NEW RECORDS AND A TAXONOMIC REVIEW OF
CALAMAGROSTIS PERPLEXA (POACEAE: POEAE:
AGROSTIDINAE), A NEW YORK STATE ENDEMIC GRASS
TIMOTHY G. HOWARD
New York Natural Heritage Program, 625 Broadway, 5th floor,
Albany, NY 12233-4757
e-mail: tghoward@gw.dec.state.ny.us
JEFFERY M. SAARELA
Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa,
ON K1P 6P4, Canada
e-mail: jsaarela@mus-nature.ca
BEATA PASZKO
W. Szafer Institute of Botany, Polish Academy of Sciences,
31-512 Krako?w, Poland
e-mail: ibpaszko@ib-pan.krakow.pl
PAUL M. PETERSON
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20013-7012
e-mail: peterson@si.edu
DAVID WERIER
30 Banks Rd., Brooktondale, NY 14817
e-mail: nakita@lightlink.com
ABSTRACT. Calamagrostis perplexa is a New York State endemic grass
species, hypothesized to be of hybrid origin, that previously was known only
from a single locality. We provide a summary of its taxonomic history, report
new New York records and ecological information for the taxon, and include a
principal components analysis and key characterizing morphological variation
in C. perplexa and its putative close relatives C. canadensis, C. porteri, C.
rubescens, and C. scribneri. Calamagrostis perplexa is now known from two
extant populations and a third population of unknown status; it remains a New
York state endemic.
Key Words: Calamagrostis, endemic, grass, hybrid, New York, Poaceae,
principal components analysis (PCA)
RHODORA, Vol. 111, No. 946, pp. 155?170, 2009
E Copyright 2009 by the New England Botanical Club
155
The grass genus Calamagrostis Adans. [Poeae (sensu Soreng et al.
2007): Agrostidinae Fr.] is characterized as having single-flowered
spikelets, one- or three-veined glumes as long as or exceeding the
floret in length (rarely slightly shorter), non-keeled lemmas that are
membranous or cartilaginous (infrequently hyaline), usually with a
single dorsally attached awn (rarely awnless), a callus with a crown
of hairs, caryopses with short hilums and without apical hairs, and
lodicules that are apically membranous (Clayton and Renvoize
1986; Watson and Dallwitz 1992). In North, Central, and South
America, 131 Calamagrostis species are reported by Soreng and
Greene (2003); 25 species are currently recognized in North
America (Marr et al. 2007). Species that occur in eastern North
America include the introduced C. epigeios (L.) Roth; the
widespread C. purpurascens R. Br., C. canadensis (Michx.) P.
Beauv., and C. stricta (Timm) Koeler; and the eastern endemics C.
coarctata Torr. ex Eaton, C. cainii Hitchc., C. porteri A. Gray, C.
pickeringii A. Gray, and C. perplexa Scribn. In this paper we review
the taxonomic history of C. perplexa, report new records and
ecological information for the species, characterize morphological
variation among C. perplexa and its putative close relatives, and
provide a brief taxonomic synopsis for the species.
Taxonomic history of Calamagrostis perplexa. Plant material
from ??Thatcher?s Pinnacles,?? Danby, in Tompkins County, New
York, was initially described as Calamagrostis nemoralis Kearney in
1898, but this name is an illegitimate homonym; a Chilean species
was named C. nemoralis Phil. [5 C. viridis (Phil.) Soreng] two years
earlier. The taxon was given a new name in 1901 as C. perplexa
Scribn., replacing C. nemoralis Kearney. Calamagrostis perplexa
(wood reedgrass) is distinguished from other North American
Calamagrostis species by its combination of densely hairy collars
and callus hairs that are 0.7?1 times as long as the lemma (Marr et
al. 2007). Hitchcock (1935) reported C. perplexa from four
additional eastern North American locations, but subsequent study
determined that these collections are other taxa (Louis-Marie 1944).
Calamagrostis perplexa has since been confirmed only at the type
location (e.g., Gleason and Cronquist 1991; Greene 1980; Hitch-
cock 1951; Marr et al. 2007). Calamagrostis perplexa was reported
for Mount Fray in Columbia County, New York (McVaugh 1958);
this record was recently mentioned as an unconfirmed report (Marr
et al. 2007). Given its highly restricted range, C. perplexa is globally
156 Rhodora [Vol. 111
ranked G1 (critically imperiled throughout its range), New York
state ranked S1 (critically imperiled in New York State), and it has
a New York state status of E (endangered species; Young 2007).
The evolutionary affinities of Calamagrostis perplexa are not clear,
although some authors have suggested that the species is closely
related to other North American species with which it shares bent
awns, including C. pickeringii, C. porteri, C. purpurascens, and some
individuals of C. stricta (e.g., Greene 1987; Louis-Marie 1944). It has
also been noted that C. perplexa is intermediate in morphology
between C. porteri and C. canadensis (Greene 1980, 1984; Kearney
1898), and Clausen (1949) treated the taxon as C. porteri subsp.
perplexa (Scribn.) R.T. Clausen. Greene (1980) found that C. perplexa
is approximately intermediate between C. canadensis and C. porteri
cytologically, with a chromosome number of 2n 5 70 (cf. 2n 5 84?104
in C. porteri and 2n 5 56 in C. canadensis var. canadensis). Greene
(1980, 1984) suggested that the species might be a hybrid of these taxa.
Indeed, the geographical distributions of these putative parental taxa
are potentially consistent with this hypothesis. Calamagrostis
canadensis is widely distributed across Canada and the United States,
whereas C. porteri has a more restricted distribution in the
northeastern and central United States (Marr et al. 2007). The closest
known population of C. porteri occurs 20 miles to the south of the
Thatcher?s Pinnacles C. perplexa population (Greene 1980), though
Marr et al. (2007) note that historically C. porteri had a larger range.
It would be worthwhile to test the hypothesis that C. perplexa is of
hybrid origin with molecular data.
In terms of morphological characteristics, Calamagrostis perplexa
shares hairy collars with C. porteri and long callus hairs with C.
canadensis. Long callus hairs also occur in many North American
Calamagrostis species (see descriptions in Marr et al. 2007). Hairy
collars are not common in North American Calamagrostis. In
addition to C. porteri, they occur only in some individuals of the
western North American taxon C. rubescens Buckley, which Marr
et al. (2007) noted appears to be closely related to C. porteri, and in
some individuals of C. canadensis var. canadensis [fide Marr et al.
(2007) and Soreng and Greene (2003)] that were originally described
as C. scribneri Beal and recognized as such in some subsequent
treatments (e.g., Hitchcock 1935, 1951; Stebbins 1930). Hitchcock
et al. (1969) placed C. scribneri in synonymy of C. canadensis, but
still recognized the form as C. canadensis var. robusta Vasey, the
earliest name for the taxon at the varietal rank. Calamagrostis
2009] Howard et al.?Calamagrostis perplexa 157
scribneri might represent a hybrid between C. rubescens and C.
canadensis in western North America?a scenario that could be
considered analogous to the putative origins of C. perplexa in
eastern North America, possibly involving one of the same parent
taxa (i.e., C. canadensis). The taxonomic status of C. scribneri
clearly deserves further study. In the absence of a robust phylogeny
for North American Calamagrostis species, the evolution of these
morphological characteristics is not clear: they could represent
convergences, parallelisms, or synapomorphies.
Despite its extremely rare status in New York State and globally,
there is little biological information available for Calamagrostis
perplexa. Other rare but relatively much more common eastern North
American Calamagrostis species have received substantially more
study, including C. porteri subsp. insperata (Swallen) C.W. Greene
(Bittner and Gibson 1998; Esselman et al. 1999; Havens and Holland
1998) and C. cainii Hitchc. (Godt et al. 1996; Wiser 1991). Greene
(1980) indicated that at the type locality, C. perplexa occurred in well-
drained, rocky soil in approximately two acres of somewhat open
forest dominated by Quercus montana, Q. alba, and Acer rubrum, and
that the abundance of flowering culms varied from year to year, with
the population persisting exclusively vegetative in some years.
Clausen (1949) noted that he observed only a single flowering
specimen in August, 1947. Greene (1980, 1984) also noted that C.
perplexa is pollen sterile and that mature fruits have not been
observed for the species. He concluded that the species probably
persists vegetatively, and that the population might represent a single
genetic individual (consistent with its putative hybrid origin) that has
attained its current distribution via extensive rhizome growth.
New ecological information for Calamagrostis perplexa at the
type locality. The Thatcher?s Pinnacles population was visited in
August 2007, and the extent of the population was observed to be
similar to that described by Greene (1980), with two flowering
individuals and about 500 vegetative ramets. Calamagrostis perplexa
occurred in unevenly scattered patches on a rim and adjacent shoulder
of a large valley adjacent to the very steep west-southwest facing slope
of the valley wall. The site varies from being flat to having a moderate
slope to the north and the soils are dry-mesic. The site was forested
with about a 70?80% canopy cover. The trees were of mixed age and
somewhat dwarfed with Quercus montana dominant. Other trees in
the canopy included: Q. rubra, Q. alba, and Carya glabra. The
158 Rhodora [Vol. 111
subcanopy included the species that were in the canopy as well as Acer
rubrum, A. saccharum, Amelanchier arborea, Fagus grandifolia,
Fraxinus americana, Hamamelis virginiana, Ostrya virginiana, Pinus
strobus, and Tsuga canadensis. The small shrub layer was sparse and
included the species in the canopy and subcanopy as well as Acer
pensylvanicum, Cornus florida, Gaylussacia baccata, Prunus serotina,
Rhododendron prinophyllum, Rosa carolina, Vaccinium angustifolium,
V. pallidum, V. stamineum, and Viburnum acerifolium. The herb layer
was patchy with Carex pensylvanica dominant. Other species in the
herb layer included: Anemone americana, Aralia nudicaulis, Asplenium
platyneuron, Comandra umbellata, Desmodium nudiflorum, Eurybia
macrophylla, Galium lanceolatum, Gaultheria procumbens, Hieracium
paniculatum, Maianthemum canadense, Mitchella repens, Monotropa
uniflora, Oryzopsis asperifolia, Poa compressa, Polygonatum biflorum,
Solidago arguta, S. bicolor, S. caesia, and Uvularia perfoliata.
Previously unconfirmed records. We have confirmed that the
record from Columbia County mentioned by Marr et al. (2007) is
indeed Calamagrostis perplexa. This specimen [McVaugh 2649 (PH);
see below] was annotated in 1936 by M. L. Fernald as C. perplexa,
and in 1980 by C. W. Greene as C. stricta subsp. inexpansa (A.
Gray) C.W. Greene var. lacustris (Kearney) C.W. Greene; the latter
name, C. stricta var. lacustris, was never published. Field work is
needed to determine if this population persists.
In their treatment of Calamagrostis canadensis for North America,
Marr et al. (2007, p. 726) noted that ??C. canadensis appears to form
hybrids with the nearly sympatric C. porteri in rocky wooded sites in
central Virginia. A few of these putative hybrids have hairy collars,
relatively long callus hairs, and short awns.?? Based on these
comments, it seemed possible that these putative hybrids might be
C. perplexa, which would represent a considerable range extension for
the taxon. We examined specimens of these putative hybrids and
concluded that they are not C. perplexa, nor do they appear to be
hybrids. Two of these Virginia specimens from near the Appalachian
Trail [Botetourt Co., C.E. Stevens 16731 (VPI-84019); Nelson Co., F.
Watson 567 (VPI-75470)] have been verified as C. porteri and 19 have
been determined to be C. canadensis [Amherst Co., Pompey Mt., C.E.
Stevens 11585 (VPI-62763), Cole Mt., T.F. Wieboldt 3326 (VPI-71724);
Augusta Co., Freezeland Flat, C.E. Stevens 17813 (VPI-85823),
Hardscrable Knob, C.E. Stevens 13608 (VPI-67030), Tearjacket Knob,
C.E. Stevens 17811 (VPI-85826); Bedford Co., Flat Top Mt., C.E.
2009] Howard et al.?Calamagrostis perplexa 159
Stevens 16330, 16331 (VPI-83966, 83965), Thunder Hill, C.E. Stevens
16696 (VPI-84018); Botetourt Co., Apple Orchard Mt., C.E. Stevens
16713 (VPI-84016); Greene Co., Bearfence Rock Scramble, T.F.
Wieboldt 3322 (VPI-69014, 71767); Nelson Co., Crabtree Meadows,
C.E. Stevens 16835 (VPI-83963), Humpback Mt., T.F. Wieboldt 3209
(VPI-77962), C.E. Stevens 18902 (VPI-86013); Page Co., Stoney Man,
C.E. Stevens 10966 (VPI-65049); Rappahannock Co., South Marshall
Mt., C.E. Stevens 19387 (VPI-86049); Warren Co., Hogback Mt., C.E.
Stevens 1570 (VPI-54329), C.E. Stevens 13977, 13978 (VPI-67054,
83207)].
Discovery of another population in New York. In August 2006, the
first author encountered a small population of an unusual
Calamagrostis species along a high ridge trail near Ellenville in
Ulster County in southern New York. We have confirmed that this
represents the second known population of C. perplexa. This newly
discovered population was 190 km (117 mi.) east of the Thatcher?s
Pinnacles population. We revisited this site in August 2007 to better
document population size and habitat information. This population
occurred in patches at a section of the ridge extending over about
400 m2 along a linear zone of about 65 meters. We counted 294
flowering culms in 2007. Patches occurred in openings, adjacent
quartzite conglomerate rock outcrops or ledges, and under low trees
with Quercus coccinea, Q. rubra, Q. alba, Q. montana, and Carya
glabra most common. Dense stands of scrub oak (Q. ilicifolia) with
low blueberries such as Vaccinium pallidum and V. angustifolium also
characterized the site. Other woody plants present included Acer
rubrum and Amelanchier spicata. The herbaceous layer was
dominated by Deschampsia flexuosa and Carex pensylvanica. Other
associated herbaceous plants included: Agrostis perennans, Aralia
hispida, Carex argyrantha, C. debilis var. rudgei, C. lucorum, C.
swanii, Corydalis sempervirens, Danthonia spicata, Dennstaedtia
punctilobula, Dichanthelium acuminatum subsp. columbianum, Erech-
tites hieracifolia, Melampyrum lineare, Poa compressa, Schizachyrium
scoparium, and Viola sagittata var. sagittata.
Discovery of this geographically distinct population of Calama-
grostis perplexa has allowed us to compare morphological
characteristics of multiple specimens from the two populations.
The two populations match in terms of the key characteristics, but
they superficially appear very different; this is likely due to
differences in canopy cover. At Thatcher?s Pinnacles, C. perplexa
160 Rhodora [Vol. 111
was almost entirely vegetative and the culms were short whereas the
Ellenville population had most patches with at least some
reproductive culms and the culms were quite tall (over one meter).
Future work should explore levels and patterns of genetic variation
within and between these populations and the Columbia County
population, if it is found to persist.
Numerical analysis of Calamagrostis perplexa and relatives. To
further characterize variation among Calamagrostis perplexa and its
putative parent taxa (C. canadensis and C. porteri), we conducted a
principal components analysis (PCA) of their morphological
characteristics. We also included the morphologically similar C.
scribneri and C. rubescens (see above) in this analysis. We examined
a total of 220 individual specimens (Appendix) for 18 quantitative
morphological characters (Table 1) using Statistica (StatSoft, Inc.
2005). A complete data set of 220 operational taxonomic units 3 18
characters is available from B. Paszko upon request. Included in the
PCA were C. perplexa (17 individuals), with hypothesized parents
Table 1. Principal component loadings on the first three components for 18
morphological characters scored for Calamagrostis canadensis, C. perplexa, C.
porteri, C. scribneri, and C. rubescens. For each component, values with
particularly large magnitudes are shown in bold.
Characters PC 1 PC 2 PC 3
1. Panicle length (cm) 20.15 20.33 20.19
2. Lower glume length (mm) 0.68 20.59 0.36
3. Lower glume width (mm) 0.66 20.14 20.14
4. Upper glume length (mm) 0.70 20.57 0.32
5. Upper glume width (mm) 0.75 20.25 20.16
6. Callus hairs length (mm) 20.68 20.62 0.02
7. Ratio: lemma to lower glume length 20.27 20.15 20.88
8. Ratio: callus hairs length to lemma length 20.86 20.35 0.05
9. Lemma length (mm) 0.56 20.72 20.17
10. Palea length (mm) 0.90 20.20 20.20
11. Ratio: palea length to lemma length 0.83 0.33 20.12
12. Rachilla extension length (mm) 0.49 20.40 20.29
13. Rachilla extension with hairs length (mm) 20.19 20.77 20.06
14. Lemma awn length (mm) 0.85 20.21 0.18
15. Ratio: distance from base of lemma to awn
insertion (character 16) to lemma length 20.79 20.11 0.07
16. Distance from base of lemma to awn insertion
(mm) 20.67 20.35 0.04
17. Ligule length (mm) 20.46 20.48 0.17
18. Number of culm nodes 20.72 20.37 20.06
2009] Howard et al.?Calamagrostis perplexa 161
C. porteri (27 individuals) and C. canadensis (102 individuals); and
C. scribneri (25 individuals), with hypothesized parents C. rubescens
(49 individuals) and C. canadensis (Figure 1). The first principal
component (PC1) had relatively high (positive or negative) loadings
for palea length, lemma awn length, upper glume width, number of
culm nodes, and upper glume length; PC2 had relatively high
loadings for rachilla extension with hairs and lemma length
(Table 1). The plot of PC1 versus PC2 shows that individuals of
C. perplexa are intermediate in many morphological features
between C. canadensis, which has shorter paleas and shorter lemma
awns, and C. porteri, which has longer paleas and longer lemma
awns (Figure 1), consistent with earlier observations (see above).
Based on the morphological characters we have scored, individuals
of C. scribneri cannot be distinguished from C. perplexa.
Individuals of C. scribneri are intermediate between C. canadensis,
which has shorter paleas, and C. rubescens, which has longer paleas
and shorter callus hairs. These data clearly distinguish individuals
Figure 1. Plot of principal components (PC) 1 (43.8% of the total variation)
and 2 (18.7% of the total variation) for Calamagrostis perplexa and relatives
using 18 morphological characters (see Table 1). Species are indicated as
follows: C. canadensis (# open circles), C. perplexa (N solid circles), C. porteri
(& solid squares), C. rubescens (% open squares), C. scribneri (n open
triangles). Arrows indicate type specimens.
162 Rhodora [Vol. 111
of C. perplexa from either C. canadensis or C. porteri, and C.
scribneri is discernable from either C. canadensis or C. rubescens.
These data argue for a detailed investigation of the variation within
the hypothesized hybrids, C. perplexa, and C. scribneri. A key
separating the five species is given below.
A KEY TO CALAMAGROSTIS CANADENSIS, C. PERPLEXA, C. PORTERI,
C. RUBESCENS, AND C. SCRIBNERI
1. Collars glabrous or scabrous, not hairy or with a few scattered
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. canadensis
1. Collars hairy, the hairs short and numerous . . . . . . . . . . . . (2)
2. Plants from west of the 100th meridian . . . . . . . . . . . . (3)
3. Callus hairs 0.2?0.6 3 as long as the lemma, the hairs 0.7?
1.5 (?2.1) mm long; paleas (2.0?) 2.2?3.4 (?4.3) mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rubescens
3. Callus hairs 0.6?1.1 3 as long as the lemma, the hairs
(1.5?) 1.6?2.7 (?3.1) mm long; paleas (1.8?) 2.0?2.7
(?3.1) mm long . . . . . . . . . . . . . . . . . . C. scribneri
2. Plants from east of the 100th meridian . . . . . . . . . . . . (4)
4. Paleas (2.0?) 2.1?2.5 (?2.6) mm long; lemma awns (2.0?)
2.1?2.5 (?2.7) mm long; callus hairs 0.65?1.0 3 as long
as the lemma, the hairs (2.0?) 2.2?2.8 (?3.1) mm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. perplexa
4. Paleas (2.5?) 2.6?3.7 (?4.6) mm long; lemma awns (2.3?)
2.7?3.8 (?4.4) mm long; callus hairs 0.4?0.75 3 as long
as the lemma, the hairs (1.2?) 1.5?2.3 (?2.7) mm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. porteri
Calamagrostis perplexa Scribner, Circ. Div. Agrostol. U.S.D.A. 30:
7. 1901. Calamagrostis nemoralis Kearney, Bull. Div. Agrostol.
U.S.D.A. 11: 26. 1898. nom. illeg. hom. Calamagrostis porteri A.
Gray subsp. perplexa (Scribner) R.T. Clausen, Cornell Univ.
Agric. Exp. Sta. Mem. 291: 11. 1949. TYPE: U.S.A. New York: on
high ground, in woods of pine, rock-oak, hickory, etc. on
Thatcher?s Pinnacle, Danby, near Ithaca, 1 Aug 1884, W.R.
Dudley s.n. (HOLOTYPE: US-556697!; ISOTYPE: GH!).
A key to Calamagrostis in North America and a description of C.
perplexa are provided in Marr et al. (2007). Illustrations of C.
perplexa are provided in Marr et al. (2007, p. 727) and Holmgren
(1998, p. 717).
2009] Howard et al.?Calamagrostis perplexa 163
ADDITIONAL SPECIMENS EXAMINED: New York: Columbia Co., 2 mi. E of Hillsdale,
Mt. Fray, 1 Jul 1934, R. McVaugh 2649 (PH-916455); Tomkins Co., high hill W of
Danby, 1 Aug 1884, W.R. Dudley s.n.(GH); near Ithaca, 1885, W.R. Dudley s.n. (US-
556697; on same sheet as the holotype); Ithaca, 1889, W.R. Dudley s.n. (US-1867170);
Thatcher?s Pinnacle, 15 Aug 1917, K.M. Wiegand & R.N. Jones 7544 (GH); Pinnacle,
Danby, near Ithaca, Aug 1889, W.R. Dudley s.n. (US-844039); summit of Thatcher?s
Pinnacles, 20 Jul 1969, R.W. Simmers 1530 (GH-four sheets); Town of Danby, on the
shoulder and rim of the east side of the Cayuga Inlet Valley, Thatcher?s Pinnacles,
about 310 meters northwest from summit of most northern pinnacle, UTM: North
American Datum 1983, zone 18, 4,686,085N 375,742E, 12 August 2007, D. Werier
3243 (BH); Ulster Co., Town of Wawarsing, Shawangunk Ridge State Forest, from
Ellenville follow Route 52 E for about 4 mi. to the Shawangunk Ridge Trail crossing,
follow trail S for about 1.2 mi, UTM zone 18, 549284mE, 4611492mN, 1780 ft., 9
Aug 2006, T.G. Howard 465 (US); approximately 1.5?2 miles S of Hwy 52 on
Shawangunk Ridge trail, 41u39913.00N, 74u24929.30W, 542 m, 15 Aug 2007, P.M.
Peterson, J.M. Saarela & T.G. Howard 20925 (US, CAN); Town of Wawarsing,
Shawangunk Mountains, along ridge just north of Sullivan County line, southeast
about 2.5 km from the village of Spring Glen, UTM: North American Datum 1983,
zone 18, 4,611,490N 549,270E, 9 August 2007, D. Werier 3242 (BH).
ACKNOWLEDGMENTS. Field work by J.M.S. and P.M.P. was
supported by a grant from the Smithsonian Institution?s Restricted
Endowment Fund and the Department of Botany, and by the
Canadian Museum of Nature. Field work by T.G.H. was supported
by the New York State Biodiversity Research Institute and the
Eastern New York Chapter of The Nature Conservancy. Troy
Weldy, Tom Rawinski, Matthew Paul, and Ethan Pierce assisted in
the field; Steve Young helped with the initial determinations of
collections from the new site, and located the Columbia County
specimen at PH. The Smithsonian Institution?s Office of Research
Training and Services is thanked for providing a Short-term Visitor
grant to B.P. We are grateful to Ken Marr (V) for helpful discussion
and Thomas Wieboldt (VPI) for specimen loans. Christopher T.
Martine and an anonymous reviewer provided helpful comments
on an earlier version of the manuscript.
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APPENDIX
SPECIMENS USED IN PRINCIPAL COMPONENTS ANALYSIS OF CALAMAGROSTIS
CANADENSIS, C. PERPLEXA, C. PORTERI, C. RUBESCENS, AND C. SCRIBNERI
Calamagrostis canadensis
CANADA. E Coast, Hudson Bay, Great Whale River, 5 mi. from mouth,
M.T. Doutl 2580 (US-1648366); James Bay, S Twin Island, M.T. Doutl a (US-
1648369). Alberta: 6 mi. N of entrance to Waterton Lakes National Park, along
dry flood-bank of Dungarvan Creek in the Foothills Parkland, W.G. Dore
12151A (US-2076127); Pigeon Lake, E of Ma-Me-O Beach, G.H. Turner 4677
(US-1938744); Fort Saskatchewan, G.H. Turner 4607 (US-1938737); 1 mi. NE of
Fort Saskatchewan, G.H. Turner 4516 (US-1938719); Jasper National Park, on
the Banff?Jasper Hwy., 4 mi. N of Athabaska Falls, rocky NW shore of Leach
Lake, W.A. Weber 2504 (US-1849264); Jasper National Park, Horseshoe Lake
trail, just below lake, P.M. Peterson, J.M. Saarela & S.F. Smith 18434 (US-
3514300); Jasper National Park, 9 mi. NW of Bubbling Springs on Hwy 93
along Sunwapta River, P.M. Peterson, J.M. Saarela & S.F. Smith 18431 (US-
3514299); The Rocky Mountain Park, Lake Wapta, J. Macoun 64818 (US-
556041); The Rocky Mountain Park, above Laggan, J. Macoun 64817 (US-
747061); Rocky Mts., Elbow River, J. Macoun 18673 (US-844121); the Northern
Rocky Mts., vicinity of Banff, Bow Valley, W.C. McCalla 2305 (US-368531);
Jasper Park, Mt. Edith Cavell, J.M. Macoun 98153 (US-1060935); Jasper Park,
Cabin Lake, J.M. Macoun 98158 (US-1060930). British Columbia: the Canadian
Rockies, the vicinity of Field, Kicking Horse Valley, S. Brown 498 (US-1085728);
Moose Lake, N. Hollister 20 (US-622657); Lake House, Skagit River, J.M.
Macoun 77224 (US-844161). Manitoba: Riding Mountain National Park, Long
Lake, W.J. Cody & W.A. Wojtas 25075 (US-3081644); Knife Lake, 120 mi. SW
of Churchill, J.M. Gillett 2275 (US-1982642). New Brunswick: Shediac Cape,
F.T. Hubbard 722 (US-726648), 724 (US-726649). Newfoundland and Labrador:
Traverspine River, J.M. Gillet & W.I. Findlay 5622 (US-2206696); N Labrador,
Saglek Bay, R.A. Bartlett 53 (US-1366430); Bay of Islands, Dark Harbor, A.C.
166 Rhodora [Vol. 111
Waghorne 10 (US-844159); A.C. Waghorne 31 (US-556698); Bay of Islands,
Humber Arm, Silurian Hills back of Birchy Cove (Curling), M.L. Fernald &
K.M. Wiegand 2540 (US-1024241); Valley of Exploits River, Grand Falls, M.L.
Fernald & K.M. Wiegand 4569 (US-1024240); Humber District, Grand Lake,
Blue Grass Brook Point, L.E. Ruoleau 2221 (US-2119734); Humber District,
Grand Lake, Hinds Brook Point, L.E. Ruoleau 2274 (US-2119737); St. Barbe
District, Bonne Bay, White Point, L.E. Ruoleau 425 (US-2119706); Port-aux-
Basques, A.S. Hitchcock 23983 (US-1388296); Grand Falls, A.S. Hitchcock
23652 (US-1388294). Northwest Territories: Fort Simpson, W.J. Cody & J.M.
Matte 8784 (US-2523928); Norman Wells, sedge-grass border of lake, W.J. Cody
& R.L. Gutteridge 7876 (US-2306101); Mackenzie District, Carcajou River, Mile
23E Canol Road, W.J. Cody & R.L. Gutteridge 7776 (US-2306091); Mackenzie
District, Yellowknife, Joliffe Island, W.J. Cody & J.B. McCanse 3110 (US-
2238582); The Region of Great Bear Lake, Head of Hornby Bay, N of the
Arctic Circle, NE end of the McTavish Arm, Great Bear Lake, H.T. Shacklette
3206 (US-2010737); Mackenzie Basin, the Wood Buffalo Park, Moose (Eight)
Lake district, Ranger Station, H.M. Raup 1752 (US-1722305), 1764 (US-
1722301); Mackenzie Basin, the Wood Buffalo Park, E shore of Lake Mamavi,
H.M. Raup 1756 (US-1722319), 1757 (US-1722304); Mackenzie District, Yellow-
knife, N of old townsite, W.J. Cody & J.B. McCanse 3022 (US-2238578). Nova
Scotia: Yarmouth Co., Yarmouth, M.L. Fernald & B. Long 19927 (US-1100087);
St. Paul Island, bog above Petrie?s Pond, L.M. Perry & M.V. Roscoe 56 (US-
1502774); Canso, J. Fowler 12 (US-844118). Ontario: Thunder Bay District,
H.E.P.C. right of way, 1 mi. E of Port Arthur, McGregor Twp., C.E. Garton
1232 (US-2075985); Wellington Co., Morristown, W.G. Dore & L.V. Bush 46-66
(US-2013398), 46-67 (US-2013399); Wellington Co., E of Morristown, Puslinch
Twp., W.G. Dore & L.V. Bush 46-60 (US-2013393); W of Kitigan, O?Brien Twp.,
C.V. Morton & E. LePage 11560 (US-2306542); Kingston, J. Fowler s.n. (US-
260837); Bruce Co., Pine Tree Point, Lake Huron, W.G. Dore 8941 (US-
1982563); The Georgian Bay Islands opposite Shawanaga Twp., Parry Sound
District, Hemlock Island (500A), E.D. McDonald Jr. 21 (US-2013764); Galt,
Herriot s.n. (US-844140); Island 74, French River, L.H. Dewey 9 (US-1814610);
Ottawa, Rideau Canal, W.G. Dore 130 (US-1648380). Prince Edward Island:
Queens Co., Hillsborough River, Mt. Stewart, M.L. Fernald, E.B. Bartram, B.
Long & H.S. John 6857 (US-1024088). Que?bec: Du Golfe Saint-Laurent,
Archipel de Mingan, Ile aux Calculeaux, Fr. Marie-Victorin & Fr. Rolland-
Germain 20541 (US-1295069); Du Bic, Comte? de Rimouski, Bic, J. Rousseau
26693 (US-1435584);Vicinity of Maniwaki, Fr. Rolland 7134 (US-1011279);
Grand Etang, along the Gaspe? Coast, Macoun 26 (US-844119); Bonaventure
Co., Little Cascapedia River, J.F. Collins & M.L. Fernald 21 (US-493639); Gaspe?
Co., Table-top Mt., M.L. Fernald & J.F. Collins 161 (US-605991); Iles de la
Madeleine, Ile du Ha?vre-aux-Maisons, Fr. Marie-Victorin & Fr. Rolland-
Germain 9032 (US-1061039, US-1116498); Kamouraska Co., Ste-Anne-de-la-
Pocatie`re, W.G. Dore 364 (US-1648381); Gaspe? Co., Mt. Albert, W.G. Dore 47-
791 (US-1935424); La Gaspe?sie, Riviere Ste-Anne des Monts, Fr. Marie-
Victorin, Fr. Rolland-Germain, J.B. Brunel & Z. Rousseau 17800 (US-1024029);
La Cote-Nord, Du Golfe Saint-Laurent, Sept Iles, Fr. Marie-Victorin & Fr.
Rolland-Germain 18206 (US-1295175); Gaspe? Co., Table-topped Mt., Head of
White Rock Slide, M.L. Fernald & J.F. Collins 377 (US-1329627); 9 mi. W of
2009] Howard et al.?Calamagrostis perplexa 167
Wakefield, F.E. Carues 81 (US-1723490); Upper slopes of Mt. Albert, J.R.
Swallen 9746 (US-1935130), 9776 (US-1935147); Rupert River and Mistassini
Lake, Northern Tesekan Lake, A. Ditilly & E. Lepage 11369 (US-2240396);
Megantic Co., Caribou Hill, Blake Lake, M.L. Fernald & H.B. Jackson 12012
(US-2011350); Fort Chimo area, J.A. Calder 2713 (US-1982637). Saskatchewan:
Lake Athabaska, small island, F. Harper 63 (US-1085365); Along the line of the
Grand Trunk Pacific Railway, Round Valley Lake, J. Macoun & W. Herriot
44152 (US-844151); Near Prince Albert, J. Macoun 13091 (US-844117); Prince
Albert, J. Macoun 13091 (US-844141). Yukon: Dawson, below summit of Dome,
along Ry, A.S. Hitchcock 4407 (US-844060); About 3/4 of a mile from NE of
Mayo, J.A. Calder & J.M. Gillet 4135 (US-2303142); Along Yukon River
between Whitehorse and Dawson, about 50 mi. below Hootalingua, A.S.
Hitchcock 4330 (US-844059).
UNITED STATES. Alaska: Dillingham, I.N. Gabrielson s.n. (US-2479007);
Katmai National Monument, Rock cliff, S side of island, Coville Lake, 2 mi.
NW from Coville River, V.H. Cahalane 752 (US-2205042); Prince William Sd.,
Hinchinbrook J., I.L. Norberg s.n. (US-2238656); Alaska Peninsula, King Cove,
W.J. Eyerdam 1564 (US-2238649); Fairbanks, Chatinika, A.S. Hitchcock 4598
(US-844096); Hot Springs on the Tanana River, A.S. Hitchcock 4520 (US-
844104); The Brooks Range, Mts. NW of Walker Lake, upper Kopuk River,
L.H. Jordal 3982 (US-2040880); Koyukuk R., Middle Fork, Bettles, E.O.G.
Hulte?n s.n. (US-2382194); Skagway, J.M. Gillett & D.A. Mitchell 3697 (US-
2303163); Mosse Pass Valley, Kenai Peninsula, J.A. Calder 6378 (US-2303169);
Lower Yerrick Creek, L.A. Spetzman 797 (US-3465309); Along the Yukon
River, A.S. Hitchcock 4690 (US-844068); Along the Yukon River, Holy Cross,
A.S. Hitchcock 4680 (US-844066), 4683 (US-844067); Nome, sandy hill, half mi.
of beach, A.S. Hitchcock 4761 (US-844070); Nome, along a dike, A.S. Hitchcock
4777 (US-844072); Yukon Valley, Yukon Flats, A.J. Collier 107 (US-379793).
Yukon Valley, White River, M.W. Gorman 1149 (US-377202).
Calamagrostis perplexa
UNITED STATES. New York: Columbia Co., 2 mi. E. Hillsdale, R. McVaugh
2649 (PH-916455); Tompkins Co., Thatcher?s Pinnacle, W Danby, near Ithaca,
W.R. Dudley s.n. (US-556697, 32); Ithaca, W.R. Dudley s.n. (US-1867170, 32);
Thatcher?s Pinnacle, Danby, Ithaca, W.R. Dudley s.n. (US-844039, 32); Danby
Twp., summit of Thatcher?s Pinnacle, R.W. Simmers Jr. 1530 (GH-244392; GH-
244393; GH-247516, 32; GH-247517), High Hill, W Danby, W.R. Dudley s.n.
(GH-261464); Thatcher?s Pinnacle, Danby, K.M. Wiegand & R.N. Jones 7544
(GH-247515); Ulster Co., Town of Wawarsing, Shawangunk Ridge State Forest,
T.G. Howard 465 (US-3483671); 1.5?2 mi. S of Hwy 52 on Shawangunk Ridge
Trail, P.M. Peterson, J.M. Saarela & T.G. Howard 20925 (CAN, US-35276461).
Calamagrostis porteri
UNITED STATES. E.S. Steele & Mrs. Steele s.n. (US-490318). New York:
Chemung Co., W side of N Peak of Maby Hill, S.J. Smith 7854 (US-2078851).
Ohio: Jackson Co., Liberty Township, F. Bartey & L. Pontius 21 (US-1645016);
Vinton Co., Vinton Furnace Experimental Forest, F. Bartley 2315 (US-2238723).
168 Rhodora [Vol. 111
Pennsylvania: unknown collector (US-556710); Bedford Co., 4 3/4 mi. NNE of
Chaneysville, D. Berkheimer 3969 (US-2116347); Centre Co., 4 mi WNW of
Bellefonte, H.A. Wahl 1538 (US-2012430); Huntingdon Co., Pulpit Rocks, T.C.
Porter s.n. (US-135195), s.n. (US-556709); T.C. Porter 440 (US-821585); Warriors
Mask, T.C. Porter s.n. (US-78269). Virginia: Blue Ridge near Luray, Stony Man
Mt., N. Hotchkiss 1612 (US-1259986); 5 mi. W of Crescent Ridge, Skyline Drive,
L.E. Hicks & F. Bartley 2168 (US-2184909); Stony Man Mountain and Vicinity
in the Blue Ridge near Luray, E.S. Steele & Mrs. Steele 103 (US-418453);
Amherst Co., Blue Ridge Mts., Punchbowl Mt., C.E. Stevens 1412 (US-
2619848); Augusta Co., Blue Ridge Mts., Turk Gap, C.E. Stevens 1429 (US-
2619849); Bath Co., vicinity of Millboro, in the Allegheny Mts., E.S. Steele s.n.
(US-726956), s.n. (US-643309); Giles Co., Salt Pond Mt., H.H. Iltis & L. Carr
1998 (US-1867935); Page Co., Luray, Blue Ridge Mts., along Appalachian Trail,
near Skyline, F.J. Herman 11401 (US-1866099); ibid., near Skyline, 7 mi. SE, F.J.
Hermann 11406 (US-2478624); Shenandoah National Park, Appalachian Trail to
Hawksbill Mt., R.C.Y. Chou 524 (US-2012488); Blue Ridge Mts., Mt. Marshall,
Skyline Drive, H.A. Allard 754 (US-1678986); Rockingham Co., Second Mt.,
C.E. Stevens 1283 (US-2619846); Shenandoah Co., Massanutten Range, Three
Top Mt., H.A. Allard 5615 (US-1787766). West Virginia: Hardy Co., Sugar
Knob, H.A. Allard 9399a (US-1872753); Pendleton Co., near Reddish Knob,
C.E. Stevens 1255 (US-2619847).
Calamagrostis rubescens
UNITED STATES. California: Mariposa, J.W. Congdon s.n. (US-844036); Mts.
about the head waters of the Sacramento River, C.G. Pringle 73 (US-821595),
254 (US-821594); Plains of Mendocino, C.G. Pringle s.n. (US-1912084), 59 (US-
723033); Santa Cruz Co., vicinity of Gazos Creek Road, J.H. Thomas 3539 (US-
2973804); Santa Cruz Portland Cement Company quarry, V.F. Hesse 1667 (US-
2973787); Santa Cruz, D. Anderson s.n. (US-844033). Colorado: N Colorado,
Honnold, F. Tweedy 4359 (US-432480); Near Hahns Peak, Wooton s.n. (US-
1502661). Nevada: Mountain City (Mill Creek), A. Nelson & J.F. McBride 2212
(US-544962). Oregon: Powder River Mts., C.V. Piper 2526 (US-843979); Canon
City, Blue Mts., D. Griffiths & B. Hunter 164a (US-844006); 9 mi. above
Wallowa Lake, C.L. Shear 1795 (US-844007); E.P. Sheldon 8637 (US-844009);
Portland, L.F. Henderson 2150 (US-844012); Hidaway Springs, Blue Mts., W.C.
Cusick 3486 (US-844015); Portland, Hd. of 25 Mile Creek, M.W. Gorman 599
(US-844017); Kerby to Tennessee Pass, Siskiyou Forest, A.S. Hitchcock 23546
(US-1299097); Meacham, M.E. Peck & J.C. Nelson 5672 (US-836541).
Washington: E Washington, Stehekin, D. Griffiths & J.S. Cotton 240 (US-
844880); L.F. Henderson 2151 (US-843989); Pullman, C.V. Piper 1919 (US-
230304); Wenatchee Valley, Sandberg & Leiberg 535 (US-843987, US-843995);
Near Bingen, Columbia River, W.N. Suksdorf 2825 (US-843991); SE Washing-
ton, Blue Mts., R.M. Horner 537 (US-843993); W Klickitat Co., Bingen, W.N.
Suksdorf 2825 (US-529618); Spokane Co., E of Rockford, J.R. Swallen 6241 (US-
2303657). Wyoming: Snowy Range, J.R. Swallen 6598 (US-2303982, US-
2434071); Grand Teton National Park, Cottonwood Creek outlet to Jenny
Lake, near park boundary, H. Bailey & V. Bailey 4161 (US-2115969); Teton
National Park, E shore of Jenny Lake, A.A. Beetle 5004 (US-2152283); Jackson
2009] Howard et al.?Calamagrostis perplexa 169
Hole, near Menors Ferry, E.D. Merrill & E.N. Wilcox 362 (US-843958); Jackson
Hole, Mosquito Creek, A.S. Hitchcock 23124 (US-1296308); Jackson, A.S.
Hitchcock 23113 (US-1296317); Yellowstone National Park, near Levis River,
E.D. Merrill 118 (US-844602); Yellowstone National Park, Mammoth Hot
Springs, E.A. Mears 2620 (US-487219); Yellowstone National Park, Mammoth
Hot Springs, road to Yancey, A.S. Hitchcock 1976 (US-844055); Yellowstone
National Park, hillside near Yancey, A.S. Hitchcock 2025 (US-843953), 2026 (US-
838801); Yellowstone National Park, near Old Faithul, A.A. Beetle 5109 (US-
2152286); Yellowstone National Park, H.L. Bolley 1897 (US-843955, US-
2150994); Yellowstone National Park, Firehole R., A.S. Hitchcock 2128 (US-
843956); Yellowstone National Park, Madison Canyon, on the bank of
Madison River, A. Nelson & E. Nelson 6760 (US-365691); Wyoming National
Forest, Grey?s River, Schofield & Woods 226 (US-1160118); Teton Co., Elbow,
Jackson Hole, Teton Mts., A.A. Beetle 5298 (US-2152298); Uinta Co., Teton
Pass, E.D. Merrill 77 (US-843888).
Calamagrostis scribneri
UNITED STATES. Idaho: Coeur d?Allene Lake, J.H. Sanberg 564 (US-844050).
Montana: Spanish Creek, T.A. Williams 2009 (US-844047); Spanish Creek, P.A.
Rydberg 3083 (US-844045), 3100 (US-844048); Belt Park, F.A. Spragg 743 (US-
844770); Elk Creek, F.L. Scribner 365 (US-135193); Dry Fork of Belt Park, P.A.
Rydberg 3346 (US-843949); F.H. Knowlton s.n. (US-844049); Flathead National
Forest, White River near falls, J.E. Kirkwood 2267 (US-1295489); Gallatin Co.,
Bridger Pass, W.N. Suksdorf 63 (US-1061911); Park Co., E of Bridger Range,
J.R. Swallen 6402 (US-2303793). Washington: Pullman, C.V. Piper s.n. (US-
843893); Mount Paddo (Adams), hillsides, W.N. Suksdorf 145 (US-844386);
Mount Paddo (Adams), W.N. Suksdorf 145 (US-844388); Kittitas Co.,
Wenatchee Mt., J.S. Cotton 1756 (US-556018); Skamania Co., Chiquash Mts.,
W.N. Suksdorf 1023 (US-821560, US-844387); High Mts., W.N. Suksdorf 1023
(US-844389). Wyoming: 14 mi. post, S from Mammoth Hot Springs, E.A.
Mearns 3718 (US-487823); Jackson?s Lake, A. Chelson 6563a (US-556705);
Yellowstone Park, G.W. Letterman 44 & 45 (US); Yellowstone National Park,
Tower Falls, E.A. Mearns 3547 (US-487723); Yellowstone Park, Slough Creek,
F. Tweedy 585 (US-556704); Albany Co., Woods Landing, Head of Wood?s
Creek, A. Nelson 3954 (US-844335); Wood?s Creek, A. Nelson 3950 (US-358953).
170 Rhodora [Vol. 111