A NEW SPECIES OF BOLTONIA (ASTERACEAE) FROM THE RIDGE AND VALLEY PHYSIOGRAPHIC PROVINCE, U.S.A

2006·11.1 MB·English

by  John F Townsend

1
NEW
A OF BOLTONIA (ASTERACEAE) FROM THE
SPECIES
AND
RIDGE VALLEY PHYSIOGRAPHIC
PROVINCE,
U.S.A.
RTownsend
John Vesna Karaman-Castro
Department and
of Conservation Recreation Louisiana State University
Division of Natural j-leritage Department of Biological Sciences
2 7 Governor 3''' Floor 107 Science Building
St., Life
1
USA
Richmond, Virginia 232 USA. Baton Rouge, Louisiana 70803,
9,
Email: [email protected]
ABSTRACT
new
Boltonia montaiia, a species from depression wetlands and riverine habitats in the Appalachian Mountains of
New
Virginia, Jersey, and Pennsylvania, described. This taxon has most often been collected under the name
is
Boltonia asteroides (L.) L'Her. due to its relatively large heads which are usually few in number, and vegetative
new
similarities. Cypselae morphology of the taxon, however, dissimilar to that of B.asiewidcs and most closely
is
resembles Boltonia caroliniana (Walt.) Fern., a species characteristic of coastal plain and piedmont bottomlands
of the Carolinas and Virginia. Boltonia montana differs from the latter taxon in terms of inflorescence structure,
head dimensions, ray color, phyllary morphology and size, plant stature, and leaf shape. Evidence of its distinc-
tiveness provided and wetland habitat discussed.
is its is
RESUMEN
Se describe Boltonia montana, una nueva especie de las tierras humedas de depresiones y habitats ribereiios de los
Apalaches de Virginia, Nueva Jersey, y Pennsylvania. Este taxon se ha colectado la mayoria de las veccs con el
nombre de Boltonia asteroides (L.) EHer. Debido a sus cabezuelas relativamente grandes que estan usualmente en
un numero
pequeno, y similitudes vegetativas. La morfologia de las cipselas del nuevo taxon, sin embargo, es
diferente de la de B. asteroides y se parecen mucho mas a Boltonia caroliniana (Walt.) Fern., una especie
caracteristica de la llanura costera y base del pie de moiite de las Carolinas y Virginia. Boltonia montana difiere
de este ultimo taxon en termmos de estructura de la inflorescencia, dimensiones de la cabezuela, color de los
radios, morfologia y tamano de los filarios, tamano de la planta, y forma de la hoja. Se aportan pruebas de su
diferenciacion y se discute su habitat de zonas humedas.
INTRODUCTION
The was
first author alerted to the presence of a taxonomically problematic Boltonia in
Augusta County, Virginia (Ridge and Valley physiographic province) by Gary FlemingJ.
P.
Christopher Ludwig (Virginia Department of Conservation and Recreation, Division of
Natural Heritage), and Thomas Wieboldt (Massey Herbarmm, Virginia Polytechnic
F.
Institute) in late 2001. This plant had been identified as BoUonia asteroides by collectors,
but certam details of the plants, particularly those of cypselae and phyllaries, indicated
the Augusta County plants were not a good that taxon.
for
fit
Subsequent taxonomic herbarium and
investigations of the literature, material, living
from ponds
populations of Boltonia led to the realization that the plants sinkhole (dolines)
in the Virginia mountains possessed a unique set oi characters. During morphological
and molecular investigations of the Virginia plants, correspondence with the second author
led to the discovery of morphologically similar herbarium specimens collected from the
New
Ridge and Valley physiographic province northwestern and eastern Penn-
of Jersey
sylvania. Field studies in the early 1980s and late 1990s by David Snyder and Kathleen
New
Strakosch Walz of the Jersey Natural Heritage Program uncovered or reinventoried
much
and
the Boltonia populations in the northwestern part of that state provided data
SIDA 873 -886. 2006
22(2):
BRIIORG/SIDA
874 22(2)
on the plants discussed here. Subsequent herbarium and field investigations of the Vir-
New
ginia, Jersey, and Pennsylvania plants confirmed that these plants belonged to the
same
taxon. described here as nev^.
It is
&
V
Niw
Boltonia inontana Tovv^nsend Karaman-Castro, 2).Tmi
sp. nov. (Figs, 1, U.S.A.
J.F.
:
W
SW
km km
Sussex Co: Muckshaw pond complex, located 3.2 of Newton and of US 206,
JliRSllY. l."^
nearly dry, herb-dominated sinkhole pond, 3 Oct 2005 J_F Ihwnscnd 3570 UiOl OTYPE: US; [bO"l^ rv-\ MO,
NY,
VPl).
Bohonia asicroidcs LI primo adspcctu maxime simile ob capitulis magnus et paucus sed acheniis glabris,
(1..) lor.
alls et pappus aristis destituris a Boltonia cavoliuiana (Wait.) Fern, similis. Phyllariis costa angustara, herbaceous
distalis viridis.
et
dm
Glabrous, perennial herbs 1.2-15 ascending to erect (often prostrate in lower por-
tall,
tion to ca. 1/2 stem length, sometimes rooting at nodes in this section). Stems glabrous,
prommcnt
light to ycUowish-grecn, ribbed with yellow or golden striations (more in up-
per 1/2 ol plants), branching above into small, simple, corymbose panicles to diffusely
and
paniculate in larger plants. Leaves basal cauline, glabrous; basal leaves 0-18, alter-
nate, in a rosette, oblanceolatc to oblanceolate-spatulate, attenuated at the base (where
often slightly widened), dark green or blue-green, often whitish or reddish colored the
at
base, with prominent, lighter-colored midrib and inconspicuous, arcuate secondary veins,
cm cm
apices bluntly rounded, mucronulate to entire, 1-17.5 long, 0.1-2 wide; cauline
leaves oblanceolatc to obovate or spatulate, bases sessile, oiten partially clasping, apices
rounded lo acute, mostly with mucronulate tips, appearing entire but minutely serrulate
cm cm
with narrow, whitish, cartilaginous margin, 2.7-11 long, 0.5-1,9 wide, sometimes
with smaller leaves clustered at nodes. Capiiulescences corymbose-paniculate, few-headed
many-headed
(1-25) in average individuals, (-100+) in large individuals. Peduncles bracte-
mm
cm
ate, 0.5-11.8 long, 0.5-Ll thick, bractcal leaves oblanceolatc to obovate or oblong,
cm cm
tips obtuse to acute, mucronulate, 0-5 per pedicel, 0.1-97 long, 0.02-1.4 wide. In-
mm mm
volucre 5.2-7.8 6.0-11.3 wide; phyllaries 30-60, lancc-attenuate, serrulate to
tall,
mm mm mm
2-4
lacerate, in series, 2.5-4.4 long, 0.4-0.9 wide, midrib 0.12-0.3 wide (1/10-
1/5 of phyllary width), proximally whitish, distally green; receptacles conic, naked, 1.9-
mm mm
Ray
3.0 1.9-3.8 wide. flowers 27-46, pale lavender to pinkish (-white), corol-
tall,
mm mm
4-5 amber
las glabrous, nerved with golden or colored veins, 8.1-12 long, 0.8-1.9
mm mm mm
wide, tubes 0.45-0.95 long; styles 1.3-2.45 long, style branches 0.25-0.4 long,
narrowly with and
cypselae glabrous, oblong, triquetrous, adaxial 2 abaxial often
ribs,
1
mm mm
curved outward, wings
slightly 1.25-2.05 long, 0.5-0.9 wide, vestigial or absent,
mm mm
0-0.05 wide; pappus awns reduced to a crown of minute scales, 0-0.1(-0.4) long.
Disk flowers 220-350, pale yellowish green to pale yellow- wdiite as cypselae mature, su-
more
ture lines pale amber, disk appearing yellowish during anthesis, corollas glabrous,
mm mm mm
1.1-2.3 long, tubes 1.0-1.85 long, lobes 0.15-0.36 long; stamens included
to
mm mm
mm,
exceeding corolla length by 0.4 anthers 0.65-1.1 long, appendages long,
0.15
mm mm
styles 1.05-2.55 long, style branches 0.2-0.4 long, cypselae glabrous, narrowly
mm
oblong, strongly flattened laterally with two marginal ribs, 1.3-1.9 long, 0.55-1.25
mm mm
wings 0-0.09 pappus awns crown
wide, wide, usually reduced
vestigial, to a of
mm
minute
0-0.1
scales (-0.6) long.
DLSfnhution. —Distributed in sinkhole pond habitats and associated riversides and
streamsides of the Ridge and Valley physiographic province in western Virginia, north-
western Ncv/ Jersey, and eastern Pennsylvania.
TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN
U.S.A 875
5
i
6
3
?
Fig. 1.fio/fo/7/om(?/?ffl/7(7j.F.Townsend &V.Karaman-Castro. Illustration based on material collected in Sussex County, New Jersey. 1.
Ray 1mm
Habit. 2. cypselae. 3. Disk cypselae. 4. Head. 5. Outer phyllary(abaxial view). 6* Ray corolla. 7. Disk corolla. Scale bar equals
except in (= dm) and 4 {=1 cm).
1 1
876 BRIT.ORG/SIDA 22(2)
FiG.2. Low-growing individual of 5o/fort/(7/?Tonfa/)(7 with f/'/oct7u/o/?a(7Uflf/fiY/n,AugustaCounty,Virginia.Photo by Gary P.Fleming.
Etymology— name montane
T\\t refers to the species' habitats.
Phcno/ag_ydnddevc/opmen(.— Flowering prnnarily occurs from early August mid-
to
m
October with precocious plants flowering in especially during dry years or dry
July,
microhabitats. Hydrologic fluctuations in the species' sinkhole pond habitat can have a
when
dramatic effect on the timing o[ flowering in a popuhition. Even water levels re-
main high enough inundate these ponds during the growing season, seeds are capable
to
germination and may develop while under water (Lassen and Hamilton
ot rosettes still
comm. may
1 003; David Snyder pers. 2006) but the plants not develop further, and
if
may comm.
inundated for successive years the plants die (David Snyder, pers 2006). Au-
m
tumnal plants produce basal offsets nearly identical appearance to the rosettes pro-
duced by young plants. In the case of reclining or ascending plants of Bohonia moniana,
may many
autumn
form nodes
adventitious roots at the as approaches. In cases, these
nodes produce complete with During
also leafy offsets root systems. collection of flower-
weak
ing or Iruiting plants, these vegetative propagulcs are often dislodged due to their
attachment
the parent
to plant.
A
concentric gradient of environmental conditions created in these sinkholes by
is
fluctuating hydrology and sloping shorelines. therefore possible to find several co-
It is
occurrmg growth moniana.^dccni Newjersey with Natu-
stages ol Bollonia field w^ork in
ral Heritage Program Botanist David Snyder and Ecologist Kathleen Strakosch Walz (au-
tumn 2005) was conducted following a long period of pond inundation. On the pond
margins that were not recently inundated, relatively well-developed plants (w^th flow-
ers, fruits, and basal offsets) were found. In more recently inundated portions of the ponds,
numbers
large of Bohonia rosettes were found. These plants were either newly germi-
TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN 877
U.S.A.
nated seedlings, rosettes persisting from a previous germination event, or vegetative off-
sets novv^ independent of the parent plant. Regardless of origin, these rosettes were unable
to f lovv^er or produce fruit before the end of the growing season due to their exceptionally
when
late emergence (September-October). In general, pondshores are exposed as ground-
new
water levels drop after such periods of inundation, plants develop from the seed bank
comm.
(David Snyder, 2006).
pers.
Observations indicate that Smith and Mettler's (2002) description of the cycle of
life
&
Wood
the Federally Endangered Boltonia decurrens (Torr. A. Gray) provides a frame-
work for understanding the life cycle of Boltonia montana as well: "Although B. decurrens
may wmter
perennial under optimal conditions, various stages behave as annu-
is a life
summer
and summer), annu-
als (winter seedlings bolt, flower, senesce the following as
als (summer seedlings bolt, flower, and senesce the same summer), or as biennials (ca, 25
and
percent of spring seedlings overwinter as rosettes that bolt, flower, senesce the fol-
lowing summer)." Such a patchwork of potential strategies makes sense for a species
Midwestern
adapted to the highly variable flood pulse of rivers (B. decurrens) as well as
the dramatic hydrological fluctuations of isolated wetland systems montana).
(B.
AND
HABITAT DISTRIBUTION
known
Boltonia montana from localized sections of the Ridge and Valley physiographic
is
New
province in three U.S. states; Virginia (one county), Jersey (two counties), and Penn-
sylvania (one county). All occurrences are found within areas of gently sloping, rolling,
m
ground between 149 and 488 (490-1600 (Walz
or nearly level ft) altitude et al. 2001,
many New
Va. Division of Natural Heritage but of the Jersey ponds are surrounded
files),
comm.
by rough terrain with extensive bedrock outcrops (Kathleen V/alz, pers. 2006).
S.
The sinkhole pond complexes which support this species span an area measurmg
km km
New
roughly 14x 15 near Newton, Jersey and 5.25 x 4.5 near Waynesboro, Vir-
known
Within from ponds and
ginia. these areas, plants are a total of 36 sinkhole lakes
New
(22 in Jersey and 14 in Virginia; Va. Division of Natural Heritage David Snyder,
files;
comm. new come
pers 2006). Although the vast majority of the records for this taxon
from isolated wetlands, a very few have been found along rivers. Besides two collections
made the same day in 1936 along the South River in Virginia, the only other such collec-
were made 1864 specimen) and 1865 specimen) along the Susquehanna River
tions in
(1 (1
Dauphin
in County, Pennsylvania— apparently the only observations of this species in
Pennsylvania. The Virginia occurrence was taken from an area extremely close and
to,
known
just downslope from, a sinkhole pond complex to contain Boltonia montana, while
pond
the disposition of the Pennsylvania material relative to potential sinkhole seed
sources unclear. assumed here that only occasional plants become established in
is It is
such riverine habitats from nearby "parent" populations. The lack of additional collec-
such
tions at least indicates that riverine habitats are rarely occupied.
many
Like Boltonia montana, species seemingly defined by their isolated wetland
community
habitat are occasionally found in other types, indicating that their affinity
these ponds not Examples include the co-occurring Helenium rginicum
for restrictive. vi
is
Blake in Virginia, which almost entirely restricted to sinkhole ponds but also occurs in
is
meadow The
a wet of the nearby South River (Va. Division of Natural Heritage files).
diminutive sedge Fimhristylis perpusilla Harper ex Small and Britt.was also thought to
be a strict pond endemic and functions as such in most localities, but discoveries in North
and South Carolina (Leonard 1981, 1987), established the species as an occupant of river-
878 BRIT.ORG/SIDA 22(2)
A
ine "draw-down" bars, banks, and sloughs as well. final example involves Boltonia
decurrens, a perennial species of floodplains in the Illinois River valley (Stoecker et
al.
which was dependent on maintenance
1995) originally natural flood pulses habitat
for
and germination. Most of the habitats occupied by this species have been described as
wet prairies or bottomland communities of various types, but McClain (1997) also
et
al.
cported the species from "sand ponds" near the Illinois River in Cass and Mason coun-
r
m
ties, Illinois. They described water levels in the ponds as fluctuatmg response to the
known human
local aquifer rather than overbank flooding, but not what degree
to
is
it
manipulation of the river has lessened the importance of this water source in the ponds.
The sinkhole pond habitats occupied by Boltonia moniana are ultimately formed by
the solution of underlying carbonate bedrock and subsequent volume loss or collapse of
&
foundation (Fleming Van Walz The found
this Alstine 1999; et 2001). habitat types
al.
New
within sinkhole ponds in Virginia and Jersey are floristically and physically differ-
owing means and
ent, to their oi origin, Pleistocene history (glacial scouring surficial
and Irom
geologic deposits), distance of calcareous materials the soil surface.
In Virginia, weathering limestones and dolomites the Cambrian Shady and
ol of
Elbrook Formations is ultimately responsible for the formation of these pond features,
but a deep overburden of relatively acidic, infertile soils covers these carbonate rocks to a
&
depth of 30 to 150m (Fleming Van Alstme 1999). This phenomenon similar to that
is
m
who
described Maryland and West Virginia by Bartgis (1992) studied ponds support-
pond endemic
ing the rare Scirpus ancislrochaetus Schuyler In Virginia, surficial depos-
its are the result of alluvial fan formation along the western slopes of the Blue Ridge
mountains. Materials range from quartzite cobbles and boulders to sand, and clay
silt,
and range from the late Pleistocene to Pliocene or Miocene in age (Whittacar et 1999).
al.
Due to the depth where the carbonate rocks reside, soil and water chemistry of the ponds
(pH 0-30
acidic 3.9-4.0 at cm). Soils are predominantly gray to blackish clays or clay
is
loams with lighter colored mottling evident below 3 10 cm. This clay weathered from
to
dowm
metasiltstone and phyllite alluvium, impedes drainage but most ponds draw as
the growing season progresses and evapotranspiration increases. Although some of these
sinkholes arc relatively stable hydrologically Boltonia moniana almost exclusively
is
m
found those ponds with highly variable and unpredictable water The Boltonia
levels.
m
montana "zone" these ponds typically doughnut-shaped and extends from the rim
is
pond
of the to the edge of the most deeply flooded portion of the basin (Fig.
3).
The community types occupied by the species in Virginia are the Qucrcuspalustris/
Panicum -Panicum
rigidiiluni verrucosum - Eleocharis acicuiaris\Woodcd Herbaceous
Vegetation Association and the CephalanihusoccidentaJis / Polygonum hydropiperoides
- Glyceria acutijlora - Proserpinaca paiusiris Shrubland Association (Grossman
et
al.
NatureServe The former community endemic and
1998; 2005). type Virginia assigned
to
is
a rarity rank ol G1G2 (NatureServe 2005). The latter type somewhat more widespread
is
(ranked G3<'/S1) but considered endemic to mid-Atlantic region. These communities
is
are open-canopied except on pond edges, where the forest canopy can provide partial
communities
shading. Associated species in these include Quercus pahistris, Nyssa
sylvaiica/^ind Cephalanthus occidentalis (mostly occurnng on pond margins), Panicum
van philadelphicum,
rigiduliim rigiduluni, ven'uco:ium, Eleocharis acicularis,
P. P.
Elcocharismdanocarpa,hriocaulonaquaticum,Lysimachia
hyhrida,Lud\\'igiapahistris,
Lecrsiaoryzoides.Hypericum K)reale,and the tederally Threatened Hefeniun] virginicum.
In addition to relatively undisturbed sinkhole ponds, Boltonia montana also found with
is
' '
TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 879
j^j
.
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1^.^^-- %.^ M rt. -:^>; '^ -^.V.^TVii>^A^-^
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.^--> :.-, P* J _1 -^* _^
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1'- '-J F?- -
9^ :^ >r
SM i.^^',-;^^^n< U 1 _
•i^. ,*: n
^ -i"-- Af'J ^Irf '
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FrJi
Fig. 3. Sinkhole pond habitat of Boltonia montona in Augusta County, Virginia. Photo by Gary Fleming.
P.
-*j^
-ito'v^'i
Fig. 4. Zone occupied by Boltonia montana along margin of "Piggyback Pond/' Warren County, New Jersey. Photo by John F.Townsend.
BRILORG/SIDA
22(2)
880
which As
most these
H. virginicum in several altered ponds, of are in pasture. a result,
ponds include a number of species with broader habitat associations such asjuncii.s cffusus,
Ambrosia Phytolacca
canademisj. Bidcnsjrondosa, artemcsiijolia,
polylcpis,
tenuis, B.
J.
Venwnia ischacmum, Echinochloa
amcricana, Daucus novclxnacensis, Digitaria
carota,
among
Datura stramonium, and Erigcron annuus, others.
Eclipta alba,
rusgalli,
C
New
The ponds decidedly calcareous, occurring within a glaciated karst
Jersey are
landscape on dolomite bedrock. These sinkholes are found within the Ordivician Allen-
composed
town Dolomite and Beekmantown Dolomite formations, with surficial deposits
may
montana populations
bedrock complex. In rare instances, Boltonia
of a glacial till/
amount
be found on glacial outwash (kettles). In all cases, the of overburden present is
pond com-
than (and more alkaline than) that found in the Virginia
significantly less
and the carbonate rocks or marl deposits are usually at or near the soil surface,
plexes,
providing a slightly acidic to mildly alkaline soil environment (pH 6.2-7.8 at 0-30 cm;
New
Walz 2001). In Jersey the presence of marl and Chara spp. are characteristic of
et
al.
ponds
the sinkhole pondshorc habitat for montana- Although soil chemistry of these
B.
much
from Virginia examples, dramatic fluctuations in the seasonal water tabic (as
differs
m
m
New
as per season Jersey) are characteristic of ponds occupied by the species in
5.5
both states. Soils in New^ Jersey are shallow silt loams, silty clay loams, silty clay, or rarely
muck
marl overlaying clay or bedrock. Marly or peat soils are also present in rare in-
silt
Deep
stances and small dolomite rocks are often found on the surface (Walz et 2001).
al.
mad deposits often occur in the center of ponds occupied by B. montana, but as with the
montana doughnut-shaped
Virginia populations, Boltonid usually occurs in a distinctive
zone or "ring" between the pond edge and the more frequently and deeply flooded pond
center
(Fig. 4).
New
new
The community type occupied by the taxon in Jersey the Boltonia
is
- Symphyotrichum raccmosum - Mentha arvensis Herbaceous
astcroidcs asteroidcs
var.
Walz com-
Vegetation Asociation (Grossman et 1998; Natureserve 2005; et 2001), a
al. al.,
G1G2 naming montana
munity type that ranked by Natureserve (the of will involve
B.
is
commu-
community
a concomitant change in the name). Characteristic species in this
Cyperus Carcx
nity type include; Asclcpias incarnata, strigosus, cryptolcpis, C. lurida, C.
C
Cuphea Eupa-
typhina, viridula, viscosissima, Eleocharis palustris, Eragrostisfrankii,
acuminatum
varJasciculatumi-Pani-
torium pcrjoliatumjris versicolor,Dichanthelium
cunilanuginosuniy-3,r.tcnncssccnse),Panicumcapillarc,Panicumrigidulumv-3ir.pubcsccns
(=Panicum Polygonum amphihium, Slum suave, the green algae Chara spp.,
longifolium),
and ihchryophytcsHypnumlindlK'rgii,CampyliunistcUatum/^ndAm^^^^
New
community which
(Walzet 2001). Species in this type are rare in Jersey include:
al.
C C C C
montana, Carcx haydenii typhina,
Boltonia hcbhii, cryptolepis, C. retrorsa,
and (Snyder
viridula, Eleocharis quadrangulata, Eragrostisfrankii 2005).
Many
endemic wetland plant species are confined to habitats with widely fluctuat-
ing hydroperiods, and has been suggested that the ability of these species to tolerate
it
such stressful conditions releases them from competition with other taxa (Knox 1997),
who
was Knox
This aspect of the history of Helenium virginicum studied by (1997),
life
concluded that the ability of plants to tolerate the twin stressors of potentially toxic soils
and fluctuating hydrology was rare and contributed to the species' endemism. Boltonia
montana and virginicum co-occur several Virginia ponds where these adverse soil
H. at
pH and aluminum and have been documented,
conditions (low elevated levels of arsenic)
with the remainder of Virginia ponds thought to have similar soil characteristics. Data
TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN
U.S.A.
881
New
on
soil relations in the Jersey ponds are incomplete, but indications high alumi-
of
num, and magnesium
calcium, have been found and
levels water
erratic level fluctua-
well-documented
tions are there (Walz et 2001).
al.
NEW
Addirional specimens examined: JERSEY: Sussex Co.: near Lake Grinnell, 17 Sep 1887 N.L
Britton, s.n.(NY);
Swarrswood
Lake, 6Jul 1907, Philip Dowell 495J (GH); low swale, Brighton, 4 Sep 1910, K.K. Machcnzie 4778 (GH,
NY); borders of pond E of Swartswood Lake, Aug 1911, K.K. Mackenzie 4922 (NY, PH); border of Muckshaw Pond,
NW
Aug K.K
Springdale, 1917, Mackenzie 7986 (NY); '^SwaUowhole Pond;' 3.1 ktn E of Swartswood, 4,7 km of
Newton, 3 Oct 2005 J,F Townsend 3556 (VPl NCU, CLEMS); Swartswood km N
''Duck Pond;'
State Park. of
1.2
WNW
km
Paulinskill, 5 of Newton., 3 Oct 2005 J.F Townsend 3557 (PH, PENN); Trog Pond;' Swartswood
State
WNW
km NNE km
Park, 1.75 of Paulmskill, 5 of Newton., 3 Oct 2005J.H Townsend 3558 (NY); Tittle Frog Pond;^
WNW
km NNE km
Swartswood
State Park, 1.75 of Paulinskill, 5 of Newton, 3 Oct 2005J.F Townsend 3559(VPI);
"Spring Lake; Swartswood State Park, 3.1 km NWof Newton, 2.1 km SE of Swartswood, 4 Oct Townsend
2005,J.E
W
3562 (BRIT, GMUI^ GH); northernmost "Muckshaw Ponds;' 1.6 km SW of Newton and 0.4 km of US 206, 4 Oct
W
J.F Townsend 3565 (MARY, WVA, "Muckshaw km SW Newton km
2005, VPI); Ponds;' 2 of and 0.6 of US 206, 4
W
Oct 2005,J.E Townsend 3566 (CHRB, PH, NY); "Muckshaw Ponds," 2.75 km SW of Newton and km US
0,6 of
Townsend 3569 NCU, GMUP). Warren Muddy
206,J.E (VPl, Co.: calcareous shores and Shyster Pond, 24
flats, Jul
muddy
1920, K.K. Mackenzie, s.n. (NY, PH); shore of limestone sink pondjohnsonburg. 4 Oct 1981, D.B. Snyder
km
551-6PH, (PH); "Piggyback Pond;' located ca. 5 NE of Blairstown and 1.6 km N of Squn-es Corner,;.^ Townsend
NW
3560 (GH, CLEMS, USCH), (Warren/Sussex Co. Ime) Tour-Angle Pond," located 1.6 km of Tranquility and
km 5SW
1.25 of Greendell, 3 Oct 2005 J.F Townsend 3561 (VPl). PENNSYLVANIA. Dauphin Co.: banks of the
Susquehanna, near Dauphm. Aug
2 1865, C.E Parfee?; SJL (NY); Harrisburg Jul 1864, Mcirfuidtde, (PH). VIR-
I.e.
s.n.
GINIA. Augusta Co.: Sandy wet soil, bank of South River, vie. Lipscomb, Jul 1936, L.L Carr 47 (GH); growing on
18
m
bank
of South River vicinity of Lipscomb, abundant Stuarts Draft area around ponds and along South
River,
Rawhnson 58A Kennedy LL
18 Jul 1936, E.S. (VPI); Mt. Meadow, (unintelligible) bog, 14 Sep Can]
1937, (PH);
sji.
moist edge of pond (open sit.), vie. Lipscomb, 5 Aug 1938, L_L. Carr 365 (GH); edge of dried up pond. Big Levels
refuge, 8 Sep 1939, A.B. Massey, s.n. (VPl); Lipscomb Pond, in water, vicinity of Lipscomb, Shenandoah Valley, 8 Sep
1940, H.P. Killip, 36069 (US)] adjacent to Spring Pond, near Stuarts Draft, bottom of dried up pond, 16 Sep 1964, R.
S. Freer 2841 (GH, VPI); adjacent to Spring Pond, near Stuarts Draft, bottom of dried up pond, 23 Sep
1964,
R.S.
Freer 2869 (VPI); edge of small pond near Spring Pond, 5 Aug 1967, A.M. Harvill 17240 (FARM, NCU); Twin Ponds,
common
near Sherando, in drying portions of pond, elev 1550 19 Sep 1970, CE. Stevens 2781 (VPI); dried up
ft,
Game
pond, open area. Twin Ponds, Big Levels Management Area, 19 Sep 1970, IE Wieboldt 271 (WILLI); around
margms drawdown
and
the areas of Spring Pond, elev 1600 26 Sep Ramsey,
ft, 1970, G. R.S. Freer, R. Bruce, etal
common
18260 (WILLI); in shallow water of open edge of Twin Ponds near Spring Pond, near Sherando, 1540
elev
Oct CE. Wood
ft, 16 1971, Stevens 4772 (VPI, WILLI); duck Pond, scattered in and around edges of shallow sink-
NW
pond mi
hole 1.2 SE of Lipscomb, elev 1440 2 Oct 1984, CE. Stevens 19078 (FARM); Campbells Pond, mi
ft, 1.7
SW
N
of Sherando, side of St. Rt. 634 1 mile of jet. with Sr. Rr, 610, elev 1470 ft. 6 Aug 1986, G. Fcnwick, s.n. (VPl);
common m
with Eriocaulon septangidare dried muddy bed of intermittent sinkhole pond CTwin Pond"), 2.3 mi
W
of Sherando, elev 1540 ft, 22 Sep 1991, G.P Fleming 6126 vv/ CE. Stevens (WILLI); Twin Ponds, located 2.26
ca.
W
mi due of Sherando, 8 Aug 2002 Townsend 2869 WILLI, Div
(VPI, Va. Natural Heritage Herbarium).
J.F.
Relationship to Boltonia asteroides var. asteroides and caroliniana
B.
The morphology
gross of Boltonia montaiia nearly
is identical to that of B.asteroides var.
m
which name New
asteroides, a fact explains the use of this both Virginia and Jersey
for
many
yc-3.rs. Boltonia asteroides v^r. asteroides will be referred to as asteroides hr the
B.
sake of brevity in the following discussion.
The
montana
cypselae of Boltonia are found to lack several key characteristics found
namely:
in B. asteroides, cypselae wings, cypselae trichomes, and pappus awns. Occasional
montana
plants of B. will posess cypselae with slender pappus awns that are longer than
low crown may
the typically of scales. In this respect, such plants have awns approach-
ing the length seen in B. asteroides, but the other key features, glabrous cypselae with
wings
absent or vestigial, readily separate them from Fernald
B. asteroides. (1940) de-
rimmed
scribed B.asteroides as having obovate, thick cypselae with awns wanting up
or
mm
and New
to 0.7 long apparently included the plants of northwestern
Jersey in his
BRIT,ORG/SIDA 22(2}
882
awn-
which probably accounts his description of
concept of aslcroides at that time, for
B.
R
many
Gray ManuaUFernald 1940 In specimens of t^sfc
astcroidesin
less s J.950).
B.
mostly sclenfied and whitish on either side of the midrib and almost to
phyllaries are
m
and
phyllary becoming greenish gradually the distal portion, have a relatively
the
tip,
large, resinous midrib (1/5 to 1/4 of phyllary width) which continues to the phyllary tip.
m
monlana phyllaries are only sclenfied and whitened their lower half or
In contrast
B.
becoming green and herbaceous in the distal portion. The midrib is relatively nar-
less,
m
row phyllary width) the lower half and anastamoses in the distal half to
(1/10 to 1/5 of
was
produce network of fine veins within the green portion of the phyllary. It at first
a
m
m
moutaua were always unlike those found
assumed that the phyllaries seen B.
B.
but several specimens of astcroides from the Susc[uchanna River in Pennsyl-
cutcroidcs. B.
Whether gene
morphology former represents past
vania have similar phyllary to the this
known.
monlana convergence not
exchange with character
or
is
B.
monlana and
Due the strong resemblance between the cypsclae of BoJlonia B.
to
cawUniana, the latter species was investigated to discern other morphological similari-
Other than cypselae morphology, these two taxa have few character states
present.
ties, if
common.
in
and most
closely
Phyllaries of B();tonicK^aro/ in icina are quite dissimilar to B.m(7nt(3?i6i,
white and midrib but are shorter
resemble asteroides in terms of their color relative size,
B.
Head and
and narrower than both asteroides and monlana. size, ray corolla length,
B. B.
than and giving
pedicel diameter of B.ainVlinianti are snialler in B.cisteroiJe.s B.moiitci/ia,
appearance the inflorescence of carolin iana. In addition, typical individu-
a delicate to B.
branched and have numerous heads. Most individuals
cawliniana profusely
are
als of
B.
monlana and have few heads (often 1-25), although plants of
asteroides relatively
of B.
B.
both have been noted produce large plants with numerous heads (Karaman-
species to
Castro and Urbatsch 2006, see below). Such large reproductive outputs are seldom seen
Ray
and may be strategy sidestep adverse meterological or edaphic conditions. corol-
a to
carohniana, addition being smaller than those of montana, are white,
las of in to B.
B.
monlana
fading to pink or lilac in age (Anderson 1987, pers. obs.X while the rays of B. are
may
usually pinkish throughout the flowering period, though they also be extremely
lilac to
pale Most plants of montana are characterized by widely oblanceolate leaves
(see Fig. B.
2).
This
whereas those of caroUniana arc linear to narrowly oblanceolate or Imear^obovate.
B.
when two compared.
most evident bracteal leaves of the species are
difference
is
montana sug-
MorphologicalsimilarityofBoIUniiti to B.dsteroiJc.sandB.caro;ini6in6/
gest that montana may have arisen as a result of an ancient hybridization between the
B.
two Phylogenetic analyses of the internal and external transcribed spacers (ITS
species.
DNA
montana
and ETS) from the nuclear ribosomal suggested sister rclationaship of B.
(Karaman-Castro
with
and while only distant relationship asteroides et
caroJiniana, B.
B.
al Thus the treatment of the awnless and wingless specimens of B. asteroides as
in prep.).
new montana was supported by molecular However, remains unclear
data.
a species, B. it
and
between caroUniana
whether montana an ancient hybridization
a result of B. B.
B. is
asteroides.
Some observations on the morphology of Boltonia montana
The monlana from ascending with most individuals
Boltonia varies to erect,
stature of
growth
exhibiting the former condition (Fig. seems probable that this pattern is a
1). It
response edaphic conditions in the ponds rather than genetics. For instance, observa-
to
^Twin Ponds" Augusta Virginia were generally of few-flowered, ascending
tions at in Co.,