CAREX REZNICEKII, A NEW WIDESPREAD SPECIES OF CAREX SECTION ACROCYSTIS (CYPERACEAE) FROM EASTERN NORTH AMERICA

2006·16.3 MB·English

by  David A Werier

NEW
CAREX A WIDESPREAD OF
REZNICEKII
SPECIES
CAREX
SECTION ACROCYSTIS (CYPERACEAE)
h
FROM EASTERN NORTH AMERICA
David A.Werier
Rood
30 Banks
New USA
Brooktondale, York 14817,
[email protected]
ABSTRACT
new
Carex reznicekii described as from mesic to dry-mesic forests of Alabama, Arkansas, Connecticut, Dela-
is
New New
ware, District of Columbia, Georgia, Kentucky, Maryland, Mississippi, Missouri, Jersey, York, North
Carolina, Pennsylvania, Rhode Island, South Carolina, Tennessee, and Virginia. Previously, C. rezniceki had mostly
i
C C
been misidentified as either umhellata or nigromarginata. distinguished from these species as well as
It is
other members of section Acrocystis by a combination of lack of basal spikes, short culms, narrow leaves, and
A
only shghtly red pistillate scales, key to the 19 taxa of section Acrocystis that occur in eastern North America is
provided.
RESUMEN
Se describe como nuevo Carex reznicekii de los boscjues mesicos a xcrico-mesicos de Alabama, Arkansas, Caro-
lina del Norte, Carolina del Sur, Connecticut, Delaware, Distrito de Columbia, Georgia. Kentucky. Maryland, Mis-
C
New New
sissippi, Missouri, Jersey York, Pennsylvania, Rhode Island, Tennessee, y Virgmia. Previamentc,
habia estadoensu mayor parte mal mterpretada como C-umbellata o nigromarginata. Se distingue
rezriicekii, C.
de estas especies asi como de otros miembros de la seccion Acwcystis por la combinacion de carcncia de espiga
basal, ciilmenes cortos, hojas estrechas, y escamas femeninas solo levemente rojas. Se proporciona una clave para
los 19 taxa de la seccion Acrocystis que ocurren en el este de Norteamerica.
INTRODUCTION
Carex section Acrocystis Dumort. contains approxmiately 35 species worldwide with 20
&
in North America (Crins Rettig 2002), ahhough the most recent comprehensive treat-
now
ment Kiikenthal's (1909) outdated revision. Species distributions Ue mostly in
is
known
and North American with one taxon from South America (Kiikenthal
Eurasia
1909). appears that Carex section Acrocystis as currently circumscribed polyphyl-
is
It
but North American species and some Eurasian species form a clade (Roalson et
etic, al.
&
2001; Roalson Friar 2004). Twenty-eight taxa, including the one described here, are
known
currently from North America, with 19 of these occurring in eastern North
&
America (Crins Rettig 2002). At least two additional undescribed species occur in east-
& &
Anton
ern North America (Crins Rettig 2002; Roalson Friar 2004; Reznicek, pers.
comm.).
Eastern North American taxa of section Acrocystis, share the following characters:
C
two and
perigynia pubescent (except tonsa (Fernald) Bicknell tonsa veined,
var.
),
abruptly narrowed to a distinct beak; stigmas three (except two and three in CJloridana
(when subtended by sub-sheathing
Schwein.); non-basal pistillate spikes present) to
sheathless bracts, approximate, the distal ones short pedunculate to sessile; and habitats
more open few
dry to mesic forests (tending to favor drier) or for a taxa.
New
During survey work in the Hudson Highlands of southeastern York, found a
I
SiDA22(2):1049-1070.2006
1050 BRIT.ORG/5IDA 22(2)
C
New
population of the York State endangered nigromarginata. While delineating and
describing this population encountered a population of a strikingly different plant,
I
C
which
at first appeared to be umhcllata because all of its culms were very short and
Upon
hidden in the bases of the plants. closer inspection, observed that these short-
I
C
culmed umhcU
plants lacked the basal spikes characteristic of These short-cuhned
at
a.
plants were uniform throughout population and appeared immediately adjacent
its to
C
and
clearly did not resemble C. nigromarginata. In contrast to the nigrorrtarginata popu-
much
had
lation, these short-culmed plants narrower leaves, uniformly very short culms,
and pale pistillate scales. Wider sampling showed that the short-culmed plants formed
akhough
discreet uniform populations, sometimes occurring mixed with, but seemingly
C
A
not intergrading with, nigromarginata. cursory examination of herbarium material
new
also suggested that the distributions of the two entities were different. This species
Carcx described here Carcx
of as rcznicckii.
is
AND METHODS
MATERIALS
C C
Specimens of rcznicckii were sought by examining specimens labeled as
all
C
nigromarginata Schw^ein. and umhcllata Schkuhr ex Willd. (two superficially similar
m
and some members from GH, MO,
species) cases other of section Acrocystis BH, BRIT,
C
NCU, me
NY, NYS, PH, and US. In addition, putative rcznicckii specimens were sent to
DOy
from MICH, and VPI by
collections at as well as Philip Additional speci-
E. tiyatt.
mens
were gathered from work. Type material was examined
field of species similar
all
C
to or confusable with rcznicckii^ including their synonyms.
Thirty-one populations were visited in the field, covering a large part of the species'
were conducted
range. Site visits to assess habitat affinities, morphological variability,
and habitat differences with other members of Carcx section Acrocystu. Soil samples were
collected at six distant sites throughout a large part of the range of the species
(Fig.
1).
C
Equal amounts of soil from four places at each of the six sites (adjacent to individual
rczn icckii plants) that were at least 10 meters apart were mixed together These were sent
NY
Cornell Nutrient Analysis Laboratories Cornell
to at University, Ithaca, for analysis.
Soils were analyzed for particle size distribution, pH, and the minerals Al, Ca, Fe, K, Mg,
Mn, and
Zn.
P,
C C
To clarify the differences between nigromarginata, the most similar species
to
specimens were compared
rcznicckii, cind rcznicckii, using eight continuous charac-
C.
Specimens
ters (see Table w^ere selected so as to have ten specimens per species from
1).
each of three geographic regions: northeast (Virginia/Kentucky north), 2) southeast
1)
(North Carolina/Tennessee south), and Arkansas/Missouri. Only adequate
3) six speci-
C
mens
were available lor rcznicckii from the latter region. Within each geographic
re-
gion (excluding Arkansas/Missouri), only two specimens and one county
per per
state
were selected to help cover the range of the species. For each species, specimens were
selected randomly given the parameters mentioned above and excluding specimens where
ANOVA)
all eight characters could not be measured. Analysis of variance (two-way and
Pearson Correlation Coefficients were calculated using SPSS Version Additionally,
13.
ranges, means, and standard deviations were calculated for both species for each region.
Nonmetric multidimensional scaling (NMDS) ordination was chosen compare
to
C C
the relationship of nigromarginata and rcznicckii. In comparison other ordina-
to
tion methods such as canonical correspondence analysis (CCA) or principle components
NMDS
analysis (PCA), does not assume an underlying measured
distribution
for vari-
NEW
WERIER, A SPECIES OF CAREX FROM EASTERN NORTH AMERICA
1051
C
Fig. 1 Geographic distribution of reznicekii. Soil samples collected in blackened counties.
.
NMDS
A
ables. ordination was created using the slow thorough mode on PC-ORD
&
m
(McCune
Grace 2002). Specimens measured for use the ordination are dehneated by
C
a single asterisk in the isotypes listed for reznicekii, in the representative specimens of
C
and specimens Appendix
reznicekii, in the cited in A.
A
scatter plot graph using a larger sample size was created for the two most explana-
C C
As and
tory characters. additional evidence that reznicekii nigromarginata are dis-
creet entities, distribution curves were created and compared for the two most explana-
C C C
and and
tory characters for nigromarginata reznicekii, as well as nigromarginata
C C
reznicekii combined, and nigromarginata specimens from outside the geographic
C C
range of reznicekii. For the latter distribution curves specimens of nigromarginata
all
C
that occur outside of the geographic range of reznicekii from BH. BRIT, MICH, and
MO,
excluding were measured. Specimens measured graph
duplicates, for the scatter plot
and
distribution curves are delineated by either a single or double asterisk in the isotypes
C
C
specimen and
listed for reznicekii, in the representative of reznicekii, in the speci-
mens
Appendix
cited in A.
A C
and members
description of reznicekii a key to all of Carex section Acrocystis
C
were from measurements measurements
created original except, for the key, for com-
C
munis and
Bailey varieties inops Bailey ssp. heliophila (Mackenzie) Crins are adapted
from Crins and Rettig (2002). Character states that were on an extreme edge or disjunct
from most measurement The
other are placed in parentheses. following notes clarify three
characters listed in the description and the key
Culm measured from culm
length the base of the to the top of the inflorescence
1. is
&
many
as interpreted or implied by other authors (LeBlond 1994; Reznicek
et
al.
Camelbeke Naczi and
1996; et 2001; others).
al.
2. Terminal staminate spikes exceeding or exceeded by the rest of the inflorescence
is
1052 BRIT.ORG/SIDA 22(2)
Table Characters measured for ordination, ANOVA, and Pearson Correlations giving the codes used through-
1.
out the paper and any specifications for choosing wliich part to measure.
Character Code Specifications
TaCuHe
culm height (cm) per specimen
Tallest Tallest
Widest width (mm) WiLeWi Widest per specimen
leaf
Anther length (mm) AnLe Mean of two per specimen
Largest length that staminate StSpExPiSp Greatest length per specimen
a
spike exceeds the distal
(mm)
most
spike
pistillate
Perigynia iength (mm) PeLe Mean of two per specimen
Perigynia width (mm) PeWi Mean of two per specimen
Taller staminate spike length (mm) StSpLe Mean of the length of two taller staminate
specimen
spikes per
Staminate spike widtli (mm) StSpWi Mean of the width of the two staminate spikes
measured
spike length
for
measured by subtracting the difference of the length from the base of the bract subtend-
ing the proximal-most non-basal spike to the apex of the terminal stammate spike, and
the length from the base of this same bract to the apex of the distal most lateral spike.
3. Perigynium body refers to the perigynium excluding the beak and stipe or stipe-
like base. The beak and stipe begin at the deepest point in the concavity formed at the
summit and base of the perigynium. Perigynium body shapes vary considerably and
al-
though their shape aids in distinguishing taxa (and used in the key), they should be
is
used cautiously
RESULTS
Carex
reznicckii Werier, sp. nov. (Fig. 2). Tyit:; U.S.A. Virginia. Camline Co.; near North Anna River,
W
Route 207 Road) from Carmel Church Oxford Road (Route mi and SE en Oxford
(Jericho to 689), 3.6 S
SW UTM
Road (Route 689) to small dirt road on S side of road; along dirt road towards North Anna River.
NAD-83
coordinates in Zone 18 419670IN 2S0996E, 10 Apr 2004, Werier 1951 (holotype: BH, isotypes:
MICH. MO, NCU,
GH"^, NY, US),
Curici nigwmarginatae simihs. scd characreribus scquentibus differt; laminis fohorum hitissimis 1.2-2.2(-2.5)
mm
cm
latis; cuhnis singulae phuitae saepe in longirudine simihbus k^ngissimis 1.9-9.91.-13.7) Longis, spicis
mm mm
stamiuabhus squamarum
3.3-8. 3(-9. 2) h.^ngis et 0.4-L6(-1.8) latis; et colore rubenti pistillatarum ad
regioncs marginales
restricto.
Densely caespitose, herbaceous, evergreen perennial. Rhizomes horizontal, ascending, or
mm mm
erect, 0.1-3.8 long betv/een shoots or branches of the rhizomes, 0.6-1.2 wide with
leaf sheaths absent, enclosed by leaf sheaths and/or cataphylls, which disintegrate into
long fibers. Vegetative shoots erect to ascending, arising from buds at the tip or side of the
rhizomes, or directly from the apical meristcm of rhizomes or previous year's vegetative
which
shoots; bases consisting of cataphylls or leaf sheathes disintegrate into long
fi-
cm
bers; pseudoculnis (represented by leaf sheaths) 1.2-7.6 long. Reproductive shoots erect,
mostly produced irom meristem
directly the apical of vegetative shoots or rhizomes, or
less frequently produced from buds at tip or side of rhizomes, bases mostly surrounded
only by leaf sheaths which disintegrate into long fibers, or less frequently surrounded
mm
cm cm
by culms
cataphylls; 1.3-9.9(-13.7) long, the tallest 1.9-9.9(-13.7) long, 0.3-0.5
wide just below the inflorescence, trigonous, with elevated vein on each angle and 2-5
1
elevated veins on each side, smooth to minutely antrorsely scabrous on angles and veins,
NEW
WERIER, A SPECIES OF CAREX FROM EASTERN NORTH AMERICA
1053
=
Bottom Bottom
FiG.2.C£7rexrez/7/feM right, plant (scale 5 cm). left. Pistillate scales, perigyni^
= =
(I to r), culm, inflorescence, and sheath (scale 1 cm).Top left, staminate scale and anther (scale 1 mm).
BRIT.ORG/SIDA 22(2)
1054
more prominently scabrous on angles and Cataphylls base of reproductive
distally. at
mm
cm
shoots 0-2, 1.6-10.8 long. Leaves of reproductive shoots 3-9. Leaf blades to 52.0
mm mm
long, 0.7-2. 2(-2. wide, the widest 1.2-2. 2(--2.5) wide, flat to V-shaped, to occa-
5)
sionally M-shaped; adaxial surface dark green and smooth, papillose, or antrorsely sca-
brous, more textured distally; abaxial surface smooth to occasionally scabrous, more tex-
on mid-vcin and margins smooth antrorsely scabrous, more scabrous
tured distally; to
cm
some
distally; leaf sheaths 0.7-5.1 long, lower portion especially on outer sheaths have
and and
red coloration; abaxial face with green to white or proximally red veins whitish
translucent proximally reddish and more opaque intervem regions; adaxial face with
to
proximally red veins and whitish, thm, and translucent intervein regions, thin-
w^hitish or
ner and more translucent than abaxial intervein regions; vein edges with ascending,
spreading, or lexed minute, broad-based deltoid hairs; sheaths disintegrate into
ref
stiff,
long fibers consisting of veins, which retain pubescence; adaxial sheath face apex con-
mm
and sometimes
cave to a depth of 0.3-0.8(-1.0) slightly thickened; ligules typically
mm mm
wider than long, 0.3-1. 0(-1.5) long (including the free portion), 0.6-1.4 wide, free
mm
with abundant minute, broad-based
portion of ligule 0.1-0.2 long, ciliate stiff, del-
mm
toid hairs. Infructcscences 5.2-12.9(-16.7) long, consisting of 3-5 approximate spikes
mm
at the summit of the culm; proximal-most internode 0.8-3. 7(-5, 2) long; bracts
mm mm
proximal-most bract with blade 3.6-23.0(-38.5) long, 0.6-1.8 wide,
sheathless;
mm mm
3.9(-4.3) shorter than to 10.9(-23.5) taller than infructcscences; distal bracts re-
mm
duced. Spikes arising singly from nodes; terminal spike staminate, 3.3-8.3(-9.2) long,
mm
mm
0.4-L6(-1.8) wide, on peduncle (0.2-)0.3-0.7 long, exceeding distal-most lateral
mm
mm; 2-
spike by exceeded by distal-most lateral spike by 1.1(-L6) lateral spikes
3.7 to
short-pedunculate, with cladoprophyll towards base of peduncle; roxi-
pistillate, a p
4,
mm mm
mal-most spikes 3.4-7.2 long, (L8-)2.2-4-4 wide, (3-)5-10(-12) flowered, on pe-
mm mm
duncles (0.3-)0.4-0.9(-L4) long, with cladoprophyll (0.7-)1.7-2.8(-3.0) long.
mm mm
apex
Staminate scales (2.5-)2. 7-4.0 long, 0.7-1.8 wide, ovate to elliptic, acute to
which
obtuse, with a green or yellowish longitudinal mid-stripe includes the mid-vein,
margins translucent, thin, whitish to occasionally yellowish-brown, often red sub-mar-
ginally forming a longitudinal stripe, more so toward apex of scales and on distal scales,
mid-vem extending toward apex more
present, further the of the scale in distal scales;
distal scales with mid-vein raised, slightly antrorsely scabrous, and ending close to apex of
awn
scale or sometimes with mid-vein projecting as a short attached just proximal to the
scale apex and projecting up to just beyond the apex of the scale; Pistillate scales 2.6-4.3
mm mm mm mm
long, 1.0-1.9 wide, 0.9(-l.l) shorter than to 0.5 longer than associated
pcrigynium, ovate to lanceolate to elliptic, apex acute to obtuse, with a green (to brownish)
which margins whitish
longitudinal mid-stripe includes the mid-vein; translucent, thin,
occasionally yellow-brown, often some reddish color sub-marginally forming a sub-
to
marginal longitudinal stripe, more so distally on the scale; sub-marginal longitudinal red
stripe present) does not extend laterally to immediately adjacent to mid-stripe except
(if
sometimes at apex of scale, mid-vein antrorsely scabrous and ending just before apex of
awn
scale or mid-vein, projecting as a short attached just proximal to the scale apex and
up beyond apex with narrow
projecting to just the of the scale; bases of scales often a red
mm
horizontal above attachment spike Anthers (1.2-)13-1.9(-2.3) long.
stripe just to axis.
3,
mm mm
Stigmas withering with age. Perigynia (2.5-)2.7-3.9 long, (0.8-)0.9-1.3(-1.5) wide,
3,
light green, occasionally sparsely red-punctate, obtusely trigonous to plano-convex in cross-
section, with two prominent nerves that extend the length of the perigynia and occasion-
WERtER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA
1055
up prominent
ally 8 nerves with up
to less at base, papillose papillae short-cylindric
to 0.01
mm
long, as well as pubescent with small than mm), predominately
(less 0.1 antrorsely
which
directed, stiff, deltoid hairs, are denser on the prominent nerves that extend mto the
mm
beak body
teeth; 1.4-L9(-2.0) long, ellipsoid, gradually tapering beak and
to stipe; stipe
mm
mm
mm
beak
0.5-L0(-l.l) long; (0.4-)0.6-1.0 long, bidentate; beak teeth 0.1-0.3 long.
mm
mm
Achenes
(1.4-)L5-1.8(-1.9) long, 0.9-L2(-L3) wide, ovoid, acutely obtusely
to
trigonous plano-convex minutely when
to in cross-section, papillose, yellow-green green
to
immature, brown when
light to chestnut mature.
Etymology —I honor Dr Anton
selected reznicekiias the epithet to Reznicek, Univer-
who me many
sity of Michigan, has inspired and others interested in carices. He has cata-
lyzed a revival in the study of Carex which has revealed tremendous amount new
a of
information. In addition, he has contributed directly to a greater understanding of these
through numerous and
plants
articles classes.
A C
242 specimens
total of (143 records) of reznicekii were examined. Of 125
these,
C
specimens were made
(88 records) collections prior recognition
to of reznicekii. Sev-
C
C
enty-one of these 125 specimens had at one point been labeled nigromarginata, 78
C
umhellata (including ahdita Bicknell and umhellata var
C. brevirostrisBoott), 6
C.
C
Jloridana [including C. nigromarginata vd^rjloridana (Schwein.) Kiikenthal], and 9
C
albicans Willd. ex Spreng. var. emmonsii (Dewey ex Torr) Rettig [including emmonsii
Dewey
ex Torr and nigromarginata var minor The sum more
C. (Boott) Gleason].
totals to
than 125 because some specimens had been annotated numerous
times.
C
Type material examined comprised
the following; unibellata, scan of holotype from
C
Dewey
B; umbellata var. vicina holotype at GH, isotype at PH; umbellata var
C.
US
examined
brevirostris, isotype at (holotype not but specimen from Saskatchewan);
is
C
C
abdita, holotype at NY; C. microrhyncha Mackenzie, holotype at NY; umhellata var
C
tonsa Fernald, lectotype and 2 isolectotypes at GH; rugosperma Mackenzie, holotype
at
C
NY; nigromarginata, holotype at PH; and Cjloridaim, holotype at PH. None of the types
examined
proved be
to reznicekii.
C.
A C C
C
comparison
detailed of reznicekii umbellata and nigromarginata made
to
is
C
below. In the past, reznicekii has been mistaken most often these two Carex
for species.
nigromarginata most similar The key compares members from
is all of section Acrocystis
eastern North America.
Carex
umbellata superficially resembles C. reznicekii, but actually quite distinct.
is
The two species can easily be fully separated by noting the presence (C umbellata) or
absence (C Mackenzie when
reznicekii) of basal spikes. (1913), clearly stated this he ex-
fl
Cfl
fact that while the spikes are on very short culms and may appear basal they are not on
basal peduncles." Fernald (1902) apparently did not understand the concept of basal
when C
spikes he stated that "the best means of distinction between umbellata [misap-
C
C
phed
to tonsa var rugosperma (Mackenzie) Grins] and nigro-marginata
[sic] is of-
by
fered the thickness of the perigynia.^^ In his 8th edition of Gray Manual, Fernald (1950)
s
C
placed nigromarginata with the basal spike members Montanae
of section Acrocystis
(as
C
and then used perigynium and
Fries) characters geographic region distinguish
to
C
nigromarginata from umbellata and its allies. This means of distinction appears to have
been followed by at least Cusick (1992) and Tucker (1995), and may have in part resulted
C
in C. reznicekii often being misidentified as umhellata.
BRIT.ORG/SIDA 22(2)
1056
followm
Cumhd/cifa from Cre2?nce/?ii include the
Other characters that separate
C
C 0-2 2-4
umbcUata has stammate spikes with approximate pistillate spikes vs. for
some spikes on elongate peduncles vs. none on elongate peduncles
reznicchii; pistillate
mm
C
up beyond approximate
staminate spikes extending to 9.0 the (if
reznicekii-
for
mm
most extendmg beyond
staminate spikes the pis-
present) pistillate spikes vs. at 3.7
mm
C
spikes reznicekii] and perigynia (2.2-)2.3-3.2(-3.3) long vs. perigynia
tillate for
mm C
C
(2.5-)2.7-3.9 long for reznicekii. In addition, leaf blades of umhellata are gener-
and wider than
ally lighter green slightly C. reznicekii.
A seems important understand-
understanding basal spikes particularly to
of
full
may
between two and enrich the understanding of sec-
ing the difference these species,
which
tion Acrocystis as a whole. Basal spikes are individual pistillate spikes arise al-
Mackenzie These
most directly from the base of the plant [subradical of (1913)]. pistillate
spikes have elongated peduncles, and as with all pistillate spikes in section Acrocystis,
Macken-
and
subtended by These bracts have sheaths blades [contrary to
they are bracts.
(Montanac
and Roalson and Friar (2004) that state that section Acrocystis of
(1935)
zie
which subsheathmg]. These authors probably
Mackenzie) has bracts are sheathlcss or
meant that bracts of non-basal spikes are sheathlcss or subsheathing. In addition, the
C
culms uniheUaia (defined as from the bases of the culms to the apices of the mflo-
of
much
Roalson
rescences) are often taller than the apices of the basal pistillate spikes.
continuous
and Friar (2004) consider basal pistillate spikes to be better described as a
While
somewhat opposed character
character with distinct states, as to a discrete in es-
two
sence could be appears that this character functions in completely dis-
this true,
it
peduncles and sheathing
with associated other characters (elongated bracts).
tinct states
C
Even more interestingly while neither nigromarginata nor C. reznicehi has basal
culm which
spikes, both often have "grouped culms.^^ These "grouped culms" have one is
m
mim-
which (much pronounced
taller and one to three are shorter less C. rezniceku),
C
culms and
ickmg and with often shorter basal
the short aspect of urn hcUata, tall
its
tall
C
culms culms
spike peduncles. In addition, the shorter in a "group" of in
pistillate
C
nigra margin at a and reznicekii have relatively shorter and less projecting terminal
staminate
spikes.
no
Carex reznicekii and nigromarginata share a few characters, including basal
C.
m
some culms and hidden
spikes, no elongated rhizomes, at least short leaf bases,
and composed 3-5 approximate
perigynia of similar length, stigmas inflorescences of
3,
C
Likely because of these similarities, reznicekii has often been misidentified as
spikes.
C
nigromarginata.
Several characters separate these two species. First, C. rc^n icekii has narrower leaves
mm mm
wide
wide
with the widest per plant 1.2-2. 2(-2. 5) vs. (1.9-)2.3-4.5 for C.
C
nigromarginata. Second, the culms per plant are shorter For reznicekii the tall-
tallest
C
cm cm
est culms are L9-9.9(-13.7) long vs. (4.5-)6.6-38.0(-51.0) long for n igromargi nata.
C
same com-
have culms often about the length
Third, individual plants of reznicekii
C C
among
pared widely different lengths individuals of nigwnuirginata. Fourth,
to
some culms droop maturity
reznicekii culms remain erect even at maturity while at for
C C
nigromarginata. Fifth, reznicekii has margins of pistillate scales without red or red
The can form submarginal longitudinal
only submarginally red coloration present) a
(if
but does not extend laterally to the green (or brown) longitudinal mid-stripe of
stripe,
the scale except the apex. Carex nigromarginata has pistillate scales either similar in
at
color reznicekii or more often with dark reddish to dark purplish/black coloration
to C.
NEW
WERIER, A SPECIES OF CAREX FROM EASTERN NORTH AMERICA
1057
that extends laterally from the submargin to the green (or brown) mid-stripe of the
scale.
C
way
In addition, the red to purple color often extends all the to the base of the scale in
C
nigromarginata, while
in reznicekii the reddish color present) does not extend
(if to
C
the base of the scale. Sixth, the apex of the staminate spike in reznicekii exceeds the
C
apex mm.
of the distal-most lateral spike by at most 0.0-3.7 In nigromarginata, the
apex which
of the staminate spike exceeds the apex of the distal-most lateral spike the
Wh
J
overlap in these two species, can sometimes be useful distmguishing
it for difficult speci-
C
mens.
Seventh, reznicekii has staminate spikes on average slightly shorter and nar-
mm
mm
measurmg x
x
rower,
3.3-8.3(-9.2) 0.4~1.6(-1.8) vs. 4.2-10,9(-12.0) 0.5-2.4(-2.8)
C
nigromarginata. Again shows
for this character significant overlap, but occasionally
it
provides useful help in distinguishing between these two
species.
Carex reznicekii and C. nigromarginata were compared for eight continuous charac-
ters (see Table Ranges, means, and standard deviations continuous
1). for all eight charac-
m
NMDS
C
measured
ters for the ordination are presented Table Comparing
2.
C
nigromarginata and reznicekii,VC-ORD recommended
two-dimensional
a solution
for
NMDS
the ordination. This resulted in a final ordination with low
(see Fig. 3) instability
(<0.001) indicating convergence of the iterations, and low stress (6.61) indicating that the
data was far from random and there was "no drawing (McCune
real risk of false inferences"
&
Grace 2002). Axis of the ordination represents decreasing TaCuHe and
(-0.961)
1
WiLeWi
the Pearson Correlation
_
m
Coefficients with Axis Axis explained 81%
(Fig. of the variability the
1 3). 1 dataset.
WiLeW
StSpW
representing the Pearson Correlation Coefficients with Axis 2 Axis 2 explained
(Fig.
3).
an 17% 98%
additional of the variability for a total of explained variability Overall, the
ordination summarizes a very structured dataset with several highly correlated variables
m
and shows
addition while two some
that the species are distinct, overlap of individual
specimens
Pearson-correlations measured
exists. illustrated that of the characters
all
width and
except length of the perigynia were strongly correlated with one another <
(p
indicating high redundancy The
0.01), a level of of the characters used. ordination pre-
is
C
C
how
sented mainly as a visual tool to show reznicekii and nigromarginata compare
with numerous
characters.
All of the characters measured except width and length of the perigynia differed
C C
significantly between reznicekii and nigromarginata The
(Table scatter plot graph,
3).
using sample two
a larger size for the strongest characters separating the two species
(tall-
est culm height and widest leaf width), shows almost individuals can be determined
all
using these two characters alone (Fig. 4). Ranges, means, and standard deviations for this
sample
larger size are presented in Table 4. Mainly depauperate or aberrant individuals
were not separable with these two characters. These specimens were determined by the
additional characters as discussed previously with scale color being the most useful.
Regional
differences in the characters were significant only width and length
for of
C
perigynia
(Table 2). Perigynia of reznicekii varied by geographical region with slightly
narrower and shorter perigynia on plants from the northern region compared the south-
to
and
ern Missouri/Arkansas These
regions. statistically significant differences represent
only minor and
variations have questionable
biological significance.
For the distribution curves widest width and culm
height
for leaf tallest a total of
1058 BRIT.ORG/5IDA 22(2)
1.5-
n
a
C/^
0.5
00
a
a
<N D
a
#
:/;
<3>
Q
00
00 -0.5
D o
H
C
o
reznicekii
a
CM
C
nigromarginata
D
D
<
a
a
a
(Z)
-1.5
-2.5 -1.5 -0.5 0.5 1.5
TaCuHe WiLeWi
Axis
(-0.961), (-0.747)
1:
Fig. 3. Non-metric multidimensional scaling ordination showing spatial relationship of C reznicekii awd C. nigromarginata based on
five explanatory characters. Symbols grade larger based on wider widest leaf blade widths. Main characters correlated to each axis
are listed with their Pearson Coefficients in parentheses.
NMDS
Table 2. Ranges, mean5,and standard deviations for characters measured for the ordination (Fig 3). Char-
= =
acter codes are defined in Table Regions listed below are as follows: NE V/irginia/Kentucky north;SE North
1
.
AR/MO
CarolinaATennessee south; and ^ Arkansas/Missouri.
Carex reznicekii Carex nigromarginata
regions regions
AR/MO AR/MO
Characters NE 5E regions N 5E All regions
All
TaCuLe 2.2-12.2 2.5-137 37-13.7 2.2-137 9.0-27.7 6.6-18.7 7.2-38.0 6.6-38_0
5.4±3.1 5.4±3.3 7.3±3.5 5.8±3.3 15.2±5.2 3.3±3.8 17.5±8.8 15.3±6.3
WiI.eW 1.3-2.2 .3-2.0 17-2.2 1.3-2.2 2.4-4.0 2.4-3.5 2.5-3.5 2.4-4.0
1.6±0.3 .7±0.3 2.0±0.2 1.7±0.3 2.9±0.5 2.7±0.4 3.0±0.3 2.9±0.4
AnLe 1.30-1.80 1.30-1.95 1.50-175 .30-1.95 1.50-2. .45-2.25 1.60-2.50 1.45-2.50
1.50±0.15 1.52±0.17 1.60±0.10 1.53±0.15 1.82±0.23 1.84±0.25 1.98±0.30 1.88±0.26
StSpExPiSp 0.2-2.5 0.2-2.2 1.1-3.7 0.2-3.7 0.7-7.3 1.1-6.0 0.5-5.5 0.5-7.3
1.4±0.7 ±0.5 ±1.2 .6±0.g 3.4±2.0 2.7±1.6 3.0±1.7 3.0±1.7
1.2 2.5
PeLe 2.75-3.45 2.80-3,70 3.35-3.55 275-3.70 2.70-3.30 2.90-3.60 3.00-3.75 2.70-3.75
3.23±0.27 3.40±0.10 3.24±0.23 3.14±0.18 3.27±0.25 3.37±0.25 3.26±0.24
3.16 :0.21
PeWi 0.95-1 .05-1 ,20 .05-1 .20 0,95-1 .20 0.95-1 ,35 0.90-1 .35 .05 ,25 0.90- .35
1 1 1 1 1 1
.
1.02±0.06 1.13±0.05 1.13±0.07 1.09±0.08 1.11±0.11 1.14±0.12 1.12±0.07 1.12±0.10
StSpHe 4.60-7.55 4.60-7.85 5.80-9.00 4.60-9,00 5.85-12.10 6.15-11.15 5.35-10.70 5.35-12.10
6.31±0.89 6.05±1.05 7.33±1.21 6.45±1.11 8.41±2.13 7.79±1.58 8.02±1.74 8.07±1.79
StSpWi 0.90-1.55 0.70-1,65 1,05-1. 0.70-1 ,65 .05-2.25 .00-2.00 .35-1 ,75 .00-2.25
1 1 1
1.27±0.20 1.20±0.28 1,31±0.22 1.25±0.23 1.62±0.36 1.50±0.31 1.52±0.15 1.54±0.28