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Contents

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and

the District of Columbia .

Christopher T. Frye and Christopher Lea 41

Instructions to Authors . 110

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The Maryland Naturalist 44(2) :4 1-108

Winter 2001

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the

District of Columbia

Christopher T. Frye1 and Christopher Lea23

1 Maryland Department of Natural Resources, Wildlife and Heritage Service, Tawes State Office
Building, E-l, 580 Taylor Ave., Annapolis, MD 21401

2 National Park Service, 7206 National Seashore Lane, Berlin, MD 21811

Abstract. — Collections and field research were conducted to document the entire
flora of the genus Carex (Cyperaceae) in Maryland and the District of Columbia as a
county-based state atlas and annotated list. One hundred seventy-five taxa, represent¬
ing 158 species, are reported. Twenty-nine of these taxa are newly reported for the area
since publication of the current Maryland flora, with eight of the new taxa first identified
by this study. Ten taxa previously reported by other authors are excluded based on
erroneous or unfounded sources. The history of floristic research on the genus in the
area is reviewed, and phytogeographic patterns are examined. County occurrence maps
and an annotated checklist of the taxa, with notes on key identification features, distri¬
bution, habitat, and rarity, are presented.

Introduction

Carex (Cyperaceae) is the largest genus of vascular plants in most areas of east¬
ern North America (e.g., Harvill 1973; Reznicek and Ball 1974; Wheeler and Ownbey
1984; Reznicek 1989). It is often not well understood in floristic studies because of its
complexity and the identification challenges presented by many taxa (Reznicek 1989). A
number of discoveries in Maryland and the District of Columbia have been made since
Brown and Brown (1984) documented Maryland’s flora. Additionally, the past two de¬
cades have seen a number of new taxonomic treatments of North American Carex (see
Literature Cited section), which, in total, represent a significant revision of those em¬
ployed by a number of state and regional floras currently in use. Together, these situa¬
tions create the need for an updated treatment and phytogeographic study of the genus
in Maryland and the District of Columbia.

History of the Study of Carex in Maryland and the District of Columbia

Any effort of this scope rests upon the foundation laid by many previous genera¬
tions of botanists and taxonomists, and we begin with a summary of historical context of
the study of Carex in our area. Collections of Maryland and District of Columbia Carex
on which most current floristic knowledge is based were begun in the 1870s. Theodor
Holm, Forrest Shreve, Howard Shriver, John Donnell Smith, Edward C. Steele, George
Vasey and Lester Ward are prominent among collectors of Carex in Maryland and the
District of Columbia in the late 1 9th century and the earliest years of the 20th century.
Ward (1 88 1) published a flora of Washington, DC and vicinity, defined approximately as
that area within 1 5 miles of the Capitol (Hitchcock and Standley 1919); he recognized 70

^GhrisLeafginps.gov

41

C. T. Frye & C. Lea

taxa within the genus. Ward’s work was eventually updated and expanded by Hitchcock
and Standley ( 1 9 1 9); the latter remains the most recent flora for the District of Columbia.
The treatment of Carex in Hitchcock and Standley ( 1 9 1 9), by G. P. Van Eselstine, lists 95
species represented by specimens, with several infraspecific taxa and hypothetical spe¬
cies also mentioned.

During an ecological survey of the vegetation of Maryland, Shreve et al. (1910)
compiled a statewide list of vascular plants. Although the authors acknowledged that
the goal of the study was primarily a description of vegetation, rather than a comprehen¬
sive floristic effort, their work is believed to be the first list of the vascular flora for the
entire state, and included 56 Carex taxa.

Significant collecting of Carex in Maryland and the District of Columbia contin¬
ued through the first half of the 20th century. Collectors include Sydney F. Blake, Eliza¬
beth Earle, Frederick J. Hermann, Ellsworth P. Killip, Emery C. Leonard, Bayard Long,
William R. Maxon, John Bitting Smith Norton, Hugh O’Neill, Paul C. Standley, Witmer
Stone, Robert R. Tatnall, Ivar Tidestrom and Rodney True. During the early part of this
period, Kenneth K. Mackenzie, who was associated with the New York Botanical Garden
and who eventually published a landmark treatment of North American Carex (Mackenzie
1931, 1935), reviewed and annotated many Maryland and District of Columbia collec¬
tions housed at the Smithsonian Institution. This period culminated with the publica¬
tion of two checklists (Hermann 1 94 1 , 1 946) of the flora of the vicinity of the District of
Columbia, the definition of which had been expanded from the area covered by Ward
(1881) and Hitchcock and Standley ( 1 9 1 9) to include virtually all of the Piedmont physi¬
ographic province and the Western Shore of the Chesapeake Bay in Maryland (Schmidt
1 993). Hermann ( 1 946) reported 1 23 Carex taxa for the expanded vicinity of Washington,
DC.

Numbers and collection dates of specimens in herbaria suggest a generally re¬
duced level of field study of Carex (and other components of the flora) in Maryland and
District of Columbia from the 1 940s through the 1 970s, as compared to the preceding 70
years. Perhaps the most prolific collector of carices from the late 1 950s to the early 1 970s
was Eduards Baltars, who contributed to much of our understanding of the genus in the
state, particularly in the northern Piedmont area. Frederick J. Hermann, a U.S. Depart¬
ment of Agriculture botanist and a prominent caricologist of the time, continued his
contributions over a period of over 30 years, primarily through examinations and anno¬
tations of countless specimens in collections.

By the early 1980s, two events had served to refocus efforts on collecting floristic
data on Carex in Maryland. First, in 1979 the Maryland Natural Heritage Program was
initiated within the Maryland Department of Natural Resources and began an effort to
determine and document the state’s rarest vascular plants. Natural Heritage staff and
their network of field biologists throughout the state were responsible for greatly ex¬
panding our knowledge of the occurrence and distribution of the state’s rarest carices.
Significant collectors of Carex during the 1980s and 1990s are D. Daniel Boone, Charles
Davis, Steven R. Hill, Wayne D. Longbottom, William McAvoy, Roderick Simmons,
Brent Steury and Edward Thompson. The second event was the publication of the first
comprehensive flora for the herbaceous vascular plants in Maryland by Melvin L.
Brown and Russell G. Brown (Brown and Brown 1 984). They recognized 1 54 Carex taxa

42

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae| of Maryland and the District of Columbia
(some of which are indicated as being hypothetical for the state).

Purpose and Development of the Study

Prior to 1997, the Maryland Natural Heritage Program listed 77 Carex species as
being of conservation concern (i.e., rare, threatened, endangered, watchlist, or status
uncertain; Maryland Natural Heritage Program 1 994), a total that represented about one-
half of the Carex taxa known from the state. The reasons for this rather high proportion
of listed taxa are several, and are similar to problems affecting the understanding of
Carex in other areas (e.g., Keznicek 1 989). First, a number of taxa had been, and contin¬
ued to be infrequently collected or under-reported because of identification challenges.
Also, compared to recent research, the treatment by Brown and Brown (1984) of some
sections of the genus had become outdated, so that taxonomic and nomenclature con¬
fusion was contributing to misunderstandings concerning the distribution and conser¬
vation statuses of some taxa. The conservation ranks of the 77 listed species were
reviewed in 1997, and nearly 60 percent of these subsequently received some level of
revision. The majority of changes involved reducing the priority of the conservation
rank of species that were found to be more common than formerly believed. For smaller
numbers of species, higher priority ranks were assigned in order to reflect evidence of
greater rarity than formerly perceived, and the ranks of several other species were
changed to reflect uncertainty about their statewide status.

While we initiated our research in 1 998, primarily as a collections review in order to
further refine Natural Heritage ranks of the rarer taxa in Maryland, this process sug¬
gested a substantial need for a revision of the treatment of the entire genus within the
state. Here, we present the summary of both our collection reviews and statewide field¬
work from 1998 through 2001 as well as substantiated reports by others. Despite four
years of research, much work remains. Some taxa remain taxonomically or floristically
problematic, often because of unstable circumscriptions or nomenclature. Others may
include local variants and perhaps inter-taxa gradations. Therefore, we welcome up¬
dates, corrections or additions (please send to the first author).

Phytogeographical Setting

For purposes of describing Carex distribution patterns in this paper, we define
five phytogeographic sections of Maryland and the District of Columbia (Figure 1),
which are based on physiographic provinces (Schmidt 1993). With the exception of one
boundary between two sections, our phytogeographical sections also correspond to
the five (vegetation) districts within three floristic zones described by Shreve et al.
(1910).

The Allegheny Plateau phytogeographic section (Figure 1) corresponds to the
physiographic province of that name as described by Schmidt (1993), and to the Moun¬
tain Zone of Shreve et al. ( 1 9 1 0). It consists of that part of Maryland west of the Allegh¬
eny Front, including all of Garrett County and extreme western Allegany County (Figure
1). Most of this section lies above 610 meters (above 2000 feet) above sea level (ASL).
It is comprised of a dissected plateau underlain by sedimentary rocks of late Paleozoic
age (Schmidt 1993). Vegetation is transitional between the northern Hemlock- White

Winter 2001

43

C. T. Frye & C. Lea

Figure 1. Map of Maryland and the District of Columbia, with counties. Phytogeographic sections are separated by boid lines and labeled
in capital letters.

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

Pine -Northern Hardwoods forest region and the Oak-Chestnut forest region character¬
istic of much of the Appalachian Mountains (Braun 1950). Accordingly, this section has
a number of floristic elements characteristic of glaciated regions to the north of Mary¬
land. It has vegetation formations unique or nearly so within the state, including north¬
ern hardwood forest associations and extensive non-tidal peatlands. Of all of the phyto¬
geographic sections in Maryland, it is perhaps the most floristically distinct.

The Ridge and Valley phytogeographic section (Figure 1), as it is used in this
paper, includes both the Valley and Ridge, and the Blue Ridge physiographic provinces
of Schmidt (1993). In Maryland, this section extends east from the Allegheny Front at
the eastern edge of the Allegheny Plateau through the folded Appalachians, which
represent the Valley and Ridge physiographic province. The Valley and Ridge province
is underlain entirely by sedimentary rocks of Paleozoic age, with resistant strata (prima¬
rily sandstones) forming elevated ridges that alternate with valleys occupied by more
easily weathered strata composed primarily of shale and limestone (Schmidt 1993). The
forest soils of the ridges tend to be somewhat acidic, while carbonate rocks in the
valleys produce a more neutral substrate supporting a characteristic flora that includes
a number of calciphilic species. Valley floor elevations generally range from 120 to 245
meters ASL, while ridge crests are generally from 365 to 610 m AST. In addition to the
(geologically defined) Valley and Ridge physiographic province of Schmidt (1993), our
Ridge and Valley phytogeographic section includes the smaller and floristically similar
Blue Ridge physiographic province to the east. The latter province is comprised prima¬
rily of resistant metamorphic rocks of early Paleozoic to Precambrian age and is repre¬
sented in Maryland by South Mountain, the Catoctin Creek valley, and the Catoctin
Mountains (Schmidt 1993). The entire Ridge and Valley phytogeographic section in¬
cludes most of Allegany County, all of Washington County, and western Frederick
County. It corresponds to the Upper Midland floristic district of the Midland Zone of
Shreve et al. (1910), with the exception of the Frederick (Monocacy) Valley and Parrs
Ridge areas (the inner Piedmont). It lies entirely within the Oak-Chestnut forest region of
Braun (1950), and is floristically transitional between the Allegheny Plateau and the
Piedmont sections, though more similar to the latter.

The Piedmont phytogeographic section (Figure 1) corresponds to the Piedmont
physiographic province of Schmidt ( 1 993), and to the entire Lower Midland district and
the eastern part of the Upper Midland district of Shreve et al ( 1 9 1 0). It extends from the
eastern base of the Catoctin Mountains to the Fall Line to include all of Carroll and
Montgomery Counties, nearly all of Howard County, most of Frederick, Baltimore, Harford,
and Cecil Counties, extreme northwestern Prince Georges County, and the western parts
of the District of Columbia and the city of Baltimore. Elevations generally range from 60
to 245 m ASL, with lower elevations occurring along Fall Line sections of rivers. The
Piedmont is the most geologically complex physiographic province in Maryland (Schmidt
1993). The eastern part (the outer Piedmont or Piedmont Plateau) is underlain by resis¬
tant metamorphic rocks of Precambrian and early Paleozoic age, while the Frederick
Valley to the west (inner Piedmont) is occupied by more easily weathered sedimentary
strata of Paleozoic to early Mesozoic (Triassic) age (Schmidt 1993). The Piedmont phy¬
togeographic section also lies entirely within the Oak-Chestnut forest region and is
floristically most similar to the Ridge and Valley section. It has considerable localized

Winter 2001

45

C. T. Frye & C. Lea

habitat diversity created by a variety of bedrock formations, including felsic, mafic, and
calcareous types, and by the high gradient reaches of rivers along the Fall Line.

The Western Shore phytogeographic section (Figure 1) includes all of the Coastal
Plain physiographic province of Schmidt (1993) that is west of the Chesapeake Bay and
the Elk River, and has the Fall Line as its western boundary (Figure 1). It is identical to
the Western Shore district of the Coastal Zone of Shreve et al. (1910) and includes all of
Anne Arundel, Charles, Calvert, and St. Mary’s Counties, nearly all of Prince Georges
County, small parts of Howard, Baltimore, Harford, and Cecil Counties, and the eastern
parts of the District of Columbia and the city of Baltimore. Elevations range from sea
level to less than 60 ASL. This section is underlain by unconsolidated sediments that
show a progression from more elevated, older (Cretaceous and Tertiary) formations of
modest topographic relief in the western part of the section to uniformly low-lying,
younger (Quaternary) sediments adjacent to the Chesapeake Bay and the larger rivers
(Schmidt 1993). The Western Shore section is floristically diverse, evidently because it
is transitional between the Oak-Chestnut forest region of the Piedmont and the Oak-
Pine forest region of the coastal southeastern United States (Braun 1950) and includes
flora elements typical of both areas. The latter region is prevalent on the Eastern Shore,
but also occupies the lower, younger formations along estuaries on the Western Shore.

The Eastern Shore phytogeographic section (Figure 1) includes all of the Coastal
Plain physiographic province of Schmidt (1993) that lies east of the Chesapeake Bay
and the Elk River. It is identical to the Eastern Shore district of the Coastal Zone of
Shreve et al. (1910). It includes all of Kent, Queen Anne’s, Talbot, Caroline, Dorchester,
Wicomico, Somerset, and Worcester Counties and southeastern Cecil County. Eleva¬
tions are mostly less than 20 m ASL. This section is underlain by relatively young
(Tertiary and Quaternary) sediments of uniformly low relief (Schmidt 1993). It is mostly
within the Oak-Pine forest region, with the northernmost counties transitional to, or
within, the Oak-Chestnut region. Significant Eastern Shore habitat features include
extensive fresh and salt tidal marshes, blackwater river swamps, barrier islands, and
Delmarva Bays (seasonal ponds).

Methods

From 1 998 to 200 1 , we examined approximately 4800 herbarium specimens held by
the following collections: BALT, GMUF, MARY, NA, PH, US (acronyms follow Holmgren
et. al. 1990), Anne Arundel Community College (Arnold, Maryland), Assateague Island
National Seashore (Berlin, Maryland), Cylbum Gardens and Arboretum (Baltimore,
Maryland), Delaware Natural Heritage Program (Smyrna, Delaware), Frostburg State
University (Frostburg, Maryland), National Capital Parks East (Washington, DC), U. S.
Geological Survey, Water Resources Division (Reston, Virginia), Tawes Herbarium (De¬
partment of Natural Resources, Annapolis, Maryland), and several private collections.
In the great majority of cases, a specimen examined by one or both of us was used as the
basis for a county record. For a small number of cases, we accepted other credible
records (e.g., from specimens examined by specialists in the group and reported to us,
credible published records of specimens not seen by us, and Natural Heritage sight
records of easily identified taxa). Collections containing vouchers used in this study but
not examined by us include some at DOV, GH, MICH, NCU, KNK, NY, OS, VPI, and WVA.

46

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

Additionally, we devoted approximately 360 person-days during this period to
floristic field study of Carex in Maryland and the District of Columbia. Although we
visited each county of Maryland and the District of Columbia on at least two trips, we
specifically allocated portions of our efforts to cover areas that our collection review
suggested were historically less well-collected (e.g., several counties on the lower East¬
ern Shore and in the northern Piedmont). We also directed special effort and attention to
taxa that appeared to be poorly represented in collections relative to their probable
abundance, often due to cryptic habits, early fruiting phenologies, and/or identification
challenges (e.g., Sections Acrocystis and Ovales). Finally, we also concentrated on
groups of taxa that were taxonomically not well known in the area (e.g.. Section
Stellulatae , the Carex amphibola complex, and infraspecific taxa).

In the field, we identified individual Carex plants, and made notes on the location,
habitat, and relative abundance of individual taxa. We collected approximately 3,500
specimens, as needed for more intensive identification work and/or as vouchers. A
dissecting microscope (10-60 x) and an ocular micrometer often assisted laboratory
examination. Specialists, particularly A. A. Reznicek, provided assistance with some
taxonomic determinations. Our collections have been deposited chiefly at the Tawes
Herbarium, MARY, MICH, US, DOV, Frostburg State University, and Anne Arundel
Community College.

From the collection research and fieldwork, we documented the presence of each
taxon verified for Maryland or the District of Columbia for each of its counties of occur¬
rence as a record in a database maintained by the Maryland Department of Natural
Resources, Wildlife and Heritage Service. The District of Columbia is treated as a county,
while records for the city of Baltimore are included under those for Baltimore County.
Each county record includes, minimally, the taxon name, county, date of collection or
observation, collector or observer name, collector number, and name of collection re¬
pository (if applicable). Details of individual county records may be obtained by con¬
tacting the first author.

To describe phytogeographical patterns of the native Carex taxa in Maryland,
we employed a principle components analysis (PC A) (Gauch 1982; Pielou 1984) of the
1 60 native taxa in counties space, using binary data in the original data matrix (taxon
presence in a county scored as 1; taxon absence scored as 0). The data were centered
by the column (county) means. We used PC Ord v. 3.04 (McCune and Mefford 1997)
for calculations.

Results and Discussion

Results Summary

We recognize 158 species and 175 total taxa for Maryland and the District of
Columbia (see annotated list). These numbers are the result of the additions of 29 taxa
(summarized in Table 1) and deletions of 10 taxa (Table 2) to the total represented by the
combined, and taxonomically reconciled, listings of Hitchcock and Standley (1919) and
Brown and Brown (1984). Our own fieldwork during this study produced six of the
additions ( Carex species 1, C arctata, C. corrugata , C. digitalis var. asymmetrica, C
haydenii, and C. laxiculmis var. eopulata ), and new determinations of specimens dur¬
ing our review of existing collections uncovered three other newly reported taxa (C.

Winter 2001

47

C. T. Frye & C. Lea

Table 1 . Additions to the Carex flora of Maryland and District of Columbia reported by Brown and
Brown (1984) and/or Hitchcock and Standley (1919), with remarks. Taxa added from fieldwork
of this study are denoted by a single asterisk (*). Taxa added from critical review of herbarium
specimens by this study are denoted by a double asterisk (**).

TAXON

REMARKS

Carex acutiformis Ehrhart
Carex aestivaliformis Mackenzie
Carex annectens (Bicknell) Bicknell
vac xanthocarpa (KiikentM) Wiegand

Carex arctata W. Boott

Carex austrina (Small) Mackenzie

Carex corrugata Femald

Carex crinita Lamarck var. hrevicrinis Femald

Carex diandra Schrank
Carex distans Linnaeus
Carex ehurnea F. Boott

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

Carex gravida Bailey var. lunelliana (Mackenzie)
Hermann

Carex haydenii Dewey
Carex species 1

Carex laxiculmis Schweinitz var. copulaia
(Bailey) Femald

Carex leptonervia (Femald) Femald

Carex lucorum Willdenow ex Link var. lucorum

Carex michauxiana Boeckeler

Carex mitchelliana M. A. Curtis

Carex oxylepis Torrey and Hooker var. oxylepis

Carex pedmcidata Muhlenberg ex Willdenow

Carex planispicata Naczi

Carex richardsonii R. Brown

Carex stipata Muhlenberg ex Willdenow var.

maxima Chapman

Carex striata Michaux var. brevis Bailey

Carex striata Michaux var. striata

Carex tonsa (Femald) Bicknell var. rugosperma

(Mackenzie) Crins

Carex tuchermanii F. Boott ex Dewey

First known Maryland collection in 1997.**

One known Maryland collection, in 1946.

Several Maryland collections, as early as 1888; infraspecific
taxa for C. annectens not recognized by Brown and Brown or
by Hitchcock and Standley.

First known Maryland collection in 2001.*

First known Maryland collection in 1998.

First known Maryland collection in 1998.*

Numerous Maryland collections found; infraspecific taxa for C.
crinita not recognized by Brown and
Brown or by Hitchcock and Standley.

First known Maryland collection in 1987.

One known Maryland collection, in 1946.

First known Maryland collection in 1986.

Several Maryland collections, as early as 1873; infraspecific
taxa for C. granularis not recognized by Brown and Brown
or by Hitchcock and Standley.

First known Maryland collection in 1997.

First known Maryland collection in 2000.*

First identified for Maryland from a collection in 2001.* A
second collection (1915) subsequently found.

First known Maryland collection in 2000.*

Brown and Brown listed taxon, but noted no Maryland
records; first known Maryland collection in 1981.

Several Maryland collections, as early as 1881 ; may have been
included in concept of C. pensylvanica by Brown and Brown.
Brown and Brown listed taxon, but noted no definite
Maryland records; first known Maryland collection in 1981.

First known Maryland collection in 1996.

First known Maryland collection in 1997.**

Brown and Brown listed taxon, but noted no Maryland
records; first known Maryland collection in 1987.

Recently described and reported for Maryland and for District of
Columbia by Naczi (1999); several collections (as C. amphiboia
or C. oligocarpa), as early as 1899.

Several Maryland collections, as early as 1951.

Numerous Maryland collections, as early as 1912; infraspecific
taxa for C. stipata not recognized by Brown and Brown or by
Hitchcock and Standley.

Several Maryland collections, as early as 1983. Listed for Eastern
Shore by Tatnall (1946), but not for Maryland by Brown and
Brown (1984).

First known Maryland collection in 1993.**

Several Maryland collections, as early as 1939; illustrated as
C. umbellata var. rugosperma (probably a nomen nudum),
but not otherwise treated, by Brown and Brown.

First known Maryland collection in 1991.

48

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Gyperaceae) of Maryland and the District of Columbia

acutiformis, C. oxylepis , and C. striata van striata ), Other additions were based on
collections of previously unreported taxa by other field workers since 1984, on speci¬
mens that we found in herbaria that had been collected before 1984 but not reported in
either flora, and on confirmed collections of taxa reported as hypothetical by Brown and
Brown (1984). All deletions are based on our critical review of herbarium collections and
other sources; in some cases, we found no basis for a report of a taxon in the herbarium
collections, and in others we found a purported presence of a taxon to be based on a
misidentified specimen.

Phytogeographical Patterns of Carices in Maryland and the District of Columbia
Figure 2 depicts a scatter plot (ordination diagram) of the first and second axis
scores generated by the PCA for individual native taxa. Phytogeographic affinities of
groups of taxa are delineated as we have interpreted them from visual examination of the
scatter plot and from supplemental knowledge gained from our field experience. Table 3
lists eigenvector loadings on each of the first two PCA axes generated by each county.
While the PCA scatter plot effectively shows general distribution patterns within the

Table 2. Carex taxa excluded for Maryland and District of Columbia. Taxa listed were
originally reported by Brown and Brown (1984), or by Hitchcock and Standley (1919).

TAXON REMARKS .

Carex alopecoidea Tuckerman

Carex bicknellii Britton
Carex foenea Willdenow

Carex intumescens Rudge var.
fernaldii Bailey

Carex nigra (Linnaeus) Reichard

Carex novae-angliae Schweinitz

Carex reirorsa Schweinitz
Carex schweinitzii Dewey

Carex tenuiflora Wahlenberg

Carex venusta Dewey var. venusta

Reported (as hypothetical) by Hitchcock and
Standley, based on a report by Holm; a specimen so
labeled at US was misidentified.

Reported by Brown and Brown, who noted no
Maryland records; no specimens found.

Reported by Brown and Brown; no specimens found;
all Maryland collections seen labeled or annotated as
C. foenea were determined to be C argyrantha, a
taxon to which the name C. foenea has been misapplied.
Reported by Brown and Brown; no specimens found.

Reported (as hypothetical) by Hitchcock and Standley
as C. goodenovii Gay, based on C. vulgaris Fries, as
listed by Ward (1881); no specimens found. Most likely
a misidentification of, or a name misapplied to, C.
emoryi.

Reported by Brown and Brown; specimen at MARY so
labeled was misidentified.

Reported by Brown and Brown; no specimens found.
Included in Brown and Brown, who noted no definite
Maryland records; no specimens found.

Reported by Brown and Brown; specimen at MARY so
labeled was misidentified.

C. venusta reported by Brown and Brown, who noted
that most Maryland plants are var. minor . All C. venusta
in Maryland found during this study are var. minor.

Winter 2001

49

C. T. Frye & C, Lea

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bullata

complanata

albicans v. emmonsii comosa stipata v maxima
albolutescens

styloflexa

louisianica
muhlenbergii v. muhlenbergii

hyalinolepis

giaucescens

mitcheliiana

venusta

annectens v. annectens

nigromarginata folliculata

. umbellata STATEWIDE typhina

tonsa v. tonsaat!antica s_ at|antica

striatula

vestita

lupuliformis

canescens v. canescen§» digitalis v. macropoda
- . . 1 corrugata

straminea )

... - . . • . r--

canescens v. disjunct^,
lacustris deBilis v. interc _ _

digits,

brevior

silicea

aestivaliformis

ntercursa striata v. striata
digitalis v. asymmetrica

blanda \

tribuloides^ crinita v. crinita

WESTERN SHORE

species

LI

caroliniana

^-.-'Tichardsonij/

radiata pensylvanica
laxicuimis v. laxicuimis ,

V in 1
glaucode?

cephalophora frankii 7
digitalis v. digitalis

rosea j

scoparia laxiflora

STATEWIDE, RARE ON la"regata /

\ mesochorea '

festucacea ) / PIEDMONT

tenera

PIEDMONT AND
WESTERN SHORE

granuiarisv. haleana .meadn . leptonervia

MONT hySteriCina
. .. JjTiolesta trichocarpa \

hitchcockiana£|fgSTO^eb‘!i® v- Pubera
buxbaumii]0 9 “ Plcabrata.

jarnesii-2 iU o.

pe,j

,bu§WU

conoideaf utncufata

aavisii/appa|achica

■antha

LOWER EASTERN SHORE \

/''APPALACPHIAN7pfEDMONT,VfranUlariSV'9ranu!anS . wopdii
! AND WESTERN SHORE-Mst.. JH8®!. albursinaemoryi

APPALACHIAN
(some with Piedmont
disjunctions)

debilis v. debilis

intumescens

laevivaginata

lupulina

lurida

stricta

swanii

vulpinoidea

hirsutella MpThlenbergi v. enervisconiuncta/TepFalea s. leptalea
prasina squarrosa gynandra / tonsa v. rugosperma

albicans v. albicans! platyphylla
wilidenowii grayi / normal!

virescens 1

I hirtifolia

stipata v. stipata, gracilescens
gracillimp/|0rta

_______ ~.-_~--comjTlur)ls

APPALACHIAN AND PIEDMONT

ALLEGHENY PLATEAU ONLY

aestivalis haydenii sartweilii

arctata lasiocarpa trisperma

baileyi michauxiana tuckermanii

debilis v. rudgei plantaginea vesicaria

diandra projects

-2

-1

0 1

Axis 1 Score (42.5% of variance)
Widespread Taxa Restricted Taxa

Figure 2. Ordination diagram (scatter plot) showing position of individual native Mary¬
land and District of Columbia Carex taxa in counties space as determined by scores
on first two RCA axes. Dash lines separate our interpretations of phytogeographically
similar groups of taxa (affinities denoted in capitals). Positions of names of some taxa
in the scatter plot have been moved minimally from the actual positions occupied by
the points representing these taxa for clarity. “Appalachian” refers to the Allegheny
Plateau and the Ridge and Valley phytogeographic sections combined, “Coastal Plain”
refers to the Eastern Shore and Western Shore phytogeographic sections combined.

50

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyper aceae) of Maryland and the District of Columbia

entire Carex flora, and suggests geographical affinities among taxa, the user of this atlas
should rely on individual taxon descriptions and the associated maps as the most
accurate representation of distribution for each taxon. The positions of a very few taxa
on the scatter plot are “misclassified” with respect to the best phytogeographic inter¬
pretation of the groups.

Axis 1 (42.5%) and 2 (16.2%) together accounted for 58.7% of the total variance in
the data. Only these axes generated eigenvalues that exceeded their broken-stick eigen¬
values, accounted (therefore) for more variation than would be expected by chance
(Jackson 1993), and were considered for interpretation. The largest positive (least nega¬
tive) eigenvector loadings on Axis 1 (Table 3) were produced by counties with fairly
distinctive Carex flora, including the Allegheny Plateau county of Garrett (particularly)
and the lower Eastern Shore counties of Somerset, Wicomico, and Worcester. The larg¬
est negative loadings for Axis 1 were represented by those of the central counties,
particularly the Western Shore counties of Anne Arundel, Charles, and Prince Georges
and the District of Columbia. The taxa groupings along Axis 1 of the PCA scatter plot
(Figure 2) show a progression from very restricted and often rare taxa on the right side
of the plot to widespread and common taxa on the left side. For Axis 2, the largest
positive eigenvector loadings were produced by the southern Eastern Shore counties
of Worcester, Wicomico, Dorchester, Caroline, and Somerset, while the counties with
the largest negative loadings are those partially or completely within in the Ridge and
Valley phytogeographic section, including Washington, Allegany, and Frederick.

The source of the greatest phytogeographical variation in Maryland’s Carex flora
defected by the PCA (represented primarily by Axis 1) is the degree of distinctiveness of
the local (county-level) flora. The relative specialties of the Allegheny Plateau and the
lower Eastern Shore produce a contrast with the widespread taxa in the flora of the
Western Shore and, to a lesser degree, the Piedmont and the upper (northern) Eastern
Shore. A second important source of floristic variation (represented primarily by Axis 2)
is the gradient of elements characteristic of the Coastal Plain in the eastern part of the
state to montane elements in the western areas. We note that much of the variance
accounted for by Axis 1 might be considered trivial for phytogeographical interpreta¬
tion. While phytogeographers generally assign greater significance to taxa co-occur¬
rences than to co-absences, the PCA is incapable of this type of [necessarily subjec¬
tive] differentiation. Two hypothetical taxa, each occurring in a single county at oppo¬
site ends of the state, would be more similar to each other (from identical frequencies of
zero in 22 of the 24 counties) than they would be to a species that occurs with them and
also in all other counties. Thus, a taxon’s position along Axis 1 is largely determined
simply by the number of counties that it is found in. Nevertheless, once this variation is
accounted for, patterns in Axis 2 that are more clearly geographical allow distinct affili¬
ations among groups of taxa to be seen in the scatter plot (Figure 2).

Several other floristic patterns are suggested by the PCA results. The progres¬
sion of eigenvector loadings on Axis 2 (righthand column of Table 3) suggests that the
Coastal Plain counties are floristically more related (with respect to Carex ) to the Al¬
legheny Plateau county of Garrett than they are to counties that lie entirely within the
less distant Ridge and Valley or the Piedmont sections. We note two phenomena that
may explain this somewhat counter-intuitive observation. First, several taxa (e.g., Carex

Winter 2001

51

C. T. Frye & C. Lea

Table 3. Eigenvector loadings for each county for first two axes of RCA (Figure 2).

COUNTY

Axis 1

Eigenvector

COUNTY

Axis 2
Eigenvector

Garrett

-0.0604

Worcester

0.3413

Somerset

-0.1374

Wicomico

0.3213

Wicomico

-0.1559

Dorchester

0.2683

Worcester

-0.1705

Caroline

0.2549

Carroll

-0.1817

Somerset

0.2475

Allegany

-0.1818

Calvert

0.1769

Dorchester

-0.1889

Saint Mary’s

0.1599

Saint Mary’s

-0.1942

Kent

0.1298

Frederick

-0.1944

Talbot

0.1212

Washington

-0.2039

Queen Anne’s

0.1157

Talbot

-0.2103

Anne Arundel

0.0829

Montgomery

-0.2114

Charles

0.0353

Calvert

-0.2144

Prince Georges

-0.0264

Caroline

-0.2176

District of Columbia

-0.1166

Howard

-0.2201

Cecil

-0.1189

Cecil

-0.2203

Baltimore

-0.1507

Harford

-0.2216

Harford

-0.1527

Baltimore

-0.2218

Garrett

-0.1856

Queen Anne’s

-0.2231

Howard

-0.1928

Kent

-0.2240

Montgomery

-0.1995

Prince Georges

-0.2341

Carroll

-0.2595

District of Columbia

-0.2358

Frederick

-0.2606

Charles

-0.2379

Washington

-0.2707

Anne Arundel

-0.2464

Allegany

-0.2789

atlantica ssp. capillacea , C. canescens var. disjuncta , C. lacustris, and C. seorsa)
show a disjunct distribution, occurring throughout the Coastal Plain and in Garrett
County, and being largely absent in the areas between. These are taxa that show moder¬
ate to strong preference for depressional wetlands, often those with some peat forma¬
tion, such as seepages and fens. These habitat features are more extensive, if not more
frequent, on both the Allegheny Plateau and on the Coastal Plain than they are in the
intervening areas. Our field observations also indicate that other taxa of these habitats
show a more or less continuous distribution across Maryland (e.g., Carex atlantica
ssp. atlantica , C. hromoides , C. folliculata , C. leptalea (both subspecies considered
together as a species), and C stricta occur more frequent on the Allegheny Plateau and
the Coastal Plain than in the intervening areas. This guild of species seems to comprise
a notable phytogeographic element in the state. On the other hand, most taxa that occur
with significant frequency on both sides of the Fall Line (i.e., that occur widely both in

52

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

the Coastal Plain and in the Ridge and Valley/Piedmont sections) are also widespread
throughout the state and occur on the Allegheny Plateau, as well. Thus, there is no
exclusively Ridge and Valley/Piedmont/Coastal Plain guild comparable to that noted for
the Allegheny Plateau/Coastal Plain.

Eigenvector loadings on Axis 2 (Table 3) suggest that the Western Shore counties
of Calvert and Saint Mary’s are floristically more affiliated (with respect to Carex) with
Eastern Shore counties than to other Western Shore counties. This probably reflects
geologic and phytogeographic similarities between the former two counties and the
Eastern Shore, including the relative predominance of lower, Quaternary age landforms
(Schmidt 1993), and of the less Carex- rich Oak-Pine forest region (Braun 1950).

The relatively high density of taxa depicted in the lower right quadrant of the
scatter plot reflects a higher contribution to taxa richness in Maryland and the District of
Columbia from taxa most affiliated with the Piedmont and Appalachian areas (primarily
within the Oak-Chestnut forest region), compared to those more associated with coastal
areas (primarily within the Oak-Pine forest region). This pattern is consistent with the
observations of Harvill (1973), who noted that Carex is predominantly a genus of tem¬
perate regions, in contrast to most other genera in Cyperaceae occurring in Virginia,
which have more southern (tropical) affinities.

Figure 3 depicts numbers of native taxa and total taxa reported for each county.
For each phytogeographic section, numbers of native taxa, total taxa, and taxa unique to
the section are shown. The eastern counties of the Piedmont section and, to a lesser
extent, the western counties of the Western Shore section exhibit the greatest taxa
richness on a county scale. This pattern appears to be largely attributable to the posi¬
tion of these counties at or near the transition between the Oak-Chestnut and the Oak-
Pine forest regions, where taxa more limited to one region or the other may be sympatric
near their respective range limits. Within this county group, the eastern Piedmont county
totals are slightly higher than those of the westernmost Western Shore counties be¬
cause of their position within the Carex- rich (as suggested by the PCA) Oak-Chestnut
forest region and to greater habitat diversity in the Piedmont due to its geologic com¬
plexity. As an example of this effect, Lea and Frye (2002) reported 90 taxa for an area of
about 1,000 hectares near the Fall Line in the eastern Piedmont. The lowest county taxa
richness totals generally occur in counties of the Eastern Shore section and in the
southeastern counties of the Western Shore section. These areas lie within the Oak-
Pine forest region, an area apparently less taxa rich with respect to Carex (Figure 2).
Restricted taxa occur most frequently on the Allegheny Plateau and the lower Eastern
Shore, which have comparatively moderate to low taxa richness but which are phyto-
geographically distinctive within Maryland, and also in the Piedmont, where specialized
habitats associated with unusual geologic situations (e.g., serpentine barrens, Fall Line
reaches of major rivers) harbor some rare taxa. Although taxa richness patterns may be
somewhat influenced by the artifact of historically greater collecting efforts in the cen¬
tral areas, much of our own recent fieldwork, in contrast, has focused on historically
poorly collected counties. Although small collecting “gaps” remain, we consider the
level of collecting effort to be now thorough enough statewide so that presence/ab¬
sence data effectively reflect true taxa richness patterns.

A number of the Maryland Carex flora are taxa at or near the limits of their known

Winter 2001

53

C. T. Frye & C. Lea

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54

The Maryland Naturalist

Volume 44

to the phytogeographic section within Maryland.

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

geographic range in the eastern United States. These include seven taxa more common
in the northern Appalachians and New England, eight taxa of the southeastern United
States Coastal Plain at the northeastern limit of their range, and two midwestem species
at their eastern range limits (Table 4).

Non-native species represent a fairly minor part of the Carex flora, with fifteen
that are not indigenous to Maryland and the District of Columbia represented (one
species native to Maryland, C. plantaginea is introduced, in part). Most non-native
species appear to be waifs, rather than well naturalized taxa, with nine of the fifteen
known only from a single site. Only one, Carex kobomugi , is considered to be competi¬
tively invasive (Small, 1954). Section Phaestoglochin , a group that includes a high
proportion of “weedy” species, accounts for seven of our fifteen non-indigenous Carex
species, and an eighth species (C. aggregata ) that is possibly introduced.

Potential and Historic Carex flora of Maryland and the District of Columbia

In order to stimulate interest in further floristic exploration, we list (Ta ble 5)
Carex taxa that have not yet been recorded in Maryland or the District of Columbia,
and which we believe to be among the most likely to be found as new occurrences for
our area. In general, these are taxa that are reported from adjoining counties of a
neighboring state or from two or more adjoining states, or both (e.g., Tatnall 1946;
Strausbaugh and Core 1978; Harvill et al. 1992; Rhoads and Klein 1993; Cusick 1996; A.
S. Weakley, personal communication). Additionally, 10 native Carex taxa historically
recorded in our area remain “lost.” To our knowledge these taxa have not been reported
since 1980. Eight of these are known from a single county each, and six are represented
by a single collection each. With year of their last known collection, the 10 taxa are:
Carex aestivaliformis (1946), C. annectens var. xanthocarpa (1965), C. cephaloidea
(1899), C decomposita (1920), C. lucorum [var. lucorum ] (1962), C. pallescens (1898),
C. polymorpha (1891), C. sartwellii (1938), C. tetanica (1925), and C. viridula (1879).
Among non-native species, we know of no recent reports for five: C. arenaria (1900),
C. caryophyllea ( 1 880s), C. distans (1948), C. divulsa( 1896), and C. muricata (1969).
All five are known from a single county each, and only C. arenaria is represented by
more than a single collection.

Table 4. Carex taxa at or nearly at their eastern United States range limits in Maryland
or the District of Columbia. Based on Tatnall (1946), Fernald (1950), Strausbaugh and
Core (1978), Grins and Ball (1983), Grins and Ball (1989b), Harvill et ah (1992),
Rhoads and Klein (1993), A. S. Weakley (personal communication), Harmon and
Ford-Werntz (2000), and Association for Biodiversity Information (2001).

MAJOR EASTERN UNITED

STATES RANGE

TAXA

Northern Appalachians/New England

cephaloidea, diandra, haydenii, lucorum var.
lucorum, michauxiana, sartwellii, viridula

Southeastern Coastal Plain

corrugata, debilis var. intercursa, digitalis var.
asymmetrica, digitalis var. macropoda,
gigantea, glaucescens , species 1 , oxylepis

Midwest

careyana, richardsonii

Winter 2001

55

C. T. Frye & C. Lea

Table 5. Carex taxa not yet recorded for Maryland and the District of Columbia, but
have the potential to occur and to be sought as possible additions to flora [neighbor¬
ing states with occurrences denoted in brackets].

TAXON

REMARKS

Carex albicans

Willdenow ex Sprengel van australis
(Bailey) Petti g

Dry woods. Specimens from Great Falls area in
Maryland have some characters of this southern
taxon (= C. physorhyncha Liebman), but are
problematic (Lea and Frye, 2002). [VA],

Carex atherodes Sprengel

Seeps and fens. Most likely to occur in Garrett
County. [PA, VA, WV].

Carex bicknellii Britton

Serpentine barrens. Most likely to occur in north
eastern Piedmont, [DE, PA].

Carex crawei Dewey

Calcareous outcrops, meadows, shores. Most likely
to occur in Ridge and Valley section. [VA].

Carex deflexa Homemann

In Appalachians, moist, high-elevation meadows.
Most likely to occur in Garrett County. [WV].

Carex limosa Linnaeus

Peaty wetlands, often with Sphagnum. Most likely
to occur in northeastern Piedmont. [DE, PA].

Carex novae-angliae Schweinitz

Sandstone outcrops; moist woods. Most likely to
occur in Garrett County. [PA, WV].

Carex oligosperma Michaux

Bogs, poor fens. Most likely to occur in Garrett
County. [PA, VA, WV].

Carex ormostachya Wiegand

Rocky woods, shale barrens. Most likely to occur in
western Maryland. [PA, VA].

Carex pauciflora Lightfoot

Bogs, poor fens, often with Sphagnum. Most likely
to occur in Garrett County. [PA, WV].

Carex praegracilis W. Boott

Western species, apparently spreading in eastern
United States along railroads and salted highways
(Reznicek and Catling 1987). Most likely to occur in
western Maryland. [PA, VA].

Carex prairea Dewey in A. Wood

Calcareous wetlands. Most likely to occur in Ridge
and Valley section. [PA, VA, WV].

Carex retrorsa Schweinitz

Swamps, stream banks, swales. Most likely to occur

in northern tier of counties. [PA].

Carex schweinitzii Dewey ex Schweinitz Calcareous wetlands. Most likely to occur in Ridge

Carex siccata Dewey

and Valley section. [PA, VA].

Sandy woods. Most likely to occur in northeastern
Piedmont. [PA].

Carex sprengelii Dewey ex Sprengel

Stream banks, moist woods, outcrops. Most likely to
occur in northern tier of counties. [DE, PA].

Carex sterilis Willdenow

Calcareous wetlands. Most likely to occur in Ridge

Carex suberecta (Olney) Britton

and Yalley section. [PA, VA, WV].

Calcareous wetlands. Most likely to occur in Ridge
and Valley section. [VA, WV].

Carex venusta Dewey van venusta

Seepage wetlands, swamps. Most likely to occur in
Charles or St. Mary’s Counties. [VA].

56

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia
General Comments on Annotated List of Taxa and Distribution Maps

Information for each individual Carex species or infraspecific taxon that we report
for Maryland and the District of Columbia is presented in an annotated list, arranged in
alphabetical order by Latin name. The entry for each taxon is accompanied by a map of
Maryland and the District of Columbia with verified county occurrences for that taxon
delineated by dots in the applicable counties. Figure 4 provides an example of how to
interpret the information presented in the annotated list.

In general, our treatments for the Latin names and authors of species and in¬
fraspecific taxa follow Kartesz and Meacham (1999), with some modifications. Where
consensus is divided on whether a taxon should be recognized individually or not, we
have tended to take a “splitter’s” approach whenever our examination of Maryland
material suggested that reasonably consistent and possibly significant phenetic, phy-
togeographical, and/or ecological differences could be detected. Such an approach
helps to ensure that information that may be valuable for future resolution of taxonomic
issues is not lost.

Common names are derived from a variety of sources, including Reed (1988),
Maryland Natural Heritage Program ( 1 994) and U.S. Department of Agriculture (200 1 ).
Where a common name was otherwise unavailable, we use an approximate translation of
the Latin specific or subspecific epithet. The use of particular common names here is not
intended to be authoritative or “standard.”

Infrageneric sectional taxonomy of Carex has undergone many recent changes in
order to incorporate the results of taxonomic research and to resolve nomenclatural
problems. Our sectional treatments follow those of Reznicek (personal communication);
some may be unfamiliar to users of older manuals (e.g, Femald 1 950; Brown and Brown
1984; Gleason and Cronquist 1991). In the annotated list, we list for each taxon the
section (our treatment), followed by a more traditional (alternative) sectional treatment
listed, if applicable (Figure 4). This should be considered a “crosswalk” for the user’s
convenience rather than a synonymy; some traditional section names may be syn¬
onyms; others may represent former or alternative sectional assignments for taxa.

In listing synonyms for taxa, we employ terms such as “in part,” “misapplied,” “in
the broad sense,” and “in the narrow sense” in their usual botanical meanings. It should
be understood that the application of these terms and other conclusions about syn¬
onymy sometimes has required our best judgement in the face of uncertainties about
other authors’ concepts.

For each taxon in the annotated list, we provide a brief description of diagnostic
characters, primarily those useful for differentating taxa from their closest relatives.
Citations of sources that describe identification, taxonomy, distribution, ecology, and /
or nomenclature are listed, when applicable. Descriptions of habitats apply to those
known for Maryland and the District of Columbia. Sources for the occurrences of taxa in
neighboring states include the state and regional references listed in Figure 4 and also
Rhoads and Klein (1993), Cusick ( 1 996), A. S. Weakley (personal communication), Harmon
and Ford-Wemtz (2000), McAvoy and Bennett (2001), and, occasionally, specimens in
collections seen by us. Generally, we report these as they have been listed by the

Winter 2001

57

C. T. Frye & C. Lea

Common Name

Section Name (Traditional Section Name)

L . “ ”J

Car ex umbellata Schkuhr ex Willdenow _ _▼

Umbellate Sedge | | Acrocystis (Montanae)

(KM, RB, T; in part only, - BR, GC, H, SC; C. abdita
Bicknell -- F; C. tonsa (Femald) Bicknell, in part — HS)

Densely cespitose sedge, with narrow (1-3 mm) leaves. Mature spikes
nearly hidden among leaf bases. Perigynium minutely pubescent, with

the beak half as long as the body. See Cusick (1992), Rettig and Crins (1996). _

Dry forests and openings. |tPE, PA, VA, WVlllCommon, PD, WS, ES; infrequent, RV.l

I

Alternative treatments

Alternative treatments are separated by
semicolons, with each treatment followed
by the source(s) that use it. Where a
treatment is not specified for a source,
refer to the immediately preceding name.

Sources

BR = Brown and Brown (1984)

F = Femald (1950)

GC = Gleason and Cronquist (1991)

H = Harvilletal. (1992)

HS = Hitchcock and Standley (1919)
KM = Kartesz and Meacham ( 1 999)

RB = Rhoads and Block (2000)

SC = Strausbaugh and Core (1978)

T = Tatnall (1946)

For the example above, Kartesz and
Meacham, Rhoads and Block, and Tatnall
also treat this taxon as C. umbellata
Schkuhr ex Willdenow, as we do; Brown
and Brown, Gleason and Cronquist,
Harvill et al., and Strausbaugh and Core
treat it as Carex umbellata Schkuhr ex
Willdenow (as do the immediately
preceding authors), but in part only;
Femald treats it as Carex abdita Bicknell;
Hitchcock and Standley treat it as Carex
tonsa (Bicknell) Femald, in part. (Author
abbreviations preceded by “infraspecific
taxa not distinguished” indicate that the
authors use the same species treatment as
we do, but do not recognize subspecies or
varieties).

Maryland-District of Columbia
Distribution Map.

Dots denote counties of occurrence

Distribution and abundance by phytogeographic section

Abundance*

Common

Frequent

Infrequent

Rare**

Naturalized or adventive***

Section Abbreviation****

AP = Allegheny Plateau
RV= Ridge and Valley
PD = Piedmont
WS = Western Shore
ES = Eastern Shore

* ordinal scale in decreasing order of abundance

** “Single station” indicates that there is one known
record of the taxon in our area (extant). “Single
Collection” indicates one known record (historical).

*** non-indigenous taxa (all can be considered rare,
where they occur); naturalized indicates that the
taxon is well established; adventive indicates that it is
likely a waif).

**** The symbol “-h” following a section abbreviation
indicates that the taxon is historical for [only] that
section (i.e., there are no known post- 1980 records).

States adjacent to Maryland where taxon occurs

DE Delaware

PA Pennsylvania

VA Virginia

WV West Virginia

none not known from surrounding states

** The symbol “-h” following a state abbreviation
indicates that the taxon is historical (not known to
be extant) for that state.

Figure 4. Example of an atlas taxon entry.

58

The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

source, as the scope of this project did not allow for our own specimen-based evalua¬
tion of reports from other states. A subjective ordinal scale (common, frequent, infre¬
quent, or rare) describes taxon abundance.

Acknowledgements

Mark Strong and other staff of the U.S. National Herbarium, Charles Delwiche of
the University of Maryland (Norton- Brown) Herbarium, Kevin Conrad of the National
Arboretum, Ted Bradley of George Mason University, Ernie Schuyler of the Academy of
Natural Sciences, and staff of the Cylbum Arboretum provided access to collections or
other support. Tony Reznicek of the University of Michigan made determinations of
some difficult specimens, shared unpublished information on Carex sectional nomen¬
clature, and generously provided invaluable advice. Bill McAvoy of the Delaware Natu¬
ral Heritage Program contributed a wealth of information on Eastern Shore records. Tom
Wieboldt of Virginia Polytechnic Institute and State University provided information on
Carex in Virginia. Rob Naczi of Delaware State University provided several determina¬
tions of and information on taxa in Sections Careyanae , Griseae , and Laxiflorae. Marcia
Waterway of McGill University, Lisa Standley, and Norbert Roeder shared information
on Carex debilis, Carex caryophyllea, and Carex distans, respectively. Alan Weakley
of Association for Biodiversity Information, P.J. Harmon of the West Virginia Depart¬
ment of Natural Resources, and Donna Ford-Wemtz of West Virginia University shared
preliminary drafts of work in progress and provided up-to-date taxa occurrence informa¬
tion for Virginia and West Virginia, respectively. Charles Davis, Frank Hirst, and Ron
Wilson made personal collections available for examination. Maryland Department of
Natural Resources volunteer Dan Siehl processed and mounted at the Tawes herbarium
a number of specimens collected during this study. Julia Mensler, formerly of the Worcester
County government, gave Geographic Information Systems guidance for producing
initial versions of the county occurrence maps. Ashton Berdine, Carol DiSalvo, Joe Kish,
Warren Steiner, Jil Swearingen, Ed Thompson, and Mary Troy are thanked for various
aspects of logistical support. Bill McAvoy, Tony Reznicek, Stan Shetler, and Rob Naczi
provided critical and useful comments to improve the manuscript. The National Park
Service, Maryland Department of Natural Resources, and the Maryland/DC Chapter of
The Nature Conservancy granted permission to collect specimens on lands they man¬
age. Funding was provided by the Washington Biologists’ Field Club Research Fund
and The Nature Conservancy’s Virginia E. Crouch Memorial Fund. Additional support
was provided by the Maryland Department of Natural Resources Wildlife and Heritage
Service and by the National Park Service, Assateague Island National Seashore.

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using achene morphology. Rhodora 88:399-403.

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C. T. Frye & C„ Lea

Carex abscondita Mackenzie
Thicket Sedge Careyanae (Laxiflorae)

(BR, F, GC, H, HS, KM, RB, T)

Forms distinct, leafy clumps. Perigynium trigonous, with sharp
angles. Similar to C. digitalis , but with lowest bract overtopping
inflorescence; upper bract spathe-like; the short (4-12 mm) terminal
(staminate) spike often hidden among the uppermost lateral (pistillate)
spikes. See Manhart (1986). Mesic to moist forests, with bottomland hardwoods or
loblolly pine; seepages. [DE, VA, WV-h]. Common, WS, ES.

Carex acutiformis Ehrhart
Lesser Pond Sedge Paludosae

(F, GC, KM)

Similar to C lacustris , but with leaves narrower; spikes more slenderly
cylindrical (6-7 mm wide); perigynia smaller (3. 5-4.0 mm long), papil¬
lose, and more compressed (flattened -tri gonous) . See Jenny et ah
(1982). Fresh marsh, [none]. Locally naturalized (European) (single station),
WS.

Carex aestivaliformis Mackenzie r — - — ; -y / i

False Summer Sedge Hymenochlaenae (Gracillimae) 1 ^ \[ \\ j

(GC, KM; C. x aestivaliformis — F, T; C aestivalis Schweinitz X X 5
x C. gracillima Schweinitz — RB) jfyH' ‘

Similar to C aestivalis , but leaves wider (2-6 mm) and spikes wider (3-4(>vAy
mm); perigynium orifice bidentate. Some authors treat as a hybrid V

between C. aestivalis and C. gracillima , but persuasive evidence for this has
not been presented. Rich forests. [PA, VA]. Rare (single collection), ES-h.

Carex aestivalis M, A. Curtis ex Gray I /aX / \ / k \A

Summer Sedge Hymenochlaenae (Gracillimae)

(BR, F, GC, H, KM, SC; C. aestivalis Schweinitz — RB) X

Lateral (pistillate) spikes relatively narrow and elongate, as in other - I , 1

members of section; terminal spike gynecandrous, as in C gracillima II

and C. prasina . Leaves 2-3 mm wide; leaf sheaths pubescent. Spikes ^ fX
loosely ascending, narrow (2-3 mm); perigynium beakless, with orifice entire.

Brown and Brown (1984) report likely based on a misidentified specimen; first known
Maryland collection was in 1984. Circumneutral mesic woodland. [PA, VA, WV]. Rare
(single station), AP

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Carex aggregate Mackenzie

Glomerate Sedge Phaestoglochin (Bracteosae)

(BR, F, HS, KM, RB, SC; C sparganioides

Muhlenberg ex Willdenow van aggregata (Mackenzie) Gleason —

GC; C. sparganioides , in part — H)

Similar to C. muhlenbergii , but leaf sheaths septate-nodulose, loose,
with inner bands firm, concave at summit. Lower spikes crowded; leaves
narrower (3-6 mm) than in C sparganioides. Open or forested habitats, often weedy.
[DE, PA, VA, WVJ. Frequent, RV, PD, WS; rare, ES. Perhaps naturalized (from farther
west).

Carex alata Torrey

Broad-winged Sedge Ovales

(BR, F, KM, GC, H, HS, RB, T)

Coarse sedge in tussocks; inflorescence crowded; perigynium broadly
winged, obovate (widest above the middle of the body); pistillate
scales acuminate to awned. See Rothrock et al (1997). Wet soils in
open areas, often along fresh tidal creeks. [DE, PA, VA]. Frequent, WS, ES.

Carex albicans Willdenow ex Sprengel var. albicans
White-tinged Sedge Acrocystis (Montanae)

(GC, KM; infraspecific taxa not distinguished — H;

C. albicans , in a narrow sense — RB; C. artitecta Mackenzie — BR, F,

SC, T; C. emmonsii Dewey, in part — HS)

Similar to C. pensylvanica , but cespitose, with narrower leaves (0.5- 1.5
mm), and shorter (0.35- 1.50 cm), sessile terminal (staminate) spikes; perigynium
body ellipsoid to fusiform-obovoid. Similar to var. emmonsii , but culms erect to
arching; midribs of staminate scales evanescent below scale apex. See Rettig (1989,
1990a, 1990b). Dry forests. [DE, PA, VA, WV], Common, RV, PD, WS; infrequent, ES.

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

Emmons’ Sedge Acrocystis (Montanae)

(GC, KM; C. emmonsii Dewey ex Torrey — BR, F, RB, T; in part only
— HS; infraspecific taxa not distinguished — H)

Distinguished from var. albicans by its decumbent culms (often hidden
in ground litter) and acuminate staminate scales, with strong, often excurrent,
midrib. See Rettig (1989, 1990a, 1990b), Cusick (1992). Dry forests, usually more acidic
than for the var. albicans. [DE, PA, VA]. Common, WS, ES; infrequent, PD.

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C T. Frye & C. Lea

Carex albolutescens Schweinitz
Green- White Sedge Ovales

(BR, F, GC, KM, RB; in part only - — H; C. straminea
Willdenow ex Schkuhr, misapplied — HS, T)

Similar to C. longii and C. festucacea. Perigyeium body obovate
(widest above the middle), with narrow beak. Style strongly contorted.

See Roth rock (1991). Moist to mesic forests; swamps; vernal pools; ditches;
meadows. [DE, PA, VA, WVj. Common, ES, WS; infrequent, PD; rare, RV(disjunct).

Laxiflorae

Carex albursina Sheldon
White Bear Sedge
(BR, F, GC, H, HS, KM, RB, SC, T)

Leaves very wide (15-40 mm), green, and overwintering. Leaf bases
green-white. Culms wing-angled. Terminal (staminate) spike overtopped
by or only slightly exceeding uppermost lateral (pistillate) spikes; uppermost
inflorescence bracts spathe-like and concealing uppermost spikes, when viewed
abaxially. See Manhart (1986). Rich forests, on calcareous or alluvial soils. [PA, VA,
WVj, Frequent, RV; infrequent, AP; rare, PD, WS.

Carex amphiboia Steudel

Eastern Narrow-leaf Sedge Griseae (Oligocarpae)

(RB; in part only: — GC, HS, SC; C. amphiboia Steudel var. rigida
(Bailey) Femald — BR, F, KM; in part — H, T)

Similar to C. grisea , C. planispicata , C. corrugata. Shoot bases green
to brown; perigynia spirally imbricate, little inflated, trigonous, 2.5-3. 1 times
as long as wide; summit of achene rounded. See Naczi (1993, 1999). Mesic forests and
meadows; floodplains. [DE, PA, VA, WV] Common, RV, PD; frequent, AP, WS;
infrequent (rare southward), ES.

Carex annectens (Bicknell) Bk: knell var. anneciens
Yellowfrait Sedge Multiflorae

(F, KM; C. annectens , in a narrow sense — SC; infraspecific taxa not
distinguished — BR, HS, RB, T; C, vulpinoidea Michaux var.
ambigua F. Boot!:, in part — GC; C. vulpinoidea , in part — H)

Similar to C. vulpinoidea , but with mature culms generally exceeding the
leaves; perigynium beak shorter than the body. Perigyeium brown-stramineous;
nerves evident. Moist soils, usually in open. [DE, PA, VA, WV]. Common, PD, WS,
ES; infrequent, AP, RV.

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

Yellowfinit Sedge Multiflorae

(F, KM; infraspecific taxa not distinguished — BR, HS, RB, T; C.
vulpinoidea Michaux var. ambigua F Boott, in part — GC; C.
vulpinoidea , in part — H; C brachyglossa Mackenzie — • SC)

Inflorescence more compact than in var. annectens ; perigynium golden yellow,
inconspicuously nerved or nerveless. Habitat as for var. annectens , possibly in less
acidic soils. [VA, WV j. Rare, PD h, WS-h.

Carex appalachica Webber and Ball
Appalachian Sedge Phaestoglochin (Bracteosae)

(GC, KM, RB; C. radiata (Wahlenberg) Dewey, misapplied — BR, F,

SC, T; C. rosea Willdenow, in part — H; C. convoluta Mackenzie, in
part — HS)

Similar to C. rosea and C radiata. Leaves narrow (0.9= 1.5 mm). Mature
perigynia ascending, 2=6 per spike. Achene base is 0. 1-0.5 mm from perigynium base.
Stigmas mostly tightly coiled. See Webber and Ball (1979, 1984). Dry to mesic forests.
[PA, VA, WV]. Rare, AP, RV, PD -h Perhaps overlooked.

Carex aquatilis Wahlenberg var. substricta /-a

Kukenthal Aquatic Sedge Phacocystis (Acutae) ' C , 'Y(\ yfl \

(F, GC; infraspecific taxa not distinguished — HS, RB; C. aquatilis f #|y 8 c

var. aquatilis , in part — KM) v t

Distinguished from similar members of section (C. stricta, C. emoryi ,

C. haydenii) in being phyllopodic and in having the lowest inflorescence
bract well overtopping inflorescence. Perigynium inflated, obovate, nerveless.
Reported for District of Columbia by Hitchcock and Standley (1919); first reported by
Terrell et ah (2000). Marshy swale. [PA, VA]. Rare (single station), WS.

Carex arctata W. Boott ex Hooker
Black Sedge Hymenochlaenae (Sylvaticae)

(F, GC, KM, RB)

Similar to C. debilis , but with basal leaves wider (6- 1 0 mm wide);
pistillate scales cuspidate or awned; perigynium prominently
trigonous and tight around the sessile achene, which is longer than the style.
See Fleming and Ludwig (1996). Dry, open oak forest. [PA, VA, WV]. Rare (single
station), AP.

Winter 2001

67

C. T, Frye & C. Lea

Carex arenaria Linnaeus
Sand Sedge Arenariae

(BR, F, GC, H, KM, T)

Coastal species with cordlike rhizomes. Spikes gynecandrous,
androgynous, or entirely staminate, the lower often subtended by
leafy bracts. Margin of perigynium forms a wide, thin wing from the
middle to the beak. See Jenny et ah (1982). Ballast; coastal sands in more
natural habitats in adjacent states. [DE, VA]. Locally adventive (European), WS-h.

Carex argyraniha Tuckerman
Hay Sedge Ovales

(BR, F, GC, H, KM, RB, SC; C.foenea Willdenow, misapplied)

Leaves pale green. Inflorescence interrupted. Perigynium silvery-
green, strongly nerved on both faces, finely granular-papillose.

Pistillate scales as long as, but narrower than, the perigynia. Sandy or rocky
forests and openings. [DE-h, PA, VA, WV]. Infrequent, AP; rare, RV-h, PD h.

Carex atlantica Bailey ssp. atlantica
Bog Sedge Stellulatae

(KM, RB; C atlantica , in a narrow sense — BR, F, H, T; C, atlantica
var. atlantica — GC; C. incomperta Bicknell — BR, F, HS, SC, T; C.
cephalantha Bicknell, misapplied, — HS)

Perigynium strongly nerved adaxially; beak less than 1 mm long. Similar to
ssp. capillacea , but with widest leaves more than 1 .6 mm wide; infractescence more
than 19 mm long. See Reznicek and Ball (1980). Swampy forests; seepages; fens;
peatlands. [DE, PA, VA, WV]. Common, AP; frequent, RV, PD, WS, ES.

Carex atlantica Bailey ssp. capillacea (Bailey)

Reznicek

Howe’s Sedge Stellulatae

(KM, RB; C. atlantica var. capillacea Bailey — GC; C. howei
Mackenzie — BR, F, H, HS, SC, T)

Similar to ssp. atlantica (intermediate plants occur), but with widest leaves
less than 1.6 mm wide, leaves typically becoming involute; infractescence less than
19 mm long. See Reznicek and Ball (1980). Swampy forests; seepages; peatlands. [DE,
PA, VA]. Frequent, WS, ES; rare, AP (disjunct), PD-h.

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The Maryland Naturalist Volume 44

Atlas and Annotated List of Carez (Cypepacea e) of Maryland and the District of Columbia

Carex austrina (Small) Mackenzie
Southern Sedge Phaestoglochin (Bracteosae)

(F, KM; C. inuhlenbergii Schkuhr.ex Willdenow var,
australis Olney ex Bailey — GC; C. muhlenbergii var. austrina Small)

Resembles C. muhlenbergii , but lowermost inflorescence bract longer
(the free portion 10-50 mm long vs. 5-20 mm) and dilated at the base;
pistillate scales longer and wider, covering most of the perigynia; perigynia
slightly larger (3.5 - 4.0 mm long, 2. 5-3.0 mm wide) and strongly ascending (not
spreading) at maturity. Meadow, [none]. Locally adventive (from south-central United
States) (single station), ES.

Carex baileyi Britton

Bailey's Sedge Vesicariae (Pseudocypereae)

(BR, F, GC, H, KM, RB, SC)

Similar to C. lurida , but leaves narrower (2-4 mm wide); lateral (pistil¬
late) spikes narrower (less than 13 mm wide in mature specimens);
perigynium abruptly contracted to a beak as long as the body. Wet soils, often
in open habitats, at high elevations. [PA, VA, WV], Frequent, AP.

Carex barrattii Schweinitz and Torrey
Barratf s Sedge Limosae

(BR, F, GC, H, KM, RB, T)

Strongly rhizomatous sedge; rarely flowering. Lateral (pistillate) spikes
drooping on slender peduncles, often staminate at tip. Perigynium
beakless and finely papillose. Pistillate and staminate scales dark purple.

Forested or open non-tidal wetlands; Delmarva Bays. [DE, PA-h, VA]. Infrequent, ES;
rare, WS.

Carex blanda Dewey
Charming Sedge Laxiflorae

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to other members of the section; perigynium obovate, asym¬
metrical, with abruptly bent beak. Distinguished from C. gracilescens
by green, brown, or white (not purple) basal sheaths and pale and subsessile
terminal (staminate) spike. See Manhart (1986). Forests, edges, and floodplains, often
on disturbed soils; somewhat weedy. [DE, PA, VA, WV]. Common, RV, PD, WS;
frequent, AP, ES (rare southward).

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Carex hrevior (Dewey) Mackenzie ex Lunell
Short-beaked Sedge Ovales

(BR, F, H, HS, KM, RB, SC, T; in part only — GC)

Perigynium body orbicular and wider (2.5-3 ,5 mm) than for most taxa of
section; the adaxial face nerveless or faintly nerved. Similar to C.
molesta; spikes less congested, less densely flowered, the terminal
often with a prolonged staminate section; mature perigynia appressed-
ascending. See Rothrock and Reznicek (2001). Dry fields and open forests. [DE-h, PA,
VA, WV]. Rare, WS, PD.

Carex bromoides Willdenow ssp. bromoides
Brome-like Sedge Deweyanae

(KM; infraspecific taxa not distinguished — BR, F, GC, H, HS, RB, SC,

T)

Distinctive sedge with narrow (1 .0-2.5 mm), stiff leaves and lance-
cylindrical spikes. Perigynium linear-lanceolate, 4 to 5 times as long as wide.

See Naczi (1990). Swampy forests; floodplains; seepages. [DE, PA, VA, WV]. Fre¬
quent, ES; infrequent, AP, RV, PD, WS.

Carex brunnescens (Persoon) Poiret var.
sphaemstachya (Tuckerman) Kiikenthal
Brownish Sedge Glareosae (Heleonastes)

(F, KM, SC; infraspecific taxa not distinguished — BR, GC, H, RB; C.
brunnescens (Persoon) Poiret ssp. sphaemstachya (Tuckerman)

Kalela)

Similar to C. canescens , but with leaves green and narrow (1.0-2. 5 mm); mature
perigynia brown and loosely spreading. Rocky, usually montane, forests and slopes.
[PA, VA, WV] . Rare, AP, PD (disjunct).

Carex bullata Schkuhr ex Willdenow
Button Sedge Vesicariae

(BR, F, GC, H, HS, KM, RB, T)

Similar to C. vesicaria , but with elongated rhizomes; lateral (pistillate)
spikes thicker (10=20 mm); perigynium beak scabrous on margins;
distributed primarily on the Coastal Plain. Delmarva Bays; seepage swamps;
other non- tidal wetlands. [DE, PA, VA]. Infrequent, WS, ES.

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Carex hushii Mackenzie

Bush’s Sedge Porocysiis (Virescenies)

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. complanata and C. hirsutella, but with perigynia
spreading, not compressed. Similar to C. caroliniana , but with leaf
sheaths hairy; pistillate scales long-acuminate, exceeding perigynia
and interrupting the outline of the spike. Dry fields, meadows, and glades,
usually over basic soils. [DE, PA, VA, WV]. Infrequent, PD; rare, WS.

Carex buxbaumii Wahlenberg
Buxbaum’s Sedge Racemosae (Atratae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Distinctive sedge, with perigynium nearly beakless, densely papillose
(conspicuous with a lOx lens). Pistillate scales aristate, brown to
purplish black, with paler midrib. Seepages; fens. [DE, PA, VA, WV]. Rare, AP,
PD WS.

Carex canescens Linnaeus var. canescens
Silvery Sedge Glareosae (Heleonastes)

(BR, F, RB, SC; C. canescens ssp. canescens — KM; infraspecific taxa
not distinguished — GC, H)

C. canescens is a distinctive sedge, with leaves very glaucous, 2-4 mm
wide; perigynia appressed ascending in ellipsoid or ovoid spikes. Var.
canescens has the inflorescence 2-1 cm long; spikes approximate or slightly remote.
Fens; seepages; other peatlands. [PA, VA, WV]. Infrequent, WS; rare, ES.

Carex canescens Linnaeus var. disjuncta Femald
Silvery Sedge Glareosae (Heleonastes)

(BR, F, HS, RB, SC, T; C. canescens ssp. disjuncta (Femald) Toivonen
— KM; infraspecific taxa not distinguished — GC, H)

Similar to var. canescens , but with inflorescence 6-15 cm long; all but
uppermost spikes remote. Fens; seepages; other peatlands. [DE, PA, VA,
WV]. Frequent, AP; infrequent, PD h, WS, ES.

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Carex careyana Torrey ex Dewey

Carey’s Sedge Careyanae (Laxiflorae)

(BR, F, GC, H, HS, KM, RB)

Distinctive within section. Leaves evergreen and wide (8-17 mm);
basal sheaths conspicuously wine-purple. Culms long and lax at
maturity. Perigynium large (5. 0-6. 5 mm long), yellow-green and
sharply trigonous. Staminate scales deep brown. See Manhart (1986). Rich
forests, on calcareous or alluvial soils. [PA, VA, WV]. Rare, AP, PD.

Carex caroliniana Schweinitz
Carolina Sedge Porocystis (Virescentes)

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. bushii , C. complanata , and C. hirsutella , but with leaves
and leaf sheaths glabrous. Perigynia glabrous, subterete and spread¬
ing. Pistillate scales ovate, shorter than the perigynia. Wet swales and
meadows; vernal pools; floodplains. [DE, PA, VA, WV]. Infrequent, WS; rare, PD, ES.

Carex cephaloidea (Dewey) Dewey

Thin-leaved Sedge Phaestoglochin (Bracteosae)

(BR, F, KM, RB; C. sparganioides Muhlenberg var. cephaloidea
(Dewey) Carey — GC)

Similar to C. aggregata , but with culms very rough on the wing-like
angles; inner bands of leaf sheaths truncate and friable; inflorescence bracts
usually lacking. Rich forests; floodplains. [PA]. Rare, RV-h.

Carex cephalophora Muhlenberg ex Willdenow

Oval-leaf Sedge Phaestoglochin (Bracteosae)

(BR, F, KM, HS, RB, SC, T; in part only — H; C. cephalophora var.
cephalophora — GC)

Inflorescence capitate. Similar to C. mesochorea , but with mature
culms equal in length to or barely exceeding leaves; perigynium smaller (1 .5
- 2.0 mm wide; 2. 0-3.0 mm long); pistillate scales (excluding awns) shorter than the
perigynia bodies. Dry forests, barrens, and fields. [DE, PA, VA, WV]. Common, AP,
RV, PD, WS; frequent (rare southward), ES.

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Carex cottmsii Nuttall
Collins’ Sedge Collins iae

(BR, F, GC, H, HS, KM, RE, T)

Distinctive sedge, with slender, weak, arching to reclining culms and
subulate perigynia that are widely spreading or reflexed at maturity,
Perigynium teeth abruptly reflexed. Seepages; Acidic seepages;
swamps; floodplain forests; often on Sphagnum . [DE, PA, VA], Infrequent,
WS,ES.

Carex communis Bailey van communis
Fibrous-root Sedge Acrocystis (Montanae)

(KM; infraspecific taxa not distinguished — BR, F, GC, H, HS, RB, SC,

V)

Similar to C. pensylvanica , but lacking long rhizomes and with wider
leaves (3-6 mm wide). Inflorescence bracts with hyaline, auriculate append¬
ages. See Cusick (1992), Rettig and Crins (1996). Rich, dry, often rocky, forests. [DE,
PA, VA, WV]. Frequent, RV, PD; infrequent, AP; rare, WS.

Carex comosa F. Boott

Bearded Sedge Vesicariae (Pseudocypereae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Distinctive sedge, with cylindrical lateral (pistillate) spikes with
reflexed perigynia having curving, divergent teeth, the inflorescence
thus appearing bristly. Marshes; seepages; sloughs; swamps; disturbed
wetlands. [DE, PA, VA, WV]. Frequent, WS, ES; infrequent, PD; rare, RV (disjunct).

Carex complanata Toney and Hooker
Flattened Sedge Porocystis (Virescentes)

(BR, F; KM, T; in part only — H, HS; C complanata var. complanata
— GC)

Leaf sheaths pubescent, perigynia glabrous, ascending, and flattened
on adaxial face, and pistillate scales ovate and shorter than the perigynia as
in C hirsutella , but leaves rigid, keeled, and glabrous to very sparsely pubescent.
Mesic to moist forests and clearings. [DE, VA]. Common, WS, ES; infrequent, RV, PD.

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Carex conjuncta F, Boot!

Soft Fox Sedge Vulpinae

(BR, F, GC, H, HS, KM, RB, SC)

Culms distinctly spongy and wing-angled; spikes compound.
Resembles C. stipata and C. laevivaginata , but perigynium lance
ovate, with shorter beak, half as long as the body or less. See Jones
and Reznicek (1995). Moist edges, open forests, floodplains, usually on
calcareous soils. [DE, PA, VA, WV]. Infrequent (locally common) RV, PD

Carex conoidea Scfrkuhr ex Willdenow
Field Sedge Griseae (Oligocarpae)

(BR, F, GC, H, KM, RB, SC, T)

Similar to C. ampin bo la, but with the axis of the inflorescence sca¬
brous, the terminal (staminate) spike elevated on long peduncle to 3 cm,
and perigynium smaller (2. 5-4.0 mm long). Wet meadows and moist forests,
usually over basic soils. [DE h, PA, VA, WV]. Rare, AP, PD, WS.

Carex corrugata Femald

Prune-fruit Sedge Griseae (Oligocarpae)

(F; C amphibola Steudel, in part — GC, KM)

Very similar to C. amphibola , but perigynium 1 .8-2.3 times as long as
wide. Similar to C grisea, but leaves generally narrower (widest less
than 5.5 mm wide) stipe generally longer (more than 0.4 mm). The most
distinctive feature is the truncate apex of the mature achene. See Naczi (1993). Rich
floodplain forests and meadows. [VA]. Infrequent, ES; rare (disjunct), RV, PD. Perhaps
overlooked.

Carex crinita Lamarck var. brevicrinis Femald
Short-fringed Sedge Phacocystis (Cryptocarpae)

(F, GC, KM, RB; infraspecific taxa not distinguished — BR, H, HS,

SC,T)

Similar to var. crinita (with apex of pistillate scale tmncate to retuse),
but with perigynia strongly inflated, obovate, the lower spikes appearing
more cylindrical and less tapering; achene symmetrical, without invaginations. See
Bruederle and Fairbrothers (1986). Wet meadows; swamps; ditches. [DE, PA, VA].
Common, WS, ES; infrequent, PD.

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Atlas and Annotated List of Carex (C yperaceae) of Maryland and the District of Columbia

Carex crinita Lamarck var. crinita
Fringed Sedge Phacocystis ( Cryptocarpae)

(F, GC, KM, RB; infraspecific taxa not distinguished —

SC, T)

Similar to C. gynandra , but with perigynia more inflated, obovate;
apex of pistillate scale truncate to retuse. Similar to var. brevicrinis,
but with perigynia ellipsoid, the lower spikes appearing long-tapering; achene
asymmetrical, invaginated on one or both sides. See Standley (1983). Wet meadows;
swamps; ditches. [DE, PA, VA, WV]. Common, AP, RV, PD; frequent, WS, ES.

Carex cristatella Britton
Crested Sedge Ovales

(BR, F, GC, H, HS, KM, RB, SC, T)

Resembles C. tribuloides (with inflorescence crowded and perigynium
thin and scale-like), but spikes globose, with recurving perigynia tips
hiding scales. Wet meadows; open swamps; floodplains of medium-sized to
large rivers. [DE-h, PA, VA, WV]. Rare, RV, PD, WS.

Carex davisii Schweinitz and Torrey
Davis’ Sedge Hymenochlaenae (Gracillimae)

(BR, F, GC, HS, KM, RB, SC, T)

Spikes elongate; terminal spike gynecandrous, as in C. gracillima and
C. prasina. Distinguished from other members of its section by having
leaf sheaths hairy, spikes relatively wide (4-6 mm), and pistillate scales with
spreading awns. Nutrient-rich floodplain forests and openings. [DE, PA, WV]. Rare,
RV, PD.

Carex debilis Michaux var. debilis
White-edged Sedge Hymenochlaenae (Sylvaticae)

(BR, F, GC, HS, KM, RB, C. debilis , in a narrow sense — SC; infraspe¬
cific taxa not distinguished — H, T )

C. debilis is distinctive, with elongate, drooping lateral (pistillate)
spikes with fusiform perigynia. Var. debilis has longer (6- 1 0 mm, as measured
at proximal end of the spike), glabrous perigynia that are widest below the middle,
with elongate, distinctly hyaline-tipped beaks. Swamps; moist forests. [DE, PA, VA,
WV]. Common, PD, WS, ES; frequent, AP, RV.

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C. T. Frye & C. Lea

Car ex debilis Michaux var. inter cursa Femald
Femald’s White-edged Sedge
Hymenochlaenae (Sylvaticae)

(F, KM; C debilis Michaux var .puhera Gray, in part — HS; infraspe¬
cific taxa not distinguished — H)

Distinguished from other varieties of C. debilis by having pubescent
perigynia and pistillate scales with the midribs evanescent below the scale
apex. Recent research (M. Waterway, pers. comm.) suggests that forms of C. debilis
with pubescent perigynia are not taxonomically distinct. See Femald (1942). Swamps;
moist forests. [VA], Rare, WS.

Carex debilis Michaux var. pubera Gray
Allegheny Sedge Hymenochlaenae (Sylvaticae)

(BR, F, GC, KM, RB; in part only — HS; C. allegheniensis Mackenzie;
infraspecific taxa not distinguished — H)

Distinguished from other varieties of C. debilis by having pubescent
perigynia and pistillate scales with the midribs generally excurrent. Recent
research (M. Waterway, pers. comm.) suggests that forms of C.
debilis with pubescent perigynia are not taxonomically distinct from var. debilis or
var. rudgei. Dry to mesic upland forests. [PA, VA]. Rare, AP, RV-h, PD.

Carex debilis Michaux var. rudgei Bailey
Rudge’s Sedge Hymenochlaenae (Sylvaticae)

(BR, F, GC, KM, RB; C.flexuosa Muhlenberg ex Willdenow — SC;
infraspecific taxa not distinguished — H)

Differs from other varieties of C debilis in having shorter (4. 5-7.0
mm, as measured at proximal end of spike), glabrous perigynia that are widest
near the middle with short beaks that are slightly or not hyaline-tipped. Northern
hardwoods, hemlock forests. [PA, VA, W V]. Frequent, AP.

Carex decomposita Muhlenberg
Cypress-knee Sedge Heleoglochin (Paniculatae)

(BR, F, GC, H, HS, KM)

Inflorescence compound, with many spikes per branch. Perigynium
obovoid, with short (0.5 mm) beak, olive to black, shiny when mature.
Differs from C. diandra in having leaf sheaths concave at the mouth (not
prolonged) and in range and habitat. Woodland pools. [DE, VA]. Rare, PD h

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Atlas and Annotated List of Carex (Cyperaceaje) of Maryland and the District of Columbia

Carex diandra Schrank

Lesser Panicled Sedge Heleoglochin (Paniculatae)

(F,GC,KM,RB)

Inflorescence compound. Perigynium lance-ovate, with prominent,
serrulate-margined beak to 1 mm long. Differs from C. decomposita in
having leaf sheaths distinctively prolonged beyond the base of the
leaf blade and in range and habitat. Minerotrophic fen at high elevation. [PA].
Rare (single station), AP. Disjunct from the glaciated Northeast.

Carex digitalis Willdenow var. digitalis

Northern Slender Woodland Sedge Careyanae (Laxiflorae)

(BR, F; infraspecific taxa not distinguished — GC, H, HS, KM,

RB, SC, T).

Leaves green, narrower (2. 7-4. 5 mm) than in most taxa in section.

Perigynium trigonous, 2.5-33 mm long. Pistillate scales acuminate to awned.

Terminal (staminate) spike with short (usually < 7 cm) peduncle, usually surpassed by
uppermost lateral spike bract. See Manhart ( 1 986), Naczi et ah (200 1 ). Dry to mesic
forests. [DE, PA, VA, WV]. Common, RV, PD, WS; frequent, AP; infrequent (rare
southward), ES.

Carex digitalis Willdenow var. macropoda Femald
Southern Slender Woodland Sedge
Careyanae (Laxiflorae)

(BR, F; infraspecific taxa not distinguished — GC, H, HS, KM, T).

Similar to var. digitalis , but leaves narrower (2. 0-2. 9 mm). Terminal
spike long-peduncled (8-16 cm), usually exceeding the uppermost lateral
bract). See Femald (1938), Naczi et al. (2001). Nutrient-rich, mesic to dry forests.
[VA]. Rare, PD, ES. Some authors question its taxonomic distinctiveness (see
Manhart, 1986).

Spirostachyae (Extensae)

Carex distans Linnaeus
Distant-flowered Sedge
(KM)

Distinctive cespitose sedge, with gray-green leaves, the terminal
spike staminate, and two to three widely-spaced lateral (pistillate)
spikes on peduncles 1 -4 cm long. Perigynium flask-shaped, with prominent
ribs and rough-edged beak. See Tutin et al. ( 1 980), Jenny et al. ( 1 982). Damp meadow;
in Europe, also brackish marshes and coasts, [none]. Locally naturalized or adventive
(European) (single collection), WS-h.

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Carex divisa Hudson

Divided Sedge Divisae

(BR, F, GC, H, KM)

Colonial sedge with coarse rhizomes with blackish scales. Leaves
narrow (1-3 mm). Spikes elongate, androgynous or entirely pistillate.
Pistillate scales brownish, largely concealing the perigynia. See Jermy
et al. (1982). Salt marshes. [VA]. Locally naturalized or adventive (European),
WS.

Phaestoglochin (Bracteosae)

Carex divulsa Stokes
Gray Sedge
(BR, F, GC, KM, RB)

Similar to C. spicata and C. muricata , with spicate inflorescence and
pistillate scales often strongly tinged with brown or reddish purple.

Basal sheaths brown to blackish (not purple-tinged, as in C. spicata ).

Perigynia appr essed -ascending (not spreading, as in C. muricata ), tapered at ends.
See Jermy et al. (1982). Disturbed sites; lawns. [PA]. Locally naturalized or adventive
(European) (single collection), WS h

Carex eburnea F. Boott
Ebony Sedge Albae

(F, GC, H, KM, RB, SC)

Distinctive sedge, cespitose from slender rhizomes, with wiry,
capillary culms and involute, filiform leaves that are less than 2 mm
wide. Terminal spike staminate, sessile to subsessile, overtopped by two
uppermost lateral (pistillate) spikes. Inflorescence few-fruited; mature perigynium
olive to black. Rocky, calcareous slope. [PA, VA, WV], Rare (single station), RV.

Carex echinata Murray ssp. echinata
Prickly Sedge Stellulatae

(KM; C. echinata var. echinata — • GC; infraspecific taxa not distin¬
guished — RB; C. angustior Mackenzie — BR, F, HS, SC, T; C.
cephalantha (Bailey) Bicknell — BR, F; C. laricina Mackenzie —

SC; C. muricata Linnaeus, misapplied — H)

Similar to C atlantica and C interior , but with longer (0.95-2.00 mm) perigynium
beak. See Reznicek and Ball (1980). Fens; seepages. [PA, VA, WV]. Infrequent, AP;
rare (disjunct), PD, WS-h.

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Carex emoryi Dewey

Emory’s Sedge Phacocystis (Acutae)

(BR, F, GC, HS, KM, RB, SC, T)

Resembles C. stricter, more robust; lower leaf sheaths not fibrillose;
summit of leaf sheaths convex; lateral (pistillate) spikes generally
longer and more overlapping; perigynium obovate, with granular
papillae restricted to apex. See Standley (1989). Rocky banks of large rivers;
rarely, river swamps. [DE, PA, VA, WV]. Infrequent, RV, PD (locally common). Mostly
along Potomac River.

Stellulatae

Carex exilis Dewey
Coast Sedge
(BR, F, GC, KM, T)

Involute leaves and solitary terminal spikes (that are either staminate,
pistillate, or gynecandrous, with the staminate part prolonged and
clavate) distinguish this species from other members of its section. See

Reznicek and Ball (1980). Atlantic white cedar swamp. [DE]. Rare (single station), WS.

Carex extensa Goodenough

Long-bracted Sedge Spirostachyae (Extensae)

(BR, F, GC, H, KM)

Leaves involute. Inflorescence bracts sheathless. Lateral (pistillate)
spikes thick-cylindrical; the upper subapproximate and the lower often
remote. See Jermy et al. (1982). Salt marshes; barrier island wash flats. [VA].
Locally naturalized (European) (coastal), WS, ES.

Carex festucacea Schkuhr ex Willdenow
Fescue Sedge Ovales

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to several other members of section; spikes usually well
separated, often with prolonged, tapering staminate bases. Body of
perigynium orbicular with abrupt transition to beak, fairly prominently nerved
on abaxial face; nerves 3 or fewer on adaxial face. See Rothrock (1991). Moist fields,
meadows, and open forests. [DE, PA, WV, VA]. Frequent, PD, WS, ES; rare, RV.

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Carex folliculata Linnaeus

Northern Long Sedge Rostrales (Folliculatae)

(HS, KM, RB, SC; in part only — H, T; C. folliculata
var. folliculata — BR, F, GC)

Distinctive sedge with inflated, lanceolate perigynia. Similar to C
lonchocarpa , but leaves wider (6-16 mm); lateral spikes usually not
staminate at the tips; pistillate scales 0.5- 1.2 times as long as perigynia. See
Wujek and Men apace (1986). Seepages; fens; other acidic wetlands. [DE, PA, VA,
WV], Common, AP; frequent, WS; infrequent, RY, PD, ES.

Squarrosae

Carex frankii Kunth

Frank’s Sedge

(BR, F, GC, H, HS, KM, RB, SC, T)

Distinguished from other members of section by having the terminal
spike staminate and the awns of the pistillate scales exceeding the
perigynia. Lateral (pistillate) spikes cylindrical; perigynia closely packed,
abruptly beaked, the bodies obovate. Open, often disturbed, wetlands; ditches;

floodplains. [DE, PA, WV, VA]. Common, RV, PD, WS; infrequent (perhaps adventive),

ES.

Carex gigantea Rudge

Giant Sedge Lupulinae

(BR, F, GC, H, KM, T)

Similar to C lupulina and C. lupuliformis . Often with multiple (2-5)
staminate spikes. Perigynia abruptly narrowed to long, horizontally
divergent beaks, 2-4 times as long as the bodies. Achene rhombic, broader
than long; summit truncate; faces concave; angles thickened. See Reznicek and Ball
(1974). Seasonal ponds; swamp forests; floodplains. [DE, VA]. Infrequent, ES; rare,
WS.

Carex glaucescens Elliott
Southern Waxy Sedge Pendulinae

(BR, F, GC, H, KM, T)

Similar to C.joorii , but with perigynium inconspicuously nerved,
glaucous, little inflated, 2. 8-3. 5 mm long; pistillate scales retuse below
the awn. Moist areas in pine woodlands; wet, acid, and/or peaty habitats
[VA]. Rare, ES.

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Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

Carex glaucodea Tuckerman ex Olney

Blue Sedge

Griseae (Oligocarpae)

(BR, HS, RB, SC, T; C. flaccosperma Dewey var. • • .. ;

glaucodea (Tuckerman) Kiikenthal — F, KM; C. flaccosperma X#vi %

Dewey, in part — GC, H) • # ?

Distinctive within section; leaves glaucous; perigynium plump and <

beakless; pistillate scales awnless or short-awned. See Naczi (1997), Rich, often
basic, forests and meadows; serpentine. [DE, PA, VA, WV]. Frequent, PD, WS;
infrequent, RV, ES.

Carex gracilescens Steudel

Slender Loose-flowered Sedge Laxiflorae

(BR, F, GC, H, KM, RB, SC, T; C. laxiflora Lamarck, mi:

HS)

Perigynium beak abruptly bent, as in C. blanda , but of
arching. Further distinguished from C. blanda by purple basal sheaths (often
obscure to absent in weathered specimens). Terminal (staminate) spike well elevated
above upper lateral (pistillate) spikes. See Manhart (1986). Rich forests; floodplains of
medium-sized streams. [DE, PA, VA, WV]. Frequent, RV, PD; rare, AP, WS.

Carex gracillima Schweinitz

Graceful Sedge Hymenochlaenae (Gracillimae)

(BR, F, GC, H, HS, KM, RB, SC, T)

A frequently encountered member of a group of sedges with • ’ J

gynecandrous terminal spikes and long, narrow pistillate lateral spikes.

Basal sheaths purple; leaf sheaths glabrous on ventral sides. Perigynium
obtusely angled, beakless. Mesic forests; floodplains. [DE, PA, VA, WV]. Common,
AP; frequent, RV, PD; infrequent, WS; rare, ES.

Carex granularis Muhl. ex Willd. var. granularis
Limestone Meadow Sedge Granulares

(F, RB; infraspecific taxa not distinguished — BR, GC, H, HS, KM, SC,

T) /*}

Distinctive sedge, with leaves somewhat glaucous. Terminal spike

staminate, sessile to short-peduncled. Lateral spikes pistillate and cylindrical,
with closely packed subglobose perigynia with short, but distinct, beaks. Rich, often
open, floodplains; minerotrophic fens. [DE, PA, VA, WV]. Infrequent, AP, RV, PD; rare,
WS,ES.

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C. T. Frye & C. Lea

Carex granularis Muhlenberg ex Willdenow
var. haleana (Olney) Porter
Hale’s Sedge Granular es

(F, RB; infraspecific taxa not distinguished — BR, GC, H, HS, KM, SC,

T; C. haleana Olney)

Similar to van granularis , but perigynium much less inflated, oblong-
ellipsoid; lateral (pistillate) spikes appearing more densely flowered. May not
be taxonomically distinct, as specimens intermediate with var. granularis occur. Rich,
often open, floodplains, usually with var. granularis. [PA]. Rare, PD.

Carex gravida Bailey var. iunelliana (Mackenzie)

Hermann

Heavy Sedge Phaestoglochin (Bracteosae)

(F, GC, KM)

Similar to C. aggregata , but perigynia brownish-yellow at maturity and
pistillate scales with a slender tip, surpassing the base of the perigynium
beak. Dry meadow. [VA]. Locally adventive (from midwestem United States) (single
station), WS.

Carex grayi Carey

Asa Gray’s Sedge Lupulinae

(GC, H, KM, RB; C. grayii , an orthographic error — BR, F, SC)

Similar to C. intumescens , but lateral (pistillate) spikes globose, with
perigynia radiating in all directions, with their bases rhombic, rather
than rounded. Most of our plants have hispidulous perigynia (var. hispidula
Gray, a dubious taxon). See Reznicek and Ball (1974). Floodplain forests. [PA, VA,
WV]. Infrequent (locally common), RV, PD, WS; rare, ES.

Carex grisea Wahlenberg

Inflated Narrow-leaf Sedge Griseae ( Oligocarpae)

(RB, HS, T; C. amphibola Steudel var. turgida (Bailey) Femald —

BR, F, KM, SC; C. amphibola Steudel, in part — GC, H)

Similar to C. amphibola and C. corrugata. Perigynia spirally
imbricate, conspicuously inflated, rounded in cross section, 1.8-2. 3 times as
long as wide; summit of achene rounded; achene stipe usually less than 0.4 mm long.
See Naczi (1993). Floodplain forests and meadows. [DE, PA, VA, WV]. Frequent, RV,
PD; rare, WS.

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Carex gynandra Schweinitz

Nodding Sedge Phacocystis (Cryptocarpae)

(BR, GC, HS, KM, RB, SC, T; C. crinita Lamarck van
gynandra (Schweinitz) Schweinitz and Tonrey ■ — F; C. crinita , in part
— H)

Similar to vars. of C, crinita , but perigynium barely inflated, ellipsoid;
apex of pistillate scale acute. Achenes invaginated on one or both sides. See
Standley (1983), Braederle and Fairbrothers (1986). Wet meadows; swamps; ditches.
[DE, PA, VA, WV]. Common, AP; infrequent, RV-h, PD; rare, WS.

Carex haydenii Dewey

Cloud Sedge Phacocystis (Acutae)

(F, GC, KM, RB; C stricta Lamarck var. decora Bailey — T)

Similar to C. stricta, but with lower leaf sheaths not fibrillose;
perigynium inflated, red-green, without nerves or papillae; pistillate
scales acuminate, longer than the perigynia. See Standley (1989).

Minerotrophic fen, with C stricta, at high elevation. [PA] . Rare (single station), AP.

Carex hirsutella Mackenzie
Hirsute Sedge Porocystis (Virescentes)

(BR, F, KM, RB, SC, T; C. complanata Torrey and Hooker var. hirsuta
(Bailey) Gleason • — GC; C. complanata Torrey, in part — H, HS)

Similar to C. complanata, but leaves flat, soft, and moderately to
densely pubescent. Dry forests, fields, and meadows. [DE, VA, WV]. Com¬
mon, RV, PD; frequent, WS; rare, AP, ES.

Carex hirta Linnaeus

Hammer Sedge Carex (Hirtae)

(BR, F, GC, KM, RB)

Vigorously rhizomatous. Leaf sheaths and pistillate scales hairy.

Although reported for Maryland by Shreve (1910) (who most likely
misapplied the name to C. hirtifolia ) and Brown and Brown (1984), the first
Maryland specimen seen was collected in 1998. See Jenny et al. (1982). Roadside;
disturbed sites. [PA]. Locally naturalized or adventive (European), AP, PD (planted at
residence).

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Carex hirtifolia Mackenzie

Pubescent Sedge Haller anae (Triquetrae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Distinctive sedge. Leaves soft and lax, hairy, with two prominent mid¬
lateral veins. Perigynium pubescent, trigonous, with slender beak.

Terminal (staminate) spike pale, sessile or short: peduncled, C x
sullivantii Boott, a purported hybrid between this species and C gracillima ,
occurs in Garrett County. Rich, dry to mesic forests, often in stream valleys. [DE, PA,
VA, WV]. Frequent, PD; rare, AP, RV-h, WS.

Carex hitchcockiana Dewey
Hitchcock’s Sedge Griseae (Oligocarpae)

(BR, F, GC, H, HS, KM, RB, SC)

Similar to C. oligocarpa; leaves wider (3-7 mm); basal sheaths
brownish; upper leaf sheaths hispidulous; perigynium larger (4-5 mm
long). Upland forests, on calcareous or mafic soils; less commonly in alluvial
forests. [PA, VA, WV]. Rare, RV, PD, WS.

Carex hormathodes Femald
Marsh Straw Sedge Ovales

(BR, F, KM, T; C. straminea Willdenow ex Schkuhr var. invisa W.

Boott — GC; C. straminea , in part — H)

Pistillate scales awned. Similar to C. straminea, but with inflores¬
cence nodding; staminate bases of lateral spikes up to 3 mm long; perigynia
more ascending, with gradually tapering beaks that are 1/3 to 3/5 as long as the lance-
ovate bodies. See Rothrock et ah (1997). Brackish to salt marshes, often with Spartina
patens. [DE, VA]. Infrequent (coastal), WS, ES.

Carex hyalinolepis Steudel
Shoreline Sedge Paludosae

(BR, F, GC, H, KM, RB; C. lacustris Willdenow var. laxiflora Dewey)

Similar to C lacustris , but phyllopodic; basal sheaths usually white,
not fibrillose; foliage glaucous; ligule shorter to slightly longer than
wide; pistillate scales with awns equal in length to the perigynia; style
straight. Fresh to slightly brackish tidal marshes (often in large stands); swamp
forests. [PA-h, VA]. Infrequent (locally common), ES (rarer northward); rare, WS.

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Care x hystericina Muhlenberg ex Willdenow
Porcupine Sedge Vesicariae (Pseudocypereae)

(GC, H, HS, KM, RB; C. hystricina , an orthographic
variant — BR, F, SC)

Similar to C. lurida, but with lateral (pistillate) spikes drooping on
short to long, fine peduncles. Perigynium less inflated than in C.
lurida , and has more (15=20) and more prominent nerves. Wet meadows on
basic soils; serpentine. [PA, VA, WV]. Rare, PD.

Car ex interior Bailey
Inland Sedge Stellulatae

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to, and often contused with, C. atlantica ssp. atlantica.

Perigynium with prominently setose-serrulate beak, nerveless adaxial
surface, with nearly straight sides that form subtle “shoulders” below the
beak. See Reznicek and Ball (1980). Seepages over serpentine. [PA, VA, WV]. Rare,
PD. Much less frequent in state than has been previously suggested.

Carex intumescens Rudge
Greater Bladder Sedge Lupulinae

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. grayi , but perigynia mostly ascending to spreading.

May resemble C. louisianica, but has inner sides of the perigynium
teeth hispid, achenes ellipsoid to obovoid. Brown and Brown (1984) report
Mdx.fernaldii Bailey, a dubious taxon, with perigynia barely inflated. See Reznicek and
Ball (1974). Seasonal ponds, floodplains, swamps, seepages. [DE, PA, VA, WV].
Common, PD, WS, ES; frequent, AP, RV.

Carex jamesii Schweinitz

James’ Sedge Phyllostachyae

(BR, F, GC, H, HS, KM, RB, SC)

Similar to C. willdenowii , with staminate scales tightly clasping, but
with perigynium body globose, abruptly contracted to a slender beak.

See Catling et al. (1993). Forests on calcareous soils; floodplains. [PA, VA,
WV]. Frequent, RV; infrequent (locally common, mostly along Potomac and
Susquehanna Rivers), PD, WS; rare, ES.

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Carex joorii Bailey

Joor’s Sedge Pendulinae

(BR, F, GC, H, KM, T)

Similar to C. glaucescens , but with perigynium strongly nerved, green
(brown when mature), well inflated, 4-5 mm long; pistillate scales
gradually tapering or rounded to the awn. Cypress swamps; swampy
meadows; edges of Delmarva Bays, [DE, VA]. Infrequent, ES; rare, WS.

Macrocephalae

Carex kobomugi Ohwi
Asiatic Sand Sedge
(BR, F, GC, H, KM)

Coarse, strongly rhizomatous and colonial, dune-forming sedge.

Dioecious; leaves serrulate. Flowering plants have not yet been found
in Maryland. An invasive species. See Small (1954). Though reported by
Brown and Brown (1984), first known Maryland collection was in 1999. Barrier island
beaches and dunes. [DE, VA]. Locally naturalized (from eastern Asia) in coastal areas,
ES.

Carex species 1

A Sedge Laxiflorae

(C. laxiflora Lamarck var. serrulata Hermann, misapplied)

An undescribed species that has been assigned to C. laxiflora var.
serrulata (a different taxon whose identity is presently obscure).

Uppermost inflorescence bract spathe-like (3-7 cm long, 2-7 mm wide and
concealing uppermost lateral (pistillate) spikes when viewed from abaxial surface).
Upper lateral spikes congested as in C. blanda; perigynium shape similar to that of C.
laxiflora. See C. laxiflora var. serrulata in Reznicek and Gonzalez-Elizondo (1999).
Floodplain or mesic forests. [VA]. Rare, WS.

Carex lacustris Willdenow
Lake-bank Sedge Paludosae

(BR, F, GC, H, HS, KM, RB, SC; C. riparia W. Curtis var. lacustris
(Willdenow) Kukenthal — T)

Similar to C. hyalinolepis , but aphyllopodic; basal sheaths pink to
deep red, fibrillose; foliage green; ligule much longer than wide; pistillate
scales, including awns, generally shorter than the perigynia; style bent. Perigynia 5.5-
7.0 mm long. Swamp forests; marshes; fens. [DE, PA, VA, WV]. Infrequent, WS, ES;
rare, AP.

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Carex laevivaginata (Kukenthal) Mackenzie
Smooth-sheathed Sedge Vulpinae

(BR, F, GC, H, HS, KM, RB, SC, T) .

Similar to C. stipata (with spongy, sharp-angled culms, compound
inflorescence, and beak of perigynium about as long as the body), but
with inner band of leaf sheaths smooth, not cross-puckered, with
cartilaginous thickened summit. Swamps; seepages; moist forests; mucky
soils. [DE, PA, VA, WV]. Common RV, PD, WS, ES; frequent, AP.

Carex lasiocarpa Ehrhart var. americana Femald
Many fruited Sedge Paludosae (Hirtae)

(BE. F, GC, KM, RB; C. lasiocarpa , in part only — H)

Perigynia pubescent. Similar to C. pellita, but with leaves narrower
(0.5-2. 0 mm) and involute. See McClintock and Waterway (1994) for
discussion of nomenclature. Minerotrophic fen at high elevation. [PA, VA,
WV-h], Rare (single station), AP.

Carex laxiculmis Schweinitz var. copulata (Bailey)

Femald

Coupled Sedge Careyanae (Laxiflorae)

(KM, RB; C. x copulata (Bailey) Mackenzie — F; infraspecific taxa not
distinguished — BR, GC, H, SC, T; C. copulata (Bailey) Mackenzie)

Similar to var. laxiculmis, but leaves green and narrower (widest blade 53-8.3
mm wide) and terminal (staminate) spike shorter (longest per plant 0. 6-2.0 mm long.
See Manhart (1986), Naczi et ah (2001). Nutrient-rich, sandy alluvial forests. [PA, VA].
Rare (single station), PD.

Carex laxiculmis Schweinitz var. laxiculmis
Lax-culmed Sedge Careyanae (Laxiflorae)

(KM, RB; infraspecific taxa not distinguished — BR, GC, H, HS, SC, T;

C. laxiculmis , in a narrow sense — F)

Distinctive within section. Leaves glaucous, 6.4-1 1 .8 mm wide; lateral
spikes on slender peduncles (the lowest drooping); proximal 1-3 scales of
lateral spikes staminate or empty; perigynium sharply trigonous. Longest terminal
(staminate) spike 1 .2-2.5 cm long. See Manhart ( 1 986), Naczi et al (200 1 ). Rich, mesic
forests. [DE, PA, VA, WV]. Common, RV, PD; frequent, WS; infrequent, AP, ES (rare
southward).

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Carex laxiflora Lamarck
Loose-flowered Sedge Laxiflorae

(BR, GC, H, RB, SC, T; C. laxiflora var. laxiflora , in a
narrow sense — F, KM; C. anceps Muhlenberg — HS)

Leaf width variable (3.5- 1 0.0 mm). Similar to C. striatula , but lateral
(pistillate) spikes narrower (2. 0-3.0 mm); perigynium smaller (3. 0-4.5 mm
long), with obovoid body and straight to slightly oblique, slender beak. See
Manhart (1986). Rich forests, often on bare soil of banks or slopes. [DE, PA, VA, WV].
Frequent, RV, PD, WS; infrequent, AP, ES (rare southward).

Carex leavenworthii Dewey

Leavenworth’s Sedge Phaestoglochin (Bracteosae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. cephalophora , but leaves narrower ( 1 -3 mm), perigynium
body cordate-deltoid, widest at or near the base. Dry to moist mead¬
ows; fields; pastures; somewhat weedy. [DE, PA, VA, WV]. Evidently locally
naturalized (from southern United States) (Mackenzie 1931), RV, PD, WS, ES.

Carex leptalea Wahlenberg ssp. harperi (Femald)

W. Stone

Harper’s Bristle-stalked Sedge Leptocephalae (Polytrichoideae)

(KM; C. leptalea Wahlenberg var. harperi (Femald) W. Stone,
misapplied — BR, F, T; infraspecific taxa not distinguished — GC, H,

HS; C. leptalea Wahlenberg var. harperi (Femald) Weatherby and
Griscom; C. harperi Femald)

Similar to ssp. leptalea; perigynia longer (3. 5-5.0 mm), more narrowly ellipsoid, and
more strongly overlapping; the staminate portion of spike more obscured by the
perigynia. Southern expression of the species. Seepages, often on Sphagnum ,
floodplains. [DE, VA]. Infrequent, WS, ES.

Carex leptalea Wahlenberg ssp. leptalea
Bristle-stalked Sedge Leptocephalae (Polytrichoideae)

(KM; C. leptalea var. leptalea — BR, F, T; infraspecific taxa not
distinguished - GC, H, HS, RB, SC)

Small, distinctive sedge, with narrow (0.5-1 .3 mm) leaves and spikes
solitary on culms. Similar to ssp. harperi; perigynia shorter (2. 5-3. 5 mm) and
more broadly ellipsoid; staminate portion of spike more prominent. Northern expres¬
sion of the species. Seepages; edges of fens; usually on Sphagnum. [DE, PA, VA,
WV]. Frequent, AP; infrequent, RV, PD.

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Car ex leptomervia (Femald) Femald
Finely-nerved Sedge Laxiflorae

(BR, F, GC, H, KM, RB, SC)

Appears much like C. laxiflora and other members of section.

Perigynium with convex sides and prominent midrib, but otherwise
faintly nerved to nerveless. Wet to mesic soil in northern hardwood
forests. [PA, VA, WV]. Frequent, AP; rare, RV (disjunct). Perhaps overlooked.

Carex lonchocarpa Willdenow ex Sprengel
Southern Long Sedge Rostrales (Folliculatae)

(KM; C. folliculata Linnaeus var. australis (Bailey) Femald — BR, F,

GC, T; C. folliculata Linnaeus, in part — H)

Similar to C. folliculata, but with narrower (3.5- 1 0.0 mm) leaves; lateral
spikes normally staminate at tips; pistillate scales 0.3-0. 5 times as long as
perigynia. See Wujek and Menapace (1986). Swamps; floodplain forests; seepages.
[DE, VA]. Frequent (less so northward), ES.

Carex longii Mackenzie
Long’s Sedge Ovales

(BR, F, GC, KM, RB, T; C alholutescens Schweinitz, misapplied
HS; in part — H)

Similar to C alholutescens (with perigynium body widest above
middle), but with perigynium beak stouter, more broadly deltoid; style
straight. See Rothrock (1991). May hybridize with C alata. Open, dry to (usually)
moist, sandy soils; ditches. [DE, PA, VA, WV]. Common, ES; frequent, WS; rare,
PD-h.

Carex iouisianica Bailey
Louisiana Sedge Lupulinae

(BR, F, GC, H, FIS, KM)

Similar to C lupulina , but with narrower (3-6 mm) leaves; lateral
(pistillate) spikes short-cylindrical, the uppermost greatly exceeded by
the iong-peduncled (3-10 cm) terminal (staminate) spike. Similar to C
intumescens , but inner sides of perigynium teeth smooth and achenes rhombic. See
Reznicek and Ball (1974). Floodplain forests; swamps. [VA]. Infrequent (locally
common), WS,ES.

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Carex lucorum Willdenow ex Link var. lucorum
Blue Ridge Sedge Acrocystis (Montanae)

(KM; infraspecific taxa not distinguished — GC, RB,

SC; C. pensylvanica Lamarck var. distans Peck — F)

Similar to C. pensylvanica , but with culms scabrous; perigynium beak
longer (1 .0-1 .5 mm), two-thirds as long as to nearly as long as the
body. See Crins and Ball (1983), Rettig (1989), Cusick (1992). Dry woodland
openings; barrens. [VA, PA, WV]. Rare, RV-h, PD-h. Perhaps overlooked.

Carex lupuliformis Sartwell
False Hop Sedge Lupulinae

(BR, F, GC, KM, RB, T; included in C. lupulina Willdenow — H)

Perigynia ascending to slightly spreading (intermediate between C.
lupulina and C. gigantea). Style strongly contorted in the middle.

Achenes rhombic, as long as broad, faces strongly concave, the angles with
prominent, nipple-like knobs. See Reznicek and Ball (1974). Seasonal ponds; swamps.
[DE, PA, VA, WV]. Rare, PD, WS, ES.

Carex lupulina Muhlenberg ex Willdenow
Hop Sedge Lupulinae

(BR, F, GC, HS, KM, RB, SC, T; in part only — H)

Lateral (pistillate) spikes cylindrical, with strongly ascending
perigynia. Style strongly contorted near the base. Achenes rhombic,
longer than broad, faces flat to shallowly concave, angles thickened.

Terminal (staminate) spike short-peduncled (0. 5-6.0 cm), shorter than to slightly
exceeding upper lateral spikes. See Reznicek and Ball (1974). Swamps; ditches; pond
edges. [DE, PA, VA, WV]. Common, RV, PD, WS, ES; infrequent, AP.

Carex lurida Wahlenberg

Yellow-green Sedge Vesicariae (Pseudocypereae)

(BR, F, GC, HS, KM, RB, SC, T)

Pistillate scales with slender, serrulate awns equal in length to or
longer than scale body. Similar to C. baileyi, but leaves wider (4-7
mm); lateral (pistillate) spikes thicker (> 14 mm wide in mature specimens);
perigynium less abruptly contracted to beak. Various wet, usually open, and, often,
disturbed habitats. [DE, PA, VA, WV]. Common throughout.

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Carex meadii Dewey

Mead’s Sedge Paniceae

(BR, F, GC, H, KM, RB, SC, T)

Rhizomatous sedge, similar to C. tetanica , but leaves gray-green (not
green); lateral (pistillate) spikes thicker (5-7 mm), with 6 rows of
perigynia. Woodland openings over basic soils (diabase, serpentine).
[DE, PA, VA, WV-h]. Rare, PD.

Carex mesochorea Mackenzie
Midland Sedge Phaestoglochin (Bracteosae)

(BR, F, HS, KM, RB, SC, T; C. cephalophora Muhlenberg var.
mesochorea (Mackenzie) Gleason — GC; C. cephalophora , in
part — H)

Similar to C. cephalophora , but with mature culms greatly exceeding leaves;
perigynium larger (2.0 - 2.5 mm wide; 3. 0-3. 5 mm long); pistillate scales (excluding
awns) as long as to longer than the perigynia bodies. Dry forests and fields. [DE, PA,
VA, WV]. Frequent, WS; infrequent, PD, ES.

Carex michauxiana Boeckeler
Andre Michaux’s Sedge Rostrales (Folliculatae)

(BR, F, GC, KM)

Similar to C. folliculata and C. lonchocarpa , but leaves narrower (1.5-
4 mm wide); terminal (staminate) spike shorter (0.6-1 .5 cm long);
inflorescence bract sheaths concave, rather than prolonged, at the mouth.

See Wujek and Menapace (1986). Seepage with Sphagnum . [none]. Rare (single
station), AP. Disjunct from northeastern United States.

Carex mitchelliana M. A. Curtis
Mitchell’s Sedge Phacocystis

( Cryptocarpae)

(F, GC, KM, RB, T; C. crinita Lamarck, in part — H; C. crinita var.
mitchelliana (M.A. Curtis) Gleason)

Similar to, but smaller overall than, C. gynandra and varieties of C. crinita.
Perigynium barely inflated, ellipsoid, densely granular-papillose throughout, with
fairly distinct nerves. See Bruederle and Fairbrothers (1986), Bruederle et al. (1989).
Floodplains; mature swamp forests. [DE, PA, VA]. Rare, ES.

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Care x molesta Mackenzie ex Bright
Troublesome Sedge Ovales

(BR, F, KM, RB, SC; C. brevior (Dewey) Mackenzie, in part —

GQ

Similar to C. brevior: ; the spikes more crowded and more densely
flowered, the terminal often lacking a prolonged staminate base.

Perigynia spreading at maturity, the bodies ovate and nerved on adazial side,

2. 0-3.0 mm wide. See Rothrock and Reznicek (2001). Dry to moist fields and other open

areas. [DE-h, PA, VA, WV]. Rare, PD-h, WS.

Carex muhlenbergii Schkuhr ex Willdenow var.
enervis W. Boott

Muhlenberg’s Sedge Phaestoglochin (Bracteosae)

(F, T; in part — GC; infraspecific taxa not distinguished — BR, H, HS,

RB, SC; C. muehlenbergii var. enervis , an orthographic variant -
KM; C. plana Mackenzie)

Similar to var. muhlenbergii , but with perigynium nerveless or with short basal nerves
adaxially. Dry forests and edges, usually on more basic soils than var. muhlenbergii.
[DE, VA]. Frequent, PD; infrequent, RV-h; rare, WS, ES.

Carex muhlenbergii Schkuhr ex Willdenow var.
muhlenbergii

Muhlenberg’s Sedge Phaestoglochin (Bracteosae)

(F, T; in part only — GC; infraspecific taxa not distinguished — BR,

H, HS, RB, SC; C. muehlenbergii var. muehlenbergii , an orthographic
variant — KM)

Inflorescence spicate, as in C. aggregata , but leaf sheets tight and not septate-
nodulose; pistillate scales as long as the perigynia and rough-awned; perigynium
body orbicular to ovate and strongly nerved on both surfaces. Dry (most often
acidic) forests and edges. [DE, PA, VA, WV-h]. Frequent, WS, ES; rare, RV, PD.

Carex muricata Linnaeus

Paira’s Sedge Phaestoglochin (Bracteosae)

(GC, KM, RB; C. pairaei F. W. Schultz — BR, F)

Similar to C. spicata and C. divulsa , with inflorescence spicate;
pistillate scales brown to blackish brown. Basal sheaths brown to
blackish, not purple-tinged, as in C. spicata. Perigynia spreading (not
appressed-ascending, as in C. divulsa ), rounded at base and rather abruptly beaked.
See Jermy et al. (1982). Roadside. [PA]. Locally adventive (European) (single collec¬
tion), PD-h.

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Carex nigromarginata Schweinitz
Black -margined Sedge Acrocystis (Montanae)

(BR, F, GC, H, HS, KM, RB, SC, T) .

Distinctive sedge, with culms of various lengths that are shorter than
the rigid, dark green leaves. Pistillate scales purple to brown-margined,
obscuring the perigynia, which have fusiform-obovoid to ellipsoid,
pubescent bodies. See Rettig (1990a). Dry forests; openings; barrens. [DE,
PA, VA, WV]. Common, WS, ES, PD; frequent, RV.

Carex normalis Mackenzie
Greater Straw Sedge Ovales

(BR, F, GC, H, KM, RB, SC, T)

Variable. Culms leafy. Sterile shoots usually numerous. Spikes some¬
what congested to remote, with perigynia beaks somewhat spreading.

Similar to C. festucacea, but with ventral band of leaf sheath prolonged at
summit; perigynium body narrowly ovate. Moist forests, fields, and meadows;
floodplains. [DE, PA, WV, VA]. Common, AP; frequent, RV, PD; rare, ES (disjunct).

Carex oligocarpa Schkuhr ex Willdenow
Few-fruited Sedge Griseae

(Oligocarpae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. hitchcockiana. Leaves narrower (2-4 mm); basal sheaths
purplish; upper leaf sheaths glabrous; perigynium smaller (3. 5-4.0 mm long).

Very similar to C. planispicata , but longest lateral (pistillate) spikes with fewer (4-8)
perigynia; mature perigynium distinctly beaked, with more globose body. See Naczi
(1999). Mesic forests, usually on basic soils. [DE-h, PA, VA, WV]. Infrequent, RV; rare,
PD, WS.

Carex oxylepis Torrey and Hooker var. oxylepis
Sharpscale Sedge Hymenochlaenae (Gracillimae)

(F, GC, H, KM)

Similar to C. gracillima, but sheaths and, often, leaves pilose. Similar
to C. aestivalis , but leaves wider (3-10 mm); spikes thicker (3-6 mm).
Perigynium sharply angled and short-beaked. Floodplain forest. [VA]. Rare
(single station), WS.

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Carex pallescem Linnaeus

Pale Sedge Porocystis (Virescentes)

(GC, H, HS, KM, RB; C. pallescem Linnaeus van
neogaea Femald — F)

The only member of section with terminal spike entirely staminate.
Leaves pilose; perigynium glabrous and beakless. Moist fields and
meadows. [PA, VA]. Rare, PD-h, WS-h. Disjunct from farther north.

Carex pedunculata Muhlenberg ex Willdenow
Long-stalked Sedge Clandestinae (Digitatae)

(BR, F, GC, H, KM, RB, SC, T)

Distinctive sedge with terminal spike androgynous and lateral spikes
pistillate and long-peduncled from green, spathe-like sheaths.

Perigynium with a basal eliasome. Pistillate scales conspicuously and abruptly
cuspidate and purple-brown with hyaline margins. Calcareous, mesic soils. [DE-h, PA,
VA, WV] . Rare, AP, RV.

Carex peilita Muhlenberg ex Willdenow
Woolly Sedge Paludosae (Hirtae)

(GC, KM, RB; C. lanuginosa Michaux, misapplied — BR, F, HS, SC, T;

C lasiocarpa Ehrhart, in part — H)

Strongly rhizomatous, with pubescent perigynia. Resembles C.
lasiocarpa , but leaves wider (1 .5-5.0 mm) and flat. Perigynium beak firm
with teeth 03-0.8 mm long. See McClintock and Waterway (1994). Rich fens; wet
meadows; ditches; pastures. [DE, PA, VA, WV]. Infrequent, PD; rare, AP, RV, WS, ES.

Carex pensylvanica Lamarck
Pennsylvania Sedge Acrocystis (Montanae)

(BR, GC, H, HS, KM, RB, SC; C. pensylvanica var. pensylvanica, in a
narrow sense — F; C. pennsylvanica , an orthographic variant — T)

Colonial by rhizomes. Leaves narrow (1-3 mm); lateral (pistillate)
spikes sessile; perigynium body globose and pubescent, with a narrow,
short (0.2-0. 9 mm) beak that is one-fifth to one-fourth as long as the body. See Crins
and Ball (1983). Dry to mesic forests. [DE, PA, VA, WV]. Common, RV, PD, WS, ES;
frequent, AP.

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Carex planispicata Naczi

Flat-spiked Sedge Griseae (Oligocarpae)

(RB; C. amphibola Steudel var. amphibola , misap¬
plied — F, KM; C. amphibola Steudel var, rigida Femald, in part ■

BR, T; C, amphibola Steudel, in part — GC, H, HS, SC)

Similar to C. amphibola , but shoot bases purple; perigynia
distichously arranged. Very similar to C. oligocarpa , but longest lateral
(pistillate) spikes with more (7-14) perigynia; perigynium beakless, trigonous. See
Naczi (1999). Rich, dry to mesic forests, often on mafic soils. [DE, PA, VA, WV].
Rare, PD.

Carex plantaginea Lamarck
Plantain-leaved Sedge Careyanae (Laxiflorae)

(BR, F, GC, H, KM, RB, SC)

Distinctive, with purple base and wide ( 1 0-30 mm) leaves with promi¬
nent parallel veins. Fertile culms leafless, with bladeless purple
sheaths. Perigynium sharply 3-angled. Staminate scales purple. See Manhart
(1986). Rich hardwood forests. [PA, VA, WV]. Rare, AP, PD-h; Piedmont plants are
locally adventive (from Appalachian region) or planted.

Carex platyphylla Carey

Broad-leaved Sedge Careyanae (Laxiflorae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Leaves wide (12-30 mm), as with C albursina and C. plantaginea, but
strongly glaucous. Basal sheaths whitish. Fertile culms unwinged,
leafy; perigynium sharply trigonous. See Manhart (1986). Rich forests, often
on calcareous or mafic soils. [PA, VA, WV], Frequent, RV, PD; rare, WS.

Carex polymorpha Muhlenberg
Variable Sedge Paniceae

(BR, F, GC, H, KM, RB, SC, T)

Colonial sedge with hard and stout rhizomes; aphyllopodic.

Perigynium with elongated beak. See Standley and Dudley (1991).

Open, sandy forests and woodlands; barrens. [DE h, PA, VA, WV]. Rare
(single collection), PD-h.

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Carex prasina Wahlenberg

Drooping Sedge Hymenochlaenae (Gracillimae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Similar to C. gracillima , but with basal sheaths brownish to greenish;
perigynium strongly trigonous, with nerves over the angles of the
achene. Seepage swamps; edges of small streams and spring runs;
occasionally on floodplains; in mucky soils. [DE, PA, VA, WV]. Frequent, AP,
RV, PD; infrequent, WS, ES.

Carex projecta Mackenzie
Necklace Sedge Ovales

(BR, F, GC, HS, KM, RB, SC; C. tribuloides Wahlenberg var. reducta
Bailey; C tribuloides , in part — H)

Much like C tribuloides; spikes generally smaller; inflorescence
moniliform; perigynia tips spreading or loosely ascending. Wet meadows and
fens, at high elevations. [PA, WV, VA]. Infrequent, AP. Reported from PD (e.g.J
Hitchcock and Standley (1919)), but all such plants seen have proven to be C.
tribuloides.

Carex radiata (Wahlenberg) Small
Eastern Star Sedge Phaestoglochin (Bracteosae)

(GC, KM, RB; C. rosea Schkuhr, misapplied — BR, F, HS, SC, T; C.
rosea , in part, misapplied — H)

Similar to C rosea , C. appalachica. Leaves 0.9“ 1.7 mm wide.

Perigynia usually 3-7 per spike, the lowest reflexed at maturity. Stigmas
straight to slightly twisted. Base of achene 0.5-0. 9 mm from perigynium base. See
Webber and Ball (1979, 1984). Moist to mesic forests; edges. [DE, PA, VA, WV].
Common, RV, PD, WS; frequent (rare southward), ES.

Carex retroflexa Muhlenberg ex Schkuhr
Reflexed Sedge Phaestoglochin (Bracteosae)

(BR, F, GC, KM, H, HS, RB, SC, T)

Similar to C. rosea , but base of perigynium bulging on adaxial surface,
distinctly nerved; beak of perigynium smooth-margined. Similar to C.
texensis , but with perigynium shorter-beaked, 2. 0-2. 5 times as long as wide.
Dry, rich forests. [DE, PA, VA, WV]. Infrequent, RV, PD; rare, WS-h, ES.

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Carex richardsonii R Brown
Richardson’s Sedge Clandestinae (Digitatae)

(F, GC, KM, RB)

Rhizoma.ro us sedge superficially resembling C. pensyivanica. Basal
leaves thick and stiff; spikes with purple sheathing bracts; terminal
spike staminate, purplish. Perigynium pubescent. Pistillate scales
obtuse to acute and purple-brown with hyaline margins. Serpentine barrens.
[PA]. Rare, PD.

Carex rosea Schkuhr ex Willdenow
Rose Sedge Phaestoglochin (Bracteosae)

(GC, KM, RB; in part only — H; C. convoluta Mackenzie — BR, F, SC,

T; in part — HS)

Similar to C. radiata and C. appalachica. Leaves 1.7-3. 0 mm wide.

Perigynia 7-14 per spike, the lower reflexed at maturity. Stigmas tightly coiled.
Achene set low in perigynium, with base 0. 1 -0.5 mm from perigynium base. See
Webber and Ball (1979, 1984). Upland forests. [DE, PA, VA, WV]. Frequent, AP, RV,
PD; infrequent, WS, ES (rare southward).

Carex sartwellii Dewey
SartwelPs Sedge Holarrhenae (Intermediae)

(BR, F, GC, KM, RB)

Rhizomes thick, black, scaly. Culms scabrous, exceeding the
scabrous leaves. Inflorescence narrowly cylindrical; spikes variable:
staminate, pistillate, or androgynous. Perigynium lance-ovate, distinctly
nerved, with serrulate, bidentate beak. Calcareous wet meadows. [PA-h]. Rare (single
collection), AP-h. Disjunct from further north.

Carex scabrata Schweinitz
Eastern Rough Sedge Anomalae

(BR, F, GC, H, KM, RB, SC, T)

Rhizomatous sedge with wide (0.6-1 .8 mm), scabrous leaves; culms
scabrous. Perigynium scabrous-puberalent, ovoid-trigonous, promi¬
nently fe w nerved, with an abrupt, distinct beak nearly as long as the body.

Banks of small to medium-sized streams; seepages. [DE, PA, VA, WV]. Frequent, AP;
rare, RV, PD.

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Car ex scop aria Schkuhr ex Willdenow var. scoparia
Pointed Broom Sedge Ovales

(KM; infraspeeific taxa not distinguished — BR, F,

GC, H, HS, RB, SC, T)

Similar to C. tribuloides , but leaves generally more narrow (1 -3 mm).

Perigynia tightly ascending and thin and scale-like as in C. tribuloides ,
but with wings continuous to base. Inflorescence arrangement variable.

Moist, or (occasionally) dry, open areas. [DE, PA, WV, VA]. Common, AP; frequent,
RV, PD, WS; infrequent, ES.

Carex seorsa Howe

Weak Stellate Sedge Stellulatae

(BR, F, GC, H, KM, RB, T; C. rosaeoides Howe — HS)

Distinguished from other members of section, by the elliptic-ovate
perigynium body that is broadest near the middle and by the short,
smooth-margined perigynium beak. See Reznicek and Ball (1980). Swamp
forests; seepages; floodplains. [DE, PA, VA, WV]. Common, ES; frequent, WS; rare,
AP (disjunct), PD-h.

Shortianae

Carex shortiama Dewey
Short’s Sedge

(BR, F, GC, H, HS, KM, RB, SC)

Distinctive sedge, with densely flowered, cylindrical spikes, the
terminal gynecandrous and the lateral pistillate. Perigynia squarrose-
obovoid and brownish at maturity. Rich, open forests, woodlands, and wet

meadows on floodplains. [PA, VA, WV]. Infrequent, RV; rare, PD, WS-h. Mostly along
Potomac River.

Carex siiicea Olney

Seabeach Sedge Ovales

(BR, F, GC, H, KM, T)

Distinctive within section. Leaves grayish-green with auricled bases.
Inflorescence moniliform, nodding. Tips of perigynia equal in length
to or exceeded by subtending scales. Open, sandy woodlands and
shrablands of barrier islands. [DE, VA]. Rare (locally frequent), coastal, ES.

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Carex sparganioides Muhlenberg ex Willdeeow
Bur-reed Sedge Phaestoglochin (Bracteosae)

(BR, F, HS, KM, RB, SC, T; in part only — H; C.
sparganioides van sparganioides — GC)

Similar to C. aggregate?, but with wider (5-10 mm) leaves and with
inflorescence elongate with lower spikes remote (separate by up to 3
cm). Rich forests and meadows, usually on calcareous or mafic soils. [DE,

PA, VA, WV-h]. Rare, RV, PD, WS, ES.

Carex spicata Hudson

Prickly Sedge Phaestoglochin (Bracteosae)

(BR, F, GC, H, KM, RB, T; C. muricata Linnaeus, misapplied — HS; C.
contigua Hoppe)

Inflorescence spicate; pistillate scales often strongly tinged with
brown or reddish purple. Similar to C divulsa and C. muricata, but basal
sheaths purplish (not brown to blackish); ligule prolonged, much longer than broad;
perigynium larger (4-6 mm long). See Jermy et al. (1982). Fields; other open, disturbed
sites. [DE, PA]. Locally naturalized or adventive (European), PD, WS-h.

Squarrosae

Carex squarrosa Linnaeus
Squarrose Sedge

(BR, F, GC, H, HS, KM, RB, SC, T)

Perigynia abruptly beaked, with obovoid bodies, and densely packed.
Terminal spike gynecandrous, as in C. typhina, but spikes
subglobose; perigynia beaks widely divergent, the lower reflexed; style

strongly contorted at base. Moist fields, meadows, and forests; floodplains. [DE, PA,
VA, WV]. Frequent, RV, PD, WS; rare, ES.

Carex stipata Muhlenber ex Willdenow van maxima
Chapman

Greater Stalkgrain Sedge Vulpinae

(F, GC, KM, RB, SC, T; infraspecific taxa not distinguished — BR, H,

HS; C uberior (Mohr) Mackenzie)

Stouter and with leaves wider (8-17 mm) than var. stipata ; perigynium longer
(5-6 mm), with beak longer (> 3 mm), as long as or longer than the body. Swamps;
floodplain forests; fresh tidal marshes. [DE, PA, VA, WV]. Frequent, WS, ES; rare, PD.

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Carex stipata Muhlenberg ex Willdenow var. stipata
Stalkgrain Sedge Vulpinae

(F, GC, KM, RB, SC, T; inffaspecific taxa not distinguished — BR,

H, HS)

Similar to C. laevivaginata , but inner band of leaf sheaths friable,
cross-puckered. Similar to var. maxima ; leaves narrower (4-10 mm);
perigynium shorter (4-5 mm), the beak shorter (under 2.5 mm). Swamps; wet meadows;
fens; floodplain forests. [DE, PA, VA, WV], Frequent, AP, RV, PD; infrequent, WS;
rare, ES.

Carex straminea Willdenow ex Schkuhr
Eastern Straw Sedge Ovales

(BR, F, GC, KM, RB, SC; in part only — H; C.
hormaihodes Femald, in part — HS; C richii Mackenzie — T)

Pistillate scales awn-tipped. Spikes well-separated (congested in C.
alata ); inflorescence often nodding . Perigynium body ovate to
suborbicular, abruptly narrowed to beak (continuously narrowed in C.
hormaihodes). See Rothrock et al. (1997). Seasonally dry pools; swamps; other non-
tidal wetlands. [DE, PA, VA, WV] . Rare, RV, PD, WS, ES.

Carex striata Michaux var. brevis Bailey
Walter’s Sedge Paludosae (Hirtae)

(GC, KM; infraspecific taxa not distinguished — H; C.
walteriana Bailey var. brevis (Bailey) Bailey — F; C walteriana
Bailey — T)

Rhizomatous. Similar to C. pellita, perigynia glabrous and with im¬
pressed nerves. See Reznicek and Catling (1986) for discussion of nomencla¬
ture. Delmarva Bays; ditches; other open, often disturbed wetlands. [DE, VA].
Frequent (rarer northward), ES.

Carex striata Michaux var. striata
Pocosin Sedge Paludosae (Hirtae)

(GC, KM; C walteriana Bailey var. walteriana —

Similar to var. brevis , but with perigynium puberulent, not glabrous
(pubescent in C. pellita). See Reznicek and Catling (1986) for discus¬
sion of nomenclature. Coastal Plain “bog.” [none]. Rare (single station),

WS. Although the range of var. striata has been said to be limited to areas well
to the south of Maryland (Femald 1950; Gleason and Cronquist 1991), forms with
puberulent perigynia actually occur sparingly north to New Jersey (A. A Reznicek,
pers. comm.)

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Care x striatula Michaux
Lined Sedge Laxiflorae

(BR, F, GC, HS, KM, RB, SC, T; in part only — H)

Very similar to and possibly not taxonomically distinct from, C.
laxiflora (see Manhart, 1986). Terminal (staminate) spike on longer
peduncle (more than 20 mm); lateral (pistillate) spikes thicker (4-8 mm),
perigynium longer (4. 0-5. 5 mm), fusiform, rather than obovoid. Dry to mesic forests.
[DE, PA, VA, WV]. Infrequent, RV, PD, ES; rare, WS.

Carex stricta Lamarck

Tussock Sedge Phacocystis (Acutae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Form somewhat variable; densely cespitose to freely rhizomatous.

Similar to C. emoryi and C. haydenii , but with lower leaf sheaths
fibrillose. Summit of leaf sheaths concave. Lower lateral (pistillate)
spikes usually remote. Perigynium ovate to elliptic, barely or not inflated,
granular-papillose above the middle. See Standley (1989). Fens; wet meadows;
swamps; alluvial forests. [DE, PA, VA, WV]. Common, AP, WS, ES; frequent, RV, PD.

Carex styloflexa Buckley
Bent Sedge Laxiflorae

(BR, F, GC, HS, KM, RB, T; included in C. striatula
Michaux — H)

Fairly distinctive within section. Basal sheaths brownish- whitish.

Terminal (staminate) spikes long-peduncled. Lateral (pistillate) spikes
densely few-flowered and remote, the lowest on elongate, filiform peduncles.
Perigynium beak arching. See Manhart (1986). Moist forests; tidal swamps; seepages.
[DE, PA, VA, WV]. Frequent, WS, ES; infrequent, PD; rare, RV.

Carex swanii (Femald) Mackenzie
Swan’s Sedge Porocystis (Virescentes)

(BR, F, GC, H, HS, KM, RB, SC, T)

Leaves and perigynia pubescent, as with C. vires cens , but spikes
subglobose to thick-cylindrical, 0.5-1 .8 cm long; perigynium less
strongly nerved; anthers smaller (0.7-1 .6 mm). Dry to mesic forests and
fields. [DE, PA, VA, WV]. Common, PD, WS, ES; frequent, AP, RV.

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Carex tenera Dewey

Slender Sedge Ovales

(BR, F, GC, H, KM, RB, SC)

Other taxa in this section are sometimes misidentified as this species
in collections. Perigynium body ovate, as in C. normalis, but leaves
narrower (1 .0-2.5 mm); entire inflorescence moniliform and nodding.
Floodplain forests. [PA, VA, WV]. Rare, PD, WS-h.

Carex tetanica Schkuhr
Rigid Sedge Paniceae

(BR, F, GC, H, KM, RB, T; in part only — HS)

Rhizomatous and similar to C. woodii, but with basal sheaths mostly
bearing blades; lateral (pistillate) spikes more densely flowered.

Similar to C. meadii , but leaves green (not gray-green); lateral spikes
narrower (3-5 mm), with only 2-3 vertical rows of perigynia. Meadow. [PA,

VA, WV], Rare (single collection), PD-h. Reports from DC area (e.g., Hitchcock and
Standley (1919)) are evidently based on C. woodii or on misidentifications.

Carex texensis (Torrey) Bailey
Texas Sedge Phaestoglochin (Bracteosae)

(BR, F, KM, RB; C. retroflexa Muhlenberg var. texensis (Torrey)

Femald — GC)

Similar to C. retroflexa , but base of adaxial surface of perigynium not
bulging and nerveless to obscurely nerved; perigynium longer beaked
and relatively more narrow (about 3 times as long as wide). Dry forests and
fields; disturbed areas. [DE, PA, VA, WV]. Locally naturalized or adventive (from
southern United States), RV-h, PD.

Carex tonsa (Femald) Bicknell var. rugosperma
(Mackenzie) Crins

Rough-seeded Sedge Acrocystis (Montanae)

(RB; C. rugosperma Mackenzie — KM, T; C. umbellata Schkuhr,
misapplied — F; in part — BR, GC, H, SC)

Similar to C. umbellata , but perigynium beak as long as the body. Similar to
var. tonsa , but leaves narrower (1 .5-3.0 mm); perigynium membranaceous, minutely
pubescent. See Cusick (1992), Rettig and Crins (1996). Dry forests; rocky openings.
[PA, VA, WV]. Frequent, AP; infrequent, RV; rare, PD.

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Carextonsa (Femald) Bicknell var. tonsa
Shaved Sedge Acrocystis (Montanae)

(RB; C. tonsa , in a narrow sense — BR, F, KM, T; in part
only — HS; C. umbeilata Schkuhr, in part — GC, H, SC; C.
rugosperma Mackenzie var. tonsa (Femald) Voss, misapplied)

Similar to var. rugosperma , but with short, rigid, wider (2. 0-4.5 mm)
leaves; perigynia glabrous and shining. See Cusick (1992), Rettig and Crins
(1996). Dry, often sandy, forests and openings; rock outcrops. [DE, PA, VA, WV].
Common, ES; frequent, RV, PD, WS; rare, AP.

Carex torta F. Boott ex Tuckerman
Twisted Sedge Phacocystis (Cryptocarpae)

(BR, F, GC, H, HS, KM, RB, SC, T)

Short-rhizomatous sedge in dense clumps. Lower lateral (pistillate)
spikes drooping; perigynium beak bent or twisted; pistillate scales
dark brown with green midrib. Along banks of and in beds of higher-
gradient small to medium-sized streams. [DE, PA, VA, WV]. Common, AP;
frequent, RV; infrequent, PD; rare, WS.

Carex tribuloides Wahlenberg var. tribuloides
B lunt Broom Sedge Ovales

(KM; infraspecific taxa not distinguished — BR, F, GC, HS, RB, SC,

T; C. tribuloides , in part only — H)

Spikes mostly crowded. Perigynia thin and scale-like, with strongly
ascending tips, as in C. scoparia , but with wings abmptly narrowed at
the base. Some Maryland plants may be var. sangemonensis Clokey. Moist
forests and open areas, floodplains. [DE, PA, VA, WV]. Common, RV, PD; frequent,
WS, ES; infrequent, AP.

Carex trichocarpa Muhlenberg ex Schkuhr

Hairy-fruited Sedge Carex (Paludosae)

(H; RB; C. trichocarpa Muhlenberg — BR, F, GC, KM,

SC,T)

Rhizomatous sedge; outwardly similar to C. lacustris, but with
pubescent perigynia, with teeth up to 2.7 mm long. Summit of inner bands of
leaf sheaths concave and purple-tinged. Wet meadows; open stream banks. [DE, PA,
VA, WV] . Rare, PD.

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Care x trisperma Dewey var. trisperma
Three-seeded Sedge Glareosae (Heleonastes)

(GC, KM; infraspecific taxa not distinguished — BR,

F, H, RB,

SC,T)

Distinctive sedge with narrow-leaves (1-2 mm) and weak, filiform,
decumbent culms. Similar to C. brunnescens , but with lowest inflorescence
bract long-linear, much exceeding the spikes, which are few-flowered (1-5). Fens;
seepages; usually on Sphagnum. [DE-h, PA, VA, WV]. Frequent, AP.

Carex tuckermanii F. Boott ex Dewey
Tuckerman’s Sedge Vesicariae

(F, GC, KM, RB)

Most similar to C. bullata and C. vesicaria , but with perigynium
wider (more than 4.5 mm) and of papery texture; achene asymmetrical
and deeply indented in the middle of one angle. Lateral (pistillate)
spikes wider (1 .2-1 .8 cm) than in C. vescaria. Grows in clumps, unlike the
rhizomatous C. bullata. Floodplains; fens. [PA, WV]. Rare, AP.

Carex typhina Michaux

Cattail Sedge Squarrosae

(BR, F, GC, H, HS, KM, RB, SC, T)

Distinctive sedge. Perigynia abruptly beaked, with obovoid
bodies, and densely packed and terminal spike gynecandrous, as in C.
squarrosa , but spikes cylindrical; perigynia beaks spreading to
ascending; style straight or curved at the base. Floodplain forests;
forested vernal pools. [DE, PA, VA, WV]. Infrequent (locally common), WS, ES;
rare, RV, PD.

Carex umbel lata Schkuhr ex Willdenow
Umbellate Sedge Acrocystis (Montanae)

(KM, RB, T; in part only — BR, GC, H, SC; C. abdita Bicknell — F; C.
tonsa (Femald) Bicknell, in part — HS)

Densely cespitose sedge, with narrow (1-3 mm wide) leaves. Mature
spikes nearly hidden among leaf bases. Perigynium minutely pubescent, with
the beak half as long as the body. See Cusick (1992), Rettig and Crins (1996). Dry
forests and openings. [DE, PA, VA, WV]. Common, PD, WS, ES; infrequent, RV.

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The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

Carex utriculata F. Boott
Northwest Territory Sedge Vesicariae

(GC, KM, RB; C. rostrata Stokes, in a broad sense — H, HS; C
rostrata Stokes var. utriculata (F. Boott) Bailey — BR, F, SC; C inflatai
Fludson var. utriculata (F. Boott) Druce — T)

Rhizomatous, forming large colonies. Leaves and sheaths septate-nodulose.
Perigynia strongly nerved, spreading, densely crowded, mostly in 8 or more vertical
rows. Fens; fresh tidal marsh. [DE-h, PA, VA,WV]. Infrequent, AP; rare, RV, WS-h
(disjunct).

Carex venusta Dewey var. minor Boeckeler
Dark Green Sedge Hymenochlaenae (Sylvaticae)

(BR, F, GC, KM; infraspecific taxa not distinguished
— H; C. oblita Steudel — HS, T)

Similar to C. debilis var. debilis, but perigynia more densely overlap¬
ping, more strongly nerved, and with shorter (0.5 mm long), stouter
beaks. Mesic to moist forests. [DE, VA], Infrequent, WS; rare, PD-h, ES.

Carex vesicaria L. var. monile (Tuckerman) Femald
Inflated Sedge Vesicariae

(F, KM; infraspecific taxa not distinguished — BR, H,

C. vesicaria L. var. vesicaria , in a broad sense — GC)

Similar to C. bullata , but cespitose (without elongated rhizomes) and
with lateral (pistillate) spikes more narrowly cylindrical (4-15 mm thick);
perigynium beak smooth; distribution primarily montane. Wet meadow. [DE,
PA, VA (possibly var. vesicaria ), WV]. Rare (single station), AP.

Carex vestita Willdenow

Velvety Sedge Paludosae (Hirtae)

(BR, F, GC, H, HS, KM, RB, T)

Strongly rhizomatous. Distinguished from other members of the
section that also have the perigynia pubescent (C. lasiocarpa, C.
pellita ), primarily by having the perigynium beak obscurely bidentate,
with soft teeth; lowermost inflorescence bract shorter and rarely reaching the
terminal (staminate) spike, which is on a relatively shorter peduncle. Open, sandy
wetlands. [DE, PA, VA]. Infrequent ES; rare, WS.

Winter 2001

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C. T. Frye & C. Lea

Carex virescens Muhlenberg ex Willdenow
Ribbed Sedge Porocystis (Virescentes)

(BR, F, GC, H, HS, KM, RB, SC, T)

Leaves and perigynia pubescent, as in C. swanii , but with spikes
linear-cylindrical, 1 .5-4.0 cm long; nerves of perigynium more pro¬
nounced, often whitened; anthers larger (1 .5-2.5 mm). Dry forests. [DE,
PA, VA, WV]. Common, RV, PD, WS; frequent, AP; rare, ES.

Carex viridula Michaux
Little Green Sedge Spirostachyae
(Ceratocystis)

(F, GC, KM, RB; C. oederi Retz, misapplied — HS)

Terminal spike staminate or pistillate at tip. Lateral (pistillate) spikes
ovoid to short-cylindrical, crowded near summit of culm. Abruptly-
beaked perigynia spreading to slightly reflexed. See Crins and Ball ( 1 989a,

1989b). Wet, often calcareous, soil; shores; springs. [PA]. Rare (single collection), PD
h or WS-h. Specimen reported by Hitchcock and Standley (1919) not found.

Carex vulpinoidea Michaux
Fox Sedge Multiflorae

(BR, F, GC, HS, RB, SC, T; in part only — H; C.

vulpinoidea var. vulpinoidea — KM)

Similar to C. annectens , but with mature culms generally exceeded by
the leaves; lowermost inflorescence bract longer (often more than 5
cm); perigynium beak about as long as body. Moist to wet, open (often
disturbed) habitats. [DE, PA, VA, WV]. Common throughout.

Carex willdenowii Schkuhr ex Willdenow
Willdenow’s Sedge Phyllostachyae

(BR, F, H, HS, SC, T; C. willdenovii , an orthographic
variant — GC, RB; C. willdenowii var. willdenowii , in a narrow sense
— KM)

Staminate scales tightly clasping, as in C. jamesii , but perigynium
body ellipsoid-fusiform, more gradually tapered to a stout, serrate beak. See
Catling et al. ( 1 993), Naczi et al. ( 1 998). Dry forests. [DE, PA, VA,

WV]. Common, PD, WS; infrequent, RV; rare, AP-h.

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The Maryland Naturalist Volume 44

Atlas and Annotated List of Carex (Cyperaceae) of Maryland and the District of Columbia

Carex wood'd Dewey
Wood’s Sedge Paniceae

(BR, F, GC, H, KM, RB; C. tetanica Schkuhr var.
woodii (Dewey) Bailey — HS)

Rhizomatous in colonies; bases purple. Resembles C. tetanica, but
with basal sheaths mostly bladeless; lateral (pistillate) spikes loosely
and alternately flowered. Colonies appear like those of C. pensylvanica, but
leaves blue-green, lateral spikes peduncled. Dry to mesic, rich forests. [PA, VA, WV].
Frequent, RV; rare, AP, PD. Evidently overlooked in past.

Winter 2001

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C. T. Frye & C. Lea

Addendum — Records for the following two taxa were discovered after the manuscript
went to press, but have been included here to maintain completeness of coverage.
(Note: Text and figures modified to accomodate these.)

Carex caryophyllea Latourette
Spring Sedge Mitratae (Praecoces)

(BR,F,GC,KM)

Rhizomatous sedge with fibrillose bases and stiff,
scabrous leaves. Terminal (staminate) spike scales
red-brown. Lateral (pistillate) spikes sessile or nearly
so. Perigynium hirsute, with short beak; pistllate

scales red-brown and cuspidate; achene capped by a dark annulus with a central
projecting apex. See Jenny et al. (1982). Cultivated area. [none]. Locally adventive
(European) (single collection),

WS-h.

Carex digitalis Willdenow var. asymmetrica Femald
Large-fruited Slender Woodland Sedge

Careyanae (Laxiflorae)

(F; infraspecific taxa not distinguished — GC, H,

KM).

Similar to var. digitalis , but perigynia longer (3. 2-4.2
mm), with the apex excurved and the angles more
obscure. SeeNaczi et al. (2001). Dry-mesic forest. [VA]. Rare (single station), WS.
Some authors question its taxonomic distinctiveness (see Manhart, 1986).

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The Maryland Naturalist

Volume 44

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The Maryland Naturalist is a peer reviewed biannual publication of the Natural
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Winter 2001

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