sphagnum strictum, was pointed out. We found
plants of Xyris torta and X. difformis, but no plants of the endangered X. carolinana. Our
attempt to relocate Rhynchospora knieskernii on solid bog-ore “pavement” also was
unproductive. No doubt, prolonged drought had impacted this seed-banking population.
At Parkdale, site of an abandoned cranberry-blueberry village, we saw a fine stand of Prunus
maritima, flowering plants of Avenaria caroliniana, Euphorbia ipecacuanhae, and in sandy
paths, long-established stations of Crotonopsis elliptica. A historical occurrence of the root-
Parasite Schwalbea americana (chaffseed) was known from this complex. We inspected a
Patch of potential habitat, a pitch pine lowland that has been cut back, drum-chopped, and
Prescribed burned by the office of Natural Lands Management (N.J. D.E.P.) in an effort to
encourage the reestablishment of the endangered chaffseed from the seedbank or rootstock.
Reintroduction of the species is also being considered. The patch included Calamovilfa
pilis, Scleria triglomerata, Aster paternus, Pyxidanthera barbulata, Ilex glabra, Kalmia
angustifolia, and many other heath shrubs. Growing within the adjacent abandoned railroad
tracks were a few plants of Viola sagittata, a violet rare in the pine barrens along railroads,
according to Witmer Stone. Our final stop was at Hampton Furnace and vicinity to check
1 two populations of Lygodium palmatum, both known to several Philadelphia botanists
me the early 1900s. The population on the west bank of the Batsto River has severely
declined; that farther to the south on the opposite bank was thriving. In a sandy expanse
Neath an open cano y on the east side of the river between Hampton Forge and the
abandoned railroad were scattered cushions of Hudsonia ericoides, H. tomentosa, and Arenarta
caroliniana, In this pioneer habitat, we conducted an intensive search over about a one-mile
Stretch for Stylisma pickeringii var. pickeringii, a state-endangered and federally threatened
‘axon. While the group retreated to their vehicles, the leader, who persisted, found a single
vine of this morning-glory. Clearly, additional vines of the species observed here in the

160 BARTONIA

1980s have vanished. Thanks to Bill Standaert for compiling a comprehensive list of species
observed. Attendance: 19. Leader: Ted Gordon.

3 August: Lobachsville, Berks County, PA. The site is a successional old field and adjacent
wet woods along Pine Creek over which Albright College holds an easement. The site
overlies Leithville and Allentown Formation limestones. Our trip was timed to coincide
with the blooming period of the cranefly orchid, Tipularia discolor, which obliged with a
number of flowering stems. Other uncommon species at the site include leatherwood, Dirca
palustris, swamp lousewort, Pedicularis lanceolata, and broom-sedge, Andropogon glomeratus.
Deer browsing is becoming an increasing problem on the drier wooded portions. Dirca is
severely affected and other shrubs and herbaceous species are becoming sparse. On a June
1996 trip club members explored the diversity of sedges (more than 30 taxa) in the old field
and forest edge and puzzled over a clump of log fern, Dryopteris celsa, or was it D. goldiana
(?), along Pine Creek. Leaders: Susan Munch and Ann Rhoads.

23 August: Ferns of the Pinelands of Burlington County, NJ. From the Atsion Ranger
Station, we traveled well into the Wharton State Forest on sand roads to see Osmunda
cinnamomea, O. regalis, Lygodium palmatum, Pteridium aquilinum, Woodwardia virginica,
and W. areolata. At our next stop along a road just off Route 206, we found a colony of
Thelypteris simulata to compare with the more common T. palustris var. pubescens. The
common Onoclea sensibilis was also found along the road. The last stop for the day was
along the Oswego River where we looked for and found Schizaea pusilla, Pseudolycopodiella
caroliniana, Lycopodiella appressa and L. alopecuroides. We looked for, but did not find, the
hybrid of the latter two clubmosses. Attendance: 14. Leader: Bill Olson.

24 August: Hawk Island, Delanco Township, Burlington County, NJ. Low tide conditions
allowed us to walk the intertidal zone about halfway around this 118-acre peninsula at the
confluence of the Delaware River and Rancocas Creek. We then explored the interior of the
site, taking note of the plant species typical of both recent and older dredge spoils as well
as of an area that had seen little disturbance in recent years. Additions to a 1996 plant list
prepared for the Citizens United to Save Hawk Island included Sium suave, Eupatorium
pilosum, Helenium autumnale, Solidago juncea, S. gigantea, Hypericum canadense, Eleochanss
tenuis, Sicyos angulatus, Pilea pumila, Scutellaria lateriflora, Panicum verrucosum, Rhynchospora
capitellata, Lycopodiella appressa, and Isoetes riparia. This brings the total list for the site to
312 species, of which 110 are non-native. A complete list is available from the trip leader.
Attendance: 11. Leader: Karl Anderson.

6 September: Hamilton and Trenton Marshes, Duck Island Mitigation Site, Mercer sare
the

parviflorum). Also found were the rare beggar-ticks (Bidens bidentoides), ee
(Heteranthera reniformis), American water-willow (Justicia americana), false-pimper -
(Lindera dubia var. dubia), water-purslane (Ludwigia palustris), purple loosestrife
salicaria), mazus (Mazus pumilus), dotted smartweed (Polygonum punctatum var. 60 ‘
florum), and New York ironweed (Vernonia noveboracensis). Our exploration at the Duc

1997 FIELD TRIPS 161

Island site, a 90-acre constructed tidal freshwater wetland along the Delaware River, focused
only on areas at the southern end accessible by foot. (Frequently visited, the entire complex
has produced over 325 species of vascular plants, some of which have been transient.) Trees
we observed included ash-leaved maple (Acer negundo var. negundo), silver maple (Acer
saccharinum), honey-locust (Gleditsia triacanthos), cork-tree (Phellodendron sp.), sycamore
(Platanus occidentalis), and wafer-ash (Ptelea trifoliata var. trifoliata). The shrub false-indigo
(Amorpha fruticosa) was in fruit. Some of the vines in bloom were virgin’s-bower (Clematis
terniflora), climbing hempweed (Mikania scandens), and wild bean (Strophostyles helvola). Vines
not in bloom were moonseed (Menispernum canadense), climbing false-buckwheat (Polygonum
scandens var. scandens), and bur-cucumber (Sicyos angulatus). Herbs in bloom included
gerardia (Agalinis purpurea), water-hemp (Amaranthus cannabinus), common ragweed
(Ambrosia artemisiifolia), giant ragweed (A. trifida), annual wormwood (Artemisia annua),
asters (Aster pilosus, A. puniceus), showy bur-marigold (Bidens laevis), bull thistle (Cirsium
vulgare), tick-trefoil (Desmodium paniculatum), bottonweed (Diodia teres), viper’s-bugloss
(Echium vulgare), the near-flowering elscholtzia (Elscholtzia ciliata), willow-herb (Epilobium
coloratum), hyssop-leaved thoroughwort (Eupatorium hyssopifolium var. hyssopifolium), white
snakeroot (E. rugosum var. rugosum), late-flowering snakeroot (E. serotinum), eyebane
(Euphorbia nutans), marsh bedstraw (Galinum palustre), fragrant cudweed (Gnaphalium
obtusifolium var. obtusifolium), sneezeweed (Helenium autumnale), sunflower (Helianthus
decapetalus), Jerusalem-artichoke (H. tuberosus), rose mallow (Hibiscus moscheutos), dwarf St.-
John’s-wort (Hypericum mutilum), common St.-John’s-wort (H. perforatum), water
horehound (Lycopus americanus), winged monkey-flower (Mimulus alatus), carpetweed
(Mollugo verticillata), forget-me-not (Myosotis laxa), ditch stonecrop (Penthorum sedoides),
smartweeds (Polygonum lapathifolium, P. pensylvanicum, P. waria, P. punctatum, P.
sagittatum), pickerelweed (Pontederia cordata), arrowhead (Sagittaria latifolia), water-parsnip
(Sium suave), goldenrods (Solidago canadensis, S. gigantia, S. juncea), bluecurls (Trichostema
dichotomum), alsike clover (Trifolium hybridum), white vervain (Verbena urticifolia), and New
York ironweed (Vernonia noveboracensis). Herbs not in bloom included water-plantain
(Alisma subcordatum), spreading dogbane (Apocynum androsaemifolium), orache (Atriplex

ta), beggars-ticks (Bidens polylepis, B. connata, B. frondosa), partridge-pea (Chamaecrista
fasciculata), Mexican-tea (Chenopodium ambrosioides), pilewort (Erechtites hieraciifolia), mud-
P lantains (Heteranthera reniformis and the rare H. multiflora), Japanese clover (Lespedeza
strata), perilla mint (Perilla frutescens), cocklebur (Xanthium strumarium var. canadense) and
the rare subulate arrowhead (Sagittaria subulata). Rushes included Juncus acuminatus and J.
effusus var. solutus. Among the sedges were umbrella sedge (Cyperus bipartitus), yellow
nutsedge (C. esculentus), C. lupulinus, straw-colored nutsedge (C. strigosus), three-way sedge

lichium arundi ), the spil hes Eleocharis acicularis and E. ovata, the rare Smith’s
bulrush (Scixpus [Schoenoplectus] smithit), common three-square (S. pungens), woolgrass (S.
‘Yperinus), and soft-stem bulrush (S. validus). Thanks to William Standaert for compiling the

plant list and Patrick Cooney for writng the report. Attendance: 18. Leaders: Mary and
Charles Leck.

7 September: Central Perkiomen Valley Park, Montgomery County, PA. We explored
Young and old fields for late summer flowering plants including many asters and goldenrods,
Bidens polylepis, Ipomoea lacunosa, and at least seven other rampant vines. Among the species
‘0 Wet spots were Juncus effusus, Lindernia anagallidea, Lysimachia nummularia, and Eclipta

162 BARTONIA

alba. A list of 182 species was produced. Attendance: 18. Leaders: Ann Newbold, Heinrich
Zoller and Val Udell.

4 October: Lakehurst area, Pine Barrens, Ocean County, NJ. Beyond the roadside litter
of a parking area at the junction of Route 539 and Route 70 was a spectacular display of
Aster spectabilis and Aster linariifolius in full bloom. Asclepias amplexicaulis, Aureolaria
pedicularia, and Chrysopsis mariana were past flowering. Lichens, the focus of this trip,
included a variety of terrestrial Cladoniae: C. clavulifera, C. chlorophaea, C. cristatella, C.
atlantica, C. coneocraea, and Cladina subtenuis. Along a sand road through upland pine
forest at the New Jersey Audubon Society’s Hovnanian Sanctuary in Berkeley Township,
we saw large areas of forest floor dominated by Cladonia uncialis, with several other species
such as C. floridana and C. rappii. Cladonia bacillaris and C. incrassata were found on
decaying wood. Some bark-loving lichens, including Flavoparmelia caperata, Punctelia decta,
P. subrudecta, Parmelia sulcata, and Parmotrema hypotropum were compared. Vascular plants
of interest included Calamovilfa brevipilis, Leiophyllum buxifolium, Pyxidanthera barbulata,
and Xerophyllum asphodeloides. The day ended with a visit to Dover Forge, where we were
treated to a magnificent display of Gentiana autumnalis, about 300 flowers were counted.
A hardwood swamp here produced Imshaugia aleurites and a few small tufts of Usnea strigosa,
growing on Acer rubrum. A stroll to the banks of Cedar Creek added Smilax pseudochina,
Eupatorium resinosum, Oenothera parviflora, Rhynchospora gracilenta, and Bartonia virginica
to the day’s list. Attendance: 13. Leader: Karl Anderson.

13 December: Exploration of Pinelands plants in their winter aspect, NJ. After a hearty
breakfast at the restaurant at Marshalls Corner on the Monmouth-Ocean County border,
we headed south on Route 539, never straying very much from this road. Our first stop was
in Pasadena along the Pasadena-Woodmansie Road. There we looked at Andropogon
virginicus, Chimaphila umbellata, Helianthemum canadense, Hypericum gentianoides, Lechea
sp., Lespedeza capitata, Polygonella articulata, and Schizachyrium scoparium. Our next stop was
at Webbs Mill and the boardwalk into an Atlantic white-cedar stand, where we saw Carex
exilis, Chamaecyparis thyoides, Chamaedaphne calyculata, Clethra alnifolia, Drosera rotundifolia,
D. filiformis, Eleocharis tuberculosa, Juncus pelocarpus, Pseudolycopodiella caroliniana,
Narthecium americanus, Rhynchospora pallida, Sarracenia purpurea, Schizaea pusilla, and
Vaccinium macrocarpon. Two liverworts common in southern New Jersey were encoun
tered, Pallavicinia lyellii and Odontoschisma prostratum. Our next stop south of Webbs Mill
was in a borrow pit to see Rhynchospora knieskernii, the federally endangered beak-rush that
is only extant in New Jersey. After a stop in Warren Grove at Lucille’s Restaurant for a
warm lunch, we continued south on Route 539 to the Lower Plains to see Corema conradit
and other pine plains vegetation such as Pinus rigida, Quercus marilandica, Q. stellata, and
Polygonella articulata. Attendance: 12. Leader: Bill Olson.

1998 Field Trips

18 April: Forsythe National Wildlife Preserve, Manahawkin Section, Ocean County, NJ ;
The federal government bought this parcel of land and made it part of Forsythe Refuge 1n
about the mid-1990s to protect a large population of swamp-pink, Helonias bullata. oe
section that we visited extends from the upper (west) end of Manahawkin Lake and up M

1998 FIELD TRIPS 163

chose the water route, and the rest went on foot. We did not take an exact count, but
numerous early Helonias blooms were seen, photographed, and smelled. Their odor is
wonderfully sweet. Thanks to William Standaert who produced a comprehensive list of plant
species observed. Attendance: 7. Leader: Rob McCombe.

26 April: Clayton Park, Upper Freehold Township, Monmouth County, NJ. The habitat
of this inner coastal-plain park is mainly upland, dissected by small westward-flowing
streams. The uplands are dominated by fine stands of Fagus grandifolia, with Betula lenta and
Quercus spp.; lowlands, by Symplocarpus foetidus and Athyrium filix-femina, with Lindera
benzoin as a common shrub. The field trip produced a list of 144 species of vascular plants,
of which 126 were native. Good finds for the region included Dryopteris cristata, Huperzia
lucidula, Cardamine concatenata, Corydalis flavula, Luzula acuminata, Collinsonia canadensis,
Goodyera pubescens, and Pyrola rotundifolia. Of some interest was the scarcity of a few
species that might have been expected; Osmunda cinnamomea was virtually missing, as were
Veratrum viride, Podophyllum peltatum, and Cornus amomum, each species being represented
by only a very few individuals. Mertensia virginica, super-abundant in wetlands only a few
miles distant, was apparently completely absent. Leader: Karl Anderson.

2 May: Goat Hill Preserve, Chester County, PA. Our group proceeded down a powerline
cut from the parking lot off Red Lion Road to the Maryland state line. We were greeted
by rafts of Cerastium arvense var. villosum and Arabis lyrata in flower; Sisyrinchium
mucronatum and Minuartia michauxii were still in bud. We also observed other typical
members of the serpentine barrens flora leafing out including Quercus prinoides, Q. ilicifolia,
Q. marilandica, and Ceanothus americanus among woody plants, and herbs such as Phlox
subulata, Asclepias verticillata (just emerging), Oenothera fruticosa, and Viola sagittata. The
upland Carex flora was disappointing, yielding only a few species such as Carex umbellata
and C. nigromarginata. However, a rich low meadow at a brook crossing yielded C. intertor
(positively identified eight days later) and C. vestita. Other unusual species seen included a
arge specimen of Amianthium muscaetoxicum along a trail in upland scrub shrub and several
populations of Lysimachia lanceolata at its only Chester County locale. It was beginning to
rain lightly, and most of the participants had turned back, when Tim Draude found the
plant the trip had been especially organized to seek out, the rare disjunct Carex richardsonit.
Several colonies of this turf-forming sedge, previously unrecorded from Goat Hill and only
at its second known Pennsylvania station, were in full fruit right at the state line. After
lunch, several trip members made a loop through Nottingham Park in an unsuccessful
attempt to locate the first-known Pennsylvania population of C. richardsonii. We were able
to check on colonies of C. meadii and C. vestita in the park. Attendance: 11. Leaders: Janet
Ebert and Jack Holt.

16 May: Meng Preserve, Montgomery County, PA. Joint trip with Valley Forge Audubon
Society. Numerous club and society members participated in a leisurely stroll along existing
trails and old logging roads through Valley Forge Audubon’s new Meng Preserve, an
excellent example of a rich woodland community on diabase. Despite recent logging atop
the ridge and the presence of a timber staging area used as a parking lot at the base of the
slope, the relative absence of alien species in the preserve is notable. The diversity of the
lush woody, herbaceous, and graminoid growth was quite high. Notable species included
Corylus cornuta, Trichophorum planifolium, Carex wildenovit, Galearis spectabilis, Smilax

164 BARTONIA

hispida, Cynoglossum virginianum, and Hypoxis hirsuta. The trip ended atop the nearly clear-
cut ridgetop, where a discussion was held comparing the surviving flora of the logged area
to that on the slopes below and regarding the prospects of recovery of the site. Leaders:
Janet Ebert and Jack Holt.

23 May: Island Beach State Park, Ocean County, NJ. In the vicinity of the Aeolium, we
found both flowering and fruiting Spergula morisonii, Morison’s sand spurry, an addition to
the park’s plant list. At the border of a parking lot near Reed Road, we saw a few plants
of Chrysopsis falcata, sickle-leaved golden aster, one of at least four small occurrences of this
state-rare aster within the park. We followed Reed Road to the bay and then walked north
along the tidal strip strewn with ribbons of Zostera marina, eelgrass. The edge of high tide
did not produce a single patch of Honckenya peploides, sea-beach sandwort. After a short
distance, we entered a coastal dune woodland dominated by Pinus rigida with interdunal
hollows sheltering Chamaecyparis thyoides. This community type is critically imperiled in the
state and apparently it is restricted to this pocket in the northwestern section of the
Northern Natural Area. In The Plants of Southern New Jersey (1911), Witmer Stone cited E.
B. Bartram’s discovery of Schizaea pusilla in back of the sand dunes near Seaside Park “in a
moist hollow with a few little White Cedars.” This passage prompted the leader to visit this
area. Pitch pine groves with canopies to 30 feet occurred in dune depressions surrounded by
extensive open Hudsonia tomentosa communities in bloom, with scattered Ilex opaca, Quercus
falcata, Q. marilandica, Q. ilicifolia, and Juniperus virginiana. Several interdunal hollows held
shallow water and contained both dead snags and living trees of Atlantic white-cedar. These
hollows were ringed by a shrub layer of Myrica pensylvanica, Vaccinium corymbosum, Kalmia
angustifolia (all three in bloom), Gaylussacia frondosa, and Ilex glabra. Salt spray injury to the
tops of a few pines and white-cedars was observed. Successive spraying events could have
resulted in the death of the latter. Days later Pete McLain and a student conducted salinity
tests of the water in one of these depressions. Their results showed 0.0 ppt for salinity. Of
the three white-cedar hollows we examined, none contained suitable curly-grass fern habitat.
Although salinity does not appear to have caused its demise, Schizaea cannot be declared
extirpated without searching all interdunal depressions harboring white-cedar. In sight of
the ocean, we next examined a foredune dominated by Carex kobomugi, Japanese sedge, first
collected on Island Beach by C.W. Townsend in 1929. Aggressive reproduction of this
dioecious sedge by rhizomes is believed to have resulted in substantial loss of native plants,
particularly in the Southern Natural Area. Control measures of this invasive plant are under
consideration. Some associated species sparsely present were Ammophila breviligulata,
Solidago sempervirens, Cakile edentula, Atriplex hastata, and a handsome patch of the
naturalized Artemisia stelleriana in flower. In other habitats, widely distributed species still
in bloom included Amelanchier canadensis, Prunus serotina, and P. maritima. A visit to the
Coast Guard Station Interpretive Center to examine the Emily de Camp Herbarium of
specimens collected in the park concluded our trip. Thanks to Cynthia Coritz and
Superintendent Bill Vibbert for arranging our visit and to Bill Standaert for compiling a list
of plants observed. Attendance: 8. Leader: Ted Gordon.

21-25 June: Johnson State College (J.S.C.), Johnson, in the Green Mountains of north
central Vermont. Joint field meeting of the Botanical Society of America, Torrey Botanical
Society, and the Philadelphia Botanical Club. On Monday we carpooled to the foot of the
toll road in Stowe, where vans took us to the 4,303-foot-high summit of Mount Mansfield.

1998 FIELD TRIPS 165

We came to observe the largest community of arctic-alpine tundra vegetation in the state.
Sighted among many more species were Arenaria groenlandica, Gaultheria hispidula,
Habenaria (Platanthera) dilatata, Ledum groenlandicum, Nemopanthus mucronatus, Potentilla
tridentata, Ribes glandulosum, Salix pyrifolia, Sambucus racemosa var. pubens, Vaccinium
uliginosum, V. vitis-idaea, V. boreale (!), and the ever-present lichen Rhizocarpon geographica.
Our leaders were Charlie Cogbill (environmental consultant) and Peter Hope (Boston
professor at St. Michael’s in Winooski, VT), both experts on the flora of the mountain. On
Tuesday, J.S.C. professor of environmental sciences John Wrazen and Peter Hope, a fern
specialist, were our guides at the Babcock Preserve, 1,000 acres of northern mixed hardwood
forest, ponds, fens, and other wetlands owned by J.S.C. A sampling of the plants seen was
Dryopteris compyloptera and its hybrids, Coptis trifolia, Drosera rotundifolia, Oxalis acetosella,
Polystichum braunit, Potentilla palustris, Scutellaria galericulata, and Streptopus amplexifolius.
On Wednesday, sphagnum specialist Cyrus McQueen of J.S.C. and Debbie Benjamin of the
Vermont Bird and Botany Club led us to several peat bog communities: South Molly Bog,
Percy Bog, and Joes Pond. A big treat was a “meadow-like” mat of sphagnum covered with
Pogonia ophioglossoides, Calopogon tuberosus, Platanthera blephariglottis (still in bud),
Eriophorum virginicum, Rhynchospora alba, and Carex trisperma. Sphagnum species were
numerous, but, due to a great deal of recent rain, many of the attendees did not follow
Cyrus into a bog that required wading hip-deep to his destination. Other plants found were
Kalmia angustifolia, Lysimachia terrestris, and Sarracenia purpurea. Three evening slide-
illustrated lectures were presented by experts in various fields. David Marvin, forester and
owner of Butternut Mountain Farm, which produces maple syrup and other maple products,
spoke on the “Production of Maple Syrup in Vermont.” Charles Johnson, a charismatic and
articulate naturalist, presented “The Nature of Vermont,” a delightful overview of the state
and its ecology. His book by the same title has just been reprinted. He is also author of the
well-known book, Bogs of the Northeast. “Aspects of Forest Decline on Camels Hump in
the Green Mountains” was the lecture presented by Hubert Vogelmann, the retired head of
the University of Vermont’s Botany Department. His trail-blazing work in the 1960s led to
the awareness of the effects of acid rain on forests. While this work was done three decades
ago, it was the foundation of much that has been studied since that time. Trip Chairperson:
Ursula Joachim.

11 July: Bennetts Mill and Vicinity, Peasle Fish and Wildlife Management Area, City of
Vineland and Maurice River Township, Cumberland County, NJ. This county has long
been a popular botanical destination. In particular, Bayard Long’s forays into Cumberland
Provide some of the best documentation of its flora. Bennetts Mill is an abandoned farm and
small industrial complex at the headwaters of the Manumuskin River. Here our work began
in a large old sandy field cultivated every other year by the N.J. D.E.P. Division of Fish and
Wildlife. In this early successional habitat we saw Panicum sphaerocarpon, P. auburne, #
depauperatum, P. scoparium, Schizachyrium scoparium, Carex tonsa, Lespedeza hirta, and
Tridens flavus. We inspected the old ruins at the intersection of Bennetts Mill and Old Mays
Landing roads. Here we found Aralia nudicaulis, Asclepias tuberosa, Solidago spp., and
Crataegus uniflora. Our second stop was 2.6 miles south of Bennetts Mill. We explored a
Power line cut from the edge of Union Road to the floodplains of the Manumuskin River.

€ uplands and wetland of the right-of-way have been repeatedly mowed, reducing woody
Vegetation to simple basal sprouts. This treatment has provided an open area for plants more
‘oOmmon on unplowed habitats devoid of a canopy. Plants of interest included Scleria

166 BARTONIA

minor, S. triglomerata, Sorghastrum nutans, Andropogon virginicus, Rhynchospora gracilenta,
Platanthera blephariglottis, Polygala brevifolia, Drosera rotundifolia, D. filiformis, Sarracenia
purpurea, and Pogonia ophioglossoides. At Cumberland Pond, we stopped at an old field on
the south side of Route 49. Here we saw Gymnopogon ambiguus, Desmodium strictum, and
large colonies of Opuntia humifusa. From this well-known site we drove to a large clear-cut
northeast of Cumberland Pond at the intersection of Hesstown Road and Main Road. In a
newly plowed area within a large oak-pine forest, we found Linum virginianum, Panicum
villosissimum, P. dichotomum, and Comandra umbellata along the forest edge. Leader: Joseph
Arsenault.

23 August: Barkwoods Pond, Hirst Pond, Goose Ponds, Egg Harbor City vicinity, Atlantic
County, NJ. Along a split-rail fence at the head of a trail leading to Barkwoods Pond, we
saw a cluster of Corallorhiza odontorhiza in full anthesis. Panicum ensifolium was along the
trail leading to the pond, which we searched unsuccessfully for a known population of
Panicum hirstii. In shallow water were the filiform leaves with cleistogamous flowers of
Utricularia geminiscapa. Yellow patches of Gratiola aurea occupied exposed pond bottom.
Also in bloom were Polygola lutea, Lindernia dubia, and Xyris smalliana. Identifiable by its
linear, lanceolate leaf segments was Bidens coronata. Among the species obviously common
to all of the intermittent ponds we visited were Eleocharis olivacea, E. microcarpa, E.
robbinsii, and Xyris smalliana. At nearby Hirst (Labounsky) Pond we compared two delicate
bladderworts in flower: Utricularia striata (U. fibrosa) and U. subulata, the latter with peltate
bracts. The pink heads with yellow disks of Coreopsis rosea were somewhat concealed within
stands of Cladium mariscoides and Carex striata. Also in bloom were Nymphoides cordata,
Lycopus uniflora, Gratiola aurea, and a few Rhexia virginica and R. mariana. We saw
scattered tiny seedlings that may have been Scleria reticularis var. reticularis, generally here
in abundance. We did not find Lobelia boykinii. The discovery of the “lost” Eleocharis
equisetoides (fruiting) in Big Goose Pond was a highlight; the state endangered knotted
spikerush was in the same spot where it was last pointed out to the leader in 1978 by the
late Gil Cavileer. Adjacent was a fine population of Rhynchospora inundata. Elsewhere
Lobelia nuttallii, Drosera intermedia, and especially Utricularia cornuta made fine displays.
At Little Goose Pond, Lobelia boykinii eluded us. However, we were rewarded by sighting
two state-endangered species: a few flowering stalks of Rhexia aristosa and a few fruiting
plants of Sagittaria teres, slender arrowhead. Associated species in bloom were Eriocaulon
aquaticum and Xyris torta. (The original specimens of awned meadow-beauty discovered in
1888 came from the Egg Harbor City vicinity.) Thanks to Bob Johnsson for compiling a
list of species observed. Attendance: 23. Leader: Ted Gordon.

27 September: Atsion and vicinity, Burlington County, NJ. The focal point of this trip was
the pine-barren gentians that have grown for many years along Route 206, but a £°
variety of other pine barrens plants was noted. Grasses were particularly abundant ani
diverse in sandy fields south of the Atsion Ranger Station; about 20 species were found,
including Chloris verticillata, Triplasis purpurea, Leptoloma cognatum, Panicum scoparium, a0
Sorghastrum nutans. Gentiana autumnalis was found in good numbers and fine bloom,
of the old Central Railroad of New Jersey tracks. Nearby were scattered plants of Spiranthes
cernua, Trichostema dichotomum, and Crotonopsis elliptica. A drying swale on the west ih
of Route 206 gave participants a chance to see some plants involved in the process of pon
succession, including Chamaedaphne calyculata, Cladium mariscoides, Carex striata, #0

1998 FIELD TRIPS 167

Vaccinium macrocarpon, along with Drosera intermedia, Proserpinaca pectinata, Dulichium
arundinaceum, Juncus pelocarpus, Eleocharis olivacea, and Rhynchospora capitellata. Lunch was
at the Atsion Recreation Area. A short walk from here along a nature trail south of Atsion
Lake produced Polygonella articulata in some abundance on dry sand, while ditches and the
lake shore produced Woodwardia virginica, Osmunda cinnamomea, O. regalis, Triadenum
virginicum, Juncus miliaris, Orontium aquaticum, and Nymphoides cordata. The trip ended
with a stroll to a stream-bank and wet area along the entrance road to the recreation area,
where a few plants of Rhexia virginica were still in bloom. Here also were Pontederia
cordata, Peltandra virginica, Juncus biflorus, J. canadensis, Woodwardia areolata, Botrychium
dissectum, Thelypteris palustris, and numerous plants of Polygala lutea, this last species still in
flower. Attendance: 20. Leader: Karl Anderson.

1999 Field Trips

25 April: Lebanon State Forest, Ongs Hat, Buffin Meadows, Burlington County, NJ. Near
the Lebanon State Forest office, we looked at a fire-protected, advanced growth oak-pine
forest dominated by chestnut, black, and white oaks with pitch pine as a minor component.
Such mature stands are rare in the Pine Barrens because of frequent fires and premature
harvesting. Here, along a path, we saw Epigaea repens in bloom. Near the fire tower was a
large patch of flowering Pyxidanthera barbulata. We noted an old Pinus resinosa plantation
exhibiting severely retarded growth. In contrast, at Ongs Hat, a Pinus taeda plantation was
flourishing. On a wet shoulder of the hardtop road were Carex barrattii and C. vestita, both
in bloom. Already in fruit, a few clumps of Carex nigromarginata occupied a dry slope. We
next visited a hardwood-cedar swamp along Company Branch to assess a Helonias bullata
population, first reported by Rusk and Svenson in 1933 as “abundant.” In 1968 the leader
saw about 50 plants in flower; in April 1989 there were at least 43 rosettes, only one in bud.
In 1999 we saw a mere 5 vegetative plants in poor condition hugging the stream bank just
west of the road that dissects the swamp. Herbivory by deer and, especially, temporary
water diversion over an extended period into a canal and the resulting water drawdown have
all but eliminated this swamp-pink occurrence. It is of interest that more than 50 scattered
large Betula alleghaniensis (B. lutea), a tree rarely encountered in the Pine Barrens, are a well
established component of this community. At Cedar Swamp Natural Area back in the
Lebanon, we were dismayed by the destruction of a roadside population of the rare Uvularia
puberula var. nitida. At their request, forest personnel had been given the precise location
of this station; yet, the maintenance crew scraped it into oblivion. Nearby was a fine stand
of Diphasiastrum (Lycopodium) digitatum, a clubmoss infrequent in the barrens. In bloom
were Lindera benzoin, Orontium aquaticum, Amelanchier canadensis, Vaccinium corymbosum,
and, with flower buds about to open, Leiophyllum buxifolium. With great difficulty we
teached more than a dozen vigorous rosettes of Helonias with some eight budding stalks six
to eight inches tall. These plants occurred on hummocks in a 100-foot stretch of stream
channel, persisting despite a relatively closed canopy of white-cedar. At Bear Hole we
examined the hard, knuckle-like, fire-scorched bases and roots of Calamovilfa brevipilis.
Added to our list of plants in bloom were Chamaedaphne calyculata, Vaccinium angustt-
folium, and Carex stricta. We monitored a 100-by-30-foot segment of a small Helonias
Population along a streamlet at Buffin Meadows, discovered by the leader in March 1991.
Fifteen clumps were seen beneath Clethva alnifolia on sphagnous hummocks surrounded by

168 BARTONIA

deep pools in an open canopy of Nyssa and Acer. The population had remained stable, and
three specimens of swamp-pink in full anthesis delighted us all. The surrounding white-cedar
forest was devoid of Helonias. We did not monitor the larger population in the segment to
the west. Thanks to Bill Standaert for compiling a list of species observed and to John
O’Herron for his input. Attendance: 16. Leader: Ted Gordon.

8 May: Sourland Mountain Preserve, Hillsborough Township, Somerset County, NJ. Joint
trip with the New Jersey Audubon Society. This 1,600-acre mostly-forested preserve has a
diverse flora, and about 100 species were identified during the day. Many woodland spring
flowers were in bloom, among them Podophyllum peltatum, Conopholis americana,
Anemonella thalictroides, Actea alba, Geranium maculatum, Asarum canadense, Arisaema
triphyllum, Claytonia virginica, Saxifraga virginiensis, Viola palmata, V. sororia, and V.
pubescens. Orchis (Galearis) spectabilis was in bloom and was quite abundant in some areas;
several white-flowered plants were noted. Aquilegia canadensis was found in bloom in partial
shade along a pipeline cut, and Matricaria matricarioides, Geranium pusillum, and several
other ruderals were in bloom around the parking area. Some noteworthy finds were
Obolaria virginica, Cynoglossum virginianum, Panax quinquefolius, and Cardamine angustata,
all of which are threatened species in New Jersey; of these, the Obolaria and Cardamine
were in bloom, and the Cynoglossum was in bud. Some identifiable plants that were not in
bloom included Menispernum canadense, Allium tricoccum, Cimicifuga racemosa, and Galium
lanceolatum. Ferns seen included Dryopteris goldiana, D. cristata, Adiantum pedatum, Deparia
achrostichoides, and Phegopteris hexagonoptera, as well as several more common species. A list
of plants seen is available from the trip leader. Leader: Karl Anderson.

25 May: Greater Forked River Mountains Region, Ocean and Lacey Townships, Ocean
County, NJ. The main purpose of this weekday trip was to seek previously unknown
occurrences of Arethusa bulbosa on sites judged earlier by the leader to contain suitable
habitat for this state-rare orchid. Along Cold Brook, a short distance northeast of Wells
Mills, we entered a cut-over (perhaps 40 years ago) sphagnum-carpeted cedar swamp with
savanna-like openings dotted with magnolia, maple, and Atlantic white-cedar saplings. Four
significant species, first recorded here in 1995, were relocated: several rough stems of Juncus
caesariensis, a clump of Calamovilfa brevipilis, a couple of sharp-pointed leaves of Narthecium
americanum, and numerous fronds of Schizaea pusilla. More than a dozen scattered plants
of Arethusa bulbosa in flower were a new discovery. These five species occupied a hummock
zone comprised in part of Carex exilis, C. atlantica including var. capillacea, C. trisperma,
Pogonia ophioglossoides, Gaylussacia dumosa, and other heaths. Two dominant mosses were
Sphagnum pulchrum and S. flavicomans. Carex collinsu occupied seeps, and near the base of
a shrubby slope was the foliage of eight Platanthera blephariglottis. Here we also saw a garter
snake. No effort was made to explore this segment of swamp more intensively; however,
we re-entered the corridor a few thousand feet upstream and added a new population of
about 80 plants of Arethusa bulbosa in bloom. Schizaea pusilla hugged the vertical cut of the
stream bank, and several Platanthera clavellata were perched on mats of sphagnum. The
adjacent fire-scorched pitch-pine lowland was dominated by a massive stand of Calamovilfa
brevipilis, and a slightly drier zone, by hundreds of grass-like clumps of Xerophyllum
asphodelloides. In the upland pine-oak forest, we encountered—all within the space of a
hundred yards—a hognose snake, a black widow spider, and a pair of mating pine snakes @
threatened species). A few miles to the northeast, we visited a young regenerating cedar

1999 FIELD TRIPS 169

swamp that runs along lower Sprague Branch, a narrow feeder stream of the North Branch
of Forked River. At the foot of a slope associated with a pine-oak forest, this sphagnous
community comprises a habitat of high species diversity, favored by harvesting, a stable
hydrological regime, and browsing of white-cedar saplings by deer. This site and that of our
first stop share a number of similarities. We relocated two species discovered in 1995: four
plants of Arethusa in bloom and Schizaea, scattered on several hummocks. Among other
species seen were Sarracenia purpurea, Platanthera blephariglottis, Aster nemoralis, A. novt-
belgii, Pogonia ophioglossoides, Drosera rodundifolia, Lilium superbum, Aronia arbutifolia,
Carex exilis, Trientalis borealis, and Utricularia subulata, the latter four in flower. Of special
significance was finding 13 plants (7 in bloom) of Uvularia puberula var. nitida, the state-
endangered pine barren bellwort. These were well concealed along the shrubby ecotone near
the base of the slope. Lastly, we concluded a banner day by briefly handling a pine barren
treefrog, Hyla andersonii. Thanks to Bill Standaert for compiling a list of species observed
and to John O’Herron for sharing his notes for this report. Attendance: 18. Leader: Ted
Gordon.

5 June: Adkins Arboretum, Caroline County, MD. Joint trip with the Muhlenberg
Botanical Society and the Maryland and Delaware Native Plant Societies. We walked the
trails of the floristically diverse grounds of the arboretum, which ranges in habitat from
upland open fields to waterlogged floodplain forest. In the upland oak-heath forests we
observed typical dry-soil, acid-loving species such as Cypripedium acaule, Goodyera pubescens,
and Chimaphila umbellata. A loblolly pine plantation yielded the spurges Euphorbia corollata
and E. ipecacuanhae along with Asclepias amplexicaulis. In richer areas near the floodplain
several species more typical of the Piedmont flora were observed, including Arabis
canadensis, A. lyrata, and Pycnathemum incanum. A surprise here was a beautiful flowering
plant of Asclepias variegata, previously unknown from the property. A few non-flowering
plants of Galearis spectabilis, along with Ophioglossum vulgatum, Desmodium glutinosum,
Aristolochia serpentaria, and Aquilegia canadensis grew in a small, rich woods pocket near a
back swamp, only a few yards downslope from Clitoria mariana and Opuntia humifusa. The
back swamp itself was host to an enormous population of Dryopteris celsa; we saw thousands
of plants of this large, uncommon fern. At the far end of the property along a hundred-year
floodplain trail grew a large flourishing colony of a shrub more typical of Appalachia and
the Midwest, Viburnum rafinesquianum. On the way back we walked along the edge of a
bare-bottomed wooded slough lined by ericaceous shrubs including Leucothoe racemosa,
Vaccinium corymbosum, and Rhododendron viscosum. Attendance: 12. Leaders: Janet Ebert
and Jack Holt.

12 June: Malaga and Vicinity, Franklin Township, Gloucester County, NJ. This trip
Concentrated on the genus Panicum, a group of grasses that has intrigued the leader. We held
a short discussion on the genus, describing the characters that are unique and similar
between the subgenera. We compared the various available keys and discussed how this grass
group is viewed today. Despite its archaic nomenclature, Gray’s Manual of Botany, 8th
Edition, was recommended over other manuals because the keys and field description were
judged superior. Our first stop was a Nature Conservancy property south of Route 40 and
east of Route 55 near Malaga. Underlain by a coarse sandy soil typical of the Outer Coastal
Plain, this site comprised a pitch pine community characteristic of the frequent fire regimes
of the Pine Barrens. We walked the sand trails and found Panicum virgatum, P. depauper-

170 BARTONIA

atum, P. clandestinum, P. columbianum, P. commutatum var. ashei, P. spretum, P. villosis-
simum, P. lanuginosum var. lindheimeri. P. meridionale. P. mattamuskeetense, P. microcarpon,
and P. lucidum. After a brief downpour, we drove to the Glassboro Fish and Wildlife
Management Area. This public area is centered on the headwaters of Still Run, a Maurice
River tributary and 5.9 miles upstream from the Malaga site. This area has soils with finer
texture, typical sandy silt of the Bridgeton Formation, a Pleistocene-era gravel cap covering
portions of the Inner and Outer Coastal Plain. Here we identified Panicum virgatum, P.
commutatum var. ashei, P. scoparium, P. sphaerocarpon, P. dichotomum, P. clandestinum, P.
lanuginosum, P. villosissimum, P. meridionale, and P. lucidum. This assemblage is similar to
the species (five common to both sites) observed in Malaga, but the presence of P. scoparium
in an upland setting with P. dichotomum reflects the moist conditions associated with the
fine-textured Bridgeton Formation. Heavy downpours during lunch and at the conclusion
of fieldwork prevented us from keying specimens as planned. Leader: Joe Arsenault.

18-20 June: Pocono Environmental Education Center, PA. Friday night we met at PEEC
and a slide show of the leader’s trip to Puerto Rico was shown. A trip to Hogback Ridge,
a limestone ridge, on Saturday in Pike and Monroe counties provided the group an
opportunity to see upland and wetland species growing on limestone. Species included
Adiantum pedatum, Arisaema dracontium, Carex granularis, C. lupulina, C. plantaginea, C.
typhina, Callitriche terrestris, Hypericum acyron (H. pyramidatum), and Quercus bicolor. The
evening was spent looking at some of the specimens at the lab back at PEEC. On Sunday,
we traveled up to Orange County, NY to explore the Rutgers Creek Conservancy of the
Paul F. Brandwein Institute. This rocky wooded area with a trout stream flowing through
it contained an interesting complex of species on limy soils. The species included Cardamine
diphylla, Carex argyrantha, C. blanda, C. bromoides, C. gracillima, C. granularis, C. pallescens,
C. squarrosa, Epigaea repens, Equisetum hyemale var. affine, Fraxinus americana, F. nigra,
Lobelia cardinalis, L. siphilitica, Trillium erectum, Viola rostrata, V. sororia, and V. labradorica
(V. conspersa). Carex davisii, a sedge seen previously on the site, was looked for, but not
found. Attendance: 11. Leader: Bill Olson.

20-24 June: Manchester College, North Manchester, northern Indiana. Joint field meeting
with the Botanical Society of America and the Torrey Botanical Society. There were three
days of field trips. The first site was the Tefft Savanna State Nature Preserve, located in the
Kankakee sand plain region in northwestern Indiana. Habitats seen here were oak savanna,
sand prairie, and wetlands. Atlantic Coastal Plain species represented an unexpected floristic
element in this area. Spinn Prairie, a Nature Conservancy project, gave participants a look
at a mesic prairie-savanna mosaic. A brief stop was made at Berns-Meyers Woods, a small
fragment of old-growth mesic forest, also a Nature Conservancy site. On the second day the
sites visited were Laketon Bog State Nature Preserve, a fen containing the southernmost
population of larch in the state, and Ginn Woods, one of the largest stands of old-growth
forest in the state, which is owned by Ball State University. The final day of field trips were
at Pigeon State Fish and Wildlife Area near Mongo. Here, participants canoed into the
Tamarack Bog State Nature Preserve and later visited a fen in the Mongoquinong State
Nature Preserve. Field trip leaders were Tom Post (Tefft and Spinn) and Lee Casebere
(Pigeon River) of the Indiana Department of Natural Resources, Paul Rothrock of Taylor
University (Laketon), and Kem Badger and Don Ruch of Ball State University eee
Woods). Evening presentations covered orchids of Indiana, presented by Mike Homoya ©

1999 FIELD TRIPS 171i

IL.D.N.R.; nature photography, by Lee Casabere; and northern Indiana vegetation, by David
Hicks of Manchester College. Attendance: only 25, primarily from the eastern states. Field
trip assistant: Betty Oldham. Meeting organizer: David Hicks.

27 June: Middle and South Branches of the Forked River, Lacey Township, Ocean County,
NJ. Participants consolidated into four-wheel-drive vehicles to penetrate to the wildest
recesses of the Forked River Mountains region. Shrubs were in bloom along the borders of
sand roads and elsewhere were Ilex glabra, Kalmia angustifolia, Lyonia mariana, and
Rhododendron viscosum. A few bushes of Vaccinium corymbosum were already bearing blue
fruit. South of the central section of the main stem of the Middle Branch and west of the
Garden State Parkway, we 1 igated a white-cedar-dotted Narthecium } savanna,
surveyed in 1998 by A.E. Schuyler and the leader. It is, in part, the subject of a paper in this
issue of Bartonia. We saw two sizeable pockets of the globally rare and state-endangered bog
asphodel, far removed from the core populations in the heart of Wharton State Forest. The
majority of these plants were sterile, but scores held aloft their yellow spikes. Associated
species also in bloom included Nymphaea, Pogonia, Sarracenia, Lophiola aurea, Utricularia
cornuta, U. striata (U. fibrosa), U. subulata, Drosera filiformis, and Calopogon tuberosus.
Among the sedges were Carex exilis, Cladium mariscoides, Eleocharis tuberculosa, E. tenuis,
Rhynchospora fusca, and immature plants of R. alba. Grasses seen were Andropogon
glomeratus, Danthonia sericea var. epilis, Mublenbergia uniflora, and Panicum ensifolium.
Young plants of Juncus pelocarpus were just beginning to emerge. Schizaea pusilla occurred
along with Pseudolycopodiella caroliniana along a white-cedar-dominated tributary farther
south. Earlier in the month the leader discovered 10 flowering stalks of Arethusa here; none
was seen this time. Farther east near the Parkway, we visited a severely disturbed habitat
whose unique flora had been previously documented. Flowering were Nymphoides cordata,
Sagittaria teres, Eriocaulon aquaticum, and Proserpinaca pectinata, while Utricularia resupinata
was still in bud. On the moist fringe were Polygala lutea in bloom, the foliage of Gentiana
autumnalis, our three Drosera, Juncus canadensis, Eleocharis olivacea, E. microcarpa, Scirpus
pungens, Cyperus dentatus, Amphicarpum purshii, and old culms of Rhynchospora knieskernii.
Occupying peaty soil were Calamovilfa brevipilis and Mublenbergia torreyana. At our final
stop, an open regenerating cedar swamp near the headwaters of the South Branch, we saw
two exotic pitcher plants, Sarracenia minor and S. rubra. Introduced many years ago, these
stands blended remarkably with the typical native species Sarracenia purpurea, Pogonia,
Utricularia striata, Juncus caesariensis, J. canadensis, J. marginatus, J. scirpoides, Eriophorum
Virginicum, Carex striata, and Vaccinium macrocarpon. Thanks to Bill Standaert for providing
a fine comprehensive list of the species observed. Attendance: 9. Leader: Ted Gordon.

10 July: Whiting and Whitesbog, Lebanon State Forest, Ocean and Burlington Counties, NJ.
Joint trip with Torrey Botanical Society. At Whiting Wildlife Management Area we stopped
at a mowed field with seepage area and associated damp swales. Here we found flowering
Calopogon tuberosus, Lobelia nuttallii, Polygala lutea, Xyris difformis, and X. torta. This site
also produced several interesting sedges, including Carex barrattii, Eriophorum virginicum,
Rhynchospora pallida, Scleria minor, and S. triglomerata. Our next stop was the ruins of a
terra-cotta factory on the eastern edge of Lebanon State Forest, where we found Chimaphila
maculata and C. umbellata flowering side-by-side. Also here were a few flowering Epipactis
helleborine, most in poor condition due to the summer’s drought. After lunching at
Whitesbog village, we found the ferns Asplenium platyneuron, Atherium filix-femina,

172 BARTONIA

Botrychium virginianum, B. matricarifolium, B. dissectum, and Ophiogossum vulgatum in the
vicinity of some abandoned buildings. Nearby was a nice clump of Linum striatum. Other
stops at Whitesbog yielded flowering Avenaria caroliniana, Asclepias tuberosa, Aster paternus,
Calluna vulgaris, Galactia regularis, Platanthera blephariglottis (with buds), Pogonia
ophioglossoides, Rhododendron viscosum, Schwalbea americana, and Stylisma_pickeringii.
Thanks to Ted Gordon, Ralph and Ellen Wilen, and Howard Wildman for help with
transportation. Leader: Linda Kelly.

14 August: Whittingham Wildlife Management Area, Fredon Township, Sussex County, NJ.
This 1,600-acre preserve has a diverse flora typical of New Jersey’s limestone areas. This trip
focused on marshes and intermittent ponds that would normally be almost impenetrable,
but that had been made accessible by several weeks of drought. A good variety of ferns was
noted, among them Dryopteris goldiana, D. cristata, D. clintoniana, several Dryopteris hybrids,
Diplazium pycnocarpon, and such limestone-loving species as Asplenium rhizophyllum, A.
trichomanes, Woodsia obtusa, and Cystopteris tenuis. A good find was a large clone of
Rhamnus alnifolia, a northern species that reaches its southern limit of range in New Jersey,
growing in a dry and rather confusing lowland dominated by Carex stricta, Toxicodendron
vernix, and Fraxinus nigra. Carex lupiliformis was seen on the edges of several dry ponds, and
several plants of Platanthera psychodes, well past bloom, were located in a dry swamp that
also contained Cypripedium calceolus, Smilacina stellata, and Equisetum fluviatile. Lobelia
cardinalis was almost the only woodland or wetland plant in bloom, but upland fields were
ornamented with flowers of Monarda fistulosa, Dipsacus sylvestris, Carduus acanthoides, and
several species of Solidago. Many thank to Ralph Wilen for pointing out some interesting
Dryopteris hybrids, and to Bill Olson for taking a second look at Carex lupuliformis. Leader:
Kar] Anderson.

29 August: Palmyra Cove Nature Park, Palmyra, Burlington County, NJ. This trip included
a 300-acre area of landfill, freshwater tidal marsh, and dredge impoundments bordered by
the Delaware River, Pennsauken Creek, and Route 73. The flora of this site consists of
about 350 species, of which about one-third are non-native. Uplands here are dominated by
such species as Ailanthus altissima and Populus deltoides, with an understory of Polygonum
cuspidatum and Urtica dioica. Also present were Strophostyles umbellata, Cycloma atriplict-
folium, Lycopus europaeus, Verbascum phlomoides, Mirabilis nyctaginea, Leptoloma cognatum,
and many other plants typical of dry, disturbed habitats. Wet ditches had Ludwigia peploides,
Spirodela polyrhiza, Potamogeton crispus, Elodea canadensis, and Ceratophyllum demersum. The
intertidal flats along the Delaware River here were surprisingly devoid of vegetation, even
though there are extensive vegetated flats just north of the site; however, we did find a few
plants that were tentatively identified as Sagittaria subulata. A fine stand of Salix exigua, @
very distinctive willow species, was found on the shore. A brief foray into a freshwater tidal
marsh was quite productive, with large stands of Nuphar advena, Peltandra virginica,
Sagittaria latifolia, Pontederia cordata, Polygonum punctatum, Amaranthus cannabinus, Bidens
laevis, Heteranthera reniformis, Zizania aquatica, and other species typical of the habitat. Old
dredge spoil produced Scirpus pungens, Cyperus erythrorhizos, C. strigosus, Eleocharis ovate,
Alisma subcordatum, and Lindernia dubia. Two trees of interest were Acer pseudoplatanus and
Ulmus procera. Leader: Karl Anderson.

1999 FIELD TRIPS 173

4 September: Cedar Lake Fish and Wildlife Management Area, Monroe Township,
Gloucester County, and Buena Township, Atlantic County, NJ. Members gathered in the
sand parking lot south of Jackson Road, immediately south of Cedar Lake’s spillway to hear
the leader give a description of the lake and its history. Because of drought, there was
reduced surface water flow into the lake, thus leaving the pond bottom exposed, providing
a nearly water-free area to survey. We were able to walk along the narrow stream channel
of the original pre-dam watercourse. Our expedition followed the eastern shoreline to the
head of the pond, returning to the parking lot briefly to explore the western, smaller arm
of Cedar Lake. The exposure of the sandy organic lake sediments provided a great
opportunity for emergent graminoids. Normally, when the pond is full of water, only the
tips of Juncus militaris are exposed. Drought provided habitat for Juncus canadensis as well.
Between the dominant rushes were numerous noteworthy species. Most of the exposed
sediments supported a dense mat of Rhynchospora (Psilocarya) scirpoides from the area of the
dam to the head of the lake. Among this immense beaked-rush population were colonies of
Xyris smalliana, Scleria reticularis var. pubescens, Rhynchospora chalarocephala, and Ludwigia
sphaerocarpa. After a brief lunch we explored the western arm of the lake, finding similar
conditions in this separate drainage. The upper edge of the lake truncates into an Atlantic
white-cedar swamp too dense for the entire entourage to visit. We ended our day in the
exposed bottom of this portion of the lake, noting the high density of Rhynchospora
chalarocephala to the exclusion of many other plants. Leader: Joe Arsenault.

18 September: Roebling Memorial Park and Duck Island of the Hamilton-Trenton Marsh,
Mercer County, NJ. Joint trip with the Torrey Botanical Society. Blooming at Roebling
Memorial Park were Ambrosia artemisiifolia, A. vulgaris, Bidens frondosa, Chenopodium
album, Chrysanthemum leucanthemum, Cirsium arvense, Commelina communis, Daucus carota,
Erechtites hieraciifolia, Eupatorium rugosum, E. serotinum, Helenium autumnale, Impatiens
capensis, Lathyrus latifolius, Linaria vulgaris, Lycopus americanus, Lythrum salicaria, Oenothera

ennis, Oxalis stricta, Plantago lanceolata, Polygonum amphibium, P. arifolium, P. caespitosum,
P. cuspitatum, P. punctatum, P. hydropiperoides, P. pensylvanicum, Pontederia cordata,
Sagittaria latifolia, Taraxicum officianale, Trifolium pratense, Verbascum thapsus, and Zizania
aquatica. At our lunch spot at the Trenton Marina boat dock north of Duck Island, we saw
in bloom the rare plant Bidens bidentoides (southern estuarine beggar-ticks) along with
Chenopodium ambrosioides, Helenium autumnale, and Vernonia noveboracensis. A sampling
of the species blooming at Duck Island, a constructed freshwater tidal wetland site, included
Aster puniceus, Bidens laevis, Helianthus tuberosus, Mikania scandens, Perilla frutescens,
Polygonum arifolium, P. punctatum, P. sagittatum, Pontederia cordata, Solanum nigrum, and
Vernonia noveborecensis. Other species were Amaranthus cannabinus, Carex grayi, Eleocharis
acicularis, the threatened mud-plantain Heteranthera multiflora, H. reniformis, Myriophyllum
Spicatum, Penthorum sedoides, Potamogeton pectinatus, Sagittaria subulata, and an alga species
of Vaucheria. Attendance: 15, 10 of whom were students in a wetlands course from the
University of Pennsylvania. Report prepared by Patrick Cooney. Leaders: Mary and
Charles Leck.

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Bartonia No. 61, 2002

Program of Meetings
2000-May 2002

Date Subject Speaker
2000
27 Jan = Phyllanthus (Euphorbiaceae) in Hepatitis Virus ...........------- David Unander
24Feb A Look at the Potomac Gorge and its Flora ................ Robert G. Johnsson
23 Mar Selected Plant Communities and Flora of the Engadine Region of Switzerland

PRL eae ORS BEANS EROME SOG Oke Ga RAR ESTE Re A Oe Ted Gordon
27 Apr Retirement Party for Alfred “Ernie” Schuyler

Peer Pee eee eres rer eT ee Morris Arboretum of the University of Pennsylvania
a taay Ware Visats of Mocks County 2... 45s se eas8¥o54 ee ess eee Ann
28 Sep Members’ Reports on Summer Botanizing
26 Oct Plants, Pests, Poisons, and Pot—Forensic Botany ...........-+--. Meredith A. Lane
16 Nov Distribution of Aquatic and Wetland Plants in Rihirion to Land Use in Mullica

Piver Dae Siar Sk cS wied Pak eee eee ee ae tert Kim Laidig

14. Dec Botanical Management Strategies in Fairmount Park

a greene et a eaehs Richard J. Horwitz, Alfred E. Schuyler, and Bradley W. Thompson
2001
25 Jan The Versatile Bartrams and Their Enduring Botanical Legacy ..... Alfred E. Schuyler
22 Feb The Sarah P. Duke Gardens: A Jewel in the University’s Crown ... Richard A. White
22 Mar Acanthaceae and Other Interesting Plants of South Africa ...... Lucinda A. McDade
26 Apr Science, Policy, and Management of an Invasive Plant, Purple Loosestrife . . . Erik Kiviat
24 May Highlights on Botanical History and Exploration of Quebec-Labrador Jacques Cayouette
27 Sep Members’ Reports on Summer Botanizing
ees. ane t peniint Ponte so as ga ek ek a ees wens 2 od James A. Macklin
1 Nov - More South Florida Goodies 2.2... 0.0 cece cee eerste nsees Stevens Heckscher
20Dec Carex polymorpha and other Botanical Treasures of the Saigon avs Creek Valley,

Luzerne County, Pennsylvania ............ F. Rhoads and Timothy A. Block

2002
24Jan A Review of Witmer Stone’s Contributions ie DOM i Sos es ware es Gerry Moore
me Feb Sex Life of Amaraniims cannabinus 02000 cece Margo Bram
28 Mar New York Metropolitan Flora Project ..........-++++++0005 Steven E. Clemants
25 Apr Evolution and Distribution of Dendrobium Orchids ...........-- Howard P. Wood
23 May Limits to Restoring Native Woodlands to Urban Lands .......... Steven N. Handel

175

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Bartonia No. 61, 2002

2001-2002 Membership List

Life Members

ARSENAULT, JOE — 961 Clark Ave., Franklinville, NJ 08322, njplants@aol.co

GREENLAND, CHRISTINE MANVILLE — 790 E. Street Rd., Warminster, PA 13974, 215-322-4105

HOLT, ROBERT J. — 3032 Taft Rd., Norristown, PA 19401, 610-584-5578

IRETON, MARY LOU — 213 4th Ave., Haddon Heights, NJ 08035, 856-547-1118

LAUER, DAVID — 49 Cornell Ave., Clanchivilic, PA 18966, 215-357-2646, dml100@aol.com

MCLEAN, ELIZABETH — 139 Cherry Ln., Wynnewood, PA 19096, 610-642-4196

MCLEAN II, WILLIAM — 139 Cherry Ln., Wynnewood, PA 19096, 610-642-4196

Moore, A. GERRY — 367 E. 201st St., Apt. 6B, Bronx, NY 10458, 718-623-7332,
gerrymoore@bbg.org

O’HERRON, JOHN — 220 Washington St., Mount Holly, NJ 08060, 609-261-0711,
joheerron@voicenet.com

OLSON, WILLIAM — 1005 Lakewood-Farmingdale Rd., Howell, NJ 07727, 732-938-3187,
bolson@maserconsulting.com

PATRICK, RUTH — 750 Thomas Rd., Philadelphia, PA 19118

ROBERTS, WILLIAM H. — 1922 Rittenhouse Sq., Philadelphia, PA 19103, 215-569-5632

RYAN, NANCY PETERS — 2355 Oakdale Ave., Glenside, PA 19038

STAILEY, HELEN M — 8701 Macon St., Philadelphia, PA 19152, 215-673-8163

STEVENS, CHARLES E. — 615 Preston Pl., Charlottesville, VA 22903, 804-293-8658

THOMPSON, SUE — 129 E. Sycamore St., Pittsburgh, PA 15211, 412-622-3295

TREADWAY, SUSAN — 1509 Monk Rd., Gladwyne, PA 19035, 610-642-8050, sptreadway@aol.com

Honorary Member

FARLEY, Anema B. — 319 Bala Ave., Bala Cynwyd, PA 19004, 610-667-0625

SCHUYLER, ALFRED E. — Academy of Natural Sciences, 1900 Benj. Franklin Pkwy., Philadelpiha,
PA 19103, 215-405-5088, schuyler@acnatsci.org

Sponsoring Members

FRICK, JULIA — Blair 213, 1400 Waverly Rd., Gladwyne, PA 19035, 610-645-8863

LIBBY, VALENCIA — 607 Cloverly Ave., Jenkintown, PA 19046, 215-576-5725

TAYLOR, LANE — 207 W. Gravers Ln., Philadelphia, PA 19118, 215-247-2890, lraylor@stradley.com

Regular Members

ALDHAM, ALBERT — 1660 Hemlock Farms, Hawley, PA 18428, 570-775-6773,
bertaldham@webtv.net

ALLEVA, DENISE — 1071 Penn Circle, Apt. G-312, King of Prussia, PA 19406, 610-783-6799,
denise.alleva@sanofi-synthelabo.com

AMOs, SANDRA — 41 Laurel, Clementon, NJ 08021, 856-346-2242

ANDERSON, KARL — Rancocas Nature Center, 794 Rancocas Rd., Mt. Holly, NJ 08060, 609-267-
2195, rancoc@bellatlantic.net

ASPELING, ROGER — P.O. Box 10, Titusville, NJ 08560, 609- pie roissy@juno.com

BAKER, BRAD — P.O. Box 662, Unionville, PA 19375, 610-347-064

BALDWIN, DON — 4240 Fairview Ave., Newtown Square, PA oom, 610-353-1550

BAUCHSPIES, JAMES — 4320 Chetwin Terr., Easton, PA 18045, 610-253-8925

BIDDLE, DORRELL — 701 Washington Ave., Palmyra, NJ 08065, 856-829-0748, dell2733@aol.com

177

178 BARTONIA

BIEN, WALTER — 144 Summit Ave., Langhorne, PA 19047, 215-752-3762, walterb988@aol.com

BLACK, ROBIN R. — 792 Woodlea Rd., Rosemont, PA 19017

BLOCK, TIMOTHY — 1234 Pine Ridge Dr., Perkiomenville, PA 18074, 215-234-0645,
block@pobox.upenn.edu

BOWELL, MICHAEL — 2148 Bodine Rd., Malvern, PA 19355, 610-827-1268, mwb@createascene.com

BOYD, HOWARD — 232 Oak Shade Rd., Tabernacle, NJ 08088, 609-268-1734

BRINTON, EDWARD — 896 Roundelay, West Chester, PA 19382, 610-793-1582,
ravinehouse@aol.com

BROTHERSON, ROBERT — P.O. Box 179, Revere, PA 18953, 610-847-1005, nellsboy@epix.net

BUCK, WILLIAM — New York Botanical Garden, Bronx, NY 10458, 718-817-8624, bbuck@nybg.org

CAIAZZA, NICHOLAS — 5 Dorothea Terr., Lawrenceville, NJ 08648, ncaiazza@gateway.net

CARR, DEBBIE — 314 E. Highland Ave., Philadelphia, PA 19118, 215-242-0734,
wechslercarr@earthlink.net

CHELSVIG, GORDANA — 308 Woodbine Ave., Narberth, PA 19072, 610-667-2581, gchelsvg@fast.net

CooK, BUD — HC 1, Box 1117, Blakeslee, PA 18619

COONEY, PATRICK — 221 Mt. Hope Blvd., Hastings-on-Hudson, NY 10706, 914-478-1803,
plcooney@aol.com

COURTNEY, JOHN — 439 Gladstone Ave., Haddonfield, NJ 08033, 856-429-4987

CRICHTON, OLIVER — 726 Loveville Rd., Ctg. 5, Hockessin, DE 19707, 302-235-0571,
crichton@magpage.

DANZENBAKER, JAY — 607 Cloverly Ave., Jenkintown, PA 19046, 215-576-5725,
vlibby@temple.edu

DAVIS, CHARLES — 1510 Bellona Ave., Lutherville, MD 21093, 410-252-4154, cadavis@bcpl.net

DECASTRO, LINDA — 1100 Concord Dr., Bridgewater, NJ 08807, Idecastr@rvcc.raritanval.edu

DENNE-HINNOV, BOEL — 1-D eee Apt.s., Faculty Rd., Princeton, NJ 08540, 609-921-1272,
boel_denne-hinnov@hmco

DENNY, GUY — 6021 Mt. Gilead Rd., Fredericktown, OH 43019, 740-694-6087, guydenny@ecr.net

DIEDRICH, ARMON — 502 Highland Terr., Pitman, NJ 08071, 856-589-8455

DOoDDs, Ju — 56 Tumble Falls Rd., Stockton, NJ 08559, 908-996-3214, jsdodds@worldnet.att.net

DORN, RUTH — 800 Trenton Rd., Apt. 244, Langhorne, PA 19047, 215-702-1433,
ruthdorn@hotmail.com

DRAUDE, TIMOTHY — 415 Poplar St., Lancaster, PA 17603, 717-393-7233

ere MOIRA CLARE — 316 Wright St., Philadelphia, PA 19128, 215-487-7515, duggan@bbs-
aw.com

EBERT, JANET — 394 Smith Bridge Rd., Chadds Ford, PA 19317, 610-459-0585

ECKERT, GREG — 137 W. 2nd Ave., 1st Fl., Conshohocken, PA 19428, 610-834-5450,
geckert@tnc.or

EDINGER, GREG — 442 Ryan Rd., Greenwich, NJ 12834, 518-692-1725, gedinger@tnc.org

EIGENRAUCH, JANE — P.O. Box 85, Red Bank, NJ 07701, 732-842-0690

EVANS, JANET — Library, Pennsylvania Historical Society, 100 N. 20th St., Philadelphia, PA
19103, 215-988-8779, jevans@pennhort.org

FARNON, CHRISTA — 1418 Surrey Ln., Wynnewood, PA 19096, 610-649-2668,
christa.farnon@drexel.edu

FARRELL, JESSIE — 366 Chatham Rd., West Grove, PA 19390, 610-869-4285,
jessie@taprootnativedesign.com

FEDERICI, ANTONIO — 911 19th St., Union City, NJ 07087, 201-864-8992,
tony_federici@golder.com

FIELD, STEPHEN — 5 Evelyn Ave., Vineland, NJ 08360, 856-691-5868

FIELD, THERESA — 5 Evelyn Ave., Vineland, NJ 08360, 856-691-5868

FINE, HELEN — 73 Nature Ln., Sewell, NJ 08080, 856-218-4809, handnfines@gateway.net

FINE, NORMAN — 73 Nature Ln., Sewell, NJ 08080, 856-218-4809, handnfine@home.com

FINN, ANDREA — 142 W. Thomas Ct., Kennett Square, PA 19348, 610-388-3542

MEMBERSHIP LIST 179

FLANIGAN, TONI — 662 W. Johnson St., Philadelphia, PA 19144, 215-951-9211,
ccooper@critpath.or

FLISSER, DANIEL — Dept. of Biology, Camden County College, P.O. Box 200, Blackwood, NJ
08012, 856-227-7200, dflisser@camdencc.edu

FOGARASI, KASIA — 317 Roxborough Ave., Philadelphia, PA 19128, 215-482-3835,
kasiankal@aol.com

FREYBURGER, HELEN — 5258 34th Ave. N., St. Petersburg, FL 33710, 747-526-1579

FUSSELL, CATHARINE — 179 Kendal Dr., Kennett Square, PA 19348, 610-388-6523

GARBACK, MARY — 3839 Janice St., Philadelphia, PA 19114, 215-332-7105

GEHRIS, ANNA — 1133 Cameo-Shell Pt. Vlg., Fort Myers, FL 33908, 941-466-6407

GLASS, AMELIA — 135 Washington Ave., Pitman, NJ 08071, 856-589-6435

GODDARD, PAUL — 127 N. Van Pelt St., Philadelphia, PA 19103, 215-557-0187

GOFF, ELINOR — 791 College Ave., Apt. 1, Haverford, PA 19041, 215-247-5777

GOOD, NORMA — 745 Redman Ave., Haddonfield, NJ 08033, 856-428-1396

GORDON, PAT — 31 Burrs Mill Rd., Southampton, NJ 08088, 609-859-3566

GORDON, TED — 31 Burrs Mill Rd., Southampton, NJ 08088, 609-859-3566

GRACIE, CAROL — 19 N. Lake Circle, S. Salem, NY 10590, 914-763-5938, cgracie@optionline.net

GRAY, DENNIS — P.O. Box 206, New Lisbon, NJ 08064, 609-894-8849, gray@imcs.rutgers.edu

GREMBOWICZ, DUANE — 100 Motor Rd., Pine Beach, NJ 08741, 732-349-0364,
polarstar@adelphia.net

GRIMES, BRUCE G. — P.O. Box 222, Sumneytown, PA 18084, 215-234-8424, taxkwaxak@cs.com

GRINER, JOSEPH — 25 Millchase Rd., Southampton, NJ 08088, 609-859-9251, joemaggi@netaxs.com

GROSS, MICHAEL F. — Biology Dept., Georgian Court College, 900 Lakewood Ave., Lakewood,
NJ 08701, 732-364-2200, gross@georgian.edu

GYER, JOHN — P.O. Box 185, Clarksboro, NJ 08020, 856-423-3889, fernhill@voicenet.com

HALLIWELL, TOM — 19 Kings Rd, Netcong, NJ 07857, 973-347-6071, tbhalliwell@worldnet.att.net

HAMILTON, MARSHALL — 18 Lakewood Dr., Media, PA 19063, mham.12741@aol.com

HAMMERSTROM, FREDERICA — 542 W. Montgomery Ave., Haverford, PA 19041, 610-649-3811

HARDESTY, GAIL — 488 Big Oak Rd., Morrisville, PA 19067, 215-295-4734, evergreenl1@aol.com

HARPEL, WILLIAM.— 150 E. Winchester Ave., Langhorne, PA 19047, 215-752-0802

HARRIS, JESSIE M. — 4401 W St. N.W., Washington, DC 20007, 202-338-9083, jes405@hotmail.com

HART, ROBIN L. — 5086 Barrington Circle, Sarasota, FL 34234, 941-351-1554

HASSEL, LLOYD V. — 45 Danbury Rd., Lancaster, PA 17601, 717-569-2368

HAUPT, ANDREA — 639 W. Ellet St, Philadelphia, PA 19119, 215-438-8175, reh215@aol.com

HAWK, JEFFREY — 16 Andrea Ln., Hamilton, NJ 08619

HECKSCHER, STEVENS — Natural Lands Trust, 1030 E. Lancaster Ave., Apt. 314, Rosemont, PA
19010, 610-527-6324, heckscher@natlands.org

HEGARTY, DAN — P.O. Box 1244, Chadds Ford, PA 19317, 302-652-4148,
dhegarty@mindspring.com

HELD, MICHAEL ~ Dept. of Biology, St. Peter’s College, Jersey City, NJ 18301, 201-915-9187

HERMELY, ALAN — 671 Moore Rd., King of Prussia, PA 19046, 610-337-3666,
alan_hermely@urscorp.com

HENRY FOUNDATION FOR BOTANICAL RESEARCH, 801 STONY LN., GLADWYNE, PA 19035, 610-
525-2037

HEWITT, DAVID A. — Harvard University, Cambridge, Massachusetts 02138,
Dhewitt@oeb.harvard.edu

» FRANKLIN S. — 5004 Little Mill Rd., Stockton, MD 21864, 410-632-1362

HUGHES, STUART W. — 721 Butler Pike, Maple Glen, PA 19002, 215-646-3873, stuhu@aol.com

HUMBERT, KAREN C. — 2401 Pennsylvania Ave., 7A6, Philadelphia, PA 19130, 215-232-0295

HUNT, LYNN F. — P.O. Box 906, Tuckerton, NJ 08087, 609-296-8022

HUTCHEON, DAVID J. — 25 Caledonia Dr., Warminster, PA 18974, 215-957-0976

INSKEEP, MICHAEL — 113 Cricket Ave., #104, Ardmore, PA 19003, 610-896-0777,

180 BARTONIA

mike@gentlehelpers.com
JANOSKI, JEFFREY F. — 1801 Buttonwood St., #1610, Philadelphia, PA 19130, 215-569-1949,
jjfeh@aol.com

JESS, ROBIN — 55 Lahiere Ave., Edison, NJ 08817, 732-572-5928, c

JOHNSON, ALAN A. — 63 Central Ave., Audubon, NJ 08106, 856-547-3498,
aajohnson@bigfoot.com

JOHNSSON, FANNY M. — 7422 Ridge Rd., Frederick, MD 21702, 301-371-5215,
johnsson@starpower.net

JUELG, J. RUSSELL —

KAISER, GEOFFREY D. — P.O. Box 222, Sumneytown, PA 18084, 215-234-8424,
taxkwaxak@enter.net

KAPLAN, PAULA WEST — 1085 Huntingdon Rd., Abington, PA 19001, 215-884-4829,
pwk@home.com

KELLER, ELIZABETH — 114 Wyomissing Blvd., Wyomissing, PA 19610, 610-374-3458,
zab12345@aol.com

KELLY, LINDA — 60 Forest Dr., Lakewood, NJ 88701, 732-363-1266, kellylcO1@aol.com

KENDIG, JAMES W. — 1212 S. Prince St., Palmyra, PA 17178, 717-832-3899

KIMELMAN, GAY — 2212 St. James Pl., Philadelphia, PA 19103, 215-563-0285

KLOTZ, LARRY — Biology Dept., Shippensburg University, 1871 Old Main Dr., Shippensburg, PA
17257, 717-477-1402, lhklot@ark.ship.edu

KOERBER, JR., WALTER A. — 1380 Valley Green Rd., Etters, PA 17319, 717-938-9618,
wakoerber@hotmail.com

KOLAGA, VAL — 186 Dilworthtown Rd., West Chester, PA 19382, 610-399-3136, sfox@axs2000.net

KRAIMAN, CLAIRE T. — 7 Chip Ln., Reading, PA 19607, 610-775-9737

KUNKLE, KARL — 507 Nina Ln., Bear, DE 19701, 302-834-2802, kkkunkle@aol.com

LADEN, MILTON — 1919 Chestnut St., #1407, Philadelphia, PA 19103, 215-568-6599

LAMONT, ERIC — 717 Sound Shore Rd., Riverhead, NY 11901, 631-722-5542,
elamont@hamptons.com

LATHAM, ROGER — P.O. Box 57, Wallingford, PA 19086, 610-565-3405, rogerlatham@dca.net

LEA, CHRISTOPHER — 202 Washington St., Berlin, MD 21811, 410-641-1443, chris_lea@nps.gov

LECK, CHARLES — 105 Kendall Rd., Kendall Park, NJ 08824, 732-821-8310

LECK, MARY — 105 Kendall Rd., Kendall Park, NJ 08824, 732-821-8310

LEFEVRE, WILLIAM M. — Historic Bartram’s Gardens, 13 Hickory Dr., Doylestown, PA 18901,
267-880-0860, wmlefevre@aol.com

LEVIN, MICHAEL H. — 414 Mill Rd., Havertown, PA 19083, 610-449-7400, mhl6841468@aol.com

LEWANDOWSKI, RICK J. — Mt. Cuba Center, P.O. Box 3570, Greenville, DE 19807, 302-239-7980

LIGHTY, RICHARD — 501 Chandler Mill Rd., Kennett Square, PA 19348, 610-444-2987

LITTLETON, JOHN E. — 331 Station Rd., Wynnewood, PA 19096, 610-642-6755,
jlittleton@home.com

LOEFFLER, CAROL — Dept. of Biology, Dickinson College, P.O. Box 1773, Carlisle, PA 17013,
717-245-1360, loeffler@dickinson.edu

siete MICHAEL — 2028 Fitzwater St., Philadelphia, PA 19146, 215-732-0849

MACKLIN, JAMES — Academy of Natural Sciences, 1900 Benj. Franklin Pkwy., Philadelphia, PA
“aes 215-405-5088, macklin@acnatsci.org

MARTIN, HARRIS — 5100 Bay Rd., Bensalem, PA 19020, 215-639-3686, martin68@atc-enviro.com

MAURICE, KEITH R. — 63 Union Ave., New Holland, PA 17557, kmaurice@normandeau.com

McAvoy, WILLIAM A. — 4876 Haypoint Landing Rd., Smyrna, DE 19977, 302-653-2880,
wmcavoy@state.de.us

MCCoMBE, ROBERT — 106 Center St., Manahawkin, NJ 08050, 609-597-7107,
rob.mccombe@worldnet.att.net

McCouRT, RICHARD — Academy of Natural Sciences, 1900 Benj. Franklin Pkwy., Philadelphia,
PA 19103, 215-299-1157, mccourt@acnatsci.org

MEMBERSHIP LIST 181

McCREA, ESTHER J. — 70 Overhill Rd., Bala Cynwyd, PA 19004, 610-664-1642

McDADE, LUCINDA — Academy of Natural Sciences, 1900 Benj. Franklin Pkwy., Philadelphia, PA
19103, 215-405-5087, mcdade@acnatsci.org

MCLAUGHLIN, WILLIAM — 5 Oak Dr., Tabernacle, NJ 08088, 609-268-1054, wmcl268@aol.com

MELENDEZ, MARISA — 1005 Lakewood Farmingdale Rd., Howell, NJ 07731, 732-961-1946,
marisabel626@hotmail.com

MELLON, RICHARD — 200 Flint Ct. S., Yardley, PA 19067, 215-493-0697, rmellon@voicenet.com

MICHAEL, DAVID — 313 Green St., Philadelphia, PA, 215-923-6163, dmichael@pennhort.org

MICKLE, ANN — Dept. of Biology, Lasalle University, 1900 W. Olney Ave., Philadelphia, PA
19141, 215-951-1254, mickle@lasalle.edu

MILNER, NORMA — 1131 eee Dr., Palmyra, NJ 08065, 856-829-3142,
normamartinmilner@juno.c

MONSHAW, HARRIET — 176B mn Ln., Cherry Hill, NJ 08003, 856-428-2342,
v.r.monshaw@ieee.org

MONTGOMERY, - — Ecology II, 804 Salem Blvd, Berwick, PA 18603, 570-542-2191,
jdm@sunlink.n

MOOBERRY, F. M_ — 111 Spotswood Ln., Kennett Square, PA 19348, 610-444-5495

MOorE, JULIA — 91 Molyneaux Rd., Camden, ME 04843

Morris, MELISSA — 234 Broughton Ln., Villanova, PA 19085, 610-525-6659, mhmorris@umich.ed

Morse, LARRY — The ap a 4245 N. Fairfax Dr., Ste. 100, Arlington, VA 27903,
og 841- 5361, lmorse@tnc.o

Moss, MIRIAM — 8120 Brookside Rd., Elkins Park, PA 19027, 215-635-0176

NACZI, ROBERT — Dept. of Agriculture & Natural Resources, Delaware State University, Dover,
DE 19901, 302-857-6450, rnaczi@dsc.edu

NEWBOLD, ANN — 50 Renninger Rd., Bechtelsville, PA 19505, 610-754-7573

NEWSTEAD, CHARLOTTE — 270 Roseland Ave., Essex Fells, NJ 07021, 973-226-7651

NEWSTEAD, EDWIN — 20 Roseland Ave., Essex Fells, NJ 07021, 973-226-7651

NICHOLS, RAY — 412 Federal City Rd., Pennington, NJ 08534, 609-737-7442, nichols@nj1.aae.com

O’BRIEN, TERRY — Dept. of Biological Sciences, Rowan University, Glassboro, NJ 08028, 856-256-
4500, obrien@rowan.edu

PALMER, MATTHEW — 1518 Millstone River Rd., Hillsborough, NJ 08844, 908-359-2073,
mipalmer@eden.rutgers.edu

PARKS, JAMES — 103 Bender Mill Rd., Lancaster, PA 17603, 717-872-5206,
james.parks@millersville.edu

PAYNE, ELISE — 643 Fernfield Circle, Wayne, PA 19087, 610-688-4377, erpayne@erols.com

LYLER, D. — 18 Bridle Path, Chadds Ford, PA 19317, plylerd@aol.com

PREUCEL, RUTH — 1147 Norsam Rd., Gladwynne, PA

QUIGLEY, PATRICIA — 1080 Quarry Hall Rd., Norristown, PA 19403, 610-584-1829,
paqinc@aol.com

RADIS, RICHARD — 69 Ogden Ave., Rockaway, NJ 07866, 973-586-0845, isotria@bellatlantic.net

RHOADS, ANN — 3 Blythewood Rd., Doylestown, PA 18901, 215-348-8139,
rhoadsaf@pobox.upenn.edu

RHODES, CHARLES — mw Stonycreek Ave., Lansdale, PA 19446, 215-368- 9591,
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RISKA, MICHAEL — te Nature Society, Box 700, Hockessin, DE 19707, 302-239-2334,

ike@dnsashland.org

ROCHE, LESLIE — 73 Hull Ave., Freehold, NJ 07728, 732-780-0121

ROMANO, GABRIELLA — 4240 Fairview Ave., Newtown Square, PA 19073, 610-353-1550

ROSENBERG, ANN — 5 Westview Rd., Bryn Mawr, PA 19010, 610-525-8683

ROSENBERG, DICK — 5 Westview Rd., Bryn Mawr, PA 19010, 610-525-8683

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182 BARTONIA

RUCH, PAMELA — 542 Liberty St., Emmaus, PA 18049, 610-966-2635, pamruch@hotmail.com

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RUSSELL, EMILY — P.O. Box 430, Mount Tabor, NI oO 973-625-3382, erussell@nac.net

RYAN, JOEY — 521 S. Narberth Ave., Merion, PA 1

SALGANICOFF, MATILDE — 556 N. 23rd St., eh uich, PA 19130

SCHAEFFER, ROBERT — 1940 W. Turner St., Allentown, PA 18104

SCHNEIDER, GEORGE W. — 345 Nursery Rd., Wellsville, PA 17365, 717-292-4035

SCHNEIDER, WILLIAM — 228 Canterbury Dr., West Chester, PA 19380, 610-431-2449,
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SCOTT, JOHN — 55 Hertzog School Rd., Mertztown, PA 19539, 610-682-2809,
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SCOTT, STELLA — 3316 Arch St., Philadelphia, PA 19104, 215-387-2698, stelsco@yahoo.com

SEAGER, KEITH — 278 Fishing Creek Rd., Cape May, NJ 08204

SEIFFERT, JUDITH — 1135 E. Oxford St., Philadelphia, PA 19125, 215-426-4201

SEIPLER, MARYJANE — 4200 Tamarack Ln., Murrysville, PA 15668, 724-325-3571,
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SHERMAN, LYNN — P.O. Box 391, Ravena, NY 12143, 518-756-5028, sherman|l@capital.net

SLANE, JOSEPH — 1207 Shackamaxon St., Philadelphia, PA 19125, 215-634-6332,
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SLATER, MIKE — R.R. 5, Box 5820, Mohnton, PA 19540, 610-775-3757

SMITH, CRAIG — 728 Seymour Rd., Bear, DE 19701, 302-324-9486, csmith6666@aol.com

ST. JOHN, MICHAEL — 4 Perry St., New York, NY 10014, misaintjon@aol.com

STAHL, STANLEY — 455 Garland Circle, Lancaster, PA 17602, 717-293-0292,
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STALTER, RICHARD — Dept. of Biology, St. John’s University, Jamaica, NY 11439, 718-990-6269

STANDAERT, WILLIAM — 45 Maltbie Ave., Apt. 19B, Midland Park, NJ 07432, 201-612-9069,
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STECKEL, CLAUDIA — 423 N. 27th St., Allentown, PA 18104, 610-740-0141

STECKEL, DAVID — 423 N. 27th St., Allentown, PA 18104, 610-740-0141

STUCKEY, RONALD — 1315 Kinnear Rd., Columbus, OH 43212, 614-292-6095

SWEENEY, BILL — Lockhouse, 393 Island Park Rd., Easton, PA 18042, 610-253-7053

SWEETMAN, HAROLD — Jenkins Arboretum, 631 Berwyn Baptist Rd., Devon, PA 19333, 610-647-
8870

TEESE, PAUL — Bowmans Hill Wildflower Preserve, 1183 Apple Rd., Quakertown, PA 18951, 215-
536-3719, teese@bhwp.org

TOFFEY, WILLIAM — 9001 Verree Rd., Philadelphia, PA 19115, 215-969-6789, wtoffey@prodigy.net

REXLER, LARRY — 7501 Germantown Ave., Philadelphia, PA 19119, 215-242-1309

TREXLER, so — 7501 Germantown Ave, Philadelphia, PA 19119, 215-242-1309

TUCKER, ARTHUR — Dept. of Agriculture & Natural Resources, Delaware State University,
Dover, DE es 302-857-6408, atucker@dsc.edu

TYNDALL, WAYNE — 15245 Oakland Rd., Goldsboro, MD 21636, wtyndall@intercom.net

UDELL, VAL — 2746 Yost Rd., Perkiomenville, PA 18074, 610-754-7163

VERLENDEN, DONALD — P.O. Box 116, Lansdowne, PA 19050

VOLLMER, JOHN — 42 Burrs Mill Rd., Southampton, NJ 08088, 609-859-2805

WHITING, GEORGE — 116 Spring Mount Rd., Schwenksville, PA 19473, 610-287-6397

WIEBOLDT, THOMAS — 155 Shady Grove Ln., Newport, VA 24128, 540-544-7967, wieboldt@vt.edu

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WILLIAMS, CARL — Box 347, Hereford, PA 18056, 215-234-0545

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WILLIAMS, DAVID — 61 Coppermine Rd., Princeton, NJ 08540, 732-297-0642, idwill@superlink.net

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WILSON, RONALD — 3740 Ridge Rd., Snow Hill, MD 21863, 410-632-3892, rwilson@dmv.com

WINDISCH, ANDREW — P.O. Box 312, Chatsworth, NJ 08019, 609-726-9054, windisch2@erols.com

WITMAN, DEANNA — 214 3rd St., East Greenville, PA 18041, 215-541-1789

WOLFF, EMILY — 295 E. Rose Tree Rd., Media, PA 19063, 610-566-6385

Woop, HOWARD — 3300 Darby Rd., C-802, Haverford, PA 19041, 610-642-9963,
howardpagewood@aol.com

WOODFORD, JEANNE — Wildlife Advocate, 2 Sawmill Rd., Medford, NJ 08055, 856-983-1617,
jeannewoodford@earthlink.net

ZAMPELLA, ROBERT — 12 Concord Dr., Shamong, NJ 08088, 609-268-3363,
zampella@njpines.statenj.us

ZAREMBA, ROBERT — 143 Hoyt St., Apt. 6-J, Stamford, CT 06905

ZOLLER, HEINRICH — 50 Renninger Rd., Bechtelsville, PA 19505

COIL

Instructions to Authors

Types of Articles Published
Research papers communicate original research in plant ecology, plant conservation biology, plant
systematics, and related topics. Other contributed papers convey the results of studies in floristics,
distribution, methods, biography, bibliography, history of botanical exploration, and other topics of
botanical interest. Short reports of one or two pages appear in “News and Notes.” Other items
include obituaries, book reviews, and field trip reports. The focus is on the mid-Atlantic region, but
contributions of interest to Bartonia readers from farther afield are welcomed.
Manuscript Style
Write in simple, clear sentences. Use the active voice where possible. Avoid redundancy. Bartonia
generally conforms to the Council of Biology Editors, Committee on Form and Style, CBE Style
Manual.
Consult recent issues of Bartonia for placement and style of main headings, subheadings, literature
cited, table and figure captions, and tables.
Double-space ail text, including tables.
Do not justify the right margin.
Arrange manuscript copy in this order:
ame, address, e-mail address, and telephone number of corresponding author
Running head (shortest intelligible version of title
Date of original manuscript submission
Title of paper
Name(s) of author(s)
Institutional address(es) of author(s)
Text of paper
tract
Introduction
Methods

Discussion (may include Conclusions)
Literature Cited
Acknowledgments
Tables (with captions)
Figure captions
igures

gu

Manuscript Submission and Review

Authors should send three, double-spaced, paper copies of the manuscript, with all tables and figures
(copies only of artwork with the first submission), to the Editor. Manuscripts with multiple authors
should be accompanied by a clear designation of the corresponding author, to whom
communication from the Editor should be addressed. The Editor will select at least two reviewers for
each manuscript and, on the basis of the reviews, decide on suitability for publication in Bartonia.

Submission of a manuscript implies that it is not under consideration in a similar form elsewhere.

The Editor will return manuscripts judged to require modification, along with suggestions for
revision, to the authors. Manuscripts not accepted for publication will be returned to the authors.

Authors who have been asked to resubmit a revised manuscript should make the requested revisions
and provide a brief rationale in a letter to the Editor for any reviewers’ comments not complied with.
Authors should send one double-spaced paper copy and an electronic copy (preferably as an e-mail
attachment; a mailed diskette or CD-ROM is also acceptable) of the revised manuscript to the Editor,
along with glossy prints or high-resolution scans in el ic f f any illustrations or photographs.
Formatted tables and illustrations must be sent as separate files, not embedded in the text

After a manuscript’s final acceptance, authors will receive page proofs with a request to proofread
and return them within 48 hours of their receipt. At this time, authors also will receive a price
schedule and order form for offprints and a page cost donation form, to be returned with payment to
the Philadelphia Botanical Club.