JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

January 1984 No. 845

Vol. 86

Che Neu Lngland Botanical Club, Ine.

Botanical Museum, Oxford Street, Cambridge, Massachusetts 02138

Conducted and published for the Club, by
NORTON H. NICKERSON, Editor-in-Chief

Associate Editors

A. LINN BOGLE GARRETT E. CROW
WILLIAM D. COUNTRYMAN RICHARD A. FRALICK
GERALD J. GASTONY NORTON G. MILLER

ROBERT T. WILCE

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Cover illustration

Ledum groenlandicum Oeder, Labrador tea, reaches its southeastern distribu-
tional limit in Concord, Mass. It was first collected by Thoreau in 1858, subsequently
regarded as extirpated by Richard Eaton in 1974, and rediscovered by Ray Angelo in
1978. Angelo has since found it in two more Concord locations.

Original artwork by Josephine Ewing.

TRhodoera

(ISSN 0035 4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 86 January 1984 No. 845

THE TYPE LOCALITIES OF THE FLORA
BOREALI-AMERICANA OF ANDRE MICHAUX

LEONARD J. UTTAL

ABSTRACT

Many habitats of the text of the Flora boreali-americana of A. Michaux (1803),
which is presumed to have been anonymously authored by L. C. M. Richard, have
been observed to vary, sometimes drastically, from the field stations on the labels of
the type specimens collected by Michaux. Since the field stations are the actual type
localities rather then those given in the text, it is important to record them to
promote precision in the use of the Michaux flora. A key to the IDC 6211 microfiche
set of the Michaux American specimens, which, are in the Paris Museum of Natural
History, has been devised because the microfiches were found awkward to use. Each
non-superfluous name proposed by Michaux is listed and its microfiche key
designation is equivalent to an annotation of the specimen as the type.

Key Words: Michaux flora, type localities, field/ text variances

INTRODUCTION

In the course of work I pursued in Senecio L. it was observed that
Barkley (1978) had not seen the type of S. romentosus and its type
locality was given as “in Carolinae loco dicto Flat-Roc.”, after the
text of Michaux (1803). The gazetteer prepared by Ewan (1974) for
the introduction to a facsimile edition of Flora boreali-americana
gives Flat Rock as near Hendersonville, Henderson County, North
Carolina, and cites it as the type locality of Sedum pusillum Michx.
and Senecio tomentosus. Knowing the montane location of Flat
Rock, North Carolina well, and the fact that these lowland species
do not occur there, this struck me as incongruous. Neither species
was dotted in distribution maps in Radford et al. (1968) for

2 Rhodora [Vol. 86

Henderson County, North Carolina. I then looked up the two types
in the IDC microfiche set of Michaux’s American specimens and
found they both indeed had Flat Rock on the labels. I then read
Michaux’s journal, as published by Sargent (1889), and found that
Michaux had never passed through Flat Rock, North Carolina, but
on 23 April 1795 and 6 April 1796 he passed Flat Rock, South
Carolina, en route northwest from Charleston, South Carolina. |
subsequently learned that this Flat Rock is a granite boss in
northern Kershaw County, South Carolina, now a quarry, on state
highway 58, the old Catawba Road, well within the range of both
species, which I found growing in the vicinity, but not at the type
locality due to habitat destruction. Flat Rock, South Carolina is the
correct type locality of these species, and the Flat Rock of North
Carolina was listed in error. The state of anthesis of the specimens
indicates that the 6 April 1796 visit was the date of collection.

In addition to the above incongruity, many examples of variation
between habitat data given on Michaux’s labels and that given in the
text of Flora boreali-americana became evident. For example, the
habitat of Vaccinium brachycerum Michx. (= Gaylussacia brachy-
cera (Michx.) A. Gray) is given in the text as Virginia, circa
Winchester, but the habitat on the label reads “Warm Spring”.
There are numerous “Warm Springs” in the Winchester area, but
Warm Spring is used in the singular on old maps only for Warm
Spring Road, present-day US 252, which passes east and west
through Winchester. Michaux passed through Winchester several
times and could well have collected at this station. Repeated
searches for the species in this region have been fruitless, It is
obvious from Michaux’s journal that he neither passed through
present Warm Springs, Bath County, Virginia, where the species
does now grow, nor old Warm Springs of Berkeley County, now
Berkeley Springs, West Virginia. The Winchester type locality of the
species remains enigmatic.

Francois-André Michaux, an eminent botanist in his own right,
who wrote the introduction to the Flora credited to his father! states
that each specimen was accompanied by a sheet of information “too
prolix” for publication. These sheets, presumably in the Paris
Museum, are not presently available to me, but a study of them

'The Flora is believed to have been anonymously authored by L. C. M. Richard - see
Stafleu and Cowan (1982),

1984] Uttal — Michaux localities 3

might further add to the precision of the type localities of Michaux’s
North American collections. This study is based on a study of
photographs of the specimens and their labels of IDC microfiche set
6211.

To use this present delineation of type localities it is necessary to
slightly augment the gazetteer prepared by Ewan (1974) of
Michaux’s collecting stations. Additional place names which appear
on Michaux’s labels which are absent from the Flora are as follows:

Florida

Northwest River; Midwest River; those portions of the St. John’s
River geographically relative to St. Augustine, the Spanish
capital of Florida, in St. John’s and Clay Counties.

Tomoka River; in northeast Volusia county, north of Ormond
Beach.

Georgia
Sunbury; in Long County, on south bank of mouth of Canoochee
River, circa 10 miles south of Savannah.

Illinois

Fort Massac; Massac County, on Ohio River between Metropolis
and Brockport, now a state park.

“Kaokia”; = Cahokia, St. Clair County.

Prairie du Rochier; Randolph County, a bluff on the Mississippi
River, 21 miles northwest of Chester.

Kentucky

Big Barren River; Barren Oaks, Barren County.

“Piercetown”, apparently transliterated from Beardstown in
Michaux’s journal, presumably now Bardstown, Nelson County.

New York

Lasinburgh; = Lansingburg, Renssalaer County, now incorporated
in Troy.

“Mountains near Lasinburg”; presumably a range of hills 10 miles
east of Troy dominated by Rock Candy Mountain.

North Carolina

Davenport; an old rest stop at foot of Yellow Mountain, near
Minneapolis, Avery County.

Fayetteville, Cumberland County.

Hill’s Iron Works; actually in South Carolina on east bank of

4 Rhodora [Vol. 86

Catawba River in York County, 32 miles north of Cane Creek
(Lancaster County), according to Michaux’s journal. Collecting
done in North Carolina, across the river from this reference point,
presumably was in Union County, near Waxhaw.

South Carolina

Fields of Caffet; not identified, believed to be in vicinity of
Charleston.

Gaillard Road; present State Road 357, in Berkeley County, 7 miles
southeast of Moncks Corner.

Garet Place; not identified, but believed to be in vicinity of
Charleston.

Flat Rock; not in Henderson County, North Carolina, but in north
Kershaw County, South Carolina, presently a quarry on State
Road 58, 0.4 mile south of Lancaster County boundary.

Goose Creek; Berkeley County.

Nine Mile Brook (Ruisseau de Neuf Milles); unidentified but
believed to be in vicinity of Charleston.2

Two Sisters: a ferry on the Savannah River north of the city of
Savannah, with the Carolina dock in Beaufort County.
Virginia

Warm Spring; believed to refer to the Warm Spring Road (present
US 252) running west out of Winchester in Frederick County.

Current taxonomic names used in this study for Michauxian
equivalents represent, in the author’s judgment, prevailing names in
current manuals; not necessarily the most recently published
combinations, but names which will be familiar to most botanists.
Rarely a name recently published, not yet in manuals but widely
accepted, is used. If an asterisk (*) follows the name it means an
alternate name is present in the National List of Scientific Plant
Names (1982). In any case, botanists can derive names they may
prefer from names I use. Manuals particularly useful in this study
are Cronquist (1980), Gleason and Cronquist (1963), Godfrey and
Wooten (1979, 1981), and Radford, Ahles, and Bell (1968).
Reference to several botanically historical works was frequent, and
reference to geography and history was indispensable. Jndex
Kewensis was virtually constantly used.

>Michaux’s plantation was at Ten Mile Station (10 miles north of old Charleston).

1984] Uttal — Michaux localities 5

In the course of this study, the microfiche set of Michaux’s North
American collections (IDC 6211) was found awkward to use. It
frequently deviated from the sequence of species in the Flora and
lacked an index so that the user often had to resort to trial and error
in locating a particular photograph of a specimen. To correct this
situation a key to the type specimens photographed for the
microfiche was devised which is very simple to use. There are
twenty-one photographs on each microfiche film card, reading from
left to right, and each film is numbered. Thus the microfiche
photograph of Senecio Iyratus Michx. is 99:13, 99 being the
microfiche card number, 13 the particular specimen. This key
appears in the upper right of each entry which follows and each such
designation is equivalent to the selection of the specimen photo-
graphed as the holotype, if single, or lectotype, if one of a series.

Michaux’s names and handwritten data are presented sic.
Superfluous names are omitted. In general, the order of species
follows the Flora text, but it is occasionally desirable to deviate
from this sequence to keep the sequence of the microfiches in
approximate order. In any case, the genus can be located in the
Flora index. The few non-vascular types in the Michaux Flora were
not considered practical to treat in this paper. There are 747 entries
of types which follow.

EXAMPLES OF FORMAT OF VARIOUS ENTRIES
(Volume and page number in Flora) (Microfiche key number)
2: 120 Senecio lyratus Michx. 99: 13

T: in nemoribus Carolinae (Habitat in Flora text).

M: pastures arides pres Charlest. Printems. (Habitat on Mi-
chaux’s specimen label, the actual type locality). Senecio
glabellus Poir. (Prevailing present name. If asterisk fol-
lows, = alternative name in National List of Scientific
Plant Names.)

In case the Michaux name is the presently accepted name and
there are no locality data on the label, or the label data are exactly
the same as the text data, the taxon will be listed (1) to give its
microfiche key number and (2) to indicate that the specimen
photographed is annotated as the type. The text habitat is accepted
for the present. Format:

2: 117. Solidago glomerata Michx. 99: 12

6 Rhodora [Vol. 86

Where the locality data situation is the same as above, but the
present name is different than the one proposed by Michaux, the
format will be the same except to show the prevailing current name,
L.;:

2: 121 Doronicum nudicaule Michx. 99: 21

Arnica acaulis (Walt) BSP

In a few cases the microfiche picture of a type was not located. In
such an entry, the microfiche key space is left blank and the Flora
text data are accepted as diagnostic.

ACKNOWLEDGMENTS

The gracious abundant counsel throughout the course of this
study by Duncan Porter is gratefully acknowledged.

This study could not have been completed without the facilities
and assistances of the Newman Library of Virginia Polytechnic
Institute and State University, generously proffered and admin-
istered.

Financial assistance toward publication was provided by the
Virginia Flora Committee.

ENTRIES OF TYPES
MICHAUX VOL. I

1:2. Salicornia ambigua Michx.
Salicornia virginica L.

1:6 Gratiola quadridentata Michx. 1:4
Gratiola ramosa Walt.

1:7 Gratiola anagallidea Michx. 133
T: in humidus Carolinae.
M: in umbrosus humidis Carolinae. Basse Carolina.

Lindernia anagallidea (Michx.) Pennell

1:7 Gratiola pilosa Michx. 2:2

1:8) Justicia humilis Michx. 2:9
Justicia ovata var. lanceolata (Chapm.) R. W. Long

1:7 Justicia pedunculosa Michx. a7
T: in littoribus hieme inundatis fluminum Ohio, S. Laurentii, Mississippi.
M: Ohio, Fl. S. Laurent.

Justicia americana (L.) Vahl

1984]

Uttal — Michaux localities

Elytraria virgata Michx.

in humidis Carolinae inferioris.

Carolina

Elytraria caroliniensis (J. F. Gmel.) Pers. var.
caroliniensis

Micranthemum orbiculatum Michx.

Micranthemum umbrosum Walt. in J. F. Gmel.

Pinguicula elatior Michx.
Pinguicula caerulea Walt.

Pinguicula acutifolia Michx.
Pinguicula villosa L.

Pinguicula pumila Michx.

Utricularia ceratophylla Michx.
Utricularia inflata Walt.

Utriculata cornuta Michx.

Utriculata setacea Michx.
Urriculata gibba L.

Cunila glabella Michx.
Satureja glabella (Michx.) Briquet var. glabella

Lycopus uniflorus Michx.
ad Lacus S-Joannis et Mistassins.
Lac St. Jean au Lac Mistassins.

Salvia angustifolia Michx.
Salvia azurca Lam.

Monarda allophylla Michx.
Monarda fistulosa L. var. fistulosa

Collinsonia tuberosa Michx.
Valeriana pauciflora Michx.

Elodea canadensis Michx.
in rivulis Canadae.
env. de Montreal.

Hertieria Gmelini Michx.
Lachnanthes caroliniana (Lam.) Dandy

Xyris brevifolia Michx.
in pratensibus udis Georgiae inferioris.
Georgia.

Commelina longifolia Michx.
Commelina virginica L.

3:3

—

ae |

32

Rhodora

Ciperus Ayvdra Michx.

in cultis Virginiae, Carolinae, Floridae.
in Virginia maritima Carolina Georgia.
Cyperus flavicornus Michx.

Cyperus strigosus L.

Ciperus virens Michx.

Avllingia pumila Michx.

Cyperus tenuifolius (Steud.) Dandy

Kyllingia maculata Michx.
Lipocarpha maculata (Michx.) Torr.

Avllingia ovularis Michx.
in Georgia et Carolina.

Jardin Basse Carol. (Charleston? - auth.)
Cyperus ovularis (Michx.) Torr. var. ovularis

Scirpus quadrangulatus Michx.
in Carolina

in Marais de la Caroline. Fleurit in May et Juin, 1794.

[Vol. 86

7:3

|

7:11

8:2

Eleocharis quadrangulata (Michx.) R. & S. var. quadrangulata

Scirpus tuberculosus Michx.
in Carolina inferiore.
in herbosis humidis Carolina inferioris.

Eleocharis tuberculosa (Michx.) R. & S.

Scirpus capillaceus Michx.

in nova Anglia.

Connecticut.

Eleocharis acicularis (L.) R. & S.

Scirpus puberulus Michx.

in Carolina et Georgia.

in Georgia.

Fimbristyvlis dichotoma (L.) Vahl

Scirpus castaneus Michx.
in Florida.

8:3

8:5

in Carolina, juxta Tomoka creek (= Tomoka River, Volusia County,

Florida—auth.)
Fimbristylis castanea (Michx.) Vahl

Scirpus mucronulatus Michx.

in montibus Carolinae.

haute et basse Caroline.

Fimbristylis autumnalis (L.) R. & S.

Scirpus maritimus var. macrostachyus Michx.

Scirpus robustus Pursh

8:21

Uttal — Michaux localities

Scirpus lineatus Michx.

Eriophorum hudsonianum Michx.

a sinu Hudsonis ad lacus Mistassins.
Lacs Mistassinis.

Eriophorum alpinum L.

Schoenus sparsus Michx.

in umbrosis syslvanum Carolinae, Georgiae.
Basse Carolina férets ombrages.
Rhynchospora miliacea (Lam.) A. Gray

Schoenus longirostirs Michx.

in Virginia et Carolina.

haute et basse Carolina.

Rhynchospora corniculata (Lam.) A. Gray

Schoenus inexpansus Michx.
Rhynchospora inexpansa (Michx.) Vahl

Schoenus rariflorus Michx.
Rhynchospora rariflora (Michx.) Ell.

Schoenus capitellatus Michx.
Rhynchospora capitellata (Michx.) Vahl.

Schoenus ciliaris Michx.
in Florida

in Florida, juxta Tomoka Creek (= Tomoka River, Volusia

County - auth.)
Rhynchospora ciliaris (Michx.) Mohr

Schoenus distans Michx.

in Carolina.

in Carolina, Georgia.

Rhynchospora glomerata (L.) Vahl var. glomerata

Schoenus fascicularis Michx.
Rhynchospora fascicularis (Michx.) Vahl

Dichromena leucocephala Michx.
in Carolina et Georgia.

Carolina, 1794

Dichromena latifolia Baldw. ex. Ell.

Fuirena squarrosa Michx.
in paludosis Georgiae et Carolinae
Basse Carol et Jardin (Charleston - auth.)

Fuirena scirpoidea Michx.
in paludosis aestate exsiccabilibus Floridae.

Le 11 Avril (1787 - auth.) vers le haut de Tomoko riv.

8:8
9:4

9:18

9:21

9:5

9:10

9:7

9:9

10:3

10:6

Rhodora [Vol. 86

Leersia lenticularis Michx. 10:12
in paludosis Illinoensibus.
in paludosis regionis Illinoensium.

Dilepyrum aristosum Michx. 10:15
in umbrosissylvis Georgiae et Carolinae.

in umbrosis Carolinae sylvis

Brachyelytrum erectum (Shreb.) Beauv. var. erectum

Dilepyrum minutiflorum Michx. 10:14
in apricis, pratensibus regionum Kentucky et Illinoensium.

in habitum praestantissinoens ad in Kentucky, Illinoensium.
Muhlengergia schreberi J. F. Gmel.

Aristida oligantha Michx. 10:19
in vastissimus pratensibus Illinoensibus.

Route des Illinois au fort Massac lieux alternativment submerger
parles Pluy.

Aristida dichotoma Michx. 10:21
in Carolina superiore, juxta Lincoln in glareosis.
in Carolina septentrionali juxta Lincoln.

Aristida stricta Michx, 11:2
in Carolina inferiore.
in Carolina.

Trichodium laxiflorum Michx. 11:4
in humidis et pratensibus a sinu Hudsonis ad Floridam.

Pensylvanie.

Agrostis hyemalis (Walt.) BSP

Trichodium decumbens Michx. 11:14
a Virginia maritima ad Floridam.

in Carolina, praesertim in umbrosis ripariis Amnium.

Agrostis perennans (Walt.) Tuckerm. var. perennans

Digitaria paspalodes Michx. 11:15
In pascuis aridis, juxta Charlston.

in pascuis aridis Carolinae.

Paspalum distichum L. var. distichum

Digitaria pilosa Michx. 11:16
in sabulosis sylvarum Carolinae, Georgiae et alibe.

in sabulosis Carolina, Georgia.

Digitaria filiformis var. villosa (Walt.) Fern.

Panicum melicarium Michx. 11:19
in Carolina, Georgia.

in Carolina ad ripas rivorum affluviente mari inundatum.
(incongruous; the species occurs in North Carolina in the

piedmont and in montane areas - auth.)

Glyceria melicaria (Michx.) F. T. Hubb.

Uttal — Michaux localities 11

Panicum anceps Michx. 12:7
in Carolinae herbosis humidis sylvaticus.

in herbosis humidis Carolinae, Virginiae, Georgiae.

Panicum anceps Michx. var. anceps

Panicum dichotomiflorum Michx. 12:8
in occidentalibus montium Alleghanis.

in regione Illinoensium.

Panicum dichotomiflorum Michx. var. dichotomiflorum

Panicum molle Michx. 12:11
in cespitosis Floridae

Lieux tres humides a 15 miles de St. Augustin (believed

to be to the south - auth.)

Frichloa michauxii (Poir.) Hitche. var. michauxii

Panicum ramulosum Michx. 12:13
in sylvis Carolinae.

in pratis, cespitosus Carolina.

Panicum angustifolium Ell.* (The basionym is questioned by
Hitchcock and Chase (1950)).

Oryzopsis asperifolia Michx. 12:20
a sinu Hudsonis ad Quebec, per tractus montium.
in praeriptus et saxosus per tractus montium a sinu hudsonis ad Canadam.

Paspalum ciliatifolium Michx.* 12:21
in Carolina.
in Carolina, Georgia.

Paspalum debile Michx.* 13:1
in maritimis Carolinae et Georgiae.
in Carolina.

Paspalum setaceum Michx. 13:2
in aridis Carolina inferioris.

in aridis apricis Carolina, Georgia. Champ a Caffet, Juiliet.

Paspalum setaceum Michx. var. setaceum

Paspalum plicatulum Michx. 13:3
Paspalum laeve Michx.* 13:4
Paspalum floridanum Michx.* 13:6

in Florida et Georgia.
Georgie et Floride.

Agrostis lateriflora Michx. 13:8
praesertim in praecipitus saxosus fluminis Mississippi et ripariis
Illinoensibus.

(same except the “et” is omitted, meaning the Mississippi shores

of Illinois - auth.)

Rhodora [Vol. 86

Agrostis racemosa Michx. 13:9
Muhlenbergia racemosa (Michx.) BSP

Agrostis dispar Michx. 13:11
in Carolina inferiore.
in Carolina maritima.
Agrostis stolonifera L.

Agrostis aspera Michx. 13:13
in collibus rupibusque regione Ilinoensis

in Illinois

Sporobolus asper (Michx.) Kunth var. asper

Alopecurus aristulatus Michx. 13i17
in paludosis Canadae.

in Canada ad ripas Lacus Champlain.

Alopecurus aequalis Sobol. var. aequalis

Erianthus saccaroides Michx. 13:19
Erianthus giganteus (Walt.) Muhl.

Erianthus brevibarbis Michx. 13:21
in collibus Tennassée et Carolinae.
in collibus desertis ab Amnio Wabash ad Ostium Missouri.

Andropogon macrourum Michx. 14:1
a Carolina ad Floridam

a Virginia ad Carolinam.

Andropogon glomeratus (Walt.) BSP

Andropogon dissitiflorum Michx. 14.5
a Carolina ad Floridam.

in Carolina, Georgia, Florida.

Andropogon virginicus L.

Andropogon avenaceum Michx. 14:6
in vastissimus pratis Illinoensibus.

in regionae IIlinoensium.

Sorghastrum nutans (1...) Nash

Andropogon scoparium Michx. 14:7
Andropogon scoparius Michx.*

Andropogon ternarium Michx. 14:8
in montosis Carolinae.

in regiona Wabash, Georgia montosa.

Andropogon ternarius Michx. var. ternarius

Chloris curtipendula Michx. 14:10
in aridis regionis Illinoensis ad Wabast et in rupibus ad prairie

du rocher.

in hauteurs du Missouri et Poste Vincenne.

Bouteloua curtipendula (Michx.) Torr. var. curtipendula

1984]

1:67

Uttal — Michaux localities 13

Chloris mucronata Michx. 14:13
Dactrvloctenium aegyptium (L.) Willd.

Chloris monostachya Michx. 14:15
Ctenium aromaticum (Walt.) Wood

Tripsacum cylandricum Michx. 14:17
in sabulosis Floridae.

in Florida.

Manisuris cylindrica (Michx.) Kuntze*

Aira ambigua Michx. 1$:2
circa lacus Mistassins et juxta anines in lacun s. Joannis

defluent es.

Riv que lombent au Lac St. Jean.

Deschampsia cespitosa (L.) Beauv.

Aira melicoides Michx. 15:3
Trisetum melicoides (Michx.) Scribn.

Aira obtusata Michx. 15:4
in aridis, a Carolinae ad Floridam.

in sabulosis Carolinae, Georgiae, Floridae; in Florida juxta

dominus Wiggin.

Sphenopholis obtusata (Michx.) Scribn. var. obtusata

Melica glabra Michx. 15:5
a Virginia ad Floridam.

a Carolina ad Floridam. Florida f. Matanca.

Melica mutica Walt.

Trachynotia polystachia Michx. 15:6
in inundatis maritimis, a Nova Anglia ad Floridam.

Basse Caroline.

Spartina cynosuroides (L.) Roth

Trachynotia juncea Michx. 15:8
in sicca maritimis Carolinae, Georgiae.

Bord des creeks sales Basse Caroline.

Spartina patens (Ait.) Muhl. var. patens

Eleusine mucronata Michx. 15:9
in cultis Ilinoensibus.

Illinois.

Leptochloa filiformis (Lam.) Beauv. var. filiformis

Bromus canadensis Michx. 15:14
in Canada.

Canada: Lac St. Jean.

Bromus ciliatus L. var. ciliatus

Festuca distichiphylla Michx. 15:18
in maritimis Carolinae.

14

Rhodora

Bord de la Mer en Carol.
Distichlis spicata (L.) Greene var. spicata

Festuca polystachya Michx.
Leptochloa fascicularis (Lam.) A. Gray

Festuca poaeoides Michx.

ad ripas maritimas fluminis S. Laurentii.
Fleuve St. Laurent

Festuca elatior L.*

Poa hirsuta Michx.

in Carolina inferiore.

Carol

Eragrostis hirsuta (Michx.) Nees

Poa reptans Michx.

in limosis ripariis amnium regionis Illinoensis.

Rivierre Kaskasia.
Eragrostis reptans (Michx.) Nees

Uniola gracilis

in umbrosis sylvarum, a Carolina ad Georgiam.
Larinburg (not to be confused with “Lasinburg”, New York:

[Vol. 86

15:19

16:12

16:14

17:20

presumably Laurinburg, Scotland County, North Carolina - auth.)

Chasmanthium laxum (L.) Yates.

Uniola latifolia Michx.

in occidentalibus montium Alleghanis.
Illinois.

Chasmanthium latifolium (Michx.) Yates.

Avena flumosa Michx.
in Pensylvania, Carolina.

Parmi les aristida Illinois. A Canada ad Carolinam.

Danthonia spicata (L.) Beauv. ex R. & S.

Briza canadensis Michx.

Glyceria canadensis (Michx.) Trin. var. canadensis

Avena striata Michx.

a sinu Hudsonis per tractus montium ad Canadam.,
Lac des Cygnes Montagn. ent. la Baye d’Hudson et le

Canada Mistassins.
Schizachne purpurascens (Torr.) Swallen

Avena mollis Michx.
in Canada.
Montreal.

Trisetum spicatum (L.) Richt. var, spicatum

16:20

17:2

17:7

Uttal — Michaux localities

Avena palustris Michx.

in Georgiae et Carolinae paludosis graminosis. Floret Maio.
Georgia et Carolina. Lieux humides.

Trisetum pensylvanicum (L.) Beauv.

Arundo canadensis Michx.

in Canada Lacunum.

a sinu Hudsonis ad Canadam praesertim ad ripas Lacunum.
Calamagrostis canadensis (Michx.) Beauv. var. canadensis

Arundinaria macrosperma Michx.

ad ripas flum. Mississipi: in Carolina, Florida, etc.

Martio florens.

a Virginia ad Floridam et in occidentibus ab Illinoensibus ad
ostium Misissipi.

Arundinaria gigantea (Walt.) Muhl.

Zizania miliacea Michx.

in aquosis Americae septentrionalis.

Lac Champlain, New Jersey, Carolines, Illinois et Lacs d’Am
Zizania aquatica L. var. aquatica

Zizania fluitans Michx.

ad lacum Champlain.

juxta Charleston, Carolines, Fleuret in Juillet.
Zizania aquatica L. var. aquatica

Zizania clavulosa Michx.
Zizania aquatica L. var. aquatica

Stipa sericea Michx.

in sabulosis Carolinae inferioris.
inferiore in Carolina

Muhlenbergia capillaris (Lam.) Trin.

Lechea thymifolia Michx.
Lechea minor L.

Lechea racemulosa Michx.

Lechea tenuifolia Michx.
Lechea tenuifolia Michx. var. tenuifolia

Galium circaezans Michx
Galium circaezans Michx. var. circaezans

Galium asprellum Michx.
in septentrionalibus Canadae.
Quebec, Lac St. Jean.

Galium claytoni Michx.

17:9

17:10

17:12

17:14

17:15

20:7

20:8

Rhodora [Vol. 86

in Canada et Nova Caesarea.

Lac St. Jean, New Jersey.

Galium tinctorium L. subsp. tinctorium

Galium latifolium Michx. 20:9
in altis montibus utrisisque Carolinae.

Caroline hautes montag.

Galium uniflorum Michx. 20:10
in Carolina

Caroline

Galium triflorum Michx. 20:11

in umbrosis Canadae sylvis.
in Canada.

Galium hispidulum Michx. 20:12
in Carolina inferiore.
in Carolina.

Galium puncticulosum Michx. 20:13
in Carolina inferiore.

Basse Carolina.

Galium pilosum Ait. var. pilosum

Rubia Brownei Michx. 20:14
in umbrosis, a Carolina ad Floridam.

in umbrosis Sylvarum Carolinae.

Galium hispidulum Michx.

Spermococe diodina Michx. 20:17
in aridis sabulosis sylvarum Carolinae. Augusto et Septembri

floret.

Lieux arides, basse Caroline.

Diodea teres Walt. var. teres

Polypremum Linnei Michx. 21:3
Polypremum procumbens L.

Oldenlandia glomerata Michx. 21:5
in humidis Carolinae inferioris.

Basse Caroline.

Oldenlandia uniflora L.

Houstonia Linnaei var. a elatior Michx. 21:11
Houstonia caerulea L.

Houstonia Linnaei var. B minor Michx. 21:10
Houstonia pusilla Schoepf.

Houstonia serpyllifolia Michx. 21:9
ad fonticulos rivosque excelsorum montium Carolinae.

1:90

1:90

Uttal Michaux localities

Maio florens.
in sommet des plus hautes montagnes.

Houstonia rotundifolia Michx.

in apricis submaritimis Floridae et Carolinae. Martio floret.
in Voyes la descript. au supplem. des Pl. de la Floride. No.

Houstonia procumbens (Walt. ex J. F. Gmel.) Standl.

Houstonia varians Michx.

in diversis locis ubrusque Carolinae et alibi. Floret Julio.
haute et basse Caroline.

Houstonia purpurea L. var. purpurea

Houstonia angustifolia Michx.
Houstonia longifolia Gaertn. var. longifolia

Ludwigia nitida Michx.

in humidis Carolinae inferioris. Floret Maio.
in Carolina, maio floret.

Ludwigia palustris (L.) EIl.

Ludwigia pedunculosa Michx.
in paludosis submaritimis Carolinae inferioris. Maio floren.

i

13:14

lem.

21:16

21:19

2257

in Carolina maritima ad ripas uliginosis. Maio floret. flos lutens.

Ludwigia arcuata Walt.

Ludwigia microcarpa Michx.
in humidis Carolina inferioris. Julio floret.
in udis graminosis Carolina. Goose Creek.

Ludwigia angustifolia Michx.

22:10

22:12

juxta fossas aquosas Carolinae inferioris. Augusto et Septembri

floret.
Carol.
Ludwigia linearis Walt.

Ludwigia jussiaeoides Michx.

22:14

in aquosis praesertim umbrosis Carolinae inferioris. Floret Julio,

Caroline.
Ludwigia decurrens Walt.

Ludwigia macrocarpa Michx.
Ludwigia alternifolia L.

Ludwigia virgata Michx.

Ludwigia mollis Michx.

in paludosis Carolinae inferioris. Julio floret.
Basse Carol

Ludwigia pilosa Walt.

Ludwigia capitata Michx.
Ludwigia suffruticosa Walt.

22:16

22:17
22:18

22:20

1:92

Rhodora [Vol. 86

Cornus tomentosula Michx. 23:7
in collibus amnium ripariis Canadae, New Yorck, Connecticut,

Junio et Julio floret.

Lac Champlain Canada et Rivierre Sagney.

Cornus rugosa Lam.

Cornus lanuginosa Michx. 23:8
in humidis Canadae et Carolinae.

Montreal. Riviere de l’'Assompt.

Cornus amomum Mill.

Cornus stolonifera Michx. 23:18
ad ripas amnium rivorumque Canadae, Novae Angliae.

des rivages Lac St. John et rivierres qui coulant aux Lacs

Mistassins,

Tres Rivierres.

Cornus stolonifera Michx. var. stolonifera

Cornus fastigiata Michx. 23:20
in dumosis, ad ripas rivolorum Virginiae, Carolinae, Aprili

floret.

Pres le ruisseau de Neuf Miles...en Caroline. Fleurit a la fin

d’Avril.

Cornus foemina Mill. subsp. foemina

Cornus asperifolia Michx.

Plantago sparsiflora Michx. 24:17
in sylvis Carolinae et Georgiae.
Basse Caroline et Georgia in aridis sylvanum.

Plantago aristata Michx. 24:19

Fraseri walteri Michx. 25:1
Swertia caroliniensis (Walt.) Kuntze

Centaurella verna Michx. 25:4
Bartonia verna (Michx.) Muhl.

Centaurella paniculata Michx. 25:5
Bartonia paniculata (Michx.) Muhl.

Ammannia humilis Michx. 25:8
in paludosis Carolinae septentrionalis. Septembri florens.

Fayetteville et Carolina.

Rotala ramosior (L.) Koehne

Allionia nyctaginea Michx. 25:10
ad ripas fluminis Tennassée.

au dessus de la maison du Docteur White des Rochers

Cumberland River.

Mirabilis nyctaginea (Michx.) MacM.

—

Uttal — Michaux localities

Pinckneya pubens Michx.

Caprifolium bracteosum Michx.
in montibus Carolinae.

Lac Champlain et Maryland.
Lonicera dioica L. var. dioica

Xylosteum villosum Michx.

in praeruptis saxosis, per tractus montium, a sinu Hudsonis ad
Canadam.

Lac des Cygnes, Mistassin et Riv des Goelands. Lonicera villosa
(Michx.) R. & S.*

Symphoricarpus vulgaris Michx.

in Virginia et Carolina.

Etat de Cumberland et Lieux arides de Kentucky.
Symphoricarpus orbiculatus Moench.

Symphoricarpus racemosus Michx.

in montanis, ad lacus Mistassins.

hauteur des terres pres Mistassin.
Symphoricarpus albus (L.) Blake var. albus

Diervilla tourneforti Michx.

in Canada, Nova Anglia et in cacumne montium excelsorum
Carolinae.

sur Black Mountain, Albany, Lasinburg, jusqu’aux Lacs de la
Baye d’Hudson.

Diervilla lonicera Mill.

Triosteum minus Michx.

in Virginia.

in Carolina septentrionali juxta Hill’s Iron Works.
Triosteum angustifolium L. var. angustifolium

Campanula acuminata Michx.

in remotis occidentalibus Pensylvania et Virginia.
hautes Montagn de Carol. et Kentucky et Illinois.
Fleurit en Juillet, Aoust.

Campanula americana L.

Campanula flexuosa Michx.
in montibus altis Carolinae.
hautes Montag.

Campanula divaricata Michx.

Campanula amplexicaulis Michx.

in cultis hortensibus Pensylvaniae, Carolinae.
in Carolina.

Specularia perfoliata (L.) A. DC.*

26:15

26:16

26:17

26:18

26:19

26:21

—

_

Rhodora

Campanula divaricata Michx.
in altis montibus Carolinae.
Montagn de la Carol.

Ribes albinervium Michx.

in Canada, ad amnen Mistassin.
Rivierre Mistassin Canada.
Ribes triste Pall.

Ribes rigens Michx.

in Canada, ad amnen Mistassin.
Rivierre Mistassin.

Ribes glandulosum Grauer

Ribes hirtellum Michx.
in saxosis, ad amnem Sagney.
Rochers de la Rivierre Sagney.
Ribes oxycanthoides L.

Ribes gracile Michx.

in montibus Tennassée.

in montibus Tennassée in itinera Nashville.
Ribes missouriense Nutt. var. missouriense

Ribes rotundifolium Michx.
in montibus excelsis Carolinae.
Tres hautes Montg Carolines.

Ribes trifidum Michx.

juxta Quebec et sinum Hudsonis.
Quebec.

Ribes glandulosum Grauer

Thesium corymbulosum Michx.

in aridis America Septentrionalis.

Lac Champlain, Pensylv, haute Caroline.
Comandra umbellata (L.) Nutt. var. umbellata

Anychia herniarioides Michx.

in arenosis Carolinae septentrionalis.

Pinieres arides, in sabulosis Carolinae.

Paronychia herniarioides (Michx.) Nutt.

Note: not listed in North Carolina today; very rare in
South Carolina-auth.

Anychia argyrocoma Michx.
in rupibus montium superioris Carolinae.
Rochers des hautes Montagn des Carolines.

Paronychia argyrocoma (Michx.) Nutt. var. argyrocoma

Anychia dichotoma Michx.

in collibus calcariis Pensylvania superioris et Kentucky.

[Vol. 86

27:3

27:11

27:12

27:13

27:14

27:12

27:18

27:19

27:20

27:21

Za5 E45

Zaz

1:119

1:120

Uttal — Michaux localities

in excelsis montib. Carolinae Montag. de Pensyl
Paronychia canadensis (L.) Wood.

Asclepias hybrida Michx.
Asclepias variegata L.

Asclepias longifolia Michx.
in sylvis Georgiae occidentalibus.
Carolines et Illinois.

Asclepias debilis Michx.

in aquosis umbrosis Carolinae.
in umbrosis Carolinae.
Asclepias perennis Walt.

Asclepias paupercula Michx.

in herbidis humidis Carolina inferioris.
in herbidis humidis Carolina.
Asclepias lanceolatis Walt.

Asclepias laurifolia Michx.

secus amnem Althamaha, in Georgia.
secus amnem Altamaha, in Georgia.
Asclepias rubra L.

Gonolobus laevis Michx.

in dumetosis Fluminis Mississipi.
Illinoense.

Cynanchum laeve (Michx.) Pers.

Gonolobus macrophyllus Michx.
Cynanchum laeve (Michx.) Pers.

Gonolobus hirsutis Michx.
Matalea carolinensis (Jacq.) Woodson

Gelsemium mitidum Michx.
Gelsemium sempervirens (L.) Ait. f.

Echites puberula Michx.

in sylvis Carolinae inferioris.

Ohio, Fort Massac, Carol. et Georgia. Partie meridionale de
la Virginie.

Trachelospermum difforme (Walt.)

Amsonia latifolia Michx.
Amsonia Tabernaemontana Walt.

Amsonia angustifolia Michx.
in sabulosis apricis Georgia.
Carol.

Amsonia ciliata Walt.

21

28:11

28:18

28:19

28:20

28:21

29:3

29:4

29:5

29:12

29:15

29:17

29:18

_

—

_—

:130

:130

Rhodora

Sideroxylon chrysophylloides Michx.
in dumetis littoralibus Carolinae.
juxta Charleston; Carol. et Georgie.
Bumelia tenax (L.) Willd.

Sideroxylon lanuginosum Michx.
in dumosis humidis Georgiae.

in Georgia.

Bumelia lanuginosa (Michx.) Pers.

Sideroxylon reclinatum Michx.
in dumetosis ripariis Georgiae.

sur la rivierre Ste Marie. Fleurit le 10 May. (1787 - auth.)

Bumelia tenax (L.) Willd.

Dodecatheon integrifolium Michx.
in montibus Alleghanis, juxta rivulos sylvaticos.

in montibus occidentalibus Caroline juxta rivulos sylvaticos.

Dodecatheon meadia L. var. meadia

Primula mistassinica Michx.
ad lacus Mistassins Canadam inter et fretum Hudsonis.
Rivierre des Goelands.

Menyanthes trachysperma Michx.
Nymphoides aquatica (Walt. ex J. F. Gmel.) Kuntze

Batschia canescens Michx.
Lithospermum canescens (Michx.) Lehm.

Batschia gmelini Michx.

Lithospermum carolinense (Walt. ex J. F. Gmel.) MacM.

Lithospermum angustifolium Michx.
ad flumen Ohio.

Pres les Rapides de |’Ohio.
Lithospermum incisum Lehm.

Lithospermum latifolium Michx.
in umbrosis sylvis Kentucky.
Bois umbrages de Kentucky.

Pulmonaria parviflora Michx.

ad littoria mari ascendente inundata imi fluminis S. Laurentii.

Malbaye.
Mertensia maritima (L.) S. F. Gray var. maritima

Cynoglossum amplexicaule Michx.
in montibus Alleghanis.

Haute Caroline, Kentucky, etc.
Cynoglossum virginianum L.

[Vol. 86

30:1

30:3

30:6

30:7

30:16

31:13

31:14

31:17

1984] Uttal — Michaux localities

1:133. Onosmodium molle Michx.
T: in Tennassée, circa Nashville.
M: Nashville, lieux arides et pierreaux.
Onosmodium molle Michx. var. molle.

1:133. Onosmodium hispidum Michx.
Onosmodium virginianum (L.) A. DC. var. virginianum
1:134 Hydrophyllum appendiculatum Michx.

T: in sylvis montanis Tennassée.
M: Forets de Cumberland et de Kentucky.

wo
BR

Phacelia bipinnatifida Michx.

i
w
wa

Phacelia fimbriata Michx.

T: in excelsis montibus Carolinae.

M: Lieux bas tres humides et riches au pied des montagnes apres
avoir tourne a main droite pour aller de Th. Yong chez

Davenport.
1:160 Ampelopsis bipinnata Michx.
Ampelopsis arborea (L.) Koene
1:159 Ampelopsis cordata Michx.
1:139° Convolvulus obtusilobus Michx.

T: in littoribus arenosis Georgiae et Floridae.
M: in littoribus arenosis Georgiae.
Ipomoea stolonifera (Cyrill.) J. F. Gmel.

136 Convolvulus stans Michx.
T: juxta lacum Champlain, Canada.
M: Lac Champlain.
Calystegia spithamaea spp. stans (Michx.) Brummitt

a
w
|

Convolvulus ciliolatus Michx.

T: circa urbem Knoxville, in Tennassée
M: environs de Knoxville.

Ipomoea pandurata (L.) G. F. W. Mey.

1:1413 | [pomoea macrorhiza Michx.

1:1414 Ipomopsis elegans Michx.
Ipomopsis rubra (L.) Wherry

1:143. Phlox triflora Michx.
Phlox glaberrima ssp. triflora (Michx.) Wherry

34On same sheet. “x” is on left side of sheet, “xy” is on right.

23

31:18

31:19

32:21

24 Rhodora [Vol. 86

1:144 Phlox aristata Michx.
var. a virens Michx. 34:3
T: in Carolina, ad ripas amnis Santee.
M: Riv. Santee, locis sabulosis
Phlox pilosa L. var. pilosa
var. B canescens Michx. 34:2
T: in Tennassée, circa Knoxville.
M: Knoxville
Phlox pilosa L. var. pilosa

1:145 Phlox reptans Michx. 34:7
T: in excelsis montibus Carolinae occidentalis.
M: Lieux umbrages des hautes Montagnes Carolina
septentrionalis.
Phlox stolonifera Sims

1:143 Phlox latifolia Michx. 34:12
T: in umbrosis humidis et solito sphagnosis sylvanum Carolinae.
M: in umbrosis Carolinae.
Phlox ovata L.

1:147. Chironia chloroides Michx. 35:11
T: in Noveboraco et Nova Caesarea.
M: New York, prairies.
Sabatia dodecandra (L.) BSP

1:146 Chironia paniculata Michx. 35:7
T: in Georgia et Carolina.
M: Georgia.
Sabatia quadrangula Wilbur
1:146 Chironia gracilis Michx. 35:9
T: in Carolina inferiore.
M: Carol.
Sabatia campanulata (L.) Torr.
1:148 Verbascum claytoni Michx. 35:18

T: in Carolina.
M: rare in Caroline.
Verbascum blattaria L.

1:149 Physalis lanceolata Michx. 35:21
T: in Carolina.
M: Carolina

Physalis heterophylla var. villosa Waterfall

1:149° Physalis obscura Michx.

var. a glabra Michx. 35:1 (a)
Physalis pubescens var. glabra (Michx.) Waterfall
var. B viscido-pubescens Michx. 35:1 (b)

Physalis pubescens var. grisea Waterfall

—

—

_

—_— —

—

_

|

<3

Uttal — Michaux localities 25

Azalea canescens Michx. 36:4
juxto rivolos Carolinae inferioris.

Basse Caroline.

Rhododendron canescens (Michx.) Sweet.

Azalea periclymenoides Michx. 36:9
in New Jersey.

New York.

Rhododendron nudiflorum L.

Azalea calendulacea Michx.

var. a flammea Michx. 36:13
ad ripas fluvii Savannah loco dicto Two Sisters.
Two Sisters.
Rhododendron flammeum (Michx.) Sargent

var. B crocea Michx. 36:15
Rhododendron calendulaceum (Michx.) Torr.

Pyxidanthera barbulata Michx. 36:18
in Carolina superiore.

in sabulosis Carolina septentrionalis juxta Wilmington.

Pyxidanthera barbulata Michx. var. barbulata

Rhamnus franguloides Michx. 36:19
ad lacum Champlain.

de Cumberland; Kentucky Lieux arides.

Rhamnus alnifolius L’Her.

Rhamnus minutiflorus Michx. 36:20
Sargeretia minutiflora (Michx.) Mohr

(reported in text from North Carolina; it is apparently not in that

state now. - auth.)

Ceanothus microphyllus Michx. 37:5
in herbosis sabulosis sylvarum Georgiae et Floridae.
le 16 Mars (1787 - auth.) Lieux arides vers Nord West river.

Claytonia caroliniana Michx. 37:17

Hydrocotyle lineata Michx. 38:21
in inundatis Carolinae inferioris. Aprili et Maio floret.

in salsis a mare inundatis Carolinae.

Lilaeopsis chinensis (L.) Kuntze

Eryngium ovalifolium Michx. 38:5
Eryngium integrifolium Walt.

Eryngium yuccifolium Michx. 38:7
in paludosis Virginiae.
in apricis sylvarum inter gramineas a Virginia ad Floridam.

Daucus pusillus Michx. 38:8

26

—_

—_

_

—_

—

[167

[167

[167

:168

Rhodora [Vol. 86
in campestribus Carolinae.
in salbulosis Carolina.
Ammi capillaceum Michx. 38:9

in campestribus Carolinae.
in cultis Carolinae.
Prilimnium capillaceum (Michx.) Raf.

Selinium canadense Michx. 38:10
ad ostium fluminis S. Laurentii.

in Canada juxta Tadoussack.

Conioselinum chinense (L.) BSP

Heracleum lanatum Michx. 38:11
in Canada.
Chicoutoume, Tadoussack.

Ligusticum actaeifolium Michx. 38:13
ad ripas fluminis S. Laurentii, juxta Tadoussack.

Tadoussack in Canada.

Ligusticum canadense (L.) Britt.

(Type locality out of range; specimen annotated by L. Constance,

1963 - auth.)

Ligusticum barbinode Michx. 38:14
Thaspium barbinode (Michx.) Nutt.

Sium lineare Michx. 38:15
Sium suave Walt.

Angelica triquinata Michx. 38:16
(Type locality (Canada) out of range - auth.)

Sison pusillum Michx. 38:17
Spermolepis divaricata (Walt.) Raf. ex Seringe

Sison trifoliatum Michx. 38:20
in Carolina superiore.

haute Caroline in Montosis.

Zizia trifoliata (Michx.) Fern

Sison marginatum Michx. 38:21
in udis Carolinae.

Caroline.

Oxypolis rigidior (L.) Raf.

Myrrhis claytoni Michx. 39:1
in montibus Alleghanis.

in Canada, Nova Anglia et in monitbus Carolinae.

Osmorhiza claytoni (Michx.) Clarke

Heuchera cortusa Michx. 39:7
in in variis locis Pensylvaniae, Carolinae, etc.

1984]

—

PU Pz

[174

SET7

Uttal — Michaux localities

frequentius in regione Illinoensis.
Illinoensi regiona Carolina etc.
Heuchera americana L. var. americana

Heuchera villosa Michx.

in altoribus montibus Carolinae borealis.
Tres hautes montagnes des Carol.
Heuchera villosa Michx. var. villosa

Ulmus alata Michx.
in Virginia et Carolina inferiore.

ra

39:8

39:10

Kentucky endroits pierreux pres Beardstown (see amended

gazetteer in introduction - auth.), Carolina Sept., Caroline Meridio-
nale et environs de Richmond 1794 (the latter station underlined,
suggesting it was collected there and is probably the actual type

locality - auth.)

Ulmus fulva Michx.

in Canada, Vermont, Connecticut, montibus Alleganis,

frigid. Americae regionibus.

Baye d’Hudson Canada, haute Carolina, N. York, etc.

Ulmus rubra Muhl.

Salsola platiphylla Michx.

in regione Illinoensium, hue alluvientibus Mississippi
aquis allata.

in regione I}linoensium.

Cycloma atriplicifollum (Spreng.) Coult.

Gentiana acuta Michx.

in altis montibus Carolinae et in Canada, prope
Tadoussack.

Partie basse pres Tadoussack, Canada.
Gentiana amarella L.*

Gentiana amerelloides Michx.

in monticulis Kentucky.

Lieux arides sur Kentuckey river.

Gentiana quinquefolia L. var. quinquefolia*

Gentiana puberula Michx.*

ad confluentum fluviorum Ohio et Mississipi, prope
propringnaculum Cheroquis.

Fort Cheroquis ad ostium fluvil Ohio.

Gentiana angustifolia Michx.
in pratis Carolinae inferioris.
in herbosis Carolinae.
Gentiana autumnalis L.

Hydrolea caroliniana Michx.

39:14
etc:,

40:1

40:5

40:6

40:8

40:9

40:10

28

<3

1:182

Rhodora

in aquis et montibus Carolinae inferioris.
in aquis Carolina.
Hydrolea quadrivalvis Walt.

Viburnum lantanoides Michx.

in rupestribus opacarum sylvarum Canadae.

des Cataractes de la Caroline Septentrionale sur les hautes
montagnes. Bois ombrages et bas vers ter Lac Champlain et
en Canada.

Viburnum alnifolium Marsh.*

Viburnum molle Michx.
in Kentucky, circa Danville.
in Kentucky, juxta Danville.

Viburnum opulus
var. @ europeanum Michx.
var. B pimina Michx.
var. y edule Michx.
in Canada
var. B Depuis Lac Champlain jusquan Lac St. Jean hautes
Montag. de Pensylv. hautes M de Caroline
(not found in Carolina - auth.)
Viburnum opulus var. americanum Ait.*

Sambucus pubens Michx.*
in altis montibus Pensylvaniae, Canadae, Carolinae.
in Canada et in altis Montib. Allegani.

Rhus toxicodendron var. B quercifolium Michx.
in Carolina inferiore et Georgia.

Virginie et Carolines.

Rhus toxicodendrom L.*

Rhus toxicodendron var. a vulgare Michx.
Rhus radicans var. rydbergii (Small) Rehder*

Rhus toxicodendron var. y microcarpon Michx.
Rhus radicans L. var. radicans*

Rhus pumilum Michx.

in Carolina superioris comitatu Mecklenbourg.

in montosis Carolina Burke comitatus Haute Carolina.
(this type locality is in error - auth.)

Rhus michauxii Sarg.

Hypoxis caroliniensis Michx.
Hypoxis hirsuta (L.) Coville

Aletris alba Michx.
Aletris farinosa L.

[Vol. 86

40:12

40:13

41:19

42:3

42:4

42:13

43:21

44:2

1:19]

1:201

Uttal — Michaux localities 29

Juncus melanocarpus Michx. 44:4
in montibus sinum Hudsonis inter et Canadam juxta lacum

Cycmorum.

hauteur des Terres Lac des Cygnes.

Luzula parviflora (Ehrh.) Desv. var. parviflora

Juncus repens Michx. 44:5
in Carolina et Georgia.

in inundatis Carolinae, Georgiae. Mayo floret; Junio-Julio

matureait.

Juncus acuminatus Michx. 44:10
in Carolina inferiore.

‘in Carolina, Georgia.

Juncus aristulatus Michx. 44:11
Juncus marginatus Rostk. var. marginatus

Juncus bicornis Michx. 44:12
in Georgia et Carolina.

“Duplicata Georgia”

Juncus tenuis Willd. var. tenuis

Juncus polycephalus Michx. 44:13
Juncus scirpoides Lam.

Allium mutabile Michx. 45:4
in Georgiae meridionalibus. Inuente Maio floret.

in Carolina et in Georgia.

Allium canadense L. var. canadense

Lilium carolinianum Michx. 45:16
Lilium michauxii Poir.

Uvularia puberula Michx. 45:20
in altissimus montibus Carolinae.

Tres hautes Montagn.

Uvularia pudica (Walt.) Fern.*

‘ Streptopus roseus Michx. 46:1

in excelsis montibus Carolinae septentrionalis et in Canada.
in sylvis a sinu Hudsonis ad Quebec et in excelsis

montib Carolinae.

Streptopus roseus Michx. var. roseus

Streptopus lanuginosus Michx. 46:3
in altis montibus Carolinae meridionalis.

in excelsis montib Carolinae septentrionalis.

Disporum lanuginosum (Michx.) Nichols.

Convallaria umbellulata Michx. 46:16
in montibus Alleghanis.

30

Rhodora [Vol. 86

Vue dans les Montagnes du Pays des Indiens Cheroquis et
cellui de la Car. Sept.
Clintonia umbellulata (Michx.) Morong.

Diphylleia cymosa Michx. 47:2

Cabomba aubleti Michx. 47:7
in aquosis Carolinae et Georgiae.

in aquis Ogechee.

Cabomba caroliniana A. Gray var. caroliniana

Chamaerops serrulata Michx. 47:9
in maritimis Georgiae-Floridaeque.

environs de Sunbury en Georgiae at Savanah (entre Savanah et Sunbury).
Serenoa repens (Bartr.) Small

Chamaerops acaulis Michx. 47:13
in Carolina et Georgia.

Environs de Charleston Carolina.

Sabal minor (Jacq.) Pers.

Triglochin triandrum Michx. 47:15
in inundatis circa Charlston.

in salsis paludosis juxta Charleston.

Triglochin striatum R. & P.

Narthecium pubens Michx. 47:17
in Carolina inferiore circa Charlston.

Basse Caroline pres Charleston.

Tofieldia racemosa (Walt.) BSP.

Narthecium glutinosum Michx. 47:18
Tofieldia glutinosa (Michx.) Pers. var. glutinosa

Narthecium pusillum Michx. 47:19
ad lacus Mistassins.

in sphagnosis a sinu Hudsonis ad Lacus Misstassinos.

Tofieldia palustris Huds.*

Xerophyllum setifolium Michx. 47:20
none given

Zinneville Mountain, haute montagne de la Caroline

septentrionale a 30 miles de Morganton.

Xerophyllum asphodeloides (L.) Nutt.

Helonias erythrosperma Michx. 47:21
in ubrosis rivulis Carolinae inferioris.

dans les lieux boubeaux et ombrages Basse Carol.

Amianthium muscaetoxicum (Walt.) A. Gray

Helonias dubia Michx. 48:1
in sabulosis Georgiae et Floridae.

1:219

1:220

1:222

Uttal — Michaux localities

Floride.
Schoenocaulon dubium (Michx.) Small

Helonias angustifolia Michx.

in herbosis et fruticetis sylvarum humidis Carolinae
inferioris.

Lieux plus eleves et moins humides Basse Caroline.
Zigadenus densus (Desr.) Fern.

Zigadenus glaberrimus Michx.
in herbosis, humidis, Carolinae inferioris.
in Carolina.

Trillium rhomboideum Michx.

var. a atropurpureum Michx.
in excelsis montibus Carolinae.
hautes Montagnes des Carolines.
Trillium erectum L.

Trillium rhomboideum var. B album Michx.

Trillium erectum L.

Trillium rhomboideum var. y grandiflorum Michx.
Trillium grandiflorum (Michx.) Salisb.

Trillium pusillum Michx.
in pinetis Carolinae inferioris.

Basse Caroline environ 35 m de Charleston, Gaillard Road.

Rumex crispatulus Michx.
Rumex obtusifolius L. var. obtusifolius

Aesculus lutea Michx.
Aesculus octandra Marsh.*

Aesculus macrostachya Michx.
Aesculus pavia L. var. pavia

Rhexia glabella Michx.
Rhexia alifanus Walt.

Rhexia ciliosa Michx.
Rhexia petiolata Walt.

Rhexia mariana, var. y exalbida Michx.
none given

-Carolines.

Rhexia mariana var. exalbida Michx.*

Rhexia mariana var. B rubella Michx.
none given

Carolines.

Rhexia mariana L. var. mariana

a1

48:2

48:3

48:5

48:10

48:9

48:11

48:16

49:2

49:5

49:7

49:9

40:11

49:12

32

1:221

Rhodora

Rhexia mariana var. a purpurea Michx.
Rhexia mariana var. purpurea Michx.*

Epilobium oliganthum Michx.

ad sinum Hudsonis et lacus Mistassins.
env. de Mistassin.

Epilobium palustre L.

Oenothera chrysantha Michx.

a Quebec usque ad sinum Hudsonis.
Quebec.

Oenothera parviflora L.

Oenothera pusilla Michx.

in rupibus, ad lacus Mistassins.
Lacs Mistassins.

Oenothera parviflora L.

‘Oenothera linearis Michx.
Oenothera fruticosa L. subsp. fruticosa

Oenothera hybrida Michx.

in Carolina superiore.

Caroline.

Oenothera fruticosa L. subsp. fruticosa

Oenothera glauca Michx.

in sylvis remotis et occidentalibus flumini Mississipi
confinibus, versus regionem IIlinoensium.

Ouest de l’Ohio Route aux Illinois.

Oenothera fruticosa subsp. glauca (Michx.) Straley

Gaura angustifolia Michx.
in Carolina inferiore.
Caroline.

Vaccinium galezans Michx.
in umbrosis Carolinae.

in Carolina.

Vaccinium tenellum Att.

Vaccinium myrtilloides Michx.
a Canada ad sinum Hudsonis.
Lacs Mistassins heauteur des Terres.

Vaccinium disomorphum Michx.

none given

Environs de Philade (Philadelphia - auth.)
Vaccinium corymbosum L.

Vaccinium erythrocarpum Michx.
in altis montibus Carolinae Septentrionalis.
Hautes montagnes de la Carol. Sept.

[Vol. 86

49:14

49:17

49:21

50:1

50:3

50:7

50:8

51:12

51:19

52:16

Uttal — Michaux localities

Vaccinium brachycerum Michx.

in Virginia, circa Winchester.

in Virginia Warm Spring. (see introduction - auth.)
Gaylussacia brachycera (Michx.) A. Gray

Vaccinium myrtifolium Michx.

in Carolina.

avant d’arriver a Wilmington sables arides.
Vaccinium crassifolium Andr.

Vaccinium caespitosum Michx.

in borealibus Americae, praesertim circa sinum Hudsonis.

au Lacs Mistassins.

Menziesia smithi Michx.
Menziesia pilosa (Michx. ex Lam.) Juss. ex Pers.

Jeffersonia bartonis Michx.
Jeffersonia diphylla (L.) Pers.

Polygonum ramosissimum Michx.

in regione Illinoensi.

Illinois, Kaskaskia.

Polygonum ramosissimum Michx. var. ramossimum

Polygonum tenue Michx.

in Canada.

in Canada et Nova Caesarea ad Snake Hill in pratensib
Illinoensum.

Polygonum clinode Michx.
in Canada.
in Canada Malbaye, Tadoussack, Riv. Sagney, etc.

Polygonum hydropiperoides Michx.
in Pensylvania, Virginia, Carolina.
Kentucky. fleurit au commencement de Juill. 1794.

Laurus diospyroides Michx.

none given

en Fleurs le 25 Mars (I have not been able to relate
this date to a locality in Michaux’s journal - auth.).
Lindera melissaefolium (Walt.) Blume

Laurus pseudo-benzoin Michx.

juxta rivulos et in udis, a Canada ad Floridam.
Pres d’Alexandrie en Virginie.

Lindera benzoin (L.) Blume

Eriogonum tomentosum Michx.

Pleea tenuifolia Michx.

52:10

52:13

§2:14

53:14

53:17

53:18

$5:4

34

1:256

1:266

1:266

Rhodora [Vol. 86
Andromeda speciosa
var. a nitida Michx. 57:18
var. B pulverulenta Michx. 57:18
Leucothoe racemosa (L.) A. Gray
Kalmia cuneata Michx. 58:20
in Carolina.

pres Cambden.

Rhododendron minus Michx. 59:5
in montibus Carolinae superioris et versus originem

amnis Savannah.

Hautes Montagnes de la Carol. Septentrionale. Aux

sources de la Rivierre Savanah.

Rhododendron catawhiense Michx. 59:6
in montibus excelsis Carolinae septentrionalis juxta

Originem amnis Catawba.

Au sommet des hautes montagnes de la Caroline Septentrionale.

Clethra acuminata Michx. 59:18
in excelsis montibus Carolinae.
Hautes montagnes de Carol.

Cassia fasciculata Michx.

in Pensylvania et Virginia

in Pensylvania et Virginia et Carolina.
Cassia fasciculata Michx. var. fasciculata

Podalyria uniflora Michx. 60:9
in Carolina et Georgia.

Route de Sunbury a Alatamah le 7 May (1787 - auth.)

Baptisia lanceolata (Walt.) Ell.

Podalyria mollis Michx. 61:2
in comitatu Mecklenburg Caroline Superiore.

Entre Burke et Lincoln.

Thermopsis mollis (Michx.) M. A. Curtis ex A. Gray var. mollis

Podalyria coerulea Michx. 60:22
in sabulosis inundatis fluminis Ohio.

Isles Bahama.

Baptisia australis (L.) R. Br. var. australis

(This taxon does not occur in the Bahama Islands but it does

in the Ohio Valley. Michaux’s localization is inexplicit

to me at present - auth.)

Monotropa lanuginosa Michx. 61:4
Monotropa hypopitys L.*

Monotropa morrisoniana Michx. 61:7

‘Monotropa uniflora L.

1:275

Uttal — Michaux localities

Saxifraga virginiensis Michx.

in rupibus Pensylvaniae, Virginiae et in montibus Carolinae.
Haute Caroline et in rupib. Virginiae Pensylvaniae ad Schuykill.
Saxifraga virginiensis Michx. var. virginiensis

Mitella prostrata Michx.

ad fines meriodionales Canadae.
Lac Champlain.

Mitella nuda L.

Jussiaea grandiflora Michx.
Ludwigia peploides var. glabrescens (Kuntze) Shinners*

Hydrangea nivea Michx.
Hydrangea arborescens ssp. radiata (Walt.) McClintock

Hydrangea vulgaris Michx.
Hydrangea arborescens L. var. arborescens

Silene pensylvanica Michx.

in Pensylvania.

Plus rare dans la Basse Caroline.
Silene caroliniana Walt.”

Stellaria pubera Michx.

in montibus sylvaticus Carolinae septentrionalis.
Hautes montagnes des Carolines.

Arenaria glabra Michx.

in rupibus Carolinae septentrionalis.

Rochers des h. Montag. de la Am. Sept.
Arenaria groenlandica var. glabra (Michx.) Fern.*

Arenaria squarrosa Michx.

in sabulosis, pinetis Carolinae, juxta amnem Santee.
Sables arides high hills Santee.

Arenaria caroliniana Walt.*

Arenaria stricta Michx.

in rupibus Novae Angliae, Canada.

Lac Champlain et Canada sur les Rochers Etat de N. York
pres Albany. Arenaria stricta Michx. var. sfricta*

Spergulastrum gramineum Michx.
Stellaria longifolia Muhl. ex Willd.

Spergulastrum lanceolatum Michx.

in borealibus Americae septentrionalis.
Riv. Sagney.

Stellaria calycantha (Ledeb.) Bong.

Spergulastrum lanuginosum Michx.

35

61:8

61:19

62:10

62:11

62:13

62:15

62:17

62:18

62:21

36

= 3

Rhodora

in meridionalibus Amercae septentrionalis.

Georgia 1790 #7 du Voyage en Georgia Insula Cumberland,
Georgia americana.

Arenaria lanuginosa (Michx.) Rohrb. ssp. lanuginosa

Sedum ternatum Michx.
in rupibus altissimus Americae septentrionalis.
hautes montagnes des Carolines et de Virginie.

Sedum telephioides Michx.

Sedum pulchellum Michx.
in rupibus, circa Knoxville.
Western Territories on the Rocks.

Sedum pusillum Michx.

in Carolina septentrionali, loco dicto Flat-rock.

Flat Rock (Kershaw County, South Carolina, not Henderson
County, North Carolina - auth.)

Asarum arifolium Michx.

in umbrosis sylvis Carolinae inferioris.
Caroline.

Hexastylis arifolia (Michx.) Small*

Befaria paniculata Michx.

in Florida arenosis.

Partie meriodionale de la Georgie et Floride.
Befaria racemosa Vent.

Decumaria forsythia Michx.
Decumaria barbara L.

Chrysobalanus oblongifolius Michx. *
in sabulosis sylvarum Georgiae et Floridae.
ad Fluvium Ste. Mary in Georgia.

Cerasus borealis Michx.
Prunus pensylvanica Lf.

Prunus chickasa Michx.
Prunus angustifolia Marsh.

Prunus sphaerocarpa Michx.
in maritimis Novae Angliae.
in maritimus Nova Caesarea.
Prunus maritima Marsh.

Prunus hiemalis Michx.
Prunus americana Marsh. var. americana

Cerasus virginiana Michx.
var. @ virginiana

[Vol. 86

63:4

63:10

63:11

63:21

64:3

64:6

64:9

64:10

64:12

64:13

1984]

Uttal — Michaux localities 37

var. B humilior Michx.
Prunus serotina Ehrh. var. serotina

Prunus acuminata Michx.

in Virginia.

none given

Prunus maritima Marsh. (occasional reports of this species
from Virginia have not been verified - auth.)

Agrimonia striata Michx. 64:20
in Canada

Chicoutoume

Crataegus apiifolia Michx. 65:1

in humidis sylvarum Carolinae.
des swamps de la Basse Caroline.
Crataegus marshallii Egg).

Crataegus spathulata Michx. 65:4
in Carolina utruque et Virginia.
haute Caroline.

Mespilus arbutifolia (L.) Michx. var. melanocarpa Michx. 65:20
in Canada, a sinu Hudsonis ad Virginiam et in Montis altis
Carolinae.

Tres hautes Montagnes de la Caroline septentrionale et Canada;

aussi Connecticut, Boston, etc.

Aronia melanocarpa (Michx.) Ell.

Mespilus canadensis var. a obovalis Michx. 65:21
Carolina inferiore.

Carolines.

Amelanchier obovalis (Michx.) Ashe

Mespilus canadensis var. 6 oligocarpa Michx. 66:1
Amelanchier bartramiana (Tausch) Roemer

Mespilus canadensis var. B cordata Michx. 66:2
a Canada ad Virginiam et in montibus Carolinae.

a Canada ad Virginiam et in montibus Carolinae

haute et basse Caroline. Rare dans la basse Caroline.

Amelanchier arborea (Michx. f.) Fern.

Mespilus canadensis var. y rotundifolia Michx. 66:3
in Canada.

Lac Champlain, Lac Mistassin.

Amelanchier canadensis (L.) Medic.

Rosa setigera Michx. 67:5
in Carolina inferiore.

Illinois, Kentucky, Georgia.

Rosa setigera Michx. var. setigera

1:300

1:303

1:303

Rhodora [Vol. 86
Rosa laevigata Michx. 67:6
Rosa pensylvanica Michx. 67:8

in paludosis Pensylvaniae et Novae Angliae.
a Nova Anglia ad Pensylvania.
Rosa virginiana Mill.

Rubus obovalis Michx. 67:12
Rubus hispidus L.*

Rubus acaulis Michx.* 67:14
in sphagnosis sinui Hudsonis adjacentibus.
in sphagnosis juxta sinum Hudsonis.

Rubus strigosus Michx. 67:15
in montibus Pensylvaniae et in Canada.

des Pensylvanie Etat de New York Canada.

Rubus strigosus Michx. var. strigosus

Rubus trivialis Michx. 67:20
in Carolina et Pensylvania.
Haute et basse Carolines.

Dalibarda fragaroides Michx. 67:21
none given

au nord au pres de |-Etang a un demiquart de mile de

Knoxville (Tennessee juxta Knoxville).

Waldsteinia fragaroides (Michx.) Tratt.

Dalibarda violaeoides Michx. 68:1
Dalibarda repens L.

Geum geniculatum Michx. 68:3
in Canada (M: also). Locality incorrect; restricted to
high mountains of North Carolina and Tennessee - auth.)

Geum radiatum Michx. 68:5
in excelsis Carolinae montibus.

Sommet de Roun Mountain.

(A. Gray is sometimes incorrectly given as the author of this

species. Topotypes are frequent in collections - auth.)

Potentilla hirsuta Michx. 68:13
in Canada, a Quebec ad ostium fluminis S. Laurentii.

Malbaye in Canada, Tadoussack in Canada a Quebec

ad ostium fluminis St. Laurentii.

Potentilla norvegica L.

Potentilla simplex Michx. 68:17
in Canada, Pensylvania, et Carolina.
Montagnes de Pensylvanae, Virginiae, etc.

1984]

Uttal — Michaux localities

Potentilla simplex Michx. var. simplex

Calycanthus ferax Michx.
in altis montibus Carolinae et Virginiae.
Montagnes de Caroline.

Calycanthus floridus var. laevigatus (Willd.) T & G

Tilia canadensis Michx.
Tilia americana L.

Tilia laxiflora Michx.

in maritimis Carolinae, Virginiae.
Tilleul de Caroline.

Tilia caroliniana Mill.

Helianthemum ramuliflorum Michx.
in Georgia et Carolina.

“Yard in Cels.” (Charleston ? - auth.)
Helianthemum canadense (L.) Michx.

Helianthemum corymbosum Michx.
in maritimis Carolinae et Georgiae.
Caroline.

Chelidonium diphyllum Michx.
Stylophorum diphyllum (Michx.) Nutt.

Sarracenia variolis Michx.
Sarracenia minor Walt.

Sarracenia psyttacina Michx.
Sarracenia psittacina Michx.

Nymphaea longifolia Michx.

in amnibus Carolinae Sept. et Merid.

Black River, Yadkin and Santee Rivers Carolines
Nuphar luteum ssp. sagittifolium (Walt.) Beal

Delphinium tridactylum Michx.

in montibus Virginiae et Carolinae.
Montagnes de Virginie et de Caroline.
Delphinium exaltatum Att.

Delphinium tricorne Michx.
in altissimis montibus Carolinae.
Tres hautes Mont.

Delphinium azureum Michx.
Delphinium carolinianum Walt.

Anemone parviflora Michx.
ad amnes in sinum Hudsonis defluentes.
Rivierre des Goelands.

39

69:5

69:8

69:12

69:19

69:20

71:17

1:320

2:2

2:2

Rhodora [Vol. 86

Anemone aconitifolia Michx.

in Canada et Nova Anglia.
Fleuve St. Laurent, Connecticut.
Anemone canadensis L.

Ranunculus filiformis Michx.

ad ripas fluminis S. Laurentii et sinum Hunsonis.

ad ripas fluminis Hudson, St. Laurent, rives dur Lac Mistassin.
Ranunculus flammula var. filiformis (Michx.) DC.

Ranunculus hispidus Michx.

in umbrosis sylvis Carolinae inferioris.

in humides Virginiae Carolinae.
Ranunculus hispidus Michx. var. hispidus

Thalictrum anemenoides Michx.

a Canada ad Virginianum.

Depuis le Canada jusque en Caroline en suivant la chaine
des Mos,

Thalictrum thalictroides (L.) Eames & Boiv.

Thalictrum laevigatum Michx.
Thalictrum dioicum L.

Cimifuga americana Michx.

Cimifuga palmata Michx.
Trautvettaria carolinensis (Walt.) Vail var. carolinensis

Ilicium parviflorum Michx.
Magnolia macrophylla Michx.

Orchidocarpum grandiflorum Michx.
in Georgia et Florida.

Georgie.

Asimina speciosa Nash*

Orchidocarpum parviflorum Michx.
Asimina parviflora (Michx.) Dunal

Orchidocarpum arientinum Michx.

a Virginia ad Floridam, juxta inundatus amnium ripas.
Pensylvania Virginie Carolines Ohio et Misissipi (river - auth.)
Asimina triloba (L.) Dunal

MICHAUX VOL. II

Mentha tenuis Michx.
Mentha spicata L.

Mentha borealis Michx.
juxta manes ad sinum Hudsonis defluentes.

71:20

72:3

72:7

70:3
70:4

73:2
73:11
73:20

73:21

74:1

74:15

74:16

1984] Uttal — Michaux localities 41

M: 48 deg de latitude. Hauteur des Terres lieux tres humides.
Mentha arvensis var. glabrata (Benth.) Fern.

2:4 Isanthus coeruleus Michx. 74:18
T: in cretaceis Virginiae, Carolinae, Kentucky.
M: Kentucky.

Isanthus brachiatus (L.) BSP var, brachiatus*
2:4 Lamium hispidulum Michx. 74:19
T: in sylvis Tennassée apacis.
M: Kentucky et foréts humides de Tennassee.

Synandra hispidula (Michx.) Britt.

2:4 Stachys hyssopifolia Michx. 74:20
Stachys hyssopifolia Michx. var. hyssopifolia

2:5 Stachys aspera Michx. 75:4

2:6 Brachystemum verticillatum Michx. 75:6
T: in montibus Pensylvaniae, adusque Carolinam superiorem.
M: Carolines, Pittsburgh (Pa. - auth.)

Pycnanthemum verticillatum (Michx.) Pers.

2:6 Brachystemum muticum Michx. 75:9
Pycnanthemum muticum (Michx.) Pers.

2:8 Pycnanthemum aristatum Michx. 75:20
T: a Marylandia ad Carolinam superiorem.
M: Carolina.

Pycnanthemum setosum Nutt.

2:8 Pycnanthemum montanum Michx. 76:1
T: in altis montibus Carolinae.
M: hautes Montagnes.

2:8 Pycnanthemum monardella Michx. 76:2
T: in altis montibus Carolinae.
M: in excelsis montibus Carolina Septentrionalis.

Monarda clinopodia L.

2:9. Thymus carolinanus Michx. 76:8
Satureja georgiana (Harper) Ahles

2:11 Scutellaria parvula Michx. 76:17
T: in regione Illinoensis et Canada.
M: Montreal - Illinois.

2:11 Scutellaria pilosa Michx. 76:19
Scutellaria elliptica Muhl. var. elliptica

2:13. Verbena bracteosa Michx. 76:19
T: in regione Illinoensi et in urbe Nash-ville.
M: Nashville sur Tennassée, St. Vincent, Kaskaskia, etc.
Verbena bracteata Lag. & Rodr.

42

2:14

2:22

Rhodora

Verbena rigens Michx.
in regione IIlinoensi.
Kaskaski, Illinois.

Scutellaria pilosa Michx.
Scutellaria elliptica Muhl. var. elliptica

Verbena bracteosa Michx.

in regione Illinoensi et in urbe Nash-ville.

Nashville sur Tennassée, St. Vincent, Kaskaskia, etc.
Verbena bracteata Lag. & Rodr.

Verbena rigens Michx.
in regione Illinoensi.
Kaskaski, Illinois.
Verbena stricta Vent.

Verbena angustifolia Michx.

in Tennessee et in comitatu Carlisle Pensylvaniae.
Cumberland et haute Pensylvaniae & western Territories.
Verbena simplex Lehm.

Melampyrum americanum Michx.

a sinu Hudsonis ad montosum Carolinam.

Canada (Tadoussack-auth., from isotypes). Hautes
montagnes de Caroline.

Melampyrum lineare var. americanum (Michx.) Beauv.

Pedicularis gladiata Michx.
Pedicularis canadensis L.

Pedicularis lanceolata Michx.
Gerardia auriculata Michx.

Gerardia auriculata Michx.

in pratis regionis Illinoensis.

Prairies des Illinois & du Misissipi.
Tomanthera auriculata (Michx.) Raf.

Gerardia afzelia Michx.

in sabuletis aridis Carolinae.

Basse Caroline.

Seymeria cassioides (Walt. ex J. F. Gmel.) Blake

Capraria multifida Michx.

in ripis arenosis fluminum amniculorumque in Tennassée
et Illinoensi regione.

Illinois, Kentucky.

Leucospora multifida (Michx.) Nutt.

Monniera cuneifolia Michx.
in locis mari inundatis Carolinae inferiores.

[Vol. 86

77:17

76:19

77:17

77:17

77:13

17:21

78:17

79:7

79:8

Uttal — Michaux localities

Lieux aquatiq inondeés par la Mer pres Charleston.
Bacopa monnieri (L.) Wettst.

Monniera rotundifolia Michx.
Bacopa rotundifolia (Michx.) Wettst.

Monniera amplexicaulis Michx.
in fossis, stagnis Carolinae.

in stagnis Carolinae.

Bacopa carolina (Walt.) Robins.

Ruellia humistrata Michx.

ad fines Georgiae et Floridae.

sur la rivierrre Ste. Marie en Georgie.

Ruellia caroliniensis (Walt. ex J. F. Gmel.) Stued,
var. caroliniensis

Ruellia oblongifolia Michx.

in Georgia.

Georgie, le 27 Avril (1787 - auth.)
Dyschoriste oblongifolia (Michx.) Kuntze

Cochlearis humifusa Michx.
Coronopus didymus (L.) J. E. Sm.

Draba arabisans Michx.

in rupibus ripariis ad lacum Champlain et in Nova Anglia.

Rochers du Lac Champlain.

Draba hispidula Michx.
Draba reptans (Lam.) Fern. var. reptans

Cardamine rotundifolia Michx.
in rivulis altissimosum montium Carolinae.
Tres hautes montagnes - Ruisseaux.

Cardamine uniflora Michx.

in rupibus circa Knoxville.

Rochers environs Knoxville.
Leavenworthia uniflora (Michx.) Britt.

Cardamine spathulata Michx.
in excelsis montibus Carolinae.
Hautes Mont

Arabis lyrata L. var. lyrata

Cardamine teres Michx.

in Nova Anglia.

Etat de Vermont Lac Champlain.

Rorippa teres (Michx.) Stuckey var. teres

(A mistake in type locality; limited to southeastern
coastal plain of U.S.; another variety in southwestern
U.S. - auth.)

43

79:9

79:10

79:17

79:16

80:12

80:15

80:16

80:17

80:18

80:19

80:20

2:20

2:50

Rhodora

Dentaria diphylla Michx.

in sylvis vastis Tennassée et montibus altissimus
Carolinae.

New York et Hautes Mont.

Cardamine diphylla (Michx.) Wood

Dentaria concatenata Michx.
Cardamine concatenata (Michx.) O. Schw.

Hesperis pinnatifida Michx.

in humidis sylvanum Tennassée.

Lieux tres humides fdrets de Tennessee.
Todanthus pinnatifidus (Michx.) Steud.

Arabis falcata Michx.

in rupestibus Canadae et Novae Angliae ad Virginianum.
Connecticut, Etat de Vermont, Lac Champlain.

Arabis canadensis L.

Sisvrinchium mucronatum Michx.
Pensylvania.
Mistassin - in Penslvania.

Halesia parviflora Michx.
in Florida, circa Matanga.
Floride.

Styrax pulverulentum Michx.
Stvrax americana Lam.

Malachodendron ovatum Michx.

in montosis a mare remotis Caroline.
Haute Caroline Catawba river.
Stewartia malacodendron L.

Sida alcaeoides Michx.

in glareosis Kentucky & Tennessee.

Barren Oaks endica de Big Barren River Kentucky.
Callirhoe alcaeoides (Michx.) A. Gray

Hibiscus grandiflorus Michx.

in maritimis Georgiae et Floridae et in regione Natchez
ad Mississipi.

Lieux maritimes.

Hibiscus hastatus Michx.

ad ripas fluviorum Ohio, Mississipi et amnium Carolinae.

Sur les Isles et les rives de la Rivierre Santee in
Carolinae de Juillet 1790.
Hibiscus militaris Cav.*

Petalostemum violaceum Michx.
Petalostemum purpureum (Vent.) Rydb.*

[Vol. 86

81:1

81:13

82:14

82:19

84:9

84:6

84:17

84:20

1984]

2:64

NX
<3$8

Uttal — Michaux localities 45

Petalostemum carneum Michx. * 84:19
in Georgia et Florida.
Lieux humides prest MidWest riv. en Floride.

Fumaria recta Michx. 85:2
ad fines Canadae et in montibus Pensylvaniae.

Montagnes pres Lasingburgh.

Adlumia fungosa (Ait.) Greene

Polygala setacea Michx. 85:5
in Carolina septentrionali.

Same as text. (The species is presently unknown in

North Carolina; it is confined to the coastal plain from

Georgia to Mississippi - auth.)

Polygala corymbosa Michx. 85:6
Polygala cymosa Walt.

Polygala uniflora Michx. 85:16
Polygala paucifolia Willd.

Dalea linnaei Michx. 86:1
none given

Kaskaskia.

Dalea alopecuroides Willd.*

Psoralea lupinellus Michx. 86:3
in Carolinae locis arides.

Carolines.

Psoralea canescens Michx. 86:4
Psoralea melilotoides Michx. 86:5
Psoralea psoralioides (Walt.) Cronq. var. psoralioides

Trifolium carolinianum Michx. 86:10
Phaseolus trilobus Michx. 86:14
in Carolina

(locality ms. faded; illegible - auth.)
Strophostyles helveola (L.) EIl.

Galactia glabella Mich. 86:21
in Carolina et Georgia.

Carolina.

Galactia volubilis (L.) Britt. var. volubilis

Amorpha pumila Michx. 87:8
Amorpha herbacea Walt.

Astragalus glaber Michx. 88:12
in sabulosis arides Georgiae.

Georgie lieux arides.

Astragalus michauxii (Luntze) Hermann

46

2:70

2:71

2:77

Rhodora [Vol. 86

Astragalus villosus Michx.

Galega villosa Michx.

a Carolina ad Floridam.

Caroline et Floride.

Tephrosia spicata (Walt.) T. & G.

Galega hispidula Michx.
Tephrosia hispidula (Michx.) Pers.

Vicia parviflora Michx.

in montibus altis Carolinae et Virginiae.
Montagnes de |’Am. sept.

Vicia caroliniana Walt.

Lespedeza sessiliflora Michx.
Lespedeza virginica (L.) Britt.

Lespedeza polystachya Michx.
Lespedeza hirta (L.) Hornem. ssp. hirta

Lespedeza capitata Michx.
Lespedeza procumbens Michx.

Hedysarum alpinum var. americanum Michx.*

in borealibus Canadae et in calaractis montium Alleghanis
(not found in the Alleghany Mountains - auth.)

Lac St. Jean et Mistassins.

Hedysarum bracteosum Michx.
Desmodium cuspidatum (Muhl. ex Willd.) DC ex Loudon var.
cuspidatum

Hedysarum glabellum Michx.

in Carolina inferiore

Dans herbois Basse Caroline
Desmodium glabellum (Michx.) DC.

Hedysarum acuminatum Michx.
Desmodium cuspidatum (Muhl. ex Willd.) DC ex Loudon var.
cuspidatum

Aeschymomene viscidula Michx.

Stylosanthes hispida Michx.

in Virginia et Carolina.

Georgie.

Stylosanthes biflora (L.) BSP var. biflora

Zornia tetraphylla Michx.
Zornia bracteata Walt. ex J. F. Gmel.

Ascyrum amplexicaule Michx.
Hypericum crux-andreae (L.) Crantz*

88:14
88:19

89:1

89:10

89:13

89:14

89:la
89:20
90:4

90:6

90:7

90:8

90:21
91:1

91:2

91:8

2:78

2:79

Uttal — Michaux localities

Ascyrum stans Michx.
Hypericum crux-andreae (L.) Crantz*

Ascyrum multicaule Michx.
Hypericum stragalum P. Adams & Robson*

Ascyrum pumilum Michx.
Hypericum suffruticosum P. Adams & Robson

Hypericum nudiflorum Michx.
in Carolina.

Goose Creek & Garet Place.
Hypericum cistifolium Lam.

Hypericum sphaerocarpum Michx.

in Kentucky.

Route de Louisville.

Hypericum sphaerocarpum Michx. var. sphaerocarpum

Hypericum frondosum Michx.

Hypericum macrocarpon Michx.
in Canada, circa Montreal.
Montreal.

Hypericum pyramidatum Ait.

Hypericum glaucum Michx.
Hypericum myrtifolium Lam.

Hypericum sarothra Michx.
Hypericum gentianoides (L.) BSP.

Hypericum axillare Michx.
Hypericum galioides Lam. var. galioides

Hypericum angulosum Michx.

in paludosis Carolinae.

in humides Carolina ad Floridam.

Hypericum denticulatum Walt. var. denticulatum

Prenanthes racemosa Michx.

in septentrionalibus Canada.

Lac St. Jean.

Prenanthes racemosa Michx. var. racemosa

Prenanthes virgata Michx.
in Virginia et Carolina.
Basse Caroline.

Prenanthes autumnalis Walt.

Prenanthes crepidinea Michx.

in regione Illinoensi et in excelsis montibus Carolinae.
(Not in mountains of the Carolinas - auth.)

none given

47

91:9

91:10

91:11

91:13

91:15

91:16
91:18

91:21

92:2

92:3

92:6

92:11

92:13

92:17

48

Rhodora

Lactuca longifolia Michx.

in Carolina superiore.

in Carolina environs de Burke.

Lactuca canadensis var. longifolia (Michx.) Farw.

Lactuca graminifolia Michx.*
in Carolina inferiore.
Floride et Basse Caroline.

Hieracium canadense Michx.

in Canada.

Lacs Mistassins.

Hieracium canadense Michx. var. canadense

Hieracium scabrum Michx.

in septentrionalibus Canadae et in excelsis montibus
Carolinae.

Lacs Mistassins.

Hieracium canadense Michx. var. scabrum

Hyoseris amplexicaulis Michx.

in Pensylvania, Tennassee et Carolina.
Knoxville.

Krigia biflora (Walt.) Blake

Hyoseris montana Michx.

in altissimis montibus Carolinae septentrionalis.
Sommet des hautes mont. de la Carol. Sept.
Krigia montana (Michx.) Nutt.

Scorzonera pinnatifida Michx.
in ruderatis Carolinae.
in ruderatis Carolinae inferioris.

Pyrrhopappus carolinianus (Walt.) DC. var. carolinianus

Cirsium repandum Michx.

Cirsium muticum Michx.
in altissimis montibus Carolinae.
in excelsis montib. Carolinae.

Liatris pychnostachya Michx.

in pratis Illinoensibus.

Illinois prairies.

Liatris pychnostachya Michx. var. pychnostachya

Liatris bellidifolia Michx.

in sabulosis Carolina septentrionalis.

Sables arides de la Caroline Septentrionale.
Carphephorus bellidifolius (Michx.) T. & G.

Liatris tomentosa Michx.
in Carolina septentrionali.

[Vol. 86

92:21

93:1

93:10

93:12

93:19

94:5

94:6

2:94

Uttal — Michaux localities 49

Fayetteville dans les bois.
Carphephorus tomentosus (Michx.) T. & G.

Liatris squarrulosa Michx. 94:9
none given
Basse Caroline.

Liatris aspera Michx. 94:10
in regione IIlinoense.

du pays des Illinois.

Liatris aspera Michx. var. aspera

Liatris sphaeroides Michx.

in pratis Illinoensibus et in excelsis montibus Caroline.
Prairies vers Misissipi.

Liatris scariosa (L.) Willd.

Liatris macrostachya Michx. 94:14
in Virginia et Carolina.

Basse Virginie et Carolina.

Liatris spicata (L.) Willd. var. spicata

Liatris cylindracea Michx. 93:21
in pratis sylvisque Illinoensibus.
Priaries et Bois du Misissipi.

Vernonia oligophylla Michx.
var. a verna Michx.
var. B autumnalis Michx.
Vernonia acaulis (Walt.) Gleason

Vernonia angustifolia Michx. 94:17
in aridis apricis sylvarum Carolinae.

Terrains arides et decouverts dans la Basse Caroline.

Vernonia angustifolia Michx. ssp. angustifolia

Vernonia fasciculata Michx. 94:18
Vernonia fasciculata Michx. ssp. fasciculata

Vernonia praealta Michx. 94:19
in Nova Anglia.

in Pensylvania.

Vernonia noveboracensis (L.) Michx.

Sparganophorus verticillatus Michx. 95:3
in inundatus, a Carolina ad Floridam.

paladosa Caroline.

Sclerolepis uniflora (Walt.) BSP.

Eupatorium falcatum Michx. 95:10
ad ripas fluviorum Ohio et Scioto.

Rives de l’Ohio.

Eupatorium purpureum L.*

49

2:100

Rhodora [Vol. 86

Eupatorium serotinum Michx. 95:14
in scirpetis Carolina maritimis.

in scirpetis Carolina et in agris neglectis Illinoensib.

Eupatorium serotinum Michx. var. serotinum

Eupatorium verbenaefolium Michx. 95:20
in humides Carolinae.

in humides Carolinae, in aridis sylvarum Caroline.

Eupatorium pilosum Walt.

Eupatorium connatum Michx. 95:7
a Canada ad Floridam.

Carolines, Virginie, Ilinois, Quebec, etc.

Eupatorium perfoliatum L. var. perfoliatum

Critonia kuhnia Michx. 96:3
none given

Haute Carol. rare dans la Basse.

Kuhnia eupatorioides L. var. eupatorioides*

Chrysocoma capillacea Michx. 96:5
in pascuis, juxta Charlstown et in Floride.

in pascuis juxta Savannah.

Eupatorium capillifolium (Lam.) Small

Crysocoma coronopifolia Michx. 96:6
Eupatorium compositifolium Walt.

Chrysocoma nudata Michx. 96:8
in humidis Carolinae.

Carolines.

Bigelowia nudata (Michx.) DC var. nudata

Personia angustifolia Michx. 96:12
Marshallia trinervia (Walt.) Trel.

Personia latifolia Michx.
Marshallia trinervia (Walt.) Trel.

Personia lanceolata Michx.
Marshallia obovata var. scaposa Channell

Melanthera hastata Michx. 95:15
in Carolina.

Caroline, Virginie.

Melanthera nivea (L.) Small

Aster solidagineus Michx. 96:16
in Virginia et Carolina.
A Charleston ad Montes Carolae. Junio floret.

Aster biflorus Michx. 96:17
Aster radula Ait.

Uttal — Michaux localities

Aster acuminatus Michx.
in Canada.

Canada et Montagnes (of North Carolina - auth.)

Aster uniflorus Michx.

in vastis sphagnosis, juxta lacus Mistassinos.

Lac des Cygnes et Rivierra des Goelands.
Aster nemoralis Ait.

Aster infirmis Michx.

a Canada ad Carolinam, per tractus montium.
in collibus Carolinae Septentrionalis et ad Lineville

Mountaenae Canada.

Aster tortifolius Michx.
in Carolina inferiore.
in Carolina.

Aster villosus Michx.

in pratis Illinoensibus.

Illinois.

Aster ericoides L. var. ericoides

Aster surculosus Michx.

Aster amplexicaulis Michx.

in Carolinae inferioris dumosis.
Basse Caroline.

Aster patens var. gracilis Hook.

Aster argenteus Michx.

in rupibus ripariis fluminis Missouri.
in rupibus ad ripas fluminis Missouri.
Aster sericeus Vent.

Aster diversifolius Michx.
Aster undulatus L.

Aster sparsiflorus Michx.

in Carolina inferiore.

Basse Carol.

Aster dumosus L. var. dumosus

Aster subulatus Michx.
Aster subulatus Michx. var. subulatus

Solidago pauciflosculosa Michx.
Chrysoma pauciflosculosa (Michx.) Greene.

5]

96:19

97:7

97:9

97:12

Rhodora

Solidago lanceolata L. var. minor Michx.
Euthamia minor (Michx.) Greene

Solidago virgata Michx.

in humidis sylvarum Carolinae inferioris.
Basse Caroline in humidis sylvariis.
Solidago stricta Ait.

Solidago retrorsa Michx.

in Carolina inferiore.

Basse Caroline. Bois pres l’habitat. (vic. Charleston -
auth.)

Solidago glomerata Michx.

Senecio lyratus Michx.

in numoribus Carolinae.
Pastures arides pres Charlest.
Senecio glabellus Poir.

Senecio pauperculus Michx.
in Canada, juxta lacus.
Lac St. Jean.

Senecio tomentosus Michx.

in Carolinae loco dicto Flat.-Roc.

Sur un Rocher. Flat Roc or hanging Roc.

(South Carolina, not North Carolina, see note under
Sedum pussillum, 1:276. 6 April 1794 - auth.)

Tussilago integrifolia Michx.
Chaptalia tomentosa Vent.

Doronicum nudicaule Michx.
Arnica acaulis (Walt.) BSP.

Inula gossypina Michx.

in maritimis Carolinae et Floridae.

Lieux arides en Basse Caroline. Fleurit en Septembre.
Chrysopsis gossypina (Michx.) EIl.*

Inula graminifolia Michx.

a Carolina ad Floridam.

Caroline.

Chrysopsis graminifolia (Michx.) Ell. var. graminifolia*

Erigeron divaricatum Michx.
in pratensibus Illinoensibus, prope Kaskaskia.

[Vol. 86

98:10

99:8

99:10

99:12
99:13

99:16

99:19

99:20

99:21

100:3

100:4

100:5

Kaskaskia, Prairie du Rocher et Kaokia (aux Pays du Illinois).

Conyza ramosissima Cronq.

Erigeron pulchellum Michx.
in Canada, Pensylvania et montibus Carolinae.

Uttal — Michaux localities

Canada et Montagn de la Caroline Septentri.
Erigeron pulchellus Michx. var. pulchellus

Erigeron hyssopifolium Michx.

ad sinum Hudsonis et juxta lacus Mistassins.
Lacs Mistassins.

Erigeron hyssopifolius Michx. var. hyssopifolius

Erigeron nudicaule Michx.

in udis herbosis Carolinae.

in humidis sylvarum Carolina inferioris.
Erigeron vernus (L.) T. & G.

Baccharis sessiflora Michx.
Baccharis halimifolia L.

Baccharis angustifolia Michx.

Conyza marilandica Michx.
a Pensylvania ad Carolinam.
Virgin et Carol.

Pluchea camphorata (L.) DC.

Conyza amplexicaulis Michx.

in humidis Carolinae.

Basse Caroline - Lieux tres humides.
Pluchea foetida (L.) DC.

Conyza pycnostachya Michx.

a Carolina ad Floridam.

Carolina.

Pterocaulon pycnostachyum (Michx.) Ell. (P. virgatum
(L.) DC. listed for this species in NLSPN (1982) is a

different species according to Cronquist (1980) - auth.)

Gnaphalium polycephalum Michx.
Gnaphalium purpureum L. var. purpureum

Artemisia caudata Michx.

ad ripas sabulosis fluminis Missouri
Bords Sablonneux du Misissipi.
Artemisia caudata Michx. var caudata

Artemisia campestris Michx.
in arena mobili secus lacus ad sinum Hudsonis.
in arena mobili lacum ad sinum hudsonis.

Artemisia campestris ssp. borealis (Pall.) Hall & Clements

Eclipta brachypoda Michx.
Eclipta alba (L.) Hassk.

Bellis integrifolia Michx.
ad ripas rivulorum et in collibus umbrosis Tennassee.

53

100:18

100:19

101:1

101:8

101:9

101:12

102:1

54

2:134

= =

x3

2:135

2:140

Rhodora

Cumberland.
Astranthium integrifolium (Michx.) Nutt. var. integrifolium

Spilanthes repens Michx.
Spilanthes americana (Mutis) Hieron. var. americana

Verbesina sigesbeckia Michx.
Verbesina occidentalis (L.) Walt.

Verbesina coreopsis Michx.
var. a lutea Michx.

in Virginia et Carolina superiore.

in Virginia et Carolina superiore et in regione Illinoensium.
var. B alba Michx.

in Carolina maritima.

in Carolina maritima, Georgia in umbrosis ripariis amnium.

Verbesina alternifolia (L.) Britt.

Verbesina helianthoides Michx.

in occidentalibus Alleghanis, territoria Tennassée, et regione
Illinoens.

Illinois & Wilderness entre Cumberland & Kentucky.

Bidens chrysanthemoides Michx.

a Pensylvania ad Carolinam, in aquosis.
Carol.

Bidens laevis (L.) BSP.

Coreopsis mitis Michx.
Bidens coronata (L.) Britt. var. coronata

Coreopsis senifolia Michx.
Coreopsis major Walt.

Coreopsis latifolia Michx.
in excelsis montibus Carolinae.
Vers le sommet de Black Mountain.

Coreopsis dichotoma Michx.

in sphagnosis umbrosis Carolinae.
Basse Caroline, lieux tres humides.
Coreopsis gladiata Walt. var. gladiata

Coreopsis aristosa Michx.
Bidens aristosa (Michx.) Britt. var. aristosa

Coreopsis trichosperma Michx.

in humidis Carolinae superioris.
Montagnes de Carol.

Bidens coronata (L.) Britt. var. coronata

Helianthus canescens Michx.
in pratensibus irrigris regionis Illinoensis et Tennassee.

[Vol. 86

102:2

102:15

102:16

102:17

103:1

103:6

103:8

103:9

103:17

103:21

104:2

104:9

Uttal — Michaux localities aS

Illinois.
Helianthus mollis Lam.

Helianthus tomentosus Michx. 104:10
in pratensibus IIlinoensibus.

Illinois.

Helianthus strumosis L.

Galardia fimbriata Michx. 104:13
in paludosis aprocis, a Carolina ad Floridam.

Lieux humides en Caroline et en Georgie.

Helenium drummondii H. Rock

Galardia lanceolata Michx.
Gaillardia aestivalis (Walt.) H. Rock

Rudbeckia chrysomela Michx. 104:16
a Pensylvania ad Carolinam.

Pensylvania, Illinois.

Rudbeckia fulgida Ait. var. fulgida

Rudbeckia pinnata Michx. 104:17
in apricis Canadae et in collibus regionis Illinoensis.

regionis Illinoensis juxta Wabash. (A Canadian specimen,

juxta Batiscan ad versus les Trois Rivierres, 104:18, is

a paratype - auth.)

Rudbeckia spathulata Michx. 105:1
Rudbeckia fulgida Ait. var. fulgida

Silphium integrifolium Michx. 105:6
Silphium integrifolium Michx. var. integrifolium

Silphium compositum Michx.

Silphium ternifolium Michx. 105:14
in montibus Virginiae et Carolinae.

Montagnes des Carolines.

Silphium trifoliatum L. var. trifoliatum

Silphium pumilum Michx. 105:18
in Florida.

in Georgia et florida.

Berlandiera pumila (Michx.) Nutt.

Viola hastata Michx. 106:15
in altis montibus Carolinae.

A l’ouest de la Caroline dans les Montagnes elevées des

Appalaches.

Viola debilis Michx. 106:20
in montibus Alleghanis.

in montib. altis America Septentrion.

Viola striata Ait.

56

tN
an
oO

2:160

Rhodora

Viola rotundifolia Michx.

Lobelia claytoniana Michx.

a Pensylvania ad Carolinam.
Pensylvania. Hab in Montibus Carolinae.
Lobelia spicata Lam. var. spicata

Lobelia crassiuscula Michx.

in paludosis herbosis Carolinae maritimae usque ad Floridam.

in paludosis Carolina, Basse Carol.
Lobelia glandulosa Walt.

Lobelia puberula Michx.

in Carolina.

in Carolina amnia maritimis.

Lobelia puberula Michx. var. puberula

Lobelia amoena Michx.

Orchis humilis Michx.

in excelsis montibus Carolinae.
Hautes Montagnes.

Orchis rotundifolia Banks ex Pursh*

Orchis lacera Michx.
Habenaria lacera (Michx.) Lodd*

Orchis cristata Michx.
Habenaria cristata (Michx.) R. Br.*

Orchis clavellata Michx.
Habenaria clavellata (Michx.) Spreng.*

Orchis quinqueseta Michx.
Habenaria lacera (Michx.) R. Br.*

Malaxis unifolia Michx.
in umbrosis sylvarum, a Carolina ad Floridam.
Basse Caroline.

Ophrys pubera Michx.
Liparis loeselii (L.) L. C. Rich.

Limodorum trifidum Michx.
Bletia purpurea (Lam.) DC.

Arethusa parviflora Michx.
Triphora trianthophora (Sw.) Rydb.

Cypripedium canadense Michx.
Cypripedium reginae Walt.

Pistia spathulata Michx.
none given

[Vol. 86

106:21
107:12

107:15

107:16

107:17

107:18

107:19

107:20

107:21

108:3

108:7

108:13

109:3

Uttal — Michaux localities

in rivulis ad Lacus Georgie, Florida.
Pistia stratiotes L.

Podostemon ceratophyllum Michx.

Eriocaulon villosum Michx.
Lachnocaulon anceps (Walt.) Morong

Eriocaulon gnaphalodes Michx.
in Carolina.

in Carolina, Georgia.
Eriocaulon compressum Lam.

Eriocaulon pellucidum Michx.

in septentrionalibus Canada.

Route de Chicoutoumé au Lac St. Jean, Chicoutoumé.
Eriocaulon septangulare With.

Scleria reticularis Michx.
Scleria reticularis Michx. var. reticularis

Scleria oligantha Michx.

Scleria triglomerata Michx.
in Carolina.
in pratis sylvaticus Carolina.

Scleria ciliata Michx.

in Carolina.

in umbrosis aridis Carolinae.
Scleria ciliata Michx. var. ciliata

Carex subulata Michx.

(Erroneously attributed to Canada by both Richard and
Michaux; normal range Rhode Island to Georgia - auth.)
Carex collinsii Nutt.

Carex militaris Michx.*

Carex viridula Michx.
in Canada.
in Canada, entre Montreal et les Tres Rivierres.

Carex paupercula Michx.

Carex typhina Michx.
in regione Illinoense.
in humidior sylvarum Illinoensium.

Carex lenticularis Michx.

Carex oligosperma Michx.
in Canada.
ad Lacus Mistassin dictos.

57

109:7
109:9

109:13

109:19

109:14
109:16

109:18

109:20

110:2
110:3

110:4
110:6

110:7
110:8

Rhodora

Carex vulpinoidea Michx.
in Canada et Nova Anglia.
Montreal.

Carex lanuginosa Michx.
Carex lasiocarpa var. americana Fern.

Carex scirpoidea Michx.
ad sinum Hudsonis.
Mistassin.

Carex debilis Michx.

none given

Basse Caroline?

Carex debilis Michx. var. debilis

Carex triceps Michx.

in Carolina.

in Carolina inferiore.

Carex complanata Torr. & Hook.

Carex striatula Michx.

Tragia cordata Michx.
in Kentucky.
entre le ville et Piercetown, a moitis de distance

(Danville, 22 miles en route to Bardstown, 15 Sept. 1793,

from Michx. journal - auth.)

Tragia urticifolia Michx.
in Georgia.
(no locality given - auth.) le 15 May 1784.

Pachysandra procumbens Michx.

Betula glandulosa Michx.
Betula glandulosa Michx. var. glandulosa

Ambrosia absynthifolia Michx.
Ambrosia artemisiifolia L. var. artemisiifolia

Ambrosia bidentata Michx.
in regione IIlinoensi.
Priaries des Illinois.

Ambrosia paniculata Michx.
Ambrosia artemisiifolia L. var. artemisiifolia

Crotonopsis linearis Michx.

in maritimis Carolinae, juxta Long-Bay, et in regione

Illinoensi.
Priaries des Illinois.

[Vol. 86

110:9

110:12

110:13

Hd:1

111:2

111:3
111:4

111:7

111:8
111:19

112:16

112:19

113:3

Uttal — Michaux localities

Calla sagittifolia Michx.
Peltandra sagittifolia (Michx.) Morong

Sparganium angustifolium Michx.
in amnibus Canadae.
in amnibus a Quebec ad Lacus Mistassins.

Sagittaria graminea Michx.
Sagittaria graminea Michx. var. graminea

Sagittaria natans Michx.
Sagittaria stagnorum Small

Myriophyllum heterophyllum Michx.
in Carolina et Georgia.
in Amne Santee (et) in amne ogechee

Myriophyllum scabratum Michx.
Myriophyllum pinnatum (Walt.) BSP.

Juglans pterocarpa Michx.

in umbrosis humidis sylvarum, juxta mare Caspium.
in humidis umbrosis juxta Angely ad mare Caspium. In

Insula senseli.
Pterocarya fraxinofolia Lam.

(Apparently inexplicable inclusion of a Eurasian species

in F. b.-a. by Richard - auth.)

Juglans mucronata Michx.

in humidis sylvarum Virginiae, Carolinae.
Pensylvania, Virginie et Caroline.

Carya lacinosa (Michx. f.) Loudon

Juncus olivaeformis Michx.
in regione Illinoensium.

Clarksville vis a vise Louisville au Kentucky, 1793.
(Indiana side of Kentucky River, in Clark County - auth.)

Carya illinoensis (Want.) K. Koch

Castanea vesca var. americana Michx.
Castanea dentata (Marsh.) Borkh.

Quercus laurifolia Michx.
in Carolina meridionali et Georgia.
in Carolina.

Quercus catesbaei Michx.

in Marilandia, Virginia et Carolinis.
in aridis, sabulosis Carolinae.
Quercus nigra L.

59

113:7

113:14

113:15

Bp lz

113:19

113:20

114:4

115:3

115:7

115:13

115:19

115:20

2:199

Rhodora

Quercus triloba Michx.

a Nova Anglia ad Georgiam.
Carol. in Pensylvania.

Quercus falcata Michx. var. falcata

Quercus falcata Michx.

a Virginia ad Floridam.

Haute et Basse Caroline.

Quercus falcata Michx. var. falcata

Quercus cinerea Michx.
in montibus Alleghanis.
in montibus Pensylvaniae et in regione Illinoensium.

Quercus obtusiloba Michx.

a Canada et Nova Anglia ad Floridam.
Pensylvania.

Quercus stellata Wang. var. stellata.

Quercus macrocarpa Michx.

in montibus Alleghanis, Kentucky, Tennassée, regione
Illinoense, ete.

Illinois, Cumberland et Kentucky.

Quercus imbricaria Michx.

Quercus banisteri Michx.
Quercus ilicifolia Wang.

Pinus mitis Michx.

in Virginia, Kentucky et Carolina.
in Carolina.

Pinus echinata Mill.

Pinus serotina Michx.

Abies denticulata Michx.

in Canada et Nova Anglia.

Baye d’Hudson et Lacs Mistassins. In monte excelsiore
Carolinae dicto Grand Father Mountain et in Canada
(The latter sentence is partly wrong as it is evident
Michaux confused Picea rubens Sarg. in the south with
this species. On his label he has “Epinette noire

des Canadiens” - auth.)

Picea mariana (Mill.) BSP.

Suillingia ligustrina Michx.
Sebastiana fruticosa (Bartr.) Fern.

Euphorbia graminifolia Michx.

in maritimis Floridae et Georgiae.

Parites maritimes et les plus meriodionales de la Georgie.
Euphorbia heterophylla var. graminifolia (Michx.) Engelm.

[Vol. 86

116:10

116:12

117:12

LPF 1S:

117:21

119:]

119:5
119:7

119:17

119:19

1984]

2215

2:214

2:214

2:215

Uttal — Michaux localities

Croton monanthogynum Michx.
Croton monanthogynus Michx.

Croton disjunctiflorum Michx.
Croton punctatus Jacq.

Croton capitatum Michx.
Croton capitatus Michx. var. capitatus

Croton argyranthemum Michx.
Croton argyranthemus Michx.

Najas canadensis Michx.
Najas flexilis (Willd.) Rostk. & Schmidt

Vallisneria americana Michx.

Ceratiola ericoides Michx.
in aridis sabulosis Georgiae et Floridae.
in Georgia, Florida.

Adelia ligustrina Michx.

in fruticetis Illinoensibus, Tennassée, etc.

in fruticetis Illinoensium et Colligus Tennassee juxta
Nashville.

Forestiera acuminata (Michx.) Poir. in Lam.

Adelia acuminata Michx.
Forestiera acuminata (Michx.) Poir. in Lam.

Adelia porulosa Michx.
Forestiera segregata (Jacq.) Krug & Urban

Salix eriocephala Michx.
Salix discolor Muhl. var. discolor

Salix cordata Michx.

Salix carolina Michx.
in Carolina et Georgia.
Caroline.

Salix incana Michx.
in Canada, juxta lacum S. Joannis.

in Canada, juxta lacum S. Joannis et Riv des Goelands.

Salix candida Fluegge ex Willd. var. candida

Salix longirostris Michx.

a Nova Anglia ad Carolinam, copiose in demetosis apricis

Tennassee.
a Canada ad Carolinam.

Ilex canadensis Michx.

in praeruptis montibus, a sinu Hudsonis ad lacum Champlain.
Quebec, Tadoussack, Lac St. Jean, Mistassins dans les

61

120:7

120:8

120:11

120:12

120:19

120:20

120:21

121:2

121:3

121:4

121:6

121:9
121:10

121:13

121:14

121:13

62

Rhodora [Vol. 86

lieux humid. et fert.
Nemopanthus mucronatus (L.) Trel.

Vitis aestivalis Michx. 122:17
in sylvis, a Virginia ad Carolinam.

a pensylvania ad Carolinam.

Vitis aestivalis Michx. var. aestivalis

Vitis riparia Michx. 122:20
ad ripas et in inusulis fluviorum Ohio, Misissipi, etc.

idem, plus Missouri.

Vitis riparia Michx. var. riparia

Vitis rotundifolia Michx. 122:21

Vitis cordifolia Michx. 123:3
a Pensylvania ad Floridam.

a Nova Anglia ad Carolinam.

Vitis vulpina L.

Pyrularia pubera Michx. 123:11
in montibus Carolinae occidentalis et in Virginia.
in Montium excelsorum umbrosis Virginia, Carolinae.

Acnida nisocarpa Michx. 123:13
none given

Carolina.

Amaranthus cannabinus (L.) Sauer

Zanthoxylum ramiflorum Michx. 123:14
a Canada ad Virginiam et in Kentucky.

in Canada juxta Montreal et in Noveboraco ad riparias

fluvii Hudsonis. Distr Canada, Pensylvania, Kentucky,

Ilinoese et raro in Carolina.

Zanthoxylum americanum Mill.

Zanthoxylum tricarpum Michx. 123:15
Zanthoxylum clava-herculis L.

Prinos ambiguus Michx. 123:18
Ilex ambigua (Michx.) Torr.

Prinos gronovil Michx. 123:19
Ilex verticillata (L.) A. Gray

Smilax pulverulenta Michx. 124:4
in Canada et Pensylvania.

a Canada ad Virginiam.

Smilax herbacea var. pulverulenta (Michx.) A. Gray*

Smilax tamnifolia Michx.* 124:5

Smilax pubera Michx. 124:13
Smilax pumila Walt.

Uttal — Michaux localities 63

Dioscorea paniculata Michx. 124:14
a Canada ad Carolinam.

N. Jersey.

Dioscorea villosa L. var. villosa

Polygonella parvifolia Michx. 124:17
in aridissimis Carolinae.

Sables arides aux environs de Wilmington dans la Caroline
Septentrionale.

Polygonella polygama (Vent.) Engelm. & Gray

Gymnocladus canadensis Michx. 124:19
none given

in regione Illinoensium.

Gymnocladus dioicus (L.) K. Koch

Populus angulosa Michx. 125:7
Populus deltoides Marsh. ssp. deltoides

Populus tremuloides Michx.* 125:12
Populus grandidentata Michx. 125:13
Planera gmelini Michx. 126:9

ad ripas fluviorum Ohio, Misissipi et Carolinae.
Caroline et Georgie.
Planera aquatica Walt. ex J. F. Gmel.

Veratrum parviflorum Michx.* 126:16
in excelsis montibus Carolinae.
Tres hautes montannes.

Melanthium racemosum Michx. 126:19
in excelsis montibus Carolinae septentrionalis.

Yellow Mountain.

Melanthium virginicum L.

Acer eeriocarpum Michx. 127:2
none given

Rives de l’Ohio

Acer saccharinum L.

Mimosa glandulosa Michx. 127:14
ad ripas fluvii Tennassée; etrain juxta Misissipi.

Isles du Misissipi et Rivierre Tenassée.

Desmanthus illinoensis (Michx.) MacMill. ex B. Rob. & Fern.

Mimosa horridula Michx. 127:11
Schrankia microphylla (Dryand.) Macbr.

Fraxinus epiptera Michx. 128:3
in Virginia et Carolina.
Il commence a 15 li. (French abbreviation of “league”,

64

Rhodora

approximately 3 miles - auth.) au dessus du Lac St Jean.

Fraxinus americana L. var. americana
Fraxinus quadrangulata Michx.

Fraxinus platicarpa Michx.
Fraxinus caroliniana Mill.

Nyssa candicans Michx.

ad amnens Ogeeche.

Rivierre Ogeeche.

Nyssa ogechee Bartr. ex Marsh.

Nyssa villosa Michx.

a Nova Anglia ad Carolinam.

a Connecticut ad Carolinam.

Nyssa sylvatica Marsh. var. sy/vatica

Nyssa tomentosa Michx.

ad amnem St. Mary et in Florida.
Riv. Altahama, St. Mary et Floride.
Nyssa aquatica L.

Nyssa angulisans Michx.
Nyssa aquatica L.

Vittaria angustifrons Michx,

in Florida, juxta amnen Aisa-hatcha.

Sur les Bords de la Riv. Aisahatcha le I Avril (1787 -
auth.) Floride.

Vittaria lineata (L.) Sm.

Preris gracilis Michx.
Cryptogramma Stelleri (S. G. Gmel.) Prantl

Woodwardia banisteriana Michx.
Osmunda cinnamomea L.

Polypodium hexagonopterum Michx.
Thelypteris hexagonoptera (Michx.) Weatherby

Nephrodium lanosum Michx.

in montibus saxosis Tennassee et Carolinae septentrionalis.

in excelsis - then same as T.
Cheilanthes lanosa (Michx.) D. C. Eaton

Nephrondium ruficulum Michx.

in rupibus Canada et Nova Caesareae.

in rupibus Canada, Nova Angliae et Nova Caesareae.
Cheilanthes lanosa (Michx.) D. C. Eaton

Polypodium connectile Michx.
Thelypteris hexagonoptera (Michx.) Weatherby

Nephrodium tenue Michx.

[Vol. 86

128:6
127:16

128:21

129:4

129:8

129:10

129:13

129:16

130:16

130:17

130:4

130:5

130:6

1984]

Uttal — Michaux localities

in Canada.
Quebec.
Cystopteris fragilis (L.) Bernh.

Nephrodium asplenioides Michx.
Athyrium asplenioides (Michx.) A. A. Eaton*

Nephrodium punctilobum Michx.
Dennstaedtia punctiloba (Michx.) Moore

Nephrodium acrostichoides Michx.

in Pennsylvania, Carolina et Tennassée.
same plus et Carolin. maritimis.
Polystichum acrostichoides (Michx.) Schott

Asplenium thelpteroides Michx.

in montibus Virginiae et Carolinae septentrionalis.

same plus Lac Champlain.
Athyrium thelypteroides (Michx.) Desv.

Asplenium angustifolium Michx.
Athyrium pycnocarpon (Spreng.) Tidest.

Ophioglossum bulbosum Michx.

in sabulosis Carolina inferioris.

in sabulosis Carolina.

Ophioglossum crotalophoroides Walt.

Botrypus lunarioides Michx.

in pascuis, circa Charlestown.

in pascuis sabulosis juxta Charleston.
Botrychium lunarioides (Michx.) Sw.

Cteisium paniculatum Michx.

in occidentalibus Virginiae ad fines Kentucky et in

Tennasseée.

Dady’s Creek et plusi. creeks a 25 miles de West Point

SW Clinch River.
Lygodium palmatum (Bernh.) Sw.

Osmunda interrupta Michx.

in Canada, Pensylvania et Kentucky.
Canada, Kentucky.

Osmunda claytoniana L.

Equisetum scirpoides Michx.

Lycopodium lucidulum Michx.
a Canada ad Carolina montosum.
New Jersey.

Lycopodium dendroideum Michx.

a Canada et Nova Anglia ad Carolinam montosam.

Montagne de Carol. Sept.

65

130:9

130:11

130:15

130:1

130:2

131:11

131:9

131:4

132:8
132:11

133:4

66 Rhodora [Vol. 86

LITERATURE CITED

BARKLEY, T. M. 1978. Senecio. N. Am. FI. Ser. I, pt. 10:50-139.

Cronaquist, A. 1980. Vascular Flora of the Southeastern United States. Vol. I.
Asteraceae. Chapel Hill, Univ. of North Carolina Press.

Ewan, J. 1974. “Introduction to Facsimile Edition of Michaux, A., Flora
boreali-americana.” ix - xlvi. New York, Hafner Press.

GLEASON, H. A., AND A. Cronquist. 1963. Manual of Vascular Plants of
Northeastern United States and Adjacent Canada. New York, D. Van Nostrand,
Inc.

Goprrey, R. K., AND J. W. WooTEN. 1979, 1981. Aquatic and Wetland Plants of
Southeastern United States. 2 vols. Athens, Univ, of Georgia Press.

Hitcucock, A. S., AND A. CHASE. 1950. Manual of the Grasses of the United
States. Rev. Ed. Washington, U.S. Government Printing Office.

INTER DOCUMENTATION COMPANY AG. 1967. Microfiche Set of the Herbarium of
A. Michaux, Flora boreali-americana. 6211, 1-145. Zug, Switzerland.

MicHaux, A. 1803. Flora boreali-americana. 2 vols. Paris and Strasbourg.

RapForD, A. E., H. E. AHLES, AND C. R. BELL. 1968. Manual of the Vascular
Flora of the Carolinas. Chapel Hill, Univ. of North Carolina Press.

SARGENT, C. S. 1889. “Portions of the Journal of André Michaux, Botanist,
Written During His Travels in the United States and Canada, 1785 to 1796,
With an Introduction and Explanatory Notes.” Proc. Amer. Philos. Soc.
21:1-146.

STAFLEU, F. A., AND R.S. Cowan. 1981. Taxonomic Literature. Vol. IIL. Lh-O.
Utrecht, Netherlands. International Bureau for Plant Taxonomy and Nomen-
clature.

U.S. Sor, CONSERVATION SERVICE. 1982. National List of Scientific Plant Names.
2 vols. SCS-TP-159. Washington, U.S. Government Printing Office.

DEPARTMENT OF BIOLOGY
VIRGINIA POLYTECHNIC INSTITUTE AND STATE UNIVERSITY
BLACKSBURG, VA 24061

ADDITIONS TO THE FLORA OF CAPE BRETON
HIGHLANDS NATIONAL PARK, NOVA SCOTIA

HAROLD R. HINDS

ABSTRACT

Ten species and one hybrid are here reported as new or from additional sites in
Cape Breton Highlands National Park. The hybrid Solidago bicolor X S. canadensis
is reported for the first time.

Key Words: vascular plants, new records, Cape Breton

While doing field work for a study of the rare plants of Cape
Breton Highlands National Park from August 1980 to August 1981,
the following vascular plants were found that had not previously
been reported from Cape Breton or had only been found once or
twice.

Three species are here reported as new to Nova Scotia and are
marked with an asterisk. The species are listed in alphabetical order
with their collection data and brief distributional notes. The
acronym, CBHNP, refers to the herbarium of Cape Breton
Highlands National Park, Ingonish Beach, Nova Scotia, BOC ILO.

Anemone multifida Poir.

Uncommon on dry loose, coarse, gravelly conglomerate ledges in
small crevices and on shelves, area of Le Buttereau and La Grande
Falaise about | km north of park entrance, Inverness County, 15
June, 1981, H. Hinds 4911 (CBHNP); gravelly bluff north of
Chéticamp, Inverness County, 8 July, 1938, A. S. Pease 26,627
(GH).

This latter collection by A. S. Pease is certainly from the same
area, but was never published. It was misplaced in the Gray
herbarium and so overlooked by previous researchers.

The nearest populations to this station on the mainland are in the
vicinity of Hartland, New Brunswick on gravel strands of islands in
the Saint John River. This species also occurs in Newfoundland
(Rouleau, 1978).

*Anemone parviflora Michx.

A small colony on north-northeast facing wet limestone cliffs
below waterfall, Corney Brook gorge south of French Lake,
Inverness County, 10 June, 1981, H. Hinds 4477 (CBHNP).

67

68 Rhodora [Vol. 86

This species occurs in Newfoundland (Rouleau, 1978) and on the
mainland the nearest population occurs on wet gypsum near Wilson
Brook, Albert Mines, Albert County, New Brunswick.

Diapensia lapponica L.

Rare on horizontal granitic shelf facing north on high shoulder of
upper Chéticamp River gorge above series of waterfalls, Inverness
County, 25 August, 1981, H. Hinds 4541 (CBHNP).

One other collection is known from Nova Scotia in Victoria
County, Lockhart Brook, tributary of Salmon River (Smith &
Erskine, 1954). Not known from New Brunswick. Closest popula-
tions in Newfoundland (Rouleau, 1978) and the Gaspé (Scoggan,
1950).

Draba norvegica Gunner var. clivicola (Fern.) Boivin

Rare on dry shelves of limestone ledges close to upper Corney
Brook south of French Lake, Inverness County, 24 July, 1981, H.
Hinds 4480 (CBHNP).

Previously reported from Cape Breton on dry, exposed shelves of
limestone cliffs, Big Southwest Brook (Smith & Schofield, 1952).
Also reported from Big Intervale, Margaree, Inverness County
(Macoun, 1898). Reported from the Gaspé (Scoggan, 1950) but
from neither New Brunswick nor Nefoundland.

*Drosera rotundifolia L. var. comosa Fern.

Rare on calcareous gravels, east shore of Paquette Lake, Victoria
County, 12 August, 1980, H. Hinds 3818 (CBHNP),.

Not previously reported from Nova Scotia. Nearest populations
are reported from marly bogs at the mouth of Grand River, Gaspe
(Scoggan, 1950) and from Ross Island, Grand Manan archipelago,
New Brunswick (Weatherby & Adams, 1945).

Minuartia groenlandica (Retz.) Ostenf.

Rare on south-southeast facing dryish, loose calcareous ledges,
Corney Brook gorge south of French Lake, Inverness County, 15
August, 1981, H. Hinds 4471 (as Arenaria groenlandica) (CBHNP).

Nearest populations in Newfoundland (Rouleau, 1978) and in
Halifax County, Nova Scotia (Roland & Smith, 1969).

Oxytropis campestris (L.) DC. var. johannensis Fern.

Frequent on open, gravelly scree, seaward side of southeast tip of
promontary north of Jumping Brook, Inverness County, 12 August,
1980, H. Hinds 3205 (CBHNP); rare in rock scree, south shoulder of

1984] Hinds — Cape Breton flora 69

La Grande Falaise about | km north of park entrance, Inverness
County, 10 June, 1981, H. Hinds 4912 (CBHNP).

Reported in Nova Scotia from Saint Paul Island off the northern
tip of Cape Breton (Perry & Roscoe, 1931) and from Cape d’Or in
Cumberland County (Schofield, 1955). This taxon is also found in
Newfoundland (Rouleau, 1978) and in New Brunswick on gravel
strands of the Saint John and Restigouche Rivers, as well as the
Gaspe and northern Maine.

Pinguicula vulgaris L.

Rare on moist south-southeast facing ledges above large water-
fall, upper Corney Brook gorge south of French Lake, Inverness
County, | August, 1981, H. Hinds 4529 (CBHNP); frequent on wet
limestone ledges, upper Chéticamp River, Inverness County, |
August 1981, H. Hinds 4529 (CBHNP).

Also reported in Nova Scotia from Saint Paul Island (Perry &
Roscoe, 1931) and from Big Southwest Brook, Inverness County
(Hounsell & Smith, 1968). This species is found in Newfoundland
(Rouleau, 1978) and on the Restigouche River in Northern New
Brunswick.

*Rhododendron lapponicum (L.) Wahl.

Rare on south facing shelves of calcareous ledge, Corney Brook
gorge south of French Lake, Inverness County, 19 June 1981, H.
Hinds 4302 (CBHNP).

Nearest populations reported from Newfoundland (Rouleau,
1978), the Gaspe (Scoggan, 1950), and from alpine mountain
summits of northern New England (Seymour, 1969). Not known
from New Brunswick.

Salix uva-ursi Pursh

Rare and local on south facing shelves of calcareous ledge,
Corney Brook gorge south of French Lake, Inverness County, 19
June 1981, H. Hinds 4305, 14 July, 1981, H. Hinds 4478 (CBHNP).

Previously reported in Nova Scotia from Saint Paul Island (Perry
& Roscoe, 1931). The nearest other populations occur in Newfound-
land (Rouleau, 1978), the Gaspé (Scoggan, 1950) and the mountains
of New York (Mitchell et a/. 1980) and northern New England
(Seymour, 1969). Not known from New Brunswick.

Saxifraga aizoides L.
Locally abundant on north facing limestone ledges dripping with
water from above and also within the spray of a large waterfall,

70 Rhodora [Vol. 86

Corney Brook gorge south of French Lake, Inverness County, 14
August 1980, H. Hinds 3755 (CBHNP).

Previously reported in Nova Scotia only from the Big Southwest
Brook, Inverness County (Smith & Schofield, 1952). Nearest other
populations occur in Newfoundland (Rouleau, 1978), the Gaspé
(Scoggan, 1950), the mountains of New York (Mitchell et a/. 1980),
and Vermont (Countryman, 1978). Not known from New Bruns-
wick.

Solidago bicolor L. < S. canadensis L.

One large plant on open gravelly roadside along the Cheticamp
River about 2 km above Cheticamp campground, Inverness
County, 11 August 1980, H. Hinds 3828 (CBHNP, UNB).

The hybrid occurred with both parents growing nearby and
combined the characteristics of both. This hybrid has apparently
not previously been reported anywhere.

LITERATURE CITED

COUNTRYMAN, W. D. 1978. Rare and endangered vascular plants in Vermont.
The New England Botanical Club, Cambridge, Massachusetts. 68 pp.

HounNSELL, R. W. AND E. C. SMITH. 1968. Contributions to the flora of Nova
Scotia. 1X: Habitat studies of arctic-alpine and boreal disjunct species. Rhodora
70: 176-191.

Macoun, J. M. 1898. Geological Survey of Canada Annual Report, Nova
Scotia I]: 194A-200A.

MITCHELL, R. S., C. J. SHEVIAK, AND J. K. DEAN. 1980. Rare and endangered
vascular plant species in New York State. New York State Museum, Albany.
38 pp.

Perry, L. M. AND B. Roscoe. 1931. Botanical Explorations of Saint Paul
Island, Nova Scotia. Rhodora 33: 105-126.

ROLAND, A. E. AND E. C. SmitH. 1969. The Flora of Nova Scotia. Nova Scotia
Museum, Halifax, Nova Scotia. 743 pp.

RouLeAu, E. 1978. List of the vascular plants of the Province of Newfoundland
(Canada). Oxen Pond Botanic Park, St. John’s, Newfoundland.

SCHOFIELD, W. B. 1955. Contributions to the Flora of Nova Scotia: V. Results
of exploration in Cumberland County. Rhodora 57: 301-310.

ScoGGan, H. J. 1950. The flora of Bic and the Gaspé Peninsula, Québec.
National Museum of Canada. Bulletin 115.

Seymour, F. C. 1969. The Flora of New England. C. E. Tuttle Co., Rutland,
Vt. 596 pp.

SmitH, E. C. anp D. S. Erskine. 1954. Contributions to the Flora of Nova
Scotia IV. Rhodora 56: 242-252.

1984] Hinds — Cape Breton flora 71

SmitH, E. C. AND W. B. SCHOFIELD. 1952. Contributions to the Flora of Nova
Scotia. Rhodora 54: 220-228.

WEATHERBY, C. A. AND J. ADAMS. 1945. A list of the vascular plants of Grand
Manan, Charlotte County, New Brunswick. Contributions from the Gray
Herbarium of Harvard University, CLVIII.

CONNELL MEMORIAL HERBARIUM,
DEPARTMENT OF BIOLOGY

UNIVERSITY OF NEW BRUNSWICK, FREDERICTON.
NEW BRUNSWICK. E3B 5A3.

STUDIES IN THE GENUS ARISTIDA (GRAMINEAE)
OF THE SOUTHEASTERN UNITED STATES.
I. SPIKELET VARIATION IN A. PURPURESCENS,
A. TENUISPICA, AND A. VIRGATA

KELLY W. ALLRED

ABSTRACT

Morphologic variation in Aristida purpurescens, A. tenuispica, and A. virgata was
assessed by principal components and step-wise discriminant analyses. Although
well-defined clusters corresponding to the three taxa were defined by principal
component analysis (PCA), extensive morphologic overlap was also evident. Both
the PCA and discriminant analyses assigned some spikelets from the same panicle to
different taxa. A single morphologic species is suggested.

Key Words: Aristida, taxonomy, principal components, discriminant analysis

North American Aristida have been classed within three sections:
1) Arthratherum, wherein the lemma is prolonged into a column or
beak that disarticulates at its base, just above the lemma;
2) Streptachne, in which the lateral awns are much reduced or
obsolete and without an articulated column; and 3) Chaetaria, with
well-developed lateral awns and also without an articulated column
(Henrard, 1929). Hitchcock (1924) further divided the section
Chaetaria into four rather informal groups, Dichotomae, Adscen-
siones, Divaricatae, and Purpureae, but left about 25 species, mostly
from the southeastern United States and West Indies, unplaced as to
group.

While working with some of these unplaced species for the
Vascular Flora of the Southeastern United States, I experienced
difficulty in distinguishing some members of the taxa A. pur-
purescens Poir., A. tenuispica Hitche., and A. virgata Trin. These
are perennial, cespitose, three-awn grasses with mostly flat blades,
narrow contracted panicles, and more-or-less equal glumes. Ap-
parently typical forms of each species were relatively distinct from
the others, and from other species as well, but there occurred
numerous specimens with confusing and seemingly random com-
binations of features from the three taxa. This situation prompted
the following queries. What is the pattern of morphologic variation
within this complex? Are the three taxa delimited by consistent
combinations of morphologic features, or does there exist an
admixture of features? How extensive is any morphologic overlap
among the taxa?

73

74 Rhodora [Vol. 86

MATERIALS AND METHODS

Over 400 herbarium specimens from throughout the geographic
range of the three taxa were examined ina preliminary study of the
morphologic features. Of these, 74 specimens were selected that
represented well the morphologic variation within the complex,
including “extremes” as well as “typical” material. These specimens
were assigned to one of four groups: three groups contained typical
purpurescens, tenuispica, or virgata based upon conformity with the
original descriptions and descriptions of the types in Henrard
(1932); the fourth group was composed of unassigned specimens
that were difficult to identify or exhibited unusual character
combinations. Each of the three typical groups contained 16
Operational taxonomic units (OTUs), one OTU from each of 16
specimens. The unassigned group contained 52 OTUs, representing
two OTUs (scorings) from each of 26 specimens. The duplicate
scoring of each of the unassigned specimens would give some
indiction of variability within a single plant. A total of 100 OTUs
was used in the analysis.

Following the survey of all herbarium specimens, and after
consulting the major reference works dealing with this group of
Aristida, 12 spikelet features were selected for analysis. They were
number of nerves on glume I, lengths of glumes I and II, lemma
length, callus length, cnetral and lateral awn lengths, central and
lateral awn angles of divergence, and awn contortion. For each of
the three typical groups of specimens, measurements were taken
from a well-formed spikelet in the upper 4 of the panicle. For the
unassigned group, with 2 OTUs per specimen, measurements were
taken from spikelets at both ends of the central 4 of the panicle.

Analysis was by principal components (PCA) using standardized
data and by step-wise discriminant analysis (SDA) from the BMDP
Statistical Software package (Dixon, 1981). For the SDA, the
typical groups were used to establish the discriminant equation, and
the members of the unassigned group were then given a phenetic
identification to one of the three groups based on this equation.

RESULTS AND DISCUSSION

Principal components analysis resulted ina fairly clear distinction
of the three typical groups of specimens (Figure 1). This, of course,
was the result of the a priori selection of morphologically separable

1984] Allred — Aristida 75

-2 | VIRGATA _ _
= = 7 oO 5 ol ~
(Oo 0 Oo 2 — O_--*s, PURPURESCENS
= an ~ re) ae , ‘
a ee s .
: So % 7”
fo) @) os t) wn 8 at
Q ® ;
E Leena aad pa Z
S TENUISPICA.-"'© @, #89 af
=) ‘s ‘ bate -
rv @ ran) ic es
® 2
3 O@ 20 @
2 ees
1 1 it i
T T T T T
e9 -] 0 1 2
Component |

Figure |. PCA of only the typical groups.

specimens. Greatest resolution was found along axes I and III,
reflecting distinctions in glume and lemma lengths (strongly
weighted on axis I) and lateral awn angle, central awn width, and
awn contortion (strongly weighted on axis III). Axis II reflected a
strong weighting with awn length, but did not distinguish the taxa.
The first three axes accounted for 54% of the variation; axes I and
III for 27% and 12%, respectively. Relative to the other groups, the
purpurescens entity was characterized by long lemmas and glumes,
the first glume longer than the second (inverse), equal awn widths,
and equal divergence of the awns with little or no contortion.
Typical virgata possessed a wide central awn reflexed or divergent
between the two erect lateral awns. Typical renuispica had markedly
contorted awns with nearly equal divergence, and subequal glumes.

Of note was the phenetic overlap of the typical purpurescens and
tenuispica groups. Specimens within the zone of overlap were
rechecked for proper identification: they did seem to fit well the
morphologic “concept” of purpurescens or tenuispica, although
differences in awn contortion were less pronounced than among the
other specimens.

76 Rhodora [Vol. 86

There were few high correlations of features. Correlations greater
than +0.50 were glume I and II lengths, glume and lemma lengths,
central and lateral awn lengths, and awn contortion and lateral awn
angle. These were all less than +0.80, however. The number of
nerves on the first glume, callus length, and central awn angle all
lacked any meaningful correlation with any other features, and
seemed to vary randomly.

Upon addition of the unassigned OTUs to the PCA grid,
distinctions between the three taxa are blurred (Figure 2). Identi-
fication of these OTUs to purpurescens, tenuispica, or virgata was
by SDA. Of the 26 pairs of OTUs in the unassigned group, 12 were
congruous, that is, both OTUs from each pair were assigned to the
same taxon by the SDA and fell roughly within that cluster on the
PCA grid. However, 14 pairs of OTUs were incongruous, with the
OTUs from each pair being assigned to different taxa by the SDA
and with generally a corresponding placement by PCA. These
incongruous pairs are linked by a solid line in Figure 2. The phenetic
similarities of the three taxa are circular, with only slightly more

It
= IT
=
e
a OT
&
°
UO
1+
~_?+ 7 . ‘s
l l l

| iT

T T T T T

-2 | 0 1 2
Component |

Figure 2, PCA of the unassigned OTUs. The dashed lines delimit the boundaries
of typical virgata, typical purpurescens, and typical tenuispica from Figure 1. The
OTUs are assigned to virgata (V), purpurescens (P), or tenuispica (T) by the SDA.
OTUs linked by a solid line are incongruous pairs.

1984] Allred — Aristida 77

incongruous pairs involving virgata and tenuispica (6 pairs) than
virgata-purpurescens or purpurescens-tenuispica (4 pairs each).

No consistent combination of features serves to distinguish the
taxa. The spikelet variability within a single plant, represented by
conformity to one of the typical groups, was just as great for some
specimens as the variability between taxa. The incongruous pairs in
particular had atypical combinations, such as inverse glumes with
strongly contorted awns on the same spikelet, or one spikelet typical
of virgata and one spikelet typical of purpurescens within the same
panicle. Although material typical of each taxon is common,
morphologic intermediates are certainly not rare, and represent a
significant phenetic shuffling of features. Clearly, only one morpho-
logic taxon is present here. The original three entities may deserve
recognition at the infraspecific level, but even this is questioned by
the presence of incongruous pairs within the same panicle. At this
point nothing is known about the chromosome numbers for these
taxa or about the reproductive mechanism. Final disposition of the
taxa must await these further investigations.

ACKNOWLEDGMENTS

Thanks to John Ludwig for comments on an earlier draft, and to
the following herbaria for their generous loans of specimens: APSC,
ARK, AUA, FLAS, FSU, KY, LAF, LSU, MARY, MO, NCSC,
NCU, NLU, NO, SMU, TENN, US, USCH, USF, VDB, VPI,
WVA. This work was supported in part by grants from the New
Mexico Agricultural Experiment Station and is Journal Article
1034.

LITERATURE CITED

Dixon, W. J. ed. 1981. BMDP Statistical Software. University of California
Press. Berkeley. 726 pp.

HENRARD, T. 1929. A Monograph of the Genus Aristida, Vol. 1. Mededeelingen
Van's Rijks Herbarium. No. 58.

1932. A Monograph of the Genus Aristida, Vol. Il. Mededeelingen Van's

Rijks Herbarium. No. 58a.

Hitcucock, A. S. 1924. The North American species of Aristida. Contr. U.S.
Natl. Herb. 22:517-586.

DEPARTMENT OF ANIMAL AND RANGE SCIENCES
BOX 3 I, NEW MEXICO STATE UNIVERSITY
LAS CRUCES, NM 88003

INFRASPECIFIC VARIATION IN GRATIOLA VISCIDULA
PENNELL (SCROPHULARIACEAE)

Davip M. SPOONER

ABSTRACT

Gratiola viscidula Pennell was separated into two subspecies by Pennell (1935): G.
viscidula subsp. viscidula, and G. viscidula subsp. shortii Durand ex Pennell. The
latter subspecies, which was distinguished by its larger corollas, calyces, and leaves,
was only known from southeastern Ohio and from an unspecified site in Kentucky.
Examinations of herbarium specimens and field populations have demonstrated that
the sizes of various plant parts are highly variable, both within individual popula-
tions and throughout the range of the species. Infraspecific categories are unwar-
ranted. The chromosome number of the species is here first reported as n = 7.

Key Words: Gratiola, Scrophulariaceae, chromosome counts

INTRODUCTION

Pennell (1935) separated Gratiola viscidula into two subspecies:
G. viscidula subsp. viscidula [as “G. viscidula typica”] and G.
viscidula subsp. shortii [as “G. viscidula shortii (Durand) Pennell”’].
The latter subspecies was distinguished by its larger corollas,
calyces, and leaves and was known only from two collections, one in
southeastern Ohio (Jackson County) and one from an unspecified
site in Kentucky.

Later workers (Cooperrider, 1976; Cusick and Silberhorn, 1977;
McCready and Cooperrider, 1978; Kartesz and Kartesz, 1980;
Roberts and Cooperrider, 1982; Spooner, 1982; United States
Department of Agriculture, 1982) accepted the validity of these
infraspecific taxa. Gratiola viscidula was collected in many new
localities in extreme southeastern Ohio and immediately adjacent
northern Kentucky and West Virginia. Almost all of these
populations are located in wetland habitats associated with pre-
glacial Teays-age valleys, and the “shortii” entity was believed to be
a disjunct taxon associated with these sites (Cusick and Silberhorn,
1977; Spooner, 1982).

Others, however, either did not accept these infraspecific taxa
(Fernald, 1950), or did not mention them (Fernald, 1937; Gleason
and :Cronquist, 1963; Radford et al., 1968; Silberhorn, 1970;
Godfrey and Wooten, 1981; Brumfield et al., 1982). The purpose of

fi,

80 Rhodora [Vol. 86

my study was to determine the taxonomic validity of the above
infraspecific taxa by assessing both inter- and intrapopulational
morphological variation of G. viscidula.

MATERIALS AND METHODS

Numerous specimens, including the types, were borrowed from 22
herbaria (see ACKNOWLEDGMENTS). Fifty-four mature individuals at
full anthesis were measured from throughout the range of the
species to ascertain the limits of morphological variability (Table 1).

Table |. Specimens measured for size variation within Gratiola viscidula (Fig. 2).

Locational Vouchers

DELAWARE: Newcastle Co., Commons 12 (PH); 2 Sep 1897, Commons s.n. (PH).
DISTRICT OF COLUMBIA: Pennell 12415 (OS); 11 Aug 1900, Steele s.n. (PH).
FLORIDA: Duval Co., Mar 1882, Smith s.n. (FP).

GEORGIA: Floyd Co., 1890, Chapman s.n. (US, two plants measured from this
sheet); McDuffie Co., Bartlett 1082 (VDB); Wilkes Co., s.d., Chapman s.n. (PH).

KENTUCKY: Estill Co., Lasseter 3054 (EKU); Knox Co., Stamper 30 (EKU).

MARYLAND: Prince Georges Co., Hotchkiss 7185 (US); St. Mary’s County, 20
Aug 1904, Chrysler s.n. (MARY).

MISSOURI: Shannon Co., Steyermark 72109 (F,GA).

NORTH CAROLINA: Anson County, Beal 4239 (DK); Bertie Co., Ahles and
Haesloop 52150 (UNC); Bladen Co., Ahles and Leisner 33341 (UNC); Biltmore
471b (US); Buncombe Co., Biltmore 461c (PH); Columbus Co., Beal 1805 (DK);
Gates Co., Correll 2249 (DK); Henderson Co., 23 Aug 1881, Smith s.n. (US);
Radford 4848 (UNC); Jackson Co., Radford 4759 (UNC); Lee Co., Beal 4284
(DK); Moore Co., Ahles 59588 (UNC); New Hanover Co., Aug 1892, Williamson
s.n. (PH); Rowan Co., Heller 135 (PH, two plants measured from this sheet);
Stokes Co., Radford 41497 (UNC); Swain Co., Aug 1891, Beardslee and Kofoid
s.n. (OS), Transylvania Co., Beal 5826 (DK); county unspecified; Aug 1841,
Rugel s.n. (US, three plants measured from this sheet).

SOUTH CAROLINA: Anderson Co., Radford 13937 (UNC); Chesterfield Co.,
Radford 15824 (TENN); Darlington Co., Smith 639 (UNC); Kershaw Co., House
2680 (US); Richland Co., 9 Jun (without year), Taylor s.n. (PH).

TENNESSEE: Blount Co., Sharp and Veloira 21520 (VDB); Carter Co., Fairchild et
al. 11741 (TENN); Cocke Co., Kearney 854 (OS, UNC, US); Morgan Co., Webb
et al. 1056 (TENN); White Co., Shaver 4228 (TENN).

VIRGINIA: Cumberland Co., Wells 38 (UNC); Dinwiddie Co., Kral 11353 (UNC);
Greensville Co., Harvill 17601 (UNC); Fernald and Long 10818 (DK);
Nansemond Co., Fernald and Long 7602 (PH); Powhatan Co., Corcoran and
Diggs 634 (UNC); Corcoran and Diggs 1147 (UNC); Southampton Co., Fernald
et al. 5917 (PH); Sussex Co., Fernald and Long 6390 (PH).

1984] Spooner — Gratiola 81

s \
fc) 100 200 300 400 |. :
asta)

Figure 1. County distribution of Gratiola viscidula. Circles and triangles repre-
sent specimens examined in this study, with the latter representing populations
measured for Figs. 2 and 3. Squares are records from literature references. Circled
records represent the previously presumed range of the “shortii” entity in wetland
habitats associated with preglacial Teays-age valleys.

These individuals were chosen to represent the morphological varia-
tion within the known range of the species (Fig. 1). Additional dis-
tributional data were obtained from Pennell (1935), Radford et al.
(1968), Harvill et al. (1981), and Medley and Thieret (in litt., 1982).
Thirty-nine mature individuals from a population within the
presumed “shortii” range [OHIO: Scioto Co., Spooner 2153 (OS)]

82 Rhodora [Vol. 86

75 Ros
| 2 eT ©

E

E

z aes oS
i of te

mi of om ce oOyee eg

ui oe oh bv

p

a
GS 2a:

Q.

BEXeN
Qa

O

ro

5 10 15 20 25 30
LEAF LENGTH mm

Figures 2 and 3. Pictorialized scatter diagrams of measurements from selected
populations of Gratiola viscidula. From throughout its range, (Fig. 2). Individuals
from a single population in Scioto Co., Ohio, Spooner 2153 (Fig. 3). Measurements
of the types of G. viscidula subsp. viscidula (TV), and of G. viscidula subsp. shortii
(TS) are included for comparison. (See Key at end of Figure 3 - facing page)

were chosen to determine the limits of intrapopulational morpho-
logical variation. The largest measurements per individual were
taken of corolla length, calyx length, pedicel length, leaf length and
leaf width, as these were the characters that Pennell (1935) used to
distinguish the infraspecific taxa. Other populations in this area
were also collected for visual comparison: KENTUCKY: Greenup
Co., Spooner 2150, 2151 (OS); OHIO: Gallia Co., Spooner 2155
(OS); Jackson Co., Spooner 2156 (OS). For chromosome number
determinations, the procedures involving conventional squash
techniques of flower buds for meiotic stages outlined in Keil and
Stuessy (1975) were used.

1984] Spooner — Gratiola 83

75 v
7 3 xe) kos
ee ts
ge & fe) ? yw re Fo
E
= bog
5 0 be °
We Yoo toy vous 0
2 oo
a
wi o Me) 0
eo}
4 fe)
os
gO O
3 We, a T T (|
5 15 15 20 25 30
LEAF LENGTH mm
KEY
Pedicel Length mm Leaf Width mm Corolla Length mm
O < 10.1 O < 49 O <94
© 10.1-14.9 OD 5.0-7.9 O 5.0-7.9
O > 149 O > 8.0 O> 10.9

RESULTS AND DISCUSSION

Measurements of individuals from throughout the range of the
species, exclusive of southeastern Ohio and immediately adjacent
northern Kentucky and West Virginia (the “shortii” range), are
presented in Fig. 2. It is evident from these data that G. viscidula
exhibits a wide range of size variation with the extremes connected
by a continuous series of intermediates. The type of G. viscidula
subsp. viscidula [NORTH CAROLINA: Forsyth Co., s.d.,

84 Rhodora [Vol. 86

Schweinitz s.n. (PH), (see Stuckey, 1979, for a discussion of this
specimen) ] falls on the small end of this size range. The type of G.
viscidula subsp. shortii [KENTUCKY: s.d., Short s.n. (PH),
(Specimen marked as Gratiola shortii by Elias Durand) ] falls on the
large end of this size range. Both specimens are marked as types by
Pennell.

Measurements of individuals of the Scioto County, Ohio
population are presented in Fig. 3. The degree of size variation in
this population is similar to that from throughout the range of the
species, with extremes encompassing measurements of the types as
above. The range of size variation of the other populations that I
collected in the field is similar.

The new distribution map (Fig. 1) fills in considerable gaps in
Pennell’s (1935) map of the species. There are now in excess of thirty
populations known for this species in southeastern Ohio and
immediately adjacent northern Kentucky and West Virginia.
Pennell (1935) was influenced by the disjunct nature of the “shortii”
populations, less disjunct now in light of the new distributional
data. Other disjunct records are known for this species: (1)
MISSOURI: Shannon Co., Steyermark 72109 (F, GA, MO, US).
These specimens were collected from Gilmore Pond, an upland
sinkhole pond. My efforts to relocate the plant at the site were
unsuccessful, and the plant is possibly extirpated. The last known
collection there was in 1975 [MISSOURI: Shannon Co., Christ s.n.
(pers. herb.)]. This region of Missouri is dotted with such habitats,
however, and the plant possibly occurs in other ponds in the area.
Other examples of Coastal Plain and Piedmont disjuncts are known
from this portion of Missouri (Steyermark, 1952).(2) FLORIDA:
Duval Co., Mar 1882, Smith s.n. (F). This specimen bears Pennell’s
annotation as G. viscidula dated 1931; it is not known why he did
not mention it in his monograph (Pennell, 1935). The Crittenden
Co. Arkansas record mentioned in Wilcox (1973) is apparently in
error, aS was suggested by Smith (1978). I requested a loan of the
specimen from MEM, where Wilcox deposited his vouchers, but it
has not been located.

Current nomenclatural errors exist with the above infraspecific
taxa of G. viscidula. Pennell (1935) treated the two taxa as
subspecies. Gleason (1952) regarded these taxa as varieties, but did
not formally treat them in a manner indicating his acceptance of

1984] Spooner — Gratiola 85

them (see Gleason, 1952, Vol. 1, p. xxxv). Cooperrider (1976)
accepted the taxonomic validity of these subspecific taxa at the
varietal level, but incorrectly cited the “‘shortii” entity as G. viscidula
var. shortii (Pennell) Gleason. This citation continues to be accepted
(McCready and Cooperrider, 1978; Roberts and Cooperrider, 1982;
Spooner, 1982; United States Department of Agriculture, 1982).
Kartesz and Kartesz (1980) incorrectly cited G. viscidula var. shortii
(Durand) Gleason. If these infraspecific taxa are recognized at the
varietal level, a new combination is required.

The chromosome number of G. viscidula is here first reported as
n = 7 from two populations: (1) KENTUCKY: Greenup Co.,
Spooner 2150 (OS); and (2) OHIO: Gallia Co., Spooner 2156 (OS).
This base number is in agreement with previous counts for the
section Gratiolaria Bentham (Lewis et al., 1962).

In view of the continuous degree of size variation represented
both within individual populations and throughout the range of the
species, recognition of infraspecific categories based on the above
size characters is unwarranted. In addition, no other consistent
differences were observed in any other features that would warrant
recognition of infraspecific categories.

The consistent and widespread occurrence of this variation seen
both in the field and on herbarium specimens indicates that this is
an inherent feature of this species, and is not due to localized
introgression with related taxa. The most closely related taxa are G.
brevifolia Raf., G. ramosa Walt., and G. aurea Muhl. ex Pursh (G.
lutea Pursh) (Pennell, 1935), but these species are largely allopatric
with G. viscidula and not notably different in size from this species.

ACKNOWLEDGMENTS

I gratefully acknowledge the Ohio Department of Natural
Resources, Division of Natural Areas and Preserves, for support for
field work, and the curators of the following herbaria for making
material available for study: ALU, DUKE, EKY, F, FSU, GA,
GFND, KE, MARY, MO, MUHW, NCSC, OS, PENN, PH,
TENN, UNA, UNCC, US, VDB, WVA. Use of the personal
herbarium of Arthur Christ, St. Louis, MO is also acknowledged.
Dr. Edward T. Browne searched MEM for the Crittenden Co.,
Arkansas voucher. Dr. John W. Thieret and Max E. Medley alerted

86 Rhodora [Vol. 86

me to recent Kentucky records of G. viscidula; Virginia Wallace and
John E. Wylie provided information about the species in Missouri.
Dr. Robert K. Jansen, Thomas G. Lammers, Dr. Ronald L.
Stuckey, and Dr. Tod F. Stuessy provided suggestions on the final
manuscript.

LITERATURE CITED

BRUMFIELD, B. M., D. K. EvANs, AND A. E. BRANT.1982. Additions to the wetland
flora of West Virginia. Castanea 47: 179-180.

COOPERRIDER, T. S. 1976. Notes on Ohio Scrophulariaceae. Castanea 41:
223-226.

Cusick, A. W., AND G. M. SILBERHORN. 1977. The vascular plants of unglaciated
Ohio. Ohio Biol. Surv. Bull. N.S. 5(4): i-x, 153 p.

FERNALD, M. L. 1937. Local plants of the inner Coastal Plain of southeastern
Virginia. Rhodora 39: 321-366, 379-415, 433-459, 465-491.

1950. Gray’s manual of botany. Ed. 8. American Book Co., New York.
i-lxiv, 1632 p.

GLeAson, H. A. 1952. New Britton and Brown illustrated flora of the north-
eastern United States and adjacent Canada. The New York Botanical Garden,
New York. Vol. 1, i-Ixxv, 482 p.; Vol. 3, i-iii, 589 p.

+» AND A. CRoNQuistT, 1963. Manual of the vascular plants of the
northeastern United States and adjacent Canada. D. Van Nostrand, New York,
i-il, 810 p.

Goprrey, R. K., AND J. W. Wooten. 1981. Aquatic and wetland plants of the
southeastern United States. Dicotyledons. University of Georgia Press, Athens.
933 p.

Harvitt, A. M., Jr., T. R. BRADLEY, AND C. E. STEVENS. 1981. Atlas of the
Virginra flora. Part II, Dicotyledons. Virginia Botanical Associates, Farmville,
Virginia. 148 p.

Kartesz, J. T., AND R. Kartesz. 1980. A synonymized checklist of the vascular
flora of the United States, Canada, and Greenland. The University of North
Carolina Press, Chapel Hill. i-xlvii, 500 p.

KeiL, D. J., AND T. F. Stugssy. 1975. Chromosome counts of Compositae from
the United States, Mexico, and Guatemala. Rhodora 77: 171-195.

Lewis, W. H., H. L. StRIPLING, AND R. G. Ross. 1962. Chromosome numbers
for some angiosperms of the southern United States and Mexico. Rhodora 64:
147-161.

McCreapy, G. A., AND T. S. CooperriIDER. 1978. The Scrophulariaceae
subfamily Scrophularioideae of Ohio, Castanea 43: 76-86.

PENNELL, F.W. 1935. The Scrophulariaceae of eastern temperate North America.
Acad. Nat. Sci. Philadelphia Monogr. |: i-xiv, 650 p.

Raprorb, A. E., H. E. AHLES, AND C. R. Bett. 1968. Manual of the vascular
flora of the Carolinas. The University of North Carolina Press, Chapel Hill.
i-Ixi, 650 p.

1984] Spooner — Gratiola 87

RoserTs, M. L., AND T. S. COOPERRIDER. 1982. Dicotyledons. 48-84. IN: T. S.
Cooperrider, ed. Endangered and threatened plants of Ohio. Ohio Biol. Surv.
Notes 16. 92 p.

SILBERHORN, G. M. 1970. A distinct phytogeographic area in Ohio: The
southeastern Allegheny Plateau. Castanea 35: 277-292.

SMITH, E. B. 1978. An atlas and annotated list of the vascular plants of Arkansas.
University of Arkansas Bookstore, Fayetteville. i-iv, 592 p.

Spooner, D. M. 1982. Wetlands in Teays-age valleys in extreme southeastern
Ohio: Formation and flora. 89-99. IN: B. McDonald, ed. Proceedings of
symposium on wetlands of the Unglaciated Appalachian Region. Morgantown,
West Virginia, May 26-28. 253 p.

STEYERMARK, J. A. 1952. New Missouri plant records (1949-1951). Rhodora 54:
250-260.

Stuckey, R. L. 1979. Type specimens of flowering plants from eastern North
America in the herbarium of Lewis David von Schweinitz. Proc. Acad. Nat. Sci.
Philadelphia 131: 9-51.

UNITED STATES DEPARTMENT OF AGRICULTURE, SOIL CONSERVATION SERVICE,
1982. National list of scientific plant names. Vol. |. List of plant names. U.S.
Govt. Printing Office, Washington, D.C. 416 p.

WiLcox, W. H. 1973. A survey of the vascular flora of Crittenden County,
Arkansas. Castanea 38: 286-297.

DEPARTMENT OF BOTANY
THE OHIO STATE UNIVERSITY
1735 NEIL AVENUE
COLUMBUS, OHIO 43210

ADDITIONS TO THE FLORA OF
ULSTER COUNTY, NEW YORK

MarY DOMVILLE

Since the publication of the “Flora of Ulster County, New York”
by Domville and Dunbar’, the following plants have been collected.
Most were collected by Karl L. Brooks, the late Stanley J. Smith
and Mary Domville. Most are deposited in the State Museum in
Albany, N. Y.

ISOETACEAE—QUILLWORT FAMILY
Isoetes tuckermanii A. Br. Quillwort
Infrequent in tidal shores

POLYPODIACEAE—FERN FAMILY
Dryopteris Xtriploidea Wherry (D. intermedia < D. spinulosa)
Triploid wood fern
Rare in mountain woods

POACEAE—GRASS FAMILY

Deschampsia caespitosa (L.) Beauv. Tufted hair-grass Natzd.
Very rare in sand plains

Digitaria filiformis (L.) Koel. Slender crabgrass
Very rare in sand plains

Panicum acuminatum Sw. Panic grass
Frequent in sterile soil

P. lanuginosum Ell. Panic grass
Frequent in Hudson Valley

Puccinellia distans (Jacq.) Parl. Goose-grass
Rare in moist soil

CYPERACEAE—SEDGE FAMILY
Carex crawfordii Fern. Sedge
Frequent in meadows and edges of woods
C. grayii Cary Sedge
Frequent in lowlands
C. lacustris Willd. Sedge
Infrequent in swampy area of Hudson Valley

\The book may be purchased from J. Donald Adams, 8 Maple Lane, Hyde Park, NY
12538. $4.00 postage paid.

89

90 Rhodora [Vol. 86

C. ormostachya Wieg. Sedge
Infrequent in woods of Hudson Valley
C. sprengelii Dew. ex Spreng. Sedge
Infrequent in open woods and damp places
C. tenuiflora Wahl. Sedge
Infrequent in bogs of mountains
Rhynchospora fusca (L.) Ait. Beak-rush
Infrequent in swamps

COMMELINACEAE—SPIDERWORT FAMILY
Tradescantia ohiensis Raf. Spiderwort
Frequent in thickets and meadows

JUNCACEAE—RUSH FAMILY
Juncus gerardi Loisel. Black grass
Rare on roadside where salt for snow removal was stored
J. greenei Oakes and Tuckerm. Greene’s rush
Rare in sandy places
J. pelocarpus Mey. Rush
Infrequent in damp places

LILIACEAE—LILY FAMILY
Allium porrum L. Leek
Spread from cultivation
Scilla hispanica Mill. Spanish bluebells Natzd.
Spread from cultivation

AMARYLLIDACEAE—AMARYLLIS FAMILY
Leucojum vernum L. Snowflake Natzd.
Frequently naturalized

ORCHIDACEAE—ORCHID FAMILY
Isotria medeoloides (Pursh) Raf. Small whorled pogonia
Collected only in 1920 by H. M. Denslow

SALICACEAE— WILLOW FAMILY
Salix pentandra L. Laurel-leaved willow
Frequent in Catskills

MORACEAE—MULBERRY FAMILY

Cannabis sativa L. Marijuana Natzd.
Frequently escapes from cultivation
Humulus japonicus Sieb. & Zucc. Japanese hop Natzd.

Frequent in waste places

1984] Domville — Ulster County flora 9]

POLYGONACEAE—BUCK WHEAT FAMILY
Rumex altissimus Wood. Pale dock
Frequent in rich soil
R. Xacutus L. (R. crispus < R. obtusifolius)
Infrequent in the Hudson Valley

CARYOPHYLLACEAE—PINK FAMILY
Gypsophila paniculata L. Baby’s-breath
Frequently escapes from cultivation
Spergularia media (L.) Presl. ex Griseb. Sand-spurrey
Infrequent in sand plains

RANUNCULACEAE—BUTTERCUP FAMILY
Ranunculus repens L. Creeping buttercup
Locally abundant

PAPAVERACEAE—POPPY FAMILY
Macleaya cordata (Willd.) R. Br. Plum poppy Natzd.
Frequently escapes from cultivation

FUMARIACEAE—FUMITORY FAMILY
Dicentra formosa (Haw.) Walp. . Bleeding-heart Natzd.
Frequent

BRASSICACEAE—MUSTARD FAMILY

Arabis hirsuta (L.) Scop. Hairy cress
Saugertie’s Point

Iberis umbellata L. Globe candytuft Natzd.
Spread from cultivation

Brassica oleracea L. var. acephala DC. Common kale _ Natzd.
Spread from cultivation

Sinapis arvensis L. Charlock Natzd.
Spread from cultivation

ROSACEAE—ROSE FAMILY

Amelanchier sanguinea (Pursh) DC. Red-branched shad
Infrequent in mountains

Crataegus pedicellata (Sarg.) Palmer. Hawthorn
Ashokan campus of New Paltz College (In herbarium of New
Paltz College)

C. pruinosa (Wendl.) K. Koch. Hawthorn
Infrequent (Collected by N. L. Britton in Shokan)

Filipendula rubra Rob. Queen-of-the-prairie Natzd.
Frequently spreads from cultivation

92 Rhodora [Vol. 86

Prunus’ nigra Sit. Canada plum
Infrequent in Hudson Valley

Pyrus prunifolia Willd. Crab-apple Natzd.
Self-seeded in Hudson Valley

Rosa centifolia L. Cabbage rose Natzd
Frequent

R. cinnamomea L. Cinnamon rose Natzd.
Spread from cultivation

R. wichuraiana Crep. Memorial rose Natzd.

Spread from cultivation

GERANIACEAE—GERANIUM FAMILY
Geranium bicknellii Britt. Bicknell’s geranium
Infrequent in disturbed soil

EUPHORBIACEAE—SPURGE FAMILY
Euphorbia epithymoides L. Cushion spurge Natzd.
Spread from cultivation

ANACARDIACEAE—CASHEW FAMILY
Toxicodendron radicans var. negundo (Green) Fern. Poison ivy
Observed in Hudson Valley and lowlands by M. Domville
T. radicans var. rydbergii Small ex Rydb. (Gillis, 1971)

Poison ivy

CELASTRACEAE—STAFF—-TREE FAMILY
Celastrus orbiculatus Thunb. Oriental bittersweet Natzd.
Frequently escapes in lowlands

HIPPOCASTANACEAE—HORSE CHESTNUT FAMILY
Aesculus hippocastanum L. Horse chestnut
Self-seeded in Hudson Valley

BALSAMINACEAE—TOUCH-ME-NOT FAMILY
Impatiens glandulifera Royle. Snapweed Natzd.
Spread from cultivation in lowlands

MALVACEAE—MALLOW FAMILY
Sida spinosa L. Prickly mallow Natzd.
Flood plains in New Paltz

HYPERICACEAE—ST. JOHN’S WORT FAMILY
Hypericum pyramidatum Ait. Great St. John’s wort
Flood plains in New Paltz

1984] Domville — Ulster County flora 93

VIOLACEAE—VIOLET FAMILY
Viola renifolia Gray. Kidney-leaved violet
(Collected by S. J. Smith)
V. Xconvicta House (V. fimbriatula < V. palmata)
Many plants at top of Hickory Hill
V. Xpeckiana House (V. palmata X V. sororia)
Locally abundant

ONAGRACEAE—EVENING-PRIMROSE FAMILY
Oenothera tetragona Roth. Sundrop
Infrequent (Collected in town of Olive)

ARALIACEAE—GINSENG FAMILY
Aralia spinosa L. Prickly ash or Hercules’ club
(Collected near Bearsville)

APOCYNACEAE—DOGBANE FAMILY
Apocynum cannabinum L. var. pubescens (Mitchell) A. DC.
Indian hemp
Frequent in thickets

ASCLEPIADACEAE—MILK WEED FAMILY
Asclepias purpurascens L. Purple milkweed
Infrequent in Hudson Valley

BORAGINACEAE—BORAGE FAMILY
Borago officinalis L. Borage Natzd.
Spread from cultivation

LAMIACEAE—MINT FAMILY
Ajuga genevensis L. Bugleweed Natzd.
Spread from cultivation
Pyananthemum verticillatum (Michx.) Pers. Whorled mountain
mint
Infrequent
Scutellaria integrifolia L. Hyssop skullcap
Open fields (Collected in town of Olive)

SOLANACEAE—NIGHTSHADE FAMILY
Solanum nigrum L. var. virginicum L. Nightshade Natzd.
Disturbed soil in Hudson Valley

SCROPHULARIACEAE—FIGWORT FAMILY
Digitalis lanata Ehrh. Grecian foxglove Natzd.
Escaped from cultivation

94 Rhodora [Vol. 86

Gratiola aurea Muhl. Golden-pert
Infrequent in wet areas

RUBIACEAE—MADDER FAMILY
Galium sylvaticum L. Scotch-mist Introduced
Observed at Saugertie’s Point by M. Domville

CAPRIFOLIACEAE—HONEYSUCKLE FAMILY
Lonicera caprifolium L. Italian woodbine Natzd.
Frequently escapes (Collected by P. Huth near Esopus)

ASTERACEAE—ASTER FAMILY

Anthemis tinctoria L. Yellow chamomile Natzd.
Frequent on roadsides

Echinacea purpurea (L.) Moench. Purple coneflower
Introduced from the west

Helianthus microcephalus T. & G. Small wood

sunflower Natzd.
Infrequent in southern part of county

LITERATURE CITED

Gittis, W. T. 1971. The systematics and ecology of poison-ivy and the poison-
oaks. Rhodora 73: 370-443.

174 CAROL WOODS
CHAPEL HILL, NORTH CAROLINA 27514

VERBASCUM DENSIFLORUM IN
SOUTHEAST WISCONSIN

JAMES A. REINARTZ

ABSTRACT

Nine of 260 species of the Eurasian genus Verbascum have been reported from
America; only two (V. thapsus and V. blattaria) are common. Verbascum
densiflorum is well established in the U.S. only in southeast Wisconsin where it is an
aggressive weed. It differs morphologically, phenologically and ecologically from
either V. thapsus or V. phlomoides, the two more common species which it
superficially resembles.

Key Words: Verbascum, adventive weed, restricted range

Verbascum is a Eurasian genus of over 260 species. Nine species
(V. thapsus L., V. blattaria L., V. phlomoides L., V. lychnitis L., V.
pPhoeniceum L., V. virgatum Stokes, V. nigrum L., V. sinuatum L.
and V. densiflorum Bertoloni) have been reported from North
America. These nine mullein species are among the most widespread
of Verbascum species in Europe and Asia (Murbeck, 1939). Two
hundred species of Verbascum are confined to a relatively small area
in Greece, southern Yugoslavia, Bulgaria, Rumania, Turkey, Syria,
Jordan, Iraq and northwest Iran. Only V. thapsus and V. blattaria
are common throughout North America. Verbascum sinuatum and
V. densiflorum have previously been reported only as rare waifs on
ballast along the east coast (Gleason, 1952).

A large, well established population of Verbascum densiflorum
(syn V. thapsiforme'! Schrader) has been found in southeast
Wisconsin. Within an area of about 30 square miles in Ozaukee and
Washington Counties, Wisconsin (Figure 1) essentially 100% of the
Verbascum is V. densiflorum. In three years I have never observed a
V. thapsus growing in this zone. Surrounding this zone of
exclusively V. densiflorum is an area of about 25 square miles in
which. both V. thapsus and V. densiflorum and some mixed
populations are found. Outside of this area V. densiflorum is not
found but V. thapsus is common.

'Verbascum thapsiforme Schrader, Monogr. gen. Verb., | (1813) is the name in more
common usage, but V. densiflorum Bertoloni, Rar. it. pl., (1810) is an older, and
therefore, the valid species name (Ferguson, 1972).

95

96 Rhodora [Vol. 86

Figure 1. Location of southeast Wisconsin population of Verbascum densi-
florum.

In an average year, flowering individuals of Verbascum densi-
florum in this area number in the thousands. The species has grown
in this area for at least 17 years as documented by a 1967 specimen
in the University of Wisconsin-Milwaukee Field Station herbarium.
The species may well have been established in this area for a long
time.

Verbascum densiflorum is distinct morphologically, phenologic-
ally and ecologically from either V. thapsus or V. phlomoides, the
two more common species which it superficially resembles (Figure
2). Verbascum densiflorum is very similar to V. phlomoides except
that the upper cauline leaves are distinctly decurrent down the stem
in V. densiflorum and are not at all decurrent in V. phlomoides. The

1984] Reinartz — Verbascum densiflorum 97

Figure 2. Population of Verbascum densiflorum at the University of Wisconsin-
Milwaukee Field Station in Ozaukee Co., Wisconsin.

98 Rhodora [Vol. 86

inflorescence of V. phlomoides is somewhat more open with more
stalk being exposed between capsules than that of V. densiflorum.
Also, the inflorescence bracts and upper cauline leaves of V.
densiflorum tend to be longer acuminate than those of V.
pPhlomoides.

Anderson (1947) described an lowa City, lowa population of
Verbascum phlomoides and concluded that V. phlomoides and V.
densiflorum (thapsiforme) are not separate species. Verbascum
Phlomoides, V. thapsus and sterile hybrids between the two species
were collected from the Iowa City population by the author during
the summer of 1983. Hybrids form readily between V. thapsus and
V. phlomoides (Wagner, Daniel & Hansen, 1980) and are inter-
mediate in the extent of leaf decurrence. Since Anderson (1947)
described V. phlomoides as more or less decurrent and did not
mention V. thapsus X V. phlomoides hybrids, the author is led to
believe that Anderson may have been interpreting these hybrids as
his V. densiflorum-like plants.

While Verbascum phlomoides and V. densiflorum are very
similar they are unambiguously separated by the extreme leaf
decurrence in V. densiflorum and complete lack of decurrence in V.
Phlomoides. Although hybrids can occur naturally between V.
densiflorum and V. phlomoides (Murbeck, 1933), the hybrids are
always infertile, indicating that these are separate species.

Verbascum densiflorum differs morphologically from V. thapsus
in having much larger flowers (25-45 mm diam.), a spathulate
rather than capitate stigma, and a longer, less crowded in-
florescence. Verbascum densiflorum tends to branch more freely
producing many inflorescence spikes, and its leaves tend to be more
dentate and more acuminate. Hybrids also occur naturally between
V. densiflorum and V. thapsus, but are completely infertile
(Murbeck, 1933).

Verbascum densiflorum differs from V. thapsus in its flowering
phenology. In southeast Wisconsin V. thapsus blooms from late
June to late August, while V. densiflorum begins to bloom in early
July and continues until the time of a hard frost (often mid- to late
October).

Verbascum thapsus and V. densiflorum in Wisconsin also differ
ecologically. V. thapsus can often be killed by cutting after it has
bolted; however, V. densiflorum is able to withstand repeated
cutting with plants branching from the base and growing many

1984] Reinartz — Verbascum densiflorum 99

shorter inflorescence stalks. Repeated cutting can delay flowering
but the plants usually survive to flowering even if mowing continues
for 3 or 4 years. Because of its ability to withstand mowing, V.
densiflorum is a much more aggressive weed in agricultural land in
Ozaukee and Washington Counties, Wisconsin than is V. thapsus.
Populations are quite common even in regularly mowed hay fields
or sparse, weedy lawns.

It is curious that this species, which occurs nowhere else in North
America, here appears to be a more aggressive weed than
Verbascum thapsus. Thousands of individuals grow in this 30-
square-mile area to the apparent exclusion of the generally more
common V. thapsus.

LITERATURE CITED

ANDERSON, W. A. 1947. Verbascum phlomoides in lowa. Rhodora. 49 (579): 67
68.

FERGUSON, I. K. 1972. Scrophulariaceae. In T. G. Tutin et al., Flora Europaea.
Vol. 3. University Press, Cambridge. 370 pp.

GLeason, H. A. 1968. The New Britton and Brown Illustrated Flora of the
Northeastern United States and Adjacent Canada. Vol. 3. Hafner Publ. Co., New
York. 496 pp.

Murskck, S. V. 1933. Monographie der gattung Verbascum. Lunds Universitets
Arsskrift. 29(2). 630 pp.

1939. Weitere Studien iiber die Gattungen Verbascum und Celsia. Lunds
Universitets Arsskrift. 35(1). 70 pp.

WaGner, W. H., Jr., T. F. Danie. & M. K. HANSEN. 1980. A_ hybridizing
Verbascum population in Michigan. Mich. Bot. 19(1): 37-45.

UNIVERSITY OF WISCONSIN-MILWAUKEE
FIELD STATION

3095 BLUE GOOSE RD.

SAUKVILLE, WI 53080

OBSERVATIONS OF POTAMOGETON HILLII
MORONG IN NORTH AMERICA

C. BARRE HELLQUIST

ABSTRACT

Potamogeton hillii Morong has been considered as either a rare or endangered
species in all states and provinces where it occurs. Currently it is under review by the
United States Fish and Wildlife Service for possible listing under the Endangered
Species Act of 1973. Prior to 1960 it was collected at 25 localities. Since 1970, 78 new
localities have been discovered. Throughout its range this species is mostly found in
regions of limestone bedrock. Presently it is most abundant in western New England
and northern Michigan, and during 1982 was relocated in Connecticut and Ohio.
Potamogeton hillii appears to be spreading in the limestone regions where it
presently occurs. This taxon is found growing in clear, cold waters of small streams,
ponds, and beaver ponds with muddy substrates. The abundant winter buds and
fruits produced apparently guarantee the continued spread of this species.
Potamogeton hillii should not be considered further for protection nationally, nor
should it be so listed in the states where locally abundant.

Key Words: Potamogeton hillii, Potamogeton porteri, Potamogetonaceae, Pusillii,
rare, endangered species

Potamogeton hillii Morong (including P. porteri Fern.) is a
distinct member of the subsection Pusillii of the Potamogetonaceae.
It was named for Rev. Ellsworth J. Hill (1833-1917) who was first
credited for-collecting it near Manistee, Michigan on August 5, 1882
(Fernald, 1932). A study of specimens produced four earlier
collections, originally identified as other taxa. The oldest known
locality is from East Dorset, Vermont in 1857. Potamogeton porteri
described by Fernald (1932) from Lancaster, Pennsylvania has been
shown by Haynes (1974) to be P. hillii.

Fernald (1932) knew of nine localities for Potamogeton hillii in
northeastern United States. Voss (1965) discussed the failure to
relocate the type locality in Manistee, Michigan, but noted that only
two new populations had been located up to 1965 in Michigan.
Haynes (1974) indicated the presence of a number of new localities
from Michigan and New York. Weber (1940) first reported P. hillii
from Massachusetts, and Hellquist (1977) reported eight new
localities for Massachusetts.

The apparent rarity of this species led to its listing by the
Smithsonian Institution (1975) and Ayensu & DeFilipps (1978) as a

101

102 Rhodora [Vol. 86

threatened species in the United States. Individual state, provincial,
and regional lists have variously classified Potamogeton hillii as
rare, threatened, endangered, or extinct. These lists are: Vermont
(Countryman, 1978); Massachusetts (Coddington & Field, 1978);
New England (Crow, 1982; Crow er al., 1981); New York (Mitchell
et al., 1980; Mitchell & Sheviak, 1981); Pennsylvania (Wiegman,
1979); Ohio (Ohio Department of Natural Resources, 1982);
Michigan (Wagner er a/., 1977); and Ontario (Argus & White, 1977).
Potamogeton hillii was omitted from the Connecticut list (Mehr-
hoff, 1978) even though only a single location was known at the
time. The results of these surveys prompted its appearance in the
Notice of Review published by the United States Fish and Wildlife
Service, Office of Endangered Species in the Federal Register (FR,
15 December 1980, Vol. 45, No. 247) as a potential candidate for
listing, but needing further study.

Observations made in the field from 1972-1976 (Hellquist, 1977)
indicated population patterns that led to increased success in
locating this species. The result of the field work was the discovery
of 54 new localities for Potamogeton hillii since 1976.

Field observations on Potamogeton hillii have shown it to be a
species found in clear, cold, alkaline water in small, slow flowing
streams, ponds, and beaver ponds with a muddy substrate. In
streams, it often appears on the upstream side of road culverts
where more marshy conditions occur. In beaver ponds and marshes,
it often grows among stumps and fallen trees, or in shallow water
among rushes and sedges. In ponds, P. hillii is occasionally found in
deeper waters up to 1.5 meters.

Potamogeton hillii has rarely been reported from lakes. Speci-
mens previously identified as being collected from Cayuga Lake,
N.Y. were studied by Fernald (1932) and found to be the Red Mills
Pond population. No other specimens have been seen from the lake.
Wiegand and Eames (1925) listed locations previously reported
from central New York in a work by Dudley (1886). These localities
were: Dryden Lake, Red Mills Pond, pools north of Ithaca, and
Myers Point. Richard Mitchell (pers. comm.) indicated that
Clausen had observed P. hillii in the marshes at the south end of
Cayuga Lake. This area has been recently filled and commercially
developed. Potamogeton hillii occurs around the Great Lakes at
Manitoulin Island, Lake Huron and Cecil Bay, Lake Michigan. The
Cecil Bay population was along the shore at the outlet to French

1984] Hellquist — Potamogeton hillii 103

Farm Creek. I observed in 1975 a few plants directly at the mouth of
the creek. In 1977 the waters of Lake Michigan had receded and |
located no plants; a subsequent check in 1983 also failed to locate
any P. Aillii.

The alkalinity of the water is an extremely important chemical
character controlling the distribution of Potamogeton hillii. Water
samples from 35 localities indicate that P. hillii occurs in waters
ranging from 53.0-290.0 mg/l CaCO; with a mean of 124.1 mg/1
CaCO:. A survey of the geological maps and reports of the states,
provinces, and regions where P. hillii is known to occur have shown
that 79% of the locations coincide with bedrock of dolomitic
limestone, 15% of calcitic marble, micacous crystalline marble, and
limestone, 3% of shale, 1% of gneiss, 1% of conglomerates, and 1%
of sandstone [Vermont (Doll, 1961); Massachusetts (Emerson, 1916;
Dale; 1923); Connecticut (Dale, 1923); New York (Dale, 1923;
Fischer et al/., 1970); Pennsylvania (Frazer, 1877; Lesley, 1885;
Pennsylvania Geological Survey, 1960; Stevenson, 1882); Ohio
(Bownocker, 1947); Ontario (Morton, 1977; Salterly, 1958; Stauffer,
1914); Michigan (Marten, 1936a; 1936b) J.

Water quality also appears to influence distribution of the taxon.
Potamogeton hillii is mostly found in clear, cold waters, often
around springs and small inlets in ponds and marshes. This taxon Is
only occasionally found encrusted with marl even though growing
in highly alkaline waters. Potamogeton hillii is rarely found in
turbid. stagnant, or polluted waters.

Potamogeton hillii most typically occurs with P. foliosus Raf., P.
natans L., P. pusillus L. var. tenuissimus Mert. & Koch., P.
amplifolius Tuckerm., and P. gramineus L. It is rarely found with
other alkaline water species such as P. strictifolius Benn., P. friesii
Rupr., and P. pectinatus L. (Hellquist, 1980) which are often
associated with more eutrophic waters.

Potamogeton hillii is characterized by having short axillary
peduncles and fruits with a small dorsal keel. The fruits of P. Aillii
are much larger than those of the closely related P. foliosus (see
Haynes, 1974). The leaf tips are mostly bristled; this trait helps to
distinguish it from sterile P. foliosus in which the leaf tips are merely
acute. Potamogeton Strictifolius typically forms bristle-tipped leaves
but may be separated from sterile P. Aillii by the presence of a bold
margin around the leaf and distinct nodal glands. The bold margin
in P. strictifolius appears similar to the more prominent leaf veins.

104 Rhodora [Vol. 86

Haynes (1974) indicated the variability which may occur in the
morphology of the leaf tips. While the bristle-tipped leaves are
common, occasionally blunt-tipped or apiculate leaf tips are
observed. These leaves appear to occur both on plants early in the
growing season and during the latter portion of the season,
especially on the winter buds.

Fernald (1932) stated that winter buds are not known for
Potamogeton hillii. Haynes (1974) indicated he had observed winter
buds on a few sheets of P. hillii. During October 1982, I observed in
Vermont many plants of P. Aillii with numerous winter buds; in
many cases only the winter buds were present. These winter buds
often had the central portion partially hardened, similar to those of
P. obtusifolius Mert. & Koch, and P. pusillus var. tenuissimus.

Potamogeton hillii is occasionally seen with up to 5 veins, not 7 as
previously reported by Hellquist (1977). When the venation is more
than 5, P. ogdenii Hellquist & Hilton, a closely related but much
rarer taxon, should be suspected.

Potamogeton hillii is known from nine states and provinces in the
United States and Canada with a total of 108 locations having been
reported (Fig. 1). The number of reported sites by states and
provinces, (known verified locations in parentheses), are: Massa-
chusetts 35(32), Vermont 24(19), Michigan 15(11), Ontario 12(10),
New York 11(6), Pennsylvania 6(0), Ohio 3(1), and Connecticut
2(1). Currently P. hillii is known to be extant in all states except
Pennsylvania. Seventy-five percent of all known localities for P.
hillii have been found since 1970. As more field work is conducted in
the proper habitats within the alkaline regions, more populations
will surely be located.

Potamogeton hillii is locally abundant at most of its locations.
The numerous winter buds and fruits produced apparently guar-
antee survival and facilitate spread of this species. Potamogeton
hillii should no longer be considered for federal protection under the
Endangered Species Act of 1973, and should likewise be dropped
from consideration by the states of Massachusetts, Vermont, and
Michigan. In these states it is well established and not in danger of
extirpation.

The following list includes site information for all populations of
Potamogeton hillii documented by herbarium specimens. In in-
stances where many different collections were made at the same site
only the earliest, or the earliest and latest, records are recorded.

1984] Hellquist — Potamogeton hillii 105

T

T ° vo
o 85° 80 75 |
GC °
454
e
aa L ave{ fe
‘ _
Qn on™ q
——
- 45° on

Figure |. Distribution of Potamogeton hillii.

VERMONT: Bennington Co., Arlington, stream in cow pasture immediately east
of Battenkill River on south side of Rt. 313, Hellquist 15139 (NASC, NEBC),
Arlington, small pond on the west side of Old Depot Rd. ca. 1/2 mile south of Rt.
7A, Hellquist 15166 (MSU, NASC, NEBC, NYS, OS, UNA, VT); Arlington, marshy
pond along Warm Brook on the east side of Old Depot Rd. south of Rt. 7A,
Hellquist 15165 (F, MO, MSU, NASC, NEBC, PH, US, VT, WAT), Arlington, in
clear cool water among sedges in small pond on west side of Old Depot Rd. ca. 3/4
mile south of Rt. 7A, Hellquist 15169 (CM, CU, DAO, NASC, NEBC, PAC, PH,
VT, WAT); Arlington, small pond SW of town at Howell’s Campground in shallow
clear water, Hellquist 15167 (NASC, PAC, VT), Arlington, small stream ca. 1/4 mile
east of Rt. 7A along connection to new Rt. 7, East Arlington, Hellquist 15175
(MICH, NASC, NEBC, NHA, VT); Dorset, in shallow water of Otter Creek, | Aug.
1934, Eames s.n. (GH, YALE); Dorset, Prentiss Pond west of town, Hellquist 15175
(NASC, NEBC, VT); Dorset, East Dorset, 28 Aug. 1857, Aames s.n. (GH); Dorset,
abundant in small pool east of railroad north of Village Rd. and immediately south
of stream under railroad, East Dorset, Hellquist 15423 (DAO, NASC, NEBC, NY,
OS, VT, WAT); Manchester, among Equisetum fluviatile along shore of Battenkill
River at old fishing access site of “Dufresne Pd.”, Hellquist 15172 (MICH, MO,
MSU, NASC, NEBC, UNA, VT); Manchester, small pond near Rt. 7 in Dellwood
Cemetery south of Manchester, Hellquist 15196 (NASC, NEBC, OS, VT, WAT);
Pownal, South Stream at fishing access, 30 July 1973, Hellquist various numbers
(MASS, MICH, NASC, NEBC, NHA, OS, NLU, UCSB, VT). Orange Co.,
Washington, bog with Rhamnus alnifolia, Seymour 29679 (VT). Rutland Co.,

106 Rhodora [Vol. 86

Danby, common on east side of Rt. 7 in small pasture stream ca. 1.5 miles south of
Danby-Wallingford town line, Hellquist 15195 (NASC, NEBC, VT, US); Danby,
common on west side of Rt. 7 in small pasture stream ca. 1.6 miles of Danby-
Wallingford town line, Hellquist 15405 (NASC, NEBC, NHA, US, VT); Hubbard-
ton, uncommon in Giddings Brook along north side of Hill Rd. just west of Parsons
School, Hellquist 15422 (MO, NASC, NEBC, VT); Hubbardton, abundant in
Giddings Brook south of Hill Rd. at jct. with Ganson Hill Rd., Hellquist 15424
(CAN, DAO, NASC, NEBC, NY, NYS, UNA, US, VT); Mt. Tabor, shallow water,
Otter Creek, Eames 11643 (YALE); Mt. Tabor, common in small pond on east side
of Rt. 7, 0.8 mile south of Danby, Hel/quist 15193 (NASC, NEBC, VT), Mt. Tabor,
east side of Rt. 7, 0.8 mile south of Danby, Hellquist 15193 (NASC, NEBC, VT); Mt.
Tabor, east side of stream that crosses Rt. 7, 0.7 mile south of Danby, Hellquist
15194 (DAO, MO, NASC, NEBC, NYS, VT); Pittsford, abundant along east shore
of Smith Pd., Florence, Hellquist 15403 (F, NASC, NEBC, NY, UNA, VT, WAT):
Pittsford, small stream ca. 1/2 mile SW of Florence on road to Butler Pd., Hellquist
15404 (MICH, MO, NASC, NEBC, OS, PH, US, VT). Windsor Co., Windsor,
Evart’s Pd. Aug. 1886, Dudley s.n. (GH, NY); Windsor, Evart’s Pd., in deep water,
27 Aug. 1933, Weatherby & Griscom s.n. (NEBC).

MASSACHUSETTS: Berkshire Co., Alford, small pond on west side of West St.
off Tom Ball Bk., 2 Aug. 1974, Hellquist various numbers (CUSC, MO, NASC,
NEBC, NLU); Alford, marsh west of small pond on west side of West St. off Tom
Ball Bk., 2 Aug. 1944, Hellquist various numbers (ALU, MASS, MO, NASC); Great
Barrington, Muddy Bk, at Blue Hill Rd., 2 Aug. 1974, Hellquist various numbers
(NASC); Great Barrington, Muddy Bk. at Stoney Bk. Rd., Hellquist 10916 (ALU,
NASC, NEBC, NLU); Great Barrington, flooded portion of Muddy Bk. north of
Stoney Bk. Rd., Hellquist 14050 (CM, NASC); Hancock, edge of pool, Hunneweill
16997; Hancock, small stream east of Rt. 43 ca. 1/2 mile north of Whitman Rd. on
land owned by Eugene Reese, Hellquist 14388 (NASC); Hancock, pond along
Kinderhook Ck. east of Rt. 43 and north of Whitman Rd., Hellquist 14389 (NASC);
Lenox, pond in bird sanctuary, Githens 208 (NASC, PH); Lenox, eighth pond in
series of beaver ponds west of Rt. 7 along Yokun Bk., Hellquist 13653 (NASC,
NEBC); Lenox, East Branch, Yokun Bk., Hellquist 13663 (NASC, NEBC); Lenox,
beaver pd. along Yokun Bk., Hellquist 13674 (CM, MICH, NASC, NEBC); Lenox,
pond west of first beaver dam, west of Rts. 7 & 20, immediately behind the Yankee
Motor Lodge ca. 1/2 mile south of Pittsfield line, Hellquist 14403 (NASC); Lenox,
pond west of second beaver dam, west of Rts. 7 & 20 behind the Yankee Motor
Lodge, ca. 1/2 mile south of Pittsfield line, Hel//quist 14385 (NASC); Pittsfield, south
end of Mud Lake at outlet, Hel/quist 15220B (NASC); Pittsfield, beaver pond along
west side of Tamarack Rd. ca. 1/4 mile south of South Mountain Rd., Hellquist
15214 (NASC, NEBC, US); Pittsfield, beaver marsh on north side of Tamarack Rd.
ca. 1/4 mile west of Bousquet Ski area, Hellquist 15217 (CONN, CU, KANU,
NASC, NEBC, NY, NYS, PH, SDC, US, VT); Pittsfield, pond south of dam at
conservation park NE of Pittsfield airport, Hellquist 15216 (NASC); Richmond,
Cone Bk. at Lenox Rd. 6 Sept. 1972, Hellquist various numbers (ALU, CM, MICH,
NASC, NEBC, NHA); Richmond, Miller’s Pd., east of jct. of Rossiter Rd. and West
Rd., Hellquist 11837 (ALU, C, DS, MO, NASC); Richmond, Fairfield Pd., Hellquist
11838 (ALU, C, DS, NASC); Richmond, Sherrill Pd. at jet. of Rossiter Rd. and Rt.
41, Hellquist 13019 (CM, NASC, NEBC); Richmond, Crystal Lake on the west side

1984] Hellquist — Potamogeton hillii 107

of Swamp Rd. ca. 1/2 mile south of Lenox Rd., Hellquist 13676 (CM, MICH,
NASC, NEBC); Richmond, pond on the north side of Summit Rd., Hel/quist 13681
(NASC, NEBC); Richmond, small pond on south side of Summit Rd., Hell/quist
13684 (CM, MICH, NASC, NEBC); Richmond, small pond along Fairfield Bk. on
the north side of Sleepy Hollow Rd. ca. 1/2 mile east of Rt. 41, Hel/quist 14382
(NASC); Richmond, small farm pond on the west side of Rt. 41 ca. 3/4 mile south of
the Pittsfield line, 1/2 mile south of North Cemetery, Hellquist 14394 (CM, MICH,
NASC); Richmond, small pond on south side of Canaan Rd., 0.7 mile east of the
New York border, Hellquist 15191 (MICH, NASC, NEBC, NYS, US), South
Egremont, in 3 feet of quiet water of the inlet to the pond fed by Karner Bk., Weber
1626 (BH, CAN, CU, F, GH, MICH, MO, NHA, NY, US); South Egremont,
common in stream along the north side of Mt. Washington Rd. ca. 1/4 mile from jet.
of Rt. 41 at SW corner of the Mill Pd., Hellquist 15199 (F, MSU, NASC, NEBC,
NYS, OS, PAC, PH, US); Stockbridge, swamp along the south branch of Lily Bk. at
Bean Hill Rd., Hellquist 11147 (NASC); Stockbridge, Kampoosa Bk. on west side of
Rt. 7, ca. 1.0 mile north of Rt. 182, Hel/quist 15391 (CAN, NASC, NEBC, US); West
Stockbridge, pond south of Wilson Rd. ca. 1/2 mile west of West Center Rd., West
Stockbridge Center, Hellquist 15389 (CAN, CM, CONN, CU, DAO, F, KANU,
MICH, MO, MSU, NASC, NEBC, NHA, NY, NYS, OS, PAC, PH, SDC, UNA,
US, VT, WAT).

CONNECTICUT: Litchfield Co., abundant in Flat Bk. at jet. of Rts. 126 & 63,
Hellquist 15200 (CAN, CONN, DAO, MICH, MO, NASC, NEBC, NYS, OS, PH,
UNA, US, VT); Sharon, Indian Pd., 9 Sept. 1909, Bissell s.n. (NCBS, NASC).

NEW YORK: Columbia Co., Canaan, Beebe Pd. south of Beebe Pd. Rd.,
Hellquist 9651 (MASS, NASC, NLU). Dutchess Co., Amenia, in deep mud and
0.7-2.0 m water of Amenia Lake, scarcely reaching surface, usually in large clumps to
near 1.0 m diameter at top, Eames 11794 (NASC, NYS); Amenia, marsh on west side
of Rt. 22 ca. 1.5 miles south of Amenia, Hel/quist & Dean 15409 (CAN, CU, GH,
MICH, MO, NASC, NY, NYS, OS, UNA, VT); Pine Plains, Hoystadt s.n. (NY).
Thompkins Co., Dryden, Red Mills Pd., July 1895, Dudley s.n. (CU) [includes those
distributed by Morong as collected by Dudley from Cayuga Lake, | Aug. 1886,
Dudley s.n., see: Fernald, 1933,(CAN, F, GH, MICH, MO, US) ]; Malloryville, near
Malloryville, July 1886, Dudley s.n. (NYS). Washington Co., Putnam, abundant in
small pool along Rt. 22 on west side near culvert, muddy bottom, ca. 2.0 miles north
of Dresden Station Rd., Haynes 3342 (GH, MICH, OS); Putnam, stream east of
Road B at jct. with Rt. 22, north of Dresden line, Hellquist 15189A (CAN, CM, CU,
DAO, MO, NASC, PH); Putnam, ditch in marsh on west side of Rt. 22 across from
Road Bca. 0.1 mile north of Dresden town line, Hel/quist 15401 (CU, GH, KANU,
NASC, NY); Putnam, Mill Creek ca. 0.5 mile south of Rt. 22 and south of Putnam
Cemetery, Hellquist 15190 (CAN, CM, CU, F, MICH, NASC, PH, WAT);
Kingsbury, east of Vaughan’s, South Beaver Creek, 17 July 1900, Burnham s.n. (CU,
GH).

PENNSYLVANIA: Bedford Co., Woodbury, densely filling large Mill Pd. south
of Woodbury, Hotchkiss 6003 (GH, US); Woodbury, edge of dam, alt. 1210 ft., 1.0
mile SSW of Woodbury, 10 Aug. 1941, Berheimer s.n. (CM, PH). Crawford Co.,
Conneaut Lake, 14 Aug. 1869, Porter s.n. (PH); Harmonburg, 1889, Whiteside s.n.
(GH). Lancaster Co., Lancaster, cold riverlets near Lancaster, 5 Oct. 1860, Porter
s.n. (P, MO, NY, PH), [Type for Potamogeton porteri Fernald]. Mifflin Co.,

108 Rhodora [Vol. 86

Newton-Hamilton, in Beaverdam Run at Brush Run School, 2 miles NE of Newton-
Hamilton, Westerfield /66/9 (NYS, PAC, PH).

ONTARIO: Bruce Co., Albermarle Twp., Hope Bay, Albermarle Bk. by road
south of Hope Bay, Bruce Peninsula, in sluggish brook, 24 Aug. 1977, Morton &
Venn s.n. (CAN, MICH, WAT); Lindsay Twp., partially dried-up ditch on east side
of Brickman’s Corner-Cape Chin Rd., 0.1 mile north of Miller Lake East Rd.
Hellquist & Hellquist 15385 (CAN, DAO, MICH, NASC, PH, UNA, US, WAT); St.
Edmunds Twp., ditch connecting into Willow Ck. on north side of Rt. 6, 1.5 miles
north of radio tower and 2.7 miles north of Hidden River Rd., Hellquist & Hellquist
15384 (CAN, CU, DAO, GH, MICH, MSU, NASC, NY, OS, US, WAT). Elgin Co.,
St. Thomas, shallow water, entirely submerged except for tips with fruit, 12 June
1951, James s.n. (DAO). Manitoulin Co., Carnarvan Twp., Mindemoya, Manitoulin
Island, road south Mindemoya ca. 1.0 mile north of jet. with Providence Bay Rd., in
sluggish stream flowing out of swamp, Morton 80 (WAT); Carnarvan Twp.,
Mindemoya, west side of stream on large open marsh ca. 1.8 km south of
Mindemoya, Hellquist & Hellquist 15380 (NASC): Tehkummah Twp., South
Baymouth, in swamp at head of small boat docking area, Morton & Venn NA14834
(WAT); Tehkummah Twp., South Baymouth, Manitoulin Island, the Slash at the
head of Leason Bay, in ditch by side of road, Morton & Venn 10095 (WAT);
Tehkummah Twp., South Bay, Manitoulin Island, the Slash, NE of Leason Bay,
west side of bridge in stream, Hellquist & Hellquist 15368 (CAN, DAO, GH, MICH,
MO, NASC, US); Tehkummah Twp., Black Creek on east side of Side Road 15 ca.
0.4 km north of Providence Bay Rd., Hellquist & Hellquist 15379 (CAN, DAO,
MICH, NASC, US, WAT). Peel Co., Caledon Twp., Green Lake between Ist and
2nd lines, east side of Hwy. 24, Grid Ref. 795544, abundant in water 5 ft. deep,
Webber & Gregory 12120 (CAN, DAO, MICH, UNA); Caledon Twp., man-made
ponds on north and south side of Rt. 24, ca. 0.1 mile east of 2nd Line Rd., Hellquist
& Hellquist 15387 (NASC).

OHIO: Ashtabula Co., Ashtabula, pools, 17 July 1877, Hill s.n. (F); Wayne Twp.,
open, impounded mud bottom ponds on west tributary to Pymatuning Ck. upstream
from beaver dams just SE of intersection between Conrail tracks and Woodworth
Rd., NE corner of Wayne Twp., occasional, forming dense mats along shoreline,
Bissell & Peskin 30 (MICH, OS). Portage Co., Garrettsville, Silver Ck., Webb 1272
(F).

MICHIGAN: Cheboygan Co., locally frequent in ca. | ft. of water of ditch in
boggy ground a few yards east of Elliot Ck. near center El/2 Sec. 35, T38N, RIW,
ca. 4 miles east of Cheboygan, Voss 12735 (MICH, MSU, UMBS); Grant Twp., in
12 ft. of water near the southern shore of the more northerly of the Twin Lakes, Sec.
34, T37N, RIE, 2 Aug. 1951, Wood s.n. (GH, UMBS); Grant Twp, north shore of
Twin Lakes ca. 5.0 miles east of Alverno, Sec. 24, T37N, RIE, Hellquist & Haynes
10667 (MASS, NASC, NLU); Grant Twp., southern-most of the two streams flowing
into the northern bay of Twin Lakes, ca. 5.0 miles east of Alverno, Sec. 34, T37N,
RIE, Hellquist & Haynes 10670 (NASC). Emmet Co., Wawatam Twp., locally
common in ca. 1.5 ft. water, marsh in Cecil Bay between mouths of French Farm
Creek, Sec. 28, ca. 4 miles SW of Mackinaw City, Voss 14061 (CAN, MICH, MSC,
NASC, NY, UMBS); Wawatam Twp., French Farm Creek east of Wilderness State
Park Rd. at Cecil Bay, Lake Michigan, Sec. 28, T39N, R4W, Hellquist & Haynes
10484 (NASC); Bliss Twp. Big Sucker Ck., Wilderness State Park, 24 July 1950,

1984] Hellquist — Potamogeton hillii 109

Sparrow s.n. (MICH); Big Sucker Ck. at eastern bridge of Sturgeon Bay Trail,
Wilderness State Park, Sec. 34, T39N, RSW, Hellquist & Hellquist 15300 (F, MO,
MSU, NASC, NHA, PAC, UMBS, US). Kalkaska Co., Little Blue Lake, Ashley 92
(MICH). Mackinac Co., Engadine, abundant at intersection of M-117 & U.S. 2;
small stream which runs under M-117 ca. | mile south of Engadine, Sec. 20, T43N,
R1O0W, Haynes 4002 (OS). Manistee Co., Manistee, 5 Aug. 1880, Hill s.n. (F, PH,
MICH, Type specimens), Manistee, pool on Garfield’s farm near Manistee, 14 Aug.
1882, Morong s.n. (NY). Missaukee Co., West Branch Twp., ditch, west side of
Nelson Rd. just north of Haymarsh Ck., 3.5 miles north by road of Star City, Sec. I,
T23N, R6W, Pringle 646 (MICH); West Branch Twp., Haymarsh Creek, Nelson
Road, 3.5 miles north of Star City, Sec. 1, T23N, R6W, Hellquist & Hellquist
15293A (GH, MO, NASC, NY, UNA). Otsego Co., locally common in muddy
shallow water along west edge of road at outlet of Grass Lake, center of eastern edge
of Sec. 5, T32N, RIW, Stuckey & Nunan 1300 (GH, MICH, NY, NYS, OS, UMBS,
US); southeastern corner of Grass lake ca. 12 miles northeast of Gaylord, Sec. 3,
T32N, RIW, Hellquist & Haynes 10467 (NASC). Presque Isle Co., Bearinger Twp.
local in shallow water of Black Mallard River near mouth, NE 1/4, Sec. 25, T35N,
R2E ca. 1.0 mile south of Grace, Stuckey 3182 (OS); Ocqueoc Twp, east side of Little
Ocqueoc River south of Rt. M-68, 4'4 miles west of Moltke, north side Sec. 25,
T35N, R3E, Hellquist, Hellquist, & Crow 15317 (CAN, F, MICH, MSU, NASC,
NY, OS, UMBS, UNA, US).

ACKNOWLEDGMENTS

I wish to thank Garrett E. Crow for his suggestions and
manuscript review. Thanks are also extended to the curators of the
following herbaria for the loan of plant specimens: CAN, CM, CU,
DAO, F, GH, MICH, MO, MSU, NASC, NEBC, NY, NYS, OS,
PAC, PH, US, VT, WAT.

LITERATURE CITED

AYENSU, E. S. AND R. A. DeFiLipps. 1978. Endangered and threatened plants of
the United States. Smithsonian Inst. and the World Wildl. Fund, Inc.,
Washington, D.C. 403 pp.

Arcus, G. W. AND D. J. Wuite. 1977. The rare vascular plants of Ontario,
Syllogeus, No. 14. Natl. Mus. of Nat. Sci., Ottawa. 63 pp.

BownockeR, J. A. 1947. Geologic map of Ohio, with revision by G. W. White,
Geol. Surv. of Ohio.

CopDINGTON, J. AND K. G. FieLtp. 1978. Rare and endangered vascular plant
species in Massachusetts. New England Botanical Club in cooperation with the
U.S. Fish and Wildlife Service [Region 5, Newton Corner, MA]. 52 pp.

COUNTRYMAN, W. D. 1978. Rare and endangered vascular plant species in
Vermont. The New England Botanical Club in cooperation with the U.S. Fish and
Wildlife Service [Region 5, Newton Corner, MA]. 68 pp.

Crow, G. E. 1982. New England’s rare, threatened, and endangered plants. U.S.
Fish and Wildlife Service, Northeast Region. 130 pp.

110 Rhodora [Vol. 86

Crow, G. E., W. D. Countryman, G. L. CHurcH, L. M. EASTMAN, C. B. HELLQUIST,
L. J. MEHRHOFF, ANDI. M.Storks. 1981. Rare and endangered vascular plant
species in New England. Rhodora 83: 259-299.

Date, T. N. 1923. The lime belt of Massachusetts and parts of eastern New York
and western Connecticut. U. S. Geol. Surv. Bull. 744.

Dott, G. G. 1961. Centennial geologic map of Vermont. Vt. Geol. Surv., U.S.
Geol. Surv. and Harvard Univ.

Duptey, W. R. 1886. The Cayuga Flora, Part I. A catalogue of the phaenogamia
growing without cultivation in the Cayuga Lake basin. Bull. Cornell Univ. (Sci.)
2: 1-123.

EMERSON, B. K. 1916. Preliminary geologic map of Massachusetts and Rhode
Island, in: U. S. Geol. Surv. Bull. 397, “Geology of Massachusetts and Rhode
Island”, 1917.

FERNALD, M. L. 1932. The linear-leaved North American species of Potamogeton
section Axillaries. Mem. Am. Acad. Arts & Sci. 17: 1-183. (also, Mem. Gray
Herb. No. 3).

Fisher, D. W., Y. W. ISAACHSEN, AND L. V. RICKARD. 1970. Geologic map of New
York. New York State Mus. & Sci. Ser., Map and Chart Ser. 15.

FRAZER, P., JR. 1877. The geology of Lancaster Co. Second Geol. Surv. of
Pennsylvania Rep. of Progr. in 1877. 350 pp.

Haynes, R. R. 1974. A revision of North American Potamogeton subsection
Pusillii (Potamogetonaceae). Rhodora 76: 564-649.

HELLquist, C. B. 1977. Observations on some uncommon vascular aquatic plants
in New England. Rhodora 76: 564-649.

1980. Correlation of alkalininty and the distribution of Potamogeton in
New England. Rhodora 82: 331-344.

Lestey, J. P. 1885. A geological hand atlas of the 76 counties of Pennsylvania:
1885. Second Geol. Surv. of Pennsylvania, Rep. of Progr. X.

MarTEN, H.M. 1936a. The centennial geological map of the northern peninsula of
Michigan. Pub. 39, Geol. Ser. 33 (part of Ann. Rep. for 1936).

———. 1936b. The centennial geological map of the southern peninsula of
Michigan. Pub. 39, Geol. Ser. 33 (part of Ann. Rept. for 1936).

MenruorF, L. J. 1978. Rare and endangered vascular plant species in Connecticut.
The New England Botanical Club in cooperation with the U.S. Fish and Wildlife
Service [Region 5, Newton Corner, MA]. 41 pp.

MITCHELL, R. S. AND C. J. SHEVIAK. 1981. Rare plants of New York State. New
York State Mus. Bull. 445. Albany. 96 pp.

. C. J. SHEVIAK, AND J. K. DEAN 1980. Rare and endangered vascular plant
species in New York State. The State Botanist’s Office, New York State Mus.,
Albany, in cooperation with the U.S. Fish and Wildlife Service [Region 5, Newton
Corner, MA]. 38 pp.

Morton, J. K. 1977. The flora of Manitoulin Island and the adjacent islands of
Lake Huron, Georgian Bay, and the North Channel. Dept. of Bio., Univ. of
Waterloo, Ont. 62 pp.

OHIO DEPARTMENT OF NATURAL REsouRCES. 1982. Rare species of native Ohio wild
plants. [list]. Ohio Dept. of Nat. Res., Div. of Nat. areas and preserves.
PENNSYLVANIA GEOLOGICAL SURVEY. 1960. Geologic map of Pennsylvania. Penn-

sylvania. Geol. Surv., Harrisburg.

1984] Hellquist — Potamogeton hillii iil

SALTERLY, J. 1958. Geological map of the Province of Ontario. Map #1958B. Ont.
Dept. of Mines.

SMITHSONIAN INSTITUTION. 1975. Report on endangered and threatened plant
species of the United States. House Document No. 94-51, Serial No. 94-A,
Washington, D.C.

STEVENSON, J.J. 1882. The geology of Bedford and Fulton Counties. Second Geol.
Surv. of Pennsylvania. 382 pp.

STAUFFER, C. R. 1914. Geological survey of southwest Ontario. Ont. Dept. of
Mines, Map I16A.

WaGNER, W. H., E. G. Voss, J. H. BEAMAN, E. A. Bourpo, F. W. Case, J. A.
CHURCHILL, AND P. W. THomMpson. 1977. Endangered, threatened, and rare
vascular plants in Michigan. Mich. Bot. 16: 99-110.

Weser, W. A. 1940. Potamogeton hillii in Berkshire County, Massachusetts.
Rhodora 42: 95.

WIEGAND, K. M. AND A. J. EAMES. 1925. The flora of the Cayuga Lake basin, New
York. Vascular plants. Cornell Univ. Agric. Exp. Sta. Mem. 92. 491 pp.
WIEGMAN, P. G. 1979. Rare and endangered vascular plant species in Penn-
sylvania. The Western Pennsylvania Conservancy in cooperation with the U.S.

Fish and Wildlife Service [Region 5, Newton Corner, MA]. 94 pp.

Voss, E. G. 1965. Some rare and interesting aquatic vascular plants of northern
Michigan with special reference to Cusino Lake (Schoolcraft Co.). Mich. Bot. 4:
11-24.

DEPARTMENT OF BIOLOGY
NORTH ADAMS STATE COLLEGE
NORTH ADAMS, MASSACHUSETTS 01247

NEW ENGLAND NOTES

GERANIUM NEPALENSE VAR.
THUNBERGII AND G. SIBIRICUM
NATURALIZED IN WESTERN MASSACHUSETTS

C. JOHN BURK

Geranium nepalense Sweet var. Thunbergii (Siebold and Zucc.)
Kudo, a Japanese variety of a species widely distributed in Asia, was
first reported in North America as an aggressive garden weed in
Wellesley, Norfolk Co., Massachusetts (Hunnewell, 1945). A second
Massachusetts population of this variety with white rather than the
usual violet petals was later found in Newburyport, Essex Co.
(Bean, 1953). Although Fernald (1950) described G. nepalense var.
Thunbergii as locally abundant in Middlesex Co., Seymour (1969)
cited only the Wellesley and Newburyport collections for New
England.

Geranium nepalense var. Thunbergii is now well established as a
weed on the Smith College campus in Northampton, Hampshire
Co. The largest campus population extends intermittently for a
distance of approximately 80 m along both sides of a recently
reconstructed gravel path leading into floodplain forest on the
northwest side of Paradise Pond. The plants, which were not
observed prior to September, 1983, have set fruit abundantly and
are spreading aggressively by rhizomes, even invading the coarse
gravel surface of the path. Individual shoots and smaller colonies
occur sporadically elsewhere on the campus. The petals of all plants
seen flowering in 1983 were white.

The only New England report of the Eurasian Geranium
sibiricum L. is based on a specimen collected on a wasteheap in
Cambridge, Middlesex Co., MA in 1885 (Knowlton and Deane,
1919). Fernald (1950) described this species as locally abundant
from New York and Pennsylvania to II|linois. In 1972, the late Harry
E. Ahles collected G. sibiricum in a woodland border on the campus
of Mount Holyoke College, South Hadley, Hampshire Co., (AhA/les
75801, Aug. 5, 1972, Ahles 76187, Oct. 12, 1972 in MASS,
duplicates sent to NEBC). At present, G. sibiricum thrives on the
Mount Holyoke College campus with a large population beginning
north of the dam which impounds Upper Pond and continuing
approximately 100 m southwest along a fence and road edge. These
plants were fruiting densely in September, 1983.

113

114 Rhodora [Vol. 86

The western Massachusetts populations of these two geraniums
are similar in both aspect and habitat and the taxa might potentially
be confused. For example, specimens of Geranium sibiricum key
out to G. nepalense var. Thunbergii in Seymour (1969), while in
Gleason and Cronquist (1963), specimens of G. nepalense var.
Thunbergii key out to G. sibiricum, with the text then differ-
entiating G. nepalense var. Thunbergii on the basis of the number of
flowers produced per peduncle. The two taxa are readily distin-
guished, nonetheless. Geranium nepalense var. Thunbergii peren-
nates and increases by vigorous rhizomes which produce upright to
decumbent shoots while G. sibiricum is nonrhizomatous and may be
annual or perennial in duration. Geranium nepalense var. Thun-
bergii usually produces two flowers per axillary peduncle, each ona
separate pedicel, while G. sibiricum usually produces a single flower
on a solitary peduncle. In addition, in the western Massachusetts
populations, the fruits of G. nepalense var. Thunbergii are more
robust and range from |.5-2.0 cm in length, as opposed to 1.0-1.5
cm in G. sibiricum.

Specimens have been deposited in SCHN, MASS, and the
herbarium of Mount Holyoke College. I thank David A. Haskell,
Marian Rohman at the University of Massachusetts and Leslie
Lovett Doust at Mount Holyoke College for their generous
assistance with this project.

LITERATURE CITED

BEAN R. C. 1953. Two unusual plants in Essex County, Massachusetts. Rhodora
55: 348.

FERNALD, M. L. 1950. Gray’s Manual of Botany, Eighth Edition. American Book
Co., New York.

Gieason, H. A. & A. Cronaquist. 1963. Manual of Vascular Plants of: North-
eastern United States and Adjacent Canada. D. Van Nostrand Co., Inc.,
Princeton, New Jersey.

HUNNEWELL, F. W. 1945. Geranium nepalense var. Thunbergii in Massachusetts.
Rhodora 47: 219-220.

KNow_ton, C. H. & W. Deane. 1919. Reports on the flora of the Boston
district. — XXX. Rhodora 21: 78-83.

SEYMOUR, F.C. 1969. The Flora of New England. Charles E. Tuttle Co., Rutland,
Vermont.

DEPARTMENT OF BIOLOGICAL SCIENCES
SMITH COLLEGE
NORTHAMPTON, MA 01063

ARISTOLOCHIA TOMENTOSA SIMS
ESTABLISHED AT TWO WESTERN
MASSACHUSETTS SITES

C. JoHN BURK

The pipe-vine or Dutchman’s pipe-vine, Aristolochia tomentosa
Sims, is a high-twining shrubby liana native to stream edge habitats
of the southeastern and south central United States (Pfeifer, 1966).
The similar A. macrophylla Lam. occurs in forests of the
southeastern Appalachians. Pfeifer (1966) noted that the natural
ranges of these species are separate, with “neither sympatry nor
hybridization” where they border in the Smoky Mountains. Both
species are widely planted northward as fast-growing covers for
trellises, fences and verandas. Aristolochia tomentosa has not been
reported outside cultivation in New England, although it Is
naturalized locally in western New York (Fernald, 1950). Aristo-
lochia macrophylla, more frequently cultivated, has become nat-
uralized at some New England stations (Fernald, 1950). Pfeiffer
(1966) reported this species from Connecticut, Massachusetts, and
Vermont, and Seymour (1969) cited Connecticut and Vermont
specimens.

In June, 1982, Constance A. Parks and I found a small colony of
Aristolochia tomentosa in the Mt. Tom State Reservation, Hamp-
den Co., near the Hampshire Co. line. The vines occur on the north
side of Christopher Clark Road approximately 30 m southwest of
the Mt. Nonotuck overlook at an elevation of about 210 m. Several
stems ascend a distance of 6 m or more to the top of a northern
catalpa (Catalpa speciosa Warder) while others scramble into a
nearby staghorn sumac (Rhus typhina L.). Smaller pipe-vines,
apparently seedlings, occur among herbaceous vegetation near the
base of the older vines. The large vines were flowering on June 22,
1982, and on the same date in 1983, but did not set fruit either year.

A second, much larger colony of Aristolochia tomentosa occurs
near the west bank of the Connecticut River in Deerfield, Franklin
Co. Aristolochia tomentosa was first observed at this site by
Roberta Poland on August 29, 1957 (personal communication). The
vines occur in the vicinity of old cellar holes on the east side of
McClellan Farm road, approximately 1.9 km from the southern
intersection of McClellan Farm Road with River Road and

115

116 Rhodora [Vol. 86

opposite the East Deerfield railroad yards. On July 4, 1967, Mrs.
Poland visited the site with the late Harry E. Ahles. Specimens
collected in 1967 are now on file in the University of Massachusetts
Herbarium and in Mrs. Poland’s personal herbarium. By summer,
1983, A. tomentosa had spread roughly 40 m along McClellan Farm
Road, extending from the road edge eastward 15 m or more past the
cellar holes and into a ravine which approaches the Connecticut
River. Individual pipe-vines range in size from seedlings of the
current year to very large plants which climb 12 m or more into
various support trees, including staghorn sumac and northern
catalpa. Several of these trees, especially the sumacs, are dead or
dying and the aspect of the colony resembles that of sites farther
south where kudzu [Pueraria lobata (Willd.) Ohwi.] or Japanese
honeysuckle (Lonicera japonica Thunb.) have overrun woody
vegetation.

The Deerfield colony of Aristolochia tomentosa almost certainly
originated from a plant or plants persistent after cultivation, along
with lilacs (Syringa vulgaris L.) and summer phlox (Phlox
paniculata L.), which are also common in the immediate vicinity.
The origin of the Mt. Tom pipe-vines is unknown. Given the widely
differing habitats in which the two western Massachusetts colonies
are established and the vigor which both A. tomentosa and A.
macrophylla show in cultivation, these species might be expected to
be spreading elsewhere in the region.

I am grateful to Roberta Poland for showing me the Deerfield
colony of A. tomentosa and sharing her knowledge of the site and to
Marian Rohman for assistance again with this project. Specimens
from both sites are in MASS and SCHN.

LITERATURE CITED

FERNALD, M. L. 1950. Gray's Manual of Botany, Eighth Edition. American Book
Co., New York.

PFEIFER, H. W. 1966. “Revision of the North and Central American hexandrous
species of Aristolochia (Aristolochiaceae).” Ann. Missouri Bot. Garden 53:
115-196.

SeEYMouR, F.C. 1969. The Flora of New England. Charles E. Tuttle Co., Rutland,
Vermont.

DEPARTMENT OF BIOLOGICAL SCIENCES
SMITH COLLEGE
NORTHAMPTON, MA 01063

NANTUCKET FIELD TRIP OF THE
NEW ENGLAND BOTANICAL CLUB

RAY ANGELO & BRUCE A. SORRIE

The New England Botanical Club held a field trip to Nantucket,
Massachusetts on September 9-11, 1983. A diversity of habitats on
this offshore island was visited.

Some of the more interesting species seen in the barrens were
Aster concolor L. (in bloom and in Massachusetts known currently
only on Nantucket), Liatris borealis Nutt., Amelanchier nantucket-
ensis Bickn. (in good quantity though not known from the
mainland), and Hypericum adpressum Bart. (at the Wigwam kettle
ponds). Along Barnard Valley Rd. acres of Corema conradii Torr.
were admired. Closer to the ocean (south of Bartlett Farm) the party
searched unsuccessfully for Agalinis (Gerardia) acuta Pennell which
had been collected in the vicinity at one time (now known from only
three sites in its range). However, at this site were found Linum
intercursum Bickn. (a few blossoms lingering), Helianthemum
dumosum (Bickn.) Fern. (a few blossoms lingering), A/etris farinosa
L. (in fruit), and Prenanthes serpentaria Pursh. These open sand
plains (or “moors”) remain one of New England’s most intriguing
plant communities but are threatened by continued advance of
Quercus ilicifolia Wang.

Unusual examples of Quercus velutina Lam. and Acer rubrum L.
with tortuous, spreading limbs were examined in low, moist woods
in company with Nyssa sylvatica Marsh. and Sassafras albidum
(Nutt.) Nees. This locality was near Pocomo Road. About 14 miles
south of this site the party was led to locally rare stations of Actaea
rubra (Ait.) Willd. and Mitchella repens L. in the vicinity of the
Windswept Bogs owned by the Nantucket Conservation Founda-
tion. A particularly large specimen of //lex opaca Ait. was growing
nearby. In a thicket beside the dirt road that provided access to this
area, one of the party found a new record for the island—Pyrus
sieboldii Reg., a species that is naturalizing in eastern Massa-
chusetts.

The disturbed site of the old Siasconset dump provided an
interesting array of species including Rhynchospora torreyana Gray
(one of two known stations in the state), Solidago elliottii T. & G.
(in bloom), Lycopodium inundatum L., Aletris farinosa L., Drosera

117

118 Rhodora [Vol. 86

filiformis Raf., Platanthera blephariglottis (Willd.) Lindl. (essential-
ly past flowering), Pogonia ophioglossoides (L.) Ker and Spiranthes
cernua (L.) Richard var. ochroleuca (Ryd.) Ames (in bloom),

A side trip to Low Beach along the southern shore of the island
made by one member of the group turned up Polygonum glaucum
Nutt. in great quantity.

In the immediate vicinity of the Nantucket Field Station
(University of Massachusetts) Baccharis halimifolia L. (in bloom)
and Rubus bifrons Vest flourished. This is one of the very few sites
in New England where this latter alien species is known to be
established.

Faunal rarities that highlighted the field trip were Egretta gularis
(Western Reef Heron), a vagrant from West Africa, and Speyeria
idalia (Regal Fritillary) which is now rare in eastern United States.
Both were seen in the general vicinity of the Field Station.

All the new records for vascular plants found on the island during
this trip are adventive species. These are Buddleia davidi Franch.,
Clematis dioscoreifolia Levl. & Vaniot, and Pyrus sieboldii Reg.
Voucher specimens have been deposited with the New England
Botanical Club herbarium (NEBC).

The Club is very grateful for the hospitality and generous
assistance of Dr. Wesley N. Tiffney Jr., Director of the Nantucket
Field Station. Thanks also go to Dr. Robert Zaremba for sharing
his knowledge of ecologically interesting sites on the island.

CONCORD FIELD STATION
OLD CAUSEWAY RD.
BEDFORD, MA 01730

86 ELM ST.
KINGSTON, MA 02364

1984] Erratum / Notice 119

ERRATUM, OCTOBER 1983 ISSUE (VOL. 85, NO. 844)

“Relationships of two isolated groups of sugar maples (Acer
saccharum Marshall ssp. saccharum) in west central Oklahoma to
eastern and western species” by Thomas C. Dent and Robert P.
Adams.

Figures 3 and 4 are transposed from respective captions.

ANNOUNCEMENT

The annual Joint Field Meeting of the Northeastern Section of
the Botanical Society of America, the Torrey Botanical Club, and
the Philadelphia Botanical Club will be held on June 17 to 20, 1984,
in Salisbury, MD. Accommodations will be at Salisbury State
College. There will be guided field trips to dunes, forests, swamps,
and marshes at various sites on the Delmarva Peninsula in
Maryland, Virginia, and Delaware. Space is limited and prior
registration is required. Full details will be available after February
1, 1984, by writing the Chairman, Dr. Larry H. Klotz, Dept. of
Biology, Shippensburg University, Shippensburg, PA 17257 (717-
532-1401).

Vol. 85, No. 843, including pages 274-396, was issued August 11, 1983.

Vol. 85, No. 844, including pages 397-473, was issued December 31, 1983.

INSTRUCTIONS TO CONTRIBUTORS TO RHODORA

Submission of a manuscript implies it is not being considered for
publication simultaneously elsewhere, either in whole or in part.

Manuscripts should be submitted in triplicate (an original and
two xerox copies) and must be double-spaced (at least 3/8 of an
inch) throughout including footnotes, figure legends, and refer-
ences. Please do not use corrasable bond. The list of legends for
figures and maps should be provided on a separate page. Footnotes
should be used sparingly. Do not indicate the style of type through
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specimens. Names of genera and species may be underlined to indi-
cate italics in discussions. Specimens citations should be selected
critically, especially for common species of broad distributions. Sys-
tematic revisions and similar papers should be prepared in the for-
mat of “A Monograph of the Genus Malvastrum”, S.R. Hill,
Rhodora 84: 1-83, 159-264, 317-409, 1982, particularly with refer-
ence to indentation of keys and synonyms. Papers of a floristic
nature should follow, as far as possible, the format of “Annotated
list of the ferns and fern allies of Arkansas”, W. Carl Taylor and
Delzie Demaree, Rhodora 81: 503-548, 1979. For bibliographic cit-
ations, refer to the Botanico-Periodicum-Huntianum (B-P-H,
1968), which provides standardized abbreviations for journals
originating before 1966. All abbreviations in the text should be
followed by a period, except those for standard units of measure
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CBE Style Manual, Sth ed. (original title: Style Manual for
Biological Journals). In preparing figures (maps, charts, drawings,
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should be calculated to reflect the actual printed size. An Abstract
and a list of Key Words should be supplied at the beginning of each
paper submitted, except for a very short article or note.

RHODORA January 1984 Vol. 86, No. 845

CONTENTS

The type localities of the Flora boreali-americana of André Michaux
Leonard J. Uttal : poe : Re re 0 ema

Additions to the flora of oe Breton Highlands National Park, Nova Scotia
Harold R. Hinds . ee ae lf 2b eae ok) Ca Se ee

Studies in the Aristida (Gramineae) of the southeastern United States I. Spikelet
variation in A. purpurescens, A. tenuispica, and A. virgata
Kelly W. Allred ; Rat en , “So! oe.” see age eee ae ee

Infraspecific variation in Gratiola viscidula Pennell (Scrophulariaceae)
David M. Spooner P et ee ery .

Additions to the flora of Ulster County, New York
Mary Domville ‘ . é : : ; 5 , : : ’ . 89

Verbascum densiflorum in southeast Wisconsin
James A. Reinartz ; : : , : F - 7 : : ‘ ;

Observations on Potamogeton hillii Morong in North America
C. Barre Hellquist : Pie wy : . ; : j ; . . 101

New England Notes
Geranium nepalense var. Thunbergii and G. sibiricum naturalized in western -

Massachusetts
C. John Burk fo et gs SAN Te ing rs,
Aristolochia tomentosa Sims established at two western Massachusetts sites
C. John Burk : ; , ‘ oo ets Sens - . ets
Nantucket field trip of the New England Botanical Club
Ray Angelo and Bruce A. Sorrie i P : : . , : Oe Bled
Erratum, October 1983 issue (vol. 85, No. 844) , : ; : ? . 119

Notice of 1984 Joint Field Meeting of BSA (Northeast dos sieas Torrey
Botanical Club and Philadelphia Botanical Club se 19

Hovova

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 86 April 1984 No. 846

Che New England Botanical Club, Ine.

Botanical Museum, Oxford Street, Cambridge, Massachusetts 02138

Conducted and published for the Club, by
NORTON H. NICKERSON, Editor-in-Chief

Associate Editors

A. LINN BOGLE GARRETT E. CROW
WILLIAM D. COUNTRYMAN RICHARD A. FRALICK
GERALD J. GASTONY NORTON G. MILLER

ROBERT T. WILCE

RHODORA,—Published four times a year, in January, April, July, and
October. A quarterly journal of botany, devoted primarily to the flora of
North America. Price $20.00 per year, net, postpaid, in funds payable at
par in the United States currency at Boston. Some back volumes and
single copies are available. Information and prices will be furnished upon
request. Subscriptions and orders for back issues (making all remittances
payable to RHODORA) should be sent to RHODORA, Botanical
Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive the
next number of RHODORA, changes of address must be received prior
to the first day of January, April, July or October.

Scientific papers and notes relating to the plants of North America and
floristically related aras will be considered by the editorial committee for
publication. Articles concerned with systematic botany and cytotaxon-
omy in their broader implications are equally acceptable. Brevity is urged
whenever possible in all papers. Short items will be published on
otherwise blank end pages as soon as possible, even if they appear ahead
of longer articles already accepted. All manuscripts should be submitted
in TRIPLICATE AND MUST BE DOUBLE (AT LEAST 3/8 OF AN INCH) OR TRIPLE-
SPACED THROUGHOUT. Please conform to the style of recent issues of the
journal. See “Instructions to Contributors to RHODORA” at the end of
each issue. Extracted reprints, if ordered in advance, will be furnished at
cost. RHODORA assesses modest page charges.

Address manuscripts and proofs to:
Joan Y. Nickerson
Managing Editor, RHODORA
Phippen-LaCroix Herbarium, Dept. of Biology
Tufts University
Medford, Mass. 02155

Second Class Postage Paid at Boston, Mass.

PRINTED BY
THE LEXINGTON PRESS, INC.
LEXINGTON, MASSACHUSETTS

Cover illustration

Ledum groenlandicum Oeder, Labrador tea, reaches its southeastern distribu-
tional limit in Concord, Mass. It was first collected by Thoreau in 1858, subsequently
regarded as extirpated by Richard Eaton in 1974, and rediscovered by Ray Angelo in
1978. Angelo has since found it in two more Concord locations.

Original artwork by Josephine Ewing.

Tbhodora

(ISSN 0035-4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 86 April 1984 No. 846

A NEW SPECIES OF PHYLLANTHUS
(EUPHORBIACEAE) FROM THE CAYMAN ISLANDS

GRADY L. WEBSTER AND GEORGE R. PROCTOR

ABSTRACT

Phyllanthus caymanensis Webster & Proctor is described from specimens collected
in the Cayman Islands. This species resembles Mesoamerican species such as P.
mevaughii and P. mocinianus rather than any of the West Indian taxa of sect.
Nothoclema. It is the first endemic species of sect. Nothoclema recorded from the
West Indies.

Key Words: Euphorbiaceae, Phyllanthus, West Indies

In the revision of the West Indian species of Phy/lanthus, Webster
(1957) recorded only 2 species of sect. Nothoclema: the widespread
P. acuminatus Vahl, and P. subglomeratus Poir. in the Lesser
Antilles. It was therefore a distinct surprise when the junior author
discovered a third species in the Cayman Islands. Although it is very
similar in aspect to the Mesoamerican P. mocinianus Baillon, the
Cayman plant clearly represents a previously undescribed species.

Phyllanthus caymanensis Webster & Proctor, sp. nov., ab aliis
speciebus sect. Nothoclemae differt ramulis glabris simpliciter
pinnatiformibus, foliis ovatis subacutis conspicue venosis,
antheris compressis subacutis, pollinis grana striato-reticulata.

Glabrous shrub c. 2.5 m high; twigs terete, 1-2.5 mm thick,
greyish. Cataphylls + scarious, indurate, subpersistent; stipules
triangular-ovate, c. 1.2-1.5 mm long and 1.2 mm broad; blade
lanceolate, c. 1-1.2 mm long, narrower. Deciduous branchlets all
simple (pinnatiform), 3-7 cm long, with 6-10 nodes; axes greenish,
angled, slender (less than | mm thick). Leaves with stipules

121

Figure

1.

Flowers

of

Phyllanthus

caymanensis.

A.

Staminate

flower.

B. Pistillate flower.

eC

e1opoyuYy

98 ‘10A]

1984] Webster & Proctor — Phyllanthus 123

lanceolate, acuminate, becoming scarious and subpersistent, 0.8~1
mm long; petioles 1-2 mm long; blades chartaceous, ovate, obtusely
to acutely pointed at tip, obtusely rounded to slightly subcordate at
base, mostly 1.5—2.7 cm long, 1.2-2.1 cm broad, above olive green
(paler beneath), with midrib, major lateral veins (c. 5~7 on a side)
and veinlet reticulum distinctly prominulous (on both faces);
margins plane.

Monoecious; cymules axillary on branchlets, bisexual, with one
central 2 and several lateral ¢ flowers. Staminate flowers: pedicel
slender, 5-10 mm long; calyx-lobes 6, biseriate, erect, elliptic to
obovate, obtuse, with paler margins, |.3-1.7 mm long, 0.7—1.2 mm
broad; disk segments 3, massive, bilobed, pitted, c. 0.6-0.8 mm
across; stamens 3, filaments completely connate into a column
0.3-0.5 mm high; anthers triangular-ovate, acute or subacute,
flattened, c. 0.4 mm long and 0.5 mm broad, fused by the
connectives, dehiscing horizontally; pollen grains subglobose, c.
18-22 um in diameter, 3-colporate, not syncolpate, colpi distinctly
marginate, exine with even and relatively fine vermiculate orna-
mentation pattern. Pistillate flowers; pedicel slender (not dilated
distally), becoming 8-12 mm long in fruit; calyx-lobes 6, biseriate,
triangular-lanceolate, acute, |1.4-1.7 mm long; disk 3-lobed, lobes c.
0.7-0.8 mm across; ovary smooth, of 3 carpels; styles free,
spreading, c. 0.4 mm long, bifid, branches slender. Capsules oblate,
prominently veiny, greenish, 4-4.5 mm across; seeds 1.9-2 mm long,
distinctly angled, light brownish, nearly smooth.

Type: Cayman Islands, Cayman Brac, Foster Land Distr., rocky
woodland c. 0.7 mi NW of Pollard Bay, alt. c. 100 ft, 7 Aug. 1975,
G. R. Proctor 35151 (JAM, Holotype). Additional collection
examined: Cayman Islands, Little Cayman, sandy woodland just N
of W end of the air strip, 7 Aug. 1975, G. R. Proctor 35145 (JAM).

This species represents an exciting novelty because it is the first
endemic species of sect. Nothoclema to be discovered in the West
Indies. In appearance, the specimen suggests a small-leaved form of
the Mexican species P. mocinianus Baillon. However, in most floral
characters (Fig. 1), especially the anther shape, it is closer to P.
mevaughii Webster of Chiapas and northern Central America. It
differs from that species, though, in its glabrous conspicuously
veined leaves, slender pistillate pedicel, and smaller seeds. Curiously,
the pollen of the Cayman plant, as seen in scanning micrographs
(Fig. 2), is much closer to P. mocinianus in its relatively fine

124 Rhodora [Vol.

Figure 2. Pollen grains of some species of Phyllanthus sect. Nothoclema.

86

A.

Phyllanthus mevaughii. B. Phyllanthus mocinianus. C. Phyllanthus caymanensis.

Scale bar = 4 yum.

1984] Webster & Proctor — Phyllanthus 125

vermiculate sculpturing than it is to the coarser, more irregular
ornamentation of P. mcvaughii (see also figures in Webster, 1967).

Overall, P. caymanensis thus shares some characters with both P.
mevaughii and P. mocinianus, but is clearly distinct from either of
them. It remains an intriguing phytogeographical puzzle that sect.
Nothoclema should be represented by an endemic Antillean species
only on the Cayman Islands.

ACKNOWLEDGMENTS

We wish to thank the Institute of Jamaica for loan of specimens,
Dr. Steven Lynch and the Botany Department of the Smithsonian
Institution for the SEM pictures of pollen, and Mr. Norman
Geesing for preparing the illustrations.

LITERATURE CITED

Wester, G. L. 1957. Phyllanthus sect. Nothoclema: A monographic study of the
West Indian species of Phyllanthus. J. Arnold Arbor. 38: 363-371.

1967. A new species of Phyllanthus (Euphorbiaceae) from Central

America. Brittonia 18: 336—342.

Gok. Ww.

DEPARTMENT OF BOTANY
UNIVERSITY OF CALIFORNIA
DAVIS, CALIFORNIA 95616

GRP:

JARDIN BOTANICO NACIONAL

AVENIDA LOS TRINITARIOS

SANTO DOMINGO, REPUBLICA DOMINICANA

126 Rhodora [Vol. 86

AN AWARD FOR THE SUPPORT OF
BOTANICAL RESEARCH
IN NEW ENGLAND, U.S.A.

The New England Botanical Club is offering an award of $1,000
in support of botanical research to be conducted in the New
England region during 1985. It is being made to stimulate and
encourage botanical research on the New England flora and to
make possible visits to the New England region by those who would
not otherwise be able to do so. The award will be given to the
graduate student submitting the best research proposals dealing
with field studies in systematic botany and plant ecology, but
proposals for research in other areas of botany will also be
considered. This award is not limited to graduate students at New
England institutions. The NEBC’s support must be acknowledged in
any publications resulting from this study. It is encouraged that
papers based on this research be submitted to RHODORA, the
Club’s journal, for possible publication—subject to standard review
processes. The New England Botanical Club hopes to be able to
make this award on an annual basis.

Applicants should submit a proposal of no more than three
double spaced pages, including a budget (the budget will not affect
the amount of the award), and their Curriculum Vitae. Two letters,
one from the student’s major professor, in support of the proposed
research are also required. Proposals and supporting letters should
be sent before 28 February 1985 to: Awards Committee, The New
England Botanical Club, 22 Divinity Avenue, Cambridge, MA
02138. The recipient of the award will be notified by 30 April 1985.

A NEW, WIDESPREAD SPECIES OF CHA PTALIA
(ASTERACEAE: MUTISIEAE) FROM MEXICO

Guy L. NESOM

ABSTRACT

Chaptalia transiliens Nesom, a new species from Mexico, is described and assigned
to Chaptalia sect. Leria; notes on its distribution and phenology are given.

Key Words: Asteraceae (Mutisieae), Chaptalia transiliens, Mexico, Guatemalan
disjunct

In order that it may properly be dealt with in Rzedowski’s
forthcoming treatment of Compositae from the Valley of Mexico, I
propose the following new species in advance of a revisionary
treatment of all 13 species from North America, Mexico, and
continental Central America (Nesom, in prep.). Burkart’s revision
of Chaptalia (1944) cited none of the collections listed below; most
of them have been made since the completion of his study.

Chaptalia transiliens Nesom, sp. nov.

Chaptaliae nutanti (L.) Polak. affinis, a qua imprimis differt foliis
crassis spathulatis petiolis distincte et anguste alatis, phyllariis
interioribus marginibus latis, ligulis erectis angustatis involutis ad
maturitatem purpureis initio albis supra, floribus pistillatis interiori-
bus paucioribus ligulis prominentibus, ramis stylorum florum
pistillatorum aliquantum complanatis, acheniis rostris brevioribus.

Plants from a simple caudex ora slender, horizontal to ascending
rhizome up to 4 cm long. Leaves thick, sometimes. slightly
coriaceous, spatulate, the blade elliptic to ovate-elliptic, sometimes
sublyrate with several shallow lobes at the base, 6-28 cm long,
petiole 1/2-2/3 as long as the leaf, winged, blade 9-55 mm wide,
densely tawny-gray pubescent below, quickly glabrescent above,
glabrous and often somewhat shiny by flowering, margins with
numerous apiculae, not revolute. Stems 12-56 cm tall at anthesis,
elongating slightly in fruit, tawny webby-pubescent, glabrescent,
ebracteate or with a linear bract within 5 mm of the head. Heads 1-2
per plant, nodding in bud; involucre campanulate to hemispheric,
9-19 mm wide (pressed); phyllaries in 3-4 imbricated series, linear-
lanceolate with acute apices, sparsely to densely tawny-villous

127

128 Rhodora [Vol. 86

except for margins and narrow midregion, inner 14-20 mm long at
anthesis, not elongating in fruit, 1.3-1.8 mm wide, sometimes purple
margined, developing wide (0.2-0.5 mm ), flange-like, scarious
margins below the upper third and often becoming involute,
margins of outer |-3 series usually minutely stipitate-glandular,
outermost 1/4—1/2 as long as the inner; receptacle flat, weakly
foveolate. Flowers trimorphic in 3 concentric zones, all fertile; outer
pistillate flowers 11-21 (32) in I series, erect, corolla 11-17 mm long,
bilabiate, the inner lip microscopic (0.5 mm long) to 2.5 mm long
and equalling the style in length, ligule white above, reddish-purple
below, usually maturing purplish above, 0.8-1 mm wide, 3—5 veined,
usually involute and often enclosing the style, apex truncate to
shallowly lobed or notched, tubular portion 5-7 mm long, style
9.2-11 mm long with somewhat flattened branches 1-2 mm long,
often purple; inner pistillate flowers rare to nearly as numerous as
the outer ones, ligulate with ligules shorter than those of the outer
but sometimes longer then the style, red to cream, bilabiate with an
inner lip up to 1 mm long; hermaphroditic (disc) corollas in center
of head, yellowish, narrowly oblanceolate in outline, 9-11 mm long,
bilabiate, longest lobes 2.2-3.5 mm long, erect to spreading, anthers
with thecae 1.5-2.3 mm long, apical appendages 0.8-1.2 mm long,
style branches narrowly obovate, 0.8-0.9 mm long. Achenes 7.5-11
mm long at maturity, the body fusiform, slightly flattened, tan to
brown, 5-6 mm long, 0.9-1.3 mm wide, 5-6 ribbed, moderately
pubescent with prominently attenuate-apiculate papillae, beak light-
colored, (0.6) 0.9-1 times as long as the body, upper part minutely
stipitate-glandular, carpopodium narrow; pappus tawny-white,
bristles weakly barbellate, 9.5-12 mm long, longer than the disc
corollas.

DISTRIBUTION AND PHENOLOGY. Scattered localities in southcentral
Nuevo Leon, Guanajuato, Hidalgo, México, and Puebla, appar-
ently disjunct to Chiapas and Guatemala; usually on steep, moist,
shaded, often rocky slopes, with pine, oak, pine-oak, or pine-oak-
juniper, in Nuevo Leon on “open, dry slopes among thorny shrubs”:
1600-2550 m; flowering (January) April through October (Novem-
ber).

TYPE) MEXICO. NUEVO LEON: NW slope of Cerro Pefia
Nevada on road to pass, ca. 1.5 km directly NW of summit, ca. 35
km ENE of Doctor Arroyo, steep slope, oak-pine-juniper with

1984] Nesom — Chaptalia 129

Agave; common but scattered, in shade, deep soil, usually under
oaks, 31 Jul 1983, Guy Nesom 4759 (HOLOTYPE: US;
ISOTYPES: [to be distributed] ANSM, CAS, ENCB, GH, K,
MEXU, MICH, MO, NY, OS, SMU, TEX, UC).

ADDITIONAL COLLECTIONS EXAMINED: GUATEMALA: Chimal-
tenango, San Martin Chile Verde, | Aug 1941, Johnston 1765 (F);
along Aguacatan road E of Huehuetenango, at KM 15, 2 Jan 1941,
Standley 81939 (F); canyon above Chiantla, Sierra de las Cuchu-
matanes, 9 Aug 1942, Steyermark 50351 (F). MEXICO. Chiapas:
mpio. Tenejapa, W of Tenejapa Center along trail to Paraiso, 5 Aug
1964, Breedlove 6870 (DS); mpio. La Independencia, 6-10 km NNE
of La Soledad along logging road from Las Margaritas to Campo
Alegre, 24 Oct 1976, Breedlove 41026 (DS); 6-10 km NNE of La
Soledad, 26 Nov 1980, Breedlove & Almeda 47775 (CAS); mpio.
San Cristébal las Casas, Cerro San Cristobal in San Cristdbal, 23
Apr 1966, Laughlin 740 (DS, MICH); Mt. Male, Porvenir, 6-12 Jul
1941, Matuda 4686 (MEXU, MO, NY); Amatenango de Valle, 12
Jun 1945, Matuda 18243 (MEXU). Guanajuato: camino a San
Miguel pasando frontera con Qro., 12 Apr 1981, Argtielles 1574
(MEXU); ca. 8 km NE of Santa Rosa [Hwy. 110], 10 Nov 1970 (past
flower), Mcvaugh 24173 (NY). Hidalgo: ca. 0.4 km N of Minas
Viejas, 3.2 km NE of Durango on Hwy. 85, ca. 32 km S of Jacala, 8
Aug 1981, Nesom 4377 (LL, MEXU, US). Mexico: 4 km N de
Atizapan, 21 Jul 1968, Rzedowski 26002 (ENCB). Nuevo Leon: 0.8
km NE of Dulces Nombres, near Tamaulipas border, 18 Jun 1948,
Meyer & Rogers 2575 (MO, US); Lower San Francisco Canyon, ca.
24 km SW of Pueblo Galeana, 18 May 1934, Mueller & Mueller 437
(GH, MICH, TEX, US). Puebla: mpio. Tecamachalco, El Ocotal,
11 Jul 1972, Ventura A. 5719 (ENCB, MICH); 8.5 km SW of San
Salvador el Seco, 13 Jul 1960, Beaman 3620 (MSC); Boca del
Monte, Tehuacan, Jun 1907, Purpus s.n. (UC).

Chaptalia transiliens possesses ebracteate scapes, nodding buds,
narrow ligules without a definite midstripe, relatively long and
narrow pistillate style branches, and fertile disc achenes; these
characters identify it as a member of Chaptalia sect. Leria and a
member of the species group that includes the widespread and well-
known C. nutans (L.) Polak., with which it has usually been
confused. It is not clear at present, however, which species within
sect. Leria C. transiliens is most closely related to. Tentative

130 Rhodora [Vol. 86

identifications of the new species can be made on the basis of the
relatively thick or slightly coriaceous, spatulate leaves with narrow,
winged petioles. Other distinctive features of this species are: 1) long
phyllaries, the outer with stipitate-glandular margins, inner with
wide, scarious, sometimes flange-like margins; 2) long, narrow,
involute ligules, white above and reddish-purple below at early
anthesis, completely purplish at maturity; 3) inner pistillate flowers
with short but prominent ligules; 4) somewhat flattened pistillate
style branches; and 5) achenes with a narrow beak about as long as
the body, pubescent with prominently attenuate-apiculate papillae.

The epithet “transiliens” refers to the “leap” shown by the new
species in geographic distribution from Mexico and Puebla south-
eastward to Chiapas and Guatemala.

ACKNOWLEDGMENTS

For their assistance I thank the curators of the following herbaria
from which specimens were borrowed: CAS, DS, ENCB, F, GH,
MEXU, MICH, MO, MSC, NY, TEX, UC, and US. Dr. G. J.
Gastony provided editorial assistance with the Latin diagnosis.

LITERATURE CITED

BURKART, A. 1944. Estudio del genero de Compuestas Chaptalia con especial
referencia a las especies argentinas. Darwiniana 6: 505—594.

DEPARTMENT OF BIOLOGY
MEMPHIS STATE UNIVERSITY
MEMPHIS, TENNESSEE 38152

NOTES ON CROOMIA PAUCIFLORA (STEMONACEAE)!
R. DAVID WHETSTONE

ABSTRACT

Croomia pauciflora (Nuttall) Torrey is an endemic to the southeastern United
States. Documented distribution includes the following states and physiographic
regions: Ridge & Valley, Cumberland Plateau, Piedmont Plateau, and Gulf Coastal
Plain of Alabama; Piedmont Plateau, and Gulf and Atlantic Coastal Plain of
Georgia; and the Gulf Coastal Plain of Florida and Louisiana. Croomia pauciflora is
a rare element of the southeastern flora and is considered an “endangered” or
“threatened” species in portions of its range. A neotype is designated due to the
apparent loss of the single specimen cited in the original description.

Key Words: Croomia pauciflora, Stemonaceae, neotype, endemic, disjunct, en-
dangered species, threatened species, rare species.

The Stemonaceae is comprised of four genera with 30-35 species
(Hutchinson, 1973; Willis, 1973; van Steenis, 1982). The genus
Croomia is highly celebrated as being disjunct from the south-
eastern United States to southeastern Asia. Three species are
currently recognized with one in the United States, Croomia
pauciflora (Nuttall) Torrey, and two in Japan, C. heterosepala
(Bak.) Okuyama and C. japonica Miq. (Ohwi, 1965). Rogers (1982)
reported the range of the latter to extend to eastern China.

Hardy Bryan Croom first discovered Croomia across the
Apalachicola River from his home. Specimens were sent to Thomas
Nuttall who named the new plant Cissampelos pauciflora (Nuttall,
1834). Apparently unaware of Mr. Nuttall’s publication of the new
name, but cognizant that Nuttall referred the plant to the
Menispermaceae, Croom (1835) provided a description and duly
noted the new taxon should be classified elsewhere. John Torrey,
friend and correspondent, recognized the distinction of the herb and

| These Notes were developed while preparing a treatment of the Stemonaceae for the
Vascular flora of the southeastern United States (Radford, in prep.). Specimens
consulted during this study are housed in the following herbaria: AUA, BH, CU,
DUKE, FSU, GA, GH, JSU (Jacksonville State University Herbarium), LSU, MO,
NCU, NY, PH, SMU, TENN, TEX, UNA, US, USF, VDB, and the University of
South Alabama. Acronyms are from Holmgren er al. (1981), unless otherwise
specified. Exsiccata are not further cited because of the rarity of this species. A list of
specimens examined is deposited in libraries at A/GH and MO.

131

132 Rhodora [Vol. 86

established the genus Croomia in honor of the discoverer (Torrey
and Gray, 1840). However, Torrey placed Croomia within the
Berberidaceae.

Nuttall (1834) mentioned a single collection made by Croom and
Loomis. No specimens labeled thus were located at the Academy of
Natural Sciences (pers. comm., Dr. A. E. Schuyler, 1983) or at the
British Museum (pers. comm., John Lewis, 1979). It should be
noted the title of the article specifically states the specimens are at
PH. Torrey (Torrey and Gray, 1840) cited two collections, “Mr.
Croom!” and “Dr. Chapman!”. Two sheets labeled “Croom” are at
the New York Botanical Garden, one with flowers and one with
buds. In the type collection at New York, a manuscript copy of the
original description of Croomia and the ensuing comb. nov. reveals
the following within the habitats section: “Under the shade of
Torreya taxifolia Arn., at Aspalaga on the Apalachicola, Florida,
H. B. Croom, Esq.—Flowers in April.” Several “Herb. Chapman”
exsiccata at NY and GH may be materials cited by Torrey. One
sheet examined by this author is labeled “Torr. and Gr. Fl. N.
Amer.” Quite possibly, this specimen represents material sent to Dr.
Torrey by Hardy Croom in 1830 (fide a letter addressed to Dr.
Torrey in the archives collection at PH). Torrey acknowledged
receipt of specimens from Mr. Croom in the subsequent publication.
Since no specimen exists at PH or at BM which is indisputably the
single specimen cited by Nuttall (1834), it becomes necessary to
designate a neotype. In the letter (mentioned above), Croom notes
he collected some specimens from along the Apalachicola River at
Aspalaga. I have chosen a collection made by Dr. Godfrey from
Torreya State Park which is ca. 8 km south of Aspalaga. The
neotype is labeled as follows: “FLORIDA, LIBERTY COUNTY.
Wooded bluffs along the Apalachicola River at Torreya State Park,
16 March 1961, R. K. Godfrey 60614, FSU 63997” (Figure 1).
Replicates of this collection which were examined are at BH (s.n.);
DUKE (147433); GA (63218 & 106380); NCU (216705); SMU (s.n.):
TENN (2 sheets, s.7.); and US (238915).

Familial placement has likewise had a rather colorful history
ranging from the Menispermaceae (Magnoliopsida) to the Croomi-
aceae (Liliopsida). Recent phylogenists (Cronquist, 1981; Hutchin-
son, 1973, Takhtajan, 1980) place the genus in the Stemonaceae
(Roxburghiaceae) although evidence (Ayensu, 1968) suggests that
this group remains rather heterogenous.

1984] Whetstone — Croomia

133

HT

m1

eS ™ x
_</

’ "

HERRARIOM OF THE UNIVERSITY OF NORTH CARCHINA
Croomie pauciflore (Muttall) Torrey

&. D. Whetetone August, 197%

HERBARIUM OF FLORIDA STATE UNIVERSITY
Tallahassee

PLANTS OF FLORIDA
COUNTY :LIBERTT March 14, 1961
Croomie pauciflora (Mutt.) Torr.
Wooded bluffe along the Apalachicola River at Torreya
State Park.

Collected by R, K, Godt
Ko, 6061) Det. mo

Figure 1. Neotype of Croomia pauciflora.

134 Rhodora [Vol. 86

Most of the known localities for Croomia pauciflora are along
watercourses that have headwaters in the southern Appalachian
Mountains and flow southward to the Gulf of Mexico. Two
collections deviate from this pattern. A specimen at NY was
collected by Pond along the Savannah River which debouches into
the Atlantic Ocean. More noteworthy is a collection by Featherman
in 1870 (LSU 34679). Label data indicate “Brashear City” which is
in St. Mary Parish, Louisiana. This parish is located west of the
Mississippi River. The primary drainage, the Red River, has
headwaters in the Ozarks and eastern Texas although there is a
confluence with the Mississippi River (some western Appalachian
drainage) near Torras. Quite possibly other localities exist in the
highlands west of the Mississippi River, perhaps in east Texas or
Arkansas.

The greatest diversity of physiographic distribution occurs in
Alabama where Croomia has been located in the Cumberland
Plateau, Valley and Ridge, and Piedmont Plateau sections of the
Appalachian Highlands Province and in the Gulf Coastal Plain
Province. Georgia localities include Gulf and Atlantic slopes of the
Coastal Plain Province. Populations in Florida and Louisiana are
within the Gulf Slope of the Coastal Plain Province. See Figure 2
for the documented county distribution.

Croomia is known from a variety of substrates but is found
chiefly in rich, sandy or rocky soils of wooded slopes and bottoms.
Most frequently, the slope is sheltered (e.g., ravines) or north-
facing. Soils are ostensibly circumneutral although not necessarily
calcareous (vide discussion in Harper, 1922). All habitats described
on labels and those visited by this author are mixed deciduous
forests which are mesic.

Presumably, Croomia pauciflora is an epibiotic, an endemic
considerably restricted from a former, much broader range.
Presently the taxon is very much disjunct in isolated localities in the
warmer temperate portions of the southeastern United States. From
personal observations, populations appear to be largely clonal.
Also, current distribution is possibly a result of a constriction of
much broader distribution owing to colder Pleistocene temperatures
and limited habitat availability south of the Appalachian Highlands
Province.

Despite the number of populations (some historical and perhaps
extirpated) throughout the range, Croomia is a rare element of the

1984] Whetstone — Croomia 135

pel OPE TEN Vt fk
|

aa
j
’

Figure 2. Documented county distribution of Croomia pauciflora.

southeastern flora. Croomia is considered “threatened” in Alabama
(Freeman ef al., 1979), “endangered” in Florida (Godfrey & Ward,
1979) and “threatened” in Georgia (McCollum & Ettman, 1977).
The “endangered” status in Florida is because of damage inflicted
on populations by feral pigs. Godfrey and Ward (1979) stressed the
importance of controlling the animals to prevent destruction of the
populations along the Apalachicola River bluffs. Much more
difficult would be the assessment of damages to populations that
were directly or indirectly affected by the impoundments created
along the numerous watercourses in Alabama, Florida, and
Georgia.

ACKNOWLEDGMENTS

I am indebted to the curators of the herbaria cited herein for the
consultation of materials under their purview. I also appreciate the
search of the manuscript collection of the Academy of Natural
Sciences by Carol M. Spawn, the type collection at PH by A. E.
Schuyler, the type collection at NY by P. K. Holmgren, and the
collections of the British Museum by John Lewis. T. A. Atkinson
and J. R. Massey provided assistance with the processing of

136 Rhodora [Vol. 86

materials as well as interesting discussions of facets of this work. In
addition, D. E. Boufford provided sound advice and bibliographic
assistance. Melanie G. Bussey very patiently and carefully typed the
manuscript. Mr. Opal R. Lovett photographed the neotype. All
conclusions and facts contained herein reflect the observations and
opinions of the author. Portions of this research were performed
while the author held a Coker Fellowship in the Department of
Botany at the University of North Carolina at Chapel Hill. | am
indeed grateful for the financial support.

LITERATURE CITED

Avenst, E. SS. 1968. Comparative vegetative anatomy of the Stemonaceae (Rox-
burghiaceae). Bot. Gaz. 129: 160-165.

Cronguist, A. 1981. An integrated system of classification of flowering plants.
Columbia University Press, New York.

Croom, H. B. 1835. Botanical communications. Amer. J. Sci. 28: 165 168.

FREEMAN, J. D., A. S. Causey, J. W. SHorT, AND R. R. Haynes. 1979. Endan-
gered, threatened, and special concern plants of Alabama. Department of Botany
and Microbiology, Agricultural Experiment Station, Auburn University, Depart-
mental Series No. 3.

Goprrey, R. K. AND D. B. Warp. 1979.) Croomia. In D. B. Ward, editor, Rare
and endangered biota of Florida, Vol. 5. Plants. University Presses of Florida,
Gainesville.

HARPER, R. M. 1922. A botanical bonanza in Tuscaloosa County, Alabama. J.
Elisha Mitchell Sci. Soc. 37: 153-160.

HoOLMGREN, P. K., W. KEUKEN, AND FE. K. SCHOFIFLD. 1981. Index herbariorum.
Part 1. The herbaria of the world. Seventh edition. Bohn, Scheltema and
Holkema, Utrecht.

HutcuHiInson, J. 1973. The families of flowering plants. Third edition. Clarendon
Press, Oxtord.

McCotiuMm, J. Lk. AND D. R. ETTMAN. 1977. Georgia’s protected plants. Georgia
Department of Natural Resources, Atlanta.

NuTTALL, T. 1834. A description of some of the rarer or little known plants
indigenous to the United States...Acad. Nat. Sci. Philadelphia J. 7: 61-115.

Ouwl, J. 1965. Flora of Japan. Smithsonian Institution. Washington, D. C.

RapDFoRD, L. S., executive editor. Vascular flora of the southeastern United States.
The University of North Carolina Press, Chapel Hill. (In preparation).

RoGers, G. K. 1982. The Stemonaceae in the southeastern United States. J.
Arnold Arbor. 63: 327-336.

TAKHTAJAN, A. L. 1980. Outline of the classification of flowering plants. Bot.
Rev. (Lancaster) 46: 225-359.

TorRREY, J. AND A. Gray. 1840. A flora of North America. Vol. I., Part 4.
(Facsimile of the 1838-43 edition, 1969). Hafner Publishing Company, New
York.

1984] Whetstone — Croomia 137

VAN STEENIS, C. G. G. J. 1982. Pentastemona, a new 5-merous genus of mono-
cotyledons from north Sumatra (Stemonaceae). Blumea 28: 151-163.

WILLIs, J.C. 1973. A dictionary of flowering plants and ferns. Eighth edition,
revised by H. K. Airy Shaw. Cambridge University Press, Cambridge.

HERBARIUM, DEPARTMENT OF BIOLOGY
JACKSONVILLE STATE UNIVERSITY
JACKSONVILLE, AL 36265

DISTRIBUTION AND ECOLOGICAL CHARACTERISTICS
OF IRONWOOD, OSTRYA VIRGINIANA
(MILLER) K. KOCH, IN NORTHEASTERN
NOVA SCOTIA

K. N. H. GREENIDGE

ABSTRACT

Ironwood, Ostrya virginiana (Miller) K. Koch, an Alleghanian species, reaches the
northeastern limit of its range in northern Nova Scotia. The species is in a vigorous
condition and widely distributed in the region, but normally reflects low density of
stocking. Notable silvical characteristics of ironwood in northern Nova Scotia
include high tolerance, wide ecological amplitude, vitality and aggressiveness.

Key Words: Ironwood, Nova Scotia, range limit, ecological amplitude, density

Ironwood, Ostrya virginiana (Miller) K. Koch, is one of a number
of tree species, mainly of Canadian or Alleghanian affinity, which
reach the northeastern limit of their ranges in northern Nova Scotia.
Other taxa reflecting this pattern of distribution include Populus
grandidentata', Quercus rubra var. borealis, Acer saccharum,
Fraxinus americana and Tsuga canadensis (Little, 1971, Scoggan,
1978.)

Available information suggests the possiblity of considerable
differences in local distribution, abundance and habitat-selection in
northern Nova Scotia among elements of the foregoing complex of
species (Roland and Smith, 1969). Accordingly, studies were
initiated with the object of gaining detailed information on the
occurrence, importance and ecological relations of Canadian-
Alleghanian species in the region. Observations on Acer saccharum
have been published previously (Greenidge, 1977). The purpose of
the present paper is to describe and discuss the local distribution
and silvical characteristics of ironwood over a broad range of sites
in the northeastern region of the Province.

METHODS

Field work was concentrated in topographically-diverse areas of
Nova Scotia bordering the southern reaches of the Gulf of St.

'Nomenclature follows Fernald (1950)

Lay

140 Rhodora [Vol. 86

Lawrence (Figure 1). This region suffered extensive Pleistocene
glaciation (Prest and Grant, 1969). However, the possibility of the
existence in late-Wisconsin time of upland and coastal refugia has
been recognized by Grant (1977).

Extensive new collections of ironwood were made to increase
knowledge of its distribution in northen and northeastern Nova
Scotia’. Reconnaissance and intensive surveys, transects, line
(strip)-cruises and sample plots were used to investigate the
occurrence, density, morphological characteristics, habitat pref-
erences, crown-class relations, tolerance, associated species, vigor,
successional tendencies and reproductive potential of ironwood
over a wide spectrum of sites.

Observations were made both in forested and partially-forested
areas of the region, with emphasis on conditions in the closed forest.
Transects utilized to investigate the occurrence and field-relations of
the species on topographically well-defined sites were oriented at
right angles to the contours on hill-slopes, and along the axes of
spurs, stream-hollows and valley-heads. Surveys and searches were
employed for the same purpose on sites of limited topographic
variability, including upland-surfaces, terraces and bottom-lands.
Quantitative data on density (trees per acre), dominance (basal area
per acre) and stand-composition were accumulated with the aid of
strip-cruises and sample plots.

RESULTS

DISTRIBUTION. Figure 2 illustrates the pattern of distribution of
ironwood in Nova Scotia. This map incorporates information on
collections housed in several herbaria: ACAD, DAL, NSPM,
NSAC, UNB, GH, CAN, and DAO. Also included in Figure 2 are
the results of the author’s collection of ironwood in the Gulf of St.
Lawrence-Cabot Strait area of Nova Scotia.

With specific reference to northeastern Nova Scotia, a study of
available collections suggests that the species becomes less frequent
northeastward. Thus far, ironwood apparently has not been
collected north of the Cheticamp River watershed, and no
collections of the species from Victoria and Richmond Counties are

*These materials have been deposited in NSPM.

1984] Greenidge — Ironwood 141

Uj

GULF

GASPE OF
PENINSULA ST. LAWRENCE
Teas OO 4 is S.
' ~ MAGDALENE Go
H ta
} ISLANDS Lf
f % mor,
/ % CAPE
£ 7 } PRINCE EDWARD BRETON
rh ' NEW ISLAND ISLAND
J | BRUNSWICK .
f H 7
i) H
| ) E/
f if" ¢ v
ra om
a8 ¢ re) 50 100
al H ow
‘ oi
ont NAUTICAL MILES
hs ¥
és —

ATLANTIC OCEAN
N

fs

Figure 1. Map of the Gulf of St. Lawrence—Atlantic Provinces Area of Eastern
North America.

known to the author. However, a report by Bulmer and Hawboldt
(1958) treating the forest resources of Nova Scotia noted the
occurrence of ironwood in both of the above-mentioned counties.

OCCURRENCE AND HABITAT PREFERENCES. In areas of discontinuous
or partial forest-cover in northern Nova Scotia, ironwood occurs as
scattered individuals or in small concentrations in riparian stands
and thickets, along roadsides and the edges of fields, and in cut-over
areas, power-line clearings and similar types of disturbed sites.
In forested areas of northern mainland Nova Scotia, the species
exploits a broad range of sites, and reflects modest, occasionally
considerable, abundance in light soils on the variable topography
immediately south of the Gulf of St. Lawrence. Scattered indi-

CHETICAMP RIVER

LEGEND (A)
8 NORTHEAST MARGAREE RIVER

BARNEY'S RIVER

7 LUNENBURG

Q Mites 36

/
YARMOUTH /
if
j SHELBURNE *

Figure 2. Distribution of Ostrya virginiana in Nova Scotia.

Chl

BIO poyYy

98 ‘10A]

1984] Greenidge — Ironwood 143

viduals or occasional small concentrations of the species occur on
saddles and upland-surfaces to elevations of approximately 750 feet.
The species also occurs on the flanks, axes and toes of spurs, on
valley-side slopes of all exposures, on flood-plains, on outwash and
alluvial-terraces, and on low, protected, off-shore islands in the
lower Barney’s River region of Pictou County.

The species is uncommon on lowland sites in the region marked
by the presence of heavy, fine-textured soils.

To the northeastward, on the approaches to the limit of its range
in Cape Breton Island (Figure 2), ironwood continues to demon-
strate modest ecological amplitude. The species is scattered and
infrequent in the topographically-varied, forested landscape of the
lower Cheticamp River, occurring on flood-plains, alluvial-terraces,
spurs, and saddles, and on east, southeast, south, southwest and
west-facing slopes. South of the Cheticamp River, in the Margaree
watershed, the species has been collected on forested alluvial-
terraces and in meadow thickets.

DENSITY AND STAND RELATIONS. Ironwood generally constitutes a
secondary species in contemporary forests of northern and north-
eastern Nova Scotia. The species is found very rarely in old-growth
forests, infrequently in disturbed or deteriorating older stands, and
more commonly in young, disturbed or undisturbed, hardwood and
mixedwood forests. Density commonly is low, with somewhat
higher levels of stocking evident in upland areas of northern Pictou
and northernmost Antigonish Counties, and north-central Col-
chester County.

Occasionally, however, ironwood reflects considerable abun-
dance over limited areas in northern Nova Scotia, and achieves
locally the status of a primary species. Illustrations of this tendency
toward greater densities are presented in Tables | and 2. These
tables summarize results obtained from sample-plots established in
northern Pictou County for the purpose of documenting high
densities of stocking in ironwood and to indicate its ecological
tolerance. Attention is invited to the occurrence and abundance of
ironwood on the alluvial-terrace in association with relatively
intolerant pioneer species, and to its considerable presence on the
upland-surface in combination with long-lived, tolerant species.

MORPHOLOGICAL FEATURES. In the closed forests of northern Nova
Scotia individual ironwood trees may reach diameters at breast

Table 1. Composition, density and basal area (BA) relations in a 0.1 acre plot established on an alluvial-terrace on the
lower reaches of Barney’s River, Pictou County, Nova Scotia.

Service

Ironwood W. birch W. ash W. spruce Grey birch berry T. aspen Totals
Species No. BA(ft2) No. BA(ft.2) No. BA(ft2) No. BA(ft2) No. BA(ft.2) No. BA(ft2) No. BA(ft.2) No. BA(ft.?)
DBH (in.)
4 3 0.26 1 0.09 1 0.09 3 0.26 4 0.35 2 0.17 1 0.09 1S) 1.31
5 7 0.95 2 0.27 2 0.27 11 1.50
6 7 1.37 1 0.20 1 0.20 9 1.77
7 4 1.07 2 0.53 6 1.60
8 1 0.35 1 0.35
9 2 0.88 1 0.44 3. 1.33
10
1] 2 1.32 2 1.32
12
Totals 23 4,53 5 1.00 5 1.17 7 1.85 4 0.35 2 0.17 1 0.09 47 9.18
Percent-
ages 49 49 1] 11 11 13 15 20 9 4 4 2 2 l - -
Ironwood: Ostrya virginiana Service berry: Amelanchier sp.
White birch: Betula papyrifera Trembling aspen: Populus tremuloides
White ash: Fraxinus americana White spruce: Picea glauca

Grey birch: Betula populifolia

prl

elo poyy

98 I9A]

Table 2. Composition, density and basal area (BA) relations in a 0.1 acre plot established on an upland-surface, Barney’s
River watershed, Pictou County, Nova Scotia.

Ironwood Sugar maple White ash Beech Totals

Species No. BA(ft.?) No. BA(ft.?) No. BA(ft.?) No. BA(ft.?) No. BA(ft.?)
DBH (in.)

4 3 0.26 9 0.79 12 1.05

5 6 0.82 4 0.55 270,27 12 1.64
6 8 1.57 5 0.98 5 0.98 2.40:39 20 3.92

7 5 1.34 5 1.34 1 0.27 1 0.27 12 3.22

8 1 0.35 1 0.35 7 2.44 9 3.14
9 1 0.44 3 1.33 4 1.77
10 1 0.55 1 0.55
I] 1 0.66 1 0.66 2° 132
12

Totals 24 5.00 26 S.11 17 5.57 5 0.93 72 16.61
Percentages 33 30 36 31 24 34 7 6 = =
Ironwood: = Ostrya_ virginiana

Sugar maple: Acer saccharum
White ash: Fraxinus americana
Beech: Fagus grandifolia

[P861

poomuol] — adptusain

Sv

146 Rhodora [Vol. 86

height of 14 inches (36 cm) and heights of 55 feet (17 m). Poorly-
formed, much-branched, damaged individuals of still greater
diameter occasionally are encountered. Vigorous, co-dominant trees
on good sites attain ages of approximately 95 years.

Tree-form in mature, forest-grown material ranges from moder-
ately good to poor. Lean and crook in varying amounts typify many
stems, although a few trees develop clear, straight boles. Individual
trees may be single or multi-stemmed, often with long, round-
topped crowns of varying width. Growth-habit is excurrent at the
outset, commonly with numerous laterals, subsequently becoming
deliquescent, but with considerable variation in branching relations
from tree to tree. Branches range in size from slender to stout and in
orientation from ascending to orthogonal, with the lowermost
branches living or dead. Epicormic branching is very common.

ASSOCIATE SPECIES. On flood-plains, alluvial-terraces and foot-
slopes ironwood occurs in association with conifers such as Abies
balsamea, Tsuga canadensis and Picea glauca. Common dico-
tyledonous associates of ironwood in valley-bottoms and coves
include Betula papyrifera, Betula lutea, Fagus grandifolia, Ulmus
americana, Acer saccharum, Acer rubrum, Acer pensylvanicum and
Fraxinus americana. Populus balsamifera and Quercus rubra var.
borealis are infrequent bottom-land associates of ironwood.

On mid-slopes and crest-slopes the complex of species associated
with ironwood is very similar to that characteristic of bottom-lands.
Two minor variations involve Ulmus americana, which is very
uncommon on valley sides, and Picea rubens, somewhat uncommon
on bottom-lands but found more frequently with ironwood on both
mid-slopes and crest-slopes. A major variant, noted only once,
involves the occurrence of ironwood with Pinus strobus in
combination with Quercus rubra var. borealis and Tsuga canadensis
on an open, south-facing crest-slope in northeast Pictou County.

Ironwood occasionally forms local concentrations on mainland
upland-surfaces at elevations below approximately 800 ft. Under
these circumstances the associated species commonly are few in
number and include Fagus grandifolia, Acer saccharum and
Fraxinus americana. More commonly ironwood occurs as a
scattered tree on mainland upland sites, in varying mixtures with
Picea glauca, Picea rubens, Tsuga canadensis, Abies balsamea,
Betula lutea, Betula papyrifera, Fagus grandifolia, Fraxinus amer-
icana, Acer saccharum, Acer rubrum and Acer pensylvanicum.

1984] Greenidge — Ironwood 147

CROWN-CLASS AND REPRODUCTIVE RELATIONS. Ironwood occurs
both as a canopy and as an understory element in the closed forests
of northern Nova Scotia. In old-growth and older second-growth
stands the species is rare, but has been noted in openings as well as
in stands reflecting disturbance or deterioration. Under the latter
circumstances the crown-class of individual ironwood stems ranges
from suppressed to intermediate or isolated. In second-growth
stands ironwood may be present in the canopy as co-dominant and
intermediate crown-class trees, and below the canopy as suppressed,
i.e. Overtopped trees.

Ironwood reproduction was noted in 59 percent of the observing
stations established in this study, the species occurring either in
advance-growth or seedling stages, and often in both phases.

DISCUSSION

Ironwood, although apparently absent from Newfoundland
(Rouleau, 1956, 1978) and Prince Edward Island (Scoggan, 1978), is
widely distributed in northeastern Nova Scotia. Density commonly
is low, the species occurring usually as scattered, single or multi-
stemmed trees. Infrequently, considerably higher than normal
densities of ironwood may be observed in north-central Nova
Scotia, the species then constituting a primary element over small
areas in young, second-growth, mixedwood and hardwood forests
in this region.

A feature of the distribution and ecology of ironwood in the
closed forests of northern Nova Scotia is its occurrence over a broad
spectrum of sites in association with a large number of broad-leaved
and coniferous elements. The species occurs both in the overstory
and understory of young, second-growth forests in the region, and
demonstrates the capacity for establishment and development under
both continuous and discontinuous canopies. This distribution-
occurrence-behaviour pattern is indicative of a tolerant species of
substantial ecological amplitude characterized by considerable
Vitality.

In partially-forested areas of the region, ironwood frequently
becomes established on disturbed sites. The species may be found in
young, developing forests, and in thickets, hedge-rows and similar
vegetation types. This capacity for establishment and continued
development on disturbed sites suggests an aggressive, vigorous
species adapted to a wide range of environments. The near absence

148 Rhodora [Vol. 86

of ironwood from closed, old-growth forests may be a reflection of a
modest life-span, whereas its presence in decadent or disturbed
older stands suggests the capacity to invade and develop in gaps and
openings.

The species appears to be in a sound condition throughout its
area of occurrence in northern Nova Scotia. Flowering and fruiting
materials have been observed and collected at numerous stations in
the region, including several on the lower Cheticamp River
watershed where the species appears to approach the northeastern
limit of its range. Regeneration in both seedling and advance-
growth phases is common in northern Nova Scotia; mature trees
attain heights and diameters approaching those reached by indi-
vidual stems in the northeastern United States (Gleason, 1952:
Hough, 1907; Hui-lin Li, 1972).

ACKNOWLEDGMENTS

The author gratefully acknowledges the financial aid and logistic
assistance provided by J. L. Martin and A. A. Wilson of the Nova
Scotia Museum in support of the field work on which this paper is
based.

My grateful thanks are extended also to W. H. Drury, D. A.
Livingstone and I. C. M. Place who read an earlier version of the
manuscript, and made helpful and useful suggestions for its
improvement.

LITERATURE CITED

Butmer, R. M., AND L. S. HAwsotptT. 1958. The forestry resouces of Nova
Scotia. Department of Lands and Forests, Province of Nova Scotia, Halifax.

FERNALD, M. L. 1950. Gray’s manual of botany. ed. 8. American Book Co., New
York.

GLEASON, H. A. 1952. New Britton and Brown Illustrated Flora of Northeastern
United States and Adjacent Canada. vol. 2. New York Botanical Garden, New
York. (Second printing, slightly revised. 1958).

GRANT, D. R. 1977. Glacial style and ice limits, the Quaternary stratigraphic
record, and changes of land and ocean level in the Atlantic Provinces, Canada.
Geogr. Phys. Quat. 31: 247-260.

GREENIDGE, K. N. H. 1977. Silvical characteristics of sugar maple, Acer
saccharum, in northern Cape Breton Island. Jour. Arnold. Arb. 58: 307-324.

HouGu, R. B. 1907. Handbook of the trees of the Northern States and Canada.
The Macmillan Company, New York.

1984] Greenidge — Ironwood 149

Li, H. L. 1972. Trees of Pennsylvania. University of Pennsylvania Press,
Philadelphia.

LitTLe, E. L. 1971. Atlas of United States trees. Vol. I. U.S. Dept. Agr. Misc.
Publ. 1146. Washington. I] pp., 202 maps, 9 overlays.

PresT, V. K., AND D. R. GRANT. 1969. Retreat of the last ice sheet from the
Maritime Provinces-Gulf of St. Lawrence region. Geol. Surv. Canada Pap.
69-33: I-15.

ROLAND, A. E., AND E. C. SmitH. 1969. The flora of Nova Scotia. Nova Scotia
Museum, Halifax.

ROuLEAU, E. 1956. A check-list of the vascular plants of the Province of
Newfoundland. Contr. Inst. Bot. Univ. of Montreal 69: 41-106.

1978. List of the vascular plants of the Province of Newfoundland
(Canada). Oxen Pond Botanic Park. St. John’s, Newfoundland.

ScoGGAn, H. T. 1978. The flora of Canada. pt. 3. National Museum of Natural
Sciences, National Museums of Canada, Ottawa. p. 547-1115.

DEPARTMENT OF BIOLOGY
SAINT FRANCIS XAVIER UNIVERSITY
ANTIGONISH, NOVA SCOTIA B2G 1CO

ANNOTATED LIST OF MINNESOTA CARICES,
WITH PHYTOGEOGRAPHICAL
AND ECOLOGICAL NOTES

GERALD A. WHEELER AND GERALD B. OwWNBEY

ABSTRACT

One hundred forty-two species of Carex are recorded for Minnesota, 4 of which
have not been previously reported. Because four floristic provinces converge in
Minnesota, certain groups of carices tend to be associated with specific geographic
areas of the state. A map is presented that divides Minnesota into 9 areas of Carex
affiliation, each of which is briefly discussed. Our carices also display other, more
subtle and local, distributional patterns, and we demonstrate, through various
examples, that many carices are greatly affected by the nature of the substrate in
which they grow. Habitat descriptions for the 142 species are given in an annotated
list, and for some carices additional taxonomic, phytogeographical and ecological
comments are made. Distribution maps are presented, and for the 4 carices new to
the state, representative collections are noted. Doubtful and excluded taxa are
indicated and discussed.

Key Words: Carex, carices, sedges, Minnesota, floristic provinces, floristics, phyto-
geography, autecology

Since the late 1940’s, the Moore and Tryon (1946) checklist of
Minnesota plants has probably been the primary source of
information used to ascertain what carices occur in the state.
Because their Carex list is now greatly outdated and unfortunately
contains some erroneous reports, we feel it highly desirable to
provide an up-to-date account of the Minnesota Carex flora, an
updating that should prove helpful to the numerous students of this
genus both in Minnesota and in surrounding areas. We also
comment here on the phytogeographical and ecological status of the
genus in the state. Four carices previously unknown from Minne-
sota, C. annectens, C. conjuncta, C. festucacea, and C. laxiculmis,
are reported for the first time. Further, we suggest here that some
carices reported for the state by Mackenzie (1931-1935) and later
workers should be excluded.

NUMBERS OF SPECIES

Prior to Mackenzie’s monograph of North American Carex, well
over 100 species of the genus had been reported from Minnesota
(Lapham, 1875; Upham, 1884, 1887; Arthur er a/., 1887; Bailey,

ed

152 Rhodora [Vol. 86

1892; MacMillan, 1892; Cheney, 1893; Sheldon, 1894, 1895, 1896;
Holzinger, 1896; Wheeler, 1900, 1901; Rosendahl, 1903). Mackenzie
recognized 116 species of Carex native to the state and deleted
several previously reported taxa. Carices reported for Minnesota
since Mackenzie’s monumental work are given in Table 1.
Presently, 142 species of Carex are known to occur in Minnesota,
and clearly they form the largest genus of vascular plants in the
state. Except for C. /urida, one or more specimens of each of the
species reported here are in the University of Minnesota Herbarium
(MIN). A specimen of C. /urida (Sheldon, s.n., Milaca, Mille Lacs
Co., July, 1892) is at the New York Botanical Garden (NY).

FLORISTICS

Gleason and Cronquist (1964) mapped three major floristic
provinces (Grassland, Northern Conifer, Eastern Deciduous) as
coming together within Minnesota. In Wisconsin, the Eastern
Deciduous Province has been divided into the Northern Hardwoods
Province to the north and the Prairie-Forest Province to the
southwest (Curtis, 1959), and the boundary between the two, which
Curtis (1959) refers to as the tension zone, meets the Minnesota
border in northern Washington, Chisago, and southern Pine
counties (Minnesota counties are shown in Figure 1). The geo-
graphical extension of this boundary zone across Minnesota is
presently under study (Wheeler, G. A., E. J. Cushing, E. Gorham,
G. B. Ownbey and T. Morley, in preparation). The approximate
areas covered by the four floristic provinces in Minnesota are shown
in Figure 2: Grassland in the west; Northern Conifer in the far
north-central and northeastern portions; Northern Hardwoods in
the north-central and east-central parts; and Prairie-Forest in
central and southeastern Minnesota.

The diversity of floristic types in Minnesota is reflected in the
large number of Carex species occurring in the state, especially
when compared to the 68 species reported for North Dakota
(McGregor er a/., 1977; Wheeler, 1983a), 74 for South Dakota (Van
Bruggen, 1976), 106 for lowa (Gilly, 1946; Hartley, 1966; McGregor
et al., 1977), and 124 for Manitoba (Scoggan, 1957). Some states
and provinces in proximity to Minnesota, however, particularly
those to the east, such as Michigan (Voss, 1972b) and Ontario
(Scoggan, 1978), have appreciably larger Carex floras than ours.

1984] Wheeler & Ownbey — Minnesota Carices 153

Table 1. Carex Species Reported for Minnesota Since Mackenzie's (1931—-1935)
Monograph of North American Carices!

. deflexa (Lakela, 1941)

. debilis (Lemon, 1943)

. crawel (Lakela, 1944)

angustior (Moore and Tryon, 1946)

. capillaris (Moore and Tryon, 1946)
cephalantha (Moore and Tryon, 1946)
eleocharis (Moore and Tryon, 1946)

formosa (Moore and Tryon, 1946)
merritt-fernaldii (Moore and Tryon, 1946)
molesta (Moore and Tryon, 1946)
oligosperma (Moore and Tryon, 1946)
xerantica (Fernald, 1950)

filifolia (Moore, 1950)

. obtusata (Moore, 1950)

katahdinensis (Lakela, 1952)
michauxiana (Butters and Abbe, 1953)

praticola (Butters and Abbe, 1953)
pallescens (Lakela, 1954)

woodii (Hartley, 1966)

. hallii (McGregor et al., 1977)

scirpiformis (McGregor er al., 1977)

. grayi (Wheeler, 1979)

typhina (Wheeler, 1979)

bromoides (Wheeler, 1981b)

gynandra (Wheeler, 1981b)

. hitchcockiana (Wheeler, 1981b)

. oligocarpa (Wheeler, 1981b)

garberi (Wheeler, 1983b)

annectens (this report)

. conjuncta (this report)

. festucacea (this report)

. laxiculmis (this report)

Cy CRY 1 Ty Oi acy Oy Pay Oey Srey Oy Oe Oy OI a

je

nly reports substantiated by annotated herbarium specimens are given here.

Because the four floristic provinces mentioned above converge in
Minnesota, certain groups of carices tend to be associated with
specific geographic areas of the state. Indeed, based on the
distribution maps presented near the end of this report, the state can
be divided into 9 areas of Carex affiliation (Figure 3). Each area is
delimited by the presence of certain species restricted to it, or by the
fact that widespread-occurring species are absent from it, or by both
of these criteria. Some of these areas correspond closely to

154 Rhodora [Vol. 86

KITTSON ROSEAU

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a

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wd A
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Figure |. Counties of Minnesota...

landscape regions already defined for the state (Yaeger and
Borchert, 1971; Wright, 1972).

Areas 1, 2, and 3 (Figure 3) harbor Carex taxa primarily
associated with the Northern Conifer (e.g. C. canescens, C. limosa,
and C. pauciflora) and Northern Hardwoods (e.g. C. arctata, C.
leptonervia, and C. ormostachya) floristic provinces; areas 5, 6 and
7 harbor taxa primarily of the Prairie-Forest Province (e.g. C.

1984] Wheeler & Ownbey — Minnesota Carices

NORTHERN CONIFER

NORTHERN
HARDWOODS

GRASSLAND .

PRAIRIE-
FOREST

Figure 2.

155

Floristic provinces in Minnesota, after Gleason and Cronquist (1964)

and Cushing (1965). Dotted line shows approximate position of the floristic
boundary that separates the Northern Hardwoods and Prairie-Forest floristic

provinces in Minnesota (Cushing, 1965).

albursina, C. cephaloidea, and C. sparganioides); and areas 8 and 9
harbor taxa primarily of the Grassland Province (e.g. C. eleocharis,
C. hallii, and C. praegracilis). Area 4 is characterized by an
intermingling of carices from these four floristic provinces. The
eastern extremity of area 4 corresponds closely to where the tension
zone of Wisconsin (Curtis, 1959) meets the Minnesota border.

156 Rhodora [Vol. 86

Figure 3. Areas of Carex affiliation in Minnesota. The number of carices known
to occur within each area is circled.

Although the Northern Conifer and Northern Hardwoods
floristic provinces are shown in Figure 2 as occupying geo-
graphically separate areas in the northern portion of Minnesota, it is
clear that the vegetation of the region, or even a small part of it,
cannot be characterized by a single species or group of only
coniferous or deciduous species (Cushing, 1965). Instead, one finds
here deciduous, pine, and boreal forest elements intermingling to
create a mosaic of forest types. In consequence, many Carex species
of northern affinity have state distributions that are relatively

1984] Wheeler & Ownbey — Minnesota Carices 157

similar, even though particular sets of species grow in clearly
dissimilar habitats (Wheeler, 1983b). For instance, the distributions
of C. arctata, C. deflexa, and C. aenea are somewhat similar, but the
species frequent different habitats, namely deciduous, boreal, and
pine forests, respectively. Owing to this intermingling of different
forest types, much of northern Minnesota has been mapped
(Kiichler, 1964; Cushing, 1965) as mixed conifer-hardwood forest.

Area |, which approximates the Ice Scoured North Shore region
of Yaeger and Borchert (1971), is a land of lakes surrounded by high
bluffs, deep gorges cut by swiftly moving streams, and rocky cliffs
and shores that border on Lake Superior. This area is notorious for
harboring plants of discontinuous range, particularly arctic-alpine
species (Butters and Abbe, 1953; Given and Soper, 1981). Of the
Carex species restricted (or nearly so) to area I, some, such as C.
supina and C. xerantica, are essentially confined to cliff-talus and
bluff-tops bordering lakes near the Minnesota-Ontario border. C.
praticola is also best known from bluff-tops in the Border Lakes
region, but it also occurs, albeit rarely, on rocky sites farther
southward. Three other carices with very restricted distributions in
area | are C. katahdinensis, C. michauxiana, and C. pallescens, the
first is known only from rocky shores of Iron Lake (St. Louis
County), the second from swampy verges near Schroeder (Cook
County), and the last from moist sites on the Lake Superior terrace
near Duluth (St. Louis County). Of the remaining carices restricted
to area 1, all of which occur more frequently than those just
discussed, C. lenticularis and C. media are best known from the
shores of Lake Superior, and C. flava and C. gynandra from springy
sites and the marshy borders of streams and “bogs” just inland from
the lake.

Issued as a caveat in a previous publication (Wheeler er a/., 1983),
throughout this paper we use bog (the unadorned word) in the sense
of true ombrotrophic conditions; we use “bog” (in quotes) in the
sense of Sphagnum-dominated areas that typically have fen
characteristics (e.g. peatland-surrounded kettle-hole lakes). Indeed,
in Minnesota most kettle-hole “bogs” are actually poor fens and
tamarack “bogs” (better termed swamps) often approach rich-fen
conditions (Wheeler er a/., 1983). Only in the upland peatland areas
in the far northern portion of the state are true ombrotrophic bogs
encountered, such as in the large mire complex situated just north of
Upper Red Lake.

158 Rhodora [Vol. 86

Area 2, which is somewhat larger than the Big Bog region of
Yaeger and Borchert (1971), is primarily an area of gentle slope and
poor drainage that contains many upland peatlands. One of these,
the Red Lake Peatland (Heinselman, 1963, 1970; Griffin, 1975,
1977; Wheeler and Glaser, 1979, 1982; Glaser et a/., 1981; Wheeler er
al., 1983), is the largest continuous mire in the contiguous United
States. Many circumpolar species of Carex (Hultén, 1962), such as
C. canescens, C. chordorrhiza, C. diandra, C. disperma, C. lasio-
carpa, C. limosa, C. pauciflora, C. paupercula, C. tenuiflora, and C.
vaginata, are widely distributed in northern Minnesota and several
of them are most common and abundant in area 2. The latter is
certainly true for the North American endemics C. oligosperma and
C. trisperma, both of which commonly occur, and are sometimes
dominants, in upland peatland.

Carices with restricted distributions in north-central Minnesota
include C. capillaris, C. exilis, C. garberi, and C. livida. Of these, C.
exilis and C. livida are primarily confined to scattered localities in
areas | and 2, where they both frequent poor- and rich-fen sites in
patterned peatlands (Glaser, 1983; Wheeler, 1983b). Of more
restricted distribution, C. capillaris occurs only in the western
portion of area 2 and small parts of 3 and 4, and C. garberi is known
only from a single station in St. Louis County. Finally, some
northern carices, such as C. arcta and C. crinita, are widely
distributed in areas 1, 3, and 4 but are uncommon or unknown in
area 2.

Areas 3, 4, and 5 harbor the greatest number of Carex species in
the state (Figure 3), with area 4, as mentioned earlier, representing a
transition zone. Nevertheless, several carices of northern affinity are
known from isolated stations south of area 4, and some species of
southern affinity north of it. Likewise, some western species, such as
C. torreyi, occur at isolated stations east of this transition area.
Carices known to be restricted (or nearly so) to one or more of these
areas include C. bromoides, C. debilis, C. festucacea, C. formosa, C.
lurida, and C. rossii. The four last-named species are known only
from a few stations in Minnesota, and they seldom occur in
abundance. C. bromoides and C. debilis, on the other hand, while
being more or less restricted to the extreme east-central portion of
the state, are both known from several localities and sometimes
occur in great abundance. Indeed, C. bromoides often provides the
major ground cover in low, swampy woodlands adjacent to rivers.

1984] Wheeler & Ownbey — Minnesota Carices 159

Wheeler (1981b) reported C. bromoides as extending no farther
west in Minnesota than wooded lowlands bordering the Kettle
River, but the authors have recently discovered several stations for
this species along the Snake River (southern Aitkin and northern
Kanabec counties), some 20 miles farther westward. Another Carex
species growing in central Minnesota, C. sterilis, until recently had
been known from very few localities in the state, but in the last few
years several spring-fed, calcareous fens have been discovered in
areas 4, 5, and 8 that harbor this apparent calciphile.

The Minnesota River, which flows primarily through the central
portions of areas 5 and 9, presently occupies a small part of the huge
valley cut out by the ancient River Warren. The river is bordered by
bluffs that are not as imposing as those in extreme southeastern
Minnesota, but these bluffs do harbor many plants that are
uncommon or unknown elsewhere in these areas. Carex hitch-
cockiana and C. oligocarpa are best known in the state from these
bluffs and, indeed, the latter provide the habitats for the north-
westernmost stations for each of these species in North America
(Wheeler, 1981b). Another sedge of frequent occurrence on these
bluffs, particularly westward, is C. saximontana. Although this
species is reported by various workers (Mackenzie, 1931-1935;
Fernald, 1950; Gleason and Cronquist, 1963) as restricted in the
state to “western Minnesota”, it does, in fact, extend as far eastward
with us as Ramsey County, with the majority of the eastern stations
situated on the bluffs of the Minnesota River.

Extreme southeastern Minnesota, the Stream Dissected region of
Yaeger and Borchert (1971), is botanically as well as geologically
one of the most interesting parts of the state. Area 7 is part of the
celebrated Driftless Area (Hartley, 1966) and, like the remaining
parts (in southwestern Wisconsin, northeastern Iowa, and north-
western Illinois), has very irregular topography resulting from the
action of various natural processes, particularly stream erosion.
Like the remainder of the Driftless Area, area 7 is notorious for
harboring plants of limited or discontinuous range, such as Montia
chamissoi (Portulacaceae), Sullivantia renifolia (Saxifragaceae),
and Talinum rugospermum (Portulacaceae).

Several species of Carex occurring in southeastern Minnesota are
unknown from elsewhere in the state, and include C. annectens
(area 6), C. conjuncta (area 6), C. crus-corvi (area 6), C.
laevivaginata (area 7), and C. laxiculmis (area 7). Furthermore, C.

160 Rhodora [Vol. 86

davisii, C. gravi, C. muskingumensis, and C. typhina are primarily
confined to the southeast, where they are characteristic species in
floodplain forests, particularly those bordering the Mississippi
River. Indeed, with the exception of C. typhina, which occurs at a
few localities along the St. Croix River in Washington and southern
Chisago counties, these species rarely occur north of Goodhue
County (Wheeler, 1979). Finally, many Carex species occurring in
area 5 and in areas farther north do not extend as far southeastward
as area 6, and fewer yet extend into area 7 (Figure 3).

It is obvious from Figure 3 that fewer species of Carex occur in
western Minnesota (areas 8 and 9) than in the eastern portion of the
state. Although the genus is clearly less well represented in prairie
than in forest, fen, or “bog”, individual species, such as C. brevior
and C. eleocharis in dry sites and C. atherodes and C. praegracilis in
wet sites, are sometimes widespread and abundant in undisturbed
prairie. Indeed, C. eleocharis sometimes forms, along with various
grasses, the major ground cover on hillsides and slopes, and C.
atherodes is often the dominant species in prairie swales. In
Minnesota, as well as elsewhere (Hudson, 1977), C. atherodes
appears to tolerate the drying up of prairie swales, unlike C.
lacustris, C. rostrata, and C. aquatilis, which seem to need more or
less persistently wet conditions. Therefore, only at very favored
locations within the prairie region of Minnesota are the last three
species able to attain dominance. Some prairie carices, including C.
hallii, C. scirpiformis, C. filifolia, and C. obtusata, have very
restricted distributions in the western half of the state, and they
seldom occur in any great abundance. The first two species seem to
be restricted to prairie swales, particularly to the moist outer
margins, and the last two taxa grow primarily in dry, sandy sites,
such as in sand barrens. A paper discussing all 9 areas in more detail
is being prepared.

Although a particular Carex species may be common in one
region of Minnesota, it may also occur, as indicated above, at
isolated stations in other parts of the state. For instance, several
carices that have their major occurrence in Sphagnum-dominated
areas in northern Minnesota, such as C. canescens, C. leptalea, and
C. paupercula, also occur in tamarack “bogs” scattered throughout
the central and, less commonly, the southern portions of the state.
Furthermore, some calcareous fens in southern and western
Minnesota harbor some carices of northern affinity, such as C,
aquatilis, C. lasiocarpa, and C. limosa.

1984] Wheeler & Ownbey — Minnesota Carices 161

While many species of Carex are geographically restricted in the
state, others are more widely distributed. A number of the latter,
such as C. hystericina, C. lanuginosa, C. stipata, C. stricta, and C.
vulpinoidea, are aquatic plants (Fassett, 1957) that grow along
shores and in adjacent wet meadows of the numerous lakes scattered
throughout Minnesota. Some woodland species, such as c. peckii,
C. pedunculata, C. pensylvanica, and C. rosea, are also widely
distributed. And some carices that grow along the banks of rivers
often have wide distributions in the state. However, some riverine
carices, such as c. alopecoidea, C. cristatella, and C. emoryi, are
essentially absent from extreme northeastern Minnesota, an area
where prolonged level riparian tracts with well-developed meander
scrolls and floodplains are mostly lacking because the rivers flow
swiftly from the upland down a steep gradient to Lake Superior
(Wheeler, 1983b).

As in the above case, the distributions of other sedges in the state
are restricted by the lack of suitable habitats. For example, several
carices, including C. atherodes, C. interior, C. praegracilis, C.
prairea, C. sartwellii, and C. tetanica, all of which grow primarily in
prairie swales and marshlands, have state distributions that
approximate that of the calcareous drift deposited by the Mankato
Substage of the Wisconsin Stage of glaciation. It appears that a lack
of low, wet, strongly calcareous habitats in southeastern Minnesota
results in the exclusion (or near exclusion) of these species from this
part of the state, an area not invaded by the Des Moines Lobe of the
Mankato Substage. Several other vascular species that frequent wet
calcareous sites, such as Juncus balticus (Juncaceae), Rumex
maritimus (Polygonaceae), /ris versicolor (Iridaceae), and Habe-
naria hyperborea (Orchidaceae), are also essentially absent from the
southeast. In Figure 3, this demarcation is illustrated by the line that
separates area 5 (covered with Des Moines calcareous drift) and
area 6 (not covered with Des Moines calcareous drift).

Because four floristic provinces converge in Minnesota, many
species of Carex reach the limits of their ranges in the state
(Wheeler, 198la). Some of these carices, such as C. brunnescens
(Northern Conifer Province), C. /leptonervia (Northern Hardwoods
Province), C. hirtifolia (Prairie-Forest Province), and C. filifolia
(Grassland Province), display more or less continuous ranges as
they approach and enter Minnesota. Other carices, however, are
represented by only a few local populations in the Midwest. As an
example of the latter, C. exilis is primarily a plant of the Atlantic

162 Rhodora [Vol. 86

coastal plain that has only limited local populations west of New
York (Reznicek and Ball, 1980). In Minnesota, it is known only
from a few patterned peatlands in the northeastern and north-
central portions of the state (Glaser, 1983; Wheeler, 1983b), with the
Red Lake Peatland harboring the westernmost stations in North
America (Wheeler and Glaser, 1979; Wheeler er al, 1983).

Lastly, a few Carex taxa, such as C. supina, represented in the
state by disjunct populations, may be relicts. However, their
migration to favorable habitats after glaciation cannot be ruled out.
The relict hypothesis is generally favored to explain the wide
discontinuities in range of arctic-alpine taxa in the Lake Superior
region (Butters and Abbe, 1953; Soper and Maycock, 1963; Given
and Soper, 1981) and is supported by some palynological data
(Cushing, 1965). The nunatak hypothesis of Fernald (1935) is no
longer considered tenable (Cushing, 1965). Probable Carex relicts
occurring in Minnesota are indicated and discussed in the listing of
species given near the end of this report.

Besides the distributions discernible for Minnesota Carex species
resulting from the convergence of the four floristic provinces within
the state, our carices display still other more subtle and local
distributional patterns. Many carices are greatly affected by the
nature of the water in which they grow (Wheeler er a/., 1983), which
in turn is influenced by the physical nature of the water source. That
some peatland Carex species are profoundly acidophilous or
basiphilous is well known (e.g. Sjérs, 1961, 1963), and this fact has
been demonstrated for some carices occurring in kettle-hole “bogs”
and upland peatland areas in northern Minnesota (Glaser er al.,
1981; Wheeler er a/., 1983).

Carex species primarily confined to Sphagnum-dominated areas
in Minnesota have state distributions that are somewhat similar, but
some differences are apparent. For example, C. oligosperma and C.
pauciflora are occasional to common in peatlands and “bogs”
throughout the northeastern, north-central, and east-central por-
tions of the state, but they are essentially absent from such habitats
in the central, west-central, and northwestern parts of Minnesota. In
contrast, many other “bog-loving” carices, such as C. lasiocarpa, C.
leptalea, and C. limosa, are present in these habitats throughout the
entire northern half of the state and, as indicated earlier, some of
them extend even farther southward and westward, growing in fens
and tamarack swamps. Significantly, other vascular “bog-loving”

1984] Wheeler & Ownbey — Minnesota Carices 163

species display similar distributional patterns: Eriophorum spissum
(Cyperaceae), E. virginicum, and Kalmia polifolia (Ericaceae) have
distributions in the state similar to those of C. oligosperma and C.
pauciflora, whereas many other species, such as Betula pumila
(Betulaceae), Equisetum fluviatile (Equisetaceae), and Menyanthes
trifoliata (Gentianaceae), have state distributions similar to C.
lasiocarpa, C. leptalea, and C. limosa. Because some oligotrophic
species, particularly obligate oligotrophs, are probably unable to
achieve ecesis in highly nutrient-rich conditions, the moderately to
highly minerotrophic nature of the fens and “bogs” in western and
southern Minnesota may well impede the growth of such species.
Indeed, C. oligosperma and C. pauciflora, as well as the other three
vascular plants mentioned above with restricted distributions in
Minnesota, are essentially confined to oligotrophic sites in the Red
Lake Peatland (Wheeler er a/., 1983), and they are more or less
restricted to kettle-hole “bogs” in the state that are “acidic” (pH
usually less than 6.5).

In the Red Lake Peatland, which is situated just north of Upper
Red Lake in north-central Minnesota, Carex species account for
approximately 15% of the vascular flora (Wheeler er a/., 1983). In
this mire, ombrotrophic bogs, areas of weak minerotrophy (poor
fens), and patterned rich fens can be distinguished on the basis of
acidity and other chemical properties of the peat and surface-waters
and by species richness. Only four Carex species (C. oligosperma, C.
pauciflora, C. paupercula, and C. trisperma) are known from
ombrotrophic sites in the mire, whereas twenty-nine carices frequent
minerotrophic sites. Some species, such as C. chordorrhiza and C.
livida, are excellent indicators of minerotrophy, inasmuch as they
separate areas of weak minerotrophy from true ombrotrophic sites.
Still other species, such as C. /eptalea and C. pseudocyperus, are
obligate rich-fen indicators. However, in the Red Lake Peatland, as
in other mires (e.g. Malmer, 1962; Sjérs, 1963), there are no vascular
taxa, including Carex, that are indicators of ombrotrophy.

In upland peatlands, variations in environmental parameters such
as in water chemistry, hydrology and shading greatly influence
whether a particular species will be present or absent from a
particular site. Figures 4 and 5 illustrate for four Carex species
common to the Red Lake Peatland the relationship between the
principal components axes, which are based on variations in the
frequency with which species occur, and environmental factors

164 Rhodora [Vol. 86

(Brush, 1982). These figures show that whereas C. oligosperma
displays a narrow range of tolerance for all environmental
parameters considered, each of the other three species displays a
relatively wide range of tolerance for one or more of these
parameters. Although Figures 4 and 5 were constructed from data
gathered from one mire complex, the results (for these four species)
seem to hold in other upland peatlands investigated in the state.

Although C. oligosperma dominates large areas in the Red Lake
Peatland, it is more or less restricted to open bogs and bog drains,
and to meadow-like, open poor-fen areas where bog drains coalesce.
In contrast, C. /asiocarpa grows over a wide range of pH and
calcium values, and it is often the dominant species in wet, open
rich-fen sites, such as flarks of patterned fens. With regard to the
other two peatland species referred to in Figures 4 and 5, C.
trisperma is restricted to wooded bogs, wooded poor-fen islands,
and hummock-tops in the wooded rich-fen islands. C. /eptalea, on
the other hand, is an obligate rich-fen indicator that grows most
abundantly on wooded rich-fen islands but also occurs in flarks and
in partially-shaded depressions between string hummocks in pat-
terned fens. A more detailed discussion of the distribution of Carex
species along various environmental gradients in the Red Lake
Peatland is given by Wheeler er al. (1983).

Another example of species of Carex being affected by differences
in water chemistry can be demonstrated from kettle-hole “bogs”. In
east-central, north-central, and northeastern Minnesota, Carex
oligosperma is almost invariably the most important species in the
formation of pioneer mats surrounding “acid” kettle-hole lakes
(actually most such lakes are weakly acid, with a pH usually less
than 6.5), as well as being the dominant vascular plant in adjacent
moss-heath zones. It is replaced by C. /asiocarpa, however, in kettle-
hole “bogs” surrounding “alkaline” lakes (pH mostly greater than
6.5). Indeed, throughout these areas the two species seldom grow
together on the same floating mats or in the same “bogs,” but C.
lasiocarpa sometimes frequents the nutrient-rich laggs of C.
oligosperma-dominated peatlands. Some species of Carex are
common to both types of Sphagnum-dominated peatlands, whereas
others are restricted to one type or the other. Species commonly
found on mats and “bogs” dominated by C. oligosperma are C.
pauciflora, C. paupercula, C. limosa, C. canescens, and C.

1984] Wheeler & Ownbey — Minnesota Carices 165

hieee . > ° eee
*

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First component ——»

Figure 4. Local environmental interactions on distribution of 4 Carex species.
The first component shows a distribution ranging from “acid” species at the left to
“circumneutral” species at the right; pH values range from 3.8 to 7. The second
component reflects tolerance to shading, with sciaphytes appearing on the top of the
axis and heliophytes on the bottom. The sizes of the circles are proportional to the
number of occurrences of a particular species in 45 releves (Wheeler er a/., 1983); the
largest circle represents 4 occurrences and the smallest circle a single occurrence. The
circles with diagonal lines represent C. oligosperma; dotted circles C. trisperma; open
circles C. lasiocarpa; and closed circles C. leptalea.

lL} o .& CON ee$o
2 € e ®
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6
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First component ——»

Figure 5. Local environmental interactions on 4 Carex species. The first
component shows a distribution ranging from calciphobic species (left on the axis) to
calciphilic species (right on the axis); calcium values range from 0.5 mg/1 to 19.6
mg/l. The second component reflects tolerance to dry sites, with “dry” species
appearing on the top of the axis and “wet” species on the bottom. See Figure 4 for
explanation of symbols and circle sizes.

166 Rhodora [Vol. 86

b]

chordorrhiza; species occurring most often on mats and in “bogs’
dominated by C. /asiocarpa are C. aquatilis, C. diandra, C. limosa,
C. canescens, C. chordorrhiza and, more rarely, C. paupercula.
Likewise, in northern Michigan (Vitt and Slack, 1975), the Carex
species of peatlands surrounding acid kettle-hole lakes are mostly
different from those in peatlands surrounding alkaline lakes.
Several instances are known from Minnesota where a particular
terrestrial species of Carex is restricted to a specific soil type and is
absent from others. For example, C. backii grows in sandy, acid
soils whereas C. albursina, C. hitchcockiana, C. laxiculmis, and C.
oligocarpa grow in calcareous sites. The four last-named species, all
of which are confined to the southern half of the state, are well
known from sugar maple-basswood forests that are situated on
steep, north- and east-facing slopes characterized by rich loamy
soils. These calciphiles or near-calciphiles (Mackenzie, 1931-1935;
Hermann, 1940; Fernald, 1950; Wheeler, 1981b) are absent from
neighboring dry hardwood forests and the more locally occurring
woodlands dominated by white pine, both of which are developed
on better-drained, more acid, sandy soils. In contrast, the acid-
ophilous species C. backii, although best known from pine forests in
northern Minnesota, is of occasional occurrence in the afore-
mentioned white pine-dominated woodlands but is rare or absent
from the neighboring, much more common, sugar maple-basswood
forests. Other examples of distributional patterns exhibited by
Carex species in Minnesota are given by Wheeler (198 la).
Quite often the same combination of carices grow in a particular
habitat even though the localities are miles apart, and at least in
some habitats, if not in most, certain species are associated with
specific microhabitats. For example, in a recent study of 17
tamarack swamps in seven counties in west-central and north-
western Minnesota, of the 15 carices found growing in these
swamps, 10 species (C. aurea, C. paupercula, C. retrorsa, C. stipata,
C. deweyana, C. disperma, C. hystericina, C. interior, C. leptalea,
and C. pedunculata) were present in at least 75% of the swamps
investigated; and the 6 last-named species were present in over 85%.
Of the remaining species, 4 of them, C. brunnescens, C. gynocrates,
C. rosea, and C. tenuiflora, occurred in more than one swamp, and
C. vaginata was limited to a single swamp in Mahnomen County.
Furthermore, one or more Carex species, such as C. atherodes, C.
lacustris, C. stricta or, less commonly, C. /asiocarpa, C. prairea, and

1984] Wheeler & Ownbey — Minnesota Carices 167

C. rostrata, invariably grow along the margins of these swamps and
sometimes in the swamp interiors.

Within these tamarack swamps, C. disperma, C. leptalea, and C.
deweyana are primarily associated with the sides and tops of
hummocks, whereas C. aurea, C. interior, and C. paupercula
generally grow in wet depressions between hummocks. C. pe-
dunculata is mostly confined to the bases of trees and windthrow
mounds, and C. hAystericina, C. retrorsa, and C. stipata are most
abundant and grow best in low, level areas. C. Jacustris often
dominates the outer edges of the swamps, sometimes growing in
nearly pure stands, and is often a common constituent in the
interiors, particularly in and around the margins of small pools and
other wet depressions.

Some Carex species have clearly expanded their ranges within
rather recent times through man’s intervention. In Minnesota,
probably the most obvious example is C. praegracilis which,
although native in western and south-central Minnesota, is now
adventive in the north-central, northeastern, and extreme east-
central portions of the state, particularly along highway verges
(Wheeler, 1983b). This species has, in fact, become adventive well
eastward of its native range (Mackenzie, 1931-1935; Hermann,
1974; Reznicek ef al., 1976), presumably through dispersal of
achenes and rhizomes along major highways and because of its
apparent high resistance to extreme conditions of road verges, such
as summer drought and high sodium levels owing to “de-icing” salt
(Reznicek et al., 1976).

Another example of area expansion of Carex species within the
state can be demonstrated from the Red Lake Peatland. Although
twenty-nine carices are known from this mire, nine of them are
clearly restricted to disturbed sites. It is assumed that these species
have only recently entered the mire, most likely following the
construction of the major highway that transects the area and the
several drainage ditches that penetrate into the peatland interior.
Although these species are most likely recent arrivals, they presently
form an integral part of the peatland flora and no doubt will
continue to spread to future disturbed areas (Wheeler ef a/., 1983).

Hybridization is known to occur in Carex (Wahl, 1940; Drury,
1956), and several hybrids and suggested hybrids have been reported
from the eastern United States and Canada (Fernald, 1950;
Scoggan, 1978). Although little is known about the hybridization of

168 Rhodora [Vol. 86

Carex in Minnesota, observations in the field and some herbarium
specimens suggest that crossing does take place between particular
species in the state. Specimens have been seen from southeastern
Minnesota (Houston and Winona counties) that appear to be
hybrids of C. tenera and C. normalis. Hybridization between these
two members of the Ova/es group is not indicated by Fernald (1950)
or Scoggan (1978), but it has been reported by Zimmerman (1976)
as possibly occurring in Wisconsin.

Specimens which display characteristics intermediate between
Carex bebbii and C. cristatella (both members of the Ovales group)
are also known from the state; these specimens come from northern
Minnesota (Lake of the Woods, Itasca, and St. Louis counties).
Hybridization between C. bebbii and C. cristatella has been
reported as possibly occurring in Michigan (Voss, 1972b), Wis-
consin (Zimmerman, 1976), and Saskatchewan (Hudson, 1977), but
it was not indicated by Fernald (1950) or Scoggan (1978).

A plant which closely resembles Carex castanea, but which is
more glabrous, is known from widely scattered localities in the
United States and Canada (Arthur er al, 1887; Bill, 1930;
Mackenzie, 1931-1935; Fernald, 1950; Scoggan, 1978), and it is
considered by some workers (Fernald, 1950; Scoggan, 1978) to be a
hybrid between C. castanea and C. arctata (both members of the
Sylvaticae group). In Minnesota, this suspected hybrid (C. Xknie-
skernii Dew.) is known only from St. Louis County. Specimens of
suspected hybrids have not been mapped for this study.

The species listed below are arranged alphabetically, and for the
most part the treatment here follows that of Mackenzie (1931-
1935), except where a departure has been forced (or at least
suggested) by certain available evidence. Section or group (see Voss,
1966, 1972a, 1972b) relationships are indicated, and synonyms
correlating this treatment with those in current manuals are noted.
Although group names are without formal taxonomic standing, the
group that includes the type species of the genus, C. hirta, is here
called Carex in accord with the spirit of the Code of Nomenclature.

Data gathered from field observations and specimens contained
in various herbaria (DUL, GH, IA, ISC, MANK, MICH, MIN,
NDA, NY, SCL, US, UWL, WIS; abbreviations after Holmgren et
al., 1981) were used to develop habitat descriptions. The areas
illustrated in Figure 3 are referred to when they are useful in

1984] Wheeler & Ownbey — Minnesota Carices 169

characterizing distribution patterns and when reporting the fre-
quency of occurrence of a species within a particular area.

On the distribution maps, which were prepared from the
herbarium specimens examined, each solid circle represents a
collection of known locality; an open circle represents a collection
without specific location within a county.

Following the annotated list are remarks on some doubtful and
excluded taxa.

ANNOTATED LIST OF MINNESOTA CARICES

C. abdita Bickn. Montanae

Cliffs and bluffs, rocky slopes and shores, rock outcrops and,
more rarely, dry prairies; infrequent to occasional in the eastern half
of the state, uncommon in the western half. Some authors (e.g.
Gleason and Cronquist, 1963) combine C. abdita, C. tonsa, and C.
umbellata under the epithet umbellata, but in Minnesota the first-
named entity is well-marked and easily distinguished from the other
two: beak of perigynium is short and the tips of the pistillate scales
are acute. In contrast, the other two entities have long beaks and
acuminate scales. Furthermore, whereas C. tonsa and C. umbellata
generally grow in sand barrens and in other sandy sites, some of
which are highly disturbed, C. abdita rarely grows in pure sand, and
particularly not in disturbed places; instead, it seems to be
invariably associated with rocky slopes, rock outcrops, and cliffs
and bluffs. For those (e.g. Mackenzie, 1931-1935, 1940; Voss,
1972b) who consider C. rugosperma Mack. to be the true C.
umbellata Schkuhr ex Willd., then this entity becomes C. umbellata
Willd. See comments under C. tonsa and C. umbellata.

C. adusta Boott Ovales
Rock outcrops, cliffs and bluffs, margins of pine forests, sandy
and rocky slopes, and sandy disturbed sites (sand and gravel pits,
roadside shoulders and embankments, clear-cut and burned-over
areas); frequent in area 1, infrequent to occasional in 2, 3.

C. aenea Fern. Ovales

Sandy and rocky slopes, margins of pine forests and pioneer
hardwood stands, cliffs and bluffs, and sandy disturbed sites (sand
and gravel pits, roadside shoulders and embankments, clear-cut and
burned-over areas); frequent to common in area 1, occasional
ied, 4:

170 Rhodora [Vol. 86

C. albursina Sheldon Laxiflorae

Steep hardwood-covered slopes, moist wooded ravines and, less
commonly, flat deciduous woodlands; occasional to frequent in area
7, infrequent to occasional in 5, 6. The type of this species was
collected near White Bear Lake (Washington County) in east-
central Minnesota by E. P. Sheldon in 1892. Some workers (e.g.
Gleason and Cronquist, 1963) treat it as C. laxiflora Lam. var.
latifolia Boott.

C. alopecoidea Tuckerm. Vulpinae
Floodplain forests, abandoned river channels and ox-bows,
woodland swales and pond margins, wooded river banks, ditches,
and moist to wet meadows adjacent to rivers and creeks: occasional
throughout the state, except in area 1, where it is unknown.

C. amphibola Steud. Griseae

Floodplain forests, mesic hardwood stands (especially in swales
and along the margins of creeks and ponds), wooded river banks,
and disturbed sites (trails and paths); frequent to common in areas
5, 6, 7, occasional in the eastern half of 4, infrequent in 9,
uncommon in 3. Our material may be referred to var. turgida Fern.
[C. grisea Wahlenb.; C. turgida (Fern.) J. W. Moore].

C. angustior Mack. Stellulatae

Wet sandy shores of lakes, margins of alder swamps, wet edges of
trails, and “bog” borders; occasional in area 1, infrequent in 2, 3. C.
angustior and C. cephalantha are fairly distinctive in Minnesota
and, as such, they are treated here as separate species. It must be
noted, however, that Reznicek and Ball (1980), in their recent
treatment of the Ste//ulatae group, place both of these entities under
C. echinata Murr. Previously, some workers (e.g. Gleason and
Cronquist, 1963) treated C. angustior and C. cephalantha as
varieties of C. muricata L., as var. angustata Carey and var.
cephalantha Baily, respectively. In Minnesota, C. cephalantha
grows in “bogs” and in upland peatlands whereas C. angustior
grows primarily in wet depressions along the edges of swamps and
trails as well as along the wet shores of lakes. In the Red Lake
Peatland, C. cephalantha is a frequently-occurring plant on strings
of patterned fens, but C. angustior is absent from this large mire
complex. A similar situation, as regards the presence of C.
cephalantha and absence of C. angustior, seems to exist in the other

1984] Wheeler & Ownbey — Minnesota Carices 171

upland peatlands that have been recently investigated in the state. C.
cephalantha, on the other hand, seems to be invariably absent from
those habitats given for C. angustior. For the majority of our
material, the morphological differences usually given to separate
these two entities (e.g. Fernald, 1950) seem to hold quite well.

C. annectens (Bickn.) Bickn. Multiflorae

Two stations are known for this species in Minnesota. It was first
collected on sand barrens at the Weaver Dunes (Wabasha County),
where it was primarily associated with C. muhlenbergii. Only a few
clumps were observed, all growing in proximity to one another.
Somewhat later, the species was collected on a mesic prairie strip in
Mower County. While the latter material is var. xanthocarpa
(Bickn.) Wieg., the former material is probably best treated as var.
annectens. This note represents the first report of this species from
Minnesota; it appears to be very rare in the state.

Mower Co.: 3 mi. NW of LeRoy on Rte. 56, mesic prairie strip, TIOIN, RI4W,
Sec. 19, Smith 4348 (MIN). Wabasha Co.: 5 mi. SE of Kellogg on Rte. 84, sand dune
area, TIOIN, ROW, Sec. 5, Wheeler 2700 (MIN).

C. aquatilis Wahlenb. Acutae

Most Minnesota material is var. altior (Rydb.) Fern., which
frequents marshy lake shores, floating peat mats surrounding
alkaline kettle-hole lakes, “bog” borders, swales that are persistently
wet, ditches, and the margins of streams; occasional to frequent in
the northern half of the state, infrequent in the southern half. The
var. aquatilis, which was reported for Minnesota by Butters and
Abbe (1953), is known only from the extreme northeast corner of
the state, where it is uncommon. These two taxa have not been
mapped separately.

C. arcta Boott Heleonastes

Floodplain forests, abandoned river channels and ox-bows,
woodland swales and pond margins, mixed conifer-hardwood
swamps, and “bog” borders; occasional in areas 1, 3.

C. arctata Boott Sylvaticae

Mesic to dry forests (deciduous, mixed conifer-hardwood and,
less commonly, coniferous), thickets, clearings, and wooded river
banks; frequent to common in areas 1, 2, 3, infrequent in 4,
uncommon in 5.

172 Rhodora [Vol. 86

C. assiniboinensis W. Boott Sy/vaticae
Floodplain forests, abandoned river channels and ox-bows and,
less commonly, mixed conifer-hardwood forests and mesic hard-
wood stands; occasional to frequent throughout the state, except in
the northeast and southeast corners, where it is rare or unknown.
The plants often bear numerous, long-arching sterile culms or
stolons, the tips of which upon reaching the ground form new
plants; at some localities these sterile culms form the major cover of
the field layer. Various workers (Tolstead, 1946; Stevens, 1950:
Bernard, 1959; Hudson, 1977; Wheeler, 198la, 1983b) have
discussed or alluded to vegetative reproduction in this species.

C. atherodes Spreng. Paludosae

Prairie swales, wet meadows, ditches, and the marshy borders of
lakes, rivers, and ponds; frequent throughout the state, except in the
northeast and southeast corners, where it is rare or unknown. The
plants often form dense, nearly pure stands in swales and other wet
depressions, even those that dry up during the summer. Easily
recognized by the pubescence on the sheaths and underside of the
leaves, but plants growing in standing water sometimes lack this
pubescence.

C. aurea Nutt. Bicolores

Moist to wet meadows, damp sandy shores, ditches, swampy
woods, marshy borders of lakes and springy banks of streams, and
disturbed sites that are artificially watered (e.g. taconite tailings
ponds), frequent in the northern half of the state, infrequent to
occasional in the southern half. The perigynia are brightly colored
(golden-yellow or brownish-orange) and fleshy at maturity, and in
all probability attract birds. See comments under C. garberi.

C. backii Boott Phyllostachyeae

Wooded slopes, cliffs and bluffs, ravines, and sandy ridges in pine
stands; occasional in the eastern half of the state, infrequent in the
western half. See comments under C. saximontana.

C. bebbii (Bailey) Fern. Ovales

Moist meadows, swales, ditches, clearings, and the marshy
margins of lakes, rivers, and ponds; occasional to frequent
throughout the state, except in the southwest, where it is un-
common. This species rarely occurs in woodlands, and it has been

1984] Wheeler & Ownbey — Minnesota Carices 173

suggested (Hudson, 1977) that its rarity in prairies is because of an
intolerance to soluble salts.

C. bicknellii Britt. Ovales

Prairies, sandy hillsides, railroad and highway embankments
(especially prairie strips), sand barrens, rock outcrops, and, less
commonly, the borders of lakes and ponds; occasional to frequent
in the southern half of the state, infrequent in the northern half.

C. blanda Dew. Laxiflorae

Deciduous woodlands of all types and disturbed sites (e.g.
clearings, paths and trails, roadway embankments); common in
areas 5, 6, 7, occasional to frequent in 4, 8, 9, infrequent in 3. One of
the most commonly-occurring carices in deciduous woodlands in
the southern half of the state. Some workers (e.g. Gleason and
Cronquist, 1963) treat it as C. /axiflora Lam. var. blanda (Dew.)
Boott. See comments under C. /eptonervia.

C. brevior (Dew.) Mack. Ovales

Prairies, sandy hillsides, railroad and highway embankments
(especially prairie strips), sand barrens, rock outcrops, sandy
disturbed sites and, less commonly, cliffs and bluffs, river banks,
and open woodlands; frequent to common in the western and
southern portions of the state, infrequent in the north-central and
northeastern portions. One of the most commonly-occurring carices
in dry, open sites in southern and western Minnesota.

C. bromoides Willd. Deweyanae

Floodplain forests, abandoned river channels and ox-bows,
woodland swales and pond margins, mixed conifer-hardwood
swamps, and the springy banks of streams; restricted to east-central
Minnesota, where it is locally abundant. Best known from low,
swampy woodlands along the Kettle (Wheeler, 1981b) and Snake
rivers.

C. brunnescens (Pers.) Por. Heleonastes

Coniferous and mixed conifer-hardwood forests and swamps,
alder swamps, clearings, and “bog” borders; frequent to common in
areas 1, 2, 3, occasional in 4, uncommon in 5, 8. Our material may
be referred to var. sphaerostachya (Tuckerm.) Kiikenth.

174 Rhodora [Vol. 86

C. buxbaumii Wahlenb. Atratae

Moist meadows, prairie swales, fens (usually confined to flarks in
patterned fens), rocky shores of lakes, and wet ditches; occasional to
frequent in the northern half of the state, infrequent to occasional in
the southern half (except in the southwest, where it is unknown). In
the Red Lake Peatland, it is infrequent in flarks of patterned fens
and in disturbed sites (Wheeler er a/., 1983).

C. canescens L. Heleonastes

Open and wooded oligotrophic and mesotrophic “bogs”, floating
peat mats surrounding acid and alkaline kettle-hole lakes, fens
(usually associated with strings in patterned fens) and, somewhat
less commonly, conifer and mixed conifer-hardwood swamps, and
alder swamps; frequent in areas |, 2, 3, occasional in 4, uncommon
in 5. In areas 4 and 5, it is mostly restricted to tamarack swamps. In
the Red Lake Peatland, it grows in wet hollows on the poor- and
rich-fen wooded islands, along the margins of string hummocks in
patterned fens, and in disturbed sites (Wheeler er a/., 1983). Most of
our material may be referred to var. disjuncta Fern., but var.
subloliacea (Laest.) Hartman also occurs occasionally. Further-
more, some material from the extreme northeast is best referred to
typical var. canescens. But because intergrades are so frequent,
particularly between the first two taxa, these three entities have not
been mapped separately.

C. capillaris L. Capillares

Moist shaded lake shores and, less commonly, thickets, wooded
beach ridges, and “bog” and swamp borders; infrequent to occa-
sional in the western portions of areas 2, 3, uncommon in 4. Best
known from the environs of Lake Itasca (Clearwater County). Our
material may be referred to var. major Drej. [C. capillaris var.
elongata Olney; C. capillaris ssp. chlorostachys (Stev.) Love, Love,
& Raymond].

C. castanea Wahlenb. Sy/vaticae

Mesic meadows, abandoned fields, “bog” and swamp borders,
lake shores, roadside embankments, and ditches; occasional in areas
| ae an

C. cephalantha (Bailey) Bickn. Stellulatae
Open mesotrophic “bogs”, fens (usually associated with strings in
patterned fens), openings in conifer swamps, and “bog” borders;

1984] Wheeler & Ownbey — Minnesota Carices 175

occasional in areas 1, 2, 3, infrequent in 4 and the northern half of 5.
In areas 4 and 5, it is mostly restricted to tamarack swamps. In the
Red Lake Peatland, it grows on strings of patterned fens and in
disturbed sites (Wheeler er a/., 1983). As treated here, our material
includes C. Jaricina Mack. [C. muricata L. var. laricina (Mack.)
Gl.]. See comments under C. angustior.

C. cephaloidea (Dew.) Dew. Bracteosae

Deciduous woodlands (wet to dry) and thickets; occasional to
frequent in areas 5, 6, 7, infrequent in 9 and the southern portion of
4. It often grows in thickets, particularly under Zanthoxylum, where
it sometimes occurs in abundance. Some workers (e.g. Gleason and
Cronquist, 1963) treat it as C. sparganioides Muhl. var. cephaloidea
(Dew.) Carey.

C. cephalophora Willd. Bracteosae

Dry deciduous woodlands and thickets, sandy hillsides, exposed
ravines, and the margins of mesic hardwood stands; occasional in
areas 5, 6, 7.

C. chordorrhiza L. f. Chordorrhizeae

Open oligotrophic and mesotrophic “bogs”, floating peat mats
surrounding acid and alkaline kettle-hole lakes, openings and edges
of conifer swamps, and fens; occasional to frequent in areas 1, 2, 3,
infrequent in 4 and the northern portions of 5, 6. In areas 4, 5, and 6,
it is mostly confined to openings in tamarack swamps. In the Red
Lake Peatland, it grows in poor- and rich-fen sites and serves as an
indicator of minerotrophy in ecotones between ombrotrophic and
minerotrophic areas (Glaser er a/., 1981; Wheeler and Glaser, 1982:
Wheeler er al., 1983). The plant seems to require at least moderate
light for best growth, and it has a most unusual and well-marked
habit of growth, where old prostrate culms of the previous year give
rise to new plants from the nodes.

C. communis Bailey Montanae

Rocky ledges in woodlands, cliffs and bluffs and, less commonly,
on rocky shores; infrequent to occasional in the eastern half of the
state. Best known from rocky ledges along the shores of Lake
Superior. According to Handel (1978a), this species is a myr-
mecochore, with the perigynia having elaiosomes that attract ants to
carry diaspores to their nests. He claims that when the fruits become
mature, the culms bend, thus placing the perigynia at ground level.

176 Rhodora [Vol. 86

C. comosa Boott Pseudocypereae

Marshes, floating peat mats of alkaline kettle-hole lakes, wet
ditches, and the borders of lakes, “bogs”, and swamps; frequent in
areas 3, 4, 5, infrequent to occasional in 1, 6, 7.

C. conjuncta Boott Vulpinae

A single station is known in Minnesota. It was collected from a
lowland forest bordering the Cannon River (Rice County), where
several clumps were observed just where the floodplain gives way to
higher ground. Of some interest, two other uncommon Minnesota
carices, C. davisii and C. grayi, were also observed in this lowland
forest, both of which were present in some abundance. Although C.
conjuncta somewhat resembles C. alopecoidea, the sheaths of the
former are conspicuously cross-puckered whereas those of the latter
are smooth. C. conjuncta is well known to the south and southeast
of us, such as in northern Missouri (Steyermark, 1963), Illinois
(Mohlenbrock and Ladd, 1978), and the southern half of Iowa
(Gilly, 1946), but it is uncommon or rare in northern Iowa (Gilly,
1946) and Wisconsin (Zimmerman, personal communication). It
appears to be very rare in Minnesota. This note represents the first
report of this species from the state.

Rice Co.: Cannon River Wilderness Area, floodplain forest, TII1IN, R20W, Sec.
34, Wheeler 5297 (MIN).

C. conoidea Willd. Griseae
Moist meadows, low prairies and, more rarely, thinly-wooded
areas; known only from a few sites scattered throughout the state.
Well known from the environs of the Twin Cities Army Arsenal
(Ramsey County), where it grows along the margins of Marsden
Marsh and also, albeit rarely, on partially-shaded wooded banks
overlooking the marsh. See comments under C. katahdinensis.

C. convoluta Mack. Bracteosae

Deciduous woodlands (mesic to dry) and thickets; frequent to
common in areas 5, 6, 7, occasional in 4, infrequent in 9, uncommon
in 3. Although some workers (e.g. Gleason and Cronquist, 1963) do
not treat this entity as distinct from C. rosea, in Minnesota these
two taxa are well-marked and easily distinguished, both morpho-
logically and in geographic distribution. Regarding the latter, C.
convoluta is restricted to the central and southern portions of
Minnesota whereas C. rosea occurs throughout most of the state.

1984] Wheeler & Ownbey — Minnesota Carices 177

Where the ranges of the two species overlap, C. rosea clearly grows
over a much wider range of soil types, with C. convoluta more or
less confined to rich sites.

C. crawei Dew. Granulares

Low prairies, moist meadows, swales, ditches, and marly shores
of lakes and, less commonly, railroad and highway embankments
(especially prairie strips); occasional to frequent in areas 4, 8,
infrequent in 2, 9, uncommon in I], 5.

C. crawfordii Fern. Ovales

Moist, open sites (e.g. meadows, ditches, clearings, roadside
embankments, borders of lakes and ponds, sandy shores and, less
commonly, the borders of “bogs” and swamps); frequent to
common in areas 1, 2, 3, infrequent to occasional in 4 and the
northern portions of 5, 6.

C. crinita Lam. Cryptocarpae

Mixed conifer-hardwood and alder swamps, woodland swales
and pond margins, wooded river banks and, less commonly, wet
ditches; occasional to frequent in areas 1, 3, infrequent in 2 and the
eastern half of 4, uncommon in the northern portions of 5, 6. In
Minnesota, only C. crinita, C. gynandra, and C. tuckermanii have
achenes that are consistently deeply invaginated on one side. See
comments under C. gynandra.

C. cristatella Britt. Ovales
Floodplain forests, abandoned river channels and ox-bows, moist
meadows, wet ditches, and the marshy margins of ponds, lakes, and
streams; occasional to frequent throughout the state, except in area
1, where it is uncommon. The plants often produce numerous
nonflowering pseudoculms along with the fertile culms.

C. crus-corvi Kunze Vulpinae

Two stations are known for this species in Minnesota: marshy
edge of the Mississippi River near Weaver, Wabasha County
[Fassett and Hotchkiss 3087 (GH)], wet site near Red Wing in
Goodhue County [Sandberg in 1885 (MIN)]. Fernald (1937) also
mapped it as occurring in east-central and extreme south-central
Minnesota, but specimens from these areas have not been seen. This
species has not been collected in the state for some fifty years, and
recent efforts to recollect it have failed. It appears to be very rare in
Minnesota (if not already extirpated from the state).

178 Rhodora [Vol. 86

C. cryptolepis Mack. Extensae

Moist to wet places, such as meadows, “bog” and swamp borders,
and the marshy margins of lakes; occasional in areas 1, 3, infrequent
in 2, 4 and the northern portion of 5. Some workers (e.g. Fernald,
1950) treat it as C. flava L. var. fertilis Peck.

C. davisii Schwein. & Torr. Gracillimae

Floodplain forests of large rivers in southeastern Minnesota; local
in areas 6, 7. It appears to be rare in the state. Probably best known
from the Cannon River Wilderness Area (Rice County), where it
grows in lowland forest (see comments under C. conjuncta).
Erroneously reported (Stevens, 1972; McGregor et al., 1977) for
North Dakota; see comments under C. formosa and also Wheeler,
1983a.

C. debilis Michx. Sylvaticae

Wet acid woods, wooded ridges bordering “bogs” and swamps
and, less commonly, sandy roadside embankments; infrequent to
occasional in the eastern portions of areas 3, 4, 5. Best known from
Anoka (Lemon, 1943) and Pine counties. Our material may be
referred to var. rudgei Bailey.

C. deflexa Hornem. Montanae

Mixed conifer-hardwood forests and swamps, rocky woods, pine
forests, cliffs and bluffs, and “bog” borders; occasional in area 1,
infrequent in 2, 3.

C. deweyana Schwein. Deweyanae

Deciduous woodlands (wet to dry), mixed conifer-hardwood
forests and swamps, conifer and alder swamps, clearings, and rocky
woods and bluffs; common in the northern half of the state,
occasional to frequent in the southern half (except in the west,
where it is uncommon).

C. diandra Schrank Paniculatae

Wet meadows, fens (usually associated with flarks in patterned
fens), floating peat mats surrounding alkaline kettle-hole lakes, wet
ditches, and “bog” and swamp borders; frequent in area 2,
occasional in 1, 3, 4, 5, uncommon in 6. In the Red Lake Peatland, it
occurs in flarks of patterned fens and on wooded islands as well as
in disturbed sites (Wheeler and Glaser, 1982; Wheeler er al/., 1983).

1984] Wheeler & Ownbey — Minnesota Carices 179

C. disperma Dew. Heleonastes

Conifer and mixed conifer-hardwood swamps, alder swamps,
“bog” borders, and the mossy shaded banks of streams; frequent to
common in areas |, 2, 3, occasional in 4 and the northern half of 5.
In the Red Lake Peatland, it is mostly confined to the sides and tops
of hummocks on the rich-fen wooded islands (Wheeler er a/., 1983).

C. eburnea Boott Albae

Steep wooded ridges and partially-shaded limestone and sand-
stone bluffs; occasional to frequent in areas 5, 6, 7, infrequent in 4,
8, 9, uncommon in 3. Best known from bluffs bordering the
Minnesota and Mississippi rivers and their tributaries, where it
often grows in dense mats under Juniperus. This species retains its
fruits for a long time after maturity (Hermann, 1940) and, indeed, it
is not unusual to find plants displaying perigynia produced from
two consecutive years.

C. eleocharis Bailey Divisae

Prairies, dry hillsides, railroad and highway embankments
(especially prairie strips), rock outcrops, and sand barrens; occa-
sional to frequent in the western half of the state, infrequent in the
eastern half. A strongly rhizomatous species sometimes occurring in
great abundance on dry hillsides and slopes in western Minnesota.
Some workers (e.g. Fernald, 1950) treat it as C. stenophylla
Wahlenb. var. enervis (C. A. Mey.) Kikenth.

C. emoryi Dew. Acutae

Along river banks, where it often forms large beds of nearly pure
stands; it also frequents, though less commonly, floodplain forests,
wet ditches, prairie swales, and the margins of ponds and lakes.
Frequent to common throughout the state, except in area |, where it
is uncommon. Clearly the most commonly-occurring and abundant
sedge along the rivers of our state. Some workers (e.g. Gleason and
Cronquist, 1963) treat is as C. stricta Lam. var. elongata (Boeckl.)
Gl.

C. exilis Dew. Stellulatae

Best known from the Red Lake Peatland (Wheeler and Glaser,
1979, 1982; Glaser ert al., 1981; Wheeler et a/., 1983), where it is
locally abundant in narrow strips of open poor-fen ecotone that
separate ombrotrophic bogs from minerotrophic fens; it also occurs,

180 Rhodora [Vol. 86

though less commonly, in flarks and on strings. Outside the Red
Lake area, the species is known only from a few patterned fens
(primarily in flarks) in the north-central and northeastern portions
of the state (Glaser, 1983; Wheeler, 1983b).

C. festucacea Willd. Ovales

A single station is known in Minnesota. It was collected from a
lowland woods bordering the Des Moines River, where a few
clumps were observed growing on slightly raised banks. The species
is distinguished from other Minnesota members of the Ovales by the
following combination of characters: spikelets with conspicuous
clavate staminate bases; inflorescence more or less straight, not
nodding; body of perigynium suborbicular and rather abruptly
narrowed to the beak. This species is well known to the south and
southeast of us, such as in Missouri (Steyermark, 1963), Illinois
(Mohlenbrock and Ladd, 1978), and the southern half of Iowa
(Gilly, 1946), but it is very uncommon or rare in North Dakota
(McGregor et al/., 1977) and Michigan (Voss, 1972b). It appears to
be very rare in Minnesota. This note represents the first report of
this species from the state.

Jackson Co.: Kilen Woods State Park, low woods, T103N, R35W, Sec. 17,
Wheeler 5828 (MIN).

C. filifolia Nutt. Filifoliae

Prairies, dry hillsides and eroded slopes, and sand barrens; occa-
sional in areas 8, 9, infrequent in the western portions of 4, 5. This
species grows in dense tufts that often appear to form rings,
apparently through some dying out of older plants at the center
‘Hudson, 1977); the old sheaths are almost invariably broken off
zvenly about 2 cm above ground level. In Minnesota, only C.
filifolia and C. obtusata consistently have a well-developed rachilla
within the perigynium. See comments under C. obtusata.

C. flava L. Extensae
Moist meadows, springy places in forests, and “bog” borders;
local in area 1. It appears to be rare in the state.

C. foenea Willd. Arenariae

Dry, open sites. In the north, the plant frequents the margins of
jack pine forests and pioneer hardwood stands, sandy ridges and
clearings, and sandy disturbed sites (roadside shoulders and
embankments, sand and gravel pits). In the southeast, it frequents

1984] Wheeler & Ownbey — Minnesota Carices 181

sand barrens, dry hillsides, railroad and highway embankments
(especially prairie strips), and sandy disturbed sites. Occasional to
frequent in the northern half of Minnesota, infrequent to occasional
in the east-central and southeastern portions of the state. A
rhizomatous species often abundant in dry, sandy soil. Some
workers (e.g. Gleason and Cronquist, 1963) still refer to this entity
as C. siccata Dew.

C. formosa Dew. Gracillimae

Known only from dry deciduous woodlands in Ramsey County,
where it is very local. It appears to be very rare in the state.
Although this species is reported by Fernald (1950) as extending no
farther west than Minnesota, it is also known from Richland
County, North Dakota (see comments under C. davisii and also
Wheeler, 1983a).

C. garberi Fern. Bicolores

One station is known for this species in Minnesota: a marshy
roadside near Cotton in St. Louis County (Wheeler, 1983b).
Although C. garberi is sometimes difficult to distinguish from C.
aurea, the Minnesota material has all the characteristics of the
former: perigynia densely crowded, strongly overlapping, granular,
whitish; pistillate scales rounded at the summits, not short-pointed.
In contrast, the pistillate scales of C. aurea are almost invariably
short-pointed and the perigynia are generally not crowded or
overlapping and the latter, when dry, are usually pale brown or
beige and not very granular. Unfortunately, a recent effort to
recollect the species near Cotton was unsuccessful. It appears to be
very rare in the state.

C. gracillima Schwein. Gracillimae

Deciduous forests (mesic to wet), alder swamps, mixed conifer-
hardwood forests and swamps, wooded banks of rivers and, less
commonly, moist meadows and highway embankments; common in
areas 1, 2, 3, 4, frequent in 5, occasional in 6, 7.

C. granularis Willd. Granulares

Moist meadows and clearings, fens, prairie swales, borders of wet
woods and, less commonly, wooded bluffs; occasional throughout
the state, except in the northeast, where it is unknown. Our material
may be referred to var. haleana (Olney) Porter.

182 Rhodora [Vol. 86

C. gravida Bailey Bracteosae
Dry hillsides, pastures and abandoned fields, railroad and
highway embankments (especially prairie strips), river banks,
woodland margins, prairies, and rock outcrops; frequent in the
southern half of the state, infrequent in the northern half.

C. grayi Carey Lupulinae

Floodplain forests of large rivers (especially the Mississippi
River); infrequent in areas 6, 7, rare in 4. Most Minnesota material
is var. grayi (with glabrous perigynia), but var. hispidula Bailey
(with hispidulous perigynia) is known from a floodplain woods near
Reads Landing in Wabasha County (Wheeler, 1979). Some workers
(e.g. Gleason and Cronquist, 1963; Reznicek and Ball, 1974) do not
recognize var. hispidula, and thus regard the presence or absence of
perigynium pubescence as part of the normal variation of the
species. These two taxa have not been mapped separately. See
comments under C. conjuncta.

C. gynandra Schwein. Cryptocarpae

Wet sites in northeastern Minnesota, especially near Lake
Superior (Wheeler, 1983b). It most often frequents the margins of
conifer swamps, wet meadows, and the banks of streams; infrequent
in area 1, uncommon in the eastern portion of 3. This entity is
sometimes treated (e.g. Fernald, 1950; Gleason and Cronquist,
1963) as a variety of C. crinita, as var. gynandra (Schwein.)
Schwein. & Torr., but these two taxa are well-marked and easily
distinguished in Minnesota and Wisconsin (Wheeler, 1983b) as well
as elsewhere (Voss, 1972b; Standley, 1983). Standley (1983) recently
found the two taxa to be distinct on the basis of morphological
differences and reproductive isolation. See comments under C.
crinita.

C. gynocrates Dre}. Dioicae

Conifer swamps, wooded mesotrophic “bogs” and, less com-
monly, alder swamps; occasional in areas 2, 3, infrequent in 1, 4,
uncommon in 5. In areas 4 and 5, it is confined to tamarack
swamps. A small, usually dioecious species that is easily overlooked,
particularly the staminate plants (however, the latter sometimes
bear one or two perigynia at the bottom of the spike).

1984] Wheeler & Ownbey — Minnesota Carices 183

C. hallii Olney Atratae
Prairie swales; local in area 8. It is a polymorphic species that
appears to be rare in the state.

C. haydenii Dew. Acutae

Wet sandy swales and moist meadows; occasional in the eastern
half of the state, infrequent in the western half. It is sometimes a
dominant species in wet sandy swales, where it often associates with
C. buxbaumii. In richer soils it is usually replaced by such species as
C. lacustris and C. atherodes.

C. heliophila Mack. Montanae

Prairies, dry hillsides, bluffs and cliffs, sand barrens, and railroad
and highway embankments (especially prairie strips); occasional to
frequent in areas 4, 5, 6, 7, 8, 9. Some workers (e.g. Fernald, 1950)
treat it as C. pensylvanica Lam. var. digyna Boeck.

C. hirtifolia Mack. Triquetrae

Mesic deciduous forests; occasional to frequent in areas 5, 6, 7,
infrequent in 4. Best known from the southeast, where the plant (all
parts of which are pubescent) sometimes occurs in abundance in
rich woodlands.

C. hitchcockiana Dew. Oligocarpae

Steep hardwood-covered slopes, wooded ravines and, less com-
monly, flat deciduous woodlands; occasional in area 5, infrequent in
6, 7. Best known from wooded bluffs of the Minnesota River
(Wheeler, 1981b).

C. houghtoniana Dew. Carex

Margins of pine stands, wet sandy shores, and sandy disturbed
sites (e.g. sand and gravel pits, railroad tracks, roadside shoulders
and embankments, clear-cut and burned-over areas); occasional to
frequent in areas 1, 2, 3, infrequent in 4, uncommon in 5, 6.
According to Hudson (1977), the rhizomes of this species grow
vigorously in response to disturbance.

C. hystericina Willd. Pseudocypereae

Conifer and mixed conifer-hardwood swamps, alder swamps,
shrub-carrs, wet meadows and ditches, marshy margins of lakes and
streams, and “bog” and swamp borders; frequent to common
throughout the state.

184 Rhodora [Vol. 86

C. interior Bailey Stellulatae

Conifer and mixed conifer-hardwood swamps, alder swamps,
fens, prairie swales, wet meadows and, less commonly, ditches,
pastures, and “bog” and swamp borders; occasional to frequent
throughout Minnesota, except in the southeast corner, where it is
very uncommon. In the southern half of the state, this species
primarily grows in tamarack swamps and in spring-fed, calcareous
fens. In the Red Lake Peatland, it grows in flarks of patterned fens,
in wet depressions on the wooded rich-fen islands, and in disturbed
sites (Wheeler er a/., 1983). In Minnesota, this species grows on
richer sites than the closely related species C. cephalantha and C.
angustior; this observation has also been reported from elsewhere
(Damman, 1964).

C. intumescens Rudge Lupulinae

Deciduous woodlands (mesic to wet), mixed conifer-hardwood
forests and swamps, alder swamps, and wooded river banks;
common in areas |, 2, 3, occasional to frequent in 4, 5, infrequent in
6. Our material may be referred to var. fernaldii Bailey.

C. katahdinensis Fern. Griseae
Known only from the environs of Iron Lake in northern St. Louis
County, where it grows on sandy beaches and amongst shore rocks
(Lakela, 1952; Wheeler, 1983b). Because in Minnesota C. katah-
dinensis and C. conoidea are well-marked and easily distinguished
both morphologically and ecologically (Wheeler, 1983b), they are
treated here as separate species. However, some workers (Ball and
White, 1982) claim that the northern populations of C. conoidea
consist of dwarf individuals with a crowded inflorescence whereas
the more southerly populations consist of tall individuals with a lax
inflorsecence. Boivin (1967) treats this entity as a form of C.
conoidea Willd., as forma katahdinensis (Fern.) Boivin.

C. lacustris Willd. Paludosae

Marshes, prairie swales, woodland ponds, ditches, wooded
swamps, and the borders of lakes, rivers, “bogs”, and swamps;
common throughout the state, except for the southeast and
southwest corners, where it is uncommon or unknown. It is often
the dominant plant in and around the margins of woodland ponds,
and it is one of the most common sedges bordering tamarack
swamps. However, in prairie swales it is often replaced by C.

1984] Wheeler & Ownbey — Minnesota Carices 185

atherodes. In the Red Lake Peatland proper it is infrequent in the
environs of ditches, but along creeks bordering the mire and in laggs
it is widespread and common (Wheeler er a/., 1983).

C. laeviconica Dew. Paludosae

River banks, abandoned river channels and ox-bows and, less
commonly, prairie swales, wet ditches, and the marshy borders of
lakes and ponds; occasional to frequent in the southern half of the
state and in the Red River Valley.

C. laevivaginata (Kiikenth.) Mack. Vulpinae

One station is known for this species in Minnesota: a wet site near
Spring Grove in Houston County [Rosendahl 456 (MIN)]. This
species appears to be very rare in the state, and it has not been
collected in Minnesota since 1902.

C. lanuginosa Michx. Carex

Meadows, marshes, swales, wet prairies, ditches, pastures and
abandoned fields, and the borders of lakes and ponds; frequent to
common throughout the state. One of the most commonly-
occurring carices in wet, open sites. Some workers (e.g. Gleason and
Cronquist, 1963) treat it as C. /asiocarpa Ehrh. var. latifolia
(Boeck]) Gilly.

C. lasiocarpa Ehrh. Carex

Open mesotrophic “bogs”, floating peat mats surrounding
alkaline kettle-hole lakes, fens, and “bog” and swamp borders;
common in areas 1, 2, 3, occasional to frequent in 4 and the
northern portion of 5, infrequent in 6, 8. Clearly the most common
and abundant sedge in rich fen in the northern half of the state. In
the Red Lake Peatland, it occurs in poor- and rich-fen sites and is
the dominant species in flarks of patterned fens (Glaser er al., 1981,
Wheeler and Glaser, 1982; Wheeler e¢ a/., 1983). Our material may
be referred to var. americana Fern.

C. laxiculmis Schwein. Laxiflorae

This sedge is confined to steep, north- and east-facing wooded
slopes of deep stream valleys in area 7, where it is locally abundant.
All of our specimens have conspicuously surrulate bract-sheaths
and sharply-angled culms, as well as gynaecandrous lateral spike-
lets. Material possessing these characters is sometimes treated
(Hermann, 1940) as var. copulata (Bailey) Fern. or even given

186 Rhodora [Vol. 86

species rank (Mackenzie, 1931-1935) as C. copulata (Bailey) Mack.
Fernald (1950), however, treats this entity as a hybrid between C.
laxiculmis and C. digitalis, and Zimmerman (personal communi-
cation) considers all of the Wisconsin material of this type to be C.
digitalis. Still others (e.g. Gleason and Cronquist, 1963) do not treat
this entity as distinct from C. laxiculmis. Of some interest, Hermann
(1940) states that var. copulata is a calciphile whereas the var.
laxiculmis grows best in neutral or only slightly calcareous soils.
The distribution of var. copulata is given by Hermann (1940) as
ranging from New Jersey to Missouri and northward to Michigan.
It seems clear that the calcareous bluffs of southeastern Minnesota
provide the habitats for the northwesternmost stations for this
entity in North America. This note represents the first report of this
species from the state.

Houston Co.: 9 mi. SW of Reno, north-facing wooded bluff overlooking
Winnebago Creek, 0.25 mi. E of Rte. 5, TIOIN, RSW, Sec. 15, Wheeler 4199 (MIN).
Beaver Creek Valley State Park, north-facing wooded bluff overlooking Beaver
Creek, TI02N, R6W, Sec. 5, Wheeler 4238 (MIN). 3 mi. SE of Caledonia, north-
facing wooded bluff overlooking South Fork Crooked Creek, T102N, R5W, Sec. 29,
Wheeler 4804 (MIN). Winona Co.: Whitewater State Park, 3 mi. SW of Alba on Rte.
74, east-facing wooded bluff overlooking Middle Fork Whitewater River, TIO7N,
RIOW, Sec. 20, Wheeler 3646 (MIN).

C. lenticularis Michx. Acutae
Rocky shores and sandy lake beaches; occasional to frequent in

area 1. Well known from the shores of Lake Superior, where it

grows amongst rocks and along the margins of rock pools.

C. leptalea Wahlenb. Polytrichoideae

Wooded mesotrophic “bogs”, conifer and mixed conifer-hardwood
swamps, alder swamps and, more rarely, fens: common in areas 1, 2,
3, frequent in 4, occasional in 5. In areas 4 and 5, this species is
mostly confined to tamarack swamps. In the Red Lake Peatland, it
occurs in open fens (flarks and string margins) and on wooded
islands, and it serves as an obligate rich-fen indicator (Glaser et al.,
1981; Wheeler e7 a/., 1983). The plants often grow in dense mats in
deeply-shaded places, and it has been suggested (Wheeler er ai.,
1983) that reduced light greatly favors asexual propagation in the
species.

C. leptonervia Fern. Laxiflorae
Deciduous forests (mesic to wet), mixed conifer-hardwood forests
and swamps, alder swamps and, less commonly, pioneer hardwood

1984] Wheeler & Ownbey — Minnesota Carices 187

stands; occasional to frequent in areas 1, 3 and the eastern portion
of 2. Whereas C. blanda is the most commonly-occurring member
of the Laxiflorae in our southern hardwood forests, C. /eptonervia
is the most commonly-occurring member of this group in our
northern hardwood and mixed conifer-hardwood forests. The
ranges of these two species in the state are more or less exclusive and
rarely, if ever, do the two species occur in the same forested area.

C. limosa L. Limosae

Open oligotrophic and mesotrophic “bogs”, floating peat mats
surrounding acid and alkaline kettle-hole lakes, fens, and openings
in conifer swamps; frequent in areas 1, 2, 3, occasional in 4,
infrequent in 5, 8, uncommon in 6, 9. In areas 4, 5, 6, 8, and 9, this
species is mostly confined to spring-fed, calcareous fens and to
Openings in tamarack swamps. In the Red Lake Peatland, the plant
frequents poor- and rich-fen sites, where it grows in flarks and, less
commonly, on strings (Wheeler e¢ a/., 1983). This species greatly
resembles the closely related C. paupercula (see comments under
this species) and, like it, has roots that are conspicuously covered
with numerous yellowish-tinged root hairs.

C. livida (Wahlenb.) Willd. Paniceae

Limy meadows and fens and, less commoniy, wet ditches and the
margins of conifer swamps (Wheeler, 1983b); infrequent to occa-
sional in areas 1, 2, but at some localities occurring in abundance. In
the Red Lake Peatland, this species grows in open poor- and rich-
fen sites (primarily associated with flarks and the margins of fen-
pools), and it is a frequent indicator of minerotrophy between
ombrotrophic and minerotrophic areas (Glaser et a/., 1981; Wheeler
and Glaser, 1982; Wheeler et a/., 1983). Our material may be
referred to var. radicaulis Paine [C. /ivida var. grayana (Dew.)
Fern. ].

C. lupulina Willd. Lupulinae

Floodplain forests, woodland swales and pond margins, wooded
swamps and, less commonly, wet meadows, ditches, and the marshy
borders of lakes and ponds; occasional to frequent in areas 3, 4, 5, 6,
7. Best known from lowland forests bordering the Mississippi and
St. Croix rivers.

C. lurida Wahlenb. Pseudocypereae
One station is known for this species in Minnesota: wet site near
Milaca in Mille Lacs County. Although collected in the state in

188 Rhodora [Vol. 86

1892, recent efforts to recollect it have failed. It appears to be very
rare in Minnesota (if not already extirpated from the state). In
Wisconsin (Zimmerman, personal communication), C. /urida is
known from a few sites along the Wisconsin and Black rivers.

C. meadii Dew. Paniceae

Prairies, sandy hillsides, railroad and highway embankments
(especially prairie strips), and the edges of rock outcrops; occasional
to frequent in areas 5, 8, 9, occasional in 4, infrequent in 6, 7. See
comments under C. tetanica.

C. media R. Br. Atratae

Rocky shores and margins of rock pools, steeply-wooded river
banks (especially near waterfalls) and, less commonly, moist
meadows, swampy ditches, and “bog” borders; infrequent to
occasional in area 1. Best known from the rocky shores of Lake
Superior. Some workers (e.g. Gleason and Cronquist, 1963) do not
treat this entity as distinct from C. norvegica Retz.

C. merritt-fernaldii Mack. Ovales

Sandy hillsides, gravelly banks, rock outcrops, cliffs and bluffs
and, less commonly, edges of pine stands; occasional in areas 1, 2,
infrequent in 3, uncommon in 4. Some workers (e.g. Gleason and
Cronquist, 1963) do not treat this entity as distinct from C. brevior
(Dew.) Mack.

C. michauxiana Boeck. Folliculatae

Known only from wet ditches near Schroeder in Cook County
(Butters and Abbe, 1953; Wheeler, 1983b). It appears to be very rare
in the state.

C. molesta Bright Ovales

River banks, pastures and abandoned fields, prairies, mesic
meadows, railroad and roadside embankments and, less commonly,
the margins of mesic hardwood stands; occasional in the southern
half of the state. Considered by some workers (Gleason and
Cronquist, 1963) to be a hybrid between C. brevior and C. normalis.
See Rothrock (1978) for a nomenclatural note.

C. muhlenbergii Willd. Bracteosae
Sandy hillsides, sand barrens, and dry prairies; occasional in the
eastern portions of areas 5, 6, 7. Probably best known from the

1984] Wheeler & Ownbey — Minnesota Carices 189

Weaver Dunes (Wabasha County) and the sand barrens of the
Anoka Sand Plain (e.g., Anoka and Sherburne counties).

C. muskingumensis Schwein. Ovales

Floodplain forests of large rivers, particularly the Mississippi
River (Wheeler, 1979); infrequent to occasional in areas 6, 7, rare in
5. The plants usually bear numerous, leafy pseudoculms along with
the fertile culms, both types of which are characteristically very stiff
and erect.

C. normalis Mack. Ovales
Deciduous woodlands (mesic to wet) and thickets; occasional to
frequent in areas 6, 7, infrequent in 3, 4, 5.

C. obtusata Lilj. Obtusatae

Sandy ridges and hillsides, sand barrens, and dry prairies; of local
occurrence in area 8 and the northwestern portion of 4. It appears to
be rare in the state but is quite abundant at some localities.
Probably best known from the Agassiz Dunes Natural Area (Polk
County), where it occurs on sand barrens. Of the two Minnesota
carices possessing a rachilla (see comments under C. filifolia), only
C. obtusata bears a scale-like appendage at the apex of the
structure.

C. oligocarpa Willd. Oligocarpae

Steep hardwood-covered slopes, wooded ravines and, more
rarely, flat deciduous woodlands; occasional in area 5S, infrequent in
6, 7. Best known from wooded bluffs of the Minnesota River
(Wheeler, 1981b).

C. oligosperma Michx. Vesicariae

Open ombrotrophic bogs, open oligotrophic “bogs”, and floating
peat mats surrounding acid kettle-hole lakes; frequent in areas 1, 2,
3, infrequent to occasional in the eastern portion of 4 and the
northeastern part of 5. It is one of the most commonly-occurring
and abundant sedges in open, oligotrophic sites in the northeastern
one-third of the state. In the Red Lake Peatland, where it is often a
dominant species, this plant frequents open ombrotrophic bogs, bog
drains, and open poor-fen areas (Glaser ef al., 1981; Wheeler and
Glaser, 1982; Wheeler ef a/., 1983).

190 Rhodora [Vol. 86

C. ormostachya Wieg. Laxiflorae

Hardwood and mixed conifer-hardwood forests (particularly
along the drier, more thinly-wooded margins), and wooded river
banks; known only from a few scattered sites in areas 1, 3, but it is
probably more common in the northeast than our present collec-
tions suggest. Some workers (e.g. Gleason and Cronquist, 1963)
treat is as C. laxiflora Lam. var. ormostachya (Wieg.) Gl.

C. pallescens L. Virescentes

Known only from moist sites on the Lake Superior terrace near
Duluth in St. Louis County (Lakela, 1954; Wheeler, 1983b). It
appears to be very rare in the state. Our material may be referred to
var. neogaea Fern.

C. pauciflora Lightf. Orthocerates

Open and partially-shaded ombrotrophic bogs, open and par-
tially-shaded oligotrophic “bogs”, floating peat mats surrounding
acid lakes and, more rarely, conifer swamps; occasional in areas I,
2, 3, uncommon in 4. In the Red Lake Peatland, this species
frequents open and partially-shaded poor-fens and, less commonly,
open and partially-shaded ombrotrophic bogs (Glaser er al/., 1981;
Wheeler and Glaser, 1982; Wheeler er al/., 1983). The plant grows
best where the water table is at or close to the peat surface, but it 1s
invariably absent from sites having a mud substrate (Wheeler er al.,
1983). The long-pointed, slender perigynia that are strongly reflexed
and easily detached at maturity are probably readily dispersed by
animals (Savile and Calder, 1953; Wheeler er al., 1983).

C. paupercula Michx. Limosae

Open and wooded ombrotrophic bogs, open and wooded
oligotrophic and mesotrophic “bogs”, floating peat mats surround-
ing acid and, less commonly, alkaline kettle-hole lakes, and conifer
and alder swamps; frequent in areas 1, 2, 3, occasional in 4 and the
northeastern portion of 5, rare in 7. In areas 4 and 5, this species is
mostly restricted to tamarack swamps. In the Red Lake Peatland, it
frequents open and wooded ombrotrophic bogs, open poor-fens,
and wooded poor- and rich-fen islands (Wheeler and Glaser, 1982;
Wheeler er al., 1983). The species seems to be more acid-tolerant
than the closely related C. /imosa (see comments under this species)
and, furthermore, it does not seem to frequent such nutrient-rich
sites as the latter (Wheeler er a/., 1983). Most Minnesota material is

1984] Wheeler & Ownbey — Minnesota Carices 19]

var. pallens Fern., but in the northeast var. irrigua (Wahlenb.) Fern.
is of occasional occurrence. Because intergrades between these two
taxa are frequent, they have not been mapped separately.

C. peckii Howe Montanae

Deciduous woodlands (mesic to dry), mixed conifer-hardwood
forests and swamps, pioneer hardwood stands, wooded river banks
and, more rarely, “bog” borders; frequent throughout the state,
except in areas 8 and 9, where it is occasional. Some workers (e.g.
Gleason and Cronquist, 1963) treat it as C. nigromarginata
Schwein. var. elliptica (Boott) Gl.

C. pedunculata Willd. Digitatae

Deciduous woodlands (wet to dry), mixed conifer-hardwood
forests and swamps, conifer and alder swamps, and pioneer
hardwood stands; occasional to frequent throughout the state,
except in areas 8 and 9, where it is very uncommon. The plant is
early fruiting (May), and its distribution in woodlands is often
clearly non-random, the majority of colonies occurring on rotting
logs, windthrow mounds, and near the bases of trees. This species 1s
a known myrmecochore (Handel, 1976, 1978b), and its “seeds”,
unlike in most species of Carex, are known to germinate the same
year they are produced (Handel, 1978b).

C. pensylvanica Lam. Montanae

Known from many habitats, particularly woodlands of all types;
it rarely occurs, however, in prairies or where the soil is extremely
wet (e.g. “bogs”, marshes, fens). Very common throughout Minne-
sota, except for the extreme northeast, where it is infrequent. This
species is probably the most commonly-occurring and widespread
sedge in the state, and it is sometimes very abundant in dry
deciduous woodlands.

C. plantaginea Lam. Laxiflorae

Steep hardwood-covered slopes and moist wooded ravines; very
local in east-central and southeastern Minnesota. It appears to be
very rare in the state.

C. praegracilis W. Boott Divisae

Low prairies, moist meadows, swales, wet depressions along
railroad and highway embankments (especially prairie strips),
ditches and, less commonly, moist places around rock outcrops; fre-

192 Rhodora [Vol. 86

quent in western and south-central Minnesota, adventive in the
north-central , northeastern, and extreme east-central portions of
the state (Wheeler, 1983b).

C. prairea Dew. Paniculatae

Low prairies, wet meadows and ditches, swales, marshes, and the
borders of lakes and streams; occasional throughout the state,
except in the northeast and southeast corners, where it is rare or
unknown. This species sometimes dominates wet meadows and
margins of lakes; its brightly-colored sheaths (yellow-brown to
bronze) are quite conspicuous. Some workers (e.g. Van Bruggen,
1976, McGregor er a/., 1977) still refer to this entity as C. prarisa
Dew.

C. praticola Rydb. Ovales

Bluff-tops, cliffs, and sandy and rocky ground; very local in area
1. Best known from bluff-tops bordering lakes of the Rove Slate
Formation, near the Minnesota-Ontario border (Cook County).
Although C. praticola can be easily confused with C. aenea, some
Minnesota specimens clearly have the characteristics of the former:
beak of the perigynium terete at the tip (the latter about 0.4 mm
long and white-hyaline or light brown), with the serrulate wing
abruptly terminating where the terete portion begins; perigynia less
than two-fifths as wide as long, 4.5-6 mm long (average about 5.3
mm ); spikelets in a flexuous inflorescence, silver-green or pale
brown. This species appears to be very rare in the state.

C. projecta Mack. Ovales

Floodplain forests, conifer and mixed conifer-hardwood swamps,
alder swamps, woodland swales and pond margins, wet clearings
and ditches, and the borders of “bogs”, lakes, and rivers; frequent in
areas 1, 2, 3, occasional in the eastern half of 4, infrequent in 5, 6.
This species often forms dense colonies by developing bud scales at
the base of sheaths on old prostrate culms that over-winter and
develop into independent plants the following year (Weatherby,
1945).

C. pseudocyperus L. Pseudocypereae

Marshy margins of lakes, “bog” borders and, less commonly,
wooded swamps, fens, and wet ditches; frequent in areas 2, 3,
occasional in 1, 4 and the northern half of 5, uncommon in 8, Inthe
Red Lake Peatland, this species serves as an obligate rich-fen

1984] Wheeler & Ownbey — Minnesota Carices 193

indicator and is best known from wooded rich-fen islands (Wheeler
and Glaser, 1982; Wheeler ef a/., 1983).

C. retrorsa Schwein. Lupulinae

Floodplain forests, conifer and mixed conifer-hardwood swamps,
alder swamps, shrub-carrs, marshes, wet ditches, and the borders of
lakes, rivers, and ponds; frequent to common in the northern half of
the state, occasional in the southern half.

C. richardsonii R. Br. Digitatae

Pine forests, dry hillsides, prairies, and sandy disturbed sites (e.g.
sand and gravel pits, roadside embankments); occasional in the
northern half of the state, infrequent in the southern half. Well
known from the Grand Rapids area (Itasca County), where it is of
rather frequent occurrence along the outer margins of red pine
stands.

C. rosea Willd. Bracteosae

Deciduous woodlands (wet to dry), thickets, mixed conifer-
hardwood forests and swamps, and conifer and alder swamps;
common throughout the state, except in the northeast, where it is
very infrequent. One of the most common sedges in deciduous and
mixed conifer-hardwood forests. Although there was a proposal
(Webber and Ball, 1979) to reject the name C. rosea (and C.
radiata), it was opposed by Boivin (1981), and, more recently, it was
not accepted by the Committee for Spermatophyta (Taxon 32:
623-624, 1983). See comments under C. convoluta.

C. rossii Boott Montanae

Two stations are known for this species in Minnesota: rocky soil
in Carlton County [Sandberg in 1891, exact location unknown
(MIN)]; an island in Lake Pokegama [Sandberg 276 (WIS)],
presumably in Pine County and mapped as such. C. rossii is a
Western Mountain element plant with disjunct populations known
from the Black Hills (Van Bruggen, 1976) and the Great Lakes
region (Fernald, 1935); the stations in northern Michigan (Kewee-
naw County) are the easternmost (Fernald, 1935). It is possible that
this species was more widespread in Minnesota during the
xerothermic period, and that it is a relict of wider eastern
distribution during that time. Presently, it appears to be very rare in
the state.

194 Rhodora [Vol. 86

C. rostrata Stokes Vesicariae

Marshy margins of lakes and rivers, “bog” borders, fens (usually
confined to flarks in patterned fens), prairie swales, shrub-carrs, and
wet ditches; common in wetlands throughout the state, but usually
replaced by C. atherodes in depressions that dry up during the
summer. In the Red Lake Peatland proper this species is infrequent
to occasional in flarks of patterned fens and in the environs of
ditches, and it is very common and abundant along creeks bordering
the mire and in laggs (Wheeler er a/., 1983). Our material may be
referred to var. utriculata (Boott) Bailey.

C. sartwellii Dew. Intermediae

Moist to wet meadows, prairie swales, fens, wet ditches and, less
commonly, swamp margins; occasional to frequent in the central
and western portions of the state, rare or unknown in the northeast
and extreme southeast.

C. saximontana Mack. Phyllostachyeae
Steep hardwood-covered slopes, wooded ravines, shaded river
banks, thickets and, less commonly, flat deciduous woodlands;
occasional in the western and south-central portions of the state.
Best known from bluffs of the Minnesota River, particularly
westward. On steep slopes, and especially where plants have
flowering culms hanging over precipitous ledges, the subglobose
fruits of this species often “roll” short distances downslope from a
parent plant; no doubt many other fruits are later washed
downslope by rain. For C. saximontana, and probably also for C.
backii, gravity (which is greatly aided by the roundish shape of the
fruits of these species) often seems to serve as an effective short-
distance dispersal mechanism for diaspores (Wheeler, 198 1a).

C. scirpiformis Mack. Scirpinae

Prairie swales; local in area 8 and the northwestern portion of 4.
It appears to be rare in the state. Some workers (e.g. Fernald, 1950)
treat it as C. scirpoidea Michx. var. scirpiformis (Mack.) O’Neill &
Duman.

C. scoparia Willd. Ovales

Moist meadows, ditches, clearings, wet sandy margins of lakes
and, less commonly, pastures and old fields, moist ledges, and “bog”
and swamp borders; frequent in the northern two thirds of the state,
occasional in the southern one third.

1984] Wheeler & Ownbey — Minnesota Carices 195

C. sparganioides Willd. Bracteosae
Deciduous forests (mesic to dry); occasional to frequent in areas
5, 6, 7, uncommon in 4.

C. sprengelii Spreng. Longirostres

Deciduous forests (dry, mesic and, more rarely, wet), river banks,
thinly-wooded hillsides and, less commonly, roadside embankments
and the margins of rock outcrops; common throughout the state,
except in the northeast, where it is very infrequent. The plants often
grow in large colonies, and they invariably have thick, pale brown
fibrous bases due to partial decomposition of old leaves (Thomas,
1982).

C. sterilis Willd. Stellulatae

Fens that are calcareous and persistently wet; infrequent to
occasional in areas 4, 5, 8, uncommon in 3, 6. Probably best known
from fens bordering the Mississippi River just southwest of
Mendota (Dakota County). Unlike some closely related species (e.g.
C. interior), the spikelets of this sedge, particularly the terminal one,
invariably lack prolonged clavate staminate bases. Some workers
(e.g. Gleason and Cronquist, 1963) treat it as C. muricata L. var.
sterilis (Carey) Gl.

C. stipata Willd. Vulpinae

Floodplain forests, conifer and mixed conifer-hardwood swamps,
alder swamps, marshes, woodland swales and pond margins, wet
ditches, shrub-carrs, moist to wet meadows and pastures, and the
borders of ponds and streams; frequent to common throughout the
state, except in the far west, where it is occasional.

C. stricta Lam. Acutae

Wet meadows, marshes, ditches, and the margins of swamps,
“bogs”, lakes, and streams; frequent to common throughout the
state, except in the southeast, where it is occasional. This sedge is
often a dominant in wet meadows, and at some localities its tussocks
are prominent over large areas. Most Minnesota material is var.
strictior (Dew.) Carey, but var. stricta also occurs at scattered
localities in the eastern half of the state; these two taxa have not
been mapped separately.

C. supina Willd. Lamprochlaenae
Known from cliff-talus at Clearwater Lake in northern Cook
County (Butters and Abbe, 1953; Wheeler, 1983b). There is also a

196 Rhodora [Vol. 86

report of the plant from South Fowl Lake (Bailey, 1892), but no
specimen has been seen. C. supina is a circumpolar, low-arctic plant
(Raymond, 1951; Given and Soper, 1981) with disjunct populations
known from southern Manitoba (Scoggan, 1957, 1978), southern
Ontario (Morton, personal communication), and northeastern
Minnesota, with the latter having the southernmost stations in
North America (Butters and Abbe, 1953). It is very possible that this
species was more common in Minnesota during late-glacial time,
and that it is a relict of wider distribution in the past (Butters and
Abbe, 1953; Wheeler, 1983b). Some support for this comes from
Michigan (Miller and Benninghoff, 1969), where macrofossil
material of this species has been reported (dated between 13,300 and
12,500 years B.P.). At the present time it appears to be very rare in
Minnesota. Our material may be referred to var. spaniocarpa
(Steud.) Boivin [C. supina ssp. spaniocarpa (Steud.) Hult. ].

C. sychnocephala Carey Ovales

Damp sandy shores of lakes and streams, swales, marshes, and
moist to wet meadows; occasional to frequent throughout the state,
except in the northeast and southeast corners, where it is un-
common. According to Hudson (1977), it appears to be a plant of
the early stages of succession, not persisting for long after a closed
perennial cover gets established.

C. tenera Dew. Ovales
Thinly-wooded hillsides, woodland margins, wooded river banks,
and moist meadows; occasional to frequent throughout the state.

C. tenuiflora Wahlenb. Heleonastes

Open and wooded oligotrophic and mesotrophic “bogs”, floating
peat mats surrounding acid and, more rarely, alkaline kettle-hole
lakes, fens (usually associated with strings in patterned fens), and
conifer swamps; occasional in areas 1, 2, 3, uncommon in 4, 5. In
areas 4 and 5, it is mostly restricted to tamarack swamps. In the Red
Lake Peatland, it grows primarily on the strings of patterned fens
but is also known from hummocks in the poor- and rich-fen wooded
islands (Wheeler er al, 1983).

C. tetanica Schkuhr Paniceae

Low prairies, swales, moist meadows, ditches, and the marshy
margins of ponds; frequent to common in areas 5, 8, 9, occasional in
4 and the western portion of 2, infrequent in 3, 6, introduced in |

1984] Wheeler & Ownbey — Minnesota Carices 197

(rare). Dried herbarium specimens of C. tetanica and C. meadii are
sometimes difficult to separate, but in the field the two entities are
fairly distinctive. C. tetanica grows primarily in wet, peaty sites and
typically has narrow leaves (2-4.5 mm wide) and pistillate spikelets
that are often loosely flowered at the base (the lower perigynia
barely or not overlapping). In contrast, C. meadii grows in much
drier sites, such as in dry prairies, and generally has broader leaves
(3-7 mm wide) and pistillate spikelets that are usually tightly
compacted, even at the base. The distributions of the two taxa in
Minnesota are somewhat similar, but some differences are note-
worthy. C. tetanica is of frequent occurrence in peaty meadows in
Lake of the Woods, western Beltrami, and Clearwater counties, but
C. meadii is unknown from these counties (they essentially lack
prairie sites). On the other hand, C. tetanica is very rare or absent
from the extreme southeastern corner of Minnesota (part of the
Driftless Area), an area lacking low, wet, calcareous sites. However,
C. meadii is known from various “goat” prairies in this area. The
same situation, as regards the presence of C. meadii and the absence
of C. tetanica, also seems to hold throughout the remainder of the
Driftless Area (Hartley, 1966).

C. tonsa (Fern.) Bickn. Montanae

Margins of pine forests, sand barrens, sandy ridges and rocky
ledges, and sandy disturbed sites (e.g. sand and gravel pits, roadside
shoulders and embankments, clearings and openings); occasional in
the eastern half of the state, particularly northward. Here we choose
to be traditional and treat C. tonsa and C. umbellata as separate
species, but because in Minnesota these two entities more or less
frequent the same habitats and sometimes grow together, one may
be justified in treating them as varieties of the same species. When
treated as a variety of C. umbellata (as the epithet umbellata is used
by Fernald (1942, 1950) and in this paper), this entity becomes var.
tonsa Fern. But for those (e.g. Mackenzie, 1931-1935, 1940; Voss,
1972b) who recognize C. rugosperma Mack. to be the true C.
umbellata Schkuhr ex Willd., then this entity, when treated as a
variety, becomes C. rugosperma var. tonsa (Fern.) E. Voss. See
comments under C. abdita and C. umbellata.

C. torreyi Tuckerm. Virescentes
Thinly-wooded hillsides and coulees and, less commonly, prairies,
mesic meadows, and sparsely-wooded river banks; infrequent to

198 Rhodora [Vol. 86

occasional in the western half of the state, uncommon in the eastern
half. Although most recent manuals (e.g. Fernald, 1950; Gleason
and Cronquist, 1963; Hermann, 1970) give this species as ranging no
farther eastward than Minnesota, it also occurs, albeit rarely, in
extreme western Wisconsin (Read, 1976; Zimmerman, 1976).

C. tribuloides Wahlenb. Ovales

Floodplain forests, woodland swales and pond margins and, less
commonly, moist meadows, roadside ditches and embankments,
and the margins of lakes and ponds; occasional to frequent in the
eastern half of the state (particularly southward), infrequent in the
western half. Like C. projecta (see comments under this species), C.
tribuloides also produces shoots on old prostrate culms that become
an effective means of vegetative reproduction (Weatherby, 1945).

C. trichocarpa Schkuhr Paludosae

River banks, abandoned river channels and ox-bows, and wet
prairies; known from scattered sites throughout the state, except in
the north-central and northeastern portions, where it is unknown.
Because in Minnesota fruiting material of this species often seems to
be scarce or absent, it may well be more common in the state than
our present collections suggest.

C. trisperma Dew. Heleonastes

Wooded ombrotrophic bogs, wooded oligotrophic and meso-
trophic “bogs”, and conifer swamps; frequent to common in areas I,
2, 3, occasional in 4, uncommon in the northern portion of 5.
Clearly one of the most frequently-occurring carices in wooded
“bogs” in the northern half of the state. In areas 4 and 5, it is mostly
restricted to tamarack swamps. In the Red Lake Peatland, where it
is often a dominant species, this plant occurs in forested ombro-
trophic bogs and on wooded poor- and rich-fen islands (Glaser ef
al., 1981; Wheeler and Glaser, 1982; Wheeler er al., 1983).

C. tuckermanii Dew. Vesicariae

Conifer and mixed conifer-hardwood swamps, alder swamps,
floodplain forests, woodland swales and pond margins, abandoned
river channels and ox-bows and, less commonly, wet meadows;
occasional to frequent in areas |, 2, 3, 4, uncommon in 5. See
comments under C. crinita.

1984] Wheeler & Ownbey — Minnesota Carices 199

C. typhina Michx. Squarrosae

Floodplain forests of large rivers, especially the Mississippi and
St. Croix rivers (Wheeler, 1979); very infrequent in areas 5, 6, 7. In
the extreme southeastern corner of the state, this species is often
associated with C. muskingumensis and C. grayi and, more rarely,
with C. davisii.

C. umbellata Willd. Montanae

Margins of pine forests, sand barrens, and sandy disturbed sites
(e.g. sand and gravel pits, roadside shoulders and embankments,
clear-cut and burned-over areas, railroad tracks, trails and paths);
occasional in the eastern half of the state, infrequent in the western
half. It is considered by Handel (1978a) to be a myrmecochore.
Some workers (e.g. Mackenzie, 1931-1935, 1940; Voss, 1972b) call
this entity C. rugosperma Mack. See comments under C. abdita and
C. tonsa.

C. vaginata Tausch Paniceae

Wooded mesotrophic “bogs” and conifer swamps; occasional in
areas 1, 2 and the northern portion of 3. Although frequenting
“bogs” and swamps, this circumpolar species seems to be uncom-
mon in upland peatlands in the state. The American plant is
sometimes distinguished (e.g. Mackenzie, 1931-1935, 1940) from
the Eurasian entity as C. sa/tuensis Bailey.

C. vesicaria L. Vesicariae

Wet meadows, marshes, margins of lakes and streams and, less
commonly, wet ditches and “bog” and swamp borders; frequent in
areas 1, 3, occasional in 2, 4, infrequent in 5, 6.

C. viridula Michx. Extensae

Peaty meadows and wet sandy margins of lakes and, less
commonly, roadside ditches and the edges of rock pools; occasional
in the northern half of the state, infrequent in the southern half. Best
known from northwestern Minnesota, where it is locally abundant
in wet meadows, particularly those dominated by C. /asiocarpa.

C. vulpinoidea Michx. Multiflorae

Moist to wet meadows, marshes, ditches, shrub-carrs, the margins
of lakes and streams and, less commonly, swampy woods; frequent
to common throughout the state. One of the most frequently-
occurring sedges in roadside verges.

200 Rhodora [Vol. 86

C. woodii Dew. Paniceae

Locally abundant in rich deciduous woodlands; known from
scattered sites in areas 3, 6, 7. Because the plant fruits very early
(about mid-May and even earlier), it is easily overlooked at other
times. Thus, it is probably more common in Minnesota than present
collections suggest. It was first correctly reported for the state by
Hartley (1966); an earlier, erroneous report (Moore and Tryon,
1946) was based on a specimen of C. blanda. Some workers (e.g.
Gleason and Cronquist, 1963) treat this entity as C. tetanica
Schkuhr var. woodii (Dew.) Wood.

C. xerantica Bailey Ovales

Known from bluff-tops bordering Watab Lake in northern Cook
County (Butters and Abbe, 1953; Wheeler, 1983b). The isolated
presence of C. xerantica in Cook County may indicate that the plant
was more widespread in the state during the xerothermic period.
However, because this species is known from stations in North
Dakota (McGregor er a/., 1977) and Manitoba (Scoggan, 1957,
1978), its migration to Minnesota in more recent times is possible
(Wheeler, 1983b). Presently, it appears to be very rare in the state.

DOUBTFUL AND EXCLUDED TAXA

Carex argyrantha Tuckerm.

Fernald (1950) and Scoggan (1978) included Minnesota within
the range of C. argyrantha, apparently based on an earlier listing for
the state by Moore and Tryon (1946). No specimen, however, has
been seen to support its presence in the state. This taxon is known
neither from Wisconsin (Zimmerman, personal communication)
nor Manitoba (Scoggan, 1957, 1978) and, based on the distribution
for the species given by Gleason and Cronquist (1963), it seems
unlikely that it occurs in Minnesota.

Carex digitalis Willd.

Reported for Minnesota by Mackenzie (1931-1935), but no
confirming specimen has been seen. It seems unlikely that this
eastern species (ranging eastward from eastern Wisconsin) occurs in
the state.

Carex lupuliformis Sartwell
Reported for Minnesota by Mackenzie (1931-1935), apparently
based on an earlier report made by Sheldon (1894). Some recent

1984] Wheeler & Ownbey — Minnesota Carices 201

authors (Fernald, 1950; Gleason and Cronquist, 1963; Scoggan,
1978) have also included Minnesota within the range of the species.
Owing to the lack of a confirming specimen, it must be excluded
from the flora at the present time. But because the species occurs in
Wisconsin (Read, 1976) and Iowa (Gilly, 1946), it is not improbable
that it does extend into southeastern Minnesota.

Carex panicea L.

This European species was reported for Minnesota by Fernald
(1950) and Hultén (1958), apparently based on an earlier listing for
the state by Moore and Tryon (1946). No confirming specimen for
Minnesota has been seen. Although this species is now naturalized
along the eastern coast of North America (Fernald, 1950; Hulten,
1958), its occurrence in Minnesota seems highly unlikely.

Carex picta Steud.

A specimen of C. picta collected by J. A. Stevenson in 1917 (June
22) and purported to be from Beltrami County (near Bemidji) is in
the Univeristy of Minnesota Herbarium. On the same herbarium
sheet is a specimen of C. willdenovii Willd., also otherwise unknown
from Minnesota. Both C. picta and C. willdenovii were reported
from Minnesota by Fernald (1950), apparently based on the Moore
and Tryon (1946) list. Neither species is known from Wisconsin
(Zimmerman, personal communication) or lowa (Gilly, 1946).
Because both of these carices would be so far from the ranges given
for them by some workers (e.g. Gleason and Cronquist, 1963), it
seems best to tentatively reject them both as part of the Minnesota
flora and to suggest confusion in place record labeling. According to
Gleason and Cronquist (1963), C. picta ranges from southern
Indiana to Alabama and Louisana, and C. willdenovii from
Massachusetts to Ontario and southern Indiana then south to
Georgia and Texas.

Carex squarrosa L.

An immature specimen of C. squarrosa collected by Rev. Z. L.
Chandonnet (no date or number given) and purported to be from
Mahnomen County (“White Earth Reservation, Fish Lake”) is in
the University of Minnesota Herbarium. Although reported from
Minnesota by various workers (e.g. Moore and Tryon, 1946;
Fernald, 1950; McGregor er a/., 1977), the locality is so far out of
range as to be suspicious. Therefore, we tentatively reject it from the

202 Rhodora [Vol. 86

Minnesota flora, awaiting further verification. The species is not
known from Wisconsin (Zimmerman, personal communication),
and it is apparently restricted to the southern one third of Iowa
(Gilly, 1946).

Carex suberecta (Olney) Britton

Reported for Minnesota by Mackenzie (1931-1935) and others
(Fernald, 1950; Gleason and Cronquist, 1963), but no confirming
specimen has been seen. Because this species is known from
Wisconsin (Read, 1976) and Iowa (Gilly, 1946), it may well occur in
southern Minnesota.

Carex torta Tuckerm.

Mackenzie (1931-1935) reported C. torta from Minnesota
apparently on the basis of a collection (MIN) made in Houston
County (Winnebago Valley) by H. L. Lyon in 1899 (June 8). Some
recent authors (Moore and Tryon, 1946; Fernald, 1950; Gleason
and Cronquist, 1963; Scoggan, 1987) have also included Minnesota
within the range of the species, apparently based on Mackenzie’s
report. However, the specimen upon which the original report was
based is actually C. emoryi Dew.

Carex willdenovii Willd.
It seems best to reject C. willdenovii as part of the Minnesota
flora. For specific details see under C. picta above.

ACKNOWLEDGMENTS

We thank Prof. Eville Gorham for reading and criticizing the
manuscript; Dr. Paul H. Glaser and Mr. Welby Smith for placing
their Carex collections at our disposal; Dr. Anton A. Reznicek, Dr.
James H. Zimmerman, and Dr. John K. Morton for information on
the status of some carices in Michigan, Wisconsin, and Ontario,
respectively; and the curators of those herbaria whose specimens
were used in the preparation of the maps. We gratefully acknowl-
edge the Hayden Fund of the University of Minnesota (Department
of Botany) for financial support to help with publication costs.

1984] Wheeler & Ownbey — Minnesota Carices 203

DISTRIBUTION
MAPS OF
142
MINNESOTA
CARICES

CAREX ADUSTA CAREX AENEA

| q
a

CAREX ALBURSINA CAREX ALOPECOIDEA

204 Rhodora [Vol. 86

CAREX ANGUSTIOR

A Ca ae

CAREX ARCTA CAREX ARCTATA

1984] Wheeler & Ownbey — Minnesota Carices 205

se

°
CAREX ASSINIBOINENSIS CAREX ATHERODES

CAREX AUREA

e : +

we
olf

| - xr o : Tt
poet | Perr

CAREX BEBBI I CAREX BICKMELLI!

206

1984] Wheeler & Ownbey — Minnesota Carices 207

CAREX CEPHALANTHA CAREX CEPHALOIDEA
i
Sy NS
3
ibe: J |
Eri Tt am

CAREX CEPHALOPHORA CAREX CHORDORRHIZA

208 Rhodora [Vol. 86

aa Lae
oak ‘

| fle
Peer La

CAREX CONVOLUTA CAREX CRAWE!

1984] Wheeler & Ownbey — Minnesota Carices 209

CAREX CRAWFORD! CAREX CRINITA

mee

CAREX CRUS-CORV!

CAREX CRYPTOLEPIS CAREX DAVISI|

210

1984] Wheeler & Ownbey — Minnesota Carices 211

SS
al |
q_ Ce]
CAREX ELEOCHARIS CAREX EMORY!
SD SS
CCCI

CAREX EXILIS CAREX FESTUCACEA

CAREX FILIFOLIA CAREX FLAVA

212 Rhodora [Vol. 86

1984] Wheeler & Ownbey — Minnesota Carices 213

SB
jBas
| |
CAREX GRAYI CAREX GYNANDRA
é e
BW
|
1 hd Sd Oe
CAREX GYNOCRATES CAREX HALLII
SS at
io
re :

e
CAREX HAYDEN! | CAREX HELIOPHILA

214 Rhodora [Vol. 86

1984]

Wheeler & Ownbey — Minnesota Carices

CAREX KATAHDINENSIS CAREX LACUSTRIS

[
[[ [1 |

CAREX LAEVIVAGINATA

CAREX LAEVICONICA

e e
CAREX LANUGINOSA CAREX LASIOCARPA

215

216 Rhodora [Vol. 86

1984] Wheeler & Ownbey — Minnesota Carices 217

CAREX MERRITT-FERNALDI! CAREX MICHAUXIANA

218 Rhodora [Vol. 86

=

CAREX MOLESTA CAREX MUHLENBERG! |

CAREX MUSKINGUMENSIS

Sse Lote

CAREX OBTUSATA CAREX OL|GOCARPA

1984] Wheeler & Ownbey — Minnesota Carices 219

ne

CAREX OLIGOSPERMA

CAREX PALLESCENS

i

‘i a aa

CAREX PAUPERCULA CAREX PECKI!

220 Rhodora [Vol. 86

CAREX PRAIREA

1984] Wheeler & Ownbey — Minnesota Carices 221

-
oh
* °
e
e
CAREX PROJECTA CAREX PSEUDO-CYPERUS

CAREX ROSEA CAREX ROSSI!

222 Rhodora [Vol. 86

1984] Wheeler & Ownbey — Minnesota Carices

223

ae

CAREX STERILIS

° aN © SON
4 - ” Y rR ° fo
. ee r bg oo e e
e e e bd
e e
e
ie e

rial Chel fe
CAREX STIPATA CAREX STRICTA

| |

CAREX SUPINA

CAREX SYCHNOCEPHALA

224 Rhodora [Vol. 86

1984] Wheeler & Ownbey — Minnesota Carices 225

DE a 2

CAREX TRICHOCARPA CAREX TRISPERMA

CAREX UMBELLATA CAREX VAGINATA

226

L] TT J

CAREX VESICARIA

1 ale
[el*] e %% pee

CAREX VULPINOIDEA

Rhodora

|

[ |

CAREX VIRIDULA

CAREX KERANTICA

[Vol. 86

1984] Wheeler & Ownbey — Minnesota Carices a27

LITERATURE CITED

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BaiLey, L. H. 1892. Notes on Carex XVI. Bot. Gaz. 17: 148-153.

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BERNARD, J. P. 1959. Le Carex assiniboinensis Boott et sa forme stolonifere. Nat.
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BiLL, J. P. 1930. Carex knieskernii Dewey. Rhodora 32: 162-166.

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1981. Should we reject Carex rosea and Carex radiata. Taxon 30:
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BrusH, G. S. 1982. An environmental analysis of forest patterns. Am. Sci. 70:
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Butters, F. K., & E. C. Aspe. 1953. A floristic study of Cook County,
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CHENEY, L. S. 1893. A contribution to the flora of the Lake Superior region.
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Curtis, J.T. 1959. The Vegetation of Wisconsin. University of Wisconsin Press,
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CusHING, E. J. 1965. Problems in the Quaternary phytogeography of the Great
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Daman, A. W. H. 1964. Key to the Carex species of Newfoundland by
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Fassett, N. C. 1957. A Manual of Aquatic Plants. Revision appendix by E. C.
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FERNALD, M. L. 1935. Critical plants of the upper Great Lakes region of Ontario
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1937. Local plants of the inner coastal plain of southeastern Virginia.

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1942. Critical notes on Carex. Rhodora 44: 281-331.

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228 Rhodora [Vol. 86

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1978a. New ant-dispersed species in the genera Carex, Luzula, and

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1978b. The competitive relationship of three woodland sedges and its
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1970. Manual of the Carices of the Rocky Mountains and Colorado

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1974, Manual of the Genus Carex in Mexico and Central America. U. S.
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1944. Notes on Minnesota plant life. Rhodora 46: 25-28.

1952. Previously unreported plants from Minnesota. Rhodora 54:

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1984] Wheeler & Ownbey — Minnesota Carices 229

1954. Previously unreported plants from Minnesota with additional
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230 Rhodora [Vol. 86

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1984] Wheeler & Ownbey — Minnesota Carices 231

1983b. Carex of northeastern Minnesota: Cook, Lake, St. Louis, and

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Peatland, northern Minnesota. Mich. Bot. 18: 137-142.

,& . 1982. Vascular plants of the Red Lake Peatland, northern

Minnesota. Mich. Bot. 21: 89-93.

, & , E. GorHam, C. M. Wetmore, F. D. Bowers, & J. A.
JANSSENS. 1983. Contributions to the flora of the Red Lake Peatland,
northern Minnesota, with special attention to Carex.Am. Midl. Nat. 110: 62-96.

WHEELER, W. A. 1900. A contribution to the knowledge of the flora of
southeastern Minnesota. Minn. Bot. Stud. 2: 353-416.

1901. A contribution to the knowledge of the flora of the Red River
Valley in Minnesota. Minn. Bot. Stud. 2: 569-600.

WRIGHT, H. E., JR. 1972. Physiography of Minnesota. Jn: P. K. Sims and G. B.
Morey (eds.). Geology of Minnesota: a Centennial Volume. Minn. Geol. Surv.,
Minneapolis. pp. 515-548.

YAEGER, D. P., & J. R. BORCHERT. 1971. Landscape regions in Minnesota (map).
In: Project 80 Staff Report No. 1, Minnesota Resource Potentials in State
Outdoor Recreation. Minnesota Dept. of Natural Resources and Minnesota
State Planning Agency, St. Paul.

ZIMMERMAN, J. H. 1976. Carex. In: N. C. Fassett. Spring Flora of Wisconsin. 4th
ed. revised by O. S. Thomson. University of Wisconsin Press, Madison. pp.
32-70.

DEPARTMENT OF BOTANY
UNIVERSITY OF MINNESOTA
ST. PAUL, MINN. 55108

NEW ENGLAND NOTES

NOTE ON THE FIRST STATION OF
DENTARIA LACINIATA MUHL. (BRASSICACEAE)
REPORTED FROM MAINE

SALLY C. ROONEY

Dentaria laciniata occurs in rich woods, wooded bottom lands,
and on calcareous rocky hillsides from western Quebec and
Vermont west to Minnesota and Nebraska, and south to Florida
and Louisiana.

In the spring of 1981, Ruth Ellis and I discovered a small clump of
Dentaria laciniata growing on a west-facing slope of Westford Hill,
Hodgdon (Aroostook Co.), Maine. This species had not been
previously reported from the State (Bean, er a/., 1966; Critical Areas
Program, 1981). Cut-leaf toothworth is rare in New Brunswick
(Hinds, 1983), New Hampshire (Storks and Crow, 1978), and
Quebec (Bouchard er al., 1983). The only site with proximity to
Hodgdon is located in upper Woodstock, New Brunswick, Canada
(ca. about 24 km to the northeast).

In June of 1982, Martin Rasmussen (pers. comm.) a botanist
from Caribou, Maine, located another small group of plants near
our original site on Westford Hill; these individuals were in poor
condition and about to die for the season.

Mapping of the population was undertaken for the Maine Critical
Areas Program in May of 1983; the D. /aciniata site was calculated
to occupy about 5 ha on the west and south slopes of Westford Hill.
The population is large, numbering thousands of individuals,
patchily distributed over the steeper rocky portions of the hill. Two
voucher specimens were collected for deposition in the herbaria of
the University of Maine (MAINE) and the New England Botanical
Club (NEBC).

Dentaria laciniata is an early blooming ephermeral, dying down
before tree leaves appear. At the Hodgdon site, D. Jaciniata
dominates the herbaceous vegetation on the disintegrating calcare-
ous slopes. It grows under a Quercus rubra/Acer saccharum
canopy, and is commonly associated with Dicentra cucullaria,
Claytonia caroliniana, Trillium erectum and Erythronium ameri-
canum.,

233

234 Rhodora [Vol. 86

D. laciniata is not rare south of Maine. After consultation with
many botanists, the site in Hodgdon remains the single known
Maine station for this species. The site is a Registered Critical Area.

LITERATURE CITED

BEAN R. C., C. D. RICHARDS, AND F. HyLAND. 1966. Revised checklist of the
vascular plants of Maine. Bull. of Josselyn Bot. Soc. of Maine.

BouCHARD, A., D. BARABE, M. DuMAIS AND S. Hay 1983. The rare vascular
plants of Quebec. National Museums of Canada. Syllogeus No. 48 Ottawa,
Canada.

CRITICAL AREAS PROGRAM. 1981. Rare vascular plants of Maine. State Planning
Office. Augusta, Maine.

Hinps, H. R. 1983. Rare vascular plants of New Brunswick. National Museums
of Canada. Syllogeus No. 50. Ottawa, Canada.

Storks, I. M. AND G. E. Crow. 1978. Rare and endangered vascular plant
species in New Hampshire. The New England Botanical Club in cooperation with
the U. S. Fish and Wildlife Service (Newton Corner, Mass.).

4 BROOK STREET
HOULTON, MAINE 04730

1984] New England Notes 235

TWO NEW LOCALITIES FOR MEDEOLARIA FARLOWII
IN NEW ENGLAND

DONALD H. PFISTER

Heretofore, only three localities have been reported for the
distinctive parasite Medeolaria farlowii Thaxter. These localities
were given by Thaxter (1922) in the paper describing the new genus
and species as Magnolia, Massachusetts; Kittery Point, Maine; and
Chocorua, New Hampshire. Despite both the widespread occur-
rence of the host, Medeola virginiana, and the peculiarity of the
fungus with its resultant hypertrophy, the fungus was known only
from these localities. In September, 1970 Richard P. Korf and I
collected it again on the hillside near William G. Farlow’s former
house where Thaxter had made a large collection in 1904. That
collection allowed a study which resulted in Korf’s (1972) delimita-
tion of the monotypic order Mediolariales and family Mediolariaceae
for the monotypic fungus genus Medeolaria. Two new localities for
this fungus are here reported: in damp mixed woods near Stevens
Corner Road, West Newfield, Maine, on the property of Reed and
Kathryn Rollins, September 1983; and in mixed woods on Parker
Trail, Mt. Monadnock, New Hampshire, October 12, 1983.

I call attention to this intriguing, seemingly rare parasite in hopes
that others will look for infected plants to help develop a more
complete picture of its distribution. Infected plants are character-
ized by having shortened, swollen internodes between leaf pseudo-
whorls. Below the lower pseudowhorl, the stem is thickened and is
at first green, later becoming brownish with olive to yellowish-green
patches. It is in these patches, which are mostly paraphyses, that asci
are found in mature specimens. The asci are thick-walled except at
the tip where they become thinned. Each ascus has eight bilaterally
asymmetrical ascospores, dark brown at maturity and longi-
tudinally striate. The above details agree with Thaxter’s description.

Ascus dehiscence was discussed by Korf (1972) who questioned
whether it might not deliquesce rather than split. Study of the large
collection from Maine showed empty asci each with a rupture of the
apical region. The apical part of the ascus is thin and apparently
breaks down at maturity. No spores were released when ascus-
bearing portions were suspended above agar.

236 Rhodora [Vol. 86

Many of the swellings had holes which may have been made by
foraging insects. This observation suggests that insects might be
involved in spore dispersal, which might explain the spotty
distribution. Incidence of infection in populations is also of note; in
some populations numerous infected plants can be found in close
proximity, while in others only single infected plants are found.
While this spottiness may have to do with systemic infection in the
vegetatively reproducing plants of Medeola (growth patterns in
Medeola were outlined by Bell in 1974), essentially nothing has been
worked out regarding the infection cycle of this parasite.

Attempts were made to establish the fungus in culture. It grew
slowly out of excised plant tissue and after two days was overtaken
by other fungi. No spores were formed by Medeolaria farlowii in
culture. The hyphae are characteristically hyaline and branch
repeatedly at nearly right angles.

ACKNOWLEDGMENTS

Work on this project was supported in part by National Science
Foundation grant DEB 80-23018. I thank Reed and Kate Rollins
for access to their property and Gennaro Cacavio for help in
collecting. To establish distributional records, all of the collections
of Medeola virginiana in the Harvard University Herbaria were
examined as well as those in the New England Botanical Club
collection. However, no infected plants were found, which is
perhaps a testimony to good collecting.

LITERATURE CITED

Bett, A. D. 1974. Rhizome organization in relation to vegetative spread in
Medeola virginiana. J. Arnold Arboretum 55: 458-468.

Korf, R. P. 1972. Synoptic key to the genera of the Pezizales. Mycologia 64:
937-994.

THAXTER, R. 1922. Note on two remarkable Ascomycetes. Proc. Amer. Acad.
Arts 57: 425-436.

HARVARD UNIVERSITY HERBARIA
22 DIVINITY AVENUE
CAMBRIDGE, MA 02138

BOOK REVIEW:
WHERE HAVE ALL THE WILDFLOWERS GONE?

Mohlenbrock, R. H. 1983. Where have all the wildflowers gone?
Macmillan Publ. Co., New York, 239 pp. (price $15.95).

This book is a popular guide to selected threatened and
endangered wildflowers of the United States. Mohlenbrock divides
the country into seven geographical areas and discusses the rare
plants that occur within each area. There are usually 15-20 species
treated in each region; each is allotted a brief, simple description
and a page or two of text. In all, 120 plants are discussed, with 40
illustrated by color photographs and 80 by excellent line drawings.
The latter were prepared by the author’s son, Mark Mohlenbrock,
himself a plant taxomomist and accomplished botanical illustrator.

The first chapter, entitled “The Disappearance of Thismia”,
unlike the others, is not a regional account of rare plants. Instead, it
details the story of the remarkable plant Thismia americana, found
only twice in a sand prairie on the south side of Chicago and never
again seen. The second half of the chapter summarizes the
important steps in the history of endangered plant protection at the
national level in the United States. As such, it is a good introduction
to the nationally endangered plants that follow in later chapters.

Chapters 2-8 are regional accounts of endangered plants.
Informative, interesting and highly readable, they do an excellent
job of telling the fascinating stories behind these plants. The more
technical information is omitted, and the text emphasizes the
humanistic aspects. One learns about many famous past and present
American botanists, and how they contributed to our knowledge of
these rare plants. The poem about Furbish’s Lousewort, written by
Charles Osgood of CBS, is particularly enjoyable. There are many
interesting anecdotes of Mohlenbrock’s personal experience with
the plants. They reveal the problems and pleasures of field botany
and the genuine excitement that occurs when something rare and
beautiful is found for the first time.

The front flap of the dust jacket promotes the book as a field
guide, but I believe that it is primarily a book to be read and enjoyed
indoors. Its use in the field to distinguish endangered plants from

Vol. 86, No. 845 including pages 1-119 was issued April 6, 1984.

237

238 Rhodora [Vol. 86

others is rather limited. Mohlenbrock has numerous qualifications
and achievements which rank him as an eminent conservationist.
The present book is a culmination of these conservation interests at
the national level.

ROBBIN C. MORAN

ILLINOIS NATURAL HISTORY SURVEY
607 E. PEABODY DR.

CHAMPAIGN, IL 61820

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inches; be sure that your illustrations are proportioned to reduce
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and a list of Key Words should be supplied at the beginning of each
paper submitted, except for a very short article or note.

RHODORA April 1984 Vol. 86, No. 846
CONTENTS

A new species of Phyllanthus (Euphorbiaceae) from the Cayman Islands
Grady L. Webster and George R. Proctor aa ee ee > See

A new, widespread species of Chaptalia (Asteraceae: Mutisieae) from Mexico
Guy L. Nesom eh a Ae. ay Gy oe ee ee ee ee

Notes on Croomia pauciflora (Stemonaceae)
R. David Whetstone . F : . , anes : ws ae

Distribution and ecological characteristics of Ironwood, Ostrya virginiana
(Miller) K. Koch, in northeastern Nova Scotia
K. N. H. Greenidge : : ie Pw) ee a ee ee

Annotated list of Minnesota Carices, with phytogeographical and ecological
notes
Gerald A. Wheeler and Gerald B. Ownbey : ee Sa at er ee

NEW ENGLAND NOTES
Note on the first station of Dentaria laciniata Muhl. (Brassicaceae) reported
from Maine

Sally C. Rooney gee Ses Ge” es

Two new localities for Medeolaria farlowii in New England
Donald H. Pfister a ee eee eT |, ae eK
Book Review: Where have all the wildflowers gone? Lad eS ee

NEBC Research Award Notice
126

Instructions to contributors to Rhodora ' ' ; P . inside back cover

ee ee nn

‘Hovdova

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 86 July 1984 No. 847

Che New England Botanical Club, Iuc.

Botanical Museum, Oxford Street, Cambridge, Massachusetts 02138

Conducted and published for the Club, by
NORTON H. NICKERSON, Editor-in-Chief

Associate Editors

A. LINN BOGLE GARRETT E. CROW
WILLIAM D. COUNTRYMAN RICHARD A. FRALICK
GERALD J. GASTONY NORTON G. MILLER

ROBERT T. WILCE

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Cover illustration

Ledum groenlandicum Oeder, Labrador tea, reaches its southeastern distribu-
tional limit in Concord, Mass. It was first collected by Thoreau in 1858, subsequently
regarded as extirpated by Richard Eaton in 1974, and rediscovered by Ray Angelo in
1978. Angelo has since found it in two more Concord locations.

Original artwork by Josephine Ewing.

Mbodora

(ISSN 0035-4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB
Vol. 86 July 1984 No. 847

TAXONOMY OF ARNICA (COMPOSITAE)
SUBGENUS AUSTROMONTANA'!

STEVEN J. WOLF AND KEITH E. DENFORD

ABSTRACT

Nine species, with no infraspecific taxa, are recognized in Arnica subgenus
Austromontana, a group primarily restricted to montane western North America.
Previous treatments, which have variously recognized several infraspecific taxa,
radiate and discoid sections, and two subgenera for these nine species, are viewed as
artificial. Chromosome counts, all based on x = 19, are reported for 74 populations
representing seven of the species. A discussion of comparative features, phylogenetic
relationships, distribution maps, descriptions, synonymies and a key to the species
are presented.

Key Words: Arnica subgenus Austromontana, taxonomy, chromosome counts,
montane western North America

Arnica L. is a circumboreal, predominantly montane genus of
about 32 species, most of which are confined to western North
America. The genus, with its yellow florets, opposite leaves,
capillary pappus and basic chromosome number of x = 19, is well
defined. However, its tribal position within the Compositae has
been the subject of considerable debate (Nordenstam, 1977;
Rokinson, 1981).

Members of the genus are extremely variable and numerous
nomenclatural variants have been recognized. Apomixis has long
been reported in Arnica (Afzelius, 1936) while Cronquist (1955)
suggested that much of the perplexing morphological variability

'This paper is based on a dissertation submitted to the Department of Botany,
University of Alberta, by S. J. W. in partial fulfillment of the requirements for the
degree of Doctor of Philosophy.

239

240 Rhodora [Vol. 86

within the genus may be due to microspecies formation via
apomixis. Cytologically, the genus is very diverse, with all ploidy
levels from diploid to octoploid being reported (Wolf, 1980). Barker
(1966) established that polyploidy within the genus always indicates
apomixis and Straley (1980) confirmed this observation for the
subgenus Austromontana.

Members of subgenus Austromontana are distinguished from the
other four subgenera of Arnica by their turbinate to campanulate
heads, white, barbellate pappus and broad leaves. The subgenus, as
circumscribed in the present study, consists of nine species
distributed in montane to alpine habitats from central Alaska
through southern California and northern New Mexico, with a few
disjunct populations of A. cordifolia occurring in Ontario and
Michigan. Four of the nine species are relatively rare and restricted
to the Klamath region of southwestern Oregon and northwestern
California.

The only worldwide monograph of the genus Arnica (Maguire,
1943) was based largely on herbarium material; only two popula-
tions were available for study of some taxa, and the consequences of
apomictic reproduction in the genus were largely unknown at the
time. More recent taxonomic treatments of the genus in North
America (Ediger and Barkley, 1978) and subgenus Austromontana
(Straley, 1980) are based largely on Maguire (1943), with added
observations on reproductive biology. The present revision of
subgenus Austromontana incorporates observations from morphol-
ogy, cytology, geography and flavonoid chemistry (Wolf, 1980;
1981; Wolf and Denford, 1983; 1984a; 1984b). More than 13,000
herbarium specimens were examined and over 250 populations were
studied in the field.

TAXONOMIC HISTORY

The concept of subgenera in Arnica originated with Maguire
(1943) when he described five subgenera: Andropurpurea, Arctica,
Austromontana, Chamissonis, and Montana. Maguire (1943) rec-
ognized thirteen species and six subspecies in subgenus Austro-
montana and placed the seven radiate species and two subspecies in
section Eulatifoliae and six discoid species and four subspecies in
section Eradiatae. Maguire (1947) later recognized four varieties of
A. cordifolia subsp. genuina: vars. cordifolia, macrophylla, pumila

1984] Wolf & Denford — Arnica 241

and humilis. Cronquist (1955), in his treatment of Arnica for the
Flora of the Pacific Northwest, included A. grayi and A. parviflora
in the new combination A. discoidea var. eradiata (A. Gray)
Cronquist and treated A. gracilis as a variety of A. Jatifolia. Shortly
thereafter, Cronquist (1958) proposed the name A. discoida var.
alata (Rydb.) Cronquist for A. alata Rydb.

In their revision of Arnica for the North American Flora, Ediger
and Barkley (1978) essentially adopted Maguire’s (1943) treatment
of subgenus Austromontana, recognizing nine species and treating
his (Maguire’s) subspecies as varieties. Exceptions include their
acceptance of Cronquist’s (1955, 1958) treatment of A. discoidea
with three varieties (var. discoidea, var. alata, and var. eradiata) and
his recognition of A. gracilis as a variety of A. latifolia. In addition,
Ediger and Barkley considered A. paniculata a possible hybrid
between A. cordifolia and A. parryi A. Gray but did not give it
formal taxonomic recognition.

In a recent systematic study, Straley (1980) retained Maguire’s
(1943) two sections and recognized seven species in a revised
subgenus Austromontana and included A. venosa and A. viscosa in
the newly erected subgenus Calarnica.

In the present revision of subgenus Austromontana nine of
Maguire’s (1943) species are recognized; however, in light of
considerable evidence, particularly with respect to flavonoid chem-
istry and the morphological consequences of apomixis, we recognize
no infraspecific taxa and reject Maguire’s sections and Straley’s
(1980) new subgenus as artificial.

CHROMOSOME NUMBERS

Chromosome numbers from the species of subgenus Austro-
montana were determined for either mitotic or meiotic material
using techniques previously outlined (Wolf, 1980).

Arnica chromosome numbers, including many species of Austro-
montana, published prior to 1980 have recently been reviewed
(Wolf, 1980). Consequently, the following discussions will be largely
limited to comments on more recent data which, in addition to the
seventy-four new counts presented in Table I, incorporates reports
by Love and Léve (1981) and Straley (1980, 1982). As previously
noted, the basic chromosome number of the genus Arnica is x = 19
and all new counts within subgenus Austromontana conform to this

242 Rhodora [Vol. 86

basic number. In the present study a new count of 2n = 57 for A.
discoidea is reported (Table 1).

Arnica cernua, A. venosa and A. viscosa are all uniformly
diploid, while A. nevadensis is uniformly tetraploid. A. spathulata
and A. Jatifolia are largely diploid; however, both have a few
tetraploid populations. A. gracilis has both triploid and tetraploid
chromosome races. A. discoidea has diploid, triploid and tetraploid
races; however, the diploids are largely restricted to the Klamath
region while the polyploids occur at the northern, eastern and
southern limits of its distribution.

Arnica cordifolia, with five chromosome races, has previously
been recognized as a mature polyploid complex (sensu Stebbins,
1971) (Wolf, 1980). Subsequent chromosome sampling, now total-
ling more than 150 populations from throughout its entire distribu-
tion, has confirmed this observation. Tetraploids are widespread,
occurring throughout the entire species’ range, triploids are found
largely in the front ranges of the Rocky Mountains, and diploids,
pentaploids and hexaploids are rare and of scattered occurrence.
Diploids are largely restricted to northeastern Oregon and southern
Yukon Territory, pentaploids are restricted to Colorado and
hexaploids have been found in Alberta and central Washington.

MORPHOLOGY AND TAXONOMIC CRITERIA

In a discussion of the morphology of the genus Arnica, Maguire
(1943) noted that there are some sharply defined species and a large
number of other loosely-knit polymorphic ones. Within subgenus
Austromontana most taxa fit into the latter category. Only A.
cernua, A. viscosa and to some extent A. venosa are morpho-
logically well defined. The remaining six species are highly
polymorphic and show a wide range of morphological intergrada-
tion. In addition, superimposed upon this interspecific variability is
a great deal of infraspecific variability. As Gustafsson (1947) and
later Cronquist (1955) suggested, apomixis and polyploidy are
probably largely responsible for this morphological variability
within the genus. That A. venosa, A. viscosa and A. cernua are the
only well defined species in subgenus Awustromontana is not
surprising since they are the only entirely sexual, diploid taxa in the
subgenus. Although the several species of subgenus Austromontana
are highly variable, a combination of several morphological features

1984] Wolf & Denford — Arnica 243

in conjunction with ecological and distributional data are sufficient
to distinguish among them. The following discussions detail
morphological variation and characters of taxonomic significance
within subgenus Austromontana.

HABIT. All species of subgenus Austromontana are herbaceous
perennials. With respect to underground parts, both Arnica venosa
and A. viscosa possess a thick woody caudex at or below the soil
surface, which gives rise to several flowering shoots. The remaining
species possess slender rhizomes which give rise to both flowering
shoots and numerous sterile basal rosettes of leaves (innovations).
Both rhizomes and caudices are frequently covered with dark scales
and old leaf bases which give them a thickened appearance. In A.
gracilis the tip of the rhizome may have several branches, thus
producing a many-stemmed crown (approximate crown) with the
flowering shoots appearing in dense clumps.

STEMS. Stems in the species of subgenus Austromontana range
from simple, as in Arnica cordifolia and A. cernua to highly
branched. A. discoidea and A. venosa are sometimes several
branched above the middle while A. viscosa is several-branched
throughout, sometimes so much that it resembles a small bush. In
addition, virtually all species are much more branched when they
occur on more exposed or disturbed sites. Stem height varies from
10-30 cm in the relatively small A. cernua and A. gracilis, up to
60-70 cm in robust specimens of A. discoidea and A. cordifolia. In
general, stem pubescence resembles leaf pubescence, although it is
usually denser above and reduced below.

VESTITURE. Virtually all parts of all species of subgenus Austro-
montana are to some degree pubescent. Both long (1-2 mm) and
short (0.1-0.2 mm) septate glandular and non-glandular hairs occur
in the subgenus. The long stipitate glandular hairs have stalks 6-8
cells long while the short stalks are 2-3 cells long. Both are two cells
thick. Arnica latifolia generally lacks glandular pubescence, A.
viscosa has only the long type, A. gracilis has only the short type
and the remaining species have both long and short. In A. viscosa
the entire plant is densely covered with long stipitate glandular
hairs, so much so that it feels slimy to the touch.

Non-glandular hairs are one cell thick and either long (1-3 mm)
or short (0.3-0.5 mm).

Table 1. New chromosome counts in Arnica subgenus Austromontana.

Taxon n= 2n

Location and voucher***

A. cernua 19
A. cordifolia 19

57

38

76

USA: CA: Humboldt Co.: Horse Mt., W471/.

USA: OR: Josephine Co.: Babyfoot Lake, W464.

CAN: YT: Canol Rd., km 16, W507; Squanga Lk, W505.

USA: OR: Wallowa Co.: Minam Park, W449; Hwy 3, 8 km S Wa. state line, W447.

CAN: YT: Skagway Rd., 19 km S. Carcross, W499*, USA: MT: Flathead Co.: 8 km E
Bigfork, W496*; Lincoln Co.: 23 km W. Libby, W444*; Missoula Co.: 34 km N Seeley Lk,
W495*.

CAN: BC: Skagway Rd., 65 km S Carcross, W500; Hwy 93: 26km N Jct Hwy 3, W441; 11 km
S Jet Hwy 3, W442; Hwy 3, 15 km E Osoyoos, B80055; Hwy between Pavillion and Clinton,
B80060. YT: Canol Rd., km 118, W508; Skagway Rd., 7 km S Carcross, W498.

USA: CA: Nevada Co.: W side Donner Lk, W476. ID: Custer Co.: 15 km W Challis, W486.
MT: Lincoln Co.: 25 km S Eureka, W443. Missoula Co.: 10 km E Bonner, W493; 22 km W
Lolo City, W491. NV: White Pine Co.: Wheeler Peak, W480. OR: Grant Co.: 72 km S
Ukiah, W452; Umatilla Co.: 14 km S Ukiah, W45/; Union Co.: 40 km E Ukiah, W450;
Wallowa Co.: 25 km N Enterprise, W448; Wheeler Co.: Ochoco Summit, W453. UT: Beaver
Co.: 23 km E Beaver, W482; Cache Co.: Beaver Mt., W484; Iron Co.: 17 km E Cedar City,
W481; Rich Co.: Bear Lake Summit, W485; Utah Co.: Mt. Timpanogos, W483. WA: Asotin
Co.: Field Springs Park, W446; Spokane Co.: Mt. Spokane, W445.

USA: CA: Plumas Co.: Gold Lake, W478. ID: Lemhi Co.: 5kmS Gibbonsville, W487. MT:
Missoula Co.: Marshall Ski Area, W492. OR: Klamath Co.: Parker Mt. Summit, W454,
UT: Duchense Co.: Hwy 33 at S border of Ashley Nat. Forest, W396; San Pete Co.: Jct
Spring City-Skyline Drive Rd., W393. WA: Okanogan Co.: 23 km W Twisp, W509. WY:
Teton Co.: Teton Nat. Park, Hidden Falls, W429.

bre

vlopoyy

98 ‘10A]

A,

A,

. discoidea

. latifolia

. spathulata

venosa

viscosa

19

38

19

38

57**

38

76

USA: CA: Humboldt Co.: 6 km W Briceland, W472; Mendocino Co.: 14 km W Boonesville,
W474; San Mateo Co.: Kings Mt., W475; Siskiyou Co.: Baldy Mt., W466; Trinity Co.: 3 km
E Burnt Ranch, W470.

USA: CA: Plumas Co.: Gold Lake, W477*.

USA: CA: Lake Co.: 13 km E Lower Lake, W473; Santa Barbara Co.: La Cumbre Peak,
W515.

CAN: ALT: Banff Nat. Park, Moraine Lake, W5/3.

USA: OR: Curry Co.: Iron Mt., W457.

USA: AK: Hatcher Pass, W503. WA: Chelan Co.: Swauk Pass, W367; Kittitas Co.:
Snoqualmie Summit, W5/0.

USA: CA: Siskiyou Co.: Bolan Lake, W465; 13 km W Etna, W467. ID: Idaho Co.: Lolo Pass,
W490; Lemhi Co.: 9 km S Lost Trail Pass, W488. MT: Missoula Co.: Seeley Lake, W494; 27
km W Lolo City, W489.

CAN: ALT: Waterton Nat. Park, Carthew Mt., W440.

USA: CA: Del Norte Co.: French Hill, W458; 10 km N Gasquet, W459. OR: Josephine Co.:
Hugo, W455; Merlin, W456; Selma, W463; Store Gulch Guard Station, W462.

USA: OR: Josephine Co.: 17 km N Patrick, W460; 20 km N Patrick, W461.

USA: CA: Shasta Co.: Hwy 5, 3 km S Gibson turnoff, W468; Shasta Bally Rd., S Brandy
Creek, W469.

USA: OR: Klamath Co.: Crater Lake Nat. Park, Garfield Peak, W5//.

*

** New count for this taxon.

Count based on meiotic cells.

*** Abbreviations of collectors: B = J. F. Bain; W = S. J. Wolf. Vouchers at ALTA.

[P8361

Duy — Plojurq F FIONA

Ss £6

246 Rhodora [Vol. 86

LEAVES. Although highly variable, characters of the leaves are
generally the most reliable in distinguishing the species of subgenus
Austromontana. Characters considered important in the present
study are leaf number, shape, margin, position and petiole width.
Within the subgenus there is a general evolutionary trend from few,
broad, coarsely dentate, narrowly petiolate leaves mostly below the
mid-stem towards many, narrower, entire margined, sessile, evenly
distributed leaves.

The number of cauline leaves varies from 2 to 4 pairs in such
species as Arnica cordifolia and A. latifolia to more than 25 in A.
viscosa. Species intermediate between these extremes include A.
spathulata (3-5), A. discoidea (3-5) and A. venosa (6-10). Addi-
tionally, in the few-leaved species the leaves tend to be mostly below
mid-stem while, in contrast, in the many-leaved Species they are
evenly distributed along both the stems and branches. This appears
to be a trend within the genus Arnica in general (Maguire, 1943).

Leaf shape varies from broadly cordate in Arnica cordifolia to
broadly lanceolate in A. discoidea. Intermediate conditions include:
ovate to ovate-elliptic in A. cernua, A. latifolia, A. venosa and A.
nevandenis,; spathulate in A. spathulata and ovate-oblong in A.
viscosa. Leaf margins vary from coarsely dentate in the primitive A.
cordifolia to entire in its derivative A. nevadensis and the highly
advanced A. viscosa. Intermediate forms include crenate in A.
cernua and serrate to serrate-dentate in the remaining species.

Within the subgenus Austromontana there is a distinct evolu-
tionary trend in petiole width from narrow to broad to completely
sessile leaves, with the latter being considered most advanced. Five
of the nine species have narrowly petiolate leaves (Arnica cordifolia,
A. cernua, A. discoidea, A. nevadensis and A. gracilis), three species
have sessile leaves (A. latifolia, A. venosa, and A. viscosa) while A.
spathulata, with broadly winged petioles, represents the inter-
mediate condition. Occasionally, the lower leaves of A. Jatifolia are
short petiolate while the lower leaves of A. cernua and A. gracilis
are infrequently narrowly winged. In addition, in all of the
rhizomatous species, including the sessile-leaved species, the leaves
of the innovations are narrowly petiolate. Leaves of innovations are
otherwise similar to the cauline leaves. Additionally, in all species of
the subgenus the upper leaves are often reduced, bract-like and not
infrequently sub-opposite.

1984] Wolf & Denford — Arnica 247

INFLORESCENCE. Within subgenus Austromontana several charac-
ters of the inflorescence are considered important in both delimita-
tion of the species and determination of evolutionary relationships.
Among these characters are head type (radiate vs. discoid), number
and shape, phyllary shape and characters of the pappus. With the
exception of Arnica parryi (subgenus Chamissonis), no discoid
species of Arnica occur outside the subgenus Austromontana.
Within the latter, A. viscosa, A. venosa, A. spathulata and A.
discoidea are discoid while the remaining are radiate. In addition, in
A. discoidea some marginal disc florets may be ampliate (elongated
and ray-like), resulting in some pressed specimens of this species
being confused with A. cordifolia. Results of the present study
support Maguire’s (1943) observation that the discoid condition
represents the advanced state in subgenus Austromontana. Indeed,
Cronquist (1977) considered the discoid condition to be derived
within the Compositae in general.

The number of ray florets varies from 5 to 15 while the number of
disc florets varies from 10 to 90. Within the subgenus there is a
general trend of reduction in the number of disc florets, particularly
in the discoid species. Arnica cordifolia and A. latifolia, with up to
90 disc florets, represent the primitive condition while A. viscosa,
with as few as 10 florets, is viewed as highly advanced.

Cronquist (1977) considered yellow-colored corollas primitive
within the Compositae in general. With the exception of Arnica
viscosa, which has cream-colored florets, all other Arnica species
possess yellow corollas. Clearly, the cream-colored florets of A.
viscosa represent the derived condition in subgenus Austromontana.

Within subgenus Austromontana heads may occur singly as in
Arnica nevadensis, A. cordifolia, and A. cernua or more commonly
3 to several heads are arranged in a corymbose inflorescence.
Species with several heads include A. viscosa (10-20), A. gracilis
(5-15) and A. discoidea (3-10, or up to 30). Maguire (1943)
considered solitary heads primitive within Arnica in general and
Cronquist (1977) noted a similar trend towards increasing head
number in the Compositae in general. In the present study, the
solitary condition is considered primitive in subgenus Austro-
montana while, in contrast, an increased number of heads is
interpreted as an advanced state.

The pappus in the genus Arnica is composed of a ring of 25-70
capillary bristles of varying length. Based on seta length, Maguire

248 Rhodora [Vol. 86

(1943) recognized three conditions within the genus: barbellate
(0.1-0.2 mm), subplumose (0.2-0.35 mm) and plumose (0.35-0.6
mm). Also, within the genus pappus color varies from white to
tawny. Maguire (1943) considered the white, barbellate pappus
primitive, while the tawny, plumose pappus was considered
advanced. Most species of subgenus Austromontana have retained
the primitive white, barbellate pappus. However, in 4. cernua, A.
discoidea, A. nevadensis and A. viscosa the pappus is infrequently
subplumose. Additionally, in both A. nevadensis and A. viscosa the
pappus is sometimes slightly tawny.

Although somewhat variable, characters of the involucral bracts
are often quite helpful in delimiting species. Phyllary shape varies
from ovate-lanceolate (Arnica cordifolia, A. cernua, A. discoidea,
A. gracilis and A. venosa) to lanceolate (A. latifolia and A. viscosa).
A. nevadensis has distinctive oblanceolate phyllaries. Phyllary
vestiture, like that of the leaves, consists of various combinations of
short and long stipitate glandular and non-glandular hairs. The
vestiture is always densest and longest at the bases of bracts, at the
point of their attachment to the peduncle.

ACHENES. Achene color in subgenus Austromontana is mostly gray
with the exception of Arnica gracilis (black), A. spathulata (black),
and A. Jatifolia (brown). Length varies from 4.5-10 mm, with A.
gracilis and A. viscosa being the shortest and A. cordifolia the
longest. Achene width is almost uniformly | mm with the exception
of A. venosa (1.5 mm). Achene vestiture, which is quite useful in
delimiting species, consists of various combinations of both short
and long glandular hairs and duplex (forked) hairs. For example, A.
viscosa has only long stipitate glandular hairs, A. gracilis has a few
short glandular hairs, A. venosa has an abundance of duplex hairs
and A. discoidea has both duplex and glandular hairs.

PHYLOGENY AND PHYTOGEOGRAPHY

Maguire (1943) considered the genus Arnica to have arisen in
Arctic or sub-Arctic western North America from where it spread
eastward, westward and southward. Hultén (1937) listed 22 species
of Arnica (some of which were not recognized by Maguire, 1943)
that are of probable northwestern North American origin. As most
species of the genus are adapted to cool montane habitats and 25 of
the 32 species recognized by Maguire (1943) are largely confined to

1984] Wolf & Denford — Arnica 249

northwestern North America this conclusion seems valid. Addi-
tional evidence to support this hypothesis is the presence of many
relictual diploid races of otherwise widespread Arnica polyploid
complexes in the unglaciated Alaska-Yukon region (Wolf, 1980).
Phytogeographical evidence suggests that the genus Arnica is
relatively old, and was part of the Arcto-Tertiary flora. The east-
west disjunct distribution of A. /ouiseana Farr and A. lonchophylla
D. C. Eaton, the circumpolar distribution of A. angustifolia Vahl
and the close relationship between A. montana of Europe and 4A.
acaulis of the eastern United States add support to this theory.

Raven and Axelrod (1978) included the genus Arnica within a
group of genera that are well-developed in California, yet wide-
spread elsewhere. They suggested that this recurrent pattern in the
Californian flora is the result of spreading aridity from Upper
Tertiary times in the western United States which culminated in the
development of a full Mediterranean climate in the late Quaternary.
A striking example of this distribution pattern occurs within
subgenus Austromontana which contains both widespread species
(e.g., A. cordifolia and A. latifolia) and several endemics (e.g., A.
cernua, A. venosa and A. viscosa). In fact, the discoid species of
Austromontana are confined almost entirely to the northern areas
of the California Floristic Province, particularly within the Klamath
region of southwestern Oregon and northwestern California. The
occurrence of many endemics in the Klamath region and its
significance in the evolution and development of the flora of the
western states, particularly California, have been discussed by
Whittaker (1960, 1961). Owing to its geological history, equable
climate and diversity of parent soils, the Klamath region contains
many endemics that probably represent both remnants of the Arcto-
Tertiary flora and more recently derived taxa (Whittaker, 1961). In
the case of subgenus Austromontana, the narrow endemics appear
to be recently derived from the more widespread A. cordifolia and
A. latifolia (Wolf & Denford, 1984b).

Speciation within the subgenus Austromontana has been ac-
companied by a number of ecological, morphological and chemical
changes, some of which include: a shift from mesic to xeric habitats;
temporal isolation of flowering periods; specialization to particular
substrates; the replacement of solitary, radiate heads by more
numerous, narrower, discoid heads with reduced numbers of disc
florets; evolution from narrowly petiolate to sessile leaves; the

250 Rhodora [Vol. 86

replacement of simple flavonol glycosides by more complex
methylated flavone aglycones, and a secondary loss of flavonoids in
several rare and geographically restricted species (Wolf, 1981; Wolf
and Denford, 1983; 1984b). Additional factors contributing to
speciation in the subgenus include hybridization, polyploidy and
changes in reproductive systems (Wolf, 1980; Wolf and Denford,
1984a; Barker, 1967; Straley, 1980).

Maguire (1943) considered Arnica cordifolia to represent the
ancestral species in subgenus Austromontana which gave rise to the
rest of the subgenus. Results of the present investigation support
this hypothesis. With respect to morphology, A. cordifolia exhibits
most features considered primitive within the genus Arnica in
general and the subgenus Austromontana in particular. Significant
primitive features of A. cordifolia include: solitary, broad heads
with white, barbellate pappus; unbranched stems with few, narrowly
petiolate, broad, dentate leaves and dark gray achenes. In addition,
the flavonoid profile of A. cordifolia, which lacks complex
methylation, is relatively primitive (Wolf and Denford, 1983).
Additionally, the mesic-montane habitat of A. cordifolia represents
the ancestral condition within the genus Arnica in general (Maguire,
1943). Indeed, within subgenus Austromontana there is a distinct
evolutionary trend from mesic-montane habitats towards drier
habitats at both lower and higher elevations.

Phytogeographical, cytological and chemical evidence suggests
that Arnica cordifolia was probably a relatively widespread diploid
species prior to the Pleistocene. Hultén (1937) included A. cordifolia
in a group of species widespread prior to the Pleistocene, but whose
ranges were interrupted by glaciation. He hypothesized that these
species survived the Pleistocene glaciations south of the ice and in
unglaciated areas of the Yukon and Alaska. Upon retreat of the ice,
these populations were rejoined to form a continuous distribution.
This hypothesis is supported by the fact that the flavonoid profiles
of populations north of the maximum limits of Pleistocene
glaciation differ from those to the south (Wolf and Denford, 1983).
Cytological evidence also supports the hypothesis that A. cordifolia
was probably a relatively widespread diploid species prior to the
Pleistocene. Barker (1966) noted that within the genus Arnica
diploids were probably more widespread in the past and that
polyploidy in the genus is a relatively recent, i.e., inter- or post-
glacial phenomenon. Stebbins (1971) noted that most mature

1984] Wolf & Denford — Arnica 251

polyploid complexes such as A. cordifolia are of Pliocene or
Pleistocene origin. The occurrence of diploid populations of A.
cordifolia in the unglaciated Yukon as well as south of the limits of
glaciation (in Oregon) suggests a former, much wider distribution of
diploid populations.

The geographical distribution, morphology, cytology and flavon-
oid chemistry of Arnica latifolia suggests that it is probably a pre-
Pleistocene derivative of A. cordifolia. A. latifolia occurs through-
out much of the range of A. cordifolia but in moister habitats at
higher elevations. Morphologically the two species are very similar
and are often difficult to distinguish on herbarium sheets. A.
latifolia and A. cordifolia share several primitive features including:
radiate heads with white, barbellate pappus; relatively few, broad
leaves; innovations; and numerous, yellow disc florets. However, A.
latifolia has several advanced features, including sessile leaves,
narrow heads and phyllaries, and brown achenes. Its flavonoid
profile, which consists largely of flavonol glycosides, is relatively
primitive and similar to that of A. cordifolia (Wolf and Denford,
1984b). However, the two species, and their putative derivatives,
differ largely with respect to the replacement of quercetin 6-
methoxy-3-0-glucoside in A. Jatifolia for kaempferol 6-methoxy-3-
0-glucoside in A. cordifolia. The largely diploid condition of A.
latifolia suggests a pre-Pleistocene divergence from A. cordifolia
prior to the elimination of the diploid level in the latter. A. /atifolia
may have been an ecological race of an archetypal “cordifolia”
adapted to moister conditions, which became established after the
advent of apomixis in the latter. The fact that A. cordifolia and A.
latifolia probably hybridized to produce A. gracilis (Wolf and
Denford, 1984a) also suggests a close relationship between the two
species.

As previously noted (Wolf and Denford, 1984a), Arnica gracilis is
probably a hybrid between A. Jatifolia and A. cordifolia. The
flavonoid profile of A. gracilis, which consists of fourteen com-
pounds, is essentially a summation of the two parental profiles.
Although A. gracilis is somewhat intermediate between A. J/atifolia
and A. cordifolia, it has several distinctive and advanced features of
its own. These characters include a much more branched habit;
narrower leaves; more numerous and smaller heads with a reduced
number of disc florets; black, glandular achenes and the dry, alpine
habitat. Since A. cordifolia is an apomictic, polyploid complex

252 Rhodora [Vol. 86

(Wolf, 1980) with no known sexual populations and A. Jatifolia is
largely sexual and diploid, A. gracilis was probably formed prior to
or during the Pleistocene, before the elimination of the sexual
condition in A. cordifolia (Wolf and Denford, 1984a).

As previously noted (Wolf and Denford, 1984b), there is little
doubt that Arnica discoidea has been derived from A. cordifolia.
Morphologically the two species are quite similar, almost to the
extent that A. discoidea appears to be little more than a rayless A.
cordifolia. However, A. discoidea is readily distinguished by several
advanced features including: more numerous, narrower, discoid
heads with a barbellate to subplumose pappus; more numerous,
narrower leaves and glandular achenes. Additionally, A. discoidea
and A. cordifolia are ecologically quite distinct. A. cordifolia is
adapted to cool, mesic, montane habitats and is generally quite rare
west of the Cascades. In contrast, A. discoidea occurs in hotter and
drier habitats west of the Sierras and Cascades. In both instances
where the two species have been observed in close association, in the
Sierran foothills and in the central Cascades, A. cordifolia had
flowered and set seed well over a month prior to the flowering of A.
discoidea. Such early flowering of A. cordifolia is no doubt a means
of escaping the relatively warm-dry summer conditions of these
areas.

Cytological and chemical evidence suggests that Arnica discoidea
was derived from A. cordifolia in the Klamath region. The
flavonoid profiles of diploid Klamath populations of A. discoidea
are strikingly similar to that of A. cordifolia while, in contrast, the
polyploid populations outside the Klamath region have more
advanced compounds and reduced flavonoid profiles (Wolf and
Denford, 1984b). This suggests that A. discoidea has been derived
from ancient diploid Klamath populations of A. cordifolia and that
migration outward from this area has been accompanied by
polyploidization and a change in flavonoid chemistry.

The Klamath region endemic Arnica spathulata has probably
been derived from A. discoidea via saltational speciation into
serpentine areas. Morphologically the two species are very similar,
differing largely in leaf and petiole shape, and degree of pubescence.
However, A. spathulata is readily distinguished by several advanced
features including: narrow phyllaries; narrower, broadly petiolate
leaves; and black achenes. Ecological distinctions between the two
species are also readily apparent. A. spathulata occurs at lower

1984] Wolf & Denford — Arnica 253

elevations and in drier habitats on serpentine soils. Additionally, the
flavonoid profile of A. spathulata, which consists of eight com-
pounds, is a subset of diploid Klamath populations of A. discoidea
(Wolf and Denford, 1984b).

Whittaker (1961) noted that the diversity of soil types in the
Klamath region has contributed greatly to the formation of many
local endemics. Kruckeberg (1954, 1969) suggested that serpentine
endemism results from an adaptation to serpentine followed by
biotype depletion and the development of isolated populations into
local endemics. Lewis (1962) noted that many serpentine endemics
originate by saltation speciation in marginal populations. As Raven
and Axel (1978) noted, marginal populations often occur in edaphic
situations that are unique for the species as a whole. What these
ideas suggest is that Arnica spathulata may have initially diverged
from marginal populations of A. discoidea that gradually adapted
to, and later became restricted to serpentine soils. The subsequent
restriction of these populations to local isolated areas may have
resulted in a gradual depletion of their flavonoid profiles. The fact
that A. discoidea and A. spathulata are still morphologically
somewhat similar suggests a fairly recent derivation of the latter.
Indeed, Raven and Axelrod (1978) suggested that most herbaceous,
localized serpentine endemics of the California Floristic Province
(such as A. spathulata) originated in late Pleistocene or more recent
time. A. spathulata is therefore probably a relatively recent
derivative of A. discoidea that gradually became adapted to, and
later restricted to serpentine areas.

The rare Klamath region endemic Arnica venosa is probably a
very recent derivation of A. discoidea. Typical populations of A.
venosa differ from A. discoidea by several advanced features
including: numerous sessile leaves, a woody caudex and a lack of
innovations. A. venosa is also distinguished by its strongly
reticulate-veined leaves, more leafy and branched habit and
preference for hotter, drier disturbed habitats. However, some
specimens of A. venosa with thin, slightly woody rhizomes and few,
broadly petiolate, weakly veined leaves tend to resemble 4A.
discoidea and suggest the two species are related. A. venosa 1s
diploid and its flavonoid profile, which consists of six compounds,
is merely a subset of the profile of A. discoidea. Since A. venosa and
A. discoidea occur sympatrically, have several flavonoids in
common and are sometimes morphologically similar, it seems likely

254 Rhodora [Vol. 86

that the former is derived from the latter. A. venosa is restricted to
the foothills of Shasta County, California in the hottest and driest
habitat of any Arnica species known. This represents considerable
divergence from the ancestral cool-montane habitat characteristic of
the genus Arnica. The very localized distribution of A. venosa, its
preference for recently disturbed habitats and derived ecology
Suggest it is very recently derived, possibly during the post-glacial
hypsithermal of 9,000 to 2,600 years ago (Flint, 1957).

Maguire (1943) and Straley (1980) considered the rare serpentine,
Klamath endemic A. cernua to be derived from A. latifolia.
However, both A. cernua and A. cordifolia share several primitive
morphological features including: long, narrow petioles; broad
solitary heads with pilose and glandular, ovate-lanceolate phyl-
laries; and dark gray achenes. In contrast, A. Jatifolia has sessile
leaves; several narrow heads with lanceolate, sparsely pubescent
phyllaries and brown achenes.

Raven and Axelrod (1978) suggested that Arnica cernua probably
evolved from a more widespread species, in late Pleistocene or
Recent times via saltation speciation in marginal populations that
gradually became adapted to and later restricted to serpentine soils.
It seems likely that A. cernua has been derived from the more
widespread A. cordifolia prior to the elimination of the diploid
condition in the latter.

Arnica nevadensis has been derived, at least in part, from A.
cordifolia. A. nevadensis is an extremely variable species which
sometimes resembles little more than a high altitude ecotype of A.
cordifolia. In fact, in the past, many high altitude Rocky Mountain
populations of A. cordifolia have been erroneously identified as A.
nevadensis. A. cordifolia and A. nevadensis share several primitive
features including: solitary, radiate heads and simple stems with
few, relatively broad leaves. Additionally, the flavonoid profile of
A. nevadensis is most similar to that of A. cordifolia (Wolf and
Denford, 1984b). However, A. nevadensis is readily distinguished
by its often tawny, subplumose pappus, entire leaves, oblanceolate
phyllaries and relatively high altitude, exposed habitat. The extreme
morphological variability of A. nevandensis, its tetraploid condi-
tion, apomictic reproductive system and resemblance to A. cordi-
folia suggests that either it is the result of introgression between the
latter and another Arnica species or it is a high altitude microspecies

1984] Wolf & Denford — Arnica 255

of A. cordifolia that has become established and more widespread
via apomictic reproduction. The evolution of A. nevadensis may
have been facilitated by the climatic cooling of the late Pliocene or
Pleistocene and/or saltation speciation of high altitude populations
of A. cordifolia.

Arnica viscosa is one of the rarest and most distinctive species of
the genus Arnica. It is known from only seven populations on high
alpine volcanic slopes, largely in the Klamath region of Oregon and
California. Its opposite leaves and chromosome number of n = 19,
among other features, clearly place it within the genus and its
pappus characters, broad leaves and flavonoid profile warrant its
inclusion in the subgenus Austromontana. A. viscosa exhibits
virtually every advanced morphological, ecological and chemical
feature of both the genus Arnica and subgenus Austromontana.
Significant derived morphological features include: numerous,
narrow discoid heads with a reduced number of cream-colored
florets; highly branched habit; numerous, sessile, entire leaves;
woody caudex; and a lack of innovations. Its restriction to very
recent volcanic soils and dry, alpine habitat are also considered
derived features. Additionally, the flavonoid profile of A. viscosa,
which includes several highly methylated flavones and a 6-
hydroxylated flavone, is considered highly advanced (Wolf and
Denford, 1984b).

The very distinctive morphology, ecology and flavonoid chem-
istry of Arnica viscosa as well as its restriction to very recent
habitats, i.e., less than 14,000 years old (McKee, 1972), makes an
evaluation of its evolutionary history quite difficult and highly
speculative at best. Maguire (1943) noted the similar leafy,
branching habit and rootstock shared by A. viscosa and A. venosa
and suggested they might be related. Straley (1980) considered A.
viscosa to be a recent derivative of A. venosa and erected the new
subgenus Calarnica to encompass the two species. However, even
Maguire (1943) noted that the two species differ significantly in
several features. In fact, except for the superficial similarity of habit
shared by the two species, they bear little resemblance to each other.
Although quite distinct, A. viscosa shares several features with A.
latifolia including: very narrow heads with lanceolate phyllaries;
sessile leaves; similarly glandular achenes which lack duplex hairs;
the cool high altitude ecology; diploid chromosome number; and

256 Rhodora [Vol. 86

several flavonoids (Wolf and Denford, 1984b). Additionally, in an
extensive artificial hybridization program among the species of
subgenus Austromontana, virtually the only successful crosses were
between A. viscosa and A. latifolia (Straley, 1980).

The derivation of Arnica viscosa from A. venosa, as suggested by
Straley (1980) also seems highly unlikely since the two species
probably represent the two greatest ecological extremes within both
the genus Arnica and subgenus Austromontana. A. venosa occurs at
the lowest elevations and occupies the hottest and driest habitat of
any Arnica while in contrast, A. viscosa is restricted to cold, high
alpine habitats on volcanic soils. The derivation of A. viscosa from
A. latifolia, a sub-alpine species, seems much more likely since it
would require only a slight ecological shift. In addition, both A.
viscosa and A. lJatifolia are diploid and occur sympatrically. Since
A. viscosa shares several morphological, ecological and chemical
features with A. Jatifolia, it seems more probable that A. Jatifolia
has been involved, at least in part, in the derivation of A. viscosa,
perhaps via saltational speciation onto very recent volcanic soils.

Assuming the above proposed phylogeny accurately represents
evolutionary relationships, it is evident that the discoid condition
has arisen independently at least twice within subgenus Austro-
montana and at least three times in the genus Arnica (cf. A. parryi
A. Gray of subgenus Chamissonis). This is not surprising since the
discoid condition has arisen independently numerous times in the
Compositae (Cronquist, 1977). Maguire’s (1943) recognition of
radiate and discoid sections within subgenus Austromontana is
therefore considered artificial and is rejected in the present study.
Morphologically, both A. venosa and A. viscosa clearly belong in
subgenus Austromontana. Additionally, both species contain two
unusual quercetin glycosides common to all species of subgenus
Austromontana, but lacking in the other subgenera of Arnica (Wolf
and Denford, 1984b). Since A. venosa and A. viscosa do not appear
to be related, as proposed by Straley (1980), his new subgenus
Calarnica is also here rejected as artificial.

TAXONOMIC TREATMENT

Arnica L. subg. Austromontana Maguire, Brittonia. 432. 1943.
Type species: A. Jatifolia Bong.
Arnica L. sect. Austromontana Maguire, Brittonia. 432. 1943.

1984] Wolf & Denford — Arnica 257

Arnica L. sect. Eulatifoliae Maguire, Brittonia. 432. 1943. Lecto-
type: A. latifolia Bong. (nom. illegit.).

Arnica L. sect. Eradiatae Maguire, Brittonia. 452. 1943. Lectotype:
A. discoidea Benth.

Arnica L. subg. Calarnica Straley, Ph.D. diss., Univ. British
Columbia. 1980. Type species: A. viscosa A. Gray. This name has
been proposed, but at present it has not been validly published.

Perennial herbs; stems simple to much branched, arising from a
scaly rhizome or woody caudex. Leaves simple, opposite, relatively
broad, 1.0-2.5 times as long as wide, cordate to narrowly ovate or
spathulate, entire to dentate or crenate, sessile to broadly or
narrowly petiolate, uppermost leaves often reduced or bract-like.
Heads solitary or numerous in a corymbiform inflorescence, radiate
or discoid, broadly campanulate to narrowly turbinate; involucral
bracts ovate to lanceolate, acute to acuminate. Ray florets yellow;
disc florets yellow or cream-colored, anthers yellow; pappus white
or rarely tawny, barbellate to subplumose. Achenes gray, brown or
black, + stipitate-glandular and/or + hirsute with duplex hairs.

KEY TO THE SPECIES OF ARNICA SUBGENUS AUSTROMONTANA
1. Heads discoid (marginal corollas sometimes ampliate in No. 3).

2. Leaves sessile.
3. Leaves toothed, veins conspicuous; achenes hirsute with duplex hairs; florets

VENOW Viaiesie a tale sen cndmteseawoeela mae Sales awe ageee’ 8. A. venosa.
3. Leaves entire, veins inconspicuous; achenes stipitate-glandular; florets cream
Le) £0) 10 ne ae ar 9. A. viscosa.

2. Leaves petiolate, the petiole sometimes broad.

4. Lowermost leaves ovate to subcordate or broadly lanceolate; petioles narrow
or scarcely winged; achenes dark gray, stipitate-glandular with duplex.
hairs; species of southern Washington to southern California, including
the Klamath regions. .4.5s.ccenesesquaweeeectncw te 3. A. discoidea.

4. Lowermost leaves spathulate; petioles broad, achenes black, stipitate-
glandular, lacking duplex hairs; species mostly restricted to serpentine soils

of Klamath region......... 0... cece cence eee eeee 7. A. spathulata.
1. Heads radiate.
5. Cauline leaves sessile; heads turbinate; achenes brown...... 5. A. latifolia.

5. Cauline leaves petiolate; heads campanulate to campanulate-turbinate; achenes
dark gray or black.

6. Leaves glabrous, thick, leathery; heads nodding in bud; plants restricted to

serpentine soils of Klamath region ..............0..005 1. A. cernua.

6. Leaves variously pubescent, thin; heads erect in bud; plants not of serpentine
soils, widespread.

258 Rhodora [Vol. 86

7. Stems several-branched, mostly clumped; heads numerous (5-15); achenes
black, mostly lacking duplex hairs ................. 4. A. gracilis.

7. Stems mostly simple, not clumped; heads solitary or few (1-3); achenes
dark gray with duplex hairs.

8. Leaves cordate, toothed; heads broadly campanulate; involucral bracts
ovate-lanceolate; pappus white, barbellate; achenes + hirsute and +
stipitate-glandular; widespread cordilleran species ...............

Uiasgistersiaiate suareie state ais va tna 476 tne Sees g's 6 Sale evo 2. A. cordifolia.

8. Leaves ovate to elliptic, entire; heads campanulate-turbinate; involucral
bracts oblanceolate; pappus white to tawny, barbellate to sub-
plumose; achenes stipitate-glandular; species of Sierra Nevada, north
less commonly to Washington ...........0.004 6. A. nevadensis.

TREATMENT OF INDIVIDUAL TAXA

1. Arnica cernua Howell, Fl. NW. Am. 373. 1900. Type: Oregon,
Josephine Co. On dry banks, base of the Coast mountains
near Waldo, July 1884, 7. Howell 166 (HOLOTYPE, ORE!).

Arnica chandleri Rydb., N. Am. Fl. 34: 339. 1927. Type: California, Humboldt
Co., Hupi (Hoopa) Indian Reservation, June, 1901, H. P. Chandler 1298
(HOLOTYPE, NY!; ISOTYPES, UC!, Us!).

Stems simple, rarely branched, often reddish-purple, 10-30 cm
high, 1.5-2.5 mm diameter, glabrous to sparsely villous; rhizomes
2-3 mm thick, 2-3 pairs reddish scales at the summit. Cauline leaves
3-4 pairs, often reddish-purple, elliptic to ovate, sometimes sub-
cordate, 1.5-8 cm long, 1.5-4 cm broad, often thick and nearly
succulent, glabrous to scabrous, acute to rounded, entire to serrate,
commonly crenate or slightly lobed; petioles narrow or infrequently
broadly winged on lowermost leaves, 2-6 cm long, 1-5 mm broad;
uppermost leaves often reduced, lanceolate and sessile; leaves of the
innovations 4-8, similar to cauline leaves. Inflorescence usually a
single head or corymb of 3-5 heads, often nodding in bud; peduncle
3-15 cm long, sparsely to densely pilose and scabrous, heads
radiate, campanulate-turbinate, 10-25 mm high; involucral bracts
8-14, ovate to broadly lanceolate, 8-16 mm long, 3-6 mm broad,
sparsely to densely pilose and stipitate-glandular, acute to acumi-
nate. Ray florets 5-10, yellow to slightly orange, broadly linear to
elliptic, 10-20 mm long, 4-6 mm broad, 1-3 dentate; disc florets
20-60, yellow, tubular, 10-15 mm long, villous below; pappus of
both ray and disc florets white, barbellate to subplumose. Achenes
dark gray, 6-8 mm long, | mm broad, sparsely to densely hirsute
with duplex hairs. Figure 1. Chromosome number 2n = 38.

1984] Wolf & Denford — Arnica 259

Figure 1. Arnica cernua.

260 Rhodora [Vol. 86

in

Figure 2. Distribution of Arnica cernua @ and Arnica venosa &.

ECOLOGY AND DISTRIBUTION: Relatively rare, but locally abun-
dant in dry, open Pinus- Pseudotsuga menziesii forests at moderate
elevations (500-1500 m). Known only from serpentine soils in Coos,
Curry and Josephine Counties, Oregon and Del Norte, Trinity and
Siskiyou Counties, California. Figure 2. Flowers late April to June.

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Del Norte Co.:
Gasquet-O’Brien Toll Rd., 7.7 mi. N. E. Patrick Creek Rd., D. Breedlove 3178

1984] Wolf & Denford — Arnica 261

(CAS); Boundary Hill, near Telephone Point, A. Eastwood s.n. (CAS); Telephone
Point, A. Eastwood 148 (US). Humboldt Co.: Hoopa, J. Davy and W. Blasdale 5645
(UC); Hoopa Indian Reservation, H. Chandler 1298 (NY, UC, US); Ruby Creek,
Willow Creek Canyon, J. Tracy 7449 (UC); Horse Mt., J. Tracy 15902 (CAS, UC), S.
J. Wolf 471 (ALTA); Willow Creek Canyon, J. Tracy 7057 (CAS, UC); Hoopa Mt.,
J. Tracy 7561 (UC). Siskiyou Co.: 15 mi. n. Happy Camp, G. L. Stebbins 3269 (UC);
Baldy Lookout, F. Hoffman 3547 (UC); Klamath River Canyon, 1/2 mi. below
mouth of Scott River, R. Barneby 11513 (CAS). Trinity Co.: w. side Backbone Ridge
on trail to Raymond Flat, E. Carter 1116 (CAS); Mt. Bally, S. Kleeberger s.n. (CAS).
OREGON: Coos Co.: Iron Mt., W. Baker 6822 (OSC, UC). Curry Co.: headwaters
Chetco River, R. J. Howell and G. True 48823 (CAS). Josephine Co.: base Coast
Mts., near Waldo, 7. Howell 166 (ORE), T. Howell 1446 (NDG, UC); Eight Dollar
Mt., L. Savage s.n. (UC); Little Rock Creek, 2 mi. s.w. O’Brien, L. Constance and R.
Rollins 2993 (CAS, MICH, UC, WTU); Cedar Creek, Deer Creek Canyon, L.
Delting 4036 (UC); Siskiyou Mts., near O’Brien, J. Thompson 1027 (CAS, NDG);
Kerby, A. Sweetser s.n. (CAS); Babyfoot Lake Trail, S. J. Wolf 464 (ALTA).

This species is easily distinguished by its glabrous, thick, ovate,
crenate, petiolate leaves that are often reddish or purple; solitary,
nodding heads, and serpentine habitat. Maguire (1943) considered
Arnica cernua a derivative of A. latifolia. However, both A. cernua
and A. cordifolia share several features including: long petiolate
leaves, broad solitary heads with pilose and stipitate-glandular,
ovate-lanceolate involucral bracts and dark gray achenes. In
contrast, A. /atifolia has sessile leaves, several narrow heads with
narrow, sparsely pubescent bracts and brown achenes. In addition,
the leaves of A. cernua are sometimes subcordate or even cordate
(T. Howell 1446, 1936 NDG) like A. cordifolia.

In the original description of Arnica cernua (Howell, 1900) no
specimens were cited and no type was designated. Maguire (1943)
designated a Howell collection (7. Howell 1466 UC, NDG) as the
lectotype for this species. However, as Rollins (1972) notes, “the
existence of a holotype in the institution where the author worked is
assumed until proven otherwise”. Examination of Howell’s collec-
tions at the University of Oregon has revealed that he did in fact
designate a type by writing “Typ[sic] specimen” on one specimen (T.
Howell 166 ORE). According to the criteria of Rollins (1972) there
would be no need to designate a lectotype since Howell 166 would
automatically be the holotype for A. cernua.

Rydberg (1927) described a smaller, more glandular form of A.
cernua as A. chandleri. Examination of the type specimens
(Chandler 1298, NY, UC, US) as well as two other collections of this
taxon (Davy and Blasdale 5645 UC and Tracy 7449 UC) indicates
that they are all referable to A. cernua.

262 Rhodora [Vol. 86

2. Arnica cordifolia Hook, Fl. Bor. Am. 1: 331. 1834. Type: Alpine
woods of the Rocky Mountains, on the east side, Drummond
s.n. (HOLOTYPE, K photo!).

Arnica macrophylla Nutt., Trans. Am. Phil. Soc. II. 7: 408. 1841. Arnica
cordifolia Hook. var. macrophylla (Nutt.) Maguire, Am. Midl. Nat. 37: 1947.
Type: Blue Mountains of Oregon, Nuttall s.n. (HOLOTYPE, K photo!).

Arnica chionophila Greene, Pittonia 4: 171. 1900. Type: Ruby Mountains,
Nevada, July 20, 1896, E. L. Greene s.n. (HOLOTYPE, NDG!).

Arnica subcordata Greene, Pittonia 4: 173. 1900. Type: on the Athabasca River,
June 26, 1898, W. Spreadborough (Geol. Surv. Can. No. 19644) (HOLOTYPE,
CAN!).

Arnica pumila Rydb., Mem. N. Y. Bot. Gard. 1: 433. 1900. Arnica cordifolia
Hook. var. pumila (Rydb.) Maguire, Madrofio 6: 154. 1942. Type: Gray’s
Peak, Colorado, Aug.-Sept., 1872, Torrey s.n. (HOLOTYPE, NY!).

Arnica parvifolia Greene, P|. Baker. 3: 28. 1901. Type: Marshall Pass, Colorado,
10,000 ft., July 19, 1901, C. F. Baker S515 (HOLOTYPE, NDG!; ISOTYPES, CAS!,
PoM!, uc!, us!, ws!).

Arnica paniculata A. Nelson, Man. Bot. Rocky Mts. 572. 1909. Type: moist
timber, Bridger Peak, Carbon Co., Wyoming, Aug. 24, 1903, L. N. Gooding
1974 (HOLOTYPE, RM!; ISOTYPES, CAS!, GH!, MO!, Ny!, RM!, UC!, Us!).

Arnica evermanii Green, Ottawa Nat. 23: 215. 1910. Type: shores of Petit Lake,
Idaho, Aug. 13, 1895, B. W. Evermann 318 (HOLOTYPE, Us!).

Arnica abortiva Greene, Leaflets 2:47. 1910. TyPE: open spaces in timber, Wind
River Mts., Wyoming, July 23, 1881 W. H. Forwood s.n. (HOLOTYPE, US!).

Arnica andersonii Piper, Proc. Biol. Soc. Wash. 33: 106. 1920. Type: Skeena,
British Columbia, Sept. 11, 1910, J. R. Anderson 677 (HOLOTYPE, Us!;
ISOTYPES, US!, WS photo!).

Arnica austinae Rydb., N. Am. Fl. 34: 340. 1927. Type: Lake City Canyon,
California, July 1898, Austin and Bruce 2165 (HOLOTYPE, NY!; ISOTYPES, NY!,
uc!).

Arnica humilis Rydb., N. Am. Fl. 34: 341. 1927. Arnica cordifolia Hook. var.
humilis (Rydb.) Maguire, Am. Midl. Nat. 37: 138. 1947. Type: on “the
saddle”, Lake Louise, Alberta, July 20, 1904, J. Macoun (Geol. Surv. Can.
No. 65504) (HOLOTYPE, NY!; ISOTYPE, CAN!).

Arnica whitneyi Fernald, Rhodora 37: 334. 1935. Arnica cordifolia Hook. var.
whitneyi (Fernald) Maguire, Brittonia 4: 452. 1943. Type: dry deciduous
woods near Copper Harbor, Keweenaw Co., Michigan, July 4, 1934, Fernald
and Pease 3579 (HOLOTYPE, GH!).

Arnica hardinae St. John, Fl. SE. Wash. 419. 1937. Type: open woods, Lake
Chatcolet, Benewah Co., Idaho, Oct. 2, 1927, G. Weitman 226 (HOLOTYPE,
ws!).

Stems mostly simple, sometimes branched, 10-40 (70) cm high,
2-3 mm diameter, glandular-puberulent to densely villous, espe-
cially above; rhizomes long, giving rise to several basal rosettes and
flowering stems, 1.5-3.0 mm thick, 2-3 pairs of thin brown scales

and often old leaf bases at the summit. Cauline leaves 2-4 (6) pairs,

1984] Wolf & Denford — Arnica 263

cordate or subcordate to narrowly ovate, 3-10 cm long, 2-10 cm
broad, puberulent to sparsely villous and sometimes stipitate-
glandular, especially above, acute to rounded, dentate to coarsely
dentate; petioles (2) 5-10 (20) cm long; uppermost leaves often
reduced, ovate-lanceolate, short-petiolate or sessile, sometimes
bract-like; leaves of the innovations 2-4, similar to cauline leaves or
often more coarsely dentate. Inflorescence usually a single head or
corymb of 3-5 (10) heads; peduncle 4-20 cm long, pilose and
stipitate-glandular; heads radiate, broadly campanulate, 1.5-3.0 cm
high; involucral bracts, 10-20, narrowly ovate to lanceolate, 10-20
mm long, 2-5 mm broad, pilose at base to puberulent above,
sometimes glandular, acute to acuminate, margins scarious. Ray
florets 6-13, yellow, elliptic-oblong, 15-30 mm long, 5-10 mm
broad, subentire to 3-dentate; disc florets numerous, 20-90, yellow,
tubular, 9-12 mm long, sparsely glandular above, sparsely to
densely villous and stipitate-glandular below; pappus of both ray
and disc florets white, barbellate. Achenes dark gray, 5-10 mm long,
1 mm broad, sparsely to densely hirsute with duplex hairs, also
sometimes stipitate-glandular. Figure 3. Chromosome number 2n =
38, 57, 76, 95, 114.

ECOLOGY AND DISTRIBUTION: Very common in mesic Picea-
Pinus-Pseudotsuga menziesii forests or occasionally sub-alpine
meadows from the central Yukon south through northern New
Mexico, northern Arizona, Nevada and northern California. Also
found in the Cypress Hills of Alberta and Saskatchewan and with
outlying populations in the Black Hills of South Dakota, the
Pasquia Hills of Saskatchewan and Riding Mt. Provincial Park,
Manitoba. Also found in Abies balsamea-Betula-Acer forests in
Keweenaw County, Michigan and Sibley Provincial Park, Ontario.
Figure 4. Elevational distribution from 500 m in the Yukon to 3000
m in Colorado. Flowers May-July.

REPRESENTATIVE SPECIMENS: Canada: ALBERTA: Mt. Park, M. Malte and W.
Watson 1969 (UC); Jasper, near Icefields, A. and R. Nelson 4889 (UC); Squaw Mt.,
F. Lewis 92130 (CAN); Mt. Norquay, B. LaSalle 45155 (CAN); Pyramid Mt., A. E.
Porsild and A. J. Breitung 16351 (CAN); Porcupine Hills, Ma/te and Watson 603
(CAN); Lake Louise, Malte and Watson 1014 (CAN); Mt. Park, Malte and Watson
1995 (CAN); Mt. Edith Cavel, J. Macoun 96072 (CAN); Cottonwood Creek, E. H.
Moss 4654 (ALTA, CAN); Mt. Eisenhower, A. E. Porsild and A. J. Breitung 15808
(CAN); Sunshine Ski Lodge, A. E. Porsild and A. J. Breitung 13604 (CAN);
Medicine Lake, M. Dumais 2816 (ALTA, CAN); Swan Dive Fire Tower, Swan Hills,
M. Dumais 4024 (ALTA, CAN); Palisades Mt., A. E. Porsild 22526 (CAN). Mt.

264 Rhodora [Vol. 86

Figure 3. Arnica cordifolia.

1984] Wolf & Denford — Arnica 265

Figure 4. Distribution of Arnica cordifolia.

Norquay, L. Jenkins 1586 (DAO); Waterton Lakes, L. Carmichael 88 (DAO);
Cypress Hills, Spring Creek, R. S., A. J. Breitung 5661 (DAO); Saskatoon Mt., L.
Jenkins 738 (DAO); mi. 21, Jasper-Banff Hwy., W. McCalla 4578 (ALTA); Lake
Louise, G. Turner 11517 (ALTA); Pyramid Lake, 7. Turner 6872 (ALTA); 20 mi.
NW Edson, /. Corns 12108 (ALTA); Ram Mt., M. G. Dumais 7767 (ALTA); Lake
George, G. La Roi s.n. (ALTA); Saskatoon Mt., Prairie and Heywood 69 (ALTA);
Swan Hills, M. G. Dumais 4024 (ALTA); Surprise Lake, M. G. Dumais 6212
(ALTA); Jarvis Lake, M. G. Dumais 2420 (ALTA); Saddle Hills, N. of Sexsmith, E.
H. Moss 8464 (ALTA); Winfield, F. Rusconi s.n. (ALTA); Mercoal, E. Woollven 23
(ALTA). Mt. Park, E. Woollven 7 (ALTA).

BRITISH COLUMBIA: Alaska Hwy., 12 mi. NW Dawson Creek, Calder and
Kukkonen 26801 (UC); Revelstoke, W. Spreadborough 64984 (NDG); Lake

266 Rhodora [Vol. 86

Osoyoos, J. Macoun 69323 (NDG); Skagit Valley, J. Macoun 69325 (NDG); SW
corner Dease Lake, T. McCabe 8726 (UC); Crowsnest Pass, 7. McCabe 6463 (UC);
Takla Landing, T. McCabe 7835 (UC); Germansen Landing, 7. McCabe 7664 (UC);
Clearwater, 7. McCabe 2078 (UC): 15 mi. SW Kleena Kleen, T. McCabe 583 (UC);
Atlin, Setchell and Parks s.n. (UC); Sinclair Pass, 7. McCabe 6219 (UC); Arnarchist
Mt., 7. McCabe 5956 (UC); Atlin, A. Eastwood 651 (CAS, UC); Bear Lake, T.
McCabe 7997 (UC); Pavilion Mt., T. McCabe 213 (UC); Pinantan, T. McCabe 2013
(UC); Skeena Crossing, T. McCabe 7027 (UC); Princeton, A. McCallum s.n. (UC);
Lake Bootahnie, J. and E. Thompson 119 (MICH); 2 mi. E. Williams Lake, J. Calder
et al 16938 (DAO, OSC); Annaham Lake, G. Bellinger 32395 (OSC); Cooper Mt.,
near Princeton, Taylor and Szczawin s.n. (OSC, UBC); Smithers, J. Menzies 6016
(UBC); Cathedral Park, Twin Buttes, R. Hainault 7879 (UBC); Mt. Edziza, Annas
and Klinka s.n. (UBC); Hedley, 7. Taylor 2089 (UBC); 10 mi. E. Summit Pass, H.
Raup and D. Correll 10738 (UBC); 49 mi. E. Teslin, H. Raup and D. Correll 11099
(CAN, UBC); Cassier, 7. Taylor et al 398 (CAN, UBC); mi. 81, Haines Rd., T.
Taylor et al 1342 (UBC). Mt. Defot, NW Dease Lake, J. Teit 99 (UBC); mi. 206,
Alaska Hwy., A. E. Porsild 9111 (GH); Mt. Finlayson, Vancouver Island, C.
Newcomb 60 (GH); Laird Hot Springs, A. E. Porsild and A. J. Breitung 22259
(CAN); Revelstoke, J. Macoun 64984 (CAN). Mt. Brilliant, H. Laing 687 (CAN);
Osoyoos Lake, J. Macoun 69323 (CAN); Telegraph Creek, Dawson s.n. 14731
(CAN); Victoria, J. Tolmie 1053 (DAO); Mt. Pope, J. Whitehorn 382 (DAO);
Blanchard River, 66 mi. S. Haines Jct., Calder and Kukkonen 28185 (DAO); mi. 625
Alaska Hwy., Calder and Gillet 25623 (DAO). Hurricane Creek, Atlin Dist., J.
Aitken 10 (DAO); Kootney Nat. Park, Sinclair Nature Trail, K. See/ 27 (DAO): 12
mi. E. Field, G. Turner 3899 (DAO); 8 mi. NE Ft. Steele, R. Taylor and D. Ferguson
629 (DAO); 32 mi. W Prince George, Mulligan and Woodbury 1688 (DAO); Kleena
Kleen P.O., Calder and Parmelee 19185 (DAO); 5 mi. N. Little Fort, Calder and
Saville 8638 (DAO); Nuttbide Lake, Quiquet and Ritcey 57 (DAO); 9 mi. E.
Williams Lake, Calder and Parmelee 17040 (DAO).

MANITOBA: Rassburn Tower Cabin, Riding Mt. Provincial Park, A. Lovaas
61-8 (DAO),

NORTHWEST TERRITORIES: Laird River between Nahanni Butte and
Simpson, C. Crickmay 114 (CAN); SW MacKenzie Mt., Laird Range, 15 mi. NW Ft.
Laird, W. Jeffrey 424 (CAN).

ONTARIO: Ravine Lake, Sibley Provincial Park, C. Garton 15485 (CAN,
MICH, UC): /5/63 (UC), 15164 (CAN).

SASKATCHEWAN: Mt. Cabin, Pasquia Hills, J. Rowe 983 (CAN); Cypress
Hills; C. Frankton 266 (DAO); R. Newsome 394-64 and 470-64 (DAO); Ledingham
and Hudson 1788 (DAO); G. Selleck 394 (DAO); R. Russell s.n. (DAO).

YUKON TERRITORY: Mi. 802, Alaska Hwy., D. Bolinger s.n. (OSC); White-
horse, J. Gillett and D. Mitchell 3862 (DAO, OSC); mi. 23, Campbell Hwy., V.
Harms 17193 (DAO, GH); mi. 116, Canol Rd., A. EF. Porsild and A. J. Breitung
10079 (CAN); St. Elias Mt., A. Pearson 67-210A (CAN); Keno, G. Potack 119369
(CAN); Canol Rd., km 15.5, S. J. Wolf 507 (ALTA); km 118, S. J. Wolf 508 (ALTA);
3 mi. S Whitehorse, J. Gillett 3259 (DAO, RM); lower Kathleen Lake, Kluane Nat.
Park, G. and G. Douglas 5884 (DAO); mi. 858, Alaska Hwy., Calder and Gillett
25734 (DAO); mi. 777, Alaska Hwy., Calder and Kukkonen 28318 (DAO); 16 mi. S.
Haines Jct., S. J. Wolf 300 (ALTA); km 196, Haines Rd., S. J. Wolf 301 (ALTA); km

1984] Wolf & Denford — Arnica 267

1479, Alaska Hwy., S. J. Wolf 307 (ALTA); Alaska Hwy., 1.6 km E Squanga Lake,
S. J. Wolf 505 (ALTA).

United States:s ALASKA: Sitka, A. Paska s.n. (UC); mi. 17.5 Haines Hwy., M.
Williams 1473 (OSC); Sitka, A. Heller 14942 (WTU),.

ARIZONA: Apache Co.: Lukachukai Mts., Goodman and Payson 2865 (GH);
rd. to Wide Lake, Lukachukai Mts., C. Mason et al 244] (UC). Coconino Co.: N.
Rim, Grand Canyon, U.S. Park Service 2007 (US); Indian Hollow, Kaibab Plateau,
L. Gooding 203 (UC); N. slope, San Francisco Peaks, J. Leiberg 5897 (US).

CALIFORNIA: Alpine Co.: Hermit Valley, F. Peirson 11593 (UC). El Dorado
Co.: Magies Peak, H. M. Hall 8810 (UC). Glenn Co.: Black Butte, V. Rattan s.n.
(CAS). Humboldt Co.: Salmon Summit, J. Tracy 14372 (UC). Lassen Co.: | mi. E
Fredonyer Pass, A. Heller (UC). Madera Co.: Shadow Lake Trail, 1 mi. fm. Agnew
Meadow, J. and C. Reveal 427 (RM, UC, WTU). Mariposa Co.: Yosemite Valley, G.
Grant 4356 (UC); Signal Peak, C. Quick 1997 (CAS). Modoc Co.: 15 mi. NE Alturas,
C. L. Hitchcock s.n. (UC); Cedar Pass, J. Weiler 61235 (UC); Emerson Creek,
Alexander and Kellogg 4668 (UC). Mono Co.: Slate Creek Basin, E. Mt. Conness, J.
Clausen 1124 (OSC); Mammoth Lakes, Lake Mary, L. Rose 42194 (WTU). Nevada
Co.: Donner Lake, S. J. Wolf 476 (ALTA). Placer Co.: Deer Park, H. Geis 38 (UC);
W. side Donner Lake, M. Denton 390] (WTU). Plumas Co.: Gold Lake Rd., S. J.
Wolf 478 (ALTA); American Valley, R. Austin s.n. (NDG). Santa Clara Co: Mt.
Hamilton, R. Pendleton 873 (UC). Siskiyou Co.: Caribou Lake, 1. Wiggins 13562
(UC); English Lake, F. Oettinger 1023 (UC); Marble Mt., H. Chandler 1615 (UC);
Salmon Mts., 10 mi. SW Etna, A. Eastwood and J. Howell 5037 (CAS). Trinity Co.:
Oregon Gulch Mt., J. Tracy 7538 (UC); Musser Hill, H. Yates 395 (UC). Tuolumne
Co.: Gaylor Lakes, H. Mason 11368 (UC).

COLORADO: Boulder Co.: Mts. between Sunshine and Ward, F. Tweedy 4893
(RM); Boulder, F. Ramaley 71 (RM). Chaffee Co.: Morass Creek, 1. Clokey 3474
(UC). Clear Creek Co.: Chicago Creek, J. Ehlers 8402 (MICH). Custer Co.:
Westcliffe, C. Erlanson 1768 (MICH). Douglas Co.: 7.3 mi. SW Sedalia, W. Weber
7440 (WTU). Eagle Co.: 2 mi. E. Tennessee Pass, /. Tidstrom 4098 (US). El Paso Co.:
2 mi. W Palmer Lake, G. Robbins 463 (DAO). Fremont Co.: Sierra Sangre de
Cristo, 7. Brandegee 241 (UC). Gilpin Co.: Tolland, E. Palmer 31255 (GH).
Fontleroy Place, H. Rodeck 58 (DAO). Grand Co.: Berthoud Pass, J. Ehlers 8452
(MICH); Berthoud Pass, F. Tweedy 582] (RM). Gunnison Co.: Bonton Mine, J.
Clokey 3009 (UC). Hinsdale Co.: W. Slumgullion Pass, J. Barrell 249-65 (US).
Huerfano Co.: Mt. SW Blue Lake, Mosquin and Gillett s.n. (UC). Jefferson Co.:
Lookout Mt., H. Shacklette 5934 (MICH). Lake Co.: Lost Man Camp. 6 mi W
Independence Pass, U. Waterfall 11623 (UC); Leadville, J. Ehlers 8218 (MICH). La
Plata Co.: Eagle Pass, Mosquin and Gillett 5430 (UC). Larimer Co.: Rocky Mt. Nat.
Park, D. McNeal 202 (RM). Mesa Co.: Grand Mesa, U. Waterfall 11653 (UC).
Mineral Co.: Wolf Creek Pass, C. Wolf 3007 (CAS). Ouray Co.: Ouray, Biltmore
1149 (US). Pitkin Co.: West Springs Creek, J. Langenhein 1399 (UC). Sanguache
Co.: Marshall Pass, J. Barrell 29-66 (US). Summit Co.: Breckenridge, K. Mackenzie
80 (RM).

IDAHO: Bannock Co.: 3 mi. above Pocatello, A. Cronquist 2302 (GH). Bear
Lake Co.: Aspen Range, Georgetown Canyon, N. Holmgren and B. Bethers 4411
(UC). Benewah Co.: SE Plummer, W. Baker 16085 (WTU). Bonneville Co.: Tie
Canyon, 6 mi. SW Victor, N. Holmgren and V. Marttala 5376 (UC). Butte Co.:

268 Rhodora [Vol. 86

Craters of the Moon Nat. Monument, Dole 49 (UC). Camas Co.: Soldier Mts.,
Ketchum-Featherville Rd., C. L. Hitchcock and C. V. Muhlick 10417 (WTU). Custer
Co.: 10 mi. W. Cape Horn, C. L. Hitchcock and C. V. Muhlick 9654 (UC). Elmore
Co.: 23 mi. NE Mountain Home, Davidse and Collotzi 453 (UC). Franklin Co.: 2 mi.
SW Franklin Basin R.S., B. Maguire 21645 (WTU). Idaho Co.: Warren Summit, R.
Davis 2555 (UC). Kootenai Co.: Albany Falls, C. Speilberg 452 (RM). Latah Co.:
Moscow Mt., L. Abrams 613 (UC). Lemhi Co.: Moccasin Creek, C. L. Hitchcock
and C. V. Muhlick 14287 (UC). Nez Perces Co.: Lake Waha, A. and E. Heller 3170
(UC). Owyee Co.: Silver City, J. Macbride 955 (RM). Rich Co.: Bear Lake Summit,
S. J. Wolf 485 (ALTA). Shoshone Co.: Bullion Pass, St. Line, W. Baker 13446
(OSC). Valley Co.: McCall, W. Boone 29 (RM); 15 mi. N. Banks, C. L. Hitchcock
and C. V. Muhlick 8578 (WTU). Washington Co.: Mann Creek, H. Tucker s.n.
(RM).

MICHIGAN: Keweenaw Co.: 3 mi. E. Agate Harbor, F. Hermann 7995 (UC);
Fort Wilkins State Park, M. Feigley and L. Nagel s.n (MICH); Copper Harbor
Cemetery C. Richards 3783 (MICH); 222] (MICH); Grand Marias Harbor, C.
Richards 2144 (DAO, MICH); | mi. W. Copper Harbor, F. Herman 776] (MICH,
RM); bluffs SE of Eagle Harbor, M. Fernald and A. Pease 3580 (GH, MICH);
Copper Harbor, Pease and Ogden 25178 (GH).

MONTANA: Deerlodge Co.: Storm Mt. S. J. Wolf 435 (ALTA). Flathead Co.: 8
km E. Bigfork, S. J. Wolf 496 (ALTA). Gallatin Co.: Targhee Pass, S. J. Wolf 433
(ALTA); Sage Creek, D. Swingle s.n. (MICH). Lake Co.: near Biological Station,
Flathead Lake, P. Smith 37 (NDG). Lewis and Clark Co.: 8 mi. W. Lincoln, C. L.
Hitchcock 17956 (UC). Lincoln Co.: Mt. Marston, S. J. Wolf 343 (ALTA); 25 kmS.
Eureka, S. J. Wolf 443 (ALTA). Madison Co.: E. of Brandon Lakes, C. L. Hitchcock
16960 (UC). Meagher Co.: 35 mi. NW White Sulphur Springs, C. L. Hitchcock 16225
(UC, WTU). Missoula Co.: Blackfoot Valley, H. LaCasse 15 (MICH); Missoula,
Nawrodcki and Neff 2 (NDG); 34 km N. Seeley Lake, S. J. Wolf 495 (ALTA):
Marshall Ski Area, S. J. Wolf 492 (ALTA). Ravalli Co.: 32 mi. E. Hamilton, G.
Hedgcock s.n. (WTU). Saunders Co.: divide between Camas and Perry Basin, F.
Barkely 2568 (NDG). Sweetgrass Co.: 1 mi. below Rainbow Lakes, C. L. Hitchcock
16506 (UC).

NEW MEXICO: Colfax Co.: Baldy Peak, P. Standley 14307 (US); Hermatite
Canyon, D. St. John 52 (GH). Rio Arriba Co.: Chama, P. Standley 6713 (US);
Brazos Canyon, P. Standley 10917 (US); Pecos River National Forest, Wuisor
Creek, P. Standley 4255 (US).

NEVADA: Elko Co.: Jarbridge Mts., Coon Creek, P. Train 67] (NDG);
Lamoille Lake, A. Holmgren 14177 (UC); Steele Creek, Ruby Mts., A. Borell s.n.
(UC); 8 mi. W. North Fork, N. Nichols 32] (DAO). Humboldt Co.: Santa Rosa
Range, J. Gentry 1581 (DAO, NY, RM). Washoe Co.: Headwaters Galena Creek, W.
Archer 6677 (DAO, UC); Hunters Creek Rd., 9-11 mi. SW Reno W. Archer 6295
(CAD). White Pine Co.: Wheeler Peak, B. Maguire 21111 (GH, UC); S. J. Wolf 480
(ALTA).

OREGON: Baker Co.: Eagle Creek, 7. Gustafson s.n. (UC). Crook Co.: Ochoco
N.F., S. Warg s.n. (OSC); 23 mi. NE Prineville, F. Chisaki 780 (RM). Curry Co.:
Summit of Pistol River Mt., J. Thompson 4565 (CAS). Deschutes Co.: Pavilina
Lake, M. Peck 9658 (OSC). Grant Co.: Malheur N.F., Fields Park, A. Kruckeberg
546 (UC). Harney Co.: Myrtle Creek Canyon, M. Peck 2846 (OSC). Steens Mts., C.

1984] Wolf & Denford — Arnica 269

Hansen 699 (OSC). Hood River Co.: Hood River, 7. Howell 477 (OSC). Jackson
Co.: Mt. Ashland, M. Peck 2934 (OSC). Jefferson Co.: Black Butte, J. Johnson 470
(OSC). Josephine Co.: 4.6 mi. S. Hugo, K. Chambers 2916 (OSC). Klamath Co.:
Fossil Lake, near Crater Lake, H. Furlong s.n. (UC). Lake Co.: 2 mi. NW Crooked
Creek, M. Loveless 77 (UC). Morrow Co.: Tupper Guard Station, E. Winn s.n.
(OSC). Umatilla Co.: 14 km S Ukaih, S.J. Wolf 451 (ALTA). Union Co.: 40 km E
Ukaih, S. J. Wolfs.n. (ALTA); Eagle Cap, G. Mason 1395 (OSC). Jarboe Creek, P.
Standley s.n. (OSC). Wallowa Co.: Lostine River, 18 mi. from Lostine, J. Murphy 89
(UC); Hurricane Creek, G. Mason 5365 (OSC); Lick Creek Rd. H. Gilkey 8 (OSC).
Wasco Co.: 2 mi. W. the Dalles, M. Peck 2791 (OSC).Wheeler Co.: Fossil, W.
Lawrence 2988 (OSC).

SOUTH DAKOTA: Lawrence Co.: Spearfish Canyon, N Black Hills, F. Bennett
941 (CAS); 10 mi. NW Deerfield, P. Johnson 527 (MICH); top of Custer Peak, E.
Palmer 37547 (GH); Mt. Roosevelt, W. Over 17639 (RM); Whitewood, H. Hayward
1207 (RM).

UTAH: Beaver Co.: 23 km E. Beaver, S. J. Wolf 482 (ALTA). Cache Co.: Logan
Canyon, B. Maguire 3881 (RM, UC). Carbon Co.: Willow Springs, | mi. E.
Sunnyvale, S. Blake 9587 (UC). Duchesne Co.: W. Mt. Agassiz, B. Maguire et al
4317 (RM); Ashley Nat. Forest, S. J. Wolf 396 (ALTA). Garfield Co.: Mt. Ellen,
Henry Mts., R. McVaugh 14652 (CAS, MICH). Iron Co.: 3 mi. N. Cedar Breaks
Nat. Monument, C. L. Hitchcock and C. V. Muhlick 4603 (UC, WTU); 17.2 km E.
Cedar City, S. J. Wolf 481 (ALTA). Juab Co.: Granite Canyon, B. Maguire and R.
Becraft 2853 (UC). Piute Co.: Marysvale, M. Jones s.n. (CAS). Rich Co.: Bear Lake
Summit, S. J. Wolf 485 (ALTA). Salt Lake Co.: Big Cottonwood Canyon, P.
Rydberg and E. Carlton 6652 (UC); City Creek Canyon, Salt Lake City, K. Brizzee
7856 (WTU). San Pete Co.: Skyline Drive, B. Maguire 20033 (WTU); S. J. Wolf 393
(ALTA). Summit Co.: Burntfork Creek, E. Jensen s.n. (UC). Tooele Co.: S. Willow
Creek, Stansbury Range, B. Maguire 21753 (GH, UC). Utah Co.: Mt. Timpanogos,
E. Applegate 8439 (CAS); 9.5 km E. Mt. Timpanogos, S. J. Wolf 483 (ALTA).
Washington Co.: Forsyth Creek, Pine Valley Mts., P. Munz 16924 (WTU).

WASHINGTON: Asotin Co.: Field Springs Park, S. J. Wolf 446 (ALTA). Chelan
Co.: Tumwater Mt., J. Thompson 6479 (WTU); Lookout Mt., J. Thompson 6479
(WTU); Wenatchee Lake, W. Dress 4228 (UC). Clallam Co.: Boulder Creek,
Olympic Nat. Forest, G. Jones 8475 (WTU). Columbia Co.: Wolf Fork, Touchet
River, H. St. John et al 6971] (UC). Garfield Co.: Blue Mts., D. Peters 385 (UC); 15
mi. S. Pomeroy, C. L. Hitchcock and C. V. Muhlick 8302 (UC, WTU). Kittitas Co.:
Virden, J. Thompson 11582 (UC, WTU); Lookout Mt., J. Thompson 14512 (MICH;
NDG). Klickitat Co.: NE Bingen, W. Suksdorf 2760 (UC). Mason Co.: Mt. Elinor,
P. Freer 371 (WTU). Okanogan Co.: Salmon Creek, C. Fiker 686 (WTU); 20 mi. W.
Winthrop, G. and G. Douglas 3514 (ALTA, DAO). Pend Oreille Co.: Calispell, F.
Kreager 351 (UC, WTU). Skamania Co.: Hamilton Mt., L. Delting 7066 (UC).
Snohomish Co.: Mt. Dickerson, R. Owen s.n. (WTU). Spokane Co.: Mt. Spokane,
S. J. Wolf 445 (ALTA). Stevens Co.: E. side Columbia River, 12 mi. above mouth of
Spokane River, H. Rogers 400 (UC). Yakima Co.: Mt. Aix, J. Thompson 15016
(WTU); Chinook Pass, J. Thompson 15136 (WTU); Bald Mt., H. St. John 7854
(UC).

WYOMING: Albany Co.: 7.9 mi. W. Centennial, S. J. Wolf 424 (ALTA); Woods
Creek Canyon, C. L. Porter and M. Porter 9810 (UC); Woods Creek, L. Goodding

270 Rhodora [Vol. 86

143] (UC); Centennial, Kauffman and Erlanson 113 (MICH); University Camp,
Medicine Bow Mts., A. Nelson 7798 (RM). Big Horn Co.: 10-15 mi. E. Kane, L. and
R. Williams 3016 (RM). Carbon Co.: South Brush Creek Campground, B. and L.
Nelson 451 (RM). Crook Co.: 6 mi. NE Hulett, M@. Ownby 599 (RM, UC, WTU).
Fremont Co.: Gannett Creek, F. Jozwik 395 (UC); 1 mi. S. Pacific Spring, C. L.
Porter 4525 (RM). Lincoln Co.: Grover Park, Afton Area, O. Harrison 55 (RM).
Natrona Co.: Casper Mt., R. Tresler 55 (RM). Park Co.: Crazy Woman Creek,
Beartooth Mts., L. and R. Williams 3518 (NDG). Saunders Co.: divide between
Camas and Perry Basin, F. Barkley 2568 (NDG). Sublette Co.: Green River Lakes,
A. Beetle 10534 (NDG); 16 km NE Pinedale, S. J. Wolf 427 (ALTA). Teton Co.:
Togowotee Pass, S. J. Wolf 430; 432 (ALTA); Teton N. P., Hidden Falls, S. J. Wolf
429 (ALTA); Jackson Hole, J. and M. Reed 2250 (RM). Washakie Co.: 3 mi. E.
Tensleep, H. Fisser 784 (RM). Yellowstone N.P.: W. Setchell s.n. (UC); Mammoth
Hot Springs, F. Burglehaus 94 (MICH).

Arnica cordifolia is probably the most widespread Arnica in
western North America, occupying fairly mesic habitats, in Pinus-
Pseudotsuga menziesii or Picea forests throughout most montane
areas of the region. In such areas as the foothills of Alberta, it forms
extensive, nearly continuous populations for many kilometers. In
the east A. cordifolia is restricted to only a few populations in
Betula-Acer forests of Keweenaw Co., Michigan and Sibley
Provincial Park, Ontario. A. cordifolia has also been recently
collected in the Pasquia Hills, east of Saskatoon, Saskatchewan
(J. Rowe 983 CAN) and in Riding Mountain Provincial Park,
Manitoba (A. Lovaas 61-8 DAO).

In its typical woodland habitat, Arnica cordifolia displays little
variability and is easily recognized by its large, solitary heads with
broad rays and its cordate, dentate leaves. However, as Maguire
(1943) noted, when it occurs in more exposed or disturbed habitats
the leaves become smaller, narrower and lose their cordate bases;
the pubescence becomes denser, harsher and more glandular; and
the plants become smaller and more branched with several heads.
This morphological form is characteristic of triploids and led to the
suggestion that it may be the result of hybridization (Wolf, 1980).
However, since recent flavonoid studies revealed no significant
systematic differences among the chromsome races of A. cordifolia,
including the triploids (Wolf, 1980), a hybrid origin can probably be
ruled out.

Higher elevation forms of Arnica cordifolia tend to be much
reduced, lack cordate leaf bases and have subentire leaves. This
form has previously been recognized as A. pumila Rydb. or as A.
cordifolia Hook. var. pumila (Rydb.) Maguire. However, when

1984] Wolf & Denford — Arnica 271

such plants are transplanted to the greenhouse they revert to typical
A. cordifolia (cf. S. Wolf 306 ALTA). The high altitude form has
often been confused with A. nevadensis, particularly in the
mountains of Utah. However, the latter can be recognized by its
narrower heads, oblanceolate phyllaries, less pubescent leaves and
darker pappus with longer setae.

According to Maguire (1943), Drummond’s explorations were
largely in the vicinity of Jasper House, Alberta, along the Upper
Athabasca River and at the headwaters of the North Saskatchewan
River. Bird (1967) also concluded that most of Drummond’s
collections from the “Rocky Mountains” probably came from
Jasper National Park. The type of Arnica cordifolia from “Alpine
woods of the Rocky Mts.”, was therefore probably collected in the
front range of the Rocky Mountains, probably in Jasper National
Park, Alberta. The type sheet consists of four specimens, the
holotype collected by Drummond and three specimens collected by
Douglas on the same expedition.

Maguire (1943) considered the rare Arnica paniculata A. Nelsona
distinct species while noting it may be a hybrid between A.
cordifolia and A. parryi A. Gray. Ediger and Barkley (1978)
considered this taxon a hybrid and therefore did not give it
taxonomic recognition. In fact, with the exception of its numerous
heads and ovate leaves, A. paniculata differs very little from A.
cordifolia as circumscribed in the present study. Additionally, a
chromosome voucher of an A. cordifolia population with 2n = ca.
97 (T. Mosquin and J. Gillett 5425 La Plata Co., Colorado; UBC)
bears considerable resemblance to the type collection of A.
paniculata (L. N. Gooding 1974, Carbon Co., Wyoming). It is very
likely that A. paniculata represents an A. cordifolia with a high
chromosome number and/or the product of introgression between
the latter and some other taxon. It is therefore reduced to synonymy
under A. cordifolia.

Arnica whitneyi of Keweenaw Co., Michigan was first described
by Fernald (1935). It occurs as one large discontinuous population
between Copper and Eagle Harbors (Straley, 1980). Recently it has
also been located in Sibley Provincial Park, Ontario (Garton 15164
CAN, 15486 MICH). Maguire (1943) noted that this taxon differed
little from western populations of A. cordifolia but retained it as a
subspecies of the latter because of its disjunct distribution.
However, Ediger and Barkley (1978) did not give this taxon formal

272 Rhodora [Vol. 86

taxonomic recognition. The facts that these eastern populations
represent very typical A. cordifolia, share similar chromosome
numbers (Wolf, 1980) and flavonoid chemistry (Wolf and Denford,
1983) with A. cordifolia, and that intervening populations of A.
cordifolia in Saskatchewan and Manitoba also exist, support Ediger
and Barkley’s (1978) treatment. In the present study A. whitneyi is
therefore reduced to synonymy under A. cordifolia.

3. Arnica discoidea Benth., Pl. Hartw. 319. 1849. Type: Monterey,
California, Hartweg 1805 (HOLOTYPE, K photo!; ISOTYPES, GH
photo!, Ny!).

Arnica parviflora A. Gray, Proc. Am. Acad. 7: 363. 1867. Type: Chaparral,
Humboldt Co., California, Geol. Surv. Calif. 1867. H. N. Bolander 6051
(HOLOTYPE, GH!; ISOTYPES, UC! K photo!, us!).

Arnica cordifolia Hook. var. eradiata A. Gray, Syn. Fl. N. Am. 1: 381. 1884.
Arnica discoidea Benth. var. eradiata (A. Gray) Cronquist, Vasc. Pl. Pac.
NW. 5: 49. 1955. Type: Hood River, Oregon, 1884, Mrs. Barrett s.n.
(HOLOTYPE, GH!).

Arnica grayi A. Heller, Muhlenbergia 1: 5. 1900. Type: Hood River, Oregon,
1884, Mrs. Barrett s.n. (HOLOTYPE, GH!).

Arnica falconaria Greene, Ottawa Nat. 23: 215. 1910. Type: Falcon Valley,
Washington, June 27, 1892, W. N. Suksdorf 1617 (HOLOTYPE, US!; ISOTYPES,
uc!, GH photo!, Ny!).

Arnica alata Rydb., N. Am. FI. 34: 342. 1927. Arnica parviflora A. Gray subsp.
alata (Rydb.) Maguire, Brittonia 4: 455. 1943. Arnica discoidea Benth. var.
alata (Rydb.) Cronquist, Contr. Dudley Herb. 5: 102. 1958. Type: Yosemite,
California, 1865, J. Torrey 258a (HOLOTYPE, NY!).

Arnica sanhedrensis Rydb., N. Am. Fl. 34: 342. 1927. Type: Foothills of Mt.
Sanhedren, Lake Co., California, Heller 5985 (HOLOTYPE, NY!; ISOTYPES, POM!,
uc!, us!).

Stems mostly simple to branched above, 15-60 cm tall, 2-5 mm
diameter, villous and stipitate-glandular throughout; rhizomes
giving rise to numerous basal rosettes and flowering stems, 2-5 mm
thick, scales and old leaf bases crowded toward the summit. Cauline
leaves 3-7 pairs, sometimes crowded toward stem base and often
reduced above, ovate to broadly lanceolate, seldom subcordate,
2-12 cm long, 1-7 cm broad, glabrate to pilose and stipitate-
glandular, serrate to coarsely dentate or crenate, rarely subentire;
petioles narrow, 1.5-8.0 cm long, often broadly winged on upper
reduced leaves; leaves of the innovations 4-10; similar to cauline
leaves. Inflorescence a corymb of 3-10 (30) heads; peduncle 2-15 cm
long, stipitate-glandular and densely pilose; heads discoid, the

1984] Wolf & Denford — Arnica 2t3

marginal corollas sometimes ampliate, turbinate-campanulate, 12-
22 mm high; involucral bracts 8-15, ovate-lanceolate to narrowly
lanceolate, 8-13 mm long, 1-4 mm broad, densely pilose and
Stipitate-glandular, acute to acuminate. Florets 20-50, yellow,
tubular, 8-11 mm long, stipitate-glandular and sparsely to densely
villous; pappus white, barbellate (to subplumose). Achenes dark
gray, 6-8 mm long, | mm broad, stipitate-glandular and hirsute
with duplex hairs. Figure 5. Chromosome number 2n = 38, 57, 76.

ECOLOGY AND DISTRIBUTION: Relatively uncommon in moder-
ately dry Quercus- Pinus forests from Klickitat County in southern
Washington, south sparingly in the Cascades through northern
California, but more common in relatively exposed chaparral in the
coast ranges of California south to Orange County. Figure 6.
Elevational distribution ranges from near sea level to 1500 m.
Flowers May-July.

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Butte Co.: Jonesville,
E. Copeland 400 (MICH, UC). Contra Costa Co.: Inner Black Hills, W. of “1970,”
M. Bowerman 2173 (UC); Meridian Peak, M. Bowerman 849 (UC). Del Norte Co.:
Black Butte, A. Eastwood 2137 (CAS); Smith River, above Rock Creek Lodge, W.
Cooke s.n. (UC). Glenn Co.: Alder Springs, M. Baker 10872 (UC). Humboldt Co.:
Croghan Hole, J. Tracy 19273 (UC); Bee Mt., C. Quick 59-31 (CAS); Hoopa Mt.,
Davy and Blasdale 5668 (UC); Trinity Summit, J. Tracy 14154 (UC); J. Tracy 18182
(UC); Horse Mt., J. Tracy 17833 (UC); Briceland, J. Tracy 6335 (CAS, UC); White
Thorn Valley, J. Tracy 500] (UC). Lake Co.: Mt. Hull, H. M. Hall 9564 (UC). Coff
Mt., H. M. Hall 9595 (UC); Ukiah, H. Yates 3735 (UC); Elk Mt., H. M. Hall 9587
(UC); Adam's Springs, R. Hoover 5347 (NDG, UC); Eel River, L. Benson 3726
(NDG). Marin Co.: Mt. Tamalpais, A. Heller 8392 (UC); K. Brandegee s.n. (UC); W.
L. Jepson s.n. (UC); F. Boiletti s.n. (NDG, UC); Blithdale Canyon, J. Howell 26621
(CAS); Lagunitas, H. M. Hall 8502 (UC). Mariposa Co.: Cathedral Rocks, C.
Sharsmith 2168 (UC); Sherlocks, J. Congdon s.n. (CAS); Yosemite Valley, L.
Abrams 4527 (UC); H. M. Hall 8867 (UC); Vernal Falls, H. M. Hall and E. B.
Babcock 3443 (UC). Mendocino Co.: 5 mi. E. Point Arena, L. Rose 39170 (UC);
Point Arena, A. Eastwood and J. Howell 6252 (CAS); Comptche, H. Walker 342
(UC). Modoc Co.: John Henry Hill, 44. Manning 370 (UC). Monterey Co.: Pacific
Grove, A. Elmer 4402 (CAS, OSC, UC); Chews Ridge, S. Jonesburg, L. Snyder 3774
(UC); Los Burros Creek, 15 mi. SW Jolon, C. Hardham 7037 (OSC); Big Sur, Y.
Mexia s.n. (CAS, UC); Point Lobos, Helley s.n. (CAS); Hanging Valley, Santa Lucia
Mts., D. Breedlove 36264 (CAS); Jolon, C. Hardham 7037 (WTU). Napa Co.:
Howell Mt., J. Tracey 2224 (UC); White Sulphur Springs, St. Helena, H. Chandler
7579 (UC); E. side of Mt. St. Helena, R. Hoover 5014 (UC). Orange Co.: Santa Ana
Mts., 4.8 mi. above gate on Silverado Canyon Truck Trail, P. Raven 17751 (UC);
Mojeska Springs, Santa Ana Mts., W. Pequegnat s.n. (WTU). Plumas Co.: S. E.
Quincy, C. Quick 41-60 (CAS); Taylorsville, M@. Glemens s.n. (CAS). San Luis
Obispo Co.: Ocean View Mine, N. of Cambria, C. B. Hardman 6828 (CAS, UCSB);

274 Rhodora [Vol. 86

Figure 5. Arnica discoidea.

1984] Wolf & Denford — Arnica Pa pe

WASHINGTON

OREGON

CALIFORNIA

Figure 6. Distribution of Arnica discoidea.

276 Rhodora [Vol. 86

Pine Mt., Santa Lucia Mts., R. Hoover 8018 (CAS). San Mateo Co.: Skyline Drive,
D. Demaree 9150 (NDG); Kings Mt. Rd., D. Keck 1775 (CAS, OSC, WTU); Portola
State Park, J. Thomas 9558 (CAS). Santa Barbara Co.: Mudulee Lookout Trail
from Big Pine Rd., E. Blakley 6056 (CAS); LaCumbre Peak, D. Breedlove 3762
(CAS); D. Breedlove 585 (CAS, UCSB); S. J. Wolf 514 (ALTA). Santa Clara Co.:
Loma Prieta, W. Price s.n. (UC); P. Covel s.n. (CAS); Mt. Hamilton, R. Pendleton
s.n. (UC); summit Santa Cruz Mts., R. Pendleton 394 (UC); Alma Soda Springs A.
A. Heller 7490 (UC); Mt. Hamilton, H. Sharsmith 1051 (UC). Santa Cruz Co.: Loma
Prieta Pk., H. Mason s.n. (UC); Eagle Rock, R. Ferri 11114 (UC); Felton, B.
Schreiber 319 (UC); Boulder Creek, 7. Kearny s.n. (CAS); Redwood Park, S. Blake
11766 (WTU). Shasta Co.: Castle Rock, H. Ripley and R. Barneby 9646 (CAS).
Montgomery Creek, E. Bethel s.n. (CAS). Siskiyou Co.: Mt. Eddy, Copeland s.n.
(CAS, MICH, NDG, UC, US); China Creek, S. Fork Salmon River, /. Wiggins
13465 (UC); Mt. Shasta, FE. Palmer 2455 (UC); trail between Taylor and Cow Creeks,
G. Butler 322 (UC); McCloud, A. Eastwood 1105 (CAS, UC); Castle Lake, A.
Eastwood 10719 (CAS). Sonoma Co.: Guernewood Park, H. M. Hall s.n. (UC);
Dohrman Creek, N. E. Jensen’s Ranch, H. Mason 8063 (UC). Tehama Co.: 1.9 mi.
qbove Whitlock Camp, M. Baker and H. Wagon 12833 (UC); Fish Ridge, F.
Hoffman 3531 (UC). Trinity Co.: Morrison Gulch Trail, E. Carter 1094 (CAS);
Ripstein Campground P. Munz s.n. (CAS); 1.5 mi. above Peanut on Hwy. 36, R.
Ferris and L. Lorraine 11693 (UC); White Rock Ranger Station, N. of Yolla Bolly,
A. Alexander and L. Kellogg 5115 (UC); Coffee Creek Canyon, J. T. Howell 1359
(GH, MICH); Dunsmuir Retreat, H. M. Hall and E. B. Babcock 8537, 8539 (UC);
Battle Creek, J. T. Howell 13590 (CAS). Tuolumne Co.: Dodge Ridge, Pine Crest, F.
Hoffman 1764 (UC). Ventura Co.: Ocean View Trail, near Divide Peak, Santa Ynez
Mts., H. Pollar s.n. (CAS).

NEVADA: Washoe Co.: Hunter Creek, P. Kennedy 1869 (US).

OREGON: Curry Co.: Iron Mt., W. Baker 5677 (UC); 3 mi. NE Brookings, M.
Peck 2803 (OSC); head of Lawson Creek, J. Leach 2250 (ORE). Hood River Co.:
base of Mt. Hood, F. Lloyd s.n. (NY); Bald Butte, M. Armstrong 471 (NY); Hood
River, L. F. Henderson 452 (NY); Mt. Defiance Trail, Columbia Gorge, L. Delting
7169 (CAS, ORE); Mt. Hood, near Tollgate, Drake and Dickson s.n. (WTU).
Jackson Co.: Ashland, M. Peck 2795 (OSC); 2395 (OSC). Josephine Co.: Noname
Creek, SW of the Caves, FE. Applegate 11917 (CAS); Siskiyou National Forest, 5 mi
E. Jct. 3941 and 3942, M. Denton 2550 (WTU). Klamath Co.: Crater Lake, A.
Coonebe s.n. (CAS), E. of Lake of the Woods, FE. Applegate 3799 (CAS); Crater
Lake, F. Colville and E. Applegate 332 (US); F. Hummewell 7876 (GH). Wasco Co.:
4 mi. E. Bear Springs, J. Thompson 4935 (CAS, WTU).

WASHINGTON: Klickitat Co.: Falcon Valley, W. Suksdorf 7301 (CAS, NY,
UC); 12047 (CAS, NY, UC, WTU); 16/7 (GH, NY, UC, US).

Arnica discoidea is an extremely variable species and occurs in a
variety of habitats from open chaparral to conifer forests from near
sea level to 1500 m. As early as 1884 Gray recognized the similarity
of this taxon to A. cordifolia and noted that “northwardly it seems
to pass into A. cordifolia.” Indeed, in southerly portions of its
range, A. discoidea is quite distinct and can easily be recognized by

1984] Wolf & Denford — Arnica 277

its narrowly ovate leaves that are mostly crowded toward the base of
the stem; broadly winged and reduced upper leaves; often numer-
ous, narrower heads and frequently branched upper stem. However,
northward it sometimes resembles little more than a rayless A.
cordifolia. Further complicating its identification is the fact that
ampliate marginal disc florets occur infrequently on some specimens
of A. discoidea (e.g., J. H. Thomas 4130 CAS and H. M. Hall 9485
UC). When pressed, these florets resemble rays, causing some
specimens to be erroneously identified as A. cordifolia. However, A.
discoidea and A. cordifolia are distinct with respect to ecology,
geographical distribution, flowering periods and a combination of
several morphological features. A. discoidea occurs in hotter, drier,
more exposed habitats at lower elevations west of the Sierras and
Cascades and flowers 1-2 months later than A. cordifolia.

Based largely on leaf shape and geographical distribution,
Maguire (1943) recognized four taxa within Arnica discoidea sensu
lato: A. grayi, A. discoidea sensu stricto, A. parviflora ssp.
parviflora and A. parviflora ssp. alata. Ediger and Barkely (1978)
recognized the latter three taxa as varieties of A. discoidea: var.
discoidea, var. eradiata (which included A. grayi) and var. alata
respectively. However, since attempts to delimit these taxa pro-
duced no significant systematic correlations between morphology,
chromosome number, geography or flavonoid chemistry (Wolf,
1981; Wolf and Denford, 1984b), A. discoidea is best treated as one
highly polymorphic species with no infraspecific taxa.

4. Arnica gracilis Rydb., Bull. Torrey Bot. Club 24: 297. Arnica
latifolia Bong. var. gracilis (Rydb.) Cronquist, Vasc. P. Pac.
NW 5: 51. 1955. Type: Spanish Peaks (Madison Range,
Montana). 6000 ft., July 14, 1896, J. H. Flodman 901
(HOLOTYPE, NY!; ISOTYPES, NY!, US!).

Arnica columbiana A. Nelson, Bot. Gaz. 30: 200. Type: Columbia Falls,
Montana, 1894, J. J. Kennedy 24 (HOLOTYPE, MONT!).

Arnica multiflora Greene, Pittonia 4: 162. 1900. Type: Lake Pend d’Oreille,
Idaho, June, 1891, J. B. Leiberg 234 (HOLOTYPE, NDG!).

Arnica lactucina Greene, Ottawa Nat. 23: 214. 1919. Type: Hamilton Mt., Banff,
Alberta, 5,800 ft., July 24, 1899, W. C. McCalla 2014 (HOLOTYPE, US!,
ISOTYPES, CU!, NY!).

Arnica betonicaefolia Greene var. gracilis (Rydb.) M. E. Jones, Bull. Univ.
Mont. Biol. 15: 48. 1910.

278 Rhodora [Vol. 86

Arnica puberula Rydb., Fl. Rocky Mts. 979. 1917. Type: Head of Lake Louise,
Alberta, July 22, 1904, J. Macoun (Geol. Surv. Can. No. 65523) (HOLOTYPE,
NY!; ISOTYPES, CAN!, US!).

Stems 3-several branched above, often in dense clumps of 5-10,
10-30 cm high, | mm diameter, mostly short stipitate-glandular
above, becoming glabrate below; rhizomes 1-2 mm broad, covered
with brown scales and old leaf bases; cauline leaves 2-3 pairs, ovate
to ovate-lanceolate, 2-6 cm long, 1-3 cm broad, stipitate-glandular
above, glandular below, acute, irregularly serrate to subentire;
petioles narrow to broadly winged, 1-6 cm long; upper pair of leaves
often reduced and connate-perfoliate; leaves of the innovations
5—14, similar to cauline leaves but narrowly petiolate. Inflorescence
a corymb of (3)5-15 heads or rarely a single head; peduncle 1.5-7
cm long, stipitate-glandular, heads radiate, turbinate-campanulate,
10-15 mm high; involucral bracts 10-16, ovate-lanceolate, 6-12 mm
long, 1-2 mm broad, short stipitate-glandular, acute to acuminate.
Ray florets 5-12, yellow, linear-elliptic, 10-20 mm long, 3-5 mm
broad, 3-dentate; disc florets 10-25, yellow, tubular, 4-6 mm long,
short stipitate-glandular; pappus of both ray and disc florets white,
barbellate. Achenes black, 4.5-7 mm long, | mm broad, short
Stipitate-glandular with few duplex hairs. Figure 7. Chromosome
number 2n = 57, 76.

ECOLOGY AND DISTRIBUTION: Dry, exposed, rocky, alpine slopes
or occasionally sub-alpine meadows in the Rocky Mountains of
Alberta, south irregularly to southern Wyoming, the Uintah
Mountains, of Utah, Wallowa Mountains of Oregon and infre-
quently in the Cascade Mountains of southern British Columbia
south to Mt. Rainier, Washington. Also known from Vancouver
Island. Figure 8. Elevational distribution from 1200-2500 m.
Flowers July-August.

REPRESENTATIVE SPECIMENS: Canada: ALBERTA: Peyto Lake, W. Weber 2445
(GH, UBC, UC); Bertha Lake, A. Breitung 16228 (UC); Lake Louise Trail, Malte
and Watson 164A (UC); Lake Louise, Butters and Holway 336 (GH, US); Tonquin
Valley, Beamish and Pindar- Moss 700151 (UC); Maligne Lake, S. Brown 1287 (GH,
WTU): Lake Louise, S. Brown 703 (GH); Bertha Lake, Gadd and Nagy 3573 (CAN);
Bow Pass, A. E. Porsild and A. J. Breitung 16208 (CAN); Mt. Bertha, Blais and
Nagy 1555 (CAN); Mt. Temple Ski Lodge A. E. Porsild and A. J. Breitung 12722
(CAN); Whitehorse Creek, M. Dumais 5248 (ALTA, CAN); Sunshine Ski Lodge, A.
E. Porsild and J. Lid 19570 (CAN); Mt. Eisenhower, A. E. Porsild and A. J. Breitung
15807 (CAN); Goat Mt., J. Macoun 96043 (CAN); Sofa Mt., A. J. Breitung 17178
(DAO); Chief Mt., A. Breitung 15867 (DAO); Sofa Mt., P. Kuchar 2730 (ALTA);

1984] Wolf & Denford — Arnica 279

A. gracilis Rydb
S.J. Wolf 1989

ORK BOTANICAL GARDEy

Flante of MONTANA

var, gracilis Ryd.) Crong,

t in granitic fill-fields al,
tock Cresk, sbout 2 iiien
ier Lake, about 23 miles

Figure 7. Arnica gracilis.

280 Rhodora [Vol. 86

Figure 8. Distribution of Arnica gracilis.

Bertha Lake, A. J. Breitung 16228 (ALTA); Maligne Lake, H. Raup 2716 (ALTA);
Red Rock Canyon, E. H. Moss 374 (ALTA); Moraine Lake, W. McCalla 4538
(ALTA); Lake Louise, W. McCalla 715] (ALTA); mi. 105, Banff-Jasper Hwy., W.
McCalla 7122 (ALTA).

BRITISH COLUMBIA: Ainsworth, 7. McCabe 5990 (UC); Kinbasket Lake, T.,
McCabe 6275 (UC); between Burton and Fauquier, 7. McCabe 6629 (UC); trail, J.
Macoun 64977 (ND); Kicking Horse Valley, §. Brown 298 (US); Mark Creek Falls,
Kimberley, F. Fedor 80 (UBC); Mt. Matier, Pinder-Moss and Schofield 894 (UBC);
Sage Creek Falls, Bell and Davidson 359 (UBC); Commerce Mt., Bell and Davidson
525 (UBC); Little Diamond Head, Garibaldi Park, E. Peterson s.n. (UBC); Boulder
Creek, F. Fedor 100 (UBC); Vancouver Island, Mt. Arrowsmith, C. Rosendahl 1644
(GH); Mt. Assiniboine, A. E. Porsild 18376 (CAN); Lake Agnes, J. Macoun 14757
(CAN); Mt. Forget-Me-Not, J. Macoun 22829 (CAN); Elk River, S. Elko, J/.

1984] Wolf & Denford — Arnica 28 |

Eastham 15629 (DAO); Flathead Summit, Be/l/ and Davidson 972 (DAO); Emerald
Lake, Yoho Nat. Park, W. McCalla 7032 (ALTA, DAO); Tuktakamin Mt., J. Grant
65-5 (DAO); Vermillion Pass, W. McCalla 2014 (ALTA).

United States: IDAHO: Clark Co.: West Camas Creek, 10 mi. above Kilgore, A.
Cronquist 1385 (ND, NY). Custer Co.: 3.5 mi. SW Stanley Lake, Hitchcock and
Muhlick 9630 (CAS, UC); 11 mi. NE Custer, W. Baker 10790 (OSC); Seafoam,
Macbride and Payson 3653 (RM, US); McKay, Nelson and Macbride 1497 (RM).
Elmore Co.: 10 mi. W. Atlanta, J. and C. Christ 19463 (OSC). Valley Co.: E. side Big
Payette Lake, P. Raven 18517 (CAS).

MONTANA: Carbon Co.: Rock Creek, 23 mi. SW Red Lodge, A. Cronquist 8102
(CAN, CAS, MICH, NY, OSC, WTU). Gallatin Co.: Spanish Basin, P. A. Rydberg
and E. Bessey 5229 (US). Glacier Co.: Glacier Nat'l. Park. B. R. and C. Maguire
15350 (UC). Lincoln Co.: Leigh Lake, Cabinet Mts., D. Woodland 882 (CAS).
Missoula Co.: Holland Creek, Holland Lake, A. Cronquist (CAS, UC); 5 mi. above
Bonner, Hitchcock and Muhlick 11432 (CAS, OSC, UC). Powell Co.: Shaw Creek
R. S., Flathead N. F., C. L. Hitchcock 18436 (WTU). Ravalli Co.: 4 mi. W.
campground, Selway Bitterroot Wilderness, D. Woodland 395 (CAS). Sweetgrass
Co.: 6 mi. E. Box Canyon, Boulder River, C. L. Hitchcock 16439 (UC, WTU).

OREGON: Wallowa Co.: Hurricane Creek, M. Peck 22631 (UC); Boy Scout
Ridge, G. Mason 5981 (ORE, OSC).

UTAH: Summit Co.: 4 mi. N. Hayden Pass, H. Bennett 8431 (CAS).

WASHINGTON: Chelan Co.: Mt. Stuart, A. Kruckeburg 2638 (CAN, UC); Hwy.
2, 6 mi. NW Leavenworth, W. Dress 428] (UC). Kittitas Co.: Cle Elum River, J.
Thompson 10418 (CAS, CAN, WTU). Okanogan Co.: Angels Pass, J. Thompson
(CAS, UC, US, WTU). Pend Oreille Co.: “Z” Canyon, 12 mi. N. Metaline Falls, C.
L. Hitchcock 2923 (UC, WTU). Pierce Co.: Mt. Rainier, H. Leschke s.n. (CAS).
Whatcom Co.: Mt. Hermann, J. Thompson 5352 (GH).

WYOMING: Albany Co.: Medicine Bow Mt., A. and R. Nelson s.n. (RM);
Laramie Creek, A. Nelson 7568 (RM). Carbon Co.: Chimneys of Pedro Mts., L.
Goodding 108 (RM, UC). Fremont Co.: Gannett Peak, F. Jozwik 404 (RM, UC);
Snow Lake, R. Scott 9576 (RM). Johnson Co.: Headwaters of Clear Creek and
Crazy Woman River, F. Tweedy 3015 (RM). Park Co.: Olson’s Meadow, E. and D.
Pearson 172 (RM). Sheridan Co.: Big Horn Mts., J. Williams s.n. (RM), A. Nelson
8501 (RM). Sublette Co.: Fremont Lake, E. and L. Payson 2834 (OSC, RM, UC,
US); canyon above New Fork Lakes, E. and L. Payson 4452 (GH); Horseshoe Lake,
SE Pinedale, C. L. Porter and B. Miller 6069 (RM).

Arnica gracilis occurs on fairly exposed, rocky, alpine slopes or
open sub-alpine meadows largely in the central Rocky Mountains.
The close relationship of this taxon to A. Jatifolia has long been
noted. Indeed, in his original description of A. gracilis, Rydberg
(1897) noted it resembles a depauperate A. Jatifolia. Maguire (1943)
considered A. gracilis a distinct species but called it “... a loose
entity which is maintained as distinct from A. Jatifolia with some
difficulty. ..”. Cronquist (1955) and later Ediger and Barkely (1978)
treated A. gracilis as a variety of A. Jatifolia, while Wolf and
Denford (1984a) recently re-elevated it to specific status.

282 Rhodora [Vol. 86

This confusion with Arnica latifolia is not surprising since Wolf
and Denford (1984a) have demonstrated that A. gracilis is a hybrid
between A. Jatifolia and A. cordifolia. Although it is somewhat
intermediate betwen these two species, A. gracilis has several
distinctive features of its own. These include a much more branched
habit; narrower leaves; more numerous and smaller heads with a
reduced number of disc florets; black, glandular achenes; and the
dry, alpine habitat. A. gracilis is relatively common, is morpho-
logically distinguishable from its two parents and is an autonomous
apomict that maintains itself quite vigorously in nature. It “behaves
as a species” (Davis and Heywood, 1963) and is therefore recognized
as such in the present study.

5. Arnica latifolia Bong., Mem. Acad. St. Petersb. VI 2: 147. 1832.
Arnica latifolia a genuina Herder, Bull. Soc. Nat. Mosc. 40:
424. 1867. Type: Sitka, Alaska, Mertens s.n. (LECTOTYPE by
Maguire, LE photo!).

Arnica menziesii Hook., Fl. Bor. Am. |: 331. 1834. Arnica latifolia B
angustifolia Herder, Bull. Soc. Nat. Mosc. 40: 424. 1867. Type: Northwest
coast of America, Menzies s.n. (HOLOTYPE, K photo!).

Arnica betonicaefolia Greene, Pittonia 4: 163. 1900. Type: Mt. Steele, Olympic
Mountains, Washington, 6000-7000 ft., Aug., 1895, C. V. Piper 2002
(HOLOTYPE, NDG!; ISOTYPE, GH!).

Arnica teucriifolia Greene, Pittonia 4: 164. 1900. Arnica latifolia Bong. var.
teucriifolia (Greene) L. Williams, Leafl. West. Bot. 1: 171. 1935. Type: Grassy
mountain slopes, divide between St. Joe and Clear Water River, alt. 1820 m,
region of the Coeur d’Alene Mountains, Idaho, July 10, 1895, J. B. Leiberg
1229 (HOLOTYPE, US!; ISOTYPES, MO!, NY!, POM!, UC!).

Arnica ovalifolia Greene, Pittonia 4: 168. 1900. TyPE: Big Horn Mountains,
Wyoming, 9000-10000 ft., July 17, 1890, Blankinship s.n. (HOLOTYPE, NDG!).

Arnica ventorum Greene, Pittonia 4: 173. 1900. Type: Union Pass, Wind River
Mountains, Wyoming, Aug. 11, 1894, 4. Nelson 836 (HOLOTYPE, NDG!;
ISOTYPES, MO!, NY!, RM!, ws!).

Arnica grandifolia Greene, Pittonia 4: 173, 1900. Type: Bridger Pass, Montana,
July 28, 1896, J. H. Flodman 896 (HOLOTYPE, NDG!; ISOTYPES, MO!, Us!).
Arnica platyphylla A. Nelson, Bot. Gaz. 31: 407. 1901. Type: moist dark fir
forests, Cascade Mts., Foothills, Hood River, Oregon, July 18, 1896. L. F.

Henderson s.n. (LECTOTYPE by Maguire, RM!).

Arnica laevigata Greene, Ottawa Nat. 15: 279. 1902. Type: by springs in woods,
Chilliwack Valley, B.C., alt. 3000 ft., Aug. 5, 1901, J. M. Macoun (Geol. Surv.
Can. No. 26926) (HOLOTYPE, NDG!; ISOTYPES, CAN!, NY!).

Arnica aprica Greene, Ottawa Nat. 15: 280. 1902. Type: open ground along
streamlets, Chilliwack Valley, B. C., alt. 3500 ft., July 10, 1901, 7. M. Macoun
(Geol. Surv. Can. No. 26284) (HOLOTYPE, NDG, photo uc!; IsoTypEs, CAN!,
No!, Ny!). The holotype is listed at NDG but has not been located by the staff.

1984] Wolf & Denford — Arnica 283

Arnica jonesii Rydb., Fl. Rocky Mts., 979. 1917. Type: Alta, Wasatch
Mountains, Utah, July 31, 1879, M. E. Jones 1119 (HOLOTYPE, NY!; ISOTYPES,
NY!, PoM!, uUTCc!).

Arnica eriopoda Gandoger, Bull. Soc. Bot. Fr. 65: 38. 1918. Type: Cascade
Mountains, Oregon, July 27, 1902, W. C. Cusick 2914 (HOLOTYPE, US photo!;
ISOTYPES, MO!, NY!, POM!, UC!).

Arnica aphanactis Piper, Proc. Biol. Soc. Wash. 33: 105. 1920. Type: Mt. Baker,
Washington, 1915, G. W. Turesson s.n. (HOLOTYPE, Us!).

Arnica flodmanii Rydb., N. Am. Fl. 34: 334. Type: Spanish Peaks, Madison
Range, Montana, July 14, 1896, /. H. Flodman 898 (HOLOTYPE, NY!; ISOTYPES,
mo!, NyY!, us!).

Arnica glabrata Rydb., N. Am. Fl. 34: 335. Type: Crater Lake, Oregon, August
1898, Austin and Bruce 1627 (HOLOTYPE, NY!).

Arnica paucibracteata Rydb., N. Am. FI. 34: 336. 1927. Type: Medicine Bow
Mountains, Wyoming, Aug. 3, 1900, A. Nelson 794] (HOLOTYPE, NyY!;
ISOTYPES, MO!, POM!, RM!, Us!).

Arnica oligolepis Rydb., N. Am. Fl. 34: 336. 1927. Type: Hazelton, Skeena
River, B. C., June 23, 1917, J. M. Macoun (Geol. Surv. Can. No. 96048)
(HOLOTYPE, NY!; ISOTYPE, CAN!).

Arnica leptocaulis Rydb., N. Am. FI. 34: 336. 1927. Type: Mt. Mark, Vancouver
Island, B. C., July 25, 1887, J. M. Macoun s.n. (HOLOTYPE, NY!; ISOTYPES,
CAN!, US!).

Arnica membranacea Rydb., N. Am. FI. 34: 338. 1927. Type: Wimmer, Jackson
Co., Oregon, June 13, 1892, E. W. Hammond 231 (HOLOTYPE, NY!; ISOTYPES,
us!, ws!).

Stems simple, sometimes sparsely branched above, 10-50 cm
high, 2-3 mm diameter, glabrate to villous throughout; rhizomes
1-3 mm thick, giving rise to several basal rosettes and flowering
stems, rhizomes with several thin brown scales, frequently covered
with old leaf bases at the summit. Cauline leaves 2—4(6) pairs,
mostly sessile to sub-sessile, the lower rarely petiolate, ovate to
elliptic-lanceolate, 2-10 cm long, 1-6 cm broad, glabrous to very
sparsely villous, obtuse to acute, serrate to dentate; lower leaves
sometimes reduced and short-petiolate, the petioles 5-15 mm long,
broadly winged; leaves of the innovations 2-10, similar to cauline
leaves, petiolate, the petioles 2-10 cm long. Inflorescence usually a
single head or corymb of 3-5(9) heads; peduncle 3-25 cm long,
glabrate to sparsely villous above; heads radiate, narrowly turbinate,
8-20 mm high; involucral bracts 8-20, lanceolate to oblanceolate,
8-15 mm long, I-3 mm broad, sparsely villous and glandular, acute
to acuminate. Ray florets 8-15, yellow, oblong-linear, 10-25 mm
long, 2-6 mm broad, 3-dentate; disc florets 20-90, yellow, tubular,
6-10 mm long, sparsely villous; pappus of both ray and disc florets
white, barbellate. Achenes dark brown, 5-9 mm long, | mm broad,

284 Rhodora [Vol. 86

sparsely short stipitate-glandular with few duplex hairs. Figure 9.
Chromosome number 2n = 38, 76.

ECOLOGY AND DISTRIBUTION: Common in relatively moist,
montane Picea-Abies forests, or more commonly sub-alpine mead-
ows from southern interior and coastal Alaska south along the coast
and through the Cascades to northern California, and south in the
Rocky Mountains from the Yukon through southern Colorado.
Also common on Vancouver Island and the Queen Charlotte
Islands. Figure 10. Elevational distribution from 500-3300 m.
Flowers July-August.

REPRESENTATIVE SPECIMENS: Canada: ALBERTA: Lake Louise, W. Serchell sn.
(UC); Banff-Jasper Hwy., W. A. Weber 2490 (UC); Banff, H. Davis s.n. (ND); Mt.
Edith Cavell, 7. McCabe 8355 (UC); Sheep Mt., J. Macoun s.n. (MICH); Vermillion
Pass, R. Ogilvie s.n. (UBC); Castlemont, M. Malte and W. Watson 488 (CAN); Mt.
Temple, A. E. Porsild and A. J. Breitung 13753 (CAN); Bow River Pass, A. E.
Porsild and A. J. Breitung 14929 (CAN); tower, Waterton Nat. Park, W. Blais and J.
Nagey 1639 (CAN); Snowshoe Cabin, Waterton Nat. Park, G. Armstrong and J.
Nagey 4554 (CAN); Crandell Lake, G. Armstrong and J. Nagey 3874 (CAN);
Cameron Lake, G. Armstrong and J. Nagey 4094 (CAN); Crows Nest Lake, Dawson
14756 (CAN); Red Rock Canyon, F. Sudol 43 (DAO), Mt. Rowe, A. J. Breitung
16979 (DAO); Bow Pass, W. McCalla 6771 (ALTA); Lake Agness, W. McCalla 3730
(ALTA); Bald Hills, P. Kuchar 521 (ALTA); Marmot Mt., W. McCalla 3105
(ALTA),

BRITISH COLUMBIA: E. end Summit Lake, Calder and Saville 10015 (UC); Mt.
Fougner at Bella Coola, Calder and Saville 20373 B (UC); Khutze Inlet, 7. McCabe
3480 (UC); Harrison Creek, 20 mi. N. Takla Landing, T. McCabe (UC); Nine Mile
Mt., 7. McCabe 8181 (UC); Mt. Revelstoke, T. McCabe 5392 (UC); Apex Mt., R.
Bowerman s.n. (UC); Green Mt., near Haylmore, J. and E. Thompson 691 (MICH);
Mt. Selwyn, H. Raup and E. Abbe 4164 (CAS); Emerald Lake, C. Shaw 993 (US);
Ft. St. James, Calder and Saville 13720 (DAO, US); Dam Mt., W. Taylor 5993
(UBC); Grouse Mt., V. Krajina s.n. (UBS); 44 mi. NNW Dease Lake, S. MacDonald
SII (CAN, UBC); 12 mi. NE Smithers, V. Krajina et al s.n. (UBC); Goodchap Mt.,
D. Martin s.n. (UBC); Gold Fish Lake, A. Szczawinski 174 (UBC); Duckling Creek,
Germansen Landing, G. Noel 158 (UBC); Mt. Semour, V. Krajina 333 (UBC);
Beatton River, H. M. Raup and D. Correll 10066 (GH); Glacier Nat. Park, E. Haber
1508 (CAN); mi. 85 Haines Rd., C. Clarke 442 (CAN); White Pass, M. Malte 364
(CAN); Tunjony Lake, R. Pilfrey 21 (DAO); Yanks Peak, Calder et al 18083 (DAO);
Alpine Mt., 12 mi. NNE Nelson, Calder and Saville 11129 (DAO); Red Rose Mine,
Calder and Saville 15190 (DAO); Mt. McLean at Lillooet, Calder and Saville 15505
(DAO); 75 mi. S. Haines Jct., Calder and Kukkonen 28154 (DAO); lake in Coast
Range, 58°41’N, 133°04’W, R. Pilfrey 79 (DAO). Queen Charlotte Islands: 20 mi. S.
Morseby Logging Camp, Calder et al 23046 (CAS, DAO, OSC, UBC, UC); Lake
Takakia, Calder and R. Taylor 36296 (DAO). Vancouver Island: Green Mt., V.
Krajina et al 5004 (UBC); Mt. Arrowsmith, G. Allen s.n. (UBC); Shaw Creek, W.
Spreadborough 96037 (CAN); Moat Lake, J. Underhill 327 (DAO); Crest Lake, A.
Young and W. Hubbard 580 (DAO).

1984] Wolf & Denford — Arnica 285

Figure 9. Arnica latifolia.

[Vol. 86

Rhodora

286

6 Ue Ch

Cpa

Distribution of Arnica latifolia.

Figure 10.

1984] Wolf & Denford — Arnica 287

NORTHWEST TERRITORIES: 62°46’N, 129° 1’W, L. Allison 29 (DAO).

YUKON TERRITORY: White Pass, A. Eastwood 936 (UC); mi. 268, Canol Rd.,
A. E. Porsild and A. J. Breitung 11374 (CAN, UC, US); mi 95, Canol Rd., A. E
Porsild and A. J. Breitung 10228 (CAN, GH, UC, US, WTU); Upper Hyland Lake,
Calder and Kukkonen 27809 (CAS, DAO, GH); Kluane National Park, Alsek River,
G. and G. Douglas 8953 (DAO); Cassiar Mts., W. Poole 49 (DAO); mt. 4 mi. W.
Upper Hyland Lake, Calder and Kukkonen 27909 (DAO); 62°11’N, 129°17’W, L.
Allison 40 (DAO).

United States: ALASKA: Hatcher Pass, S. J. Wolf 503 (ALTA); Craig, /. Norberg
s.n. (UC, US); Yakutat Bay, F. Funston 79 (CAN, ND); Olga Bay, Kodiak Island, S.
Blake 553 (ND); Deer Mt., F. Went 80 (UC); Yes Bay, T. Howell 1634 (UC); Harris
Peak, Prince of Wales Island, D. Jaques 1569 (OSC); Indian River, Sitka, L. Smith
s.n. (OSC); Duchess Mine, Latouche Island, H. Shacklette 4704 (MICH); Cairn
Ridge, near Juneau, H. Shacklette (MICH); Curry Lookout, L. Jordal 2555
(MICH); Mt. Roberts, Juneau, M. Williams 1392 (OSC); Kenai Lake, J. Calder 6089
(CAS, DAO), Kuiu Island, £. Walker 774 (CAS, US); 3 St’s. Bay, Kodiak Island, W.
Eyerdam 602 (CAS, US); Eyak Lake, Cordova, M. Hanna s.n. (CAS); Skagway, A.
Eastwood 818 (CAS, US), Haines, E. Scheuber s.n. (US); Sitka, C. Piper 4245 (US),
Stikine Glacier, W. Cooper 72 (US); Mt. Marathon, J. Calder 5899 (UBC); Juneau,
E. Scamman 1123 (GH); Mt. Roberts, A. and R. Nelson 4440 (GH); Mt. Crillion, R.
Bates 160 (GH), Barren Island, /. Gabrielson s.n. (GH); Alaska Range, 62°40’N,
152°30’°W. L. Viereck 5259 (CAN); Ketchican, J. Anderson 24691 (CAN); Orca, 1.
Norbert s.n. (CAN); Burma Rd., mi. 74, Richardson Hwy., L. Spetzman 3319
(CAN); Palmer Creek Valley, SE Hope, J. Calder 6233A (DAO); Chugach Mts.,
Anchorage, Dutilly et al 21143 (DAO).

CALIFORNIA: El Dorado Co.: S. side Echo Lake, A. Heller 12544 (UC); Sugar
Bowl Mt., L. Kildale s.n. (UC). Nevada Co.: Donner Pass, A. Heller 7029 (MICH).
Trinity Co.: Salmon Mts., Union Creek, H. M. Hall 9648 (UC). Siskiyou Co.:
Jackson Lake, A. Alexander and L. Kellogg 183 (UC); Bolan Lake, C. L. Hitchcock
and J. Martin 5233 (UC), S. J. Wolf 465 (ALTA); Hancock Lake Trail, Marble Mt.
Wilderness, F. Oettinger 460 (UC). Caribou Basin, Trinity Alps, J. Howell 13380
(CAS); 8.3 mi. from Etna on road to Sayer’s Bar, F. Balls 13942 (WTU), S. J. Wolf
467 (ALTA).

COLORADO: Chaffee Co.: Monanos Creek, /. Clokey 3500 (RM, UC). Clear
Creek Co.: Loveland Pass, S. J. Wolf 416 (ALTA). Gunnison Co.: Queen’s Basin, J/.
Langenheim 3908 (RM, UC). Lake Co.: Lake Creek, /. Clokey 3515 (UC). Larimer
Co.: Rocky Mt. National Park, Rainbow Curve Trail, U. Waterfall 14958 (UC),
Lock Vale, Estes Park, J. Clokey 3962 (CAN, MICH, UC); Cameron Pass, G.
Osterhout 3795 (RM); Lake Helene Trail, Rocky Mt. National Park, R. Ashton 70-
g-7 (RM). Routt Co.: Hahn’s Peak, W. Weber 6929 (RM, WTU).

IDAHO: Bear Lake Co.: Bloomington Lake, R. Davis 1613 (UC). Benewah Co.:
Bald Mt., W. Baker 13388 (OSC). Blaine Co.: 5 mi. from Alturas Lake, C. L.
Hitchcock and C. V. Muhlick 10534 (UC). Boise Co.: Jackson Peak, C. L. Hitchcock
and C. V. Muhlick 10026 (CAN, UC). Bonner Co.: Queen Mt., W. Eggleston 9770
(US). Bonneville Co.: E. Payson and G. Armstrong 3511 (RM). Custer Co.: | mi. E.
Castle Peak, C. L. Hitchcock and C. V. Muhlick 10913 (UC); Mt. Mogg, C. L.
Hitchcock and C. V. Muhlick 11236 (UC, WTU); 10 mi. S. Atlanta, C. L. Hitchcock

288 Rhodora [Vol. 86

and C. V. Muhlick 10247 (RM, UC, WTU). Franklin Co.: 2 mi. SW Franklin Basin
R.S., B. Maguire 21643 (CAN, UC). Fremont Co.: Ponds Lodge, Targhee N. F., W.
Baker 9831 (OSC). Idaho Co.: Lolo Trail, 65 mi. E. Pierce, C. L. Hitchcock and C.
V. Muhlick 21921 (UC); Pilot Knob, W. Elwood 36 (UC); Burnt Knob Lookout, W.
Baker 12638 (OSC). Lemhi Co.: 9 km S. Lost Trail Pass, S. J. Wolf 488 (ALTA).
Shoshone Co.: St. Maries River, G. Wilson 103 (UC); 14 mi. E. Clarkia, W. Baker
16208 (WTU). Valley Co.: Lick Creek Summit, S. J. Wolf 357 (ALTA); Gold Fork
Lookout, J. Thompson 13745 (MICH, UC); Upper Payette Lake, W. Baker 10357
(OSC).

MONTANA: Beaverhead Co.: Pintlar Falls, C. L. Hitchcock and C. V. Muhlick
12784 (UC); Lake Waukena, C. L. Hitchcock and C. V. Muhlick 13105 (OSC).
Deerlodge Co.: Storm Lake, C. L. Hitchcock and C. V. Muhlick 14830 (RM); Storm
Lake Rd., S. J. Wolf 435 (ALTA). Fergus Co.: Big Snowy Mts., 31 mi. SW Lewiston,
G. and F. Ownby 2417 (RM, UC). Flathead Co.: Bowman Lake, R. Turley 212 (UC).
Gallatin Co.: 8 mi. E. Eldridge, C. L. Hitchcock and C. V. Muhlick 15149 (UC).
Glacier Co.: Glacier National Park, N. Carlson s.n. (UC). Granite Co.: Burnt Fork
Trail, C. L. Hitchcock and C. V. Muhlick 14502 (WTU). Lake Co.: 10 mi. NE
Polson, J. Thomas 11051 (CAS). Lewis and Clark Co.: 25 mi. NW Agusta, C. L.
Hitchcock 17997 (RM, UC). Lincoln Co.: Mt. Marston Rd., S. J. Wolf and P. and
D. Wolf-Thompson 344 (ALTA) Madison Co.: Upper Brandon Lake, C. L.
Hitchcock 17045 (RM). Missoula Co.: 2 mi. E. Holland Lake, C. L. Hitchcock 18357
(UC, WTU). Park Co.: 5 mi. E. Cooke City, J. Witt 1755 (WTU). Powell Co.: 3 mi.
W. Big Salmon Lake, C. L. Hitchcock 17162 (WTU). Ravalli Co.: St. Mary’s Creek,
C. L. Hitchcock and C. V. Muhlick (CAN, UC). Stillwater Co.: Mt. Haystack, C. L.
Hitchcock and C. V. Muhlick 13429 (CAN, OSC, UC, WTU). Sweetgrass Co.: Crazy
Mts., Big Timber Creek, C. L. Hitchcock and C. V. Muhlick 13288 (OSC, UC).

OREGON: Clackamas Co.: SW. slope Mt. Hood, H. and J. Thomas 248 (UC).
Clatsop Co.: Saddle Mt., S. J. Wolf 379 (ALTA); Onion Peak, L. Heckard 1606
(UC), K. Chambers 3149 (CAS, OSC, WTU); Sugarloaf Mt., K. Chambers 3764
(OSC, WTU). Curry Co.: Iron Mt., S. J. Wolf 457 (ALTA); above Agness, FE.
Applegate 7153 (CAS). Douglas Co.: Black Rock Lookout, D. Overlander s.n. 1944
(OSC). Grant Co.: Strawberry Mt., W. Cusick 3565 (WTU). Harney Co.: Stein’s Mt.,
P. Train s.n. (OSC). Hood River Co.: Mt. Hood, P. Munz 1446] (UC). Jackson Co:
Mt. Ashland, M. Peck 2934 (OSC). Jefferson Co.: J. Johnson 462 (OSC). Josephine
Co.: Big Meadow, SE Oregon Caves, E. Applegate 11243 (UC); Bolan Lake, J.
Thompson 12510 (UC, WTU); Sexton Mt., L. Savage s.n. (UC), Lane Co.: Fairview
Mt., L. Constance s.n. (UC); Horse Pasture Mt., M. Peck 23841 (OSC); Fairview
Mt., W. Baker 5565 (OSC, WTU). Linn Co.: Breitenbush, M. Peck 18718 (UC); Mt.
Jefferson, M. Peck 9109 (OSC); Monument Peak, A. Aller 812 (OSC). Marion Co.: |
mi. E. Breitenbush, M. Peck 18718 (OSC); House Mt., M. Peck (OSC). Union Co.:
Anthony Creek, Blue Mts., W. Cusick 3820 (WTU). Wasco Co.: 5 mi. W. Mosier, J.
Thompson 4224 (WTU). Washington Co.: Tillamook Burn, N. of Wilson River
Hwy., K. Chambers 4052 (OSC).

UTAH: Cache Co.: Tony Lake, A. Holmgren and C. Biddulph 8172 (UC); Mt.
Naomi, B. Maguire et al 14154 (GH, UC, US). Duchesne Co.: Blind Stream Rd., NW
Hanna, S. J. Wolf 397 (ALTA). Salt Lake Co.: Big Cottonwood Canyon, W. Cooper
329 (RM); A. Garrett 1509 (RM). Summit Co.: Stillwater Ford, Uintah Mts., E. and
L. Payson 4995 (RM). Utah Co.: Mt. Timpanogos, B. Maguire 17507 (UC).

1984] Wolf & Denford — Arnica 289

WASHINGTON: Chelan Co.: Mt. Stuart, J. Thompson 7685 (CAS, UC).
Clackamas Co.: Mt. Hood, J/. Thompson 3403 (WTU). Clallam Co.: Hurricane
Ridge, W. and M. Muenscher 10004 (UC); Mt. Angeles, J. Thompson 7522 (CAS,
UC, WTU). Columbia Co.: 1.5 mi. E. Table Rock, Umatilla N. F., A. Kruckenberg
2514 (UC). Grays Harbor Co.: Colonel Bob L. O., J. Thompson 7245 (WTU).
Jefferson Co.: Olympic Mts., Mt. Constance, R. Rollins and T. Chambers 2654
(UC). King Co.: Goldmeyer Hot Springs, J. Broadbent s.n. (WTU). Kittitas Co.:
Mission Peak, J. Thompson 14913 (CAS, MICH, UC, WTU). Klickitat Co.: Mts.
NE Bingen, W. Suksdorf 2760 (WTU). Mason Co.: Mt. Ellinor, W. Eyverdam 1276
(UC). Pierce Co.: Yakima Park, Mt. Rainier N. P., B. Maguire 17260 (UC); Mt.
Rainier, L. Benson 2337 (UC); Chinook Pass, W. Eyerdam s.n. (UC); Cowlitz Pass,
J. Thompson 11102 (CAS, WTU). Skamania Co.: Big Lava Beds, J. Franklin 448
(OSC); Mt. St. Helens, F. Coville 747 (US, WTU). Snohomish Co.: Mt. Pugh, J.
Thompson 14351 (CAS, MICH, UC). Whatcom Co.: Mt. Baker, W. Muenscher 8030
(UC). Yakima Co.: Chinook Pass, J. Thompson 15136 (CAS, MICH, UC, WTU);
Mt. Aix, J. Thompson 15016 (CAS, MICH, UC, WTU).

WYOMING: Albany Co.: 7.4 mi. W. Centennial, S. J. Wolf 422 (ALTA). Lincoln
Co.: Jackson’s Hole, E. and L. Payson 2276 (UC). Sublette Co.: 26 mi. W. Big
Pinney, F. and L. Meyer 2369 (UC). Teton Co.: Skyline Trail, Teton N.P., L.
Wehmeyer et al 5450 (MICH); | mi. E. Togwotee Pass, S. J. Wolf 431 (ALTA).
Yellowstone National Park: Obsidian Creek, A. and E. Nelson 6108 (UC).

Arnica latifolia is one of the most polymorphic and widely
distributed of western arnicas. This taxon is common in relatively
cool, montane Picea-Abies forests or sub-alpine meadows from
Alaska through Colorado and northern California. In its most
typical form A. Jatifolia is easily recognized by its sessile, ovate,
glabrous leaves; very narrow heads with narrow phyllaries and rays;
and glabrous, brown achenes. However, both environmentally
induced morphological variability and plants with petiolate lower
cauline leaves are sometimes encountered. Consequently, this taxon
is sometimes confused with both A. cordifolia and A. gracilis.
Plants of shaded forests represent the typical form of the species
while plants of more exposed areas are usually much reduced; have
thicker, smaller, more glandular leaves; have broader heads with
more glandular phyllaries and are often confused with A. gracilis.
However, they are readily separable from the latter by their broader,
sessile, petiolate leaves and fewer heads. Pressed specimens of A.
latifolia with petiolate lower leaves are sometimes confused with A.
cordifolia; however, they can be readily distinguished by leaf shape,
margin and pubescence, head and phyllary shape and achene color.

The type sheet of Arnica betonicaefolia consists of two collec-
tions: C. V. Piper 2202 and 2002. Both were collected on Mt. Steele,
Washington, in August, 1895; however, the former was collected at

290 Rhodora [Vol. 86

7000 ft. while the latter was collected at 6000 ft. Maguire (1943)
reduced A. betonicaefolia to synonymy under A. /atifolia and cited
2002 as the holotype while Ediger and Barkley (1978), who reduced
this taxon under A. gracilis, cited 2202. Both collections represent
the reduced high alpine form of A. /atifolia and have ovate, serrate,
sessile leaves and solitary narrow heads typical of this taxon. There
is really no question as to which collection represents the holotype
of A. betonicaefolia since Greene (1900) clearly designated 2002.

6. Arnica nevadensis A. Gray, Proc. Am. Acad. 19: 55. 1883. TYPE:
Lassen’s Peak, California, R. M. Austin s.n. (LECTOTYPE by
Rydberg, GH!; SYNTYPE, Summit Valley, California, Sept. 25,
1882, C. G. Pringle s.n. NY!).

Arnica tomentella Greene, Pittonia 4: 166. 1900. Type: open woods in Middle
Tule River, California, alt. 5500 ft., April-Sept. 1897. C. A. Purpus 5625
(HOLOTYPE, US!; ISOTYPES, GH!, MO!, UC!).

Stems simple, 10-50 cm high, 1.5-2.5 mm diameter, short
stipitate-glandular throughout and puberulent above, rhizomes 1-2
mm thick, with several brown scales and old leaf bases at the
summit. Cauline leaves 2-3 pairs, ovate to elliptic, 3-8 cm long, 2-4
cm broad, short stipitate-glandular throughout, acute to rounded,
entire to denticulate; petioles narrow to broadly winged, |.5-4.0 cm
long; upper pair of leaves sometimes reduced, sessile and lanceolate;
leaves of the innovations 4—6, similar to cauline leaves. Inflorescence
a single head or corymb of 3 heads, peduncle 4-15 cm long,
stipitate-glandular and somewhat villous, heads radiate, campan-
ulate-turbinate, 15-20 mm high; involucral bracts 10-16, oblance-
olate, 10-17 mm long, 2-4 mm broad, stipitate-glandular, acute to
acuminate. Ray florets 6-14, yellow, linear to broadly elliptic, 15-25
mm long, 4-6 mm broad, 3-dentate; disc florets 20-60, yellow,
tubular, 8-10 mm long, short stipitate-glandular; pappus of both
ray and disc florets white to tawny, barbellate to subplumose.
Achenes dark gray, 6-9 mm long, | mm broad, stipitate-glandular
throughtout. Figure 11. Chromosome number 2n = 76.

ECOLOGY AND DISTRIBUTION: Relatively uncommon in fairly dry
Tsuga- Pinus forests or exposed rocky slopes of the Sierra Nevada
Mountains from south of Yosemite National Park, California and
adjacent Nevada, northward irregularly to the north Cascades and
Olympic Mountains of Washington. Also known from the Ruby
Mountains of Nevada. Figure 12. Elevational distribution 1500-
3000 m. Flowers July-August.

1984]

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Wolf & Denford — Arnica

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Lectotype of Arnica nevadensis.

292 Rhodora [Vol. 86

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Figure 12. Distribution of Arnica nevadensis.

1984] Wolf & Denford — Arnica 293

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Truckee River, above
Shingle Mill, C. Sonne s.n. (UC); open moist woods, middle Tule River, C. Purpus
1540 (UC); Coburn Mill, 7. Brandegee s.n. (UC); Middle Tule River, C. A. Purpus
5625 (GH, UC, US). Amador Co.: G. Hansen 416 (UC). Butte Co.: Jonesville, A.
Heller 12861 (OSC). El Dorado Co.: Fallen Leaf Lake, G. L. Stebbins 2032 (UC);
Angora Peaks, H. M. Hall 8796 (UC); Velma Lakes Trail, above Eagle Lake, G.
Robbins 2054 (UC); Red Peak, G. Robbins 1806 (CAS, UC); Fallen Leaf Lake, M.
Baker s.n. (UC). Fresno Co.: Vidette Meadows, J. Howell 24955 (CAS); Fish Camp,
P. Raven 5786 (CAS); Mills Creek, P. Raven 5674 (CAS); Kip Camp, P. Raven 7361
(CAS); Pocket Meadow, P. Raven 6006 (CAS). Glenn Co.: Black Butte, J. Howell
19250 (CAS). Inyo Co.: Flower Lake, S. Austin 558 (UC); Big Pine Lakes, J. Howell
23815 (CAS); Box Lake, J. Howell 22445 (CAS); Rock Creek Lake, J. Howell s.n.
(CAS), Madera Co.: Garnet Lake, J. Howell 16479 (CAS); Shadow Lake, P. Raven
3390 (CAS); Lake Ediya, P. Raven 3527 (CAS). Mariposa Co.: Yosemite Park:
Glacier Point, H. M. Hall 9149 (UC); Fletcher Lake, B. Schreiber 2013 (UC);
Donohue Pass, B. Schreiber 1787 (UC); Mt. Dana, H. M. Hall and E. B. Babcock
3599 (UC); Lost Lake, H. M. Hall 9068 (UC). Mono Co.: Mt. Lyell, A. Hawbecker
s.n. (UC); Tiogo Crest, H. Mason 11469 (UC); Harvey Monroe Hall Natural Area, J.
Clausen 1124 (UC), P. Stockwell 1346 (CAS); Sonora Pass, A. Eastwood and J.
Howell 7565 (CAS). Nevada Co.: Stanford Peak, A. Kellogg s.n. (UC); Summit-
Soda Springs, Kennedy and Doten 274 (UC); Donner Pass, J. Howell 18662 (CAS);
S. side Donner Lake, A. Heller s.n. (CAS). Placer Co.: Mt. Anderson, C. Sonne s.n.
(CAS, UC); Truckee River, C. Sonne 7 (CH, UC); Truckee, C. Sonne 27 (GH).
Plumas Co.: Gold Lake Road, J. Ewan 8206 (UC). Shasta Co.: Helen Mt., G. Gillett
1078 (CAS); Lassen Peak, R. M. Austin s.n. (GH); Lassen National Park, F.
Hermann 11956 (UC). Sierra Co.: Gold Lake, H. Baker 82 (CAS). Siskiyou Co.:
Taylor Lake, D. Barbe 129 (UC); Medicine Lake, H. Baker 502 (UC); Upper English
Lake, F. Oettinger 1082 (UC); Wildcat Peak, Alexander and Kellogg 204 (UC);
Marble Mt., H. Chandler 1615 (CAS). Tehma Co.: Brokeoff Mt., G. Gillett 1066
(CAS). Tuolumne Co.: Elizabeth Lake, H. Mason 692 (UC); Johnson Peak, C.
Sharsmith 217 (UC); peak between Matterhorn and Whorl Mts., C. Sharsmith 3826
(UC). Trinity Co.: Grizzly Creek, Trinity Alps, E. Carter 10/1 (CAS).

NEVADA: Elko Co.: Three Lakes, Ruby Mts., A. Borell s.n. (UC). Clover Mts.,
near Deeth, A. Heller 9242 (UC). Ormsby Co.: Fall Creek, C. Baker 1432 (CAS,
MICH, RM, UC, US). Washoe Co.: White Creek, E. slope Mt. Rose, P. Train 4420
(UC).

OREGON: Douglas Co.: Mt. Bailey, C. Geddes 18670 (OSC). Klamath Co.:
Crater Lake National Park: Union Peak, J. Simpson 9 (UC); Garfield Peak, W.
Baker 6184 (OSC, UC); E. Applegate 9851 (CAS, OSC); W. Baker 7203 (WTU);
Hillman Peak, W. Baker 6401 (UC); H. Sullen s.n. (OSC). Lane Co.: N. Sister Mt.,
M. Peck 14473 (CAS, OSC).

WASHINGTON: Chelan Co.: Nason Creek valley between Berne and Cascade, /.
Otil s.n. (CAS). Clallam Co.: Mt. Angeles, J. Flert 3325 (US). Jefferson Co.: Marmot
Pass, Olympic National Forest, J. Thompson 9907 (WTU).

Arnica nevadensis occurs mostly at high elevations in open
Tsuga- Pinus forests or open rocky slopes of the Sierra Nevada and

294 Rhodora [Vol. 86

sparingly northward in the Cascades and eastward into Nevada. In
the southern part of its range this species is quite distinct and readily
recognized by its entire, elliptic to ovate leaves; oblanceolate
phyllaries; white-tawny, barbellate-subplumose pappus and rela-
tively open, high altitude habitat. However, in the northern part of
its range it is often confused with dwarf, high altitude forms of A.
cordifolia. However, the entire leaves, darker pappus with longer
seta and narrower heads of A. nevadensis distinguish it from the
latter.

In his original description of Arnica nevadensis, Gray (1883) did
not designate a type; however, he cited two specimens he had
examined: R. M. Austins.n., Lassen’s Peak, California (GH) and C.
G. Pringle s.n. 1882, Summit Valley, California (NY). In his Flora
of North America, Rydberg (1927) designated the first specimen
cited (Austin) as the lectotype for this species. This choice was also
later accepted by Maguire (1943). However, recently Ediger and
Barkely (1978) rejected Rydberg’s choice as arbitrary and desig-
nated the second specimen cited (Pringle) as the lectotype “... in
order to preserve the traditional application of the name.”
According to the rules of the International Code of Botanical
Nomenclature (Stafleu et a/, 1978) this practice cannot be accepted
without proper justifiction. Both Article 8 and the Guide to the
Determination of Types specifically state that the first choice of a
lectotype must be followed by subsequent workers unless it can be
shown that the choice was based on a misinterpretation of the
protologue or if the choice was made arbitrarily and without
understanding the group concerned.

Ediger and Barkley (1978) based their decision on Recommenda-
tion 7B which states ‘Whenever the elements on which the name of
a taxon is based are heterogeneous, the lectotype should be be
selected as to preserve current usage ...” Although poorly pressed,
the ovate, entire leaves, as well as the broad rays and tawny,
subplumose pappus of the Austin specimen are typical of A.
nevadensis. Further, it seems clear that Gray’s (1883) description
was based on the Austin specimen. He made specific reference to the
cinereous color in both the type description and the discussion of
the Austin specimen, while noting that the Pringle specimen was a
“greener form”. Although Rydberg (1927) gave no reason for his
choice of the Austin specimen at the Gray Herbarium, he must have
given it close examination since it would have been much easier for
him to cite the Pringle specimen at the New York Botanical Garden.

1984] Wolf & Denford — Arnica 295

Rollins (1972) has stressed the importance of selecting a lectotype
from the institution where the author worked. Since both specimens
cited by Gray (1883) are referable to A. nevadensis, but the first
cited (Austin s.n.) was originally chosen as the lectotype by Rydberg
(1927) and later accepted by Maguire (1943), it must be retained as
the lectotype for this species.

Both Maguire (1943) and Ediger and Barkley (1978) recognized
the rare Arnica tomentella of the Sierra Nevada. Maguire (1943)
considered A. tomentella a close relative of A. nevadensis. This
decision is not surprising since even a casual comparison of his
(Maguire, 1943) description of the two taxa indicated they are
nearly identical in most respects including: stem pubescence, leaf
shape and margin, head shape, phyllary and ligule shape and all
characters of the pappus. Only the taller stature and tuft of hairs on
the phyllary tips distinguish A. tomentella from A. nevadensis. A.
nevadensis is an apomictic, polyploid complex (Barker, 1967;
Straley, 1980; Wolf, 1980) and the form previously recognized as A.
tomentella probably represents an apomictic microspecies of the
former.

Three specimens (J. P. Tracy 19273 UC, C. F. Sonne s.n. UC.
#193450 and G. D. Butler 643 UC) previously recognized as Arnica
tomentella are A. cordifolia. Other specimens previously recognized
as A. tomentella (Lemmon s.n. UC #337194; C. F. Sonne s.n., June
6, 1886, GH, UC; C. A. Purpus 1540 UC; T. S. Brandegee s.n. UC
#91026; C. A. Purpus 5625 GH, MO. UC, US) are all treated here as
A. nevadensis. Another collection previously identified as A.
tomentella (Bolander 4937 UC) is A. mollis Hook.

7. Arnica spathulata Greene, Pittonia 3: 103. 1896. Type: Glendale,
Oregon, June 30, 1887, 7. Howell s.n. (HOLOTYPE, NDG!;
ISOTYPES, CAN!, US!).

Arnica eastwoodiae Rydb., N. Am. FI. 34: 343. 1927. Arnica spathulata Greene
subsp. eastwoodiae (Rydb.) Maguire, Brittonia 4: 458. 1943. Arnica spathu-
lata Greene var. eastwoodiae (Rydb.) Ediger and Barkley. N. Am. FI. II. 10:
43. 1978. Type: Gasquet, French Hill, Del Norte Co., California, Sept, 14,
1912, A. Eastwood 221 (HOLOTYPE, NY!; ISOTYPES, NY!, US!).

Arnica cusickii Rydb., N. Am. Fl. 34: 343. 1927. Type: dry western slopes,
Cascade Mountains, southern Oregon, July 11, 1902, W. C. Cusick 2873
(HOLOTYPE, NY!; ISOTYPES, MO!, ORE!, POM!, UC!, Us!).

Stems simple to several branched, 15-50 cm high, 2-3 mm
diameter, sparsely to densely villous and stipitate-glandular through-

296 Rhodora [Vol. 86

out; rhizomes giving rise to several basal rosettes and flowering
stems, 2-3 mm thick, covered with scales and old leaf bases at the
summit. Cauline leaves 3-5 pairs, sometimes crowded towards stem
base and reduced above, spathulate to elliptic-ovate, 2-8 cm long,
1-4 cm broad, sparsely to densely villous and stipitate-glandular,
acute, sub-entire to mostly irregularly dentate; petioles mostly
broadly winged, 1-9 cm long, 2-15 mm broad; leaves of the
innovations 4-10, similar to cauline leaves. Inflorescence a solitary
head or corymb of 3-9(25) heads; peduncles 2-20 cm long, sparsely
to densely villous and long stipitate-glandular; heads discoid,
turbinate-campanulate, 15-28 mm high; involucral bracts 8-15,
broadly to narrowly lanceolate, 5-15 mm long, 1-4 mm broad,
sparsely to densely villous and stipitate-glandular, acute to obtuse.
Florets 15-50, yellow, tubular, 8-11 mm long, sparsely villous and
glandular below; pappus white, barbellate. Achenes black, 5-10 mm
long, | mm broad, sparsely short stipitate-glandular. Figure 13.
Chromosome number 2n = 38, 76.

ECOLOGY AND DISTRIBUTION: Relatively rare and forming small
populations in dry, open Pinus-Quercus-Pseudotsuga menziesii
forests or such disturbed areas as roadcuts. Largely restricted to
serpentine soils in Curry, Douglas, Jackson and Josephine Coun-
ties, Oregon and Del Norte and Siskiyou Counties, California.
Figure 14. Elevational distribution 200-1500 m. Flowers April-
July.

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Del Norte Co.:
Douglas Park, on Smith River, J. Thompson s.n. (CAS, NY); Smith River Canyon,
15S mi. E. Crescent City, Ripley and Barneby 6798 (NY); Gasquet, A. Eastwood 22/1]
(NY, US); French Hill, 2 mi. S. Gasquet, J. Tracy 1146] (UC); Smith River at 18 mi.
Creek, 3 mi. E. Gasquet, J. Tracy 12284 (UC); Old Gasquet Toll Road, J. Tracy
11208 (UC); State Line, N. Monumental, J. Tracy 19423 (UC); Hayne’s Flat Rd. on
Coon Mt., J. Tracy 18921 (UC); Grade from Patrick Creek to Shelly Creek, A.
Eastwood and J. Howell 3667 (CAS); Gasquet, M. Peacock s.n. (CAS); French Hill,
A. Eastwood 2211 (CAS); Patrick Creek, A. Eastwood 12120 (CAS); Gasquet Mt.,
A. Eastwood 12155 (CAS); along Hwy. 99, 3.2 mi. N. Gasquet, D. Breedlove 3146
(CAS); Gasquet, Parks and Tracy 11208 (UC); French Hill, 8. J. Wolf 458 (ALTA);
10 km N. Gasquet, S. J. Wolf 459 (ALTA); 5.8 km NW Patrick, S. J. Wolf 387
(ALTA). Siskiyou Co.: Humbug Mt. G. Butler 985 (UC); Raspberry Lake, D.
Kildale 8706 (CAS).

OREGON: Curry Co.: 13 mi. SE Port Orford, M. Peck 8933 (GH, OSC); Agness,
M. Peck 2794 (OSC); Iron Mt., W. Baker 5677 (UC); Snow Camp, J. Thompson 31
(CAS). Douglas Co.: Glendale, T. Howell s.n. (CAN, NDG, US). Jackson Co.:
Wimer, £. Hammond 230 (US). Josephine Co.: Caves City, L. Rose 34218 (CAS,

1984] Wolf & Denford — Arnica 297

Figure 13. Arnica spathulata.

298 Rhodora [Vol. 86

e %
, a
o rk € 6G o IN 120°
&
| ee
® oe a ae) ee ee a |
e
*
* er *
a |
0 50km
on

Figure 14. Distribution of Arnica spathulata @ and Arnica viscosa *.

1984] Wolf & Denford — Arnica 299

MICH, UC); 2 mi. S. Union Mt. Lookout, 12 mi. W. Waters Creek, C. L. Hitchcock
and J. Martin 5125 (CAS, UC, WTU); Oregon Mt., A. Kruckeberg 1871 (UC), A.
Sweetser s.n. (UC); Kerby, L. F. Henderson s.n. (OSC, UC); Grants Pass, T. Howell
s.n. (MICH, OSC); Rough and Ready Creek, E. Meola 99 (OSC), hwy. 99,1 mi. N.
Cave Jct., K. Chambers 2912 (CAS, OSC); Hellgate, Rogue River, M. Peck 8933
(OSC); Wonder Post Office, M. Peck 23777 (OSC); 12 mi. W. Waldo, M. Peck 2916
(OSC); Merlin, L. Smith s.n. (CAS); Finch Ranch, near Kirby, L. F. Henderson 5900
(CAS); 2 mi. S. Wolfcreek, Ripley and Barneby 9552 (CAS, NY); Waldo Jct., D.
Kildale 9623 (CAS); Selma, N. Gale 14 (CAS), H. and S. Parks 5947 (UC), Grants
Pass, T. Howell 131 (US), C. Piper s.n. (US), Hammond 250 (NY), Caves Jct., near
Kirby, L. Rose 34218 (NY); 10 mi. S. Waldo, J. Tracy 4617 (CAS); 17 km. N.
Patrick, S. J. Wolf 460 (ALTA); 20 km N. Patrick, S. J. Wolf 461 (ALTA); Hugo, S.
J. Wolf 455 (ALTA); Merlin, S. J. Wolf 456 (ALTA); Store Gulch Guard Station, S.
J. Wolf 462 (ALTA); Babyfoot Lake, S. J. Wolf 384 (ALTA).

Arnica spathulata is a relatively rare, predominantly serpentine
endemic, and occurs in dry, open forests at mid elevations in the
Coast Ranges of the Klamath region. As previously noted (Wolf
and Denford 1984b), this species is almost certainly derived from A.
discoidea. However, it is readily distinguished by its spathulate,
broadly winged, petiolate leaves, broader phyllaries and larger,
black achenes which lack duplex hairs.

In more exposed habitats specimens of Arnica spathulata are
often smaller, less hairy, with narrower, frequently reddish leaves
crowded towards the base. Plants of this form have previously been
recognized as A. eastwoodiae by Rydberg (1927). Maguire (1943)
tentatively recognized this taxon as a subspecies of A. spathulata
but noted it may, in fact, only represent an environmentally reduced
form. These characters are probably environmentally induced and
no correlations between morphology, chromosome number, geog-
raphy or flavonoid chemistry could be discerned in this form (Wolf,
1981; Wolf and Denford, 1984b). Therefore it has not received
formal taxonomic recognition in the present study.

In his original description of Arnica spathulata Greene (1896) did
not designate a type or refer to any specimens examined. Maguire
(1943) later designated a Howell specimen at the U.S. National
Herbarium as the lectotype for this taxon (7. Howell s.n. Glendale,
Oregon, June 30, 1887 (erroneously cited as June 3)). However, a
specimen of A. spathulata on which Greene had written “Actual
type of my A. spathulata, Pitt. iii, 103!" has been located in his
personal herbarium at Notre Dame University. According to Article

300 Rhodora [Vol. 86

8 of the International Code of Botanical Nomenclature (Stafleu er
al., 1978) this specimen supersedes Maguire’s (1943) lectotype and is
here treated as the holotype of A. spathulata. Both the U. S.
National Herbarium specimen, previously chosen as lectotype, as
well as a specimen at the National Museum of Canada are of the
same collection as the holotype and are here recognized as isotypes.

8. Arnica venosa H. M. Hall, Univ. Calif. Publ. Bot. 6: 174. 1915.
Type: Salt Creek, Shasta Co., California, alt. 430 m H. M.
Hall and E. B. Babcock 4013 (HOLOTYPE, UC!; ISOTYPES, GH!,
NY!, RM!, uc!, us!).

Stems simple to 3-several branched, prominently ribbed, 20-60
cm high, 2-5 mm diameter, densely pilose and stipitate-glandular
above to less so below; woody caudex 3-5 mm broad, covered with
dark scales and old leaf bases. Basal rosettes lacking; cauline leaves
6-10 pairs, the middle largest, becoming reduced and bract-like
above and scale-like below, broadly sessile or rarely short-broadly
petiolate, ovate-elliptic to broadly lanceolate, 3-7 cm long, 1.5-4 cm
broad, firm, 3-5 nerved above, strongly reticulate-veined below,
glabrate to stipitate-glandular above, pilose and stipitate-glandular
below, especially on veins below, acute to obtuse, irregularly and
coarsely serrate. Inflorescence a solitary head on each branch, 1-7:
peduncle 2-5 cm long, densely pilose and stipitate-glandular toward
summit; heads discoid, turbinate-campanulate, 15-22 mm high;
involucral bracts 8-19, 8-16 mm long, 3-5 mm broad, ovate to
broadly lanceolate, pilose and stipitate-glandular, acute to obtuse.
Florets 30-60, yellow, tubular, 8-10 mm long, densely pilose below;
pappus white, barbellate. Achenes dark gray, angled and ribbed,
6-8 mm long, 1.5 mm broad, densely hirsute with duplex hairs. Fig-
ure 15, Chromosome number 2n = 38.

ECOLOGY AND DISTRIBUTION: An extremely rare species of very
dry, open Pinus-Quercus forests or, more commonly, of such
disturbed sites as road cuts. Known from about thirty small
populations, largely in western Shasta County and adjacent Trinity
County, California. Figure 2. Elevational distribution 400-1400 m.
Flowers May-June.

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Shasta Co.: Salt
Creek, Hall and Babcock 4013 (GH, NY, RM, UC, US); Castella, L. E. Smith 348
(CAS); Iron Mt., L. E. Smith s.n. (CAS, UC); road to Shasta Bally, 2.2kmS. Brandy

1984]

Wolf & Denford — Arnica

Figure 15. Arnica venosa.

301

302 Rhodora [Vol. 86

Creek, S. Whiskeytown Lake, G. Straley 1791 (UBC), S. J. Wolf 469 (ALTA); bluff
above mining road, E. of town of Iron Mt., W. Barker 227 (WTU). Lamoine Quad:
Baker Pine Plantation, W. of I-5, T36N, RSW, sec. 2, S. Horner 112 (STNB), G.
Straley 1793 (UBC), S. J. Wolf 468 (ALTA); Shell Mtn. Quad: 0.5 mi inside National
Forest Boundary, Trinity Mt. Rd., T34N, R7W, sec. 10, B. Williams 25] (STNF); W.
of Dog Creek Rd. between Tollhouse and Grouse Springs, 7 mi. W. Delta, T35N,
R6W, sec. 3, B. Williams 369, 370, 371 (STNF), M. Taylor 3195 (STNF); above
forest service road, | mi. NW Damnation Peak, T36N, R6W, sec. 22, M. Taylor 3196
(STNF). Trinity Co.: Swift Creek Rd., 0.7 mi. W. Hwy. 3, W of Trinity Center,
T36N, R7W, sec. 18, B. Williams 363 (STNF).

Arnica venosa is probably one of the rarest and most geo-
graphically restricted species of Arnica. It is known from about
thirty populations, all within a 25 km radius, largely in western
Shasta County, California. It is restricted largely to north-facing
slopes, at elevations of 400-1400 m, in open Pinus- Quercus forests
or more commonly on such disturbed sites as road cuts. Until very
recently A. venosa was known from only six populations and,
consequently, appeared on the California list of rare and endan-
gered species (Smith er al., 1980). However, based largely on the
efforts of Ms. Barbara Williams of the Shasta-Trinity National
Forest, many more populations have recently been discovered. The
authors are greatly indebted to Ms. Williams for providing
considerable information on A. venosa including both herbarium
specimens and highly detailed and complete ecological observa-
tions.

In its typical form Arnica venosa is readily recognized by its
rather stout, leafy stem; woody caudex; absence of innovations and
broadly sessile, ovate-elliptic, reticulate-veined, coarsely dentate,
very firm leaves. Since few specimens were available for exami-
nation, previous studies have concluded that this taxon exhibits
little variability (Maguire, 1943; Straley, 1980). However, the
availability and study of many newly collected specimens have
revealed that A. venosa, like most other Austromontana species,
exhibits considerable morphological variability and it appears to
intergrade with A. discoidea. As previously noted (Wolf and
Denford, 1984b), this taxon is probably derived from A. discoidea.
At one extreme are typical forms of A. venosa with leafy stems and
broadly sessile, veined leaves such as the type collection Hall and
Babcock 4013 (GH, NY, RM, UC, and US); S. J. Wolf 468, 469
(ALTA) and B. L. Williams 251 (STNF). At the other extreme are
such specimens as B. L. Williams 371 (STNF) with weakly veined,

1984] Wolf & Denford — Arnica 303

long, narrowly petiolate leaves more characteristic of A. discoidea.
This specimen is an otherwise typical form of A. venosa and is
characteristically highly branched above with very reduced leaves.
Additionally, it was collected near a population of A. discoidea and
may represent some introgression from that species. However,
without further evidence a hybrid hypothesis would be difficult to
support. Another seemingly intermediate form is represented by B.
L. Williams 250 (STNF) which has very typical upper leaves, leafy
stems and floral characters but has narrowly petiolate leaves below
the mid-stem.

Whether these intermediate forms represent introgression be-
tween Arnica discoidea and A. venosa or natural variability in the
latter is unclear. It is clear, however, that these two species are more
similar than previously demonstrated. The flavonoid profile of A.
venosa is a subset of that of A. discoidea and it is hypothesized that
the former is a relatively recent derivative of the latter (Wolf and
Denford, 1984b).

9. Arnica viscosa A. Gray, Proc. Am. Acad. 13: 374. 1878. TYPE:
Mt. Shasta, California, 8000’, Sept. 1877, J. D. Hooker and A.
Gray s.n. (HOLOTYPE, GH!).

Raillardella paniculata Greene, Erythea 3: 48. 1895. Type: near the limit of trees
on Mt. Shasta, California, Aug. 4, 1894, W. L. Jepson s.n. (HOLOTYPE, NDG!!.

Chrysopsis shastensis Jepson, Man. Fl. Pl. Cal. 1037. 1925. Type: Horse Camp,
Mt. Shasta, California, 1000 ft., W. L. Jepson 59i (HOLOTYPE, JEPS!). In his
original publication Jepson cited number 5/i as the holotype; however,
according to his notes (Robbins, annotation on type sheet) as well as his
designation of “Type” on number 59i, this latter specimen is the holotype, and
a typographical error occurred on publication.

Stems usually several branched, prominently ribbed, 20-50 cm
high, 3-5 mm diameter, strongly stipitate-glandular, also becoming
densely pilose above; woody caudex 3-5 mm broad, covered with
dark scales, lacking basal rosettes; leaves numerous, 5-10 pairs on
main stem, 2-6 pairs on branches, sessile, ovate-oblong to obovate-
oblong, (1)2-4(5) cm long, 1-3 cm broad, sparsely to densely pilose
and densely stipitate-glandular, more or less acute, entire. Inflores-
cence of 10-20 heads, peduncles 0.5-5 cm long, stipitate-glandular
and pilose; heads discoid, narrowly turbinate, 1-2 cm_ high;
involucral bracts 10-20, 6-10 mm long, 1-3 mm broad, broadly
lanceolate, stipitate-glandular and pilose below, acute. Florets
10-30, cream colored, tubular, 6-10 mm long, stipitate-glandular;

304 Rhodora [Vol. 86

pappus white, rarely tawny, barbellate to subplumose. Achenes
dark gray, ribbed 4.5-6.5 mm long, | mm broad, stipitate-glandular.
Figure 16. Chromosome number 2n = 38.

ECOLOGY AND DISTRIBUTION: A very rare species of dry,
exposed, pumice slopes at elevations of 1750-2500 m. Known
localities in Oregon include three small populations in Crater Lake
National Park, Klamath County and a single collection from the
Three Sisters area of Deschutes County. Also known from four
populations in Siskiyou County, California, a large population on
Mt. Shasta, two populations in the Marble Mountains and a single
collection from Preston Peak. An additional population is known
from the Trinity Alps, Trinity County, California. Figure 14.
Flowers August-September.

REPRESENTATIVE SPECIMENS: United States: CALIFORNIA: Siskiyou Co.: Mt.
Shasta, A. Eastwood 2055 (CAS, GH, UC), W. B. Cooke 9228 (UC), W. L. Jepson
s.n. (ND), R. Bohmannson s.n. (CAS); Horse Camp, Mt. Shasta, J. D. Hooker and
A, Gray s.n. (GH), W. B. Cooke s.n. (UC), 11501 (CAS, DS, GH, OSC, UC), 13833
(CAS, DS, ND, NY, OSC, UC), 17828 (CAS, WTU), P. Kamb 1488 (UC), W. L.
Jepson 59i (JEPS), W. Dress 3735 (UC), A. A. Heller 13519 (CAS, DS, NY, US,
WTU), W. Barker 232 (WTU), G. Straley 1411 (UBC), S. J. Wolf 391 (ALTA); S.
slope above ski lodge, Mt. Shasta, R. Thorne and F. Oettinger 39010 (NY, RSA,
UC); near Wagon Camp, Mt. Shasta, M. De’Evelyn s.n. (CAS); between Panther
Meadow and ski lift, Mt. Shasta, P. Hutchinson 938 (JEPS, K, US); South Gate, Mt.
Shasta, W. B. Cooke 25603 (GH, NY, WTU); Medicine Mt., 41° 33’48”, 121° 36’30”,
G. L. Clifton, s.n. (PUS); Devils Punchbowl, 41° 48’24”, 123° 40’36”, D. V. Hemphill
s.n. (PUA), Preston Peak, C. A. Ground s.n. (PUA); Upper English Lake,
41°24’36.2”, 123° 12’53”, F. W. Oettinger 668 (HSC, PUA, UC); Cliff Lake, G. Muth
s.n. (PUA); Avalanch Gulch, 41° 22’10”, 122° 13’39”, W. B. Cooke 2000 (UC). Trinity
Co.: Trinity Alps: Boulder Creek, W. J. Ferlatte 1286 (HSC, NY, UC); W. J. Ferlatte
484 (HSC); Mirror Lake, J. P. Smith 2361 (HSC).

OREGON: Deschutes Co.: Three Sisters area, Moraine Lake, E. Rock Mesa and
S. of South Sister, G. Van Vechten 219 (GH, OSC). Klamath Co.: Crater Lake
National Park: Union Peak, J. Mees s.n. (CLNP), F. Colville 1420 (RM, UC), E.
Applegate 10090 (CLNP); Hillman Peak, E. Applegate 10126 (CLNP), 10134 (CAS);
shore under Watchman Peak, E. Applegate 9218 (CAS, CLNP); Wizard Island, A.
A. Heller s.n. (CAS), 13820 (US); Garfield Peak, W. Baker 7201 (NY, WTU), G.
Straley 1946 (UBC), S. J. Wolf 511 (ALTA).

Arnica viscosa is one of the rarest and probably the most
distinctive species of the genus Arnica. This species is restricted to
volcanic soils and occurs on very open, rocky slopes at high
elevations in the Cascades of northern California and southern

1984] Wolf & Denford — Arnica 305

nO HID) }O HUY
DINVLO8 YWNY VINVS OHONVY

Figure 16. Arnica viscosa.

306 Rhodora [Vol. 86

Oregon. It is known from a few populations in Crater Lake
National Park, Oregon; Mt. Shasta, the Trinity Alps, Marble
Mountains and Preston Peak, California. An additional collection
was once made in the Three Sister Area of the central Oregon Cas-
cades (G. van Vechten 219 OSC, GH); however, repeated attempts
by several workers, including the senior author, have failed to
relocate this population.

Arnica viscosa 1s quite distinctive and easily recognized by its
woody caudex; leafy branching habit; small, sessile, entire leaves;
and narrow heads with cream-colored florets. In addition, virtually
all parts of the plant are densely covered with long glandular hairs,
so much so that it feels slimy to the touch. Additionally, it has a very
distinctive odor which is retained almost indefinitely on herbarium
sheets. All known collections of A. viscosa have been examined and
this species appears to exhibit virtually no interpopulational
variation. In fact the only atypical specimens examined were from
Upper English Lake, Siskiyou Co., California (F. Oettinger 668
HSC, UC). These plants were less viscid and the upper leaves and
branches had a tendency to be sub-opposite to alternate.

The underground parts of Arnica viscosa, including the caudex
and root sytem, are quite woody. This character is probably an
adaptation to its rocky, relatively disturbed habitat on very steep
slopes. Much of the root system is exposed, probably due to rock
movement associated with heavy winter snows and runoff.

EXCLUDED TAXA

A. latifolia Bong. var. viscidula A. Gray, Syn. Fl. N. Am. 1: 381.
1884. Type: Sierra Nevada Mts., California, Sept. 25, 1882, C. G.
Pringle 2 (HOLOTYPE US!) = A. diversifolia Greene, Pittonia 4:
171. 1900.

A. granulifera Rydb., Fl. Rocky Mts., 978. 1917. Type: Long Baldy,
Little Belt Mt., Montana, Aug. 19, 1896, J. H. Flodman s.n.
(HOLOTYPE, NY!) = A. mollis Hook., Fl. Bor.-Am. 1: 331. 1834.

A. ovalis Rydb., N. Am. Fl. 34: 338. 1927. Type: Crowsnest Pass,
Canadian Rocky Mts., J. M. Macoun (Can. Geol. Surv. No.
72719) (HOLOTYPE, CAN!) = A. mollis Hook., Fl. Bor.-Am. 1: 331.
1834.

1984] Wolf & Denford — Arnica 307

ACKNOWLEDGMENTS

We thank John Bain for providing additional collections and
Barbara Williams for sharing ecological and locality data for A.
venosa. Financial support from the Boreal Institute for Northern
Studies, California Native Plant Society and NSERC Canada is
gratefully acknowledged. We also thank the curators and staff of the
following herbaria for supplying loans and/or accommodating
visits: ALA, ALTA, BM, BRY, CAN, CAS, DAO, DS, GH, HSC,
JEPS, K, LCU, LE, MICH, MO, MONT, ND, NDG, NY, ORE,
OSC, POM, PUA, RENO, RM, RSA, UBC, UC, UCSB, US, UTC,
WISC, WS, WTU, Crater Lake National Park (here designated
CLNP), Shasta-Trinity National Forest (here designated STNF).

LITERATURE CITED

AFZELIUS, K. 1936. Apomixis in der Gattung Arnica. Sv. Bot. Tidskr. 30:
§27-579.

BARKER, W. 1966. Apomixis in the genus Arnica (Compositae). Ph.D. Disserta-
tion. University of Washington, Seattle.

Birp, C. D. 1967. The mosses collected by Thomas Drummond in western
Canada 1825-1827. Bryologist 70: 262-266.

Cronaqulist, A. 1955. Compositae. /n C. L. Hitchcock, A. Cronquist, M. Ownbey
and J. W. Thompson. Vascular plants of the Pacific Northwest. Princeton Univ.
Press, Princeton, N. J.

1958. Arnica. In Ferris, R., Taxonomic notes on western plants, Cont.
Dudley Herb. 5: 102.

1977. The Compositae Revisited. Brittonia 29: 137-153.

Davis, P. AND V.H. HEywoop. 1963. Principles of angiosperm taxonomy. Oliver
and Boyd, Edinburgh.

EpIGeR, R. I. AND T. M. BARKLEY. 1978. Arnica. In C. T. Rogerson (ed.), North
American Flora. Series II, Part 10, N. Y. Botanical Garden.

FERNALD, M. L. 1935. Critical plants of the upper Great Lakes region of Ontario
and Michigan. Rhodora 37: 324-341.

Fiint, R. F. 1957. Glacial and Pleistocene geology. John Wiley, New York.

Gray, A. 1883. Contributions to North American Botany. Proc. Am. Acad. 19:
1-96.

1884. Arnica. In Synoptical flora of North America, Vol. I, Part 2,
Caprifoliaceae-Compositae. Smithsonian Institution, Washington, D. C.

GREENE, E. L. 1896. New or noteworthy species XVII. Pittonia 3: 1-149.

1900. A series of papers relating to botany and botanists. Pittonia 4:
104-226.

GusTAFsson, A. 1947. Apomixis in the higher plants, II. The casual aspect of
apomixis. Lunds Univ. Arsskr. 43: 71-178.

308 Rhodora [Vol. 86

Howe Lt, T. J. 1900. A flora of northwest America.

HuLtTén, E. 1937. Outline of the history of arctic and boreal biota during the
Quaternary period. Bokforlags Akiebolaget Thule., Stockholm.

KRUCKEBERG, A. R. 1954. The plant species in relation to serpentine soils.
Ecology 33: 267-274.

1969. Soil diversity and the distribution of plants, with examples from
western North America. Madrofio 20: 129-154.

Lewis, H. 1962. Catastrophic selection as a factor in speciation. Evolution 16:
257-271.

Love, A. AND D. Love. 1982. IOPB Chromosome number reports. LXXV.
Taxon 31: 344-360.

MaGulIRE, B. 1943. A monograph of the genus Arnica. Brittonia 4: 386-510.

1947. Great Basin Plants—IX. Compositae. Am. Midl. Nat. 37: 136-145.

McKeg, B. 1972. Cascadia, the geological evolution of the Pacific Northwest.
McGraw-Hill, New York.

NORDENSTAM, B. 1977. Senecioneae and Liabeae: Systematic Review. Jn Hey-
wood, V. H., J. B. Harborne and B. L. Turner (eds.), The biology and chemistry
of the Compositae. Academic Press, New York.

RAVEN, P. R. AND D. I. AXELROD. 1978. Origin and relationships of the Cali-
fornia flora. University of California Press, Berkeley.

Rosinson, H. 1981. A revision of the tribal and subtribal limits of the Heli-
antheae (Asteraceae). Smithson. Contrib. Bot. 51.

Ro.iins, R. C. 1972. The need for care in choosing lectotypes. Taxon 21: 635-
637.

RypperG, P. A. 1897. Rarities from Montana III. Bull. Torrey Bot. Club 24:
292-299.

1927. North American Flora (Carduales) Carduaceae, Liabeae, Neuro-
laeneae, Senecioneae, 34, Part 4. N. Y. Botanical Garden.

SmiTH, J.. R. CoLE AND O. Sawyer. 1980. Inventory of rare and endangered
vascular plants of California. California Native Plant Society, Special Publi-
cation No. I, Berkeley.

STAFLEU, F. A., (ed.). 1978. International code of botanical nomenclature. Reg.
Veg. Vol. 97. Bohn, Scheltema and Holkema, Utrecht.

STEBBINS, G. L. 1971. Chromosomal Evolution in Higher Plants, Arnold Ltd.
London.

STRALEY, G. B. 1980. Systematica of Arnica, subgenus Austromontana and a new
subgenus Calarnica (Asteraceae: Senecioneae), Ph.D. Dissertation. University of
British Columbia.

1982. IOPB Chromosome number reports. LXXVI. Taxon 31: 579.

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and
California. Ecol. Monogr. 30: 279-338.

1961. Vegetation history of the Pacific Coast States and the “central”
significance of the Klamath Region. Madrofio 16: 5—23.

Wotr, S. J. 1980. Cytogeographical studies in the genus Arnica (Compositae:
Senecioneae). I. Amer. J. Bot. 67: 300-308.

1981. A biosystematic revision of Arnica L. (Compositae) subgenus
Austromontana Maguire, Ph.D. Dissertation. University of Alberta.

1984] Wolf & Denford — Arnica 309

, AND K. E. DENFoRD. 1983. Flavonoid variation in Arnica cordifolia: an
apomictic polyploid complex. Biochem. Syst. Ecol. I}: 111-114.

AND 1984a. Arnica gracilis (Compositae), a natural hybrid
baveei A. latifolia and A. cordifolia. Syst. Bot. 9: 12-16.

, AND 1984b. Flavonoid diversity and endemism in Arnica
subgenus Austromontana. Biochem. Syst. Ecol. (in press.)

S. J. W.

MISSOURI BOTANICAL GARDEN
P.O. BOX 299

ST. LOUIS, MISSOURI 63166
U.S.A.

KBD:

BOTANY DEPARTMENT
UNIVERSITY OF ALBERTA
EDMONTON, ALBERTA T6G 2E9
CANADA

A SYNOPSIS OF THE GENUS HALENIA
(GENTIANACEAE) IN MEXICO

ROBERT L. WILBUR

ABSTRACT

An abbreviated revision of the Mexican representatives of the genus Halenia
Bérckh. (Gentianaceae) is provided. Descriptions, comments and distribution data
with citation of some of the examined specimens is presented for the twelve species
known to occur in Mexico. Two new species, Halenia alleniana and H. crumiana, are
validated.

Key Words: Halenia, Gentianaceae, Mexico

The genus Halenia Borckh. is an almost exclusively American
genus belonging to the Gentianaceae—Gentianeae and comprised of
perhaps as many as seventy species according to the most recent
revision of the genus (Allen, 1933). Allen’s studies indicate that
more than seventy percent of the species are restricted to Andean
South America and most of the remainder are found in the
mountains extending from the southwestern United States to
western Panama. Apparently a single species ranges across most of
the northern United States and Canada from Newfoundland to
British Columbia. The type of the genus, H. corniculata (L.)
Cornaz, is one of the very few non-American species in the genus
and that Asian species ranges from the Urals to eastern Siberia and
south into Manchuria and Mongolia. A second Asiatic species, H.
elliptica D. Don ex G. Don, was described from the Himalayas and
apparently extends north into Soviet Central Asia and China.

The genus has proven to be a difficult one owing at least in part to
the considerable plasticity of the plants involving even the most
fundamental diagnostic characters used in distinguishing the
various taxa, the corolline spurs. Opportunity for extensive
observations on the plants in their natural habitats seems a
prerequisite for understanding much of the puzzling variation
encountered within the genus and both the present study and the
previous study by Allen (1933) suffer from the fact that they have
been almost exclusively based upon herbarium investigation. It
would also seem that the genus would yield results of greatest
biological interest from an investigation of pollination biology of
the various species since much of the most conspicuous variation is

311

312 Rhodora [Vol. 86

based in nectariferous spurs which vary strikingly in size, shape and
posture. Unfortunately this paper is not a report of such a study.
Instead of these promising approaches to the considerable and
puzzling variation that exists within the genus, the present report is
the result of study of approximately two thousand specimens of
Halenia, from Mexico and Central America. In the more than half
century that has elapsed since Allen revised the American species in
her doctoral dissertation, Mexico has been the site of considerable
collecting and as a consequence there is now a far more ample series
of specimens than what was available to her. Still, it would be very
much a mistake to conclude that botanical collecting in Mexico is
approaching the point of adequacy, and that collectors might be
well advised to abandon general collecting and concentrate their
efforts almost exclusively upon their own special group. In fact the
evidence seems to me conclusive that there has been far too much
emphasis by “collectors” on the intensive study of their narrow
special interest and a neglect of general collecting, except by those
who are preparing regional or state floras. Careful, critical
collecting is very much needed in most parts of Mexico and will be
necessary for decades to come. The present synopsis is offered to
provide a more realistic account of the genus and consequently
prove useful to those who might be tempted to work with these most
interesting plants living as they do in some of the most scenic areas
in Mexico.

GENERIC DESCRIPTION

Halenia Bérckh., Arch. Bot. (Leipzig) 1: 25. 1796. nom. cons. TYPE:
Halenia sibirica Bérckh., nom. illeg. [= Swertia corniculata L.,
Halenia corniculata (L.) Cornaz].

Tetragonanthus Gmel., Fl. Sibirica 4: 114, pl. 53. 1769. nom. illegit. (Art. 34.14).

Ceratia Pers., Syn. Pl. 1: 287. 1805, non Adans., Fam. 2: 319, 535. 1763 (=
Ceratonia L.)

Exadenus Griseb., Gen. Sp. Gentian. 322. 1838. LECTOTYPE: E. brevicornis (H.B.K.)
Griseb.

Annual, biennial or perennial, glabrous, caulescent herbs. Leaves
decussate, opposite or rarely whorled, entire, membranous to fleshy,
sessile or petiolate, usually 3-5-veined. Inflorescence a terminal or
axillary, subumbellate or rarely racemose to spicate cyme. Calyx
deeply 4-parted with the lobes only basally united and often bearing
squamellae internally at the base of each lobe, the lobes somewhat

1984] Wilbur — Halenia 313

inconspicuous to foliaceous, linear, lanceolate or ovate to spatulate,
often marginally papillate. Corolla 4-parted, usually greenish yellow
or rarely whitish or purplish, marcescent; lobes dextrorsely con-
volute, elliptic to ovate, obtuse to acute or acuminate, entire to
erose, often marginally papillate. Stamens 4, inserted on the corolla-
tube and alternating with the lobes; filaments linear or occasionally
basally dilated; anthers versatile, ovate, oblong or subtriangular.
Pistil bicarpellary; stigma sessile with 2, oblongish lobes 3-4 times
as long as thick and these receptive on the inner surface; style
lacking; ovary sessile, l-celled with two parietal placentae bearing
numerous ovules. Capsule compressed, septicidally dehiscent from
the apex; seeds globose to elliptic and slightly flattened with a
granular to reticulate surface.

A genus of perhaps 70 species with most of the American species
found in Andean South America with a smaller center of variation
in Central America and Mexico.

Allen (1933) recognized two formal sections within the American
species of the genus Halenia: Swertiella and Haleniastrum (=
Halenia). These were distinguished by her diagnostic key as follows:

Plants usually coarse with fleshy leaves, rarely slender with thin,

herbaceous leaves; stem usually leafy; spurs absent, or present
as very small inconspicuous protuberances, frequently ob-
scured by calyx; distribution chiefly South America.......

ities ieee ed eeee eee hee ek ehed eR aee Es 1. Swertiella

Plants usually slender, with thin, herbaceous leaves, stems leafy

or scapose; spurs present; distribution North and South
A. KC SEGE HES ENS ENERO RARE ES 2. Haleniastrum

According to Allen, Halenia alata and H. brevicornis are the two
species from North of South America belonging to her section
Swertiella, while the remaining species belong to what we would call
section Halenia and which she designated as section Haleniastrum. |
am not convinced that the late Caroline K. Allen has delimited
natural subgeneric groupings within the genus Halenia but am
unable to suggest a more meaningful grouping or infrageneric
classification at this time.

KEY TO THE MEXICAN SPECIES OF HALENIA

| Pimaie annual OF DiCnNIG) saw ec de bee ee RO EEE He PERERA (2)
2. Corolline protuberances either lacking or less than 2 mm long

314 Rhodora [Vol. 86

and not developed into conspicuous nectariferous spurs or
WOU INS sana saat oeyoe cawuee se eesans 1. H. brevicornis.
2. Corolline protuberances well-developed and usually repre-
sented by nectariferous spurs 3 mm long or longer.
3. Corolline spurs either strongly divergent or at least spread-
ing outwardly and distally arching outwardly.

4. Corolline spurs so strongly divergent as to be horizontal
or nearly so; corolla lobes + acute; midcauline leaves
mostly linear, usually less than 4 mm wide........
MGUCRUEECEETERRETAES Ke ota a ean es 2. H. recurva.

4. Corolline spurs diverging especially distally but not more
than 30°; corolla lobes rounded to obtuse; midcauline
leaves elliptic, usually 4 mm or more wide ........
PERREGH EA's G OUAREE Sheu ak Bea R be Ee 3. H. crumiana.

3. Corolline spurs + pendent and often distally inwardly

curved.
5. Midcauline leaves linear, mostly 10 or more times as long
|: | Sa ee re 4. H. palmeri.

5. Midcauline leaves oblong to lanceolate or broadly elliptic
or ovate, much broader than 6 times as long as wide.
6. Calyx lobes spatulate, obtuse............... eee ee
eT Tee Cee eee eee 5. H. conzattii.

6. Calyx lobes oblong to lanceolate or elliptic, acute.
7. Midcauline leaves ovate to broadly lanceolate, clearly
slenderly petiolate with the petioles 3-12 mm long;
basal leaves with petioles 10-30 mm long and
about as long or even longer than the blades:
capsules 15 mm long or less ...............4.
(iRee Neh eee aeedaereriaases 6. H. schiedeana.
7. Midcauline leaves oblong to oblanceolate; tapering
to a rather broad base or at most very indistinctly
winged-petiolate; capsules (12) 15-25 mm long .
eee eT Te rer eT ee re ree 7. H. alleniana.

1. Plants perennial.
8. Corolline nectaries merely pouch-like or at least not developed
into spurs more than 1.5 mm long.

9. Calyx lobes obtuse; corolla 6 mm long or less; corolline
nectaries pouch-like, drying dark brown .............

1984] Wilbur — Halenia 315

9. Calyx lobes acute to acuminate; corolla 6 mm long or
longer; corolline nectaries never drying as dark brown
CICUIAT DAICHOS i+. 14-4 s2555,050 eae ees 9. H. pringlei.

8. Corolline nectaries typically spur-like and longer than 2 mm.
10. Corolline spurs strongly divergent or outwardly spreading.
11. Plants usually 1-5 dm tall or less; corolline spurs
horizontally spreading or at least very strongly
divergent; cauline leaves mostly linear to oblanceo-

late, mostly 6 mm wide or less..... 9. H. pringlei.

11. Plants 1.5 dm tall or more; corolline spurs slender and
more than 4 times as long as the diameter and these

more or less descending or if moderately spreading

never approaching the horizontal; cauline leaves

mostly broadly elliptical, about 1 cm wide........
bpiciee cae Ee de AEs ORS eee eas 10. H. hintonii.

10. Corolline spurs + pendulous and neither divergent nor
conspicuously outwardly curved, often distally straight.

12. Basal rosette absent or at least the leaves mostly
CRUE hs oer he 2 ROSE OS 11. H. decumbens.
12. Basal rosette present and cauline leaves few or none.
13. Spurs distally incurved, less than half the length of the

corolla.

14. Flowers more than 10 mm long; corolline spurs
usually | /3-1/2 the length of the corolla, 3-7 mm
FONE Sc icee sexs seuss cashes 12. H. plantaginea.
14. Flowers less than 10 mm long; corolline spurs
rudimentary and usually 2.5 mm long or less .
pS en Saeed eR Raa RAT 13. H. nudicaulis.
13. Spurs + straight or distally slightly outwardly curved,
about half the length of the corolla............
(itusesee tines cake aoseke es 11. H. decumbens.

TREATMENT OF INDIVIDUAL TAXA

1. Halenia brevicornis (H.B.K.) G. Don, Gen. Hist. 4: 177. 1838.

Swertia brevicornis H.B.K., Nov. Gen. et Sp. Pl. 3: 174. 1818. Type: Ecuador;
Quito, Humboldt & Bonpland s.n.

Swertia parviflora H.B.K., Nov. Gen. et Sp. Pl. 3: 174. 1818. Type: Mexico;
Guanajuato, Humboldt & Bonpland, photo of type at Paris seen at Us.

Swertia parviflora var. a angustifolia Schlect. & Cham., Linnaea 5: 122. 1830.

316 Rhodora [Vol. 86

Swertia parviflora var. B latifolia Schlect. & Cham., Linnaea 5: 122. 1830.

Halenia parviflora (H.B.K.) G. Don, Gen. Hist. 4: 177. 1838.

Exadenus brevicornis (H.B.K.) Griseb., Gen. et Sp. Gent. 323. 1839.

Exadenus parviflorus (H.B.K.) Griseb., Gen. et Sp. Gent. 322. 1839.

Exadenus parviflorus var. B latifolius (Schlecht. & Cham.) Griseb., Gen. & Sp.
Gent. 322. 1839.

Halenia multiflora Benth., Pl. Hartw. 24. 1839. Mexico: in pinetis Bolafios,
Hartweg 210 (K, HOLOTYPE, photo!; Ny! ISOTYPE).

Exadenus paucifolius Mart. & Gal., Bull. Acad. Brux. 11: 372. 1844. Type: Se
trouve avec l’espete précédente [E. alatus Mart. & Gal.] de 9 a 12,000 pieds
[“du pic d’Orizaba”], Galiotti 7219 (BR, HOLOTYPE, photo!).

Halenia paucifolia (Mart. & Gal.) Hemsl., Biol. Centr. Amer. Bot. 2: 352. 1882.

Halenia parviflora var. latifolia (Schlect. & Cham.) Hemsl., Biol. Centr. Amer.
Bot. 2: 351. 1882.

Tetragonanthus paucifolius (Mart. & Gal.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Tetragonanthus parviflorus (H.B.K.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Tetragonanthus brevicornis (H.B.K.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Swertia cucullata Sess¢ & Mocino, Fl. Mex. ed. 2 p. 73. 1894. Type: Mexico,
Sesse & Mocino s.n. (MA, HOLOTYPE, photo at MO!).

Halenia erythraeoides Gilg, Engler Bot. Jahrb. 54: Beibl. 118, p. 105. 1916. TyPE:
Venezuela: “Auf den Hochanden von Merida”, Linden 456 (B, HOLOTYPE).

Halenia micranthella Brig., Candollea 4: 320. 1931. Type: Mexico; Hidalgo, wet
meadows, Sierra de Pachuca, alt. 2450 m, Pringle 6964 [as 1964] (G, not seen,
HOLOTYPE; ENCB! F! MICH! MO! MSC! NY! PH! US! ISOTYPES).

Halenia tuerkheimii Brig., Candollea 4: 317. 1931. Type: Guatemala; Alta
Verapaz, “Fichtenwalder bei San Joaquin” 1000 m, von Tuerckheim 2041 (G,
HOLOTYPE; F! GH! NY! US! ISOTYPES).

Halenia brevicornis var. multiflora (Benth.) C. K. Allen, Ann. Missouri Bot.
Gard. 20: 142. 1933.

Halenia brevicornis var. micranthella (Briq.) C. K. Allen, Ann. Missouri Bot.
Gard. 20: 143. 1933. Hylenia brevicornis var. divergens C. K. Allen, Ann.
Missouri Bot. Gard. 20: 144. 1933. Type: Mexico; Michoacan, vicinity of
Morelia, near La Huerta, 1950 m, | Sept. 1910. Arsene s.n. (MO! HOLOTYPE; US!
ISOTYPE).

Halenia brevicornis var. chihuahuensis C. K. Allen, Ann. Missouri Bot. Gard.
20: 144. 1933. Type: Mexico; Chihuahua, pine plains, base of the Sierra
Madre, Pringle 1664 (MO! HOLOTYPE; CAS! MSC! ISOTYPES),

Halenia brevicornis var. ovata C. K. Allen, Ann. Missouri Bot. Gard. 20: 145.
1933. Type: Mexico; Nayarit, Tepic, Jan 5-Feb 6, 1892, Palmer s.n. (us!
HOLOTYPE).

Halenia brevicornis var. tuerckheimii (Briq.) C. K. Allen, Ann. Missouri Bot.
Gard. 20: 145. 1933.

Annual herbs (0.5)1.5—4(9) dm high from a slender tap root; stems
slender, strongly angled to even quadrate, striate, usually branched
only above the base. Leaves mostly cauline, thinly membranous,
sessile or only the lowermost petiolate (the petioles in these
0.5-1.5(3.0) cm long), the blades linear to lanceolate or elliptic,

1984] Wilbur — Halenia 317

mostly 2-3.5(5) cm long and (1)2-10(18) mm wide. Inflorescence
cymosely racemiform to paniculate, compactly congested to loosely
arranged; pedicels 1—-12(25) mm long. Calyx tube shallowly turbin-
ate to campanulate, 0.3-0.6(0.8) mm _ high, the lobes linear to
lanceolate or more rarely oblong or narrowly spatulate, | / 2-3/4 the
length of the corolla, mostly (1.5)2-4 mm long and 0.2-0.6 mm
wide, usually faintly 3-nerved, marginally often minutely papillate;
corolla green to greenish yellow, 4-8 mm long, the tube 3-4.5 mm
long, the lobes broadly to narrowly ovate to deltoid-oblong, apically
acute to acuminate and basally somewhat auriculate, 2-3 mm long,
spurs lacking or marked by slight protuberances or represented by
readily detected, pendant to divergent spurs 0.5—1.0(2.0) mm long;
stamens 2-4 mm long, the filaments attached near the middle of the
corolla-tube. Capsule compressed, lanceolate, 5-9(-14) mm long;
seeds ovoid to subglobose, 0.4-0.6 mm long, reticulate.

DISTRIBUTION. Dry, grassy slopes and open woods from n.
Mexico south through Central America into northwestern South
America.

This species has been divided by Allen (1933) into eight varieties
thought by Williams (Fieldiana Bot. 24: 317. 1969) “to have slight if
any systematic importance.” My own findings are in agreement with
those of Williams. Allen, although stating “that environment is ina
large measure responsible for the variation found within the
species,” keyed what she treated as varieties as follows:

a) Leaves not ovate, longer than 1.2 cm long.
b) Pedicels stouter than filiform, shorter than 1.2 cm long.
c) Corolla without distinct spurs.
d) Inflorescence compact.
e) Leaves linear, slender .....csceseees var. brevicornis
¢) Leaves ovate to lanceolate, coarse 6666 6s sc deee caw
ee var. latifolia (Schlecht. & Cham.) C. K. Allen
Gh TUTORS RCE BIE oh ase eee ee R44 2S SEER ESS
eer Te ee ee ee ee var. micranthella (Briq.) C. K. Allen
c) Corolla with small but distinct spurs.
f) Spurs thick, conical, more or less pendulous .........
(oubedatado ae nee var. multiflora (Benth.) C. K. Allen
1) Spurs Glent, spreading Squarrose....csvwessn see cendns
ieguaweweeeneoweetae ty var. chihuahuensis C. K. Allen
f) Spurs slender, divergent.... var. divergens C. K. Allen
b) Pedicels filiform, slender, elongate; habit decidedly

318 Rhodora [Vol. 86

eet ee ee ere ee var. tuerckheimii (Briq.) C. K. Allen
a) Leaves ovate, shorter than 1.2 cm... var. ovata C. K. Allen

Naturally with so much described variation and so many formally
named varieties of Halenia brevicornis, the suspicion is great that
the easiest path has been followed in not recognizing any of them in
this synopsis. Considerable opportunity to observe the plants in the
field in a wide variety of habitats and throughout its range ought to
help sort out at least some of the environmentally induced plasticity.
The opportunity to grow the plants under controlled environmental
conditions also gives promise of resolving this problem. But until
such steps are taken, I believe it best to treat the species broadly as
the problem seems not to lend itself well to analysis based upon the
more traditional morphological-geographical analysis. I have not
found Allen’s treatment helpful in sorting out the populations into
meaningful taxa and believe that much of the difficulty is traceable
to the extreme plasticity of individuals responding to differences in
light, available moisture and seasonality.

REPRESENTATIVE MEXICAN COLLECTIONS: Baja California: locality uncertain,
‘“Lower California”, Orcutt s.n. (NY). Sonora: pine slopes, La Mesa Colorado, 14 Oct
1933, Gentry 542M (DS, MICH, US); open pine forests, Saguaribo, Rio Mayo, 5500 ft,
2 Nov. 1935, Gentry 2099 (F, GH, MO, PH, US); slopes in oak forest on W slope of the
Sierra Madre, 35 miles SW of Chuhuichupa on trail to Rio Bonito, 2 Oct 1939,
Muller 3606 (GH, LL, MICH). Chihuahua: along arroyo just NW of Cusarare church,
2200 m, 14 Oct 1977, Bye & Weber 8112 (LL, GH); Sierra Madre Mts., Mesa, W of
Hop Valley, 6500 ft, 17 Sept 1903, M. E. Jones s.n. (F, MICH, MO, US); stony pineland,
Madera, 2150-2200 m, Sierra Madre Occidental, 25 Sept 1934, Pennell 19235 (PH,
us); pine plains, base of the Sierra Madre, 4 Oct 1887, Pringle 1330 (F, MICH, NY, PH,
us). Nuevo Leon: open chaparral, upper west slope of Sierra de la Cebolla, 21 Aug
1939, Muller 2898 (GH—a mixed sheet). Sinaloa: Sierra Surutato, 0.5 mi N of Los
Ornos, 5800 ft, | Nov 1969, Breedlove & Kawahara 16742 (CAS, MICH); steep moist
slope, Sierra Surutato, 3 miles N of Los Ornos along road to Ocurahui, 2 Oct 1970,
Breedlove & Thorne 18338 (CAS, MICH). Durango: steep moist ravine 2 miles W of
Revolcaderos along Mexican Hwy 40, 7100 ft, 8 Nov 1970, Breedlove 18921 (cas,
MICH); open oak-pine woodland Sierra Madre Occidental about 50 km W of Ciudad
Durango, 2550 m, 27 Sept 1962, McVaugh 21661 (CAS, ENCB, LL, MICH, NY); dryish
cliffs, 11.2 miles NE of El Paraiso (Sinaloa) on road between Villa Union and El
Salto, 7800 ft, 29 Sept 1953, Ownbey 1970 (F, GH, MICH, NY, US); rocky pineland
canyon, El Salto (Aserraderos), 2500 m, 31 Aug 1934, Pennell 18510 (GH, NY, PH, US);
pumice gravel in dry pine-oak forest, 8600 feet, Sierra Madre, W slope, 17.5 miles, W
of El Salto, 18 Oct 1965, Ripley & Barneby 14176 (CAS, NY, US). Zacatecas: Sierra de
Los Morones, near Plateado, | Sept 1897, Rose 2732 (GH, US); about 38 km al W de
Jalpa, sobre la carretera a Tlaltenango, bosque de Quercus, 21-23 Oct 1973.

1984] Wilbur — Halenia 319

Rzedowski & McVaugh 975 (ENCB, MICH). Aguascalientes: moist N facing slopes near
summits, Sierra del Laurel, about 10 miles SE of Calvillo, 2500 m, 4 Nov 1959,
McVaugh & Koelz 206 (MiIcu). San Luis Potosi: Alvarez, 28 Sept-3 Oct 1902, Palmer
160 (F, GH, MO, NY, US); San Luis Potosi, 6000-8000 ft, 1878, Parry & Palmer 600 (F,
GH, MO, NY, PH, US). Nayarit: Tepic, Pa/mer s.n. (US). Jalisco: grassy slopes 12 km
NW of Los Volcanes, 1900 m, 30 Oct 1973, Breedlove 35772 (CAS, MICH); pine forest,
Sierra de Manantlan, 15-20 miles SE of Autlan, about 1700 m, McVaugh 13962
(MICH); in pine-oak woodland 5 miles NE of San Miquel de la Sierra, 1950 m, 2 Nov
1962, McVaugh 22014 (MICH, NY); Rio Blanco, Oct 1886, Palmer 680 (GH, MICH, NY,
us); banks of ravines near Guadalajara, 5000 ft, 21 Oct 1903; Pringle 11636 (F, GH,
us). Guanajuato: second growth oak forest on steep rocky mountainsides about 8 km
NE of Santa Rosa, 2400 m, 10 Nov 1970, McVaugh 24178 (micH). Hidalgo:
mountain meadows, El! Chico near Pachuca, Sept 1905, Purpus 176] (F, GH, MO, NY,
us). Veracruz: ladera de cerro en parcela de cultivo abandonado, 1700 m, 6 July 1970,
Ventura 1526 (DS, F, MICH, MO). Michoacan: llano about 4 km SW of Cerro San
Andres about 10 km N of Ciudad Hidalgo, about 2930 m, 6 Sept 1960 Beaman 4251
(GH, MSC): Zitacuaro—Las Cafioas, 2600 m, 4 Nov 1938, Hinton 13412 (GH, LL, MICH,
NY, PH, TEX, US); Zitacuaro—Guanoro, 1875 m, I1 July 1938, Hinton 13424 (GH, ILL,
LL, NY, PH, TEX, US); pine forest, Tancitaro, 2250 m, 22 Nov 1940, Hinton 15558 (Ds,
F, MICH, NY, US); pine-covered slopes and meadows about 18 miles S of Patzcuaro,
8900-9000 feet, 20-25 Nov 1961, King & Soderstrom 5156 (MICH, NY, TEX, US).
Mexico: grassy hill, Tequesquipan, Dist. Temascaltepec, 2800 m, 28 Oct 1932,
Hinton 2316 (F, GH, LL, NY, PH, US); hill, Ocotepec, Dist. Temascaltepec, 1500 m, 9
Dec 1932, Hinton 2905 (F, GH, LL, MO, NY, PH, US); pine forest, Nanchtitla, Dist.
Temascaltepec, 16 Dec 1938, Hinton 5354 (ENCB, F, MO, NY, US); llano, Sierrita, Dist.
Temascaltepec, Hinton 8311 (ENCB, F, GH, LL, MO, NY, PH, TEX, US); pine forest,
Mez6n Viejo, Dist. Temascaltepec, 11 Oct 1935, Hinton 8345 (F, GH, MO, NY, US);
near Ozumba, 8000 ft, 3 Nov 1902, Pringle 11329 (CAS, ENCB, F, GH, MICH, MO, MSC,
us); cool slopes, Sierra de las Cruces, 9500 ft, 12 Sept 1904, Pringle 13120 (CAs, F, GH,
MICH, US); open woods, Salto de Agua, Nov 1905, Purpus 1762 (F, GH, MO, NY, US);
Distrito Federal: open grassy roadbank, at La Cima Station between Mexico and
Cuernavaca, 3035 m, 25 Sept 1961, Beaman & Andresen 4534 (GH, TEX, US); Sierra de
Ajusco, 9 Nov 1903, Pringle 11842 (CAS, F, GH, MICH, MO, MSC, Us); alrededores de la
Estacién La Cima, Serrania del Ajusco, 3000 m, 25 Nov 1966, Rzedowski 23192 (Ds,
ENCB, MICH, MSC, TEX) Morelos: pine forest, Sierra de Morelos near Cuernavaca,
2100 m, 25 Dec 1969, Hinton 17427 (DS, ENCB, MICH); Tlaxcala: Cerro La Hoyanca,
cerca de Calpulalpan, 9 Sept 1956, Paray 2/02 (ENCB). Puebla: open volcanic slopes,
San Manuel de la Sierra, 9300 ft, 19 Aug 1938, Balls 5296 (GH, MSC, Us); Esperanza,
Aug. 1907, Purpus 2697 (F, MO, US); roadside bank between La Venta and San
Martin, 8200 ft, 15 Nov 1944, Sharp 441638 (GH, MO, NY). Guerrero: forests N slope
of Cerro Alquitran, 10-14 km by road W of Mexican Hwy 95 and Mazatlan,
2250-2450 m, 6 Dec 1966, Anderson & Laskowski 4407 (DUKE, ENCB, GH, MICH, NY,
US); pine forests, Pilas, Dist. Mina, 1500 m, 22 Nov 1936, Hinton 9886 (GH, NY, US);
pine forests, San Antonio—Buenos Aires, Dist. Montes de Oca, Hinton 11695 (Gu,
LL, NY, PH, US); Open pine forests, Teotepec, Distr. Mina, 3600 m, 17 July 1939,
Hinton 14463 (DS, ENCB, F, GH, MICH, NY); pine forest slope, Petlacala, Distr. Mina,
1820 m, Mexia 8963 (CAS, F, GH, MO, NY, US); Chichihualco, El Asoleadero, 15 km al
oeste de Camotla, 2650 m, 2 Dec 1963, Rzedowski 18055 (DS, ENCB, MICH, MSC).

320 Rhodora [Vol. 86

Oaxaca: grassy ridge and meadow in pine forest, 26 miles SSE of Miahualtan, 2600
m between Oaxaca and Suchixtepec on road to Puerto Angel, 7 Nov 1966, Anderson
& Laskowski 4156 (DUKE, ENCB, GH, MICH); SW slope of Cerro Zempoaltepetl along
trail from Tlahuitoltepec to Santo Domingo Alberradas, oak forests, about 2000 m,
14 Aug 1950, Hallberg 967 (ENCB, MICH, US); Sierra de San Felipe, 8000 feet, Smith
665a (F, MICH, MO, NY, US). Chiapas: on steep moist slope along Mexican Hwy 190, 2
km W of Navenchauk, 6000 ft, 19 Dec 1964 Breedlove 7974 (DS, ENCB, F, MICH); steep
slope, barrio de Tuk paraje of Matsab, Municipio Tenejapa, 7500 ft, 30 Sept 1965,
Breedlove 12517 (DS, ENCB, LL, MICH, US); Mt. Tacana, 1000-2000 m, Aug 1938,
Matuda 2473 (Ff, GH, LL, MICH, NY); grassy slope S of the center of Amatenango del
Valle, 6100 feet, 11 Nov 1966, Alush Shilom Ton 1508 (DS, ENCB, MSC, NY, US).

2. Halenia recurva (J. E. Sm.) C. K. Allen, Ann. Missouri Bot.
Gard. 20: 161. 1933.

Swertia recurva J. E. Smith, Rees’ Cyclopedia 34: [under Swertia.] 1819. Type:
Mexico: locality unknown, Escalante s.n. by Mutis 38 (LINN, not seen).
Halenia rothrockii A. Gray, Proc. Amer. Acad. I1: 84. 1876. Type: Arizona;

Mount Graham, at 9000 ft, Rothrock s.n. (GH, HOLOTYPE, not seen).
Tetragonanthus rothrockii (A. Gray) Heller, Catalogue N. Amer. Pl. 6. 1898.

Annual herbs with quadrate, narrowly winged stems basally
unbranched although often branched above, (1.2)2.5-6.5 dm tall.
Basal leaves (0.8)1-2.5(3.0) cm long, 3-6 mm wide, elliptic or
elliptic-lanceolate to spatulate; cauline leaves remote, lance-linear
to linear, 1.5—4(—7) cm long and |-3(4) mm wide, obscurely 3-nerved
above and only the midvein prominently elevated beneath.
Inflorescence a loosely flowered, sub-umbellate cyme; pedicels
slender, 0.5-3 cm long. Calyx-tube campanulate to turbinate, 1-1.4
mm high; calyx-lobes lanceolate to linear, acute, 4-8 mm long and
0.8-1.2(-1.5) mm wide, marginally very minutely papillate; corolla
bright yellow, about 0.8-1.2 cm long, tube less than half the length
of the corolla, lobes ovate, acute to subacuminate, delicately
veined, papillate, spurs strongly outwardly curved, mostly hori-
zontal but distally strongly ascending, 0.8-1.6 cm across from spur-
tip to tip or each spur mostly (4-)6-8 mm long; filaments slightly
obovate, anthers broadly oblong, mucronate, papillate. Capsule
lance-ovoid (8-)10-16 mm long; seeds yellow-brown, subglobose-
ovoid, granular.

DISTRIBUTION: mountains of the southwestern United States
(Arizona and New Mexico) and Mexico (Sonora, Chihuahua,
Coahuila and Durango) apparently above 7500 ft (= c. 2300 meters).

One is startled to read in Allen’s revision of the genus (1933, p.
162) that the type of Swertia recurva J. E. Smith is a “specimen

1984] Wilbur — Halenia 321

collected by Mutis and sent to Linnaeus, now preserved in the
herbarium of the Linnaean Society...” Since there is no evidence
that Mutis ever collected in Mexico, suspicion is aroused that the
Mutis collection upon which Swertia recurva is based belongs in all
probability to a South American species. There is, however, a
specimen “Mutis 38” at US which was obtained from the Madrid
Botanic Garden in 1932. J. E. Smith originally reported that the
plants were collected in Mexico by Escallon and a specimen sent to
Linnaeus by Mutis. This seems to be a plausible explanation of how
a specimen from northwestern Mexico was described by J. E. Smith
as Swertia recurva at such an early date.

REPRESENTATIVE MEXICAN COLLECTIONS: Coahuila: 25 km NW of Fraile on top of
mountain covered with Abies, Pseudotsuga and Pinus...3550 m, 16 July 1941,
Stanford et al 453 (DC, GH, MO, NY). Chihuahua: open pine slope, 69 miles from
Parral along road to El Vergel, 8200 feet, 7 Oct 1959, Correl] & Gentry 22882 (ENCB,
LL, MO); pine woods, Cajurichi, Rio Mayo, 7200 ft, 13 Sept 1936, Gentry 27/1 (F, GH,
MO); stony pine woods near First Meadow, Sierra Madre Occidental, 2250-2300 m,
23 Sept 1934 (F, GH, MICH, NY, US); cool slopes, Sierra Madre, 24 Sept 1887, Pringle
1329 (DS, F, GH, NY, PH, US); Sierra Madre near Colonia Garcia, 8000 ft, 6 Sept 1899,
Townsend & Barber 309 (F, GH, MO, NY, US). Sonora: pine zone, Las Tierritas del
Temblor, region of Rio de Bavispe, 20 Aug 1940, Phillips 648 (GH, LL, MICH).
Durango: barranca below Sandia Station, 6500 ft, 13 Oct 1905, Pringle 13588 (CAS, F,
GH, LL, MICH, MSC, TEX, US).

3. Halenia crumiana Wilbur, sp. nov.

Herba annua vel biennis, 3-4.5 dm alta. Radix palaris. Folia
basalia + rosulata; lamina lance-elliptica vel elliptica, 3-4 cm longa
et 6-10 mm lata, petioli 1-1.5 cm longi. Folia caulina elliptica
1.5-3.5 cm longa et ca. 4 mm lata, internodiis |—5-plo longioribus.
Lobi calycis lineares vel anguste oblanceolati, acuti, 4-6 mm longi et
0.8-1 mm lati. Lobi corollae oblongi vel ovati vel orbiculares, 4-6
mm longi et 0.8-2(2.2) mm lati; calcaria corollae 5-7 mm longa, +
pendula, paulo divergentia.

TYPE: MEXICO: Jalisco; cypress-pine forest in mountains E of
Manantlan about 15 miles SSE of Autlan by way of Chante. 30 July
1949. R. L. and C. R. Wilbur 1981 (MICH; 6 duplicates were also
collected but were not encountered during this study).

Probably an annual herb from a strong taproot 4-5 mm in
diameter and with a weakly 4-angled stem 3-4.5 dm tall and this
either little-branched or branching from near the base or even
sparingly branched throughout with the stems very slightly winged.
Basal leaves apparently at least sometimes rosulate or with

322 Rhodora [Vol. 86

internodes only 2-3 mm long and these leaves lance-elliptic to
elliptic with a petiole 1-1.5 cm long and a blade 3-4 cm long and
6-10 mm wide; cauline leaves rather remote with the internodes 1-5
times as long as the leaves, narrowly elliptic, 1.5-3.5 cm long and
about 4 mm wide, 3-nerved but only the midvein prominently
elevated beneath. Inflorescence of subumbellate cymes terminating
the main stem and the lateral branches (and these often very much
foreshortened); pedicels stiff, + 4-angled and very narrowly winged,
ascendent, 5-10 mm long. Calyx-tube campanulate, about 1-1.5
mm high; calyx-lobes linear to narrowly oblanceolate, acute, 4-6
mm long and 0.8-1 mm wide, microscopically marginally papillate;
corolla greenish distally but yellowish below including the spurs,
8-12 mm high, the tube about | /3 the height of the corolla, the lobes
broadly oblong to ovate or almost orbicular, broadly rounded to
obtuse but occasionally shortly apiculate, marginally erose, the
spurs outwardly divergent especially distally but still + pendent,
slender, medially usually less that 0.5-0.8 mm in diameter, mostly
5-7 mm long; anthers yellow, broadly oblong, about | mm long, the
filaments green, linear, slender, spreading ciliate for the basal half.
Capsule lance-ovoid, 9-12 mm long; seeds yellowish, subglobose to
oblongoid, smoothish.

DISTRIBUTION. Known only from the higher mountains of the
state of Jalisco, Mexico at an elevation of above 2500 m in rather
open forests of pine, fir, cypress, oaks and various other hardwoods.

This species has in the past been identified with Halenia recurva
(J. E. Sm.) C. K. Allen (= H. rothrockii A. Gray), a species found
high in the mountains of southern New Mexico and Arizona as well
as the Mexican states of Sonora, Chihuahua, Coahuila and
Durango. Halenia recurva differs in its narrowly linear leaves and
widely divergent corolline spurs that are often + horizontal. The
cauline leaves of H. crumiana are narrowly elliptic and the spurs,
although divergent, never approach being horizontal.

The species is named in honor of the accomplished bryologist
Howard A. Crum, a much-admired friend. He was a companion on
a collecting trip in the summer of 1949 spent in the vicinity of
Autlan in the Mexican state of Jalisco. The summer apparently was
not bryologically richly rewarding but he endured it in good humor
or at least what passed for such among those gifted complainers, the
Wilbur brothers. The mountains in which this species of Halenia
grows are (or were) clothed in some of the most handsome forests in

1984] Wilbur — Halenia 32

ios)

that part of Mexico with large fir, cypress and pines. We spent a
most delightful week collecting in then relatively undisturbed forest.

SPECIMENS EXAMINED: Jalisco: Volcan Tequila, due S of Tequila, woods of
Quercus, and also Pinus and Arbutus, 1.1 miles from summit on road from Tequila,
2610 m, occasional in shade, 11 Aug 1968, W. R. & C. Anderson 5125 (DUKE, ENCB,
MICH); Sierra de Manantlan (15-20 miles SE of Autl4n) near Aserradero El Cuartén,
2500 m; steep slopes near summits, in pine-oak-fir forests, 2 Nov 1952, McVaugh
13844 (MICH); Sierra de Tequila, 8000 ft, 5 July 1893, Pringle 5465 (GH); hardwood
pine-fir forest in mountains E of Manantlan about 15 miles SSE of Autlan by way of
Chante; about 8300 ft, 25 July 1949, R. L. & C. R. Wilbur 1834 (mic), 1872 (MICH),
30 July 1949, R. L. & C. R. Wilbur 198] (micH).

4. Halenia palmeri A. Gray, Proc. Amer. Acad. 21: 401. 1886.
TYPE: MEXICO; Chihuahua, mountain summits above Bato-
pilas, 8850 feet, Palmer 359 (GH, HOLOTYPE, not seen; NY! PH!
US! ISOTYPES).

Tetragonanthus palmeri (A. Gray) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Annual herb; stems simple or little branched and then
mostly above and only occasionally from the base, striate,
more or less terete, 3-7.7 dm tall. Cauline leaves linear, acute,
sessile, faintly 3-nerved, mostly 2-4.5(7) cm long and 2-4 mm
wide; lower leaves oblanceolate to linear-lanceolate, acute to
obtuse. Inflorescence racemosly cymose, loosely few- to many-
flowered. Calyx lobes (4-)7-12 mm long and 1-2.5 mm wide,
lanceolate to long-triangular, acute, 3-nerved with the midrib
prominent, marginally minutely papillate; corolla 1.0—1.8(—2.2)
cm long, deeply yellowish, the tube 7-10 mm long, the lobes
broadly ovate, acute, slightly auriculate, papillate and the
spurs pendulous, tapering, distally incurved, 1/4 length of the
corolla, about 4~-5(-7) mm long. Filaments linear; anthers
oblong. Capsules 12—15(-22) mm long, ellipsoidal, attenuate,
subfalcate; seeds globose, dark brown, granular.

DISTRIBUTION: mountains of northern and central Mexico
(Chihuahua and Durango).

Halenia palmeri is one of the most easily recognized and
distinctive endemic Mexican species.

REPRESENTATIVE MEXICAN COLLECTIONS: Chihuahua: La Rocha, along tributary of
Rio del Soldado, on N-facing conifer slope, Sierra Mohinora, 14-15 Oct 1959,
Correll & Gentry 23120 (ENCB, LL); Memilichi, Rio Mayo, on drier slopes, 11 Sept
1936, Gentry 2686 (F, GH, MO, US); Marsh Lake, Sierra Madre Mts., 7000 ft, 19 Sept
1903, M. E. Jones s.n. (DS, GH, MO, US); Madera Municipio, scattered in open pine

324 Rhodora [Vol. 86

forest, Arroyo Negro, 7 miles SW of Chuhuichupa, 8 Oct 1939, Muller 3705 (GH, LL,
MICH); Sierra Madre 60 miles S of Guadelupe y Calvo, 7500-8500 ft, 20 Aug 1898,
Nelson 4798 (Gu, Us); Sierra Madres near Colonia Garcia, 7500 ft, 4 Sept 1899,
Townsend & Barber 303 (F, GH, MICH, MO, NY, US). Durango: common in wet
meadow and into pine woods about 2 km E of La Ciudad, 8300 ft, 5 Oct 1970, Bates,
Blanchard & Fryxell 1527 (CAS, ENCB, MICH); high mountain meadows 6 miles E of
La Ciudad, W of El Salto, ca. 9000 feet, 20 Oct 1964, Bratz M680 (GH); steep slope
with Pseudotsuga, Abies, Pinus, Quercus, Arbutus and Juniperus, 54 miles N of
Estacién Coyotes along a lumber road just NW of Guachichilas, 9000 ft, Breedlove
18799 (CAS, MICH); steep moist slope with Pinus, Quercus, Arbutus and Juniperus
along Mexican Hwy. 40, 6 miles W of La Ciudad, 8800 ft., 7 Nov 1970, Breedlove
18866 (CAS, MICH, MO); Sierra Madre Occidental, Mex. Highway 40 12.9 km WSW of
El Salto, 2.4 km W of Lecherias (23° 43’ N, 105° 29’ W), 2520 m, 6 Sept 1975, N. H.
Holmgren & Lowrey 8073 (NCU, NY); Sierra Madre Occidental, about 10 mi W of El
Salto on the Durango-Mazatlan Hwy., rocky rhyolitic hillsides and wet depressions
in open rolling pine woodlands, abundant in meadows and occasional in forests 2650
m, 2 Oct 1962, McVaugh 21741] (CAS, LL, MICH, NY); Common in wet meadows 2 miles
E of La Ciudad on road from Durango to Mazatlan, 8400 feet, Mason 2934 (F, NCU,
NY, PH, TEX); in the more moist openings of the pine forest, about 9100 ft, 30 Sept
1953, Ownbey 199] (F, GH, MICH, NY, US); grassy edge of marsh, El Salto
(Aserraderos) Sierra Madre Occidental, 2530-2540 m, 28 Aug 1934, Pennell 18286 (F,
GH, MICH, NY, PH, US); marshy glade in pineland, El Salto (Aserraderos), 2600-2650
m, | Sept 1934, Pennell 18551 (GH, NY, PH, US); Moist grassy flats along a brook in the
forest belt of the Sierra Madre 5 mi W of E! Salto, 7800 ft, Ripley & Barneby 13987
(CAS, NY).

5. Halenia conzattii Greenm., Fieldiana Bot. 2: 335. 1912. LEcTo-
TYPE: MEXICO: Oaxaca, Cerro San Felipe, Distrito del Centro,
2000 m, 20 Sept 1918, Conzatti 2295 (F!).

Erect, usually branching above, probably biennial herbs with
rather coarse angular, striate or very slightly winged stems
2.0-3.5(5.0) dm tall. Basal leaves ovate-elliptic with petioles nearly
as long as the blade or even longer; cauline leaves shortly winged
petiolate or sessile, elliptic, lanceolate, ovate, acute to obtuse, 1-4
cm long and 0.5-1.3 cm wide, 3-nerved, marginally papillate.
Inflorescence terminal or terminating axillary branches; pedicels
3-15(-20) mm long, 4-angled, smooth to inconspicuously but
copiously papillate. Calyx lobes spatulate, obtuse to rounded, 3-
nerved, marginally papillate, 4-6.5(8.0) mm long and 2-3 mm wide;
corolla 8-12 mm long and 4-7 mm in diameter, green or yellowish
green, the lobes 3-5 mm long, ovate, acute, papillate; spurs slender,
pendulous and distally incurved, 1.5-3 mm long. Stamens 2-5 mm

1984] Wilbur — Halenia 325

long, the filaments linear, the anthers broadly ovate. Capsule 1-1.8
cm long, lanceolate, subfalcate; seeds globose to ovoid, yellow-
brown, granular.

DISTRIBUTION: Common in the higher mountains of Oaxaca,
Mexico.

This distinctive species is readily identified by its broadly
spatulate calyx lobes.

REPRESENTATIVE SPECIMENS: Oaxaca: pine-oak forest along road from Oaxaca to
Guelatao de Juarez and Tuxtepec, 11.2 miles N of intersection with Mexican Hwy.
190, 2410 m, 18 July 1968, W. R. & C. Anderson 4826 (ENCB, MICH); Cerro Grande de
Huancliela, Dist. de Nochixtlan, 2520 m, 13 Oct 1921, Conzatti 4265 (us); between
Mitla and Cuesta, 30 Jan 1966, Ernst 2365 (us); S facing slopes along Hwy from
Oaxaca to Tuxtepec in pine and madrone forest with Lupinus, 12.3 miles N of Jct.
190 & 175 on 175, 15 Aug 1975, LeDoux, Dunn & Wallace 2233 (ENCB, MO, NY);
vicinity of Cerro San Felipe, 9500-11000 ft, Ne/son 1115 (Gu, us); 18 miles SW of the
city of Oaxaca, 7500-9500 ft, 10-20 Sept 1894, Nelson 1340 (us); Sierra de San
Felipe, 10,000 ft, 15 Sept 1894, Pringle 4908 (ENCB, GH, MICH, MO, MSC, NY, PH, US);
Sierra de San Felipe, 10,000 ft, | Sept 1894, C. L. Smith 236 (Mo) and also 664 (F, NY)
and 665 in part (MO).

6. Halenia schiedeana Griseb., Gen. & Sp. Gent. 327. 1839.

Swertia Michauxiana sensu Schlecht. & Cham., Linnaea 5: 122. 1830 but not
Schultes, Syst. Veg. 6: 130. 1820.

Tetragonanthus Schiedeanus (Griseb.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Halenia chlorantha Greenm., Proc. Amer. Acad. 41: 240. 1905. Type: Mexico;
Hidalgo, wet woods near Trinidad Iron Works, 5700 ft, Pringle 8939 (F!
HOLOTYPE; CAS! ENCB! MO! MSC! NY! PH! US! ISOTYPES).

Annual herb with erect, striate to narrowly winged, terete to
weakly 4-angled stems 2-6.5 dm tall, frequently branched above but
simple below. Basal leaves with ovate to broadly elliptic blades
about 2-3 cm long and 10-18 mm wide with petioles |—-3 cm long or
about as long or longer than the blades; cauline leaves ovate to
broadly lanceolate, 3-6 cm long and 1.5-2 cm wide, acute to
strikingly apiculate, 3-5 nerved, shortly to moderately petiolate with
the petioles mostly 3-12 mm long. Inflorescence cymose, terminat-
ing the main stem and the axillary branches; pedicels erect, 7-15 mm
long, narrowly winged and minutely papillate. Calyx lobes lance-
elliptic, marginally densely but microscopically papillate, about 3-5
mm long and 1.5-2.5 mm wide, 3-nerved, acute to even strikingly
apiculate, usually strongly reflexed; corolla 8-11 mm long, greenish,
the tube almost equaling the obovate, abruptly acuminate, often

326 Rhodora [Vol. 86

papillate lobes, the spurs 2-3 mm long, pendulous, tapering, nearly
parallel with the tube, distally slightly incurved. Anthers 2 mm long;
filaments linear. Capsule oblong, subfalcate, about 10-15 mm long;
seeds globose, yellow-brown, granular.

DISTRIBUTION: Moist montane forests of Central Mexico.

REPRESENTATIVE SPECIMENS: Durango: steep slopes at base of Espinazo del Diablo,
4 km NW of Los Angeles along road between Mazatlan and Durango, 2500 m, 28
Oct 1973, Breedlove 35749 (cas); Arroyo del Infierno, deep, well-watered, rough
rocky canyon west of Santa Barbara, about 20 km § of El Salto, 2550-2650 m, 23
Aug 1963, Gordon 54 (MICH); 78.8 miles E of Villa Union, El Espinazo del Diablo,
rocky hillsides with pines, about 7400 feet, 3 Sept 1967, Oliver, Austin, Mac Bryde
807 (MO); Metates, N of Cueva, Sierra Madre Occidental, along stream, 2600-2650
m, 29-30 Aug 1934, Pennell 18448 (GH, PH, US). Hidalgo: wet woods near Trinidad
Iron Works, 5700 ft, 11 July 1904, Pringle 8939 (CAS, ENCB, F, MO, MSC, NY, PH, US).
Veracruz: in the pedrigal of Las Vegas but near La Joya, 7000 ft, 25 June 1945, Sharp
45563 (GH); Plan de Cedefio, Municipio de Acajete, 1750 m, 25 Feb 1981, Ventura A.
18240 (ENCB).

7. Halenia alleniana Standl. ex Wilbur, sp. nov.

Herba annua vel biennis, 3-5 dm alta. Folia basalia oblonga vel
oblanceolata, 3-5 cm longa et 8-18 mm lata; petioli indistincti, +
alati, |-1.5 cm longi. Folia caulina oblonga vel oblanceolata; lamina
3-5 cm longa; petioli indistincti, alati, 1-1.5 cm longi. Lobi calycis
lance-lineares vel late elliptici, 4-7 mm longi et 2-4 mm lati, acuti,
erecti. Lobi corollae late oblongi, acuti; calcaria corollae 2-4 mm
longa pendula, distaliter incurva.

TYPE: MEXICO: Nuevo Leon; Cerro Potosi, NE side of mountain at
abandoned sawmill site, about 2800 m, in open pine forest, 13 Sept
1960, Beaman 4481 (GH, HOLOTYPE; MSC & US, ISOTYPES).

Annual herb with an erect, ridged to very narrowly winged,
weakly 4-angled stem 3-5 dm tall, usually branched above but not
below. Basal leaves with oblong to oblanceolate blades 3-5 cm long
and 8-18 mm wide and gradually tapering to the stem or with rather
indistinct winged petioles 1-2 cm long; cauline leaves oblong to
oblanceolate with the blade 3-5 cm long and tapering to the rather
broad base or very indistinctly petiolate and the petiole strongly
winged and about 1-1.5 cm long, the blade apically acute and
conspicuously veined when dry with 3-5 veins arising from near the
base. Inflorescence cymose, terminating the main stem and the
axillary branches; pedicels strongly ascendent, mostly 1-3 cm long,
narrowly winged and microscopically papillate. Calyx lobes lance-

1984] Wilbur — Halenia 327

linear to more typically broadly elliptic, marginally minutely
papillate, about 4-7 mm long and 2-4 mm wide, acute, erect; corolla
10-14 mm long, broadly cylindric, the tube about half as long as the
entire corolla, the lobes broadly oblong, acute, the spurs 2-4 mm
long, pendent to,somewhat divergent, conical, occasionally distally
slightly incurved. Anthers |.5-2 mm long; filaments 5-7 mm long,
slender. Capsule ellipsoidal, subfalcate, about (12)15-25 mm long,
tapering to the 2-3 mm apex from the 6-8 mm base; seeds
oblongoid, turgid, about 0.8-1 mm long and slightly less, yellowish
to pale reddish brown, smooth.

DISTRIBUTION. Oak and pine forests in the high mountains of
the Mexican states Nuevo Leon, Morelos and Michoacan.

Halenia alleniana has had a confused history in the Mexican flora
as the relatively few collections made have been attributed to H.
brevicornis var. latifolia, Halenia deflexa (J. E. Sm.) Griseb. which
is otherwise known only as a transcontinental species across
southern Canada and the northern United States, and to an
indicated but unpublished species by Paul Standley. More recent
collections, especially those made by John Beaman on Cerro Potosi
in Nuevo Leon, have convinced me that Standley was correct.
Careful and extensive collecting is very much needed in much of
Mexico.

This species is named in memory of Dr. Caroline K. Allen who
published a revision of this most difficult genus fifty years ago and
thus provided the basis for this and all subsequent investigations.

SPECIMENS EXAMINED: Nuevo Leon: Cerro Potosi; NE side of mountain at
abandoned sawmill site, about 2800 m, in open pine forest, 13 Sept 1960, Beaman
448] (GH, HOLOTYPE; MSC & US, ISOTYPES); On moist gravelly arroyo bank, the Cafion
below Las Cafioas on Cerro Potosi, Municipio de Galeana, 20 July 1935; Mueller
2233 (F, GH, MICH); in ponderosa pine forest near microwave station on Cerro Potosi
N of Galeana, Sept 1970, Norris 17610 (cas); Morelos: Valle del Tepeite, 16 Oct 1937,
Lyonnet & Elcoro 1796 (us); Tres Marias Mts, 9500 ft, 16 Dec 1907, Pringle 13971
(GH, Us); Michoacan: Morelia, Rincén, 2300 [meters], IV 1909, Arsene 37 (us).

8. Halenia alata (Mart. & Gal.) Hemsl., Biol. Centr. Amer. Bot. 2:
351. 1882.
Exadenus alatus Mart. & Gal., Bull. Acad. Brux. 11: 372. 1844. Type: Mexico;
Vera Cruz, “Se trouve dans les foréts et sur les rochers trachytiques du pic
d’Orizaba, de 9 a 11,000 pieds Galeotti 722] (BR, HOLOTYPE, not seen; photo of

holotype seen at MO).
Tetragonanthus alatus (Mart. & Gal.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

328 Rhodora [Vol. 86

Perennial herbs from a thickened caudex with |-several, simple,
erect, slightly winged and often quadrate stems (0.3)1-1.5(2.4) dm
tall. Basal leaves numerous and crowded, mostly 2-5 cm long and
3-4 mm wide, obtuse, very narrowly oblanceolate, 3-nerved,
tapering basally into an elongate, slender petiole; lower cauline
leaves subsessile and almost twice as long as the basal, upper cauline
leaves linear, obtuse, sessile, 1-1.8 cm long and mostly 1-3 mm
wide. Inflorescence an umbellate cymose cluster of slightly nodding
flowers; pedicels 4-winged or angulate, (1)4—12(15) mm long. Calyx
tube turbinate to shallowly campanulate, 0.4-0.6(1.0) mm long, 4-
angled; calyx lobes slightly shorter than the corolla, oblong, 3-
nerved, occasionally minutely but inconspicuously papillate, acute,
3-4.2(5.5) mm long and |-1.8(-2.2) mm wide; corolla yellow, almost
rotate, 4.5-6 mm long, the tube about 1/3-1/2 as long as the
corolla, the lobes ovate to oblong, obtuse but often apiculate, erose,
3-4.5 mm long; spurs lacking but represented by short pouch-like
glandular protuberances and these basal nectariferous pouches
slightly protruding and drying darkish brown; filaments linear, the
anthers ovate to oblong. Capsule broadly ovoid to lance-elliptic,
(4.5)6-7(9) mm long; seeds yellow-brown, globose, granular.

DISTRIBUTION. High alpine meadows of Mexico and Guatemala

Halenia alata is a most distinctive species with short pouch-like
nectariferous protuberances instead of spurs.

MEXICAN COLLECTION: Mt. Orizaba, 3000-3125 m, Aug 1839, Linden 934 (MICH).

9. Halenia pringlei B. L. Robinson & Seaton, Proc. Amer. Acad.
28: 113. 1893. Type: Mexico, springy meadows, Sierra de las
Cruces, Pringle 4209 (GH! HOLOTYPE; CAS! F! G! GH! Mo! Msc!
NY! PH! US!. ISOTYPES).

Halenia crassiuscula B. L. Robinson & Seaton, Proc. Amer. Acad. 28: 113. 1893.
Type: Mexico; bare alpine summits, Nevado de Toluca, 14,000 ft, Pringle 4229
(GH! HOLOTYPE; ENCB! MSC! PH! US! ISOTYPES).

Halenia candida Ramirez, Inform. Secret. Foment. Mexico 34. 1895. TYPE:
Sierra de Las Cruces, June 1895, Altarmirano s.n. (US! ISOTYPE).

Perennial herbs with simple to much-branched, + narrowly 4-
angled, erect, slender to compactly bushy stems (2)5-—15(27) cm tall
from a fleshy taproot 3-4 cm long and 3-6 mm in diameter. Basal
leaves present with the blade from narrowly to broadly elliptic to
narrowly oblanceolate to even somewhat oblong, about 1-3.5 cm
long and mostly 4-9 mm wide, apically acute to obtuse, faintly to

1984] Wilbur — Halenia 329

distinctly 3-nerved, tapering into a petiole from about as long as the
blade to as short as | cm; cauline leaves 1-3 pairs, + linear to
narrowly oblanceolate to oblong or elliptic, sessile to basally
narrowed, mostly 1.5-3 cm long and 2-5(8) mm wide. Inflorescence
a terminal or occasionally also lateral, loose to dense umbellate
cyme with + ascendent flowers occasionally slightly nodding after
anthesis; pedicels 2~10(20) mm long. Calyx lobes linear to narrowly
oblong to oblanceolate (3)4—6 mm long, acute to obtuse; indistinctly
3-nerved and marginally microscopically papillate; corolla white
6-10(15) mm long with the tube 3-4 mm long and the lobes elliptic
to oblong-elliptic, acute to acuminate, occasionally slightly erose,
the spurs when present 1.5-7 mm long usually strongly divergent
and often mostly horizontal with distally upturned tips but often
completely lacking or very nearly so especially in late-developing
flowers; stamens 2-3.2 mm long, the filaments linear, the anthers
ovate to oblong, 0.3—1(1.2) mm long. Capsules lanceolate, frequent-
ly subfalcate, acute, exserted beyond the marcescent corulla; seeds +
globose, light yellowish brown, granular.

DISTRIBUTION. Bare volcanic central Mexican alpine summits
and meadows.

Within my concept of H. pringlei, as is shown by its very brief
synonymy, two long-recognized species have been combined in the
present treatment. Both were described by the same authors at the
same time and the numerous specimens in the type collections are so
dissimilar that one can certainly understand their original decision
to recognize two species. The extremes, as is admirably demon-
strated by the fine series of specimens collected by Pringle, are so
unlike that no one would place them together unless confronted by
the specimens that have since accumulated. It must be admitted that
the number of recent collections is far less than would be desirable
to resolve the question and an opportunity to carefully observe the
plants in the field is very much needed. Still I believe that there is no
other course now but to combine the two names since I am unable
to find even strong tendencies let alone consistent differences to
distinguish them.

Halenia crassiuscula was described from specimens gathered on
“bare alpine summits, Nevado de Toluca, 14,000 ft.” and the plants
were as described i.e. much-branched, depressed (“2-4 inches in
height”) and altogether of the dwarfed aspect that alpine plants
often possess. In contrast the type collection of H. pringlei is +

330 Rhodora [Vol. 86

unbranched and slender. The basal leaves of H. crassiuscula are
described as oblanceolate and 3-nerved while those of H. pringlei
are narrowly elliptic to lanceolate and apparently I-nerved. The
difference in venation is one of degree and the distinction disappears
when one examines a larger series of specimens. The original
descriptions and Allen’s key depended greatly upon the supposed
obtuseness of the calycine lobes of H. crassiuscula in contrast to
their acuteness in H. pringlei but this distinction is completely
blurred when a larger series of specimens is examined. In summary,
none of the differences noted or observed holds up. In spite of the
striking differences in the extremes, my study has convinced me that
at present there is no way in which the two species can be
maintained. A larger series of specimens is very much needed and it
is surprising how little additional material has accumulated in the
past fifty years or since Allen examined the available specimens for
her dissertation.

The Flora of Guatemala attributes Halenia crassiuscula to the
high volcanoes of western Guatemala but I believe that the
specimens upon which its presence in that country rests are all
stunted or grazed specimens of Halenia decumbens, a species which
is common there. The corolline spurs of H. crassiuscula tend to be
much more strongly divergent than the + pendent spurs of H.
decumbens. The stunted alpine specimens annotated as H. cras-
siuscula in the Flora of Guatemala and illustrated there (Fieldiana
Bot. 24(8): 318, fig. 87. 1969) possess the normal corolla of H.
decumbens which differ markedly from the strongly divergent spurs
of H. pringlei (including H. crassiuscula). The variation of corolline
spurs within H. pringlei is remarkable. Even in the type of H.
pringlei the corollas vary from spurless to so strongly divergent as to
be mostly horizontal with distally upturned tips but its variability
does not extend to the elongate pendent spurs of H. decumbens (s./.)
so aptly illustrated in the above mentioned figure.

REPRESENTATIVE MEXICAN COLLECTIONS: Veracruz: in turf, Cofre de Perote, 14000
ft, 25 May 1938, Balls 4610 (GH, US); open slopes, Apitza Ixtaccihuatl, 12700 ft, Balls
5123 (Us); in wet meadow E side of Cofre de Perote about 3860 m, 6 Aug 1958,
Beaman 2184 (GH, MSC, US). Mexico: Ojos de Agua, Nevado de Toluca, 12500 ft, 9
July 1938, Balls 4964 (GH, MICH, MSC, US); bog in open sunny glades in forest of
Abies, El Ricén de Selgado, Nevado de Toluca, 10,000 ft, 13 July 1938, Balls 5200
(Us); shore of large lake in the crater of the Nevado de Toluca, about 4140 m, 26 July
1958, Beaman 1883 (GH, MICH, MSC, TEX, WIS, US); alpine meadow, S side of

1984] Wilbur — Halenia 3

ioe)

Ixtaccihuatl, 3950-4000 m, 30 July 1958, Beaman 1953 (MSC); in gravelly soil on rock
slide of S wall of the crater of Nevado de Toluca about 4230 m, 2 July 1960, Beaman
3454 (DUKE, GH, MSC, TEX, US); grassy alpine meadow on W slope S side of
Ixtaccihuatl, 6 July 1960, Beaman 3485 (GH, MSC, TEX); Amecameca, slopes of
Popocatepet! between 10,000 and 12,000 ft, 4 July 1943, Gilly & Dodds 23 (MICH,
MSC); alpine zone on SW slope of Volcan Ixtaccihuatl 5-6 km N of Paso de Cortez,
Iltis, Koeppen & Iltis 1009 (mic, wis); springy alpine meadows, Sierra de las Cruces,
9800 ft, 28 Aug 1904, Pringle 13121 (GH, MO, US); wet meadows, Ixtaccihuatl,
12-13000 ft, 1903, Purpus 318 (Mo, Us); Palomas, Municipio de Iturbide, 3400 m, 18
July 1968, Rzedowski 25930 (ENCB, MICH, MSC). Tlaxcala: turfy slopes, Mt. Malinche,
above San Francisco, 22 June 1938, Balls 4887 (us). Distrito Federal: in alpine
meadow top of Cerro Ajusco, 3937 m, 12 July 1959, Beaman 2776 (Msc).

10. Halenia hintonii Bullock, Hooker’s Icon. Plant. 34: tab. 3399.
1939. [as Hintoni]. Type: MEXICO: Mexico; Distr. de Temas-
caltepec, Cumbre Trojes in Pinus and Alnus forest. Hinton
8273 (K, HOLOTYPE; ENCB! F! GH! UTD! MO! Ny! PH! ISOTYPES).

Perennial herb 1.2-3 dm tall with 4-sided, striate to narrowly
winged stems. Basal leaves scarcely rosulate, the lowermost long
petiolate, spatulate to oblanceolate or elliptic with the blades about
1.5-3(4.5) cm long and 1-1.5(2) cm wide with a rounded apex and
tapering into a winged petiole 2~3.5 cm long; cauline leaves either
sessile or tapering into an indistinct winged petiole, oblanceolate-
elliptic or elliptic or somewhat lanceolate, (2)3-4.5 cm long and
about 1 cm wide, apically acute to obtuse or somewhat rounded.
Inflorescence a terminal or axillary, 3-7-flowered, umbellate cyme
with pedicels quadrate and slightly wing-angled, 0.5-1.5 cm long.
Calyx lobes erect, oblong or oblong-spatulate, 3-4 mm long and
1.5-2 mm wide, apically rounded or obtuse, marginally minutely
papillate, 3-nerved; corolla white, about (6-)7-9 mm long, the tube
3-4 mm long, the lobes erect, ovate to oblong, 3-5 mm long, 2-2.5
wide, apically often shortly apiculate or occasionally broadly
rounded, the spurs about 5-7 mm long, acute, slender, outwardly
spreading or very strongly divergent and distally outwardly curved.
Stamens included; filaments linear, about 1.5 mm long; anthers
reniform-sagittate, about | mm long. Capsule thinly cartilaginous,
about 1.2 cm long and 4 mm wide, apically often somewhat arcuate,
apiculate; seeds ovoid, 16-20 in number, about 1.5 mm long and 1
mm in diameter, smooth, pale yellow.

DISTRIBUTION. Known only from the type locality in the state of
Mexico.

332 Rhodora [Vol. 86

11. Halenia decumbens Benth., Pl. Hartw. 67. 1840. Type: MEXICO;
Oaxaca, “in monte Pelado,” Hartweg 494 (K, HOLOTYPE; G,
photo! Ny! w, photo! IsoTYPEs).

Halenia longicornu Mart. & Gal., Bull. Acad. Brux. 11: 370. 1844. Type:
Mexico; Oaxaca, Croit dans les endroits humides des foréts de pins, chénes et
arbousiers du Cerro de San Filipe, pres d’Oaxaca, de 8,500 a 9,500 pieds,”
Galeotti 7166 (BR, HOLOTYPE, photo!, Mo-fragment!).

Halenia apiculata Mart. & Gal., Bull, Acad. Brux. 11: 371. 1844. Type: Mexico;
Oaxaca, Se trouve avec |’Halenia longicornu au Cerro San Felipe, de 8 a
9,000 pieds, Galeotti 7166 (G, HOLOTYPE).

Halenia plantaginea [var.] apiculata (Mart. & Gal.) Griseb., Linnaea 22: 45.
1849,

Tetragonanthus decumbens (Benth.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Tetragonanthus longicornis (Mart. & Gal.) Kuntze, Rev. Gen. Pl. 2: 431. 1891.

Halenia guatemalensis Loesener, Verhandl. Bot. Vereins. Brandenb. 55: 182.
1913. Type: Guatemala; Huehuetenango, Todos Los Santos, road near
Chiantla, 3000 m, Seler & Seler 2728 (B, not seen, probably destroyed).

Halenia plantaginea var. latifolia Loesener, Verhandl. Bot. Vereins. Brandenb.
55: 182. 1913. Type: Guatemala; Huehuetenango, Todos Los Santos,
Bergwald Oberh., 2800-3000 m, Seler & Seler 3086 (B, not seen, probably
destroyed).

Halenia shannonii Brig., Candollea 4: 320. 1931. Type: Guatemala; Sacatepe-
quez, Volcan de Agua, 12,400 ft. Shannon [J. Donnell Smith #] 36/3 (G,
HOLOTYPE; GH! MO! Ny! US! ISOTYPES).

Halenia shannonii f. compacta C. K. Allen, Ann. Missouri Bot. Gard. 20: 178.
1933. Type: Guatemala; Huehuetenango, mountains above Chinantla, Cook
45 (US! HOLOTYPE).

Halenia guatemalensis var. latifolia (Loesener) C. K. Allen, Ann. Missouri Bot.
Gard. 20: 180. 1933.

Halenia caleoides C. K. Allen, Ann. Missouri Bot. Gard. 20: 173. 1933. Type:
Guatemala; vicinity of Agua, 2700-3000 m, Maxon & Hay 3675 (us!
HOLOTYPE).

Halenia platyphylla C. K. Allen, Ann. Missouri Bot. Gard. 20: 173. 1933. TyPE:
Guatemala: Sacatepequez, Volcan Agua, 2875 m, J. Donnell Smith 2170
(GH! HOLOTYPE; US! ISOTYPE).

Perennial herb (0.5)1.5—3(4.5) dm tall with | to several terete to
quadrate, often slighlty winged stems commonly arising from a
basal cluster of leaves and occasionally with elongate leafy offsets.
Basal leaves elliptical to broadly oval or lanceolate to oblanceolate
or even linear, long petiolate, acute to obtuse, typically 3-nerved but
usually only the midvein prominent, (2-)3-6(12) cm long and
(3-)6—12(-18) mm wide, acuminate to more typically acute to even
obtuse; petiole usually sharply delineated from the blade and often
comprising half the length of the leaf; cauline leaves 1-6 pairs,

1984] Wilbur — Halenia

Ww
no
Ww

sessile to more typically the lower tapering into a petiole, 3-nerved
although only the midvein prominent, (1.5-)2—4(-12) cm long and
5-15(-20) mm wide, elliptic to oblong-lanceolate or lanceolate,
acute to acuminate. Inflorescence an axillary and terminal, umbelli-
form cyme with usually ascendent to erect pedicels 1-2 cm long.
Calyx lobes erect, mostly oblong-elliptic to oblong or rarely
oblanceolate, acute to acuminate and occasionally even apiculate,
marginally minutely papillate, mostly half to three-fourths as long
as the corolla but occasionally as long, 4-7 mm long and 2-2.5 mm
wide; corolla mostly (1.0—)1.2-1.8(2.0) cm long, the lobes 1.5-3 mm
long, yellowish to yellowish green or even green, broadly ovate to
deltoid, usually acute but occasionally acuminate or sometimes
obtuse, about as long as the tube, entire to erose, the spurs pendent
to moderately divergent, slender, 4-7(10) mm long. Capsule
exserted, elliptic to more typically lanceolate, 8-12(-18) mm long
and (3-)4-6 mm wide.

DISTRIBUTION. grassy alpine slopes, talus slides and open
montane woods from southern Mexico (Guerrero, Oaxaca and
Chiapas) and western Guatemala (Chimaltenango, Huehuetenango,
Quetzaltenango, Quiche, Sacatepéquez, San Marcos, Solola, Totoni-
capAn) reportedly ranging from 2000-4000(4200) m.

REPRESENTATIVE MEXICAN COLLECTIONS: Guerrero: open pine forest, Mina,
Teotepec, 3600 m, 17 July 1939, Hinton 14463 (MICH, Us). Oaxaca: abundant on
steep hillside in open Pinus hartwegii forest on Cerro Pelong, 25.5 miles E of Ixtlan
along the Oaxaca-Valle Nacional Hwy about 2950 m, 21 July 1960, Beaman 3672
(DUKE, GH, MSC, TEX, US); middle to upper slopes, Zempoaltepetl, Feb 1937, Camp
2631 (ENCB, F, MICH, NY); in open pine woodland, about 3396 m, vicinity of Cerro
Zempoaltepetl, 23 July 1950, Hallberg 738 (Micu), Cerro de Humo at 10,000 ft,
Comaltepec, 19 May 1971, MacDougal s.n. (F, NY); in pine and oak forest, Cerro
Zempoalteptl, 9500 ft, 6 Aug 1963, Molseed & Hallberg 300 (Mick); above road on
rocky hill, 40 miles S of Valle Nacional, 22 Mar 1978, Poole, Bain & Kerr 1287 (TEX);
floor of dry oak forest, Cerro Cusimulco on northern slope near San Pedro Yolox,
2100 m, 24 June 1939, Schultes 676a (GH, MO); rich cloud forest about 29 miles N of
Ixtlan de Juarez on Hwy 175 about 2800 m, 18 July 1976, Stevens, Donoghue &
Scott 2475 (ENCB, GH, MSC); Sierra Madre del Sur, cloud forest near top of Cerro
Pilon c. 70 mi from Oaxaca, 9000-9100 ft, 20 June 1962, Webster, Hallberg & K. &
L. Miller 11558 (CAS, DUKE, F, GH, MSC, TEX). Chiapas: flat meadow and steep slopes
on the SE side of the summit of Volcan Tacana, 3600 m, 3 Mar 1972, Breedlove
24313 (Ds); small meadow near the summit of Volcan Tacana, 30 July 1972 Breedlove
26707 (ps); steep dry slope on the SE side of Volcan Tacana above Talquian, 3500 m,
11 Nov 1972, Breedlove 29406 (ENCB, DS, LL); steep slope on the N and W slope of
Cerro Mozotal below microwave tower along road from Huixtla to El Porvenir and

334 Rhodora [Vol. 86

Siltepec, 3000 m, 19 Sept 1976, Breedlove 40311 (Ds); slopes 5km N of Mexican Hwy
190 on a logging road from Laguna Chamula microwave station, 2400 m, 15 Oct
1976, Breedlove 40808 (Ds); Siltepec, 9 Aug 1937, Matuda 1596 (F, GH, LL, MICH, MO,
NY, US); in pinelands, Mt. Tacana, 2000-4038 m, Aug 1938, Matuda 2346 (F, GH, LL,
MICH, MO, NY, US); Mt. Male near Porvenir, 3200 m, 6 July 1941, Matuda 4706 (GH,
LL, MO, NY).

12. Halenia plantaginea (H.B.K.) G. Don, Gen. Hist. 4: 177. 1838.

Swertia plantaginea H.B.K., Nov. Gen. & Sp. Pl. 3: 175. 1818. Type: Mexico; in
Andibus Mexicanis, Humboldt & Bonpland s.n. (Pp, photograph seen at MO,
US).

Halenia elongata D. Don ex G. Don, Gen. Hist. 4: 177. 1838. Type: Mexico, (K,
not seen. Described as an annual).

Halenia nudicaulis Mart. & Gal., Bull. Acad. Brux. 11: 371. 1844. Type:
Mexico, Veracruz, Croit dans les forets humides du haut pic d’Orizaba, de 9a
11,000 pieds d’elevation absolue Orizaba, Galeotti 7220 (BR, HOLOTYPE; G, W,
ISOTYPES, not seen).

Halenia nutans Mart. & Gal., Bull. Acad. Brux. 11; 371. 1844. TyPE: Mexico;
Vera Cruz, Croit dans les foréts humides du haut Pic d’Orizaba, de 9 a 11,000
pieds d’elévation absolue Galeotti 7222 (BR, HOLOTYPE; photograph seen, MO).

Tetragonanthus plantagineus (H.B.K.) Kuntze, Rev. Gen. Pl. 2: 431. 1891,

Halenia purpusi Brandegee, Zoe 5: 235. 1906. Type: Mexico; meadows above
timberline, Ixtaccihuatl, Purpus 1760 (CAS, HOLOTYPE not seen; F! Mo! Ny! Us!
ISOTYPES).

Halenia scapiformis Briq., Candollea 4: 322. 1931. Synrypes: Mexico: Sierra
San Pedro Nolasco, Talla, etc. 1843-44, C. Jurgensen 811 & 812 (G, not seen;
photo of 8/2 seen (MO).

Halenia plantaginea f. grandiflora C. K. Allen, Ann. Missouri Bot. Gard. 20:
176. 1933. Type: Mexico: Nevado de Toluca, Pringle 4224 (Mo! HOLOTYPE;
CAS! F! GH! MO! PH! Ny! Us! ISOTYPES).

Perennial herb with 1-several, striate to very narrowly winged, +
quadrangular, erect stems (0.8)1.5—3.0(4.5 dm tall, usually simple
below and often with short, floriferous branches above, arising from
a thick fleshy caudex. Basal leaves numerous, rosulate, the blades
lanceolate or elliptic to ovate, 3-nerved, 3-10(15) cm long and
0.5—1.5 cm wide, obtuse to acute or even apiculate and with slender
petioles (1)2-5 cm long, the cauline leaves 1-2 pairs, sessile, linear to
lanceolate or broadly elliptic, (1-)2-3(-4.5) cm long. Inflorescence a
terminal or axillary umbellate cyme with slightly curved or nodding
slender pedicels (2-)8-18(—22) mm long. Calyx lobes lance-elliptic to
oblong to even narrowly spatulate, 2.5-6 mm long, obtuse to acute
or rarely even abruptly acuminate, |/3-1/2 as long as the corolla,
3-nerved, microscopically marginally papillate; corolla yellow,
(0.7-) 1-1.7 cm long, the tube slightly shorter than the ovate to

1984] Wilbur — Halenia 335

oblong, obtuse to acute lobes, the spurs very slender, pendulous,
distally slightly incurved, mostly 1 / 3-1/2 length of corolla, (2)3-5(7)
mm long. Filaments linear; anthers ovate. Capsule elliptic to
lanceolate in outline, subfalcate, (0.7—) 1.2-1.8 cm long and about 5
mm wide, exserted; seeds subglobose, brown, granular.

DISTRIBUTION. Mountains of central and southern Mexico
(Vera Cruz, Hidalgo, Mexico, Tlaxcala, Puebla, Michoacan,
Morelia and Oaxaca).

The merit of H. nudicaulis as a distinct species separate from H.
plantaginea, although not accepted in this treatment, is still
deserving of critical study. The differences noted to differentiate H.
nudicaulis from its nearest congener are all quantitative. The size of
the plant as well as its leaves, corolla and spur length and the size of
the capsules are all smaller in H. nudicaulis than in H. plantaginea
and the suspicion is natural that this might well be environmentally
induced. Specimens of Halenia nudicaulis all come from the upper
altitudinal limits of H. plantaginea s.l. Opportunity to study
populations of these plants in the field might do much to resolve this
question. In any event the two are very closely related to one
another and my expectation is that further study will demonstrate
that H. nudicaulis is not specifically distinct.

Specimens thought to be representative of H. nudicaulis seem-
ingly were separable from H. plantaginea by the following key:

a) Corolla less than 10 mm long; corolline spurs less than 2.5(-3)

mm long; capsule less than 10(-13) mm long ..........
ere eee Te ee ee eee Tr re H. nudicaulis.

a) Corolla 10 mm long or longer; corolline spurs (2-)3-5(7) mm

long; capsule 1.2-1.8 mm long ......... H. plantaginea.

I believe the available collections suggest that the specimens
referred to H. nudicaulis are merely smaller plants usually found in
less favorable sites usually at the upper altitudinal range of H.
plantaginea s.1.

Allen (1933, p. 177) stated in regard to f. grandiflora that it
“appears to be only a variation due merely to habitat, moisture, or
some nutritional factor” and that “it has no distinctive geographical
distribution.” From her own observations one would conclude that
this variant did not deserve a formal rank. This apparently
represents the opposite extreme of variation within H. plantaginea
than H. nudicaulis. Specimens of f. grandiflora are seemingly from
the more favorable habitats within the range of H. plantaginea.

336 Rhodora [Vol. 86

REPRESENTATIVE SPECIMENS: Tamaulipas: open rocky slopes of Cerro Linadero
[near] Dulces Nombres, Nuevo Leon and just E of border of Tamaulipas-Nuevo
Leon, 2450 m, 10 Aug 1948, Meyer & Rogers 2917 (Mo). Nuevo Leon: open pine
woods, Cerro del Viejo, 15 mi W Dulces Nombres, 18 Aug 1948, Meyer & Rogers
2971 (Mo). Queretaro: bosque de Abies, | km al SW de la cumbre, Cerro Zamorano,
3100 m, 13 Nov 1971, Rzedowski & McVaugh 404 (ENCB, MICH). Hidalgo: Mt.
Orizaba among thin grass in open forests of Pinus, 10,500 ft, 18 Aug 1938, Balls 5281]
(GH, MICH, MSC, US). Sierra de Pachuca, 10,000 ft, 22 Aug 1902, Pringle 11033 (F, Gu,
MICH, MO, NY, US); S slopes Cerro Jihuingo (19° 49-50’ N, 98° 33-34’ W), 2750-3250
m, 26 July 1966, West T-16 (MICH, WIs). Veracruz: Los Pescados, Cofre de Perote,
9500 ft, 12 Sept 1938 Balls 5434 (msc, Us); pine forests, Citlaltepetl, 11-12000 ft,
Purpus 2766 (F, GH, MO, NY). Mexico: open pine woods, Ojos de Agua, Nevado de
Toluca, 12500 ft, 9 July 1938, Balls 4963 (ENCB, GH, MICH, US); upper pine belt,
Volcan de Zinantecatl near Toluca, about 10,000 ft, 23 Aug 1947, Barkely, Webster
& Westlund 44 (F, TEX, US); open pine forest, NW side of Nevado de Toluca between
Loma Alta and Cerro Gordo, 3450 m, 7 Sept 1957, Beaman 1678 (GH, MSC, TEX, US);
in grassy meadow under open pine forest, Telapon N of Iztaccihuatl, S side of
mountain, 3450-3650 m, 4 Sept 1958, Beaman 2435 (GH, MSC, US); forest of pine and
fir, Las Cruces, Temascaltepec, 3350 m, 13 July 1932, Hinton 1034 (F, LL, MO, NY, PH,
us); under thick pine, Monte de Rio Frio, km 49 road from Mexico City to Pueblo,
4000 m, 31 July 1929, Mexia 2693 (CAS, MICH, MO, NY, US); springy alpine meadows,
Sierra de las Cruces, 9800 ft, 28 Aug 1904, Pringle 1312] (CAS, MICH, MO); subalpine
meadow, Popocatepetl, Sept 1908, Purpus 3070 (F, GH, MO, NY, US). Distrito Federal:
Rio Frio, 27 Aug 1930, Russell & Souviron 74 (CAS, GH, US). Tlaxcala: in pine and fir
forest on N slope of Cerro Matlalcueyetl, 26 Sept 1953, Sohns 680 (MICH, US).
Michoacan: in open pine forest, summit of Cerro San Andres, about 12 km N of
Ciudad Hidalgo, 3589 m, 6 Sept 1960, Beaman 427] (GH, MSC, US); pine forest,
Zitacuaro-Cacique, 3325 m, 29 Aug 1938, Hinton 13175 (F, GH, ILL, LL, MICH, NY, PH,
TEX, US); only under pines, summit of Cerro Tancitaro, 12,600 ft, 17 July 1940,
Leavenworth 277 (F, GH, ILL, NY). Morelos: pine-fir forest, Lagunas de Zempoala, 29
July 1957, Straw & Gregory 107] (MicH). Puebla: open forest of Pinus, Tesmala-
quilla, Mt. Orizaba, 10,500 ft, 18 Aug 1938, Balls 528] (GH, MICH, MO, Us); in grassy
pine forest about 3 miles SE of Villa Hidalgo, Pico de Orizaba, about 3780 m, 15 July
1960, Beaman 3648 (GH, MSC, TEX, US); in pine forest about 2 km NE of the Paso de
Cortez Monument, about 3700 m, 10 Sept 1960, Beaman 4445 (GH, MSC, US); Mt.
Orizaba, 12,000 ft, 6 Aug 1891, Seaton 205 (F, GH, NY, US). Oaxaca: vicinity of Cerro
Zempoaltepet! on SE slopes of peak, 2900-3100 m, 10 Aug 1950, Hallberg 919
(MICH); vicinity of Cerro San Felipe, 9500-11000 ft, Ne/son 1096 (Gu, Us); Sierra de
San Felipe, 10,000 ft, 23 June 1894, Pringle 4720 (ENCB, GH, MICH, MO, MSC, NY, PH,
US).

ACKNOWLEDGMENTS

Grateful acknowledgment is made to the National Science
Foundation for a grant to Duke University (DEB-76-10185) that
made this study possible. I am also very grateful to the curators of

1984] Wilbur — Halenia 337

the herbaria listed below whose loans provided the basis for this
study: CAS, DUKE, DS, ENCB, F, GH, ILL, MICH, MO, MSC,
NY, PH, TEX, US, WIS.

LITERATURE CITED

ALLEN, CAROLINE K. 1933. A monograph of the American species of the genus
Halenia. Ann. Missouri Bot. Gard. 20: 119-222. pl. 8-12 + f. 1-7.

DEPARTMENT OF BOTANY
DUKE UNIVERSITY
DURHAM, N. CAROLINA 27706

A REVISION OF THE MEXICAN AND
CENTRAL AMERICAN SPECIES OF
CERASTIUM (CARYOPHYLLACEAE)

Davip A. Goop!

ABSTRACT

Eighteen species of Cerastium are known to occur in Mexico and Central America,
including 16 native and two alien species. The limits of these species are described
and their taxonomy is revised. Three species are newly described.

Key Words: Cerastium, Caryophyllaceae, Mexico, Central America, systematics,
taxonomy

INTRODUCTION

Cerastium, known commonly as “mouse-eared chick weed”, is of
virtually worldwide distribution with its center of diversity in
Eurasia. The number of species included varies (Pax and Hoffmann,
1934; Lawrence, 1951; Willis, 1973). The genus is represented in
Mexico and Central America by 18 species, all but two of which are
native.

While the taxonomy and relationships within Cerastium are not
well understood anywhere, this situation is particularly true in Latin
America. It was in order to improve the understanding and facilitate
the identification of species in Latin America that this study was
conducted. In looking for diagnostic characters, only externally
visible characters were used. Neither cytological nor biochemical
characters were investigated; work in these areas may further
improve the systematics of Cerastium.

Since only morphological features were available for analysis, the
species concept employed is necessarily typological. I have tried to
be conservative in assigning taxonomy. Hence, in such variable
species as Cerastium nutans and C. sinaloense, | have not tried to
give specific or varietal status to forms which may warrant it but for
which insufficient material is available.

Cerastium is in many ways a difficult genus. Species are often
poorly delimited, resulting in the publication of many names which
have since been reduced to synonymy. Relationships among species

'Present address: Museum of Vertebrate Zoology and Department of Zoology,
University of California, Berkeley, CA 94720

339

340 Rhodora [Vol. 86

have also been poorly understood. Pax and Hoffmann (1934)
described a system of subgeneric classification based primarily on
style number and capsule morphology, and included the Mexican
and Central American species in their subgenus Cerastium. While
they listed none of these species in their discussion of sections, it is
clear from character descriptions that all of them, with the possible
exceptions of C. sinaloense, C. sordidum, and C. texanum, belong
to the section Orthodon. Robinson (1894) included C. texanum in
the section Strephodon (C. sinaloense and C. sordidum were as yet
undescribed), but the presence of revolute capsule teeth makes this
inclusion unlikely since Pax and Hoffmann (1934) stated that its
members never have this character.

The literature concerning Mexican and Central American species
of Cerastium consists primarily of lists of species collected on a
single expedition, by a particular collector on several expeditions, or
by several collectors from a single region (Schlechtendal and
Chamisso, 1830; Presl, 1831; Schlechtendal, 1838; Schultz, 1862;
Hemsley, 1878; Britton, 1888; Watson, 1888; Sessé and Mocifio,
1894; Robinson, 1900, 1904; Greenman, 1904; Briquet, 1911;
Standley, 1937; Standley and Steyermark, 1940, 1944). More
recently floras have appeared, some of which treat species of
Cerastium. These include Sanchez S. (1968) and Beaman (1979) for
the Valley of Mexico, Shreve and Wiggins (1964) for the deserts of
Sonora and Baja California, Standley (1937) for Costa Rica, and
Standley and Steyermark (1946) for Guatemala.

CHROMOSOME VARIATION IN CERASTIUM

Published chromosome counts for species of Cerastium range
from 2n = 34 to 2n = 180. By far the majority of these are multiples
of n = 18 (2n = 36, 54, 72, 90, 108, 126, 144, 162, and 180). Brett
(1952) stated that 2n = 36 is probably tetraploid, although 2n = 18 is
unknown, and she concluded that, for the section Orthodon, the
basic chromosome number is probably n = 9. She also stated that
the species in the section Strephodon are mostly 2n = 38 or
multiples thereof and that the basic number for these is probably n
= 19. Many of the species from Mexico and Central America seem
to be 2n = 34 (Beaman et al., 1962), but she made no mention of any
of these.

1984] Good — Cerastium 34]

Known chromosome numbers for the Mexican and Central
American species are listed following the individual species de-
scriptions below. The native species can readily be separated into
two groups, those with 2m = 34 and those with 2n = 36. Some
question arises as to the position of Cerastium nutans in this scheme
since Sdllner (1952, 1954) listed the chromosome number as 2n =
35-36, while Beaman et al. (1962) counted 2n = 34 for a Cerastium
sp. which, upon further examination of the voucher specimen,
turned out to be C. nutans. Séllner, however, did not use Mexican
material for his count and it is possible that the species is
geographically variable in chromosome number.

Beaman et al. (1962) published the only chromosome counts for
several of the Mexican and Central American species of Cerastium.
After study of the voucher specimens for these counts, it has become
apparent that some revisions of their determinations are in order.
Beaman 3745 and Beaman 3896, recorded as C. brachypodum, are
specimens of what is here recognized as C. cuchumatanense.
Beaman 3436 and Beaman 3508, reported as C. orithales, are C.
ramigerum. Beaman 3711, listed as Cerastium sp., is C. nutans.

KEY TO THE SPECIES

1. Capsules straight, teeth revolute; rosette leaves absent, lower
leaves large, crowded and broadly spatulate, upper leaves few
(2-25, usually fewer than 10), smaller and lanceolate .. (2)

2. Petals large, 10.0-17.5 mm long, usually greater than 14.0
mm; Pacific side of the Sierra Madre Occidental.......

Jib wee hea a Reed ens eRe eo ae 13. C. sinaloense

2. Petals smaller, not more than 8.0 mm long; Chihuahuan-
Sonoran Sierra Madre and Baja California......... (3)

3. Stems very slender; stems, leaves and sepals soon turning
light orange-brown with age; cyme 8-25 flowered; flowers

small, petals 4.0-5.5 mm long; capsules short, 4.0-7.0 mm

long, barely exserted beyond the calyx; Baja California

TCT COLT CLT OC LETC ET Ce Te ee TTT 15. C. texanum

3. Stems not particularly thin; stems, leaves and sepals
remaining green; cyme 3-10 flowered; flowers larger,

petals 5.5-8.0 mm long; capsules longer, 8.0-13.5 mm

long, well exserted beyond the calyx; mainland ......

ee TOL TT CRT CET eT eee TET 14. C. sordidum

342 Rhodora [Vol. 86

1. Capsules curved, teeth not revolute; spatulate lower leaves
sometimes present, but usually as rosette leaves....... (4)

4. Lanate pubescence present (at least some), usually best seen at
the nodes and on the rosette leaves (when present) and

lower cauline leaves (this character is sometimes hard to see

in old specimens of C. vulcanicum)........0.0 eee (5)
5. Sepals, pedicels and usually the upper parts of the stem
DISCUIAl=PUDCSCONG cies 80545 oe eens eeee ceed (6)

6. Plants regularly branching above the base; basal rosette
usually lacking; cymes usually more or less many
flowered; petals usually less than, equal to, or only
slightly exceeding the sepals in length; annual .. (7)

7. Plants usually fairly large (over 25 cm tall); internodes
longest just below the inflorescence, becoming
shorter toward the base; pedicels fairly long, 18.0-
30.0 mm long; petals usually equal to or only slightly
exceeding the calyx in length, white; seeds 0.7-1.0
MM: 1. GISMCLED 2c ioe cease eecce as 9. C. nutans

7. Plants smaller (6-30 cm tall); internodes more or less
equal throughout; pedicels short, 5.0-15.5 mm long;
petals shorter than the sepals (except on Pico de
Orizaba and Cofre de Perote and sometimes on
Ixtaccihuat] and Popocatepetl), white or more often
pale green; seeds 0.5-0.7 mm in diameter .......
eee ee rr ere kere eee 18. C. vulcanicum

6. Plants branching usually only at the base; basal rosette
almost always present; cyme relatively few (1-13)
flowered; petals usually well exceeding the sepals in
ION Gt PEPONNIE) oyt iad vsaee nv eeen sone ers (8)

8. Plants usually fairly large (15-35 cm tall); cauline
leaves few (usually 2-3 pairs); pedicels longer, the
lower ones 18.0-32.0 mm long; sepals lanceolate;
seeds 0.9-1.3 mm in diameter; Sierra Madre of
Chihuahua and Durango ........ 8. C. madrense

8. Plants smaller, usually less than 20 cm tall; cauline
leaves numerous, closer together (though still usually
shorter than the internodes); pedicels shorter, though
sometimes to 25.0 mm long; sepals elliptic; seeds
0.7—-0.9 mm in diameter; volcanoes of central Mexico

1984] Good — Cerastium 343

(Distrito Federal, México, Michoacan and Puebla)

eer ce da Rann s 16. C. tolucense

5. Plants nowhere glandular-pubescent................ (9)
9. Plants low, caespitose; internodes usually shorter than
the leaves; cymes dense, flowers crowded at the apex of

each fertile branch; pedicels usually shorter, less than

10.0 mm long; capsules broader .... 11. C. purpusii

9. Plants not very caespitose; internodes usually longer than
the leaves; cymes looser; pedicels usually longer, to 25.0

mm long; capsules less broad ..... 16. C. tolucense

4) Danae Puesce nse ACRE fs cess asarer pisses eds (10)
10. Plants glabrous or only very sparsely pilose, hairs on the
(OCICS SUDIETIENES, 25 coc eeeeeeawes 2. C. barberi

10. Plants more or less densely pubescent............ (11)
11. Pedicels short, the lowermost (longest) usually less than

TOD W0 IONE 4. oa cis ce oc kee ee aue ees (12)

12. Glandular hairs few or lacking; flowers usually many,
very densely crowded at the apex of each fertile
DYENCh: MNOCUCEd SHCCIES «c2.0+ cin nedne cas (13)

13. Bracts scarious margined; flowers larger, sepals
usually 4.5-6.5 mm long; capsules longer, 7.0-
11.5 mm long; seeds larger, 0.5-0.7 mm in
diameter; perennial............ 17. C. triviale
13. Bracts herbaceous; flowers smaller, sepals 3.0—4.5
mm long; capsules shorter, 5.0-9.0 mm long;
seeds smaller, 0.3-0.4 mm in diameter; annual
Maleiieseine sseee sence eas 5. C. glomeratum

12. Glandular hairs predominant, at least on the sepals
and pedicels; flowers not densely crowded, or if
crowded, then flowers few; native species... (14)

}4,. Petals: shorter ‘than the sépals iicaxwassccns (15)
15. Plants larger, usually over 10.0 cm tall; leaves
more or less similar throughout; lowermost

flower well above the ground .............

Ha eaws een eeeeiaeee nes 3. C. brachypodum

15. Plants smaller, usually less than 10.0 cm tall;
lower leaves often more or less spatulate;
lowermost flowers near ground level.......

ieee e ee ES wea HERES Ae Sees 1. C. axillare

344 Rhodora [Vol. 86

14. Petals longer than the sepals.............. (16)
16. Leaves often more or less erect, close to the stem;
pedicels with short, dense, subreflexed hairs;

flowers nodding; petals larger, 6.5-10.5 mm

long, well exserted beyond the calyx; peren-

nial; volcanoes of central Mexico (México,
Puebla, Tlaxcala, and Veracruz)...........

Lica eee te alee wea wees 12. C. ramigerum

16. Leaves more spreading; pedicel hairs not re-
flexed; flowers not nodding; petals shorter,
capsules 4.0-7.0 mm long, often barely ex-

serted beyond the calyx; annual; Sierra de los
Cuchumatanes and various Guatemalan vol-

PAUOCS «cube esa cde 4. C. cuchumatanense

11. Pedicels longer, usually more than 10.0 mm long ...
ee ee eT Ce rT ae (17)
17. Cymes few (1-8) flowered; flowers large, sepals
6.0-7.5 mm long, petals 9.5-18.5 mm long; capsules
longet,. 12,0=] 7.0 mm ION co ov senkvewaees (18)

18. Plants very slender, propped up by the surrounding
vegetation; pedicels with short, dense, subre-
flexed hairs; volcanoes of central Mexico (Méxi-

co, Puebla, and Veracruz) ....10. C. orithales

18. Plants more or less free-standing; pedicel hairs not
reflexed; Sierra de los Cuchumatanes, Guate-

MANA. sucrak ctiawe eens 7. C. juniperorum

17. Cymes usually many (to 40 but usually somewhat
fewer) flowered; flowers smaller, sepals 3.0-6.0 mm
long; capsules shorter, 6.0-10.5 mm long... (19)

19. Leaves acuminate; leaves more or less similar in size
throughout; perennial; volcanoes of Guatemala
(pPEELELAA SAAR OER POTS 6. C. guatemalense

19. Leaves not acuminate (except in a form found in
central Mexico); leaves largest at the base,
smaller upward; annual......... 9. C. nutans

TAXONOMIC TREATMENT

GENERIC DESCRIPTION
Cerastium Linnaeus, Sp. Pl., ed. 1, p. 437. 1753.

1984] Good — Cerastium 345

TYPE SPECIES: Cerastium arvense L., lectotype of Britton and
Brown (1913).

Centunculus Adanson, Fam. 2: 256. 1763.
Prevotia Adanson, Fam. 2: 256. 1763.
Moenchia Ehrhart, Beitr. 2: 177. 1788.
Quaternella Ehrhart, Beitr. 4: 149. 1789.
Doerriena Borkhausen in Rhein, Magaz. 2: 528. 1793.
Myosotis Tournefort ex Moench, Meth., p. 224. 1794.
Esmarchia Reichenbach, Fl. Germ. Excurs., p. 793. 1832.
Dufourea Grenier, Act. Soc. Linn. Bord., p. 25. 1837.
Doerriera Steudel, Nom., ed. 2, 1: 522. 1840.
Prevotia Steudel, Nom., ed. 2, 2: 394. 1840.
Dichodon Bartling ex Reichenbach, Nom., p. 205. 1841.
Pentaple Reichenbach, Ic. Fl. Germ. 37: 227. 1841.
Leucodonium Opiz, Seznam, p. 59. 1852.

(Synonymy according to Jackson, 1895; and Pax and Hoffman, 1934).

Plants herbaceous, low and caespitose to fairly tall and erect,
often decumbent; annual or perennial; stems almost always
pubescent, either glandular or not; leaves variously linear to
elliptical or spatulate, opposite, exstipulate, entire, acuminate, acute
or obtuse, variously pubescent, usually without a petiole; basal
rosette present or lacking; cauline leaves few to many, usually
shorter then the rosette leaves when the latter are present;
inflorescence a cyme, either compact or loose; flowers various in
size, from very small (sepals 2.5 mm long) to larger (sepals over 8
mm long); petals shorter than to more than twice as long as the
sepals; sepals 5, rarely 4, lanceolate to elliptic, acute, scarious
margined, usually pubescent; petals 5, rarely 4 or absent, usually
white, variously bifid; stamens 10, rarely 5 or 4, shorter than the
petals; ovary superior, carpels 5, rarely 4 or 3; styles 5, rarely 4 or 3,
opposite the sepals; capsules usually well exserted beyond the calyx
at maturity, opening apically by twice as many teeth as there were
styles; seeds many, small (0.3-1.2 mm in diameter), variously
tuberculate.

In Mexico and Central America only the most frequent numbers
of flower parts are found: 5 sepals, 5 petals, 10 stamens, and 5 styles.

The name Cerastium is derived from the Greek “cerastes”
meaning “horned”, in reference to the shape of the capsule.

346 Rhodora [Vol. 86

TREATMENT OF INDIVIDUAL TAXA

1. Cerastium axillare Correll, Brittonia 18: 308. 1966. Type:
UNITED STATES. Texas: Jeff Davis Co., Little Aguja Canyon,
Buffalo Trail Scout Camp Area, Davis Mountains, Correll &
Ogden 25069 (HOLOTYPE: LL).

Plant annual; branches few, usually erect; stems short (usually
less than 10 cm tall), often more or less caespitose, glandular-pilose;
lower leaves somewhat spatulate, those above lanceolate to elliptic,
7.0-20.0 mm long, 1.0-6.0 mm wide, acute to obtuse, more or less
glandular-pilose; basal rosette lacking; cymes 5-20 flowered, bracts
not scarious margined, lowermost flowers often close to the ground;
pedicels short, 2.0-5.0 mm long, glandular-pilose; sepals lanceolate,
3.0-5.0 mm long, 1.0—1.5 mm wide, acute, scarious-margined except
at the apex, viscid-glandular; petals shorter than the sepals, 3.0-4.5
mm long, bifid about 1/4 of their length, white; filiments 2.5-3.0
mm long, anthers 0.3 mm long; styles 1.0-1.2 mm long; capsules
6.0-11.7 mm long, 2.0-2.6 mm wide; seeds 0.4—0.7 mm in diameter,
light brown, tuberculate; chromosome number unknown.

DISCUSSION: Cerastium axillare is known in the United States
only from the Trans-Pecos of Texas. In Mexico it has been collected
in Chihuahua, Coahuila, and Durango (Figure 1A). No habitat
information is given on any of the Mexican specimens but Correll
and Johnson (1970) listed “open forested slopes, rocky hills and
grasslands in the mountains.”

SPECIMENS EXAMINED: Mexico. CHIHUAHUA: 10-15 mi southwest of Nueva
Casas Grandes, Correll & Johnston 21694 (LL); Puerta de San Diego, alt. ca. 6500 ft,
Hartman 637 (GH, NY, US). COAHUILA: 45 mi east of Saltillo, Palmer s.n. (GH).
DURANGO: Otinapa, Palmer 375 (F, GH, MO, NY, UC, US). United States:
TEXAS: Jeff Davis Co., Little Aguja Canyon, Buffalo Trail Scout Camp Area,
Davis Mountains, Correll & Ogden 25069 (LL, holotype).

2. Cerastium barberi Robinson, Proc. Boston Soc. Nat. Hist. 31:
266. 1904. Type: MExICO. Chihuahua: In the Sierra Madres
near Colonia Garcia, 27 June 1899, Townsend & Barber 453
(HOLOTYPE: GH; ISOTYPES: MO, US).

Plant perennial; stems 10-35 cm tall, sometimes branched basally;
branches erect, glabrous or only extremely sparsely pilose; inter-

1984] Good — Cerastium 347

nodes quite long just below the inflorescence, grading to very short
at base; leaves linear to lanceolate, 8.0-20.0 mm long, 1.0-3.5 mm
wide, acute, virtually glabrous; cymes 1-6 flowered, bracts not
scarious-margined; pedicels 2.0-14.6 mm long, hooked in fruit,
almost glabrous or with only very short reflexed hairs; sepals
lanceolate to ovate, 4.5-5.0 mm long, 1.1-1.7 mm wide, acute,
scarious-margined, glabrous or with a few ciliate hairs on the
margin at the base; petals 5.5-6.5 mm long, bifid about 1/4 of their
length, white; filaments 5.0-6.5 mm long, anthers 0.7 mm long;
styles 2.0-2.2 mm long; capsules 8.0-10.0 mm long, 2.3-3.1 mm
wide, curved; seeds 0.8-1.0 mm in diameter, tuberculate; chromo-
some number unknown.

Figure |. Distribution in Mexico and Central America of Cerastium. A. C.
axillare (closed stars), C. guatemalense (open stars), and C. brachypodum (closed
circles); B. C. barberi (closed stars), C. cuchumatanense (open stars), and C.
madrense (closed circles); C. C. juniperorum (triangle), C. sinaloense (closed stars),
C. texanum (open stars), and C. sordidum (closed circles); and D. C. nutans. There is
also a single specimen of C. guatemalense from Costa Rica.

348 Rhodora [Vol. 86

Discussion: Cerastium barberi is known primarily from two
collections both apparently made on the same expedition and in the
same place; in the Sierra Madre Occidental near Colonia Garcia,
Chihuahua (Figure 1B). The elevation was about 2290 m, but no
other habitat information is available. Another specimen, collected
at Yepachic, Chihuahua, also appears to be referable to Cerastium
barberi although it is slightly more pubescent.

SPECIMENS EXAMINED: Mexico. CHIHUAHUA: Near Colonia Garcia, Nelson 6167
(GH, US); Yepachic, Pennington 49 (TEX); In the Sierra Madres near Colonia
Garcia, alt. ca. 7500 ft, Townsend & Barber 453 (GH, holotype; MO, US, isotypes).

3. Cerastium brachypodum (Engelmann ex Gray) Robinson, Proc.
Amer. Acad. Arts 29: 277. 1894. Type: “w. Illinois and
southwestward” (not seen).

Cerastium nutans var. brachypodum Engelmann ex Gray, Man., ed. 5, p.
94. 1867. Type: same as above.

Cerastium nutans var. genuinum (lusus 2) Rohrbach, Linnaea 37: 289, 1873.
Type: “Habitat in media parte totus fere Americae borealis usque ad finis
Mexicanos” (not seen).

Plant annual; usually diffuse, branches few to many, usually more
or less erect but sometimes decumbent; stems to 40 cm high but
usually less than 20 cm, very small in alpine and other harsh
environments, glandular-pilose, often very viscid; leaves usually
similar throughout, lanceolate to elliptic, 7.0-21.0 mm long (usually
shorter than the adjacent internode), 1.2-5.0 mm wide, acute, rarely
obtuse, more or less glandular-pilose; basal rosette lacking; cymes
5-30 flowered, bracts not scarious-margined; pedicels short, 2.1-5.8
mm long, glandular-pilose; sepals lanceolate, 3.2-5.8 mm long,
1.0-1.8 mm wide, acute, scarious-margined except at the apex,
viscid-glandular; petals shorter than the sepals, 3.0-4.5 mm long,
bifid about 1/4 of their length, white; filaments 2.6-3.2 mm long,
anthers 0.3 mm long; styles 1.0-1.2 mm long; capsules 6.3-12.8 mm
long, 2.0-2.8 mm wide; seeds 0.4-0.7 mm in diameter, light brown,
tuberculate; chromosome number unknown.

DISCUSSION: Cerastium brachypodum is a species of varied
habitat. In Mexico it has been collected at elevations from 2000 to
over 3800 m in such habitats as Pinus and Populus woodlands and

1984] Good — Cerastium 349

alpine and subalpine meadows. It is often a species of disturbed
areas such as trails and roadsides. Found in the United States and
Canada north to southeastern Virginia, Illinois, North Dakota,
Alberta, and Washington (Fernald 1950), it ranges into Mexico as
far south as the state of México. It has been collected in the states of
Chihuahua, Coahuila, Durango, Hidalgo, Mexico, Nuevo Leon,
Queretaro, San Luis Potosi, Tamaulipas, and Zacatecas (Figure
1A).

Cerastium brachypodum is often considered to be a form of C.
nutans and many recent floras and checklists have listed it as such.
However, at least in the area covered by the present work, the two
forms are sympatric and easily distinguishable. I therefore retain
their specific status.

REPRESENTATIVE SPECIMENS: Mexico. CHIHUAHUA: Majalca, LeSueur 624 (F,
GH, MO, TEX, UC, US). COAHUILA: Municipio de Ocampo, Sierra Maderas del
Carmen, at Ojo del Negro below and west of Campo 0, alt. ca. 2100 m, Riskind &
Patterson 1801 (LL). DURANGO: Tejamen, Palmer 542 (GH, US). HIDALGO:
Sierra de Pachuca, alt. ca. 10,000 ft, Pringle 11321 (ENCB, F, GH, US). MEXICO:
Cafiada Alcalican, La Joya side of Ixtaccihuatl, alt. ca. 3850 m, Murry 47 (MSC); on
road to Nevado de Toluca, alt. ca. 3540 m, Murray 52 (MSC). NUEVO LEON: Pefia
Nevada, alt. ca. 3200 m, Good 1006 (MSC); Cerro Potosi, just above timberline,
Mueller 2245 (F, GH, MICH, MO); QUERETARO: Cerro Zamorano, | km
southwest of the cumbre, alt. ca. 3100 m, McVaugh 466 (ENCB). SAN LUIS
POTOSI: Sierra de Alvarez, cerca de Puerto Huerta, alt. ca. 2300 m, Rzedowski 4118
(ENCB, MSC). TAMAULIPAS: between Marcella and Hermosa, Stanford et al.
2650a (NY, US). ZACATECAS: Alt. 7000-8000 ft, Purpus 415 (MO, UC, US).

4. Cerastium cuchumatanense D. A. Good, sp. nov. TYPE:
GUATEMALA. Huehuetenango: Sierra de los Cuchumatanes,
immediately north of Tojiah at km. 322 on Ruta Nacional 9N,
alt. ca. 3200 m, | August 1960, Beaman 389] (HOLOTYPE: MSC;
ISOTYPES: GH, TEX, US) (Figure 2).

Planta annua. Caules usque ad 25 cm alti sed plerumque minus
quam 10-15 cm, plerumque basi ramosi, rare superne. Rami erecti
vel saepius ascendentes, glanduloso-pilosi. Folia omnia similaria,
anguste ad late lanceolata, foliis in ramis sterilibus saepe ovatis vel
leviter spathulatis, 6.0-11.2 mm longa (internodiis contiguis lon-
giora vel breviora), 1.8-3.0 mm lata, acuta vel obtusa, interdum plus
minusve ad apicem rotundata, plus minusve dense pilosa. Rosula

Figure 2. Illustrations of Cerastium. (Left to right) C. tolucense, C. sinaloense, and C. cuchumatanense. The scale in the

upper right represents 5 cm.

OSE

elopoyy

98 10A]

1984] Good — Cerastium 351

basalis absens. Cymae plerumque 3-8 floribus, floribus ad apicem
uniuscujusque rami fertilis aggregatis. Pedicelli brevissimi,!.2-5.0
mm longi, glanduloso-pilosi. Sepala lanceolata usque ovata, 3.0-5.0
mm longa, |.1-1.6 mm lata, glandulifera, praeter apicem anguste
scarioso-marginata. Petala 4.2-6.2 mm longa, quam sepala lon-
giora, bifida per circa 1/4 longitudinem, alba. Filamenta 3.0 mm
longa. Antherae 0.3 mm longae. Styli 1.3 mm longi. Capsulae
plerumque tantum vix ultra calycem exsertae, 4.0-7.0 mm longae,
1.6—2.2 mm latae, curvae. Semina 0.6-0.9 mm diametro, aurantiaca
usque rubro-brunnea, dense tuberculata. Chromosomatum nu-
merus 2n = 34 (Beaman et al., 1962).

Cerastium cuchumatanense is characterized by the following set
of characters: basal rosette lacking; cauline leaves not ascending;
pubescence present, glandular, not lanate; pedicels short, with hairs
not reflexed; flowers not nodding; petals longer than the sepals;
capsules curved, usually only barely exserted beyond the calyx.

DISCUSSION: Cerastium cuchumatanense is a species primarily of
the high Ilanos of the Sierra de los Cuchumatanes, Guatemala, but
is also found on the Tecum Uman Ridge and above timberline on
Volcan de Agua and Volcan Acatenango, also in Guatemala (Figure
1B). It is restricted to the subalpine meadows above about 2700 m.
There may be some preference for slightly moist sites.

The consensus to date (Standley and Steyermark, 1946; Beaman
et al., 1962; Beaman, 1979) has been that the specimens here referred
to as Cerastium cuchumatanense are C. brachypodum. This
consensus seems to be based primarily on the fact that both C.
brachypodum and cuchumatanense have small flowers and un-
usually short pedicels. Standley and Steyermark, however, stated
that “more ample specimens may show the Guatemalan plant to be
an undescribed species, since it does not appear to be referable to
any other species known from Mexico.”

There is a gap of approximately 980 km between the southern
limit of Cerastium brachypodum near Nevado de Toluca, México,
and the Sierra de los Cuchumatanes, Guatemala, the northermost
locality for C. cuchumatanense. Ecologically, while the distributions
of the two species overlap with both being found in high mountain
meadows, C. brachypodum has a much broader range of habitats,
often occurring in woodlands and other situations in which C.

352 Rhodora [Vol. 86

cuchumatanense has never been collected. For these reasons and
because of the morphological differences obvious when descriptions
of the two species are compared, there is no doubt that the
Guatemalan specimens are indeed distinct from Cerastium brachy-
podum.

SPECIMENS EXAMINED: Guatemala. CHIMALTENANGO: Volcan Acatenango,
northwest side of peak, alt. ca. 3825 m, Beaman 3271 (MSC). HUEHUETENANGO:
Sierra de los Cuchumatanes, at Chémal at km. 316.8 on Ruta Nacional 9N, alt. ca.
3310 m, Beaman 3089 (GH, MSC); Sierra de los Cuchumatanes, between Tojiah and
Chémal at km. 319.5 on Ruta Nacional 9N, alt. ca. 3380 m, Beaman 3745 (MSC);
Sierra de los Cuchumatanes, immediately north of Tojiah at km. 322 on Ruta
Nacional 9N, alt. ca. 3200 m, Beaman 3891 (GH, MSC, TEX, US); Sierra de los
Cuchumatanes, immediately north of Tojiah at km. 322 on Ruta Nacional 9N, alt.
ca. 3200 m, Beaman 3896 (MSC, holotype; GH, TEX, US, isotypes); Sierra de los
Cuchumatanes, large meadow 7 mi north of Santa Eulalia along road to San Mateo
Ixtatan, Municipio de Santa Eulalia, alt. ca. 9100 ft, Breedlove 11521 (DS, LL,
MICH, US); Sierra de los Cuchumatanes, on road from Huehuetenango to San Juan
Ixcoy, south of road to Todos Santos Cuchumatanes, alt. ca. 3450 m, Good 1022
(MSC); Sierra de los Cuchumatanes, open llano to the west of road to San Juan
Ixcoy, alt. ca. 3600 m, Good 1026 (MSC); Sierra de los Cuchumatanes, between
Paquix and Llanos San Miguel, road to San Juan Ixcoy, alt. ca. 3300 m, Molina R.
21242 (F, NY); Chémal, Sierra de los Cuchumatanes, alt. ca. 4000 m, Molina R. &
Molina 26415 (F); between Capzin and km. 143 on way to San Juan Ixcoy, Sierra de
los Cuchumatanes, alt. ca. 3100 m, Molina R. & Molina 26446 (F); Region of
Chémal, Sierra de los Cuchumatanes, alt. ca. 3300 m, Standley 81085 (F); between
Tojquia and Chémal, Sierra de los Cuchumatanes, alt. 3700-3750 m, Steyermark
50237 (F, US). SACATEPEQUEZ: Volcan de Agua, summit of the south rim of
crater, alt. ca. 3750 m, Beaman 2918 (MSC);Volcan de Agua, on floor of crater, alt.
ca. 3670 m, Beaman 2946 (GH, MSC); upper slopes of Volcano Agua, near crater,
alt. ca. 3600 m, Harmon 3669 (ENCB). TOTONICAPAN: on the Tecum Uman
Ridge at km. 154 on Ruta Nacional |, ca. 20 km east of Totonicapan, alt. ca. 3340 m,
Beaman 4183 (GH, MSC, UC, US); 4.5-5 mi southeast of Totonicapén, alt.
9600-9800 ft, Webster et al. 11781a (F).

5. Cerastium glomeratum Thuillier, Fl. Paris, ed. 2, p. 226. 1799.
TYPE: FRANCE: “Se treuve dans le bois de Boulogne; a
Vinncennes et ailleurs” (not seen).

Cerastium viscosum Linnaeus (nom. ambig.), Sp. Pl., ed. 1, p.437. 1753. Type:
“Habitat in Europae pratis macilentis” (photograph seen).

Plant annual; stems decumbent or erect, sometimes caespitose,
2.5-30 cm tall, very much branched at the base, little otherwise;
branches pilose; internodes usually longer distally than basally,
longer or shorter than the leaves; leave more or less similar

1984] Good — Cerastium 353

throughout, perhaps slightly smaller above, broadly elliptic to
orbicular or spatulate, 5.0-20.0 mm long, 3.0-12.0 mm wide,
obtuse, pilose; basal rosette lacking; cymes many flowered, flowers
very crowded terminally; pedicels very short, 1.0-5.0 mm long,
pilose, sometimes slightly glandular; sepals lanceolate, 2.8-5.1 mm
long, 0.5-1.6 mm wide, acute, scarious margined, pilose, sometimes
glandular; petals shorter than or equal to the sepals, bifid about 1/4
of their length, white; filaments 1.8-2.3 mm long, anthers 0.1 mm
long; styles 0.8—1.1 mm long; capsules 5.0-8.8 mm long, 1.3-1.8 mm
wide, curved; seeds 0.3-0.4 mm in diameter, finely tuberculate;
chromosome number 2n = 72 (Rohweder, 1937, 1939; Heiser and
Whittaker, 1948; Brett, 1952, 1955; Séllner, 1952, 1954; Love and
Léve, 1956; Blackburn and Morton, 1957; Huynh, 1965; Gadella
and Kliphius, 1966; Favarger, 1969; Love and Kjellquist, 1974). No
counts have been published for Mexican or Central American
material.

DISCUSSION: Cerastium glomeratum and C. triviale are the only
two introduced species of Cerastium in Mexico or Central America.
While native to Europe, C. glomeratum has become established
almost worldwide and is found in North America from Florida,
Texas, and California north to southeastern Massachusetts, New
York, Ohio, Illinois, South Dakota, and British Columbia (Fernald,
1950), as well as in central Mexico (Distrito Federal, Hidalgo,
Mexico, Michoacan, and Veracruz), southern Mexico (Chiapas),
Guatemala (Alta Verapaz, Chimaltenango, Guatemala, Huehue-
tenango, Jalapa, Quetzaltenango, Quiche, Sacateépequez, San
Marcos, and Solola), Honduras (Itibuca), Nicaragua (Matagalpa),
Costa Rica (Alajuela, Cartago, and San José), and Panama
(Chiriqui) (Figure 3A). It is found in a variety of habitats at
elevations from 1200 to 3700 m. Such habitats include roadsides,
fields, woodlands, volcanic rocks, cornfields, gardens, etc.

Cerastium glomeratum was originally described in 1753 by
Linnaeus as C. viscosum. However, examination of photographs of
the type specimen and that of C. vulgatum, described by Linnaeus in
1762, indicate that these specimens were somehow reversed in the
Linnaean herbarium (i.e. the type description of one matches the
type specimen of the other). This mixup has resulted in enough
confusion to warrant declaring both names ambiguous. Many
recent workers (cf. Jalas et al. 1964) have taken this position.

354 Rhodora [Vol. 86

Figure 3. Distribution in Mexico and Central America of Cerastium. A. C.
glomeratum; and B. C. triviale.

1984] Good — Cerastium 355

Cerastium viscosum then becomes C. glomeratum and C. vulgatum
becomes C. triviale (or C. fontanum subsp. triviale; see below).

REPRESENTATIVE SPECIMENS: Costa Rica. ALAJUELA: Region of Zaracero,
Guadeloupe de Zaracero, alt. ca. 4500 ft, Smith A366 (F). CARTAGO: Volcan de
Turrialba, alt. ca. 2600 m, Pittier 7556 (NY, US); Birris, south slope of Volcdn de
Irazu, Standley 35438 (US); SAN JOSE: La Palma, alt. ca. 1600 m, Standley 38043
(US); near Laguna de Escuandra, northeast of El Copey, alt. 2000-2200 m, Standley
41996 (US). Guatemala. ALTA VERAPAZ: Coban, von Tuerckheim 1334 (GH, NY,
US). CHIMALTENANGO: Cerro de Tecpam, region of Santa Elena, alt. ca. 2700
m, Standley 58679 (F). GUATEMALA: Volcan de Pacaya, above las Calderas, alt.
1800-2400 m, Standley 58364 (F). HUEHUETENANGO: Sierra de los Cuchu-
matanes, between Paquix and Llanos San Miguel, road to San Juan Ixcoy, alt. ca.
3300 m, Molina R. 21235 (F, NY). JALAPA: on the summit of Montafia
Miramundo, between Jalapa and Mataquescuintla, alt. 2000-2500 m, Steyermark
32611 (F). QUETZALTENANGO: Cerro la Pedrera, south of Quetzaltenango, alt.
ca. 2400 m, Standley 65520 (F); Olintepeque, alt. ca. 2415 m, Standley 66002 (F).
QUICHE: south of Chichicastenango, alt. 1830-1880 m, Standley 62390 (F).
SACATEPEQUEZ: Volcan de Agua, above Santa Maria, alt. 7000-8000 ft, Bell &
Duke 16969 (MO). SAN MARCOS: Barrancas south and west of town of
Tajumulco, northwest slopes of Volcan Tajumulco, alt. 2300-2500 m, Stevermark
36535 (F); Steyermark 35673 (F). SOLOLA: near Maria Tecum, Sierra Madre
Mountains, ca. 10-12 km northwest of Los Encuentros, alt. ca. 3000 m, Williams et
al. 27319 (F). Honduras. INTUBUCA: vicinity of La Esperanza and Intubuca, alt.
1500-1600 m, Standley 25434 (F). Mexico. BAJA CALIFORNIA NORTE:
Guadeloupe Island, alt. ca. 700 m, Moran 17304 (ENCB, LL). CHIAPAS: north and
west slope of Cerro Mozotal below the microwave tower along the road from Huixtla
to El Porvenir and Siltepec, Municipio de Motozintla de Mendoza, alt. ca. 3000 m,
Breedlove 40319 (DS,MP); north end of San Cristébal las Casas, alt. ca. 7100 ft,
Breedlove & Raven 8264 (DS). DISTRITO FEDERAL: ladera oriental del Cerro
Ajusco, alt. ca. 3250 m, Arrendondo 69 (ENCB); along the road up the Cafiada del
Rio Magdelina near the 4th dynamo, alt. ca. 3000 m, Murry 1] (MSC); MEXICO:
Ocotepec, Distrito Temascaltepec, alt. ca. 1500 m, Hinton 2888 (GH, MO, NY, US).
MICHOACAN: near the microwave tower on Cerro Burro, 7 km south of Opopeo,
alt. ca. 3170 m, Murry 58 (MSC). VERACRUZ: La Joya, alt. ca. 2100 m, Ventura A.
11061 (ENCB). Nicaragua. MATAGALPA: road to La Fundadora, cloud forest area
north of Santa Maria de Ostuma, Cordillera Central de Nicaragua, alt. 1300-1500 m,
Williams et al. 24948 (F, NY). Panama. CHIRIQUI: vicinity of Boquete, Finca
Collins, “El Vilo”, alt. ca. 6150 ft, Stern et al. s. n. (MICH, US).

6. Cerastium guatemalense Standley, Field Mus. Nat. Hist., Bot.
Ser. 17: 244. 1937. Type: GUATEMALA. Chimaltenango:
Volcan de Agua, 22 July 1937, Johnson 816 (HOLOTYPE: F).

Plant perennial; stems to 45 cm tall, often more or less much
‘ranched basally and above; branches erect or decumbent, densely

356 Rhodora [Vol. 86

glandular-pilose; internodes more or less equal except at the base,
where shorter; leaves more or less similar throughout or the lower
ones somewhat shorter, linear to lanceolate, long, widest at the base,
narrowing to an attenuate tip, 15.0-48.0 mm long, 2.0-5.0 mm wide,
glandular-pilose, particularly on the lower surface; basal rosette
lacking; cymes 3-40 flowered, usually 3-15; bracts not scarious-
margined; lowermost (longest) pedicels 14.0-35.0 mm long, upper
ones shorter, all more or less hooked in fruit, glandular-pilose;
sepals lanceolate, 4.4-6.0 mm long, 1.0-1.7 mm wide, acute,
scarious-margined except at the apex, glandular; petals 6.0-7.1 mm
long, bifid about 1/8 of their length, white; filaments 3.2-3.5 mm
long, anthers 0.3 mm long; styles 1.9-2.1 mm long; capsules 7.8-11.8
mm long, 2.0-2.8 mm wide, curved; seeds tuberculate, 0.8-1.0 mm
in diameter; chromosome number unknown.

DISCUSSION: Cerastium guatemalense is known from open pine
forests and adjacent subalpine meadows between 2130 and 4600 m
elevation on the following Guatemalan volcanoes: Acatenango,
Agua, Fuego, Santa Maria, Santo Tomas, Tacana, and Tajumulco.
Outside of Guatemala three collections have been made: Stevens et
al. 2427 from the Chiapas side of Volcan Tacana, Breedlove 40315
from nearby Cerro Mozotal, Chiapas, and Burger & Gomez P. 8216
from the Department of San José, Costa Rica. This last specimen is
considerably disjunct from the known range of Cerastium guate-
malense (Figure 1A).

REPRESENTATIVE SPECIMENS: Costa Rica. SAN JOSE: along the trail to the Valle de
los Conejos along the upper Rio Talari, alt. 3250-3450 m, Burger & Gomez P. 8216
(F, MO). Guatemala. CHIMALTENANGO: Volcan de Fuego, north side of
mountain on Meseta, alt. ca. 3500 m, Beaman 4047 (GH, MSC); slopes of Volcan de
Acatenango, above Las Calderas, alt. 2700-2900 m, Standley 61893 (F). QUETZAL-
TENANGO: summit of Volcan Santa Maria, alt. ca. 12,400 ft, Skutch 834 (F, GH,
US); Volcan Santo Tomas, alt. 3000-3300 m, Steyermark 34882 (F). SACATE-
PEQUEZ: Volcan de Agua, Johnson 816 (F, holotype). SAN MARCOS: Volcan
Tajumulco, east side of peak, at timberline, alt. ca. 4050 m, Beaman 3176 (GH,
MSC); between Sibinal and summit of Volcan Tacana, lower slopes above ridge of
La Vega, alt. 2500-4400 m, Steyermark 36088 (F). Mexico. CHIAPAS: on the north
and west slope of Cerro Mozotal below the microwave tower along the road from
Huixtla to El Porvenir and Siltepec, Municipio de Motozintla de Mendoza, alt. ca.
3000 m, Breedlove 40315 (DS); on southeast slope of Volcan Tacané, alt. ca. 3550 m,
Stephens et al. 2427 (MSC).

1984] Good — Cerastium 357

7. Cerastium juniperorum Standley & Steyermark, Field Mus. Nat.
Hist., Bot. Ser. 23: 51. 1944. Type: GUATEMALA. Huehue-
tenango: alpine areas in the vicinity of Tumina, Sierra de los
Cuchumatanes, alt. 3400-3500 m, 7 July 1942, Sreyermark
48413 (HOLOTYPE: F).

Plant perennial; stems to 40 cm tall with little branching except
sometimes at the base; branches densely glandular-pilose, almost
villous in places; leaves more or less similar throughout, lanceolate
to almost ovate, 10.0-33.0 mm long, 1.5-9.0 mm wide, acute,
densely pilose tending toward villous when young; upper few
internodes much longer than the leaves, gradually shortening to
much shorter below, creating, in some cases, a basal cluster; basal
rosette lacking; cymes 3-8 flowered, bracts not scarious-margined;
pedicels 10.2-36.0 mm long, the lower ones being the longer, very
densely glandular-pilose, somewhat hooked in fruit; sepals lanceo-
late to ovate, 5.8-7.6 mm long, 1.8-2.2 mm wide, acute, scarious
margined except at the apex, sparsely glandular-pilose; petals large,
9.5-14.5 mm long, bifid about 1/4 of their length, white; filaments
5.5-6.0 mm long, anthers 0.9 mm long; styles 3.7-4.0 mm long;
capsules 12.0-16.1 mm long, 2.0-2.4 mm wide, curved; seeds dark
red-brown, tuberculate, 1.0-1.2 mm in diameter; chromosome
number 2n = 34 (Beaman et al., 1962).

DISCUSSION: Cerastium juniperorum has perhaps the most limited
range of any species of Cerastium in Mexico or Central America. As
far as is known, it is restricted to a few square kilometers in the
Sierra de los Cuchumatanes, in west-central Guatemala, where it is
not uncommon in pine and juniper woodlands at elevations of
between 3050 and 3700 m (Figure IC.)

Standley and Steyermark (1944), in their original description of
the species, stated that it is perhaps only an extreme form of
Cerastium guatemalense but that the latter has a much smaller calyx
and that its cauline leaves are much narrower. Indeed, these are two
outstanding differences between the species, but they are only two of
many. Others can be seen when the respective descriptions are
compared.

SPECIMENS EXAMINED: Guatemala. HUEHUETENANGO: Sierra de los Cuchu-
matanes at km. 311 on Ruta Nacional 9N (between Paquix and Chémal), alt. ca. 3360

358 Rhodora [Vol. 86

m, Beaman 2973 (GH, MSC, TEX, UC, US); Sierra de los Cuchumatanes, between
Tojiah and Chémal at km. 320 on Ruta Nacional 9N, alt. ca. 3365 m, Beaman 3748
(ENCB, GH, MSC, TEX, UC, US); Sierra de los Cuchumatanes, ca. 3 km south of
road between Llano San Miguel and Todos Santos, from a point 2.5 km west of
Llano San Miguel, near highest point in Cuchumatanes, alt. ca. 3680 m, Beaman
3974 (MSC); Sierra de los Cuchumatanes, side of cliff 2 km north of km. 299 on road
to San Juan Ixcoy, alt. ca. 3350 m, Good 1024 (MSC); Sierra de los Cuchumatanes,
along road to Todos Santos, alt. ca. 3050 m, Good 1025 (MSC); along road in region
of Chémal, Sierra de los Cuchumatanes at km. 36, alt. ca. 3300 m, Standley 81687
(F); near Tunima, Sierra de los Cuchumatanes, alt. 3300-3500 m, Steyermark 48262
(F); alpine area in vicinity of Tunima, Sierra de los Cuchumatanes, alt. 3400-3500 m,
Steyermark 48413 (F, holotype); between Tojquia and Caxin bluff, summit of Sierra
de los Cuchumatanes, alt. ca. 3700 m, Steyvermark 50192 (F).

8. Cerastium madrense Watson, Proc. Amer. Acad. Arts 23:
269. 1888. Type: MExICcO. Chihuahua: On cool summits of
the Sierra Madre, 7 October 1887, Pringle 1504 (HOLOTYPE:
GH).

Plant perennial; stems 15-35 cm tall, much branched from the
base, not so above, erect, glandular-pilose, often more or less lanate
near the base; basal rosette prominent. Basal and sterile leaves
oblanceolate, obovate or spatulate, 10.0-60.0 mm long, 4.0-12.0
mm wide, acute, lanate, particularly basally and beneath; cauline
leaves much smaller, 8.0-20.0 mm long, 2.0-5.0 mm wide, lanceo-
late, acute, pilose, tending toward lanate particularly toward the
base of the plant, much shorter than the rather long internodes
(upper internodes 29.0-85.0 mm long); cymes 4—13 flowered, large,
open; bracts not scarious-margined; pedicels very long, the lower-
most (longest) 18.0-32.0 mm long, the upper ones 5.0-10.0 mm
long, glandular-pilose, apically hooked when in fruit; sepals lanceo-
late, 4.5-6.2 mm long, 1.2-1.8 mm wide, acute, scarious-margined
except at the apex, glandular-pilose; petals large, 7.8-11.1 mm long,
bifid about 1/5 of their length, white; filaments 6.0-6.2 mm long,
anthers 0.8 mm long; styles 4.0-4.2 mm long; capsules 7.6-11.9 mm
long, 2.1-3.0 mm wide, curved; seeds dark brown, 0.9-1.3 mm in
diameter, densely tuberculate; chromosome number unknown.

Discussion: As far as can be discerned from the specimens ex-
amined, Cerastium madrense is found in moist Pinus and Quercus
forests at elevations of between 2900 and 3200 m in the states of
Chihuahua and Durango (Figure 1B).

1984] Good — Cerastium 359

SPECIMENS EXAMINED: Mexico. CHIHUAHUA: cool summits of the Sierra
Madres, Pringle 1504 (GH, holotype). DURANGO: north slopes of Cerro Huehueto
(Huehuento), south of Huachicheles, ca. 75 mi west of C. Durango, alt. 2900-3150 m,
Maysilles 7241 (MICH); north slopes of Cerro Huehueto (Huehuento), ca. 75 mi
west of C. Durango, alt. 2900-3150 m, Maysilles 7250 (MICH); north slopes of Cerro
Huehueto (Huehuento), south of Huachicheles, ca. 75 mi west of C. Durango, alt.
2900-3150 m, Maysilles 7276 (DS, MICH, TEX); San Luis del Rio, 51 road miles
northwest of Coyotes, Maysilles s.n. (F, GH, NY, US).

9. Cerastium nutans Rafinesque, Prec. Decouv., p. 36. 1814.
TYPE: UNITED STATES. “en Pensylvanie” (not seen).

Cerastium nutans var. genuinum Rohrbach (lusus 1), Linnaea 37: 289. 1873.
TYPE: same as above.

Cerastium longepedunculatum Muhlenberg (nom. nud.), Cat., p. 47. 1813.
TYPE: UNITED STATES. “Pens.” (not seen).

Cerastium apricum Schlechtendal (with varieties angustifolium and brachy-
carpum), Linnaea 12: 208. 1838. Cerastium nutans var. apricum Rohr-
bach, Linnaea 37: 1873. Cerastium longepedunculatum var. apricum
Briquet, Ann. Conserv. Jard. Bot. Geneve 13 & 14: 381. 1911. Type:
MEXICO. state unknown: “Jalapam”, May, June (year not known), Schiede
S.n. (HOLOTYPE: HAL?), (not seen).

Cerastium ripartianum Schultz, Flora 45: 458. 1862. Type: MEXICO. state
unknown: “Hab. in Mexico, unde cl. Schaffner, absque nomine, misit”, (not
seen).

Cerastium cuspidatum Hemsley, Diag. Pl. Nov., p.21. 1878. Type: MEXICO:
state unknown: “in Convalli Mexici”, Schaffner 60 (HOLOTYPE: K) (not seen).

Cerastium sericeum Watson, Proc. Amer. Acad. Arts 20: 354. 1885. Type:
UNITED STATES. Arizona: “Huachuca Mountains, 8000 ft”, Lemmon &
Lemmon s.n., 1882, and “Santa Rita Mountains”, Pringle s.n., 1884 (not
seen).

Plant annual; stems 15-50 cm tall, often much branched;
branches erect or decumbent, pilose, usually more or less glandular-
pilose, sometimes more or less lanate below; internodes longest just
below the inflorescence, gradually shorter toward the base, usually
longer than, though sometimes shorter than, the leaves; leaves
largest near the base of the plant, gradually smaller upward,
lanceolate to ovate or spatulate, 7.0-60.0 mm long, 2.0-25.0 mm
wide, acute or obtuse, often somewhat thin, pilose or glandular-
pilose; basal rosette lacking; cymes 2-25 flowered, bracts not
scarious-margined; the lowermost pedicels the longest, 11.0-37.0
mm long, upper ones shorter, distally hooked when in fruit,
glandular-pilose; petals shorter than to much longer than the sepals

360 Rhodora [Vol. 86

(the petal/sepal ratio being 0.95-1.95), 3.6-10.3 mm long, bifid
about 1/4 of their length, white; filaments 2.5-4.0 mm long, anthers
0.3 mm long; styles 2.0-3.0 mm long; capsules 6.0-13.2 mm long,
1.5-3.5 mm wide, curved; seeds 0.7-1.0 mm in diameter, tubercu-
late; chromosome number 2n = 34 (Beaman et al., 1962) or 2n =
35-36 (Sdllner, 1952, 1954).

DISCUSSION: Cerastium nutans is the most widely distributed
species of Cerastium in Mexico. It has been collected throughout
the upland areas (1700-3660 m) except in the northeastern part of
the country. Collections have been made in the states of Chiapas,
Chihuahua, Distrito Federal, Durango, Guerrero, Hidalgo, Jalisco,
México, Michoacan, Morelos, Oaxaca, Puebla, San Luis Potosi,
Sonora, and Veracruz. To the south of Mexico, the species is known
only from Guatemala (Chimaltenango and San Marcos), and to the
north it is found as far as southwestern Quebec and British
Columbia (Fernald, 1950) (Figure 1D). The habitat includes such
areas as roadsides, woodlands, and rocky hillsides, almost always
below timberline.

Not only is Cerastium nutans the most widely distributed
Cerastium in Mexico, it is also the most variable. Some attempt
might be made at using this variation to define varietal or even
specific boundaries, but in most cases the material available for this
study showed too much intergradation between extremes. There
was also too little material on which to base sound taxonomic
judgement. | will therefore content myself here with describing the
variation without assigning any taxonomy to it.

The typical Cerastium nutans is a rather large, robust, much
branched plant with fairly thin, lanceolate to ovate leaves. It has
fairly long pedicels (usually well over 17 mm long) and moderately
sized sepals (4.0-5.0 mm long). The petals are usually equal to or
slightly longer than the sepals. This form is found from western
Guatemala north through the central highland of Mexico and into
the United States and Canada. From this typical form, specimens
vary in several ways, as described below:

a. Inthe Sierra Madre Occidental of western Mexico and in the
trans-Mexican volcanic belt (Clausen, 1959) there is a tendency for
the petals to increase considerably in size, reaching almost twice the
length of the sepals. This tendency is also seen in some parts of the
United States (Correll and Johnston, 1970).

1984] Good — Cerastium 361

b. In central and southern Mexico, particularly Michoacan,
specimens have been collected which have generally shorter pedicels
(13.0-19.0 mm long) and slightly smaller flowers than the typical.
These characteristics combine to give the plants to some extent the
aspect of Cerastium brachypodum with which these specimens have
often been confused. However, they are clearly not of that species
since, even in these characters, they much more closely approach C.
nutans.

c. Apparently restricted to the high mountains of Jalisco and
Guerrero is a variant which is usually fairly small and has rather
small, pale, rhomboidal leaves covered with a distinctively long,
glandular-pilose pubescence. Its pedicels are short (12.0-23.0 mm
long) and it has fairly large petals (5.2-7.1 mm long).

d. In northern Oaxaca a similar set of specimens has been
collected but these have elliptical leaves and lack the distinctive
pubescence described above.

e. Also in central Mexico can be found plants with very narrow,
attenuate leaves and small flowers.

f. A form found in western Chihuahua and eastern Sonora, is
characterized by very few, small flowers (sepals 2.9-4.0 mm long,
petals 3.9-4.0 mm long), short, broad capsules (6.0-8.6 mm long,
1.8-2.8 mm wide) and long, linear to lanceolate leaves (the length to
width ratio averaging 11.5, as opposed to 4.4 for all other specimens
of C. nutans).

g. The last variant bears varying amounts of lanate pubescence
on the lower parts of the stem and lower leaves. This group is
probably referable to the binomial C. sericeum, described from the
Huachuca and Santa Rita Mountains of Arizona (Watson, 1885).
Aside from the lanate pubescence, Watson, and later Robinson
(1897), separated it from C. nutans because its “seeds are twice
larger and more coarsely tuberculate.” Kearney and Peebles (1939),
however, stated that “intergradation in pubescence [between C.
sericeum and C. nutans] is complete in Arizona specimens” and
therefore desribed the specimens as a variety of C. nutans, C. nutans
var. obtectum. This situation is also seen in the Mexican specimens
which vary from extremely lanate basally (Knobloch 5915) through
less lanate (bana G. 416 and Knobloch 5750) to only slightly lanate
(Townsend & Barber 160, Phillips 672, and Nelson 6122). Kearney
and Peebles also stated that “the two forms are not constantly
distinguishable by the seed characters mentioned by Robinson

362 Rhodora [Vol. 86

(1897).” In Mexico, specimens showing lanate pubescence are found
in Chihuahua, eastern Sonora, and Durango.

REPRESENTATIVE SPECIMENS: Guatemala. CHIMALTENANGO: Sierra Santa
Elena, bei Tecpam Guatemala, Seler 2362 (GH). SAN MARCOS: Volcan Tacana,
east side of mountain at La Haciendita, alt. ca. 3375 m, Beaman 3188 (GH, MSC).
Mexico. CHIAPAS: slopes on southeastern side of Zontehuitz near summit,
Municipio de Chamula, alt. ca. 9400 ft, Breedlove 6703 (DS, F); near summit of
Chuchil Ton, northeast of Bochil, Municipio de San Andres Larranizar, alt. ca. 2700
m, Breedlove 26786 (DS, LL, MO). CHIHUAHUA: Mojarachic, Knobloch 5720
(MSC); Mojarachic, Knobloch 5915 (MSC); Majalca Cafion, LeSueur 466 (F, GH,
TEX); foothills of the Sierra Madres near Colonia Juarez, Nelson 6122 (GH, US);
Cerro Mohinora, 10 mi south of Guadeloupe y Calvo, alt. 2300-2400 m, Straw &
Forman 1960 (MICH); near Colonia Garcia in the Sierra Madres, alt. ca. 7300 ft,
Townsend & Barber 160 (F, GH, MICH, MO, MSC, NY, TEX, UC, US).
DISTRITO FEDERAL: Cafiada de Contreras, alrededores del 4° dinamo, alt. ca.
3000 m, Rzedowski 20411 (ENCB, MSC). DURANGO: ca. 5 mi north of railroad at
Coyotes (45 airline mi west of C. Durango), west facing slopes of broad arroyo,
tributary to Rio del Presidio, alt. ca. 2400-2500 m, Maysilles 7118 (NY).
GUERRERO: near Toro Muerto, Distrito Mina (Galeana), Hinton 11231 (GH,
MICH, NY, US). HIDALGO: Distrito Pachuca, Municipio Mineral del Chico,
below Parque Nacional El Chico, Moore 1554 (GH, UC). JALISCO: Nevado de
Colima (Nevado de Zapotlan), a few mi. south of Ciudad Guzman (Zapotlan), alt.
ca. 3080 m, Gregory & Eiten 287 (GH, MICH, MO, MSC, NY). MEXICO: Salto de
Agua, Purpus 1668 (F, GH, NY, UC, US); northwest slopes of Nevado de Toluca, 10
km (by road) southwest of junction of roads to Sultepec and Temascaltepec on Hwy.
130 to Temascaltepec or 27 km (by road) southwest of Toluca, alt. ca. 3000 m, Roe,
Roe & Mori 273 (ENCB, MICH, US, WIS). MICHOACAN: at the southwest side of
Cerro San Andres, ca. 12 km (straight line distance) north of Ciudad Hidalgo, alt. ca.
3100 m, Beaman 4318 (GH, MSC, NY, TEX, UC, US). MORELOS: along Rte. 95,
12 mi north of Cuernavaca toward Mexico City, Powell & Edmondson 732 (F,
MICH, TEX); Lagunas de Zempoala, alt. ca. 2775 m, Villamar s. n. (ENCB, MSC).
OAXACA: Llano de las Flores, on the Oaxaca-Valle Nacional highway 20 km east of
Ixtlan, alt. ca. 2870 m, Beaman 3711 (GH, MSC, US). PUEBLA: vicinity of San Luis
Tutitlanapa, Purpus 2720a (F, GH, MO, NY, UC, US). SAN LUIS POTOSI: region
of San Luis Potosi, alt. 6000-8000 ft, Parry & Palmer 47 (GH, NY, US). SONORA:
between Las Lierritas and El Tigre, region of the Rio de Bavispe, Phillips 672 (GH,
MICH). VERACRUZ: Mt. Orizaba, Sierra Negra, alt. ca. 11,800 ft, Balls & Gourlay
B4436 (MICH, UC).

10. Cerastium orithales Schlechtendal, Linnaea 12: 209. 1838.
TYPE: MEXICO, state unknown: “in regione subnivale montis
Orizaba”, September (year not known), Schiede s._n.
(HOLOTYPE: HAL?) (not seen).

Cerastium arvense var. orithales (Schlechtendal) Rohrbach, Linnaea 37:
305. 1873. Type: same as above.

1984] Good — Cerastium 363

Cerastium mutabile var. arvense f. angustatum Grenier, in part, Monogr.
Cerast., p. 68. 1841. Type: “Hab. in America boreali; in Siberia (DC.
herb); in Pyrenaeis (Grenier); in alpibus; (ex nonullis bot. hortis etiam
habui)” (not seen).

Plant perennial; stems 15-50 cm tall, branched at the base or not,
never branching above; branches very slender, usually held up by
surrounding grasses, more or less short glandular-pilose; internodes
much longer than the adjacent leaves near the inflorescence,
gradually becoming shorter basally until the leaves become densely
clustered; basal rosette lacking. Leaves more or less similar
throughout, sometimes smaller just below the inflorescence, linear
to lanceolate, 15.0-25.0 mm long, 2.0-4.0 mm wide, acute, densely
short glandular-pilose; cymes 1-6 flowered, bracts not scarious-
margined; pedicels various, 7.2-30.0 mm long, densely pubescent
with short glandular sub-reflexed hairs; sepals lanceolate to broadly
lanceolate or ovate, 6.0-7.5 mm long, 2.0-3.0 mm wide, acute,
scarious-margined except at the apex, glandular-pilose; petals large,
12.0-18.5 mm long, bifid about 1/4 of their length, white; filaments
6.1-7.3 mm long, anthers 0.8 mm long; styles 4.0-4.9 mm long;
capsules 13.5-17.0 mm long, 2.6-3.2 mm wide, curved; seed 0.9-1.2
mm in diameter, apparently only sparingly tuberculate, dark red-
brown; chromosome number 2n = 36 (Beaman et al., 1962).

DIscussIoN; Cerastium orithales is found at elevations between
3600 and 4100 m on Ixtaccihuatl, Pico de Orizaba, Sierra Negra,
and Cofre de Perote (all in central Mexico) (Figure 4A). The species
is restricted in habitat to grassy floors of open forest (almost
invariably Pinus harfwegii forests) just below timberline. Specimens
were collected by the author on Cofre de Perote in open meadows
(Good 1019), but these meadows were the result of the recent clear-
cutting of the native pine forest.

SPECIMENS EXAMINED: Mexico. MEXICO: Joya de Alcalican, extremo sur del
Ixtaccihuatl, alt. ca. 3900 m, Aldanda A. 47 (ENCB); Ixtaccihuatl, northwest side of
mountain above San Rafael, alt. ca. 3810 m, Beaman 2844 (GH, MSC, TEX, UC,
US). PUEBLA: Sierra Negra (adjacent to Pico de Orizaba), west side of mountain,
alt. ca. 3880 m, Beaman 2523 (F, GH, MSC, UC); Pico de Orizaba, north of
Alberque Piedra Grande, alt. ca. 3950 m, Beaman 3643 (GH, MSC, UC, US); Pico de
Orizaba, north side of mountain, ca. 3 km southeast of Villa Hidalgo, alt. ca. 3780 m,
Beaman 3649 (ENCB, GH, TEX, UC, US); along the road down the east side of the
Paso de Cortés, alt. ca. 3660 m, Good 1018 (MSC); north side of Pico de Orizaba,

364 Rhodora [Vol. 86

Figure 4. Distribution in central Mexico of Cerastium. A. C. orithales (closed
circles) and C. purpusii (open circles); B. C. ramigerum; C. C. tolucense; and D. C.
vulcanicum. There is also a single specimen of C. vulcanicum from Guatemala.

Murry 64 (MSC); Mt. Orizaba, alt. ca. 12,000 ft, Pringle 8551 (ENCB, F, GH,
MICH, MO, MSC, NY, UC, US). VERACRUZ: Cofre de Perote, east side of
mountain, alt. ca. 3930 m, Beaman 2159 (F, MSC, UC); ladera este del Cofre de
Perote, alt. ca. 4000 m, Dorantes L. 331 (CAS, GH); Cofre de Perote, Nelson 28
(US). STATE UNKNOWN: Mt. Orizaba, alt. 13,000-13,400 ft, Nelson 283 (US);
Ixtaccihuatl, alt. 11,000-12,000 ft, Purpus 268 (GH, MO, UC, US); Citlaltepetl,
Purpus 2803 (F, GH, MO, NY, UC, US). Mt. Orizaba, Rose & Hay 5756 (US); Mt.
Orizaba, alt. ca. 13,000 ft, Seaton 236 (F, GH, NY, US).

11. Cerastium purpusii Greenman, Zoe 5: 183. 1904. Type:
MEXICO, state unknown: Mt. Ixtaccihuatl, 1903, Purpus 472
(HOLOTYPE: GH; ISOTYPES: MO, UC, US).

Cerastium molle Bartling in Presl, Rel. Haenk. 2: 17. 1831. (non C. molle
Villars, Hist. Pl. Dauphine 3: 644. 1789). Type: MEXICO. state unknown:
Haenke s. n. (HOLOTYPE: PR; photograph of holotype in MICH).

Cerastium lanuginosum Sessé & Mocifio, Fl. Mex., ed. 2, p. 118. 1894. (non
C. lanuginosum Willdenow ex Reichenbach, Fl. Germ. Excurs., p. 797.
1832.) TYPE: MEXICO. Mexico: “habitat in Vulcano Tolucae” (not seen).

Plant a short-lived perennial, low, dense, caespitose; stems to 20
cm tall, usually less than 10 cm, much branched at the base, not

1984] Good — Cerastium 365

otherwise; branches pilose to lanate; internodes usually, though not
always, shorter than the leaves; cauline leaves broadly lanceolate to
ovate, 5.0-15.0 mm long, 2.0-6.0 mm wide, often largest just below
the inflorescence, usually acute, pilose or lanate; basal rosette often
present; basal leaves larger, to 35.0 mm long and 10.0 mm wide,
lanate, acute, lanceolate to spatulate; cymes 1-10 flowered, flowers
more or less densely crowded at the apex of each branch; bracts not
scarious-margined; pedicels less than 10 mm long, pilose; sepals
4.2-7.8 mm long, 1.3-1.8 mm wide, lanceolate, acute, pilose, never
glandular, margins narrowly scarious except at the apex; petals
6.0-10.6 mm long, bifid about 1/8 of their length, white; filaments
4.5-6.7 mm long, anthers 0.7 mm long; styles 3.0-3.3 mm long;
capsules 7.2-10.3 mm long, 2.5-3.0 mm wide, curved; seeds 0.7—0.9
mm in diameter, densely tuberculate; chromosome number 2n = 34
(Beaman et al., 1962).

DISCUSSION: Cerastium purpusii is one of the species of Cerastium
living at the highest elevations of any in Mexico, being found only in
the high, wet alpine meadows between 3650 and 4610 m on Nevado
de Toluca and Ixtaccihuatl, central Mexico (Figure 4A).

The binomial Cerastium molle Bartling in Presl, commonly used
for the species here described as C. tolucense, was originally used for
this species. However, since Villars used the name C. molle in 1789
(42 years before Bartling used it), Bartling’s name is not valid.

The name Cerastium lanuginosum was published in 1894 by Sessé
and Mocifio for a plant bearing the description of C. purpusii; this
binomial antedates C. purpusii by ten years. However, the fact that
C. lanuginosum Willdenow ex Reichenbach (probably synonomous
with C. alpinum Linnaeus) was published in 1832, 62 years before
the Sessé and Mocifio publication, prohibits its use for the Mexican
species.

REPRESENTATIVE SPECIMENS: Mexico. MEXICO: Nevado de Toluca, near large
lake in the crater, alt. ca. 4140 m, Beaman 1882 (F, GH, MICH, MSC, NY, TEX,
UC, US). PUEBLA: Ixtaccihuatl, alpine region, Purpus 472 (GH, holotype; MO,
UC, US, isotypes).

12. Cerastium ramigerum Bartling in Presl, Rel. Haenk. 2: 16.
1831. TYPE: MEXICO, state unknown: Haenke s. n. (HOLOTYPE:
PR; photograph of holotype in MICH).

Cerastium lithophilum Greenman, Zoe 5: 183. 1904. Type: MEXICO. state
unknown: Mt. Ixtaccihuatl, July 1903, Purpus 23] (HOLOTYPE: GH;
ISOTYPES: MO, UC, US).

366 Rhodora [Vol. 86

Plant perennial; stems 5S—25 cm tall, often much branched at the
base, sometimes above; sometimes more or less caespitose; branches
glandular-pilose; internodes very long just below the inflorescence,
very short basally, grading in between; leaves more or less similar
throughout, lanceolate to broadly lanceolate, often erect and close
to the stem, 10.0-30.0 mm long, 3.0-6.0 mm wide, acute, glandular-
pilose; basal rosette lacking; cymes 1-8 flowered, bracts not
scarious-margined; pedicels short, 2.2-10.6 mm long, densely
pubescent with short glandular subreflexed hairs; flowers nodding;
sepals lanceolate, 4.1-5.8 mm long, 1.6-2.2 mm wide, acute,
scarious-margined except at the apex; petals 6.4-10.3 mm long,
bifid about 1/3 of their length, white; filaments 3.6-4.0 mm long,
anthers 0.3 mm long; styles 1.2-1.6 mm long; capsules 7.0-10.8 mm
long, 2.0-2.4 mm wide, curved; seeds 0.5-0.7 mm in diameter, light
brown, tuberculate; chromosome number 2n = 36 (Beaman et al.,
1962).

DISCUSSION: This is a species of very high elevations (3660 to 4720
m) on Nevado de Toluca, Ixtaccihuatl, Popocatepetl, Tlaloc,
Malinche, Pico de Orizaba, and Cofre de Perote in the states of
Mexico, Puebla, Tlaxcala, and Veracruz (Figure 4B). It is restricted
to the high alpine meadows and barrens on those mountains and is
sometimes found approaching the upper limit of vascular plants.
One collection from Orizaba (Swan s. n.) bears the phrase “in
forests only” but this is atypical.

At present this species is referred to (in floras, etc.) as Cerastium
lithophilum, However, the binomial C. ramigerum has precedence.

REPRESENTATIVE SPECIMENS: Mexico. MEXICO: Nevado de Toluca, shore of large
lake in the crater, alt. ca. 4140 m, Beaman 1878 (F, GH, MICH, NY, TEX, UC, US);
Tlaloc, near summit of mountain, alt. 4100-4140 m, Beaman 2333 (GH, MSC, NY,
TEX, US); Ixtaccihuatl, south side of mountain, alt. ca. 4410 m, Beaman 2550
(MICH, MSC, NY); Municipio Amecameca, slopes of Popocatepetl between 10,000
and 12,000 ft, Gilly & Dodds 20 (MICH, MSC). PUEBLA: north side of
Popocatepetl, above timberline, Beaman 1727 (GH, MSC); Ixtaccihuatl, south side
of mountain, ca. 7 km north of Paso de Cortes, alt. ca. 4300 m, Beaman 2875 (MSC).
Pico de Orizaba, north side of mountain at Alberque Piedra Grande, alt. ca. 4275 m,
Beaman 3631 (GH, MSC); Ixtaccihuatl, south side of mountain on the south side of
Cerro Amacuilecatl, alt. ca. 4300 m, Beaman 4233 (MSC). TLAXCALA: Malinche,
crest of north rim of crater, alt. 4400-4450 m, Beaman 2241 (MICH, MSC, TEX,
UC, US). VERACRUZ: Pico de Orizaba, south side of mountain, north of Cueva del
Muerto, Beaman 1765 (GH, MSC, US); Cofre de Perote, east side of mountain, alt.
ca. 3930 m, Beaman 2158 (GH, MSC, NY, TEX, US).

1984] Good — Cerastium 367

13. Cerastium sinaloense D. A. Good, sp. nov. TyPE: MEXICO.
Sinaloa: Los Pucheros, Sierra Surotato, alt. 5500-6500 ft,
17-24 March 1945, Gentry 7224 (HOLOTYPE: GH; ISOTYPES: F,
US). (Figure 2).

Planta perennis. Caules usque ad 40 cm alti, basi ramosi. Rami
erecti, glanduloso-pilosi. Internodia basi brevissima, inflorescent-
iam versus longissimascentia. Rosula basalis absens sed folia infima
aggregatissima, magna, 30.0-65.0 mm longa, 7.2-18.2 mm lata,
spathulata, acuta. Folia superiora pauca (1-3 paria per ramum),
parviora, 4.0-28.7 mm longa, 1.2-7.6 mm lata, lanceolata usque
ovata, acuta. Omnia folia plus minusve pilosa. Cymae 2-8 floribus,
apertae. Pedicelli longi, glanduloso-pilosi, non uncati ubi fructifi-
cantes. Sepala lanceolata ad late lanceolate, 4.0-7.2 mm longa,
1.1-2.0 mm lata, praeter apicem scarioso-marginata, glanduloso-
pilosa. Petala maxima, 10.0-17.5 mm longa, bifida per circa 1/8
longitudinem, alba. Filamenta 6.4-7.0 mm longa. Antherae 1.1 mm
longae. Styli 4.7-5.5 mm longi. Capsulae 11.2-14.2 mm longae,
2.8-3.3 mm latae, rectae, dentibus recurvatis. Semina 0.5—-0.6 mm
diametro, tuberculata. Chromosomatum numerus non cognitus.

Cerastium sinaloense is characterized by having broad, spatulate
leaves crowded at the base of the plant (not in a rosette), very large
flowers, and straight, revolute-toothed capsules.

DISCUSSION: Cerastium sinaloense has been collected only in the
pine-oak forests of the Sierra Surotato in northern Sinaloa, near
Rosario in southern Sinaloa, and in the Sierra de Manantlan in
Jalisco (Figure 1C). It has been collected only at relatively low
elevations, between 1680 and 2250 m.

Three specimens, one from northern Sinaloa (Gentry 7234), one
from southern Sinaloa (Norris et al. 20466), and one from Jalisco
(McVaugh 23172), while obviously allied with Cerastium sinalo-
ense, C. sordidum and C. texanum by the presence of straight
capsules with revolute teeth, are atypical of C. sinaloense in that
they have far smaller flowers. They do, however, match C. sinalo-
ense more closely in general habit and in distribution than they do
either C. sordidum or C. texanum. Whether they merely represent
variation within C. sinaloense or are in fact specimens of a fourth
and as yet undescribed species is not discernible from the limited
material at hand.

368 Rhodora [Vol. 86

SPECIMENS EXAMINED: Mexico. JALISCO: Sierra de Manantlan (25-30 km
southeast of Autlan), along lumber roads 5 km east of the road crossing called “La
Cumbre” between El Chante and Cuzalapa, alt. 2000-2250 m, McVaugh 23172
(ENCB, MICH). SINALOA: east slope of Sierra Madre Occidental, 2.2 mi east of La
Palmita, ca. 47 mi east of Concordia on Mexico 40, Municipio de Rosario, alt. ca.
6450 ft, Breedlove 1710 (DS, MICH); Sierra Surotato, near settlement of Los Ornos
along the road to Surotato, 53 mi east of Mocorito, alt. ca. 5800 ft, Breedlove 15559
(MO, MSC); near settlement of El Triguito along the road from Los Ornos to
Surotato, alt. ca. 6200 ft, Breedlove 16474 (MSC); Sierra Surotato, below Buenas
Juntas, 5 mi northwest of Los Ornos along the road to Mocorito, Municipio de
Sinaloa y Vela, alt. ca. 5800 ft, Breedlove 19194 (MSC); Sierra Surotato, 5 mi
northeast of La Cienenga along the road to Santa Rita, Municipio de Badiraguato,
alt. ca. 7000 ft, Breedlove 19261 (MO, MSC); Los Pucheros, Sierra Surotato, alt.
5500-6500 ft, Gentry 7224 (GH, holotype; F, US, isotypes); Los Pucheros, Sierra
Surotato, alt. 5500-6500 ft, Gentry 7234 (GH); along Hwy. 40, ca. 6 mi west of Las
Palmitas, alt. ca. 7500 ft, Norris et al. 20466 (CAS, MO); 4-8 mi west of Palmito on
the Durango-Mazatlan Highway, Oatman & Rowlett s. n. (TEX).

14. Cerastium sordidum Robinson, Bot. Gaz. 30: 58. 1900. TyPE:
MEXICO, Chihuahua: Sierra Madre 8 km southeast of Colonia
Garcia, alt. ca. 2310 m, 30 May 1899, Townsend & Barber 40
(HOLOTYPE: GH; ISOTYPES: F, MO, MSC, TEX, UC, US).

Cerastium longepedunculatum var. sordidum Briquet, Ann. Conserv. Jard.
Bot. Genéve 13 & 14: 381. 1911. Type: same as above.

Plant perennial; stems to 40 cm tall, usually much less, erect or
decumbent, branches primarily at the base but sometimes above;
branches erect or decumbent, sparsely glandular-pilose; leaves
mostly crowded basally with very short internodes, these becoming
longer toward the inflorescence; lower leaves 18.0-35.0 mm long,
6.0-14.0 mm wide, ovate to spatulate, acute, glandular-pilose; upper
leaves few, much shorter than the adjacent internodes, 9.0-19.0 mm
long, 1.0-2.5 mm wide, linear to lanceolate, acute, glandular-pilose,
basal rosette lacking; cymes more or less open, but relatively small,
3-10 flowered; lower pedicels 10.0-20.0 mm long, upper ones
4.0-10.0 mm long, glandular-pilose, not hooked when in fruit,
sepals ovate, 3.0-5.7 mm long, 1.2-1.8 mm wide, acute, narrowly
scarious-margined; petals 5.5-7.8 mm long, bifid about 1/8 of their
length, white; filaments 4.6-6.3 mm long, anthers 1.0 mm long;
styles 3.0-3.3 mm long; capsules 8.0-13.4 mm long, 2.2-3.0 mm
wide, straight, teeth revolute; seeds red-brown, densely tuberculate,
0.6-0.8 mm in diameter; chromosome number unknown.

1984] Good — Cerastium 369

Discussion: The only specimen of Cerastium sordidum seen
during this study bearing any mention of habitat was Spaulding et
al. s. n. which listed “spruce stand”. The species has been collected
from Mexico only in Chihuahua (Figure IC). It is also known from
the pine forests of the Chiricahua and Santa Rita Mountains of
Arizona (Kearney and Peebles, 1969).

Briquet (1911) stated that Cerastium sordidum is not separable
from C. longepedunculatum (a synonym for C. nutans) and
therefore coined the name C. longepedunculatum var. sordidum. He
stated that “il existe en effet tous les intermediares a corolle reduite,
parfois meme nulle” (“in fact there exist all intermediate forms in
regard to reduced corolla, occasionally even none at all”). While I
have never seen a specimen with no corolla, the rest of this
statement is quite true, as far as it goes. Cerastium nutans, in fact,
varies considerably in corolla length (see above). If this were the
primary distinguishing character between the two species, | would
agree with Briquet’s diagnosis. However, since other characters such
as the straight, revolute-toothed capsule and the shape and
distribution of leaves are of greater importance, there is no reason to
consider C. sordidum and C. nutans conspecific.

Briquet (1911) mentioned three specimens from Oaxaca (Galeotti
4410, Galeotti 4428 and Jurgensen 15) as being ascribable to
Cerastium longepedunculatum var. sordidum. Although I have not
seen these specimens, no other indication of any plant resembling C.
sordidum has been found south of Chihuahua. It is therefore likely
that Briquet was mistaken about the identity of these three
specimens and that they were actually C. nutans.

SPECIMENS EXAMINED: Mexico. CHIHUAHUA: Mojarachic, Knobloch 5037 (F,
MSC); El Rialito spruce stand, 5 km south-southwest of San Juanito, alt. ca. 2400 m,
Spaulding, Martin & Wiseman s. n. (ENCB); in the Sierra Madre 8 km southeast of
Colonia Garcia, alt. ca. 2310 m, Townsend & Barber 40 (GH, holotype; F, MO,
MSC, TEX, UC, US, isotypes).

15. Cerastium texanum Britton, Bull. Torrey Bot. Club 15:
97. 1888. Type: “Hills, Blanco”, March, April (year not
known), Wright 69 (HOLOTYPE: NY?) (not seen).

Stellaria montana Rose, Contr. U. S. Natl. Herb. 1: 93. 1891. Type:
MEXICO. Sonora: Alamos Mountains, Palmer s. n. (HOLOTYPE: US) (not
seen).

370 Rhodora [Vol. 86

Plant perennial; stems very slender, erect or decumbent, much
branched basally, not so above, 15-35 cm tall, sparsely glandular-
pilose; internodes nowhere long (to 60 mm) but longest toward the
inflorescence, almost nonexistent at the base; basal rosette lacking
but lower leaves very crowded, large, 8.0-55.0 mm long, 3.0-16.0
mm wide, broadly spatulate, acute or obtuse, sometimes acuminate,
very sparsely pilose, often turning pale orange-brown with age,
upper leaves few or lacking, small, 4.0-8.0 mm long, 1.0-2.0 mm
wide, linear to lanceolate, pilose; cymes very open and loose, 8-25
flowered; pedicels 5.1-18.0 mm long, the lowest ones the longest,
very slender, glandular-pilose; sepals lanceolate to ovate, 3.0-5.1
mm long, !.0-2.0 mm wide, glandular, scarious-margined except at
the apex, turning light orange-brown when in fruit; petals 4.1—-5.4
mm long, narrow, bifid about 1/8 of their length, white; filaments
4.5 mm long, anthers 0.4 mm long; styles 1.2 mm long; capsules
small, only slightly exserted beyond the calyx, 4.2-6.8 mm long,
1.8-2.1 mm wide, straight, teeth revolute; seeds red-brown, 0.4—0.6
mm in diameter, densely tuberculate; chromosome number un-
known.

DISCUSSION. Cerastium texanum is found near watercourses in
canyons (Tidestrom and Kittel, 1941) or in open oak woods (Correll
and Johnston, 1970). The only specimens from Mexico seen in this
study with habitat data say “igneous rocky canyon slope in pine
forest” (Gentry 7991) and “talus slope” (Moran 20425). Cerastium
texanum is found at lower elevations than any other native
Cerastium species in Mexico or Central America, having been
collected at between 1275 and 1980 m in Baja California,
Chihuahua, and Sonora (Figure IC). North of Mexico this species is
found north to Coconino and Apache counties, Arizona (Kearney
and Peebles, 1969) and the Edwards Plateau of Texas (Correll and
Johnston, 1970).

SPECIMENS EXAMINED: Mexico. BAJA CALIFORNIA SUR: San Julio Cafion,
Brandegee s. n. (UC); lower north slope of Volcan las Tres Virgenes, alt. ca. 1275 m,
Moran 20425 (ENCB, LL, MO). CHIHUAHUA: Arroyo Hondo, Sierra Charuco,
alt. 4500-5500 ft, Gentry 7991 (US); Puerta de San Diego, alt. ca. 6500 ft, Hartman
593 (CAS, F, GH, NY, UC). SONORA: San Bernardo, Rio Mayo, Gentry 1253
(GH); Sierra de los Alamos, Palmer 293 (MICH, UC, US). arroyo in Sierra de
Alamos, in vicinity of Alamos, Rose et al. 12975 (NY, US).

1984] Good — Cerastium 371

16. Cerastium tolucense D. A. Good, sp. nov. TYPE: MEXICO.
Mexico: Nevado de Toluca, north side of mountain, 0.7 mi
east of point where road goes above timberline, alt. ca. 3985
m, 28 July 1958, Beaman 192] (HOLOTYPE: MSC; ISOTYPES: GH,
MICH, TEX, US). (Figure 2).

Planta perennis. Caules usque ad 35 cm alti sed plerumque minus
quam 20 cm, erecti vel ascendentes, basi profuse ramosi, non
superne. Rami pilosi usque lanati, praesertim basi et ad nodos
lanati. Internodia proxima infra inflorescentiam longissima, ple-
rumque (praeter aliquot specimina juvenia) multo longiora quam
folia. Folia caulina plerumque parva, 5.0-30.0 mm longa, 1.0-5.5
mm lata, linearia usque lanceolata, acuta, plus minusve lanata.
Rosula basalis praesens. Folia basalia majora, usque ad 60.0 mm
longa et 10.0 mm lata, lanceolata usque late elliptica, acuta vel
obtusa, plerumque praesertim basi dense lanata. Cymae 1-12
floribus. Bracteae non scarioso-marginatae. Pedicelli usque ad 25.0
mm longi, plerumque breviores, pilosi, interdum glanduloso-pilosi.
Sepala 3.4-6.5 mm longa, 1.0-2.0 mm lata, elliptica, acuta, praeter
apicem scarioso-marginata. Petala 4.5-11.6 mm longa, bifida per
circa 1/8 longitudinem, alba. Filamenta 4.0-7.0 mm longa. An-
therae 0.7 mm longae. Styli 3.0-4.0 mm longi. Capsulae 6.0-9.0 mm
longae, 2.1-2.8 mm latae, curvae. Semina 0.7-0.9 mm diametro,
tuberculata. Chromosomatum numerus 2n = 34 (Beaman et al.,
1962).

Cerastium tolucense is characterized by its non-caespitose habit,
the presence of a basal rosette, usually extensive lanate pubescence,
many cauline leaves, relatively large flowers, and curved capsules.

DISCUSSION: Cerastium tolucense is a species of fairly high
mountains (3000 to 4000 m) in central Mexico, having been
collected in the mountains of Distrito Federal, México, Michoacan,
and Puebla (Figure 4C). Its habitat includes both alpine and
subalpine meadows and, at slightly lower elevations, open Pinus
forests.

Although the name Cerastium molle Bartling in Presl (1831) has,
since its publication, been used exclusively for this species,
examination of a photograph of the type indicates that the name is
instead referable to what is here recognized as C. purpusii. Because

372 Rhodora [Vol. 86

of this confusion, no description has been published for this species;
I therefore describe it here as C. tolucense.

REPRESENTATIVE SPECIMENS: Mexico. DISTRITO FEDERAL: top of Cerro
Ajusco, alt. ca. 3937 m, Beaman 2773 (GH, MSC, US); Cerro Coyotes, cerca de
Contreras, Paray 300 (ENCB). Llano Grande, cerca del Desierto de los Leones, alt.
ca. 3250 m, Rzedowski 20496 (ENCB). MEXICO: Nevado de Toluca, north side of
mountain, 0.7 mi east of point where road goes above timberline, alt. ca. 3985 m,
Beaman 1921 (MSC, holotype; GH, MICH, TEX, US, isotypes); Telapon (north of
Ixtaccihuatl), south side of mountain, alt. 3450-3650 m, Beaman 2431 (GH, MSC,
TEX, UC, US); Ixtaccihuatl, south side of mountain between Altzomoni and La
Joya, 0.7 km south of La Joya, alt. ca. 3980 m, Beaman 3495 (GH, MSC); park area
at junction of road to Temascaltepec and road to Nevado de Toluca, alt. ca. 10,500 ft,
Dunn et al. 22518 (MO), vertiente oeste del Ixtaccihuatl, alt. ca. 3800 m, Espinosa 6
(CAS, ENCB, MSC); Cerro Tlaloc, Municipio de Tecaltitlan, alt. ca. 3000 m, Garcia
S. sn. (MSC); Lerma, alt. ca. 3000 m, Pina C. 79 (ENCB); Nevado de Toluca, alt.
13,000 ft., —-Palomas, Municipio de Iturbide (Santiago Tlazala), alt. ca. 3400 m,
Rzedowski 25912 (MSC). Rzedowski 28562 (ENCB). MICHOACAN: summit of
Cerro San Andres, ca. 12 km (straight line distance) north of Ciudad Hidalgo, alt. ca.
3589 m, Beaman 4278 (GH, MSC, UC, US); Municipio Tancitaro, alt. ca. 12,600 ft,
Leavenworth 278 (F, GH, MO, NY). PUEBLA: Ixtaccihuatl, south side of
mountain, ca. 6 km north of Paso de Cortés, alt. ca. 3900 m, Beaman 287] (MSC).

17. Cerastium triviale Link, Enum. Hort. Berol. 1: 433. 1821.
TYPE: not seen.

Cerastium vulgatum Linnaeus (nom. ambig.), Sp. Pl., ed. 2, p. 627. 1762.
Type: “Habitat in Scandiae et Europae australioris pratis, areis” (photo-
graph seen).

Cerastium caespitosum Gilibert (nom. ambig.), Fl. Lithuan. 2: 159. 1781.
TYPE: LITHUANIA: not seen.

Cerastium holosteoides Fries (nom. ambig.), Novit., ed. 2, p. 126. 1823.
TYPE: not seen.

Cerastium fontanum subsp. triviale (Link) Jalas, Arch. Soc. Zool.-Bot. Fenn.
‘Vanamo’ 18: 63. 1963. TYPE: not seen.

Plant weakly perennial; much branched, mostly basally; often
caespitose; braches 10-25 cm long, decumbent, more or less long-
pilose; internodes longest just below the inflorescence; leaves more
or less similar throughout, ovate to spatulate, 7.0-30.0 mm long,
3.0-10.0 mm wide, acute, sometimes obtuse, pilose; basal rosette
lacking; cymes usually quite dense, few to many flowered; bracts
with scarious margins; pedicels short, 2.2-8.4 mm long, long-pilose;
sepals lanceolate, 4.3-6.2 mm long, 1.0-1.6 mm wide, scarious-
margined, acute, pilose; petals slightly shorter to slightly longer than
the sepals, bifid about 1/3 of their length, white; filaments 3.7-4.0

ios)

1984] Good — Cerastium 37

mm long, anthers 0.3 mm long; styles 1.8-2.2 mm long; capsules
7.0-11.6 mm long, 2.1-3.0 mm wide, curved; seeds 0.5-0.7 mm in
diameter, red-brown, tuberculate; chromosome number 2n = 72
(Blackburn and Morton, 1957), 110 (Heitz, 1926), 126 (Hagerup,
1944, Blackburn and Morton, 1957), 136 (Brett, 1950), 137-147
(Brett, 1955), 140 (Taylor and Mulligan, 1968), 144 (Tischler, 1937;
Heiser and Whittaker, 1948; Blackburn and Morton, 1957; Fav-
arger, 1969; Live, 1972), 160 (Favarger and Kiipfer, 1968) or 180
(Blackburn and Morton, 1957). No counts have been published for
the Mexican or Central American populations.

Discussion: Although native to Europe, Cerastium triviale has
become established in North America throughout temperate and
subarctic Canada and the United States (Hitchcock et al., 1964) and
in central Mexico (Distrito Federal, México, and Veracruz),
southern Mexico (Chiapas), Guatemala (Alta Verapaz, and Baja
Verapaz), Honduras (Morazan), Costa Rica (Alajuela, Cartago,
Heredia, and San José), and Panama (Chiriqui). In habitat, it
ranges from roadsides and open meadows through brushy and
wooded areas to cloud forests at elevations of about 1400 to 3700 m
(Figure 3B).

As has already been discussed (see Cerastium glomeratum), the
oldest name for this species, C. vulgatum, is ambiguous and
therefore invalid. Two of the other four names listed above have
been used interchangeably for C. triviale and C. glomeratum and
have therefore also been considered ambiguous (C. caespitosum and
C. holosteoides). This series of eliminations leaves the names of C.
triviale and C. fontanum subsp. triviale. I here use the older and
shorter name, C. triviale, since in the absence of extensive
hybridization studies such a question of taxonomic rank is largely a
matter of personal preference.

REPRESENTATIVE SPECIMENS: Costa Rica. ALAJUELA: in and around Zaracero,
Canton Alfaro Ruiz, Hwy. 15, Weston et al. 2110 (UC). CARTAGO: cerca de la
cima del Volcan Irazu, Jimenez 140 (F); south slope of Volcan de Turrialba, near
Finca del Volcan de Turrialba, alt. 2000-2400 m, Standley 35276 (US). HEREDIA:
Cerro de Zurqui, northeast of San Isidro, alt. 2000-2400 m, Standley & Valerio
50601 (US). SAN JOSE: Cerro de Piedra Blanca, above Escasu, Standley 32481
(US). Guatemala. ALTA VERAPAZ: mountains east of Tactic, on road to Tamaht,
alt. 1500-1650 m, Standley 71180 (F). BAJA VERAPAZ: region of Patal, alt. ca.
1600 m, Standley 69597 (F, NY). Honduras. MORAZAN: Montana de la Tigra, al
sudoeste de San Juancito, alt. ca. 2000 m, Molina R. 14490 (F). Mexico. CHIAPAS:

374 Rhodora [Vol. 86

northeast slope of Zontehuitz near summit, Municipio de San Cristobal las Casas,
alt. ca. 9300 ft, Breedlove 14031 (DS, F, LL, MICH, US). DISTRITO FEDERAL:
Puerto de las Cruces, alt. ca. 3100 m, Rzedowski 34297 (ENCB). MEXICO: between
kms. 76 and 77 on the Amecameca-Popocatepetl road, alt. ca. 3250 m, Beaman 2062
(MSC). VERACRUZ: Las Vigas, Ne/son 14 (US). Balsequillo, Municipio de Perote,
alt. ca. 2350 m, Ventura R. 7386 (ENCB). Panama. CHIRIQUI: Volcan de Chiriqui,
alt. 3500-4000 m, Woodson & Schery 472 (GH, MO, US).

18. Cerastium vulcanicum Schlechtendal, Linnaea 12: 208. 1838.
TYPE: MEXICO. state not known: “in regione subnivale montis
Orizaba”, September (year not known), Schiede 508 (HOLO-
TYPE: HAL; MSC photograph no. 3868).

Cerastium micropetalum Greenman, Zoe 5: 183. 1904. TYPE: MEXICO. state
not known: Mt. Ixtaccihuatl, 1903, Purpus 473 (HOLOTYPE: GH; ISOTYPES:
MO, UC, US).

Plant annual or short-lived perennial; stems 6-30 cm tall, very
much branched, often tangled, sometimes more or less caespitose;
branches erect or decumbent, usually leafy, glandular-pilose above,
sparsely to densely lanate or villous toward the base; internodes
longer than to shorter than the leaves, mostly more or less similar
throughout except at the extreme base, where shorter; leaves more
or less similar throughout, linear to broadly lanceolate, 10.0—40.0
mm long, 2.0-7.0 mm wide, acute, pilose to lanate, the latter
particularly at the leaf bases and on the margins, leaves generally
more lanate toward the base of the plant, often lost in very old
plants; basal rosette usually lacking; cymes many flowered, plants
often mostly inflorescence; bracts not scarious-margined; pedicels
5.2-15.4 mm long, slender, densely glandular-pilose; sepals lanceo-
late to ovate, 3.5-4.8 mm long, 0.8-1.2 mm wide, acute, scarious-
margined, glandular-pilose; petals usually shorter than the sepals,
3.0-4.8 mm long, sometimes longer than the sepals (particularly on
Pico de Orizaba), to 5.2 mm long, bifid about 1/4 of their length,
white or pale green; filaments 2.6-3.4 mm long, anthers 0.2 mm
long; styles 1.3-1.6 mm long; capsules 5.0-9.0 mm long, 1.8 mm
wide, curved; seeds brown, 0.5-0.7 mm in diameter, lightly
tuberculate; chromosome number 2n = 34 (Beaman et al., 1962).

DISCUSSION: Cerastium vulcanicum 1s primarily a species of
alpine and subalpine meadows and disturbed sites in pine and fir
forests at elevations from 2900 to 4210 m in the Distrito Federal and

1984] Good — Cerastium 375

the states of México, Puebla, Tlaxcala, and Veracruz, Mexico
(Figure 4D). There is one collection of what appears to be C.
vulcanicum from northwestern Guatemala (Skutch 1219).

REPRESENTATIVE SPECIMENS: Guatemala. HUEHUETENANGO: Sierra Cuchu-
matanes, alt. ca. 10,800 ft, Skutch 1219 (F, GH). Mexico. DISTRITO FEDERAL:
Volcan Xitle, Matuda s. n. (CAS); Ajusco, Orcutt 3702 (F, GH, MO, US).
MEXICO: Nevado de Toluca, north side of mountain 2.0 mi east of point where road
goes above timberline, alt. ca. 4020 m, Beaman 1937 (F, GH, MICH, MSC, TEX,
UC, US); 3 km north of Paso de Cortés on road to Ixtaccihuatl, alt. ca. 3800 m,
Beaman 3610 (GH, MSC, TEX, UC, US); Llano Grande, Municipio de Zequiapan,
cerca de Rio Frio, alt. ca. 3200 m, Cruz C. 126] (ENCB, MICH, MSC). PUEBLA:
Pico de Orizaba, west side of Cerro Colorado, alt. ca. 3860 m, Beaman 2486 (GH,
MSC, TEX, UC, US); ca. 1.5 km east of the Paso de Cortés, alt. ca. 3580 m, Beaman
2897 (MSC). TLAXCALA: Ladera noreste de la Malinche, entre Apizaco y Hua-
mantla, alt. ca. 3750 m, Ern 95 (ENCB). VERACRUZ: west slope of Barranca de
Mala Cara, south-southeast of peak of Orizaba, alt. ca. 4210 m, Clausen s. n. (NY);
ladera este del Cofre de Perote, alt. ca. 3750 m, Dorantes L. 346 (CAS, GH).

EXCLUDED SPECIES

Two type specimens seen during this study, those of Cerastium
fasciculatum Bartling in Presl and C. stellarioides Mociiio ex
Seringe in de Candolle, were labelled as having been collected in
Mexico (the type of C. stellarioides is a drawing). However, in its
type description (de Candolle 1824), C. stellarioides is listed as being
found “in American bor. circa Nutka” so that either the label on the
specimen is in error or it is not the type of this species and is actually
an illustration of some Mexican species, probably a large flowered
C. nutans.

No other specimen even remotely resembling the type of
Cerastium fasciculatum was seen in this study; it is therefore
unlikely that the species occurs in Mexico. Although both the type
specimen and the original description (Pres! 1831) list “Mexico” as
the collection locality, Fenzl, on the type specimen, noted “vero-
similiter pl. chilensis non Mexicana!” (probably a Chilean plant, not
Mexican).

ACKNOWLEDGMENTS

For advice and aid in preparing this paper, I thank the following
people: John H. Beaman, Stephen N. Stephenson, and Richard K.
Rabeler, all of the Department of Botany and Plant Pathology,

376 Rhodora [Vol. 86

Michigan State University, and Donald L. Beaver and Richard W.
Hill of the Department of Zoology, Michigan State University. I
thank also the curators and staff of the following herbaria from
which specimens were borrowed: CAS, COLO, DS, ENCB, F, GH,
LL, MICH, MO, MSC, NY, TEX, UC, US, and WIS. (Abbrevi-
ations follow Holmgren and Kueken, 1974.)

Finally, I would like to dedicate this paper to the memory of Dr.
William T. Gillis of the Natural History and Botany Departments of
Michigan State University. He will be very much missed by all those
who knew and, in knowing, loved him.

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DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
MICHIGAN STATE UNIVERSITY
EAST LANSING, MI 48824

A NEW SPECIES OF Z/ZIPHUS (RHAMNACEAE)
FROM FLORIDA!

WALTER S. JUDD AND DAVID W. HALL

ABSTRACT

A new endemic, Ziziphus celata, is described from the xerophytic scrub and
pinelands of the Lake Wales Ridge in peninsular Florida. This distinctive geniculate-
thorny shrub shows clear affinity with the arid, southwestern North American
“Condaliopsis group” of Ziziphus, and represents a significant eastward range
extension for this group. The native vegetation of the Lake Wales Ridge has been
extensively destroyed, and the plant is represented only by a single specimen collected
by Ray Garrett in 1948 near Sebring, Highlands County, Florida. The species is very
likely now extinct.

Key Words: Endemism, extinction, flora of Florida, Rhamnaceae, Ziziphus

Ray Garrett, a local botanist, collected the type (and only)
specimen for the species described herein near Sebring, Florida in
March 1948. Correspondence at FLAS indicates that Garrett lived
in Sebring. He collected extensively in the surrounding scrub and
pinelands, and sent numerous specimens to the Herbarium of the
University of Florida. Garrett consulted with Erdman West and
Lilian Arnold at the U. F. Herbarium; neither could identify the
plant but both thought it to be in the Rhamnaceae. The specimen
was placed unmounted at the end of the family in an undetermined
folder. Over the past thirty-five years no other specimen of this plant
has been found, to the authors’ knowledge. During this time efforts
were made to identify this unusual plant; visiting botanists were
shown the specimen, but none was familiar with it. Efforts were also
made by several researchers to find the type locality and collect
other specimens. On a recent visit to the Field Museum of Natural
History (Chicago, Il.) the senior author noticed the similarity of the
undetermined specimen to Condalia Cav. and Ziziphus Mill. The
taxonomic diversity of these genera in the southwestern United
States and Mexico further suggested the possibility that they could
be reasonable generic determinations, since Florida has a number of
disjuncts of western taxa. Examination of material of many genera
and species of the family Rhamnaceae has convinced the authors

'This paper is Florida Agricultural Experiment Station Journal Series No. 5179.

381

382 Rhodora [Vol. 86

that the Sebring material is indeed referable to Ziziphus, and that it
was unnamed. An interesting and prophetic remark was made by
Ray Garrett, the collector of this specimen. Ina letter written during
the summer of 1948, he mused the J. K. Small had found many
novelties in the arid scrub habitat, and wondered why he could not
also find something rare. The specimen described below gives clear
evidence that he did.

The specific epithet “celata” (meaning hidden) reflects the
frustrating taxonomic history of the plant.

Ziziphus celata Judd and Hall, sp. nov. (Figure 1).

Frutex; rami primarii geniculati et glabri, ferentes brachyblastos
et spinas inaequales binatas ad nodos; spinae minores ca. 1-4 mm
longae, longiores (0.8) 1.3-3.7 cm longae cum (2)3 vel 4 brachy-
blastis secundariis et spinis tertiariis parvulis. Cortex grisea;
internodia 3-12 mm longa. Folia decidua, alterna (vel fasciculata ad
brachyblastos), glabra; lamina ca. 4.5-10 mm longa, 3-5 mm lata,
oblonga-elliptica vel leviter ovata, ad apicem rotundata vel leviter
emarginata cum mucrone minuto, ad basin cuneata vel attenuata;
margo integer; nervatura brochidodroma; petiolus 1.5-3 mm
longus; stipulae minutae. Flores axillares (fasciculati ad brachy-
blastos), vernales; pedicellus 2-3 mm longus. Sepala 5, ovata-
triangularia, 1.1-1.6 mm longa, glabra, viridia. Petala 5, spathulata,
1-1.3 mm longa, glabra, alba. Stamina 0.8-1 mm longa. Discus
laevis, incrassatus, ovarii basin cingens. Ovarium 2-3 loculare;
stylus contractus, bifurcatus vel trifurcatus. Fructus non visus.

Shrubs; primary branches geniculate and arcuate, to 20 cm long,
glabrous, without thorn-tips, bearing numerous short shoots from
the lowest nodes of which arise one or usually two distinctly unequal
thorn-tipped secondary branches [the smaller ca. 1-4 cm long and
the larger (0.8)1.3—3.7 cm long] the larger of which in turn bear from
(2)3 to 4 short shoots. These secondary short shoots each usually
produce a single small [0.3-4(-6.5)mm] tertiary thorn. Bark gray;
internodes 3-12 mm long. Leaves deciduous, alternate (fascicled on
short shoots); blade oblong-elliptic to slightly ovate, ca. 4.5-10 mm
long, 3-5 mm wide (possibly becoming larger with age), glabrous,
eglandular, coriaceous; apex rounded to slightly emarginate with a
small mucro; base cuneate to slightly attenuate; margin entire;
venation brochidodromous with midvein prominent, impressed
above and raised beneath, secondary veins inconspicuous; petiole

1984] Judd & Hall — Ziziphus 383

1.5-3 mm long; stipules narrowly-triangular, minute. Flowers
small, perfect, perigynous, solitary and axillary but seemingly
fascicled on the short shoots, vernal (appearing with leaves): pedicel
2-3 mm long. Cup ca. 1-1.3 mm in diameter, glabrous. Sepals 5,
ovate-triangular, 1.1-1.6 mm long, glabrous, green. Petals 5,
spatulate and clasping the stamens, I-1.3 mm long, glabrous, white.
Stamens 5, 0.8-1 mm long; anther opening by 2 longitudinal slits.
Disc smooth, thickened and closely surrounding but free from base
of ovary. Ovary 2-3 loculed, glabrous, tapering to style; style ca.
0.7-1 mm long, bifid or trifid to ca. one-half its length. Fruit not
seen, likely a drupe.

TYPE: UNITED STATES. Florida. Highlands Co., on sand dunes,
vicinity of Sebring. 18 March, 1949. Ray Garrett s.n. (FLAS).
Ziziphus celata is reminiscent of both Z. obtusifolia (Hook.) A.
Gray and Z. parryi Torr. It consistently differs from the former
highly variable species in that it (1) has its flowers solitary in the
axils of leaves of the short shoots, (2) lacks unicellular hairs on its
twigs, leaves, and flowers, and (3) has branchlets lacking a grayish
to whitish wax-like bloom. The taxon is separated from the latter by
its (1) usually paired unequal secondary thorns, (2) several-noded
secondary thorns (these bearing several short shoots and often small
tertiary thorns), and (3) floral cup and sepals greenish (not purplish-
green). These three species, along with Z. //oydii (Standley) M. C.
Johnston, Z. pedunculata (Brandg.) Standley and Z. mexicana
Rose, form a complex intermediate between the widespread tropical
genus Ziziphus and the xerophytically adapted southwestern U.S.—
Mexican—South American genus Condalia. This intermediate
complex has the floral characters of Ziziphus, i.e., disc thickened
around ovary, ovary base broadly attached to receptacle, style
tapered and forked, petals present, along with some (to all) of the
xeromorphic features of Condalia. The latter group includes scrubs
of arid habitats with small, + obovate, entire-margined, pinnate-
veined leaves fascicled on short shoots borne on thorn-tipped
branches with very short internodes. In addition, stipular spines are
lacking (see Johnston, 1962). These intermediate species have been
treated as Condalia subgenus Condaliopsis by Weberbauer (1895)
or as a distinct genus Condaliopsis (Weberb.) Suessenguth (1953).
However, Johnston (1962) noted that species of “Condaliopsis share
characters of ovary, disc and style which set them off from Condalia

[Vol. 86

Rhodora

384

1984] Judd & Hall — Ziziphus 385

and at the same time show their similarity to Ziziphus.” He added
that although these species approach “Condalia in several charac-
teristics which can be put under the general heading of ‘xero-
morphy’”, they are “not separable from the large diverse genus
Ziziphus.” Following Johnston (1962, 1963) we consider the species
described above as a Ziziphus.

Previously known species of the “Condaliopsis group” are limited
to arid habitats of the southwestern United States (California to
Texas and Oklahomay and Mexico (Johnston, 1963). Ziziphus
celata, collected only in the xerophytic sand scrub of Highland Co.,
Florida, thus represents a significant eastward range extension.
However, as mentioned above, other Florida taxa show similar
patterns, either occuring as disjuncts in mesic to xerophytic habitats
of Florida and the southwestern U.S./Mexico or having closely
related species to the west. Some examples include: Callirhoe
papaver (Cav.) A. Gray, Ceanothus microphyllus Michx., Erio-
gonum longifolium Nutt. var. gnaphalifolium Gand. (E. floridanum
Small), Forestiera pubescens Nutt., Krameria lanceolata Torr.,
Lyonia ferruginea (Walt.) Nutt., Lyonia fruticosa (Michx.) G. S.
Torrey, and Rudbeckia nitida Nutt. var. nitida. The ancestors of
Ziziphus celata may have reached Florida from the west during past
periods of increased aridity.

The species has also been compared with various small-leaved
Caribbean species of Ziziphus (see Johnston, 1964). Ziziphus celata
is clearly distinct from these taxa (compare leaf margin, venation,
inflorescence structure), and probably is not closely related. In
addition, the Caribbean species are typically plants of limestone or
serpentine habitats whereas Z. celata is found on acid white sand.

Ziziphus celata is likely a member of the characteristic and highly
endemic flora occurring on the sterile white sands of the Central
Florida Ridge. The most common plant communities of this region
are known as the Sand Pine - Xerophytic Oak Scrub and the High
Pinelands/Sandhills. The former is dominated by Pinus clausa
(Chapm. ex Engelm.) Vasey ex Sarg., Quercus geminata Small, Q.

Figure 1. Ziziphus celata: a, habit (drawn from specimen), X 1; bed, leaves
(drawn from material re-expanded by boiling), X 2; e, flower (drawn from material
re-expanded by boiling), X 15; f, flower (drawn from dried material), X 15; g, petal
clasping stamen, X 15; h, stamen, X 15.

386 Rhodora [Vol. 86

myrtifolia Willd., and Q. chapmanii Sarg., while the latter is
dominated by Pinus palustris Mill. However, the area is now mainly
occupied by citrus groves and urban developments. This area
supports numerous other endemics: Asclepias curtissii A. Gray,
Bonamia grandiflora (A. Gray) Heller, Bumelia lacuum Small,
Carya floridana Sarg., Chapmannia floridana Torr. & Gray,
Chionanthus pygmaeus Small, Clitoria fragrans Small, Conradina
canescens (Torr. & Gray) A. Gray, Dicerandra frutescens Shinners,
Eryngium cuneifolium Small, Garberia fruticosa (Nutt.) A. Gray,
Hypericum cumulicola (Small) P. Adams, H. edisonianum (Small)
Adams & Robson, //ex opaca Ait. var. arenicola (Ashe) Ashe,
Lechea cernua Small, Liatris ohlingerae (Blake) Robins., Lupinus
aridorum McFarlin ex Beckner, L. cumulicola Small, Nolina
brittoniana Nash, Osmanthus megacarpus Small, Paronychia char-
tacea Fern., Persea humilis Nash, Polygala lewtonii Small, Poly-
gonella ciliata Meisn. var. basiramia (Small) Horton, P. myriophylla
(Small) Horton, Prunus geniculata Harper, Quercus inopina Ashe,
Sabal etonia Swingle ex Nash, Stylisma abdita Myint, Warea
amplexifolia (Nutt.) Small, and W. carteri Small (see also James,
1961; Harper, 1949; Ward, 1979). Even the striking geniculate-
thorny habit of Ziziphus celata is present in several other char-
acteristic scrub taxa, such as Prunus geniculata, Bumelia lacuum,
Ximenia americana L. and a distinctive local variant of Crataegus
flava Ait.

The authors suggest that systematic studies of the endemic species
of the Central Florida Ridge are urgently needed. The natural plant
communities of the region are rapidly being destroyed by agri-
cultural and urban development, with the only significant remnants
of native flora existing in the Ocala National Forests in the north
and Archbold Biological Station in the south. Very likely Ziziphus
celata is now extinct; most of the natural vegetation near Sebring
has been destroyed, and the species has never been collected since,
despite repeated searches by D. Burch, D. Ward and the authors. It
is feared that many additional taxa of this area (e.g., Warea
amplexifolia, Lupinus aridorum, Stylisma abdita, Nolina_brit-
toniana, N. atopocarpa Bartl., Eryngium cunifolium, and Hyperi-
cum cumulicola) may soon become extinct thus rendering future
studies of their ecology and systematics impossible or very difficult
to accomplish.

1984] Judd & Hall — Ziziphus 387

ACKNOWLEDGMENTS

We thank Dr. Marshall C. Johnston for his several helpful
suggestions and comments concerning the manuscript. Thanks are
also extended to the curator of the Field Museum (F) for loan of
material, and to Drs. Derek G. Burch, Daniel B. Ward, and Erdman
West who have puzzled over the plant in the past 30 years.

LITERATURE CITED

Harper, R. M. 1949. A preliminary list of the endemic flowering plants of
Florida. Jour. Fla. Acad. Sci. 2: 39-57.

James, C. W. 1961. Endemism in Florida. Brittonia 13: 225-244.

JoHNsTON, M. C. 1962. Revision of Condalia including Microrhamnus (Rham-
naceae). Brittonia 14: 332-368.

1963. The species of Ziziphus indigenous to United States and Mexico.

Amer. J. Bot. 50: 1020-1027.

1964. The fourteen species of Ziziphus including Sarcomphalus (Rham-
naceae) indigenous to the West Indies. Amer. J. Bot. 51: 1113-1118.

SUESSENGUTH, K. 1953. Rhamnaceae. Jn: Die Naturlichen Pflanzenfamilien. Ed.
2., 20d: 7-173.

Warp, D. B. (ed.) 1979. Vol 5, Plants. Jn: P. C. H. Pritchard (ed.), Rare and
endangered biota of Florida. 175 pp. Univ. Presses of Florida, Gainesville.
WEBERBAUER, A. 1895. Rhamnaceae. /n: Engler, A. & Prantl, Die Naturlichen

Pflanzenfamilien. 3(5): 393-427.

W.S. J.

DEPARTMENT OF BOTANY
UNIVERSITY OF FLORIDA
GAINESVILLE, FL 32611

D. W. H.

DEPARTMENT OF NATURAL SCIENCES
FLORIDA STATE MUSEUM
UNIVERSITY OF FLORIDA
GAINESVILLE, FL 32611

THE EFFECT OF POWER UTILITY RIGHT-OF-WAY
CONSTRUCTION ON CAT-TAIL
(TYPHA LATIFOLIA L.) MARSH

F. R. THIBODEAU AND N. H. NICKERSON

ABSTRACT

Vegetation in an eastern Massachusetts freshwater cat-tail marsh was measured
over a five-year period, two years before and three years after construction of a 345
kv transmission line; no change in vegetation occurred because of the construction.

Key Words: Inland wetland, power-line construction, vegetative cover

Darnell (1976) suggested that power utility rights-of-way will
have a significant long-term effect on the vegetation of a wetland if
the natural cover is disturbed. Thibodeau and Nickerson (in
preparation) and Nickerson and Thibodeau (1984) questioned this
assumption based on field studies of wooded swamp and bog which
were subject to both long-term management and new right-of-way
construction. Both areas did show evidence of perturbation, but
recovery was rapid. We now report that marsh dominated by the
cat-tail Typha latifolia L. showed no evidence of disturbance even
during the first growing season after power line construction.

During the winter of 1977-1978 a 345 kv transmission line was
built across an area of marsh covering approximatley 1.5 square
miles along the Saugus River in Wakefield, MA. Construction
equipment was driven directly across the frozen marsh and no other
alterations to the substrate, such as filling, were made. Because the
vegetation was uniformly herbaceous, the construction company
did not purposefully remove any plants. Any alteration was at-
tributable to incidental effects of the construction equipment itself.

From 1976 to 1980, stem counts of all vegetation were taken
during June and July at eight 1 m? stations along a transect under
the lines and at eight other | m? stations a parallel distance of 50 m
from them. The stations were spaced along the transects using a
random digit multiplied by 3 m as the distance from one station to
the next. 1976 and 1977 growing season measurements were taken
before construction; 1978, 1979, and 1980 measurements were taken
after construction. These stem counts were converted to measures of
diversity and evenness (Shannon and Weaver, 1949) and richness

389

390 Rhodora [Vol. 86

Table 1. Plant cover comparisons between right-of-way and undisturbed marsh
for years immediately before and after construction.
Year #Species #Stems Diversity Evenness Richness
1977 9.56+.83 55.30+7.44 1.21+.01 0.63+.01 1.30+.05
1978 10.63+.72 61.02+8.39 1.20+.01 0.61+.01 1.32+.04

Data shown are grand means for each year + the deviation between group means
under and away from the lines. None of these deviations is significant.

(Margalef, 1957). In addition, analyses were made of the total
number of species and of the stem count itself. None of these
measures distinguished the two sets of plots in any year-pair using
analysis of variance to discriminate between them (SPSS, 1979; p >
-05 in all cases). The 1977 and 1978 growing seasons, which should
mirror the greatest changes, showed no significant differences
(Table 1). The vegetative cover ranged from 10% to 80% of each m?.
Typha made up 50% to 95% of the total cover per m?. Table 2 lists
the major species comprising the plant cover. Names and authorities
are those of Fernald (1950).

It appears that in eastern Massachusetts, at least, frozen substrate
protects Typha rhizomes and other living plant tissues sufficiently
from the compaction and mixing which might be expected to occur
when construction equipment is used in such a fresh-water cat-tail
marsh, Power transmission line construction carried out in this
fashion, with no change in the water regime of the wetland, had no
demonstrable effect on the plant association of the marsh.

Table 2. Species comprising plant cover!

Typha latifolia L.
Sagittaria latifolia Willd.
Impatiens capensis Meerb.

Cicuta bulbifera L.

Galium palustre L.

Lemna sp.

Dryopteris thelypteris (L.) Gray

Lythrum salicaria L.
Cephalanthus occidentalis L.
Rosa palustris Marsh.

'The first three species comprised 80% or more of the cover on any | m? quadrat; the
second four, while often present, never comprised, even together, more than 5% of
the cover on any | m? quadrat; the last three were only sporadically encountered.

1984] Thibodeau & Nickerson — Power lines/ Typha marsh 391]

ACKNOWLEDGMENTS

We thank the many students who were our conscientious
employees over the summers of field work. This investigation was
supported by a research grant from the New England Power
Company to the second author.

LITERATURE CITED

DARNELL, R. M. 1976. Impacts of Construction Activities in Wetlands of the
United States. U. S. Environmental Protection Agency. EPA-600/3-76-045.

FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th Edition. American Book
Co., Boston. 1632 pp.

MARGALEF, R. 1977. Information Theory in Ecology. General Systems Bulletin
31:36-71.

NICKERSON, N. H. AND F. R. THIBODEAU 1984. The Impact of Power Utility
Rights-of-Way on Wooded Wetlands. Environmental Management: (In press).

SHANNON, C. E. AND W. WEAVER 1949. The Mathematical Theory of Communi-
cation. U. Illinois Press, Urbana, 117 pp.

SPSS (Statistical Package for the Social Sciences) 1979. Documentation by C. H.
Hull and N. H. Nie, McGraw-Hill, N.Y., N.Y.

E.R: T.

CENTER FOR PLANT CONSERVATION
ARNOLD ARBORETUM

JAMAICA PLAIN, MA 02130

N. H.N.

DEPARTMENT OF BIOLOGY
TUFTS UNIVERSITY
MEDFORD, MA 02155

REDISCOVERY OF STYLISMA HUMISTRATA
(CONVOLVULACEAE) IN TENNESSEE

VERNON BATES AND PAUL LEwIS

Recent field work has uncovered a native species, Stylisma
humistrata (Walter) Chapman, that has remained undetected in
Tennessee for more than a century. Previously, its presence in
Tennessee was known from only one collection (‘Lookout Mt.,
Tenn.’, Hamilton County) made by George Vasey in 1878. Our col-
lection is from west Tennessee, approximately 200 miles west of
Vasey’s original collection site. There has been no attempt as yet to
relocate this species in the vicinity of Lookout Mountain.

Vasey’s collections of Stylisma humistrata were deposited in
herbaria outside Tennessee and, until the revision of Stylisma
(Myint, 1966), had remained unknown to Tennessee botanists. We
examined one specimen that was originally deposited in the
Princeton University Herbarium and later transferred to the New
York Botanical Garden in 1945. According to Myint, a duplicate of
the same collection was seen by him at the Philadelphia Academy of
Science. However, a search of Stylisma material at the Philadelphia
Academy of Science did not reveal its presence. It is interesting to
note that even though Augustin Gattinger spent about 15 years
botanizing in southeastern Tennessee (Oakes, 1932), he apparently
did not collect this species nor did he hear of Vasey’s collections
(Gattinger, 1901). In the most recent state checklist (Sharp et al.,
1960), this species continued to be unrecognized as a member of the
flora of Tennessee.

Of the eight taxa currently recognized in Stylisma (Myint, 1966),
S. humistrata has the widest distribution. Its range extends from
coastal Virginia to northern Florida and west to eastern Texas. It
appears that the southern border of Tennessee (35° N lat.) approxi-
mates its northernmost limits in the Mississippi River Valley. In
fact, our collection site is less than one mile from the Mississippi
border. Interestingly, Smith (1978) also gives the northernmost
distribution for this species in Arkansas at approximately the same
latitude.

The recent collection of Stylisma humistrata reported here is from
west Tennessee, an area we are currently studying. The collection
data are: Tennessee, McNairy Co., deep sand areas along the

393

394 Rhodora [Vol. 86

Tuscumbia River, at dirt road crossing about 1.0 mi E of its
confluence with the Hatchie River, V. Bates & P. Lewis 5189, 13
August 1983. Voucher specimens have been deposited at APSC,
GH, MEM, TENN, and VDB.

ACKNOWLEDGMENTS

We would like to thank Robert Kral and Eugene Wofford for
checking their Tennessee records, and Alfred Schuyler for searching
for the Vasey specimen at Philadelphia Academy of Science.

LITERATURE CITED

GaTTINGER, A. 1901. The flora of Tennessee and a philosophy of botany. Gospel
Advocate Publishing Co., Nashville.

MyInT, T. 1966. Revision of the genus Sry/isma (Convolvulaceae). Brittonia 18:
97-117.

Oakes, H. N. 1932. A brief sketch of the life and works of Doctor Augustin
Gattinger. Cullom and Ghertner Co., Nashville.

SHARP, A. J., R. E. SHanks, H. L. SHERMAN, & D. H. Norris. 1960. A
preliminary checklist of dicots in Tennessee. University of Tennessee, Knoxville.

SmitTH, E. B. 1978. An atlas and annotated list of the vascular plants of Arkansas.
University of Arkansas, Fayetteville.

V. B.

HARVARD UNIVERSITY HERBARIA
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PL:

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Applicants should submit a proposal of no more than three
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02138. The recipient of the award will be notified by 30 April 1985.

Vol. 86, No. 846, including pages 121-238, was issued June 28, 1984.

395

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RHODORA July 1984 Vol. 86, No. 847
CONTENTS

Taxonomy of Arnica (Compositae) subgenus Austromontana

Steven J. Wolf and Keith E. Denford Re, ee cone a a Fe
A synopsis of the genus Halenia (Gentianaceae) in Mexico
Robert L. Wilbur. : : Bey eat es... Sane
A revision of the Mexican and Central American species of Cerastium
(Caryophyllaceae)
David A. Good Be ee al aig Ge

A new species of Ziziphus (Rhamnaceae) from Florida
Walter S. Judd and David W. Hall ; : ; eae ; . 381

The effect of power utility right-of-way construction on cat-tail (Typha latifolia

L.) marsh
F. R. Thibodeau and N. H. Nickerson ; F F : : ‘ . 389

Vernon Deies and Paul pn : A ; : ; : , ‘ . 393
NEBC Research Award Notice . . . . . . . . .. . 395

Instructions to contributors to Rhodora ; . , ‘ . inside back cover

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

October 1984 No. 848

Vol. 86

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Cover illustration

Ledum groenlandicum Oeder, Labrador tea, reaches its southeastern distribu-
tional limit in Concord, Mass. It was first collected by Thoreau in 1858, subsequently
regarded as extirpated by Richard Eaton in 1974, and rediscovered by Ray Angelo in
1978. Angelo has since found it in two more Concord locations.

Original artwork by Josephine Ewing.

Tbhodora

(ISSN 0035-4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 86 October 1984 No. 848

THE HERB STRATA OF
THREE CONNECTICUT RIVER
OXBOW SWAMP FORESTS

MARJORIE M. HOLLAND! AND C. JOHN BURK

ABSTRACT

The herb floras of swamp forests in three of four large Connecticut River oxbows
in western Massachusetts have been described and compared. The herb stratum at
Hatfield includes a lush growth of annual and perennial herbs, lianas, and emergent
marsh species, but no seedlings of the dominant canopy tree, Populus deltoides.
Composition of the herb stratum in Ned’s Ditch, Northampton, forest was found to
vary markedly during four years of sampling. Seedlings of major canopy species at
Ned’s Ditch increased steadily through successive years while abundance of floating
hydrophytes fluctuated strongly. Ferns and spring-blooming herbs predominated in
the herb stratum at Whately which also contained seedlings of all canopy species. Of
the herbaceous species sampled, 70% occurred only at one site. An argument is made
that floodplain forest. severely limited in New England, should be more stringently
protected.

Key Words: Swamp forest, floodplain vegetation, herbaceous flora, wetlands, Con-
necticut River

INTRODUCTION

Published accounts of floodplain and swamp forest vegetation in
New England are chiefly limited to the perceptive comments of
Nichols (1915, 1916) on lowlands and stream banks in Connecticut.
Floodplain forests elsewhere have been investigated more inten-
sively. However, much of this work has concentrated on overstory

'Present address: Department of Biology, College of New Rochelle, New Rochelle,
NY 10801.

397

398 Rhodora [Vol. 86

species with only a few studies examining herbaceous strata in
detail (Wistendahl, 1958; Lindsey et al., 1961; Bell, 1974; Keam-
merer et al., 1975; Barnes, 1978; Hardin and Wistendahl, 1983).

Within an 18 km stretch of the Connecticut River in western
Massachusetts, four large oxbow lakes occur. Three are in the
active floodplain while the fourth now lies on a higher terrace. We
have recently (Holland and Burk, 1982) discussed the relative ages
(time since cut-off) of these oxbows. The “youngest” was separated
from the main stream in 1840 and remains in large part open water.
The others contain varying amounts of open water, marsh, shrub
swamp, and forest.

The object of the present study is to describe and compare the
floras of the herbaceous strata in the forests of the three older
western Massachusetts oxbows. More detailed accounts of the
geology, history, and marsh vegetation of these oxbows are found
in Robinton and Burk (1971); Burk (1973, 1977), Sackett (1974,
unpublished Master’s thesis, Smith College, Northampton, Mass.;
1977, unpublished Ph.D. dissertation, U. Mass, Amherst), Burk
and Lauermann (1977) and Holland and Burk (1982).

LOCATION AND AGE OF THE STUDY SITES

Figure | indicates the location of the study sites. In order of
increasing age they are (A) the Hatfield oxbow, which includes two
adjacent segments known as Great Pond and Cow Bridge Brook,
(B) Ned’s Ditch, the northeastern half of a prehistoric oxbow in
Northampton, and (C) the Whately oxbow.

The Hatfield oxbow was formed sometime prior to the late
1700’s. Depicted as two large ponds on the earliest maps of the area,
it was subject to full erosive flow from the Connecticut River during
heavy flooding as recently as 1936 (Collins and Schalk, 1937).

Stratigraphy and radiocarbon dating indicate that the prehistoric
Northampton oxbow was cut off from the main stem around 710
(+130) yr. B. P. (Holland and Burk, 1982). The Ned’s Ditch section
is largely forested. Hulbert’s Pond, the lower southeastern portion
of this oxbow, contains an extensive marsh complex kept open by
the Mill River which flows through it into the 1840 oxbow.

The Hatfield and the prehistoric Northampton oxbows tend to be
flooded yearly or more often (Brower, 1971). The Whately oxbow is
for the most part above the present floodplain. The lower eastern

1984] Holland & Burk — Oxbow herb strata 399

The Connecticut
River Oxbows

Key:

1 Whately Pond
2 Great Pond Co
3 cow er ——
4 Ned's Ditch or. 4 ws
5 Hulbert's Pond

6 1840 Oxbow

Vy ?
Connecticut River

==S5 Former Oxbow

Hadley

awe

a ae Sit Ae
L<l sy , Extracted from USGS quadrants:
f& Mt. Toby 1971, Easthampton 1964,
Williamsburg 1964, Mt. Holyoke 1964.

Contours: 50 feet

ye J

Figure 1. Map of the study area and location of the study sites (from Holland
and Burk, 1982. Reprinted with permission from Northeastern Geology, Vol. 4.
No.1.)

400 Rhodora [Vol. 86

portions of this oldest site were flooded in 1936, but the western
portions have not been flooded at all within historic times (Jahns,
1947).

METHODS

Since each of the oxbows contains a diversity of habitats, sam-
pling techniques were planned to assure that only swamp forest was
included. The course of the old river bank can usually be identified.
After a general reconnaisance at each site, belt transects approxi-
mately 300 m long and | m wide were laid out, crossing the former
channel at regular intervals.

At Hatfield, 10 belt transects were established at intervals of 575
m. On four of these, swamp forest occurred adjacent to one side of
the old river bank in sufficient quantity to lay out a 10 m X 10 m
quadrat beginning 3.3 m in from the edge of the forest closest to the
bank. Within each 10 m X 10 m quadrat, ten | m X | m quadrats
were laid out, five at regular intervals along the transect and five at
regular intervals along a baseline perpendicular to the transect.

Ned’s Ditch encompasses approximately half of the total North-
ampton oxbow. Five transects were laid out at 285 m intervals in
Ned’s Ditch. Swamp forest occurred adjacent to both sides of the
old river bank on all of these. Ten 10 m X 10 m quadrats were
established in Ned’s Ditch, two on each transect beginning 3.3 m in
from the edge of forest at each end of the transect. Within each 10 m
x 10 m quadrat, | m X | m quadrats were laid out as at Hatfield.

Five transects were laid out at 575 m intervals at the Whately site.
Sufficient swamp forest was present at Whately to establish five 10
m X 10 m quadrats, one adjacent to the river bank on one transect
and two adjacent to the river bank on either end of two other
transects. Ten | m X | m quadrats were laid out within each larger
quadrat, as in the other sites.

Presence and coverage as determined by visual estimate were
noted for all herbs, vines and woody seedlings under 60 cm height in
the | m X | m quadrats. During 1975, 40 | m X | m quadrats were
sampled at Hatfield, 100 at Ned’s Ditch, and 50 at Whately. Differ-
ent numbers of quadrats sampled at each site reflect the different
amounts of swamp forest present. The proportion of each oxbow
supporting swamp forest was estimated from aerial photographs,
USGS topographic maps, and other maps available from local

1984] Holland & Burk — Oxbow herb strata 401

sources. A comparison of ratios of total area sampled to overall
swamp forest present indicated that roughly equal proportions of
vegetation were sampled at each oxbow.

Within the study area, few floodplain forest. herbs break dor-
mancy before early May, and many begin seasonal development in
late May or early June. Therefore sampling was conducted during
the last two weeks of July and the first three weeks of August, the
period when the herbaceous stratum is most fully developed.
Because preliminary observations indicated that the herb stratum of
Ned’s Ditch varied markedly from year to year, Ned’s Ditch was
sampled in 1973, 1974, 1975, and 1977. At Hatfield and Whately,
belt transects and 10 m X 10 m quadrats were situated during 1974,
but sampling at these sites that year was limited to five quadrats
along the transect in each 10 m X 10 m quadrat. Full sampling at all
sites was conducted in 1975. General collecting was conducted each
season to determine species present in the swamp forest at any site
but not represented in the quadrats.

For purposes of analysis, a table was prepared listing all species
sampled in any quadrat during 1973, 1974, and 1975. In a few
instances, specimens could be identified only at the generic level
and, in the case of some immature grasses, only to family. These
identifications are included in the list but omitted in later floristic
comparisons. Species occurring at a study site but not within any
quadrat in 1975 are indicated (a) if they appeared in any quadrat at
the site during any prior year of sampling or (b) if they were col-
lected elsewhere in swamp forest at the site during 1975. Cover and
frequency data are included only for 1975. Species collected at a site
but never found within a quadrat are not included.

To document changes in the composition of the flora of Ned’s
Ditch during the four years of sampling, a table was prepared listing
cover values and frequencies for all species encountered each year.
This list and maps indicating the locations of the belt transects at
each site are available from the authors upon request. Precipitation
data used in the discussion of vegetational changes were obtained
from Philip T. Ives (personal communication through the study
period.)

Unless otherwise indicated in Table 1, nomenclature follows Fern-
ald (1950). Voucher specimens of all species collected have been
deposited in SCH.

402 Rhodora [Vol. 86

Table |. Composition of the herb strata. Percent cover (C) and frequency (F) of
species sampled, total number of species present, and total vegetative cover of her-
baceous strata of three western Massachusetts oxbow forests in 1975. P indicates
sampling prior to 1975 but not that year, X indicates presence in swamp forest
elsewhere at the site but not in quadrats. Total cover is the sum of actual cover values
for each species.

Hatfield Ned's Ditch Whately

Species %C %F %F WF FC WF

Herbs and vines

Agrostis alba L. 4 12.5 2 17

Amphicarpa bracteata (L.) Fern. <5 25° -<5 2 -

Apios americana Medic. - I 6

Aralia nudicaulis L. : -- - -- l 14

Arisaema triphyllum (L.) Schott <5 5 <5 I <5 2

Aster divaricatus L. <.5 2

A. vimineus Lam. - P P

Aster spp. <5 5

Athyrium filix-femina (L.) Roth l 2.5 x x 13 30

Bidens tripartita L. 2 47.5 <5 8 -

B. frondosa L. - l 4

Boehmeria cylindrica (L.) Sw.! 7 85 <— 9

Botryvchium multifidum (Gmel.) Rupr. <5 l

Callitriche heterophylla Pursh <5 7.5 --

Cardamine pensylvanica Muhl. P P

Carex tribuloides Wahlenb. <5 10

Carex spp. 4 22.5 <5 I <5 6

Cicuta maculata L. <5 5

Circaea quadrisulcata (Maxim.) l 7.5
Franch. & Sav.

Clematis virginiana L. <<) 2.5

Clintonia borealis (Ait.) Raf. <§ 2

Commelina communis L. P P

Convolvulus sepium L. <5 2.5

Coptis groenlandica (Oeder) Fern. - - P P

Cuscuta gronovil Willd. <.5 15 x xX

Cyperus strigosus L. P P

Dennstaedtia punctilobula 11 34
(Michx.) Moore

Drvopteris spinulosa <.5 5) P P
(O. F. Muell.) Watt

D. thelypteris (L.) Gray | 5 Xx xX P P

Dulichium arundinaceum (L.) Britt. 7 16

Echinocystis lobata <5 2.5

(Michx.) T. + G.

'May include some immature Pilea pumila

1984] Holland & Burk — Oxbow herb strata 403
Table |. Continued

Hatfield Ned's Ditch Whately
Species %C KF FC FF FC HF
Eleocharis acicularis (L.) R. & S. -- 2 14 -- -
E. obtusa (Willd.) Schultes P P = - a
Equisetum fluviatile L. 2 22:5 - -- -
Eupatorium rugosum Houtt. <5 7.5 -- -
Galium aparine L. | 37.5 <5 8 -- -
Gaultheria procumbens L. ps 40
Geum canadense Jacq. - P P
G. virginianum L. | 15 - -
Glechoma hederacea L. <5 10 -
Gramineae (unidentified) 7 17.5 <5 2 --
Hypericum virginicum L. - <5 l
Impatiens capensis Meerb. 13 80
Leersia oryvzoides (L.) Sw. 4 35 P P -
L. virginica Willd. <2 6
Lemna minor L. 34 88
Ludwigia palustris (L.) Ell. <5 30 <.5 16
L. polyvcarpa Short & Peter <3 l
Lycopodium complanatum L. 4 28
L. obscurum L. 6 30
Lycopus uniflorus Michx. <.5 |
L. virginicus L. P P xX xX
Lysimachia ciliata L. | Ue Bee) xX xX
L. nummultlaria L. 7 10 <.5 3
L. terrestris (L.) BSP. P P
Maianthemum canadense Desf. <.§ 2.5 xX xX 3 24
Matteuccia struthiopteris (L.) Tod. 5 10
Medeola virginiana L. | 14
Mitchella repens L. Xx X 3 24
Monotropa uniflora L. P P
Myosotis scorpioides L. <.5 10
Onoclea sensibilis L. | 7.5 l 3
Osmunda cinnamomea L. 18 46
O. claytoniana L. 2 6
O. regalis L. P 3 12.5 <.§ 2
Oxalis europaea Jord. <5 12:5
Parthenocissus quinquefolia | 12.5 x x 2 18

(L.) Planch.
Peltandra virginica 2 22.5
(L.) Schott & Endl.

Penthorum sedoides L. <.$ |
Pilea pumila (L.) Gray Xx X xX xX
Polygonatum pubescens <2 2

(Willd.) Pursh

404 Rhodora [Vol. 86

Table |. Continued
Hatfield Ned’s Ditch = Whately

Species %C %F FC YF FC EF
Polygonum sagittatum L. <5 7.5 -- _-

P. scandens L. <9. -:2;5 P P --
Pontederia cordata L. P P -- -- --
Potamogeton pectinatus L. <5 |

Ranunculus abortivus L. P P ' =

R. flabellaris Raf. 2 22 - ~~
Rhus radicans L. <5 2.5 Xx Xx -
Sagittaria latifolia Willd. P P -- : -- --
Scrophularia lanceolata Pursh 3 7.5 - -- --
Scutellaria lateriflora L. l 20. <.$ l - --
Sium suave Walt. <.§ 6 --
Smilacina racemosa (L.) Desf. ~ P P
Smilax herbacea L. <.§ 5 -- --
Solanum dulcamara L. <.5 | -- --
Solidago spp. <5 10 --

Spirodela polyrhiza (L.) Schleid. 3 78 -- --
Streptopus roseus Michx. -- -- l 16
Syvmplocarpus foetidus (L.) Nutt. 3 10 -- ~-
Thalictrum polygamum Muhl. P P X xX --
Trientalis borealis Raf. - l 36
Trillium undulatum Willd. -- - l 10
Utricularia vulgaris L. - 2 13 -

Viola conspersa Reichenb. | 17.5 - P P
Vitis spp. <5 15 <5 5 <.5 2
Woody seedlings

Acer negundo L. | 22.5 -- - --
A. rubrum L. P P -- I 42
A. saccharinum L. l 37.5 4 35 - -
Alnus rugosa (DuRoi) Spreng. l 5 P P -- -
Amelanchier laevis Wieg. -- - <5 2
Betula lutea Michx. f. -- <.5 6
Betula spp. <5 l -- --
Cephalanthus occidentalis L. <5 2.5 4 31 -- --
Cornus alternifolia L. f. - -- -- <.5 16
C. amomum Mill. 3 25 : -- --
C. stolonifera Michx. 2 12 7 --
Fagus grandifolia Ehrh. -- -- - <5 2
Fraxinus pennsylvanica Marsh. -- -- l 4 ~ ~

*Not all specimens of Vitis could be identified in quadrats. V. riparia Michx. was
present at all sites, V. /abrusca L. at Hatfield and Whately, and V. aestivalis Michx.
at Ned’s Ditch.

1984] Holland & Burk — Oxbow herb strata 405

Table |. Continued
Hatfield Ned's Ditch Whately

Species %C OF AF FF AWC AMF

Hamamelis virginiana L. - - <.5 16

Ilex laevigata (Pursh) Gray =<) 2.5 - -

I. verticillata (L.) Gray -- l 11 <5 2

Kalmia latifolia L. - 4 14

Nyssa sylvatica Marsh. - FP P

Populus deltoides Marsh. - <.5 | -

Prunus serotina Ehrh. l 5 2 44

Quercus bicolor Willd. - <5 I

Q. palustris Muenchh. <5 3

Q. velutina Lam. : | 12

Rhododendron viscosum (L.) Torr. l 8

Robinia pseudo-acacia L. P P

Rubus hispidus L. P P

Rubus spp. | 14

Salix spp. <5 2

Sambucus canadensis L. <5. 2:5

Spiraea latifolia (Ait.) Borkh. <5 2.5 <5 |

Ulmus rubra Muhl. - <5 l

Vaccinium atrococcum | 6
(Gray) Heller

Viburnum acerifolium L. <.§ 4

V. cassinoides L. - | 12

V. recognitum Fern. <5 | 8 36

Total in 1975 55 47 40

% Cover 79% 71% 89%

RESULTS AND DISCUSSION

The Hatfield Oxbow

The forest at the Hatfield oxbow shares aspects of floodplain
forests described early this century on the lower Connecticut River
(Nichols, 1916). These include “rank and luxuriant” undergrowth, a
“wealth of lianas”, few shrubs, and a trend toward replacement of
Populus deltoides by shade-tolerant trees.

Anestimated 16 ha of swamp forest occupies approximately 10%
of the oxbow. Populus deltoides dominates an open canopy in
association with Acer saccharinum and unusually large specimens
of Alnus rugosa. Acer negundo, Fraxinus pennsylvanica, Ulmus
rubra, and Rhus typhina occur as small trees. Shrubs are poorly
developed with A/nus rugosa and Cornus amomum most common,

406 Rhodora [Vol. 86

By late summer, the herb stratum supports lush vegetation.
Although fewer quadrats were sampled at Hatfield, more herb-
aceous species were encountered there than at other sites (Table 1).
Perennial herbs contribute most to overall cover. However, annuals
are also common, with /mpatiens capensis ranked first in individual
coverage. Boehmeria cylindrica, Lysimachia nummularia, and Mat-
teuccia struthiopteris are important perennial herbs. Vines, includ-
ing Amphicarpa bracteata, Clematis virginiana, Convolvulus
sepium, Cuscuta gronovii, Echinocystis lobata, Parthenocissus
quinquefolia, Polygonum scandens, Rhus radicans, Vitis labrusca
and V. riparia form tangled masses on other growth. Typical emer-
gent marsh species of both the Hatfield and Northampton oxbows,
including Equisetum fluviatile, Leersia oryzoides, Ludwigia palus-
tris, Peltandra virginica, and Scutellaria lateriflora, possess high
frequency but low cover.

Few species occur as woody seedlings. Populus deltoides is not
regenerating, but seedlings of Acer saccharinum, A. negundo, and
Cornus amomum are well distributed.

Ned’s Ditch

Swamp-forest is more extensive and better developed in Ned’s
Ditch, covering 30 ha, an estimated 66% of the upper half of the
Northampton oxbow. The forest was intermittently logged prior to
1973 but not since. Acer saccharinum, Quercus palustris, and Frax-
inus pennsylvanica are the most important components of the can-
opy, with Ulmus rubr., Betula lenta, B. papyrifera, Acer rubrum,
and large specimens of Cephalanthus occidentalis. Populus del-
toides, Salix nigra, S. fragilis, and S. rigida occur as small trees on
one site where coarse sand fill was deposited during highway con-
struction. Cephalanthus occidentalis, Cornus stolonifera, and Ilex
verticillata are important shrubs.

Over the first three years of sampling at Ned’s Ditch, species
richness (total number of species present) and coverage in the herb
stratum increased strikingly. The relative contribution of individual
species to cover also varied (Table 2). In 1973, coverage was only
14% with 27 species encountered in 100 quadrats. The depauperate
vegetation and sparse flora resulted in large part from an atypical
early summer flood wich followed a very wet spring (Table 3). The
herb stratum was completely inundated for at least 10 days in early
July (Sackett, 1974 unpublished Master’s thesis, Smith College,

1984] Holland & Burk — Oxbow herb strata 407

Table 2. Yearly variation in Ned’s Ditch. Relative cover (percentage of total
vegetative cover) represented by species occurring with a total of 5% or more during
any sampling period.

Year
Species 1973 1974 1975 1977
Acer saccharinum L. 7 11 6 12
Cephalanthus occidentalis L. l 8 6 10
Dulichium arundinaceum (L.) Britt. 29 17 10 <5
Eleocharis acicularis (L.) R. & S. l 3 3 8
Lemna minor L. 7 3 48 7
Onoclea sensibilis L. 14 11 l 8
Osmunda regalis L. 14 14 4 7
Ranunculus flabellaris Raf. l 3 3 11

Table 3. Precipitation measurements. Monthly April-August, total April
August, annual 1973-1977, and mean data 1948-1983. Data were recorded in inches
at Amherst, MA.

Month Total

Year April May June July August April-August Year
1973 6.40 5.45 4.48 3.33 2.24 21.90 46.65
1974 3.81 4.01 3.46 3.65 3.97 18.90 44.71
1975 2.87 2.10 4.68 10.56 6.13 26.34 59.97
1976 3.40 4.49 2.97 1.58 6.06 18.50 39.63
1977 4.9] 357 3.83 4.04 5.94 22.29 55.79
Mean Data from 1948-1983

4.00 3.57 3.88 3.55 3.86 18.86 44.06

Northampton, Mass.). By August, Dulichium arundinaceum, Ono-
clea sensibilis, and Osmunda regalis together accounted for 8%
coverage. Lemna minor and Spirodela polyrhiza contributed little
cover but occurred at high frequencies.

Relatively dry conditions during the frost-free period of 1974
allowed successful establishment or re-establishment of numerous
species. Cover increased to 36% with 36 species present. Dulichium
arundinaceum, Onoclea sensibilis, and Osmunda regalis were again
the most prominent herbs, contributing, with Dryopteris spinulosa,
19% coverage. Lemna minor and Spirodela polyrhiza continued
with low cover but high frequency.

408 Rhodora [Vol. 86

The !975 season was unusually wet and, following heavy rainfall
in July (Table 3), shallow standing water was present in most of the
quadrats through the sampling period. Cover reached 71% with 47
species encountered. One invader, Ludwigia polycarpa, is included
in the Massachusetts list of rare and endangered species (Crow et
al., 1981). Lemna minor, intermixed with Spirodela polyrhiza, was
most important, contributing 34% cover. Other frequent “errant
hydrophytes” (Mueller-Dombois and Ellenberg, 1974) were Ranun-
culus flabellaris and Utricularia vulgaris. Dulichium arundinaceum
increased while ferns as a group declined.

Some woody seedlings present before the August, 1973 sampling
may have been destroyed by the July flood. Woody seedlings
increased from 5 to 13 species from 1973 to 1975, with coverage by
woody seedlings increasing from less than 2% to over 12%. All
canopy trees were represented in the herb stratum in 1975,

During 1977, another wet season with precipitation more evenly
distributed than in 1975, coverage was 73%, slightly higher than in
1975 with 41 species present. Woody species contributed more than
23% cover (Table 2). Ranunculus flabellaris and Eleocharis acicula-
ris increased and Dulichium arundinaceum declined to very low
levels. Lemna minor decreased but was still very frequent, as was
Spirodela polyrhiza. Seedlings of Fraxinus pennsylvanica occurred
in 51% of the quadrats.

During the four years of sampling at Ned’s Ditch, 61 taxa were
identified at the specific level within quadrats. Of these, 16 occurred
every year. In addition to the species listed in Table 2, these
included Agrostis alba, Apios americana, Dryopteris spinulosa,
Ilex verticillata, Pilea fontana, Quercus palustris, Spirodela poly-
rhiza, and Vitis riparia. Their life forms include six herbaceous per-
ennials, four woody seedlings, two “errant hydrophytes”, one
annual, and two vines. In addition, immature specimens of Bidens
spp., probably mostly B. cernua and B. tripartita were sampled
every year. Ferns as a group contributed about 5% of the total
coverage during 1973 and 1975 and more than twice that amount in
1974 and 1977. The canopy species, well represented by seedlings,
are apparently replacing themselves, and, as long as flooding con-
tinues, the forest might be expected to continue ina persistent state
of “hydric disclimax” (Daubenmire, 1968) or “pulse stability”
(Odum, 1969).

1984] Holland & Burk — Oxbow herb strata 409

The Whately Oxbow

The lower eastern portions of the Whately oxbow support a
poorly drained complex of forest, marsh, and open water. The
upper western portions are almost entirely forested; much of this
woodland occurs on slopes leading up to a higher river terrace.
Only about 25% of the Whately oxbow, an estimated 18 ha, sup-
ports swamp forest suitable for sampling.

The forest has been occasionally logged but apparently never
clearcut. Acer rubrum and Prunus serotina are the most important
trees in association with Quercus alba and Nyssa sylvatica, Ame-
lanchier laevis, Castanea dentata, and Tsuga canadensis are present
as small trees; some specimens of Hamamelis virginiana are also
large enough to transgress the canopy. Viburnum dentatum and
saplings of all overstory species form a dense shrub stratum.

The herb stratum is very well developed with a total coverage of
89%. Perennials and woody seedlings are the most important life
forms. Vines are scarce while annuals and “errant hydrophytes” are
lacking. Ferns are more abundant at Whately than at any other site,
with Osmunda cinnamomea, Athyrium filix-femina and Denn-
staedtia punctilobula predominant. These, with Lycopodium com-
planatum and L. obscurum, contribute 52% cover. Spring-blooming
perennial herbs, generally absent at other sites, are abundant. These
include Aralia nudicaulis, Clintonia borealis, Coptis groenlandica,
Medeola virginiana, Polygonatum pubescens, Smilacina racemosa,
Streptopus roseus, Trientalis borealis, and Trillium) undulatum.
Epigaea repens is also present as is Monotropa uniflora. Maianth-
emum canadense occurs at all three sites and Mitchella repens at
Ned’s Ditch and Whately.

Woody seedlings account for more than 20% of the cover with all
canopy species represented. Acer rubrum and Prunus serotina
occur with highest frequencies; although the prevalence of these
species may reflect disturbance, there is no reason to believe the
general composition of the overstory will change under present
conditions.

FLORISTIC TRENDS AND COMPARISONS

Overstory strata of floodplain forests are generally characterized
by few wide-ranging species with similar life histories (White, 1979).
Herb strata of these forests differ markedly, however. In our study,

410 Rhodora [Vol. 86

excluding taxa identified only above the species level, 94 species of
herbs and vines were sampled. Of these, 70% were found at a single
site, with seven species occurring in all three oxbow forests. Most of
the 27 species shared by two sites occurred jointly in Hatfield and
Ned’s Ditch. Similarly, of the 32 species represented by woody
seedlings in the oxbow forests, 75% were limited to one site and
none occurred at all three sites. Hatfield and Ned’s Ditch shared
half the woody species occurring at two\sites; one of these, Acer
saccharinum, was a potential overstory tree. More species of woody
seedlings were found at Whately than elsewhere and 75% of these
occurred only there.

Comparable data from other river systems are few; some of the
most interesting come from Wistendahl’s (1958) study of the flood-
plain of the Raritan River in New Jersey. On one stretch of the
Raritan, three forest types can be distinguished running more or
less parallel to one another along one side of the channel (Buell and
Wistendahl, 1955; Wistendahl, 1958). These are (1) outer floodplain
forest characterized by a diverse overstory including Fagus grandi-
folia, Acer saccharum, Liriodendron tulipifera, Ulmus rubra, and
Tilia americana, (2) inner floodplain forest with Acer rubrum,
Ulmus americana, and other wetland trees, and (3) a forest on a
high terrace characterized by Acer saccharum in association with
tree species more frequent on drier sites.

Wistendahl’s methods were generally similar to ours except that
he sampled twice during the summer. Excluding taxa identified
only to genus level, 56 species of herbs and vines were recorded in
the three Raritan forests. Of these, 50% occurred at a single site,
while II species occurred in all three forests. As in the western
Massachusetts oxbows, the greatest number of species shared by
two sites occurred in the more frequently flooded sites, and the
higher (terrace) site had more species represented as woody seed-
lings. Herbs and vines were poorly represented in the Raritan
terrace, and most of those sampled (83%) were also present in one
or both of the other sites. Of the 14 species represented by woody
seedlings in the Raritan forests, 57% were limited to one site and
29% occurred at all sites. Possible explanation for the greater sim-
ilarity of the herb strata within the three Raritan forests is that the
sites lie immediately adjacent to one another and that the geological
processes which formed them are more gradual than the cut-off of

1984] Holland & Burk — Oxbow herb strata 411

an oxbow, which often takes place in the course of a single major
flood event.

A comparison of the various floras of the Massachusetts and
New Jersey forests shows that of the 56 species of herbs and vines
sampled on the Raritan, 16 were recorded in at least one western
Massachusetts site: 12 at Hatfield, 8 at Ned’s Ditch, and 4 at
Whately. In both cover and shared species, the inner floodplain
forest on the Raritan most closely resembles the Hatfield site, while
the outer floodplain and terrace forests show little resemblance to
any western Massachusetts site.

The herbaceous floras of the western Massachusetts oxbows bear
less floristic resemblance to those of bottomland forests along the
Missouri River in North Dakota (Keammerer et al., 1975). The
North Dakota forests occur near the western limit of “northern
floodplain forest” (Vankat, 1979) and have not been flooded since
1954 because of dam and reservoir construction. Of the 216 species,
both woody and herbaceous, found in the North Dakota forests,
only 20 are represented in the forests of the western Massachusetts
oxbows. No species restricted to cottonwood (Populus deltoides)
forest in North Dakota occurs in the western Massachusetts sites.
Twenty species found in North Dakota bottomland forests gener-
ally, or restricted to more mesic sites, also occur in one or more
western Massachusetts oxbow forests: 15 at Hatfield, 12 at Ned’s
Ditch, and | at Whately.

Only four herbaceous species occur jointly in the Raritan, North
Dakota, and western Massachusetts sites: Amphicarpa bracteata,
Circaea quadrisulcata, Ranunculus abortivus, and Scutellaria late-
riflora. All four of these occur at Hatfield, two occur in Northamp-
ton, and none at Whately.

Ferns are important at all the western Massachusetts sites and
several fern species are also included by Nichols (1915, 1916) as
characteristic of floodplains and swamp forests in Connecticut.
Ferns are scantily represented at sites described elsewhere within
the northern floodplain forest from New Jersey (Wistendahl, 1958)
through Ohio (Hardin and Wistendahl, 1983), Indiana (Lindsey et
al., 1961), Illinois (Bell, 1974), and Wisconsin (Barnes, 1978) west
into North Dakota (Keammerer et al., 1975).

An increasing abundance of non-native species in cultivated
fields near the three western Massachusetts oxbows was recognized

412 Rhodora [Vol. 86

by the mid-Nineteenth Century (Judd, 1863). Non-natives are now
of considerable importance in various stages of old field succession
there (Walker, 1980, unpublished Master’s thesis, Smith College,
Northampton, Mass.). The land surrounding the oxbows has been
heavily cultivated since the Colonial period or earlier, but non-
natives are scarce in the oxbow forests. At the Hatfield oxbow.
Lysimachia nummularia contributes 7% cover and Glechoma hede-
racea, Myosotis scorpioides, and Oxalis europaea occur at low
abundance. Commelina communis, Lysimachia nummularia, and
Solanum dulcamara are present though not abundant at Ned’s
Ditch. No non-natives at all were sampled at Whately.

Non-native species are also relatively unimportant in the herb
strata of the floodplain and terrace sites along the Raritan (Wisten-
dahl, 1958). Although these areas are surrounded by agricultural
land, only five of the 56 herbs reported are introduced and only
two, Lysimachia nummularia on the inner floodplain and Alliaria
officinalis on the outer floodplain, contribute coverage greater than
1%

In contrast, Keammerer et al. (1975) found that 18.6% of the
flora of the Missouri River bottomland forests in North Dakota
were non-natives, primarily of European or Eurasian origin. Some
of these occurred only on disturbed sites or were persistent after
cultivation; others were common or abundant. Johnson et al. (1982)
reported recent changes in the composition of the North Dakota
forests, including poor reproduction of major Overstory species,
resulting from altered stream flow and flooding regime. The higher
representation of non-natives in the flora may also reflect disrup-
tions of this sort.

CONSERVATION

Floodplain habitats along major streams in New England are
severely restricted. They are also increasingly threatened by urban
development and by plans to reduce or eliminate flooding (Brower,
1971; Franz and Bazzaz, 1977; Sackett and Nagazina, 1978). The
entire Northampton oxbow, which includes Ned’s Ditch, is owned
by the Massachusetts Audubon Society and is largely preserved.
Both the Hatfield and Whately oxbows, however, suffer multiple
private ownerships which might allow unrestricted logging at any
time. In addition, proposed alterations of flow on the Connecticut

1984] Holland & Burk — Oxbow herb strata 413

River may result in trends on the floodplain similar to those
reported by Johnson et al. (1982), whether logging is restrained or
not. The Massachusetts oxbows are of particular interest because
they support vegetation which has developed on sites available for
colonization during differing lengths of time within a single river
system. They support floristic assemblages substantially different
from those described elsewhere and therefore deserve more rigorous
legal protection than that afforded them at present.

ACKNOWLEDGMENTS

Much of the data on which this paper is based are included as
part of a Ph.D. dissertation written at Smith College and submitted
by M. Holland to the Five-College Ph.D. program at the University
of Massachusetts, Amherst. M. Holland has also published as Mar-
jorie Holland Sackett. We thank Lincoln P. Brower, H. Allen Cur-
ran, David Mulcahy, and Marshall Schalk for helpful discussions
on several phases of this work, Carolyn Crane and Paul Griffin for
help with the map, William Clapham, Wendy Coleman, Florence
Holland, Rich Kesselli, Kevin Richardson, Russell Sackett, Karen
Stone and Willow Zuchowski for invaluable assistance with the
field work, and Robie Hubley, former director of Arcadia Wildlife
Sanctuary, for encouragement and logistic support throughout the
course of this and previous studies in the area.

LITERATURE CITED

BARNES, W.J. 1978. The distribution of floodplain herbs as influenced by annual
flood elevation. Trans. Wisc. Acad. 66: 254-266.

BELL, D.T. 1974. Studies on the ecology of a streamside forest: composition and
distribution of vegetation beneath the tree canopy. Bull. Torrey Bot. Club 101:
14-20.

Brower, L. P. 1971. Biology of the annual flow cycle of the Connecticut River.
Connecticut River Ecology Action Corporation I: 14-24.

BuELL, M. F. AND W. A. WISTENDAHL. 1955. Floodplain forests of the Raritan
River. Bull. Torrey Bot. Club 82: 463-472.

BurK, C. J. 1973. Partial recovery of vegetation in a pollution damaged marsh.
Water Resources Research Center. Univ. Mass. Amherst Publ. No. 27.

1977. A four year analysis of vegetation following an oil spill in a fresh-

water marsh. J. Appl. Ecol. 14: 515-522.

AND S. D. LAUERMANN. 1977. Catalpa speciosa naturalized in western

Massachusetts. Rhodora 79: 305-308.

414 Rhodora [Vol. 86

CoLtins, R. F. AND M. SCHALK. 1937. Torrential flood erosion in the Connecti-
cut Valley, March, 1936. Amer. Jour. of Science 34; 293-307.

Crow, G. E., W. D. Countryman, G. L. Cuurcn, L. M. EASTMAN, C. B. HELL-
Quist, L. L. MEHRHOFF, AND I. M. Storks. 1981, Rare and endangered
plant species in New England. Rhodora 83: 259-299.

DAUBENMIRE, R. 1968. Plant Communities, a Textbook of Plant Synecology.
Harper and Row, New York.

FERNALD, M. L. 1950. Gray's Manual of Botany, Eighth Ed. American Book
Company, Boston.

FRANZ, E. H. AND F. A. Bazzaz. 1977, Simulation of vegetation response to
modified hydrologic regimes: a probabilistic model based on niche differentia-
tion in a floodplain forest. Ecology 58: 176-183.

Harbin, E. D. AND W. A. WISTENDAHL. 1983. The effects of floodplain trees on
herbaceous vegetation patterns, microtopography and litter. Bull. Torrey Bot.
Club 110: 23-30.

HoLLanp, M. M. AND C. J. BuRK. 1982. Relative ages of western Massachusetts
oxbow lakes. Northeastern Geology 4: 23-32.

Jauns,R.H. 1947. Geologic features of the Connecticut Valley, Mass., as related
to recent floods. Geol. Survey Water-Supply Paper 996. U. S. Government
Printing Office, Washington, D. C.

JOUNSON, W.C., P. W. ReiLy, L. S. ANDREWS, J. F. MCLELLAN, AND J. A. BRopHy.
1982. Altered hydrology of the Missouri River and its effects on floodplain
forest ecosystems. Virginia Water Resources Research Center, Virginia Poly-
technic Inst. and State Univ. Blacksburg Bull. 139.

JUDD, SYLVESTER. 1863. History of Hadley. Metcalf and Co., Northampton,
MA.

KEAMMERER, W.R., W. C. JOHNSON, AND R.L. BURGESS. 1975. Floristic analysis
of the Missouri River bottomland forests in North Dakota. Can. Field Nat. 89:
5-19,

Linpsry, A. A., R. O. Petty, D. K. STERLING, AND W. VAN ASDALL. 1961. Vege-
tation and environment along the Wabash and Tippecanoe Rivers. Ecol.
Monogr. 31: 105-156.

MUELLER-Dompols, D. AND H. ELLENBERG. 1974. Aims and Methods of Vegeta-
tion Ecology. John Wiley & Sons, New York.

Nicuois, G. E. 1915. The Vegetation of Connecticut - 1V. Plant societies in low-
lands. Bull. Torrey Bot. Club 42: 169-217.

Nicnors, G. E. 1916. The vegetation of Connecticut - V. Plant societies along
rivers and streams. Bull. Torrey Bot. Club 43: 235-264.

OpuM, E. P. 1969. The strategy of ecosystem development. Science 164: 262-270.

Rowinton, E. D. AND C. J. BuRK. 1971. The Mill River and its floodplain in
Northampton and Williamsburg, Mass.: a study of the vascular plant flora,
vegetation, and the presence of the bacterial family Pseudomonadaceae in rela-
tion to patterns of land use. Water Resources Res. Center. Univ. Mass. Amherst.
Completion Report 72-4.

SackeTT, M. H. (HOLLAND) AND J. NAGAZINA. 1978. Northfield Diversion Pro-
ject. Appalachia Journal 12: 104-111,

1984] Holland & Burk — Oxbow herb strata 415

VANKAT, J. L. 1979. The Natural Vegetation of North America. John Wiley and

Sons, New York.
Wuite, P.S. 1979. Pattern, process, and natural disturbance in vegetation. Bot.

Rev. 45: 229-299.
WISTENDAHL, W. A. 1958. The floodplain of the Raritan River, New Jersey.

Ecol. Monogr. 28: 129-153.

DEPARTMENT OF BIOLOGICAL SCIENCES

SMITH COLLEGE
NORTHAMPTON, MASS. 01063

A NEW BASIC CHROMOSOME NUMBER IN THE GENUS
STELLA RIA (CARYOPHYLLACEAE)

J. K. MORTON

ABSTRACT

Chromosome numbers are reported for two closely related species from the decid-
uous forest region of eastern North America, Ste/laria pubera Michx. (2n = 30) and
S. corei Shinners (2 = 60). These counts represent a new basic chromosome number
of 15 for the genus Sre/laria.

Key Words: Stellaria, basic chromosome number, North America

INTRODUCTION

Stellaria is a large, very variable and cosmopolitan genus of the
Caryophyllaceae with about 120 species. Its main centers of diver-
sity are in the temperate and sub-temperate regions of Europe, Asia
and North America, but representative species occur in the mon-
tane floras of the tropics. Basic chromosome numbers of x = 10, II,
12 and 13 have been reported for the genus, but almost all the native
North American species that have been examined have a base
number of 13 (Federov, 1969; Moore, 1973; Goldblatt, 1981). As
part of a survey of chromosome numbers in the North American
Caryophyllaceae, Stellaria pubera and S. corei were examined.
Their respective chromosome numbers of 2n = 30 and 60 are here
reported for the first time and provide a new basic chromosome
number of 15 in the genus Sre//aria.

MATERIALS AND METHODS

Rooted material was transplanted into the greenhouse and grown
in clay pots in a sterile potting-soil mixture. Root-tips were re-
moved from actively growing plants, pre-treated for about two
hours in a saturated aqueous solution of paradichlorobenzene,
fixed in 1:3 acetic acid:ethanol, hydrolyzed for 15 mins. at 60°C. in
N-hydrochloric acid, rinsed in water and squashed in aceto-
carmine. A few drops of ferric chloride solution were added to the
rinse water before squashing, to intensify staining. Also, flower-
buds were fixed in the field in Carnoy’s solution and the pollen
mother cells subsequently squashed in aceto-carmine. The plant

417

418 Rhodora [Vol. 86

Table |. Source of materials of Srellaria

n 2n
S. pubera
Morton NA4435 Asheville, N.C. 30
Morton NA4460 Cheoah Reservoir, Fontana, N.C. 30
Morton NA5710 Cherokee, N.C. 15 -
Morton & Venn NAI5014 Whitley to Honeybee, 30

McCreary Co., Ky.

S. corei
Morton NA4444 Gatlinburg, TN 60
Morton NA5705 Cherokee, N.C. 60
Morton & Venn NA14969 Linville, Avery Co., N.C. 60

materials used in this study are listed in Table |. Voucher specimens
have been deposited in the following herbaria: WAT and JKM,
with incomplete sets in CAN and MICH.

RESULTS

Four populations of Sre/laria pubera were examined, three using
root-tips and one using flower-buds. They had chromosome
numbers of 27 = 30 and n = 15 respectively. Pollen mother cells
showed a regular configuration of 15 bivalent chromosomes at dia-
kinesis. Three populations of the much rarer S. corei each had 2n =
60 chromosomes in root-tip preparations.

DISCUSSION

Stellaria pubera and S. corei are part of the endemic flora of the
Carolinian region in the deciduous forest zone of eastern North
America. This is an ancient flora and characterizes the mixed decidu-
ous mesophytic forests which have had a continuous existence in
this part of North America since the mid-Tertiary era (Graham
1972). These two species are closely related and morphologically
very similar. S. corei appears to be a polyploid derivative of the
diploid S. pubera. Classification within the genus usually follows
that proposed by Fenzl in Endlicher (1840). Under this system S.
pubera and S. corei belong to Section Eu-Stellaria, along with most
of the familiar North American and European species. The division
of this section into subsections is based on leaf characters and

1984] Morton — Stellaria 419

appears to be unsatisfactory. Neither of the species under consider-
ation fits clearly into any of the five subsections.

LITERATURE CITED

ENDLICHER, S. L. 1840. Genera Plantarum 13: 961-1040. Wien.

FeperRoV, AN. A., editor. 1969. Chromosome numbers of flowering plants.
Leningrad. 927 pp.

GOLDBLATT, P., editor. 1981. Index to plant chromosome numbers 1975-1978.
Missouri Botanical Garden Monographs in Systematic Botany No. 5. 553 pp.

GRAHAM, A., editor. 1972. Floristics and paleofloristics of Asia and Eastern
North America. Elsevier Pub. Co. New York. 278 pp.

Moore, R. J., editor. 1973. Index to plant chromosome numbers. 1967 1971].
Regnum Vegetabile 90.

DEPARTMENT OF BIOLOGY
UNIVERSITY OF WATERLOO
WATERLOO, ONTARIO
CANADA N2L 3G!

A FIRST REPORT OF THE FERN GENUS VITTA RIA
IN NEW YORK

JAMES C. PARKS AND DONALD R. FARRAR

ABSTRACT

The first New York State records for gametophytes of the fern genus Visraria are
reported from Cattaraugus and Chautauqua Counties. The Chautauqua Co. station
is only the third in a glaciated area and is the northernmost location known, The
stations cited also provide the first county records in western New York for Tricho-
manes gametophytes.

Key Words: Vittaria, gametophytes, state record, Trichomanes

The morphology, ecology, and biogeography of sporophyteless
gametophytes of the fern genera Vitraria and Trichomanes have
been described by Farrar (1967, 1978) and McAlpin and Farrar
(1978). Farrar et al. (1983) established state records for these genera
in Pennsylvania and expanded considerably the known range of
Trichomanes in New England. Noting that all sites reported for
Vittaria were south of the Wisconsinan terminal moraines, these
workers suggested that its pre-Pleistocene distribution may have
been truncated by glaciation. Cusick (1983) subsequently reported
Vittaria gametophytes from 80 km north of the Wisconsinan termi-
nal moraine in Geauga County, Ohio.

The first reports of Vitraria for New York State are recorded
herein as follows:

1. Cattaraugus Co. August 3, 1983. Rock City Park, Rt. 646.

Scattered in large NW-facing rock houses of Pottsville sandstone

conglomerate, in woods. Parks 4295.

2. Cattaraugus Co. August 3, 1983. Three miles W of Rt. 646

on Nichols Run Road. Very rare in shaded crevice of SW-facing

outcrop of Pottsville sandstone conglomerate, in woods. Parks

4296.

3. Chautauqua Co. August 3, 1983. Panama Rocks Park. Scat-

tered populations of gemmiferous plants in massive 10 m high,

SE-facing rock houses of Pottsville sandstone conglomerate, in

woods. Area glaciated. Parks 4298.

Trichomanes gametophytes were also located at sites one and
three above (Parks 4294 and 4297 respectively) and at Bear Cave

421

422 Rhodora [Vol. 86

Rocks in Allegany Park, Cattaraugus Co., on August 2, 1983,
Parks 4293. Vouchers of Parks’ cited collections are on deposit at
MVSC.

The Chautauqua County station at Panama Rocks lies in a gla-
ciated region (Muller, 1963) and was itself ice covered (Muller,
personal correspondence, 1983). The Wisconsinan-Kent terminal
moraine lies about 19 km to the southeast and the Illinoian-Maple-
dale terminal moraine lies an additional 6.2 km further away (Berg,
1980). Panama Rocks is the northernmost station reported for Vit-
‘aria and is quite different from the surrounding area of Chautau-
qua County. It is part of an outlying ridge of Pennsylvanian
sandstone conglomerate (Pottsville formation or equivalent).
Though the Cattaraugus County site to the southeast is inan ungla-
ciated portion of New York knownas the Salamanca reentrant, it is
strikingly similar geologically and ecologically to Panan Rocks.
Further, judging from Cusick’s (1983) description of Vittaria habi-
tat in glaciated Ohio, that site is also similar to the Panama Rocks
station. These observations support Cusick’s argument that the
occurrence of Vittaria gametophytes probably is closely related to
appropriate physical and geological factors.

Post-Pleistocene reestablishment. in glaciated areas of these
sporophyteless fern gametophytes has been discussed by Farrar et
al. (1983). The report of Vittaria from glaciated western New York
is COnsistent with this discussion. Dispersal mechanisms and other
factors which might influence the differential distribution of
Vittaria and Trichomanes in the glaciated NE United States and
adjacent Canada remain enigmatic. Workers are encouraged to
examine carefully other appropriate sites norti of the terminal
moraines for Vittaria gametophytes. Though less likely to be
encountered there than the gametophytes of Trichomanes, those of
Vittaria clearly may be found.

ACKNOWLEDGMENT

Grateful acknowledgment is expressed to Dr. Eugene Williams,
Professor of Biology, Edinboro University of Pennsylvania, who
provided invaluable field assistance and kind hospitality to the
senior author.

1984] Parks & Farrar — Vittaria 423

LITERATURE CITED

BERG, T. (chief compiler). 1980. Geologic Map of Pennsylvania. Harrisburg: PA
Geological Survey Map |.

Cusick, A. 1983. Vittaria gametophytes discovered in a new physiographic pro-
vince. Amer. Fern J. 73: 33-38.

FarRAR, D. 1967. Gametophytes of four tropical fern genera reproducing inde-
pendently of their sporophytes in the southern Appalachians. Science 155:
1266-1267.

1978. Problems in the identity and origin of the Appalachian Vitraria

gametophyte, a sporophyteless fern of the eastern United States. Amer. J. Bot.

65: 1-12.

J. Parks AND B. MCALPIN. 1983. The fern genera Virtaria and Tricho-
manes in the northeastern United States. Rhodora 85: 83 92.

MCALPIN, B. AND D. FARRAR. 1978. Trichomanes gametophytes in Massachu-
setts. Amer. Fern J. 68: 97-98.

MULLER, E. 1963. Geology of Chautauqua County, New York, Part II, Pleisto-
cene Geology. New York State Museum and Science Service Buil., v. 392.

J; €. Ps

DEPARTMENT OF BIOLOGY
MILLERSVILLE UNIVERSITY
MILLERSVILLE. PA 17551

DR: F.

DEPARTMENT OF BOTANY
IOWA STATE UNIVERSITY
AMES, IA 50011

LOMATOGONIUM ROTATUM (GENTIANACEAE) AND
PRIMULA LAURENTIANA (PRIMULACEAE)
IN MAINE: NEW LOCALITIES AND
GENERAL DISTRIBUTIONS

NORMAN C. FAMOUS AND CHRISTOPHER S. CAMPBELL

ABSTRACT

Two herbaceous plant species, Lomatogonium rotatum and Primula laurentiana,
are known to occur in the United States only on the offshore islands of eastern
Maine. Both species grow in full sun and in thin, circumneutral soil just above the
upper splash zone of the ocean. We report new localities and discuss the overall
geographic distribution for these two species.

Key Words: Lomatogonium rotatum, Primula laurentiana, Maine, range limit, off-
shore islands

Several plant species reach the southern limit of their range along
the coastal headlands and islands of the west coast of the Bay of
Fundy (Fernald and Wiegand, 1910; Hodgdon and Pike, 1964;
Olday et al., 1982). A maritime climate of cool summer tempera-
tures, extensive rainfall and fog, and low evapotranspiration rates
account, in part, for this phytogeographic pattern. In this region,
two boreal species, Lomatogonium rotatum (L.) Fries, the star gen-
tian or marsh felwort, and Primula laurentiana Fern., the bird’s-eye-
primrose, are now known from only one location each in New
Brunswick and a total of 16 offshore islands in eastern Maine. We
report discovery of eight of these stations during our field work ona
more detailed study of the reproductive biology of P. /aurentiana.
We also characterize the local habitat and overall distribution of
these two taxa.

Stebbins (1929) first discovered Lomatogonium rotatum in
Maine at Schoodic Point. This population has not been seen
recently, but this species has since been found on nine islands in
eastern Maine (Table 1; Figure 1). In 1982 and 1983 we discovered
five of them and verified those previously reported. In all of these
localities, L. rotatum grows in full sun and in thin, granitically
derived, mineral soil and organic duff in a narrow belt between the
forest edge and the upper splash zone above the intertidal zone. It
also occurs in areas of high soil moisture near brackish pools and in
rock crevaces serving as drainage tracks from the upland. This plant

425

426 Rhodora [Vol. 86

Table I. Historic, current and new localities of Lomatogonium rotatum! and
Primula laurentiana in eastern Maine. (Numbers correspond to map locations in-
dicated in Figure 1.)

Estimated number of
individuals of

Island L. rotatum P. laurentiana
1. Sand Island (outer) extant, new
2. Seal Rock 0
3. Curlew Rock extant, new 15, new
4. Crumple Island extant 100°s
5. Great Wass Island extant 100,000
6. Unnamed Island North

of Water Island extant, new 400, new
7. Water Island (inner) extant, new 400, new
8. Water Island (outer) extant, new
9. Mistake Island extant 1,000
10. Knight's Island unknowns
11. Head Harbor Island unknown?
12. North Scabby Island 100,000;
13. Little Scabby Island no estimate?
14. Ram Island 03
15. South Libby Island 10,0004
16. North Libby Island 04

'Also extant, but not shown on Figure |, on Little Moose Island (Olday, Gawler &
Vickery, 1982); also known historically from Schoodic Point (Stebbins, 1929).
?Odlay, et al., 1982.

‘Lewis (1983).

‘Lewis (pers. comm.)

‘Norton (pers. comm.)

has probably been overlooked because it is small and rather incon-
spicuous, occupies a specialized habitats, and flowers after the peak
of summer field work by botanists. Undoubtedly it grows on other
islands east of Frenchman’s Bay. Species associated with L. roratum
include: Agrostis stolonifera L., Aster novi-belgii L., A. nemoralis
Ait., Campanula rotundifolia L., Carex canescens L., C. viridula
Michx., Deschamsia flexuosa (L.) Trin., Empetrum nigrum L.,
Euphrasia canadensis Townsend, E. randii Robins.. Festuca rubra
L., Iris hookeri Penny, Juncus filiformis L. (vel aff.), Plantago jun-
coides Lam. var. decipiens (Barnéoud) Fern., Prenanthes trifoliata
(Cass.) Fern., Primula laurentiana, Sagina nodosa (L.) Fenzl. ssp.

1984] Famous & Campbell — Lomatogonium 427

Cy?
¢ YY 0,
G2, \
JONESPORT : 4 a
a)
. . _- @-» g (8)
r (3 4 ey! ao @© ©
7 “ey nm MACHIAS BAY
& (7) e Y
A - ‘ o : © 6{km___44° 30
e eo A é 7 :
oY @&I6

& 67° 25°
oc

be
@Go® GULF OF MAINE

Figure 1. Map of Islands off Jonesport and in Machias Bay, Maine. See Table |
for reference to numbered locations.

borealis Crow, Solidago bicolor L., Triglochin maritima L., and
Viola septentrionalis Greene. Authorities are those of Fernald
(1950).

At the two inland localities discovered in the nineteenth century,
Primula laurentiana has not been seen in this century (Critical Areas
Program, 1981). Cushman (1907) was the first to find coastal popu-
lations of the bird’s-eye-primrose, and Norton (1913), Pike (1963)
and Lewis (1983) added to our knowledge of its ecology and local
distribution. We located three new island sites (Table |; Figure 1),
uncovered one historical record (Knight’s Island) and determined
that this species no longer grows on Seal Rock. The plant is there-
fore now known to occur on nine islands and to be absent from
three historic sites; Its status for two additional localities is
unknown (Table |). The bird’s-eye-primrose is more likely to have

428 Rhodora [Vol. 86

attracted people’s attention because of its striking, early summer
flowers and light-green leaves than is the star gentian.

Primula laurentiana occupies the same habitats as Lomatogo-
nium rotatum, although the primrose appears to prefer better soil
drainage. The soils are derived from granitic bedrock, but soil tests
from five Primula localities indicate circumneutral pH (from 5.9 to
7.5, mean = 6.6) and oftentimes an excessive level of calcium. Prox-
imity to salt spray and shell clasts may account for this relatively
high pH. These tests verify the generally held view that this species is
a calciphile (Fernald, 1928; Lewis, 1983; Pike, 1963; Scoggin, 1979).

In the eastern United States, these two species are limited to
eastern Maine and are threatened or endangered in New England
(Crow et al., 1981). In New Brunswick, the star gentian is known
only from South Wolf Island (Hodgdon and Pike, 1963) and the
bird’s-eye-primrose only from one recently located population of
about 150 individuals in the coastal town of Alma (H. R. Hinds,
pers. comm.). Primula laurentiana is also rare in Nova Scotia
(Maher er al., 1978). Globally, Lomatogonium is a circumpolar
halophyte while Primula occupies meadows, ledges and cliffs chiefly
near the ocean over much of northeastern North America (Scoggin,
1979).

ACKNOWLEDGMENTS

This work was supported by grants from the Maine Chapter of
The Nature Conservancy and the Critical Areas Program of the
Maine State Planning Office. We thank the Maine Soil Testing
Laboratory for the soil tests.

LITERATURE CITED

CRITICAL AREAS PROGRAM. 1981 Rare vascular plants of Maine. State Planning
Office, Augusta, Maine 04333. 656 pp.

Crow, G. E., W.S. CouNTRYMAN, G. L. CuurcH, L. M. EASTMAN, C. B. HELLQUIST,
L. J. MEHRHOFF AND I. M. Storks. 1981. Rare and endangered vascular
plant species in New England. Rhodora 83:259- 299.

CusuMan, J. A. 1907. Primula farinosa var. macropoda on the Maine coast.
Rhodora 9:217, 218.

FERNALD, M.L. 1928. Primula section Farinosae in America. Rhodora 30:59-77,
85-104.

1950. Gray’s Manual of Botany. Eighth Edition. American Book Co.,

Boston. 1632 pp.

1984] Famous & Campbell — Lomatogonium 429

.AND K.M. WIEGAND. 1910. A summer's botanizing in eastern Maine and
western New Brunswick, Part I. Rhodora 12:101-121.

Hopcpon, A. R. AND R. B. Pike. 1963. The Flora of the Wolf Islands, New
Brunswick, Part I. Rhodora 65:82-96.

and . 1964. Flora of the Wolf Islands, New Brunswick, Part 2. Some
phytogeographic considerations. Rhodora 66:140-155.

Lewis, A.J. 1983. Distribution of three rare plants on islands in Machias Bay, Maine.
Rhodora 85:385-388.

Mauer, R. V., D. J. Wuite, G. W. ArGus AND P. A. Keppy. 1978. The rare
vascular plants of Nova Scotia. Syllogeus No. 18. National Museums of Can-
ada. Ottawa.

Norton, A. H. 1913. Some noteworthy plants from the islands and coast of
Maine. Rhodora 15:137-143.

Otpay, F. C., S. C. GAWLER, AND B. Vickery. 1982. Seven unusual sub-arctic
plants of the Maine Coast. Planning Report No. 78. Critical Areas Program,
Maine State Planning Office, Augusta.

Pike, R. B. 1963. Note on Primula laurentiana in Maine. Rhodora 65:286- 188.

ScoGGIN, H. J. 1979. The Flora of Canada. Part 4 - Dicotyledoncae (Loasceae to
Compositae). National Museum of Natural Sciences Publications in Botany,
No. 7(4).

STEBBINS, G. L. 1929. Lomatogonium rotatum (L.) Fries, in Maine. Rhodora
31:143.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
UNIVERSITY OF MAINE
ORONO, MAINE 04469

POPULATION STRUCTURE AND ENVIRONMENTAL
COROLLARIES OF PANAX QUINQUEFOLIUM
(ARALIACEAE) IN DELAWARE COUNTY,
NEW YORK

WALTER H. LEwIs

ABSTRACT

A large population of Panax quinquefolium from New York is characterized by
age structure, morphology, and flowering. Population dynamics are correlated with
local climate, the most interesting association being the dramatic increase in popula-
tion size during 1982 when monthly precipitations between January and June com-
pared favorably with 20-year averages at a time when the need for protective snow
cover for seeds and optimum conditions for seed germination would apparently be
paramount.

Key Words: Panax quinquefolium, American ginseng, population, environment.
New York

INTRODUCTION

Panax quinquefolium L. (American ginseng) is a rare or extir-
pated species in northeastern United States and adjacent Canada
where it once thrived. To locate a large population of some age that
is expanding in a natural ecosystem is uncommon. Thus, the discov-
ery by Arthur W. Rashap and Edmund Millar of such a population
near Roxbury, Delaware County, in the Catskill Mountains of east-
ern New York, where the species at one time must have been fre-
quent, was important. It prompted me to initiate a study of
population dynamics similar to one underway in Missouri (Lewis
and Zenger, 1982), which could serve not only as a comparison with
this geographically different site, but could apply to programs for
re-establishing the species in New York. The first of these goals is
partially fulfilled by this report, insofar as population structure 1s
concerned, but the second proved impossible. Within weeks of
recording the data on June 30, 1982, the population was eradicated
by root diggers.

MATERIALS AND METHODS

The population was found in a well-drained, sloping, deciduous
woods with deep humus accumulated from leaf litter. Plants were

431

432 Rhodora [Vol. 86

described, labeled, numbered, and located on a map for future ref-
erence. Several were photographed for permanent record and prints
are deposited at MO (Lewis 9778). Rhizomes were carefully exposed
to count bud scars in order to determine ages and then re-covered.
U.S. climatological data for New York include only four stations
with precipitation and two having temperature records in Delaware
County during a 20-year period from 1963-82. These reports are
summarized monthly for a 6-year period, 1977-82, together with
20-year means and standard deviations. The station near Roxbury,
which is situated very close to the population, is not included, for
recording of data terminated in 1972. Nevertheless, the other county
stations are sufficiently near and similar in their situation to present
approximate environmental conditions for the population site.

RESULTS AND DISCUSSION

Age structure and morphology

For a population of 211 individuals, ages in 1982 ranged from | to
21 years (Table 1), plus an exceptional plant 48 years old. This plant
not only was 27 years older than the next oldest plant in the popula-
tion, but the aerial stem was strongly flattened and the leaflets were
very coarse and thick; it does not figure further in this analysis. A
second exceptional plant had two aerial stems, one having three
prongs (15 leaflets) and an inflorescence with nine flowers, and the
other with two prongs (10 leaflets) and seven flowers. For analytical
purposes, these stems are considered representative of two plants.
The most predominant age class, representing 56.9% of the whole
population, was of seedlings that had germinated during the spring
of 1982. Plants. of two and three years of age were next most com-
mon, but each represented only about 10% of the population. Those
between 4 and 11 years of age varied from 1.8 to 3.8%, whereas all
plants established between 1963 and 1971 were represented by a
total frequency of only 3.8%. Thus, annual recruitment represented
by surviving plants was gradual for 18 years, followed by modest
expansions during 1980 and 1981, and culminated by an explosive
increase during 1982. There is, of course, no certain way of knowing
whether these frequencies represent nearly equivalent numbers of
plants established annually or not, but based on the high annual
survival rate of 97% for a large Missouri population (Lewis and
Zenger, 1982), it isa reasonable assumption. The interesting ecolog-

1984] Lewis — Panax 433

Table 1. Population structure of 211 plants of Panax quinquefolium from
Roxbury, NY.
Flowers
Prongs Leaflets Flowering flowering
Frequency plant plant plants plants
Age (%) (average) (average) (%) (average)
l 56.9 1.0 3.2 0 0
2 10.4 1.5 6.2 13.6 2.7
3 9.5 22 10.1 60.0 13
4 3.8 2.4 11.4 87.5 8.4
5 1.8 2.8 12.8 75.0 12.7
6 1.8 1.8 12.8 100 14.0
7 3.3 2.6 12.7 100 8.2"
8 2.8 3.0 15.0 100 16.7
9 1.4 2.3 12.0 100 225"
10 1.8 3.0 15.0 100 22.0"
1] 2.4 3.0 14.8 100 15.4
12-21 3.8 3.0 15.0 87.5 20.7*

*One inflorescence broken or chewed and flower number could not be determined.

ical observations of Huet al. (1980) involving a quadrat of 24 plants
in Connecticut showed a death rate of 58.3% over an interval of
seven years, but this high mortality may be biased because of small
sample size.

Morphologically the population consisted of four classes based
on prong and leaflet numbers (a prong consists of 3-6 palmately
compound leaflets and a petiole without an axillary bud at the
juncture of the aerial stem typical of leaves; most flowering plants
have 2-4 prongs and an inflorescence in a whorl). One-pronged
plants had 3-5 leaflets, predominantly with 3 leaflets and only infre-
quently 4 (0.8%) or 5 (3.3%), two-pronged plants had 6— 10 leaflets,
three-pronged plants had 14-16 leaflets, and the one four-pronged
plant had 19 leaflets. Five leaflets per prong were most common,
those with 3 or 4 occasional, and those with 6 rare among plants
with two or more prongs.

Most one-pronged plants were one year old, although a few were
two (9.0%) or 3 years (0.8%) of age. Two-pronged plants varied in
age from 2 to 9 years, with an exceptional one being 19 years old, for
an average of 3.9 years. The majority were two (26.0%), three
(40.5%), or four (13.5%) years old. Three-pronged plants also varied
markedly in age from 3 to 21 years. They averaged 8.6 years, but

434 Rhodora [Vol. 86

their numbers were rather constant between years 3 and I1 (min-
imum of 3 plants or 7.5% and maximum of 6 plants or 15.0%), and
consistently very few (0 or 1) between ages 12 and 21. The single
4-pronged plant was 17 years old. There was a strong correlation
between age and morphological class as measured by both prong
number (r = 0.798, P<0.001) and leaflet number (r = 0.799,
P<0.001), and thus the population illustrated an orderly increase in
size and leaf area with age, as found in Missouri (Lewis and Zenger,
1982).

Flowering

All one-pronged plants were juveniles (nonflowering), 45.9% of
two-pronged plants were also juveniles, but all three-pronged and
the one four-pronged plants formed inflorescences. Of the two-
pronged flowering plants, those two years old averaged 2.7 flowers
(or floral buds) per plant, while older ones averaged 4 to 5 flowers
depending on age. They ranged from 1-12 per plant with an overall
average of 4.4 flowers. Interestingly, the 19 year old two-pronged
plant had no discernable inflorescence. Flowers of three-pronged
plants ranged from 5-32 with an average of 16.2 flowers per plant.
The four-pronged plant had 31 flowers. There were highly signifi-
cant correlations between flower numbers and morphological class
(prong number r = 0.817, P<0.001, leaflet number r = 0.822,
P<0.001) and also age (r = 0.724, P<0.001), as reported for pop-
ulations in Missouri (Lewis and Zenger, 1982) and Wisconsin (Car-
penter and Cottam, 1982). However, two-pronged plants rarely
flowered in Wisconsin in contrast to those in Missouri and New
York where flowering reached 55.8% and 54.1%, respectively.

Environmental Corollaries

Based on survivorship in 1982, only 15.6% of the population was
established in the 5-year period from 1962 to 1976. During the
subsequent five years the number of plants increased by 25.6%, a
modest rise perhaps reflecting additional propagules in an aging and
slowly increasing population. Not until 1982 was there a large
increase in plants when 56.9% of the population was established ina
single year. To explain this dramatic rise solely by small population
increases and propagule accumulations seems unlikely based on the
modest reproductive performance in the immediate past.

1984] Lewis — Panax 435

As growth and development patterns were altered markedly ina
Missouri population of P. quinquefolium during adverse environ-
mental conditions (Lewis and Zenger, 1982), it is of interest to com-
pare the New York population structure during the past few years
with local climatic conditions during the same period. The latter are
summarized for monthly precipitation and temperature measure-
ments in Table 2, as well as for 20-year averages. For the 5-year
period prior to 1982, one striking aspect of the precipitation data

Table 2. Precipitation and temperature records for Delaware Co., NY.

20-yr

1982 1981 1980 1979 1978 1977 Mean + S.D.
Precipitation (inches)!
January 3.32 0.64 1.05 6.69 5.72 1:25: : Dal Tice 1.63
February 2.21 5.78 0.95 1.56 1.20 2.65 2.41 + 1.20
March 2.47 0.74 5.59 3.42 2.42 6.55 3.22 + 1.31
April 2.22 3.28 3.96 3.70 1.70 3.38 3.08 + 0.86
May 3.84 4.22 1.05 5.56 3.48 4.03 3.88 + 1.45
June 3.89 3.53 4.74 1.27 3.59 3.02 3.94 + 1.75
July 3.47 2.95 3.77 3.63 2.55 3.83 + 1.39
August 3.16 4.21 2.50 5.14 5.49 3.82 + 1.20
September 5.54 3.44 5.03 3.36 10.21 3.74 + 2.07
October 4.30 3.17 5.22 3.95 7.32 3.28 + 1.91
November 3.96 3.28 2.62 1.45 4.98 3.54 + 1.37
December 1.55 1.59 2.38 3.25 480 3.35 + 1.61
Total 40.15 35.97 43.69 38.88 56.22 40.87 + 6.34
Temperature (°F)
January 14.95 14.65 24.35 22.80 20.15 13.10 20.3 + 4.8
February 24.75 31.60 20.15 13.55 15.60 23.50 21.8 + 4.5
March 31.25 34.60 31.90 37.35 29.55 36.70 32.8 + 3.2
April 42.35 47.85 46.30 4450 42.75 4640 43.9 + 3.5
May 57.45 56.20 57.05 57.25 55.80 57.95 §5.2 + 3.1
June 59.75 64.30 60.50 62.65 62.45 61.15 63.8 + 2.1
July 68.40 68.60 68.75 66.70 67.95 68.4 + 1.4
August 63.60 70.25 66.70 69.20 66.85 66.5 + 2.3
September 59.15 61.25 58.50 58.10 60.90 59.7 + 2.0
October 48.80 46.05 45.60 47.45 47.30 48.7 + 3.4
November 42.90 34.80 39.05 38.75 42.00 39.0 + 3.4
December 31.10 21.05 25.80 27.65 25.00 27.1 + 3.3

!Arkville (42° 08’, 74° 39’, 1310 ft), Deposit (42°04’, 75° 26’, 1000 ft), Downsville Dam
(42°05’, 74°58’, 1300 ft), and Kortright (42°25’, 74°48’, 1720 ft).
?Deposit and Downsville Dam.

436 Rhodora [Vol. 86

was the irregular monthly averages during the first part of each year.
Thus, very low precipitation occurred either in January or February
each year from 1977 to 1981 when much moisture would be in the
form of snow. If adequate, snow would serve as an important insu-
lation for seeds during the coldest months of the year (averages
13.1-20.2°F), but when markedly reduced, as from 1977 to 1981,
this lack of cover might prove deleterious to seed viability. Also,
during these years, precipitation was sometimes much lower than
average in April or May when seed germination and seedling estab-
lishment would have been paramount; reduced moisture at this time
would clearly hinder maximal recruitment. In 1981, for example,
January precipitation totaled only 0.6 in. (20-year average 2.8 in.)
during one of the coldest months in Delaware County (average
14.7°F). In 1980, the situation was even more extreme, for not only
was there below average precipitation (0.95 in. versus 20-year aver-
age 2.41 in.) during the coldest month of the year (February, aver-
age 20.2°F), but May proved low in precipitation (1.05 in. versus
20-year average 3.88 in.) during an important month for germina-
tion and seedling development. For the first six months of 1982,
however, monthly precipitation averages compared favorably with
the 20-year averages. This presumed norm had not prevailed in
Delaware County since 1974-76. That no appreciable population
increase occurred during this period is perhaps explained by the fact
that only 20 to 33 plants existed in the population, many being
juveniles, and thus few propagules and little recruitment would have
been possible even under ideal conditions.

These corollary data suggest that local environmental factors are
very significant for the recruitment and survival of American gin-
seng populations. It is unfortunate, but because of the destruction of
this important population, such a relationship cannot be tested
further.

ACKNOWLEDGMENTS

| appreciate the assistance of Arthur W. Rashap, Ginseng
Research Institute, and Edmund Millar, Cold Mountain Farm,
both of Delaware Co., New York for locating the population and
helping in the field, and the O'Connor Foundation and its Executive
Secretary, Donald Bishop, Hobart, New York, fora grant-in-aid to

1984] Lewis — Panax 437

study American ginseng in Delaware County. Special thanks go to
Dr. Vincent E. Zenger, Washington University School of Medicine.

LITERATURE CITED -

CARPENTER, S. G. AND G. CoTTAM. 1982. Growth and reproduction of American
ginseng (Panax quinquefolium) in Wisconsin, U.S.A. Canad. J. Bot. 60:
2692-2696.

Hu. S. Y., L. RUDENBERG, AND P. D. Trepici. 1980. Studies of American gin-
sengs. Rhodora 82: 627-636.

Lewis, W. H. AND V. E. ZENGER. 1982. Population dynamics of the American
ginseng Panax quinquefolium (Araliaceae). Amer. J. Bot. 69: 1483-1490.

DEPARTMENT OF BIOLOGY
WASHINGTON UNIVERSITY
ST. LOUIS, MISSOURI 63130

A 24-YEAR COMPARISON OF THE VEGETATION
OF AN ADIRONDACK MOUNTAIN SUMMIT

E. H. KETCHLEDGE AND R. E. LEONARD

ABSTRACT

A permanent 300-foot triangular transect was installed by H. E. Woodin in 1957
immediately above timberline on the isolated west slope of the summit cone of
Mount Marcy, elevation 5334 feet, highest peak in the Adirondack Mountains of
northern New York State. Re-measurement of the transect in 1981 revealed only
slight change in either the floristic composition of the transect or in the percentage of
arctic species present. Minor differences in data appear to be caused by different
personnel performing the measurements or by sampling error inherent in the line-
intercept procedures.

Key Words: Adirondack Mountains, permanent transect, arctic species

The value of permanent vegetation sampling sites for long-term
ecological studies is increasingly evident; their importance is partic-
ularly great in evaluating the impact of recreation upon the natural
environment. Fragile ecosystems such as alpine meadows are often
subject to intense trampling by hikers visiting scenic summits. More
recently, vegetational stability of alpine peaks in the northeast is of
concern because of increasing evidence of vegetational damage due
to acid deposition, especially in the high-elevation spruce-fir forests
of New York and New England. In the Adirondack Mountains, the
only permanent study site designed for alpine investigations was
established by H. E. Woodin (Woodin, 1959) on Mount Marcy, el.
5334 ft., highest peak in the region. Woodin selected an isolated
section of the 15-acre open summit zone extending above timberline
on the unfrequented western slope of the peak, far from the public
hiking trail and thus free from trampling by summit visitors. The
transect, an equilateral triangle 100 feet on a side and spanning the
elevations of 4,958 ft. to 4,984 ft., was intended to allow © uture
investigators to determine whether timberlines in the eastern United
States are ascending or descending”. The transect now serves
equally well as a baseline for studies of vegetational stability in
general. In 1981, we remeasured the transect to ascertain any floris-
tic changes or vegetational shifts that may have occurred over the
past twenty-four years.

439

440 Rhodora [Vol. 86

Comparison of the data from 1957 with those of 198] (Table 1)
showed little change in either flora or vegetation along the transect
during the 24-year period. Minor differences are attributable to
sampling error inherent in use of a line-intercept procedure in a
floristically diverse community which contained approximately
ninety-three species of vascular plants. Similarly, some variation
can be expected whenever different personnel remeasure the same
population. The most striking result was the apparent absence of
significant change in the virgin alpine meadow when contrasted
with the severe deterioration observed along the public hiking trail
several hundred feet upslope and out of sight of the study area.

FLORISTIC CHANGES

We note two additions to the list of species. Trientalis borealis
was a single plant. The species is frequent in the krummholz transi-
tion into the spruce-fir zone downslope but is rarely successful on
the wind-swept open summit. The occurrence of Vaccinium boreale
is not unexpected either. Its presence in the alpine zone of the
Adirondacks was first pointed out to us over fifteen years ago by
the late Stanley Jay Smith, Senior Curator, New York State Herba-
rium, Albany, who identified the 1957 plants for Dr. Woodin. Cur-
rently we have records for the species from seven alpine summits
and from five exposed stations at lesser elevations in New York
State. We report also that Vaccinium myrtilloides and V. angustifo-
fium occur sporadically near the timberline in the Adirondack
high country.

We are uncertain of the significance of absence of the Mountain
Alder and Cordate Birch from the 1981 data. The alder covered
eleven inches in 1957 and was likely a single plant. The birch, how-
ever, which covered forty-one inches, may have been two or more
plants. Both species are still present in the transect area but neither
had branches intercepting a vertical projection through the inter-
cept line. Plants nearby seemed otherwise healthy and vigorous at
the time of measurement.

VEGETATIONAL SHIFTS

The living plant cover decreased 1.43% during the 24-year period,
from 77.67% in 1957 to 76.24% in 1981. This minor shift may be

1984] Ketchledge & Leonard — Mt. Marcy 44]

Table |. Comparison of 1957 and 1981 permanent transect line-intercept data
on Mt. Marcy.

1957
(data of Woodin, 1959) 1981
Species Recorded Inches Cover Inches Cover
SPHAGNACEAE
Sphagnum pylaesii Brid.* 25 0.69% 110 3.05%
Sphagnum spp. 80 2.22 95 2.64
ANDREAEACEAE
Andreaea rupestris Hedw. 42 1.16 1D 2.08
GRIMMIACEAE
Racomitrium spp. 55 1.53 2 0.06
Other Bryophytes & Lichens** 113 3.14 95 2.63
LYCOPODIACEAE
Lycopodium annotinum L. var.
pungens Desv.*** 5 0.14 l 0.03
Lycopodium selago L. 7 0.19 10 0.28
PINACEAE
Abies balsamea (L.) Mill. 210 5.83 257 7.14
Picea mariana (Mill.) BSP. 280 7.77 374 10.39
CYPERACEAE
Carex bigelowii Torr. 8 0.25 16 0.44
Scirpus caespitosus L. 290 8.05 268 7.44
BETULACEAE
Alnus viridis (Ait.) Pursh
ssp. crispa (Ait.) Turrill 11 0.31 0 0.00
Betula cordifolia Regel 41 1.14 0 0.00
CARYOPHYLLACEAE
Minuartia groenlandica
(Retz.) Ostenf. 7 0.19 4 0.11
ROSACEAE
Potentilla tridentata Solander
ex. Ait. 67 1.86 7 0.19
EMPETRACEAE
Empetrum nigrum L. 85 2.36 63 1.75

CORNACEAE
Cornus canadensis L. 6 0.17 10 0.27

442 Rhodora [Vol. 86

Table |. Continued

1957
(data of Woodin, 1959) 1981
Species Recorded Inches Cover Inches Cover
ERICACEAE
Chamaedaphne calyculata (L.)

Moench 18 0.50 66 1.84
Kalmia polifolia Wang. 16 0.44 15 0.42
Ledum groenlandicum Oeder 110 3.06 43 1.19
Vaccinium boreale Hall &

Aalders 0 0.00 6 0.17
Vaccinium oxycoccus L. 5 0.14 13 0.36
Vaccinium uliginosum L. 1241 34.47 1107 30.75
DIAPENSIACEAE
Diapensia lapponica L. 78 2.16 107 2.97
PRIMULACEAE
Trientalis borealis Raf. 0 0.00 | 0.03
DEAD LEAF LITTER --- 125 3.47
BARE SOIL 93 2.58 81 2.25
BARE ROCK 706 19.61 649 18.02
Totals 3600 99.96% 3600 99.98%

*Nomenclature of bryophytes follows Ketchledge, 1980.

**Includes 113 inches of lichens missing by printers’ error from Woodin, 1959, but
subsequently supplied to us (personal correspondence, Woodin, 1982).
***Nomenclature of vascular plants follows Kartesz, John T. and Rosemarie
Kartesz, 1980.

attributed either to sampling error or to our recognition of a new
cover category, Dead Leaf Litter, which was visible through dis-
continuities in the living plant canopy. Whatever the cause, the shift
is too slight to indicate major vegetational change.

On the other hand, the 2.36% increase in Sphagnum pylaesii and
1.59% decrease in Bare Rock is in our opinion, a notable shift. Our
observations of the ecology of summit bryophytes, begun in 1949,
repeatedly demonstrate the pioneering nature of Sphagnum pylaesii
spreading over the wet bedrock wherever water seeps out from the
vegetational mat. Lateral expansion of the vascular plant commun-
ity on the open Adirondack summits is mediated by the Sphagnum
mat which engulfs other bryophytes and in time provides a sub-
Strate permitting the establishment of herbaceous and woody
plants.

1984] Ketchledge & Leonard — Mt. Marcy 443

The lateral spread of branches of the two nano-phanaerophytes,
Picea mariana, up 2.6%, and Abies balsamea, up 1.31%, seemed to
be at the expense of their chief canopy competitor, Vaccinium ull-
ginosum, down 3.72%. These three species characteristically occur
in intermixtures where the upward growth of spruce and fir is trun-
cated every two or three years by winter-killing above the protecting
snow field. The basal branches of spruce, less so fir, may slowly
grow outward many feet, whereas the central trunks are ecologic-
ally restricted to a height of one or two feet; some of these dwarfed
trees, we find, are over 200 years old.

ARCTIC VS. NON-ARCTIC SPECIES

Woodin considered only four species found along the transect to
be “non-arctic’: Cornus canadensis, Abies balsamea, Betula cordi-
folia and Carex bigelowii. He suggested future investigators should
note any change in arctic vs. non-arctic percentages and should
evaluate the invasion or loss of species on the transect. Using his
criteria, non-arctic species totalled 7.66% cover in 1957 and 7.85%
in 1981. We do not agree, however, with Dr. Woodin’s inclusion in
the “arctic” category of a number of species which reach their great-
est abundance and vigor in non-arctic environments at lower eleva-
tions but which also survive in lesser numbers in alpine situations.
We place the following additional summit species in that non-arctic
category: Picea mariana, Alnus viridis ssp. crispa, Chamaedaphne
calyculata, Kalmia polifolia, Ledum groenlandicum, Vaccinium
oxycoccus and Trientalis borealis. On this revised basis, the non-
arctic plants totalled 19.88% in 1957 and 22.08% in 1981, an
increase of 3.20% during a quarter century.

The apparent stability over time of the virgin, alpine community
on Mount Marcy stands in sharp contrast to the severe damage
occurring along public hiking trails elsewhere on this peak.

ACKNOWLEDGMENTS

We gratefully acknowledge the assistance of Dixie Sipher in data
analysis and Debbie Berrier and Jean Ketchledge in preparation of
the manuscript. We particularly thank Dr. Woodin for supplying us
with the lichen data inadvertently omitted from his 1959 report.

444 Rhodora [Vol. 86

LITERATURE CITED

KARTESZ, JOHN T. & ROSEMARIE KARTESZ. 1980. A Synonymized Checklist of the
Vascular Flora of the United States, Canada, and Greenland. Volume II, The
Biota of North America. The University of North Carolina Press, Chapel Hill.
498 pp.

KETCHLEDGE, EDWIN H. 1980.Revised Checklist of the Mosses of New York State.
New York State Museum Bulletin No. 440. Albany. 19 pp.

WoopIn, H. E. 1959. Establishment of a Permanent Vegetational Transect
Above Timberline on Mt. Marcy, New York. Ecology 40: 320-322.

E.-H..K;

DEPARTMENT OF ENVIRONMENTAL & FOREST BIOLOGY
SUNY COLLEGE OF ENVIRONMENTAL SCIENCE & FORESTRY
SYRACUSE, NY 13210

R.E.L;

USDA FOREST SERVICE

FORESTRY SCIENCES LABORATORY
PO BOX 640

DURHAM, NH 03824

A REAPPRAISAL OF THE ORCHID GENERA
BROUGHTONIA R. BR., CATTLEYOPSIS LEM.
AND LAELIOPSIS LINDL.

RUBEN P. SAULEDA AND RALPH M.’ ADAMS

ABSTRACT

The orchid genera Broughtonia R. Br., Cattleyopsis Lem. and Laeliopsis Lindl.
are re-established on the basis of observations of natural populations or living mate-
rial from them. The six species that comprise these genera were previously placed in
the genus Broughtonia. A new combination, Cartlevopsis cubensis (Lindl.) Sauleda
and Adams and a new natural hybrid, Broughtonia Xjamaicensis, are reported.

Key Words: Orchids, Broughtonia, Cattleyopsis, Laeliopsis, monograph

The taxonomic status of the orchid genera Broughtonia R. Br.,
Cattleyopsis Lem. and Laeliopsis Lindl. has been the subject of
much controversy. There appears to have been three major, but
often overlapping, taxonomic approaches to these three genera.
One approach retains the three genera as distinct entities (Correll,
1941: Leon, 1946). Another approach incorporates Laeliopsis into
Broughtonia, ultimately recognizing the existence of only two gen-
era, Broughtonia and Cattleyopsis (Rolfe, 1889; Cogniaux, 1910;
Schlechter, 1915; Acufia, 1939; Fowlie, 196la and 1961b). The third
approach incorporates the genera Laeliopsis and Cattleyopsis into
the genus Broughtonia, lumping all species into a single genus
(Dressler, 1966; Liogier, 1969; Adams, 1970 and 1971).

Much of this taxonomic confusion was caused initially by the
floral similarities of some of these species which resulted in gross
misidentifications. Once these problems were resolved and the six
species that comprise the group were established, their differentia-
tion into genera was impeded because of: 1) a failure to examine
natural populations of all species in the group as well as their distri-
butional patterns; 2) a startling misunderstanding of the morphol-
ogy of the nectary; and 3) a disagreement on the number and
significance of pollinia in certain species in the group.

All the botanists who have made taxonomic decisions concerning
this group of orchids neither saw natural populations of each of the
six species nor possessed living material of them. Most decisions
were based on scanty collections of dried herbarium specimens.
Adams (1970) recognized this failure when he stated that taxonomi-

445

446 Rhodora [Vol. 86

cally, “almost every error as we now see it can be attributed to the
extreme difficulty of obtaining reliable factual information from a
small number of inadequate herbarium specimens...”.

The terms “spur”, “nectary-spur”, “sepaline tube” and “external
adnate spur” have all been used as a principal character in the
separation of the genus Broughtonia from Cattleyopsis and Laeli-
opsis. The use of this character in this group requires that its precise
morphological identity be determined and its presence or absence in
each of the species in the alliance be ascertained. In no case has the
morphology of the nectary been carefully studied, presumably
owing to the lack of fresh material available for dissection.

The number and symmetry of the pollinia in species of this
alliance also has been a source of confusion. As Adams (1970)
pointed out, even in good dried herbarium specimens, “the pollinia
are often missing altogether, either having been removed naturally
or stuck to the drying sheets”.

We have examined natural populations or living specimens col-
lected from natural populations of all six species in the alliance as
well as extant herbarium specimens and types. Their distinctive
distributional patterns strongly suggest evolutionary divergence as
a function of isolation. The three species comprising the genus Cat-
tleyopsis are restricted to Cuba, with the exception of C. lindenii
which extends northward into the Bahama Archipelago. The two
species of Broughtonia are endemic to Jamaica, while the mono-
typic genus Laeliopsis occurs on Hispaniola and Mona Island. The
floral similarities among certain members of this alliance are attrib-
utable in part to what we perceive as secondary adaptations to
ornithophily or to melittophily. The. vegetative similarities of the
four species that comprise.the genera Cartleyopsis and Laeliopsis
are attributable in part, to secondary adaptations to the more xeric
habitats in which they oceur. In addition, we have noticed that
Laeliopsis appears to make vegetative phenotypic adjustments to
environmental conditions more readily than does either Cattleyop-
sis or Broughtonia. For example, vegetatively, Laeliopsis resembles
Broughtonia in mesic habitats and Cartleyvopsis in xeric habitats.

The confusion concerning the terms “spur”, “nectary-spur”,
“sepaline tube” and “external adnate spur” was easily resolved when
we made cross-sectional dissections of living flowers of each species.
Unequivocally, none of these structures occurs. Instead, each spe-
cies has a well-defined nectary embedded within the ovary. In all six

1984] Sauleda & Adams — Orchids 447

species of the alliance the nectary lumen is wide at its distal opening,
gradually narrowing proximally, except in the two species of
Broughtonia, in which the proximal end of the nectary lumen swells
to form a nectar-filled chamber. The externally visible proximal
swelling has been confused with a spur or sepaline tube.

Arditti (1969) and Arditti and Fisch (1977) were the only
researchers to recognize the true nature of the nectary in this group
although they failed to note the prominent proximal nectary
chamber present only in Broughtonia. This understanding of the
nature of the nectary, when coupled with their results of pigment
analyses, led Arditti and Fisch (1977) to question the lumping of all
six species into the genus Broughtonia by Dressler (1966).

We agree with Dressler (1966) that the number of pollinia is not
an inviolable generic character. However, differences in distribu-
tional patterns, and in vegetative and floral morphology, combined
with differences in the number and symmetry of pollinia, are com-
pelling reasons to maintain separation of the three genera Brought-
onia, Cattleyopsis and Laeliopsis (Table 1).

KEY TO THE GENERA

1. Leaves coriaceous, margin entire; column short relative to pedicel, wings short
and thick; nectary lumen swollen proximally ...........4 1. Broughtonia

1. Leaves fleshy-rigid, margin erose; column elongate, relative to pedicel, narrowly-

winged; nectary lumen not swollen proximally ........ 0.0 e cece e ences 2
2. Pollinia 8, unequal; column with basal appendages..... 2. Cattlevopsis
2. Pollinia 4, equal; column without basal appendages ...:.. 3. Laeliopsis

1. Broughtonia Robert Brown in Aiton, Hort. Kew, ed. 2, 5: 217.
1813.

Epiphytic plants; rhizomatous; roots velamentous; stems modi-
fied into ellipsoid to ovoid pseudobulbs; leaves coriaceous, margin
entire; inflorescence terminal; ovary pedicellate, with a proximally
swollen nectary; labellum entire, obovate or obovate to orbicular;
column short, blunt, wings near apex short and thick; anther termi-
nal, incumbent, operculate; pollinia 4, equal, with a caudicle; cap-
sule smooth, ellipsoid.

Type: Epidendrum sanguineum Sw.
This genus contains two species, endemic to Jamaica.

Table I. Major characteristics which differentiate the genera Broughtonia, Cattleyvopsis and Laeliopsis.

GENUS LEAVES POLLINIA COLUMN DISTRIBUTION
Length relative Apical Basal Nectary
Margin Rigidity Number Symmetry to pedicel wings appendages lumen
Cattlevopsis — erose ___ fleshy- 8 unequal elongate narrow present distally Bahama
rigid swollen, Islands,
gradually Cuba
narrowing
proximally
Laeliopsis erose _ fleshy- 4 equal elongate narrow absent distally Hispaniola
rigid swollen, and
gradually © Mona Island
narrowing
proximally
Broughtonia entire coriaceous 4 equal short short absent distally Jamaica
and swollen,
thick mesally
narrowing,
proximally
swollen

8bP

e1OpoyYy

98 ‘10A]

1984] Sauleda & Adams — Orchids 449

Figure |. Broughtonia sanguinea (Sw.) R. Br. A. Flower, frontal view. B. La-
bellum, frontal view. C. Petal. D. Dorsal sepal. E. Lateral sepal. F. Column,
dorsal view. G. Column, ventral view. H. Column and ovary, lateral view and
serial cross sections (a-f) showing nectary lumen.

KEY TO THE SPECIES

1. Basal margins of labellum not involute, not enclosing column, not forming a tube;

center of labellum glabrous ...... 00.0000 cece eee rece ene |. B. sanguinea
1. Basal margins of labellum involute, enclosing column, forming a tube; center of
labellum with yellow villous lamellae ........ 000s eee eee 2. B. negrilensis

1. Broughtonia sanguinea (Sw.) R. Br. in Aiton, Hort. Kew ed. 2,
5: 217. 1813. (Figure 1)

Epidendum sanguineum Sw., Prodr. Veg. Ind. Occ. 124. 1788.
Dendrobium sanguineum (Sw.) Sw., Nov. Act. Ups. 6: 82. 1799. Type: No holo-
type indicated. A search of the major herbaria known to contain collections of

450 Rhodora [Vol. 86

Swartz failed to uncover any specimens that can be ascertained to have been
collected or examined by Swartz. In the absence of a holotype, a lectotype is
here chosen from material cited by Swartz in the protologue. LECTOTYPE
designated here: Plate 121, figure 2, in H. Sloane, A Voyage to the Islands
Madera, Barbados, Nieves, S. Christophers and Jamaica, Vol. 1, 1707.

Broughtonia coccinea Hook., Bot. Mag. 63, t. 3536. 1836. Based on Broughtonia
sanguinea (Sw.) R. Br.

Plant epiphytic, rhizomatous, to 72 cm tall; roots numerous,
thick, velamentous; rhizome short, stout, creeping or ascending,
enclosed by imbricating scarious sheaths; stem modified into pseu-
dobulb, erect or ascending, clustered, ellipsoid to ovoid, flattened,
to 5 cm long, 4 cm wide, 3 cm thick, basally enclosed by scarious
sheaths, to 4-leaved at apex; leaves coriaceous, elliptic to oblong or
ligulate, subacute, margin entire, to 18 cm long, 3 cm wide; inflores-
cence to 67 cm tall, peduncle slender, erect, distantly several-
sheathed, simple or paniculate raceme, to 18 flowers; floral bracts
ovate, acute, to 0.4 cm long, 0.2 cm wide; ovary slender, to 4 cm
long, sepals and petals pink, white, yellow or usually reddish-
lavender to red; sepals elliptic to lanceolate, acute, to 2.7 cm long,
0.8 cm wide; petals oblong to orbicular, abruptly acute, to 2.4 cm
long, 1.6 cm wide; labellum pink, white, yellow or usually reddish-
lavender to red, with a central basal yellow spot and 8-12 radiating
red veins, to 3 cm long, 2.7 cm wide, broadly obovate to orbicular,
center of labellum glabrous, basal margins not involute, not com-
pletely enclosing column, margin erose, shallowly emarginate;
column white, occasionally tinged with red, to 0.8 cm long, 0.5 cm
wide, anther white; capsule pendent, to 2.4 cm long, 0.8 cm thick.

REPRESENTATIVE SPECIMENS: Jamaica: Parish of St. Elizabeth, Pepper, 13 Mar
1931, Miller 1349 (US); coastal region E of Montego Bay, 28 Mar 1920, Maxon &
Killip 1626 (AMES, NY); Great Goat Island, 19 Apr 1906, Harris 9209 (NY); Lover's
Leap, Santa Cruz Mts., 4 Sep 1907, Britton 1148 (NY).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from March to June.

Vegetatively, this species may be confused with Broughtonia
negrilensis Fowlie. Florally they differ significantly in color and in
the position of the basal margins of the labellum. Broughtonia
sanguinea is usually reddish-lavender to red, while B. negrilensis is
pink to lavender. In B. negrilensis the basal margins of the labellum
are involute, enclosing the column to form a tube and the center of

1984] Sauleda & Adams — Orchids 451

the labellum has yellow villous lamellae, while in B. sanguinea the
basal margins are not involute, do not enclose the column and the
center of the labellum is glabrous.

A rare autogamous dwarf form occurs with flowers that remain
partially closed. Hooker illustrated this dwarf form as Broughtonia
sanguinea (Bot. Mag. 58, t. 3076, 1831) and later, after seeing the
more common fully-opened form, mistakenly named it B. coccinea
Hook. (Bot. Mag. 63, t. 3536, 1836). Inexplicably, Hooker based B.
coccinea on B. sanguinea (Sw.) R. Br. making B. coccinea a super-
fluous name at the time of publication.

Although this species is endemic to Jamaica, the name has been
misapplied to Cartlevopsis oritgiesiana (Reichb. f.) Cogn. (Grise-
bach, 1866; Cogniaux, 1910; Acuna, 1939), a species endemic to
Cuba.

In southwestern Jamaica, on Negril Ridge, where Broughtonia
sanguinea and B. negrilensis are sympatric, a natural hybrid occurs
which is intermediate between the two parental species. Introgres-
sive hybridization appears to occur between this natural hybrid and
B. sanguinea, accounting for the high degree of variability in shape
and color of the floral parts of B. sanguinea in the areas of sympa-
try. A series of experimental crosses verified the introgression.

2. Broughtonia negrilensis Fowlic, Orch. Digest 25: 417-418. 1961.
(Figure 2) Type: JAMAICA, Negril Ridge, 2 mi SW of Old
Hope, elev. 250 ft., epiphytic on Ceiba, collected Nov. 1959,
flowered in cultivation at the Los Angeles State and County
Arboretum at Arcadia, California, Fowlie s.n. (HOLOTYPE:

LASCA).

Broughtonia domingens!s auct. non (Lindl.) Rolfe: Fawcett & Rendle, Fl. Jam. 1:
24, 1910; Cogn. in Urban, Symb. Antill. 6: 543, 1910.

Laeliopsis domingensis auct. non (Lind!.) Lind!.: Grisebach, Fl. B. W. I., 621,
1864.

Plant epiphytic, rhizomatous, to 75 cm tall; roots numerous, vel-
amentous; rhizome short, stout, creeping or ascending, enclosed by
imbricating scarious sheaths; stem modified into pseudobulb, erect
or ascending, clustered, ellipsoid to ovoid, flattened, to 6.0 cm long,
2.8 cm wide, 1.8 cm thick, basally enclosed by scarious sheaths, to
3-leaved at apex; leaves coriaceous, elliptic to oblong, subacute,
margin entire, to 12 cm long, 2.4 cm wide; inflorescence to 69 cm

452 Rhodora [Vol. 86

Figure 2. Broughtonia negrilensis Fowlie. A. Flower, frontal view. B. Label-
lum, frontal view. C. Petal. D. Dorsal sepal. E. Lateral sepal. F. Column, dorsal
view. G. Column, ventral view. H. Column and ovary, lateral view and serial cross
sections (a-f) showing nectary lumen.

tall, peduncle slender, erect, distantly several-sheathed, simple or
rarely paniculate raceme, to 12 flowers; floral bracts ovate, acute, to
0.6 cm long, 0.3 cm wide; ovary slender, to 3.0 cm long; sepals pink
to lavender, linear-oblong, subacute, to 2.8 cm long, 0.8 cm wide;
petals pink to lavender, broadly-oblanceolate to obovate, acute, to
3.0 cm long, 1.3 cm wide; labellum basally yellow, pink to lavender
towards apex, with purple lines from center radiating to near mar-

1984] Sauleda & Adams — Orchids 453

gin, to 3.6 cm long, 2.9 cm wide, obovate, basal margins involute,
enclosing column, forming a tube, center of labellum with yellow
villous lamellae, margin crenate-dentate, undulate, emarginate;
column white, occasionally tinged with pink, to 0.9 cm long, 0.5 cm
wide, anther white; capsule pendent, to 2.8 cm long, 1.0 cm thick.

SPECIMEN EXAMINED: JAMAICA: Negril Ridge, Jun 1967, Gauntlerts.n., flowered in
cultivation (USF).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from October to December.

Broughtonia negrilensis is similar vegetatively to B. sanguinea R.
Br. but florally can be easily distinguished by floral color and by
shape and ornamentation of the labellum.

This species was incorrectly referred to Laeliopsis domingensis
(Lindl.) Lindl. by Grisebach (1864) and to Broughtonia domingen-
sis (Lind].) Rolfe by Fawcett & Rendle (1910) and Cogniaux (1910).
Laeliopsis domingensis is a different species endemic to Hispaniola
while Broughtonia domingensis is a synonym of it.

NATURAL HYBRID

Broughtonia Xjamaicensis Sauleda & Adams, Aybr. nov. (Figure 3)
Broughtonia sanguinea (Sw.) R. Br. X Broughtonia negrilensis
Fowlie

Hybrida floraliter intermedia. Plantae epiphyticae, rhizomato-
sae; pseudobulbi ellipsoidei usque ovoidei; folia coriacea, elliptica
usque oblonga; inflorescentia erecta, simplex vel paniculata; flores
rosel usque rubeo-lavendull.

Plants epiphytic, rhizomatous, to 68 cm tall; roots numerous,
velamentous; rhizome short, stout, creeping or ascending, enclosed
by imbricating scarious sheaths; stem modified into pseudobulb,
erect or ascending, clustered, ellipsoid to ovoid, flattened, to 4.8 cm
long, 2.7 cm wide, 1.9 cm thick, basally enclosed by scarious
sheaths, 2-leaved at apex; leaves coriaceous, elliptic to oblong, sub-
acute, margin entire, to 9.6 cm long, 2.3 cm wide; inflorescence to
63 cm tall, peduncle slender, erect, simple or paniculate raceme, to 8
flowers; floral bracts ovate, acute, to 0.5 cm long, 0.3 cm wide;
ovary slender, with proximally swollen nectary, to 3.2 cm long;
sepals and petals pink to reddish-lavender, elliptic, acute, to 2.8 cm

454 Rhodora [Vol. 86

Figure 3. Broughtonia Xjamaicensis Sauleda & Adams. Flower, frontal view,
x 2.

long, 0.6 cm wide; petals obovate to oblong, acute, to 2.8 cm long,
1.3 cm wide; labellum pink to reddish-lavender, basally usually
yellow, with purple lines from center radiating to near margin, to
3.4 cm long, 2.7 cm wide, obovate, basal margins slightly involute
but not completely enclosing column, center of labellum occasion-
ally with a few yellow villous lamellae, margin crenate-dentate,
undulate, emarginate; column white, to 0.9 cm long, 0.5 cm wide,
anther white.

Type: JAMAICA: Negril Hill, Jun 1967, Gauntleti s.n. (HOLoTyPE:
NY). Flowered in cultivation September 1968,

1984] Sauleda & Adams — Orchids 455

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from August to October.

This natural hybrid has been referred to in the literature (Adams,
1970 and 1971; Fowlie, 196la and 1961b) on several occasions but
has not been named.

Florally, the hybrid is intermediate between the parental species
and occurs commonly where they are sympatric. The labellum has
the characteristic yellow villous lamellae of Broughtonia negrilensis
while the basal margin of the labellum is slightly involute but does
not enclose the column, a characteristic of B. sanguinea. This
hybrid is named for the type location.

2. Cattleyopsis Lemaire, Jard. Fleur. 4, Misc. p. 59. 1853.

Epiphytic plants; rhizomatous; roots velamentous; stems modi-
fied into cylindric, ovoid or pyriform pseudobulbs; leaves fleshy-
rigid, margin erose; inflorescence terminal; ovary pedicellate;
labellum entire, oblong or obovate to orbicular; column elongate,
slender, with two auricle-like appendages near the base, narrowly
winged towards apex; anther terminal, incumbent, operculate; pol-
linia 8, unequal, with a caudicle; capsule smooth, ellipsoid.

Type: Cattleyopsis delicatula Lemaire
This genus contains three species endemic to xeric regions of
Cuba and the Bahama Archipelago.

KEY TO THE SPECIES

1. Basal margin of labellum enclosing column, forming a tube; center of labellum
with yellow villous lamellae; margin of labellum incised to crenate-dentate or

CRENALE a iscsi o Wire ee oe lane ate ett Laisa sees ihe Hae a ol eet Gee 2

1. Basal margin of labellum not enclosing column; center of labellum glabrous:
margin of labellum entire ........ 2.0.0 cece eee eee eee 3. CG. ortgiesiana

2. Labellum oblong; leaves obtuse... 0... ccc cece eee eee eee 1. C. cubensis
2. Labellum broadly-obovate to orbicular; leaves acute ......... 2. C. lindenii

1. Cattleyopsis cubensis (Lindl.) Sauleda & Adams, comb. nov.
(Figure 4)

Epidendrum cubense Lind|., Bot. Reg. 29, Misc. p. 17. 1843.
Laeliopsis cubense (Lindl.) Lind]. ex Cogn. in Urban, Symb. Antill. 6: 543. 1910,
pro syn.

456 Rhodora [Vol. 86

Broughtonia cubensis (Lind|.) Cogn. in Urban, Symb. Antill. 6; 542. 1910. Type:
CUBA, imported to England by Messrs. Loddiges (HOLOTYPE: K-L, photograph
seen).

Plant epiphytic, rhizomatous, to 33 cm tall; roots few, thick,
velamentous; rhizome short, stout, creeping or ascending, enclosed
by imbricating scarious sheaths; stem modified into pseudobulb,
erect or ascending, clustered, ovoid to pyriform, to 3 cm long, | cm
wide, enclosed by scarious sheaths, to 2-leaved at apex; leaves

\\

\ 4 \
\\
AN

|

NN

Figure 4. Cattleyopsis cubensis (Lindl.) Sauleda & Adams. A. Flower, frontal
view. B. Labellum, frontal view. C. Petal. D. Dorsal sepal. E. Lateral sepal.
F. Column, dorsal view. G. Column, ventral view. H. Column and ovary, lateral
view and serial cross sections (a-d) showing nectary lumen.

1984] Sauleda & Adams — Orchids 457

fleshy-rigid, oblong, obtuse, margin minutely erose, to 8 cm long, |
cm wide; inflorescence to 30 cm tall, peduncle slender, erect, dis-
tantly several-sheathed, simple raceme, to 8 flowers; floral bract
ovate, acute, to 0.2 cm long, 0.1 cm wide; ovary slender, to 1.5 cm
long; sepals pink to yellowish-pink, linear oblong to ligulate,
obtuse, to 1.8 cm long, 0.5 cm wide; petals pink to yellowish-pink,
oblong, obtuse, to 1.8 cm long, 0.7 cm wide; labellum pink to
yellowish-pink with reddish-pink radiating lines near base extend-
ing to margin, to 2.0 cm long, 1.1 cm wide, oblong, basal margins
involute enclosing column, forming a tube, center of labellum with
yellow villous lamellae, margin crenate, shallowly emarginate;
column pink, to 0.7 cm long, 0.3 cm wide, anther pink to reddish-
pink; capsule pendent, to 2.5 cm long, 1.2 cm thick.

SPECIMEN EXAMINED: CUBA: Prov. de Pinar del Rio, Peninsula de Guanahacabibes,
2-3 mi E of Caleta Piojo, near Sawmill La Posa de Juan Claro, Jul 1959, Osment s.n.
(USF).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from December to March.

Cattleyopsis cubensis was first described by Lindley (1843) as
Epidendrum cubense Lindl. and later (Lindley, 1853) referred it to
the genus Laeliopsis Lindl. without formal transfer. Although Lind-
ley examined living material, he failed to recognize that the anthers
contained eight unequal pollinia. Cogniaux (1910) also failed to
note the number of pollinia and transferred this species to the genus
Broughtonia R. Brown. Correll (1941 and pers. comm.), after exa-
mining a photograph of the type, concluded that the exact taxo-
nomic status of this species was not clear and in the absence of
living material suggested that it should be retained in the genus
Laeliopsis, to which Lindley (1853) had originally referred it.
Dressler (1966) examined the type and considered it to be conspe-
cific with Epidendrum chinense Lindl. and placed it under synon-
ymy with Barkeria chinensis (Lind].) Thien ex Dressler.

We have examined three living plants of this species which were
collected by William Osment in Prov. de Pinar del Rio, Cuba, July
1959. Additionally, we have examined the progeny of selfing crosses
from the original material. In every case, the specimens have eight
unequal pollinia, elongate and narrowly-winged columns with basal
appendages, no proximally swollen nectaries and erose leaf mar-

458 Rhodora [Vol. 86

gins. Taken together, these characters unequivocally place these
specimens in the genus Cattleyopsis Lemaire.

Furthermore, an examination of a photograph of the holotype of
Barkeria chinensis convinces us that Cattleyopsis cubensis is not
conspecific with it.

2. Cattleyopsis lindenii (Lindl.) Cogn. in Urban, Symb. Antill. 6:
544. 1910. (Figure 5)

Laelia lindenii Lindl., Orch. Lind. 10. 1846.

Bletia lindenii (Lind|.) Reichb. f., Walp. Ann. Bot. 6: 431. 1862.

Laeliopsis lindenii (Lind|.) Lind]. ex Cogn. in Urban, Symb. Antill. 6: 545. 1910,
pro syn.

Broughtonia lindenii (Lind|.) Dressler, Taxon 15: 241. 1966. Type: cuBA, Linden
1805 (HOLotTyPE: K-L, photograph seen).

Cattleyopsis delicatula Lem., Jard. Fleur. 4, Misc. p. 59. 1853. Type: No holotype
indicated and a search of the major herbaria of Europe failed to uncover any
specimens that can be ascertained to have been collected or examined by
Lemaire. Lectotype designated here: Figures 1-3 in Lemaire’s Jard. Fleur. 4,
p. 60, 1853.

Cattleyopsis northropiorum Cogn. in Urban, Symb. Antill. 6: 545. 1910. TypE:
No holotype indicated. Lectotype here designated from syntypes: Bahama
Islands, Andros, Northrop & Northrop 437 (AMES).

Cattleyopsis guanensis Acuna, Cat. Descr. Orquid. Cub., Estac. Exper. Agron.
Bol. Tec. 60: 109-110, 1939, nom illeg., lacking Latin diagnosis. TyPE: CUBA,
Pinar del Rio, Guane, Fors 4837 (HOLOTYPE: Present location of specimen in
Cuba not known, photograph seen at AMES).

Cattleya domingensis auct. non Lindl.: Britton & Millspaugh, Bahama FI. 95,
1920; Richard, in Sagra’s Hist. Fisica, Polit. Nat. Isl. Cuba, pt. 2, 11: 243, 1850.

Laeliopsis domingensis auct. non (Lindl.) Lindl.: Britton & Millspaugh, Bahama
Fl. 95, 1920; Northrop, Mem. Torr. Bot. Cl. 12: 30, 1902.

Broughtonia domingensis auct. non (Lind!.) Rolfe: Cogn. in Urban. Symb.
Antill. 6: 543, 1910; Britton & Millspaugh, Bahama FI. 95, 1920.

Broughtonia lilacina auct. non Henfr.: Northrop, Mem. Torr. Bot. Cl. 12: 30,
1902.

Plant epiphytic, rhizomatous, to 68 cm tall: roots numerous,
thick, velamentous; rhizome short, stout, creeping or ascending,
enclosed by imbricating scarious sheaths: stem modified into
pseudobulb, erect or ascending, clustered, cylindric to narrowly
ovoid, to 8 cm long, 2 cm wide, enclosed by scarious sheaths, to
3-leaved at apex: leaves fleshy-rigid, oblong to linear-oblong, acute
to apiculate, subcanaliculate, margin erose, to 11 cm long, 2.5 cm
wide; inflorescence to 60 cm tall, peduncle slender, erect, distantly

1984] Sauleda & Adams — Orchids 459

Figure 5. Cattlevopsis lindenii (Lind|.) Cogn. A. Inflorescence. B. Plant. C. La-
bellum, frontal view. D. Petal. E. Dorsal sepal. F. Lateral sepal. G. Column,
dorsal view. H. Column, ventral view. I. Column and ovary, lateral view and serial
cross sections (a-d) showing nectary lumen.

460 Rhodora [Vol. 86

several-sheathed, simple or paniculate raceme, to 12 flowers: floral
bracts ovate, acute, to 0.5 cm long, 0.3 cm wide: ovary slender, to
2.5 cm long; sepals pink to lavender, linear-oblanceolate, acute, to
2.8 cm long, 0.8 cm wide; petals pink to lavender. narrowly obovate
to oblanceolate, subobtuse to acute, to 2.4 cm long, 1.0 cm wide;
labellum pink to lavender with purple lines radiating from center to
near margin, to 3.5 cm long, 3 cm wide, broadly-obovate to
orbicular, basal margins involute enclosing column, forming a tube,
center of labellum with yellow villous lamellae, Margin incised to
crenate-dentate, undulate, deeply emarginate; column white, to 1.2
cm long, 0.3 cm wide, arcuate, anther greenish-white: capsule
pendent, to 3.0 cm long, 1.5 cm thick.

REPRESENTATIVE SPECIMENS: Bahama Islands: GREAT ABACO, near North Harbour,
20 Jul 1904, Barbour 773 (AMES); Cherokee Sound, 29 Dec 1904, Brace 1942 (F,
NY): Abaco Heights road, 11 Apr 1979, Sauleda & Correll 2262 (FAU); Guiana
Schooner Bay, 12 Apr 1979, Sauleda & Correll 2304 (FAU): Hole-in-the-Wall, 28
Jun 1980, Sauleda & Sauleda 3667 (FA U). LITTLE ABACO, near Fox Cay settlement, 4
Aug 1979, Sauleda, Adams & Adams 2864 (FAU). ANDROS ISLAND, mangrove
swamp, SE edge of Lake Forsythe, 6 Nov 1976, Sauleda /530(FAU); 8 mi S of Fresh
Creek, 5 Feb 1977, Sauleda 1834 (FAU);: Deep Creek, 2 May 1979, Sauleda &
Correll 2465 (USF). BIMINI GRoUP, South Bimini, 15 Apr 1904, Millspaugh 2397 (F,
NY). ELEUTHERA, N. of James Cistern, 17 May 1975, Correll & Hill 45248 (FTG).
GREAT EXUMA, 6 mi N of George Town, 22 Jan 1979, Sauleda, Correll & Correll 2200
(FAU). GRAND BAHAMA, Garden Cay, West End, 16 Apr-8 May 1905, Brace 3660 (F,
NY). LONG ISLAND, Gordons, trail to lighthouse, 7 Jul 1974, Hill 2426 (FTG). MAYA-
GUANA, Dykes s.n. (AMES). Cuba: Prov. de Oriente, southern Baracoa region, 17
Jul-4 Aug 1924, Leon 11761 (NY): Prov. de Oriente, Guantanamo Bay, 17-30 Mar
1909, Britton 2230 (NY); Prov. de Pinar del Rio, Pan de Azucar, 5 Feb 1956, Morton
9834 (US); Isle of Pines, top of Sierra de los Caballos, 12 May 1910, Jennings 230
(NY).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from May to September.

Cartleyopsis lindenii is a highly variable species distributed only
in the Bahama Islands and Cuba. The Cuban populations are char-
acteristically larger florally and vegetatively, and the flowers open
more fully than those in the Bahamas. While these characteristics
appear to predominate in the two areas, occasional variants can be
found.

Cattleyopsis lindenii was first described by Lindley (1846) as Lae-
lia lindenii, based on a Cuban specimen. Cattleyopsis northropi-
orum was first described by Cogniaux (1910) based mainly on

1984] Sauleda & Adams — Orchids 461

collections from the Bahama Islands. A comparison of the types
cited by these authors convinced us that C. northropiorum repres-
ents the smaller-flowered and more closed form of C. /indenti and
should be considered a synonym of it.

Cattleyopsis guanensis was first described by Acuna (1939) based
on a specimen collected in Cuba. We compared a photograph of the
type of C. lindenii with a photograph of the type of C. guanensis.
Additionally, we examined a color photograph of the type plant of
C. guanensis in flower, taken by Acuna. We are convinced that C.
guanensis is in reality the small-flowered C. /indenii, but a pecul-
iarly dark lavender form, and should therefore be considered a
synonym of it.

Cattleyopsis delicatula was first described by Lemaire (1853) as
the type of the genus. An examination of an illustration published
with the protologue (lectotype) leads us to the conclusion that it is
the Cuban form of C. /indenii and should be considered a synonym
of it.

The floral similarities between Cart/eyopsis lindenii and Laeliop-
sis domingensis (Lind].) Lindl. has lead to a great deal of confusion
in the literature. These two species can be easily distinguished by an
examination of the column. C. /indenii has eight unequal pollinia
and two basal appendages on the column, while in L. domingensis,
the column bears four equal pollina and lacks basal appendages.

Specimens of Cartleyopsis lindenii were incorrectly referred to
Laeliopsis domingensis by Britton & Millspaugh (1920) and to
Broughtonia domingensis (Lindl.) Rolfe by Cogniaux (1910). Laeli-
opsis domingensis is a different species endemic to Hispaniola while
B. domingensis is a synonym of it. In addition, Northrop (1902)
referred specimens of C. lindenii to B. lilacina Henfr. which is also a
synonym of L. domingensis.

3. Cattleyopsis ortgiesiana (Reichb. f.) Cogn. in Urban, Symb.
Antill. 6: 546. 1910. (Figure 6)

Bletia ortgiesiana Reichb. f.. Hamb. Gartenz. 420. 1860.

Broughtonia ortgiesiana (Reichb. f.) Dressler, Taxon 15: 241. 1966. Ty Pe: Local-
ity not known. Cultivated in Botanical Garden in Zurich, by Mr. E. Ortiges.
(HoLotypr: W).

Broughtonia sanguinea auct. non (Sw.) R. Br.: Cogn. in Urban, Symb. Antill. 6:
542, 1910; Griseb., Cat. Pl. Cub. 263, 1866.

462 Rhodora [Vol. 86

OOSe-

Figure 6. Cartlevopsis ortgiesiana (Reichb. f.) Cogn. A. Flower, frontal view,
B. Labellum, frontal view. C. Petal. D. Dorsal sepal. E. Lateral sepal. F. Col-
umn, dorsal view. G. Column, ventral view. H. Column and ovary, lateral view
and serial cross sections (a—d) showing nectary lumen.

Plant epiphytic, rhizomatous, to 58 cm tall: roots thick, vela-
mentous; rhizome short, stout, creeping or ascending, enclosed by
imbricating scarious sheaths; stem modified into pseudobulbs, erect
or ascending, clustered, cylindric to narrowly pyriform, to 5 cm
long, 1.4 cm wide, enclosed by scarious sheaths, to 2-leaved at apex;
leaves fleshy-rigid, oblong to linear-oblong, obtuse, margin erose,

1984] Sauleda & Adams — Orchids 463

to 18 cm long, 2.4 cm wide; inflorescence to 53 cm tall, peduncle
slender, erect, distantly several-sheathed, simple or paniculate
raceme, to 20 flowers; floral bracts narrowly ovate, acute, to 0.4cm
long, 0.2 cm wide; ovary slender, to 2 cm long; sepals reddish-
purple to pink, oblong, acute, to 1.7 cm long, 0.7 cm wide; petals
reddish-purple to pink, obovate, obtuse, to 1.8 cm long, 0.9 cm
wide; labellum reddish-purple, with a central basal white spot and
with 8-12 radiating red veins, to 1.8 cm long, 1.8 cm wide, obovate
to orbicular, basal margins involute, but not completely enclosing
column, margin entire, undulate, emarginate; column reddish-
purple to pink, to 0.7 cm long, 0.3 cm wide, anther reddish-purple
to pink; capsule pendent, to 2.2 cm long, 1.1 cm thick.

REPRESENTATIVE SPECIMENS: CUBA: Prov. de Oriente, Playa de Punto Padre, 24 Jul
1959, Lopez 10 (US); Prov. de Camaguey, near Camaguey. 2 7 Apr 1912. Britton,
Britton & Cowell 13143 (AMES, NY. US); Prov. de Pinar del Rio. vicinity of
Coloma, 2 Mar 1911, Britton & Cowell 9692 (NY): Prov. de Camaguey, 3 km E of
Florida, 31 Mar 1950, Cutler 12204 (AMES), Isle of Pines, Loma la Daguilla, 3 Mar
1916, Britton, Britton & Wilson 15165 (AMES, NY. US).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from December to March.

This species has been referred to Broughtonia sanguinea (SW.)
R. Br. by Cogniaux (1910), Grisebach (1866) and Acufia (1939).
These misidentifications account for the incorrect reports of B. san-
guinea occurring in Cuba. Broughtonia sanguinea is a different
species endemic to Jamaica.

Florally, Broughtonia sanguinea 1s similar to Cattleyopsis ortgie-
siana but is distinctive in column morphology. The column of C.
ortgiesiana bears eight unequal pollinia and has two basal appen-
dages, while the column of B. sanguinea bears four equal pollinia
and lacks basal appendages.

3. Laeliopsis Lindley in Paxton, Flow. Gard. 3: 155. 1853.

Epiphytic plants; rhizomatous; roots velamentous; stems modi-
fied into ovoid pseudobulbs; leaves fleshy-rigid, margin erose; inflo-
rescence terminal; ovary pedicellate; labellum entire, narrowly
obovate; column elongate, clavate, narrowly winged towards apex;
anther terminal, incumbent, operculate; pollinia 4, equal, with a
caudicle; capsule smooth, ellipsoid.

464 Rhodora [Vol. 86

Type: Cattleya domingensis Lind.
This is a monotypic genus occurring on Hispaniola and Mona
Island.

1. Laeliopsis domingensis (Lind].) Lindl., Paxt. Flow. Gard. III:
156, t. 105. 1853. (Figure 7)

Cattleya domingensis Lindl., Gen. & Sp. Orch. 118. 1831.

Bletia domingensis (Lindl.) Reichb.f., Walp. Ann. VI: 432. 1862.

Broughtonia domingensis (Lindl.) Rolfe, Gard. Chron. Ser. 3, V: 491. 1889.
TYPE: SANTO DOMINGO, MacKenzie s.n. (HOLOTYPE: K—L, photograph seen).

Broughtonia lilacina Henfr. in Moore & Ayres, Gard. Mag. Bot. 3: 201. 1851.
Type: No holotype designated. Lectotype designated here: Plate in Moore &
Ayres, Gard. Mag. Bot. 3, 1851, drawn from a plant collected on Hispaniola.

Broughtonia violacea Henfr. in Moore & Ayres, Gard. Mag. Bot. 3: 201. 1851.
Published in synonomy.

Plant epiphytic, rhizomatous, to 110 cm tall; roots numerous,
velamentous; rhizome short, stout, creeping or ascending, enclosed
by imbricating scarious sheaths; stem modified into pseudobulb,
erect or ascending, clustered, ovoid, to 6 cm long, 5 cm wide,
enclosed by scarious sheaths, to 3-leaved at apex; leaves fleshy-
rigid, oblong, obtuse to subacute, subcanaliculate, margin erose, to
18 cm long, 3 cm wide; inflorescence to 104 cm tall, peduncle
slender, erect, distantly several-sheathed, simple or paniculate
raceme, to 15 flowers; floral bracts ovate, acute, to 0.3 cm long, 0.3
cm wide; ovary slender, to 2.5 cm long; sepals pink to lavender,
linear-oblong, acute, to 3.3 cm long, 0.6 cm wide; petals pink to
lavender, broadly-oblanceolate to obovate, acute, to 3 cm long, 1.2
cm wide; labellum pink to lavender, to 4 cm long, 2.7 cm wide,
narrowly obovate, basal margins involute, enclosing column, form-
ing a tube, center of labellum with yellow villous lamellae terminat-
ing near apex, frontal margin with purple veins terminating near
apex, margin minutely incised to crenate-dentate, undulate, shal-
lowly emarginate; column pink, to 1.6 cm long, 0.5 cm wide,
slender, anther pink to reddish-purple; capsule pendent, to 5.2 cm
long, 2.4 cm thick.

REPRESENTATIVE SPECIMENS: Hispaniola: DOMINICAN REPUBLIC, Prov. La Romana,
NE of La Romana, 19 Aug 1982, Sauleda, Sauleda, Ragan & Dod 7450 (USF);
Prov. Peravia, Bani, 19 Feb 1966, Lavastre 2070 (NY); Prov. Altagarcia, SE of Playa
El Macao, 13 May 1980, Mejia & Zanoni 6263 (SDM): Prov. Monte Cristi, NW of
Villa Sinda, 21 May 1980, Mejia & Zanoni 6398 (SDM). HAITI, Ile de la Tortue,

1984] Sauleda & Adams — Orchids 465

Figure 7. Laeliopsis domingensis (Lind!.) Lindl. A. Flower, frontal view. B. La-
bellum. frontal view. C. Petal. D. Dorsal sepal. E. Lateral sepal. F. Column,
dorsal view. G. Column, ventral view. H. Column and ovary, lateral view and
serial cross sections (a-d) showing nectary lumen.

vicinity of La Vallee. 3-10 May 1929, Leonard & Leonard 15612 (NY, US); Dept.
Du Nord. NW of Pedro Santana (D. R.), 27 Aug 1982, Sauleda, Sauleda, Ragan &
Dod 7617A (USF).

FLOWERING PERIOD: Flowers sporadically throughout the year,
mainly from April to June.

This species has been reported to occur in the Bahama Islands
and Cuba based on misidentifications of Cartlevopsis lindenii

466 Rhodora [Vol. 86

(Lindl.) Cogn., by Richard (1850), Britton & Millspaugh (1920) and
Cogniaux (1910). Additionally, this species has been reported from
Jamaica, based on misidentifications of Broughtonia negrilensis
Fowlie, by Fawcett & Rendle (1910), Cogniaux (1910) and Grise-
bach (1864).

Laeliopsis domingensis is florally similar to C attleyopsis lindenii
and Broughtonia negrilensis. These three species can be distin-
guished easily by examining column morphology. Cartleyopsis lin-
denii has eight unequal pollinia and two basal appendages on the
column, while B. negrilensis and L. domingensis both have four
equal pollina and lack basal appendages. Broughtonia negrilensis is
distinguished from L. domingensis by having a short column rela-
tive to the pedicel, thick wings on the column anda nectary which is
swollen proximally. Laeliopsis domingensis has an elongate column
relative to the pedicel, narrow column wings and a nectary not
swollen proximally.

ACKNOWLEDGMENTS

We thank the Directors, Curators and staff members of the her-
baria cited throughout the text for their cooperation and help. Spe-
cial thanks are extended to Drs. Peter Taylor (K) and Robert Read
(US). We acknowledge the generous field assistance given by
Donald and Tudy Dod of the Dominican Republic, Gladys and
Kenneth Fehling of the Bahama Islands and Marv Ragan of
Orange Park, Florida. Our thanks are also extended to William
Osment of Hollywood, Florida for his donations of plant material,
to Dr. Helen B. Correll for preparing the Latin description, and to
Rebeka Sauleda for some of the illustrations. This research was
supported in part by donations from the Tropical Orchid Society of
Palm Beach, Florida.

LITERATURE CITED

ACUSA GALE, J. 1939. Catalogo Descriptivo de las Orquideas Cubanas. Boletin
no. 60, pp. | 221, Estacion Experimental Agronomica, Santiago de las Vegas,
Cuba.

Apams. C.D. 1970. Broughtonia - A Brief Review. The Florida Orchidist 13:
8 11.

1971. Broughtonia Again. The Florida Orchidist 14: 101 10S.
Arpitti, J. 1969. Floral Anthocyanins in Species and Hybrids of Broughtonia,
Brassavola and Cattleyvopsis (Orchidaceae). Amer. J. Bot. 56: 59 68.

1984] Sauleda & Adams — Orchids 467

AND M. H. Fiscu. 1977. Anthocyanins of the Orchidaceae: Distribu-
tion, Heredity, Functions, Synthesis and Localization. /n: Orchid Biology.
Reviews and Perspectives I, ed. J. Arditti. Cornell University Press, Ithaca,
New York.

Britton, N. L. AND C. F. MILLSPAUGH. 1920. The Bahama Flora. Published by
the authors. Reprinted 1962, Hafner Publishing Co., New York.

CoGNiaux, A. 1910. /n: Urban, I.. Symbolae Antillanae (Orchidaceae), 6:
293-696. Reprinted 1964, A. Asher & Co., Amsterdam.

CorRELL, D.S. 1941. Notes concerning Some West Indian Orchids I. The Genus
Broughtonia and Some of its Allies. Bot. Mus. Leaflets, Harvard Univ. 10:
41-S8.

DresscerR, R. L. 1966. Nomenclatural Notes on the Orchidaceae III. Taxon 15:
241-243.

Fawcett, W. AND A. B. RENDLE. 1910. Flora of Jamaica. Vol. 1, Orchidaceae.
London, England.

Fow ik, J. A. 196la. Ecological Notes: Natural Hybridization in the Genus
Broughtonia. Amer. Orchid Soc. Bull. 30: 707-710.

1961b. Obscure Species. The Orchid Digest 25: 416-418.

GrRIseBACH, A. H. R. 1864. Flora of the British West Indian Islands. Lovell
Reeve & Co., London, England.

1866. Catalogus Plantarum Cubensium, Lipsiae.

Hooker, W. J. 1831. Bot. Mag. (Curtis) 58, t. 3076.

1836. Bot. Mag. (Curtis) 63, t. 3536.

Lemaire, C. 1853. Jardin Fleuriste, Vol. 4, Misc. p. 59.

Leon, Her. 1946. Flora de Cuba. Contribuciones Ocasionales del Museo de His-
toria Natural del Colegio de la Salle, No. 8. Orquideas, pp. 341 404, Habana,
Cuba.

LINDLEY, J. 1843. Edwards's Botanical Register, Vol. 29, Misc. p. 17.

1846. Orchidaceae Lindenianae. Bradbury and Evans, Whitefriars,

London, England.

1853. Paxton’s Flower Garden, Vol. III. Bradbury and Fvans. London,
England.

LioGieR, A. H. 1969. Flora de Cuba. Suplemento. Editorial Sucre, Caracas,
Venezuela.

NortHRop, A. R. 1902. Flora of New Providence and Andros. Mem. Torry Bot.
Club, 12: 1-98.

RicHaRD, A. 1850. /n: Sagra’s Historia Fisica, Politica y Natural de la Isla de
Cuba, Orchidaceae, Pt. 2, 11, Botanica, pp. 234 253, tab. 74 86.

Roirr, R. A. 1889. List of Garden Orchids. Gard. Chron. Ser. 3, 5: 491.

SCHLECHTER, R. 1915. Die Orchideen. Paul Parey. Berlin.

DEPARTMENT OF BIOLOGICAL SCIENCES
FLORIDA ATLANTIC UNIVERSITY
BOCA RATON, FL 33431

A RE-EVALUATION OF SPIRANTHES
XSTEIGERI CORRELL

PAuL M. CATLING

ABSTRACT

The hybrid Spiranthes Xsteigeri Correll has been re-examined and found to be
referable to S. ochroleuca (Rydb.) Rydb., a recently re-established species. Synon-
ymy of the hybrid with S. ochroleuca is proposed.

Key Words: Spiranthes, putative species hybrid, extant synonymy

Spiranthes Xsteigeri Correll was proposed as a hybrid of S. cer-
nua (L.) L. C. M. Rich. and S. romanzoffiana Cham. parentage in
1941, based on material collected on 22 September 1940 by Theo-
dore L. Steiger in a “dense grassy marshland” near Warner, Merri-
mack County, New Hampshire (Correll, 1941). It has become the
most frequently recognized and widely accepted hybrid in nor-
theastern Spiranthes, having been reported over a broad geographi-
cal area and included in various manuals (e. g. Correll, 1950; Case,
1964: Voss, 1972: Luer, 1975).

Correll (1941) made reference to several plants, four of which he
referred to Spiranthes cernua and three which he considered to
represent the new hybrid, S. Xsreigeri. The largest plant of the three
was recognized as a monstrosity, the flowers having multiple
aborted lips with adnate anther tissue. This plant was reterred to the
hybrid since its abnormality suggested a hybrid origin and it was
found growing with the hybrid. The smallest of the three plants had
rather small calli at the base of the lip suggesting S. romanzoffiana.
However, it was apparently the plant of intermediate size (of these
three) upon which Correll (1941) based his description, /. e. “the
plant with intermediate characters that typifies the hybrid. ..”. The
flowers of this plant were described as having the characteristic
ascending appearance of the flowers of S. romanzoffiana, and the
basal calli of the lips “approached most closely those of S. cernua”.
Lips from the flowers of this plant were reported to be highly varia-
ble, ranging from distally pandurate to pandurate on one side to not
at all pandurate (Correll, 1941, Figures 3, 4, and 5).

The type sheet (AMES 59049) contains three plants. These are
definitely the same three plants on which Correll based his taxon

469

470 Rhodora [Vol. 86

since the largest (left hand) plant has abnormal flowers with multiple
pollinia and viscidia and anther tissue developed in some perianth
parts, as described and illustrated (Correll, 1941).

All three plants have glandular pubescence, ascending flowers,
leafy stems, and rather long inflorescences accounting for 22-30%
of the plant height. These features and the general appearance sug-
gest the recently re-established Spiranthes ochroleuca (Rydb.)
Rydb. (Luer, 1975; Sheviak and Catling, 1980). The ascending
flowers noted by Correll (/oc. cit.) as a feature of S. romanzoffiana
are also characteristic of §. ochroleuca.

With the permission of L. A. Garay, curator of AMES, a flower
was removed from each of the three plants on the type sheet, sof-
tened in warm strong saline and drawn with the aid of camera
lucida (Figure 1, a-o). The left-hand plant (the largest) has flowers
that are characteristic of Spiranthes ochroleuca with respect to the
relatively well developed basal calli, relatively long claw, cuneate
base, median constriction and fleshy centre (Figure |, a). The
column (Figure I, f) is abnormal, having two viscidia, and anther
tissue is apparent in the lateral petals (Figure 1, c, d).

The middle plant (the smallest) on the type sheet has flowers
approaching Spiranthes ochroleuca with respect to the separation of
dorsal and lateral sepals (Figure |, g), the basal curvature of the lip
stalk (Figure |, h), and the central fleshiness of the lip (not shown).
The basal calli and claw are also relatively well developed (Figure 1,
i). However, the truncate base of the lip and relatively wide perianth
parts (Figure I, j, k, 1; Correll, 1941, Figure 6) are unusual in S.
ochroleuca.

A flower from the plant of intermediate size (i. e. the right-hand
plant and the one Correll apparently considered most typical of the
hybrid) has a lip with well-developed basal calli, a cuneate base. a
slight central constriction and a conspicuously fleshy centre (Figure
1, 0). Itis therefore clearly referable to Spiranthes ochroleuca. While
it may be that some flowers on this plant have a median constriction
in the lip, they are not pandurate as in S. romanzoffiana.,

All three plants are lacking in intermediacy in certain characters
that would allow confidence in the hybrid concept. For example,
lateral sepals are not basally connate and distally confluent with the
dorsal sepal, the pubescence is not shorter and sparser, there is not
strong venation in the lateral sepals, the pattern of lip venation is not

1984] Catling — Spiranthes 471

j

10 mm

ho /

Figure |. Camera lucida drawings of flowers from the type sheet of Spiranthes
Xsreigeri Correll (AMES 59049). a-f, left-hand plant: a, lip: b, dorsal sepal: ¢, d.
lateral petals; e, lateral sepal; f, column from below. g-l, middle plant; g, flower in
lateral view; h, basal portion of flower with lateral sepal removed; i, lip; j, lateral
sepal,; k, dorsal sepal; 1, lateral petal: m-o, right-hand plant: m, lateral sepal: n,
lateral petal; o, lip.

472 Rhodora [Vol. 86

intermediate, nor are the lengths of the basal calli. Pollen from each
of the three flowers examined was normal, i. e., there was no indica-
tion of disturbed meiosis through the presence of dyads or monads, as
one might expect from a hybrid involving plants as different as
Spiranthes romanzoffiana and either S. cernua or S. ochroleuca (c.f.
Simpson & Catling, 1978).

Although a first generation hybrid may show only the dominant
expression of a particular trait, most hybrids are more or less inter-
mediate and demonstrate features of both parents. This intermediacy
is not the case with the putative Spiranthes Xsteigeri. The only
characters in the type material which could be regarded as atypical of
S. ochroleuca are the disorganized floral structure in the left hand
plant, and the relatively wide perianth parts and truncate lip base (the
latter a feature of S. cernua instead of S. romanzoffiana) in the
middle plant. These few abnormalities do not seem to be sufficient to
assume a hybrid.

The four plants assigned to Spiranthes cernua which were col-
lected with S. Xsteigeri on the same date (AMES 59203) are also
referable to S$. ochroleuca. Correll’s misidentification of Steiger’s
collections as S. cernua and as a S. cernua X S. romanzoffiana
hybrid is understandable given the lack of information on various
diagnostic characters at the time. Furthermore, S. ochroleuca was
poorly understood then and considered trivial until quite recently
(cf. Correll, 1950; Luer, 1975: Sheviak and Catling, 1980).

As for the reports of Spiranthes Xsteigeri from Yarmouth
County, Nova Scotia (Correll, 1950), Chippewa County, Michigan
(Case, 1964) and Caledonia County in northern Vermont (Luer,
1975), it appears that there are no documenting specimens (for list
of herbaria checked, see Sheviak and Catling, 1980). Luer’s (1975)
photograph of S. Xsreigeri from Vermont appears to represent only
an abnormal specimen of §. romanzoffiana. A plant labelled S.
xsteigert collected from the shoreline of Deep Lake in Barry
County, Michigan by F. W. Case on 4 September 1967 (MICH), a
site with both putative parents present (MICH), has a weakly pan-
durate lip. This characteristic and its unusually late flowering may
suggest a hybrid, but otherwise the plant resembles S. romanzoffi-
ana, which does occasionally bloom late. The pollen of this plant is
normal. Both of the Michigan reports of S. Xsteigeri have recently
been rejected (Case and Catling, 1983). In view of the points dis-

1984] Catling — Spiranthes 473

cussed above, I formally propose that Spiranthes Xsteigeri Correll
be considered a synonym of S. ochroleuca (Rydb.) Rydb.

LITERATURE CITED

Cask, F. W.. uk. 1964. Orchids of the western Great Lakes region, Cranbrook
Institute of Science. Bloomfield Hills. 147 pp.

AND P.M. CATLING. 1983. The genus Spiranthes in Michigan. Michigan
Bot. 22: 79-92.

Corrett, D.S. 1941. A new Spiranthes hybrid from New Hampshire. American
Orchid Society Bulletin 9:241.

1950. Native orchids of North America north of Mexico. Chronica
Botanica Co., Waltham, Massachusetts. 399 pp.

Luer,C. A. 1975. The native orchids of the United States and Canada excluding
Florida. New York Botanical Garden. 361 pp.

Sneviak, C.J. AND P.M. CatTLiInG. 1980. The identity and status of Spiranthes
ochroleuca (Rydberg) Rydberg. Rhodora 82: 525 562.

Simpson, R. C. AND P.M. CATLING. 1978. Spiranthes lacera var. lacera * S,
romanzoffiana, a new natural hybrid orchid from Ontario. Can. Field-Nat.
92(4): 350-358.

Voss, E.G. 1972. Michigan Flora. part 1. Gymnosperms and Monocots. Cran-
brook Inst. Science Bull. 55: 488 pp.

BIOSYSTEMATICS RESEARCH INSTITUTE
WILLIAM SAUNDERS BUILDING
CENTRAL EXPERIMENTAL FARM
OTTAWA, ONTARIO

CANADA KIA 0C6

TAXONOMY OF THE V/C/IA LUDOVICIANA
COMPLEX (LEGUMINOSAE)!

J. STUART LASSETTER

ABSTRACT

The annual vetches Vicia ludoviciana Nutt. ex T. & G.. V. leavenworthii T. & G.,
V. exigua Nutt. ex T. & G., and their varieties compose the native North American
V. ludoviciana complex. The center of diversity is Texas: distribution is from Ala-
bama to California, and from northern Mexico into the Rocky Mountains of Colo-
rado. Karyotypes, artificial hybridizations. characteristics of the breeding system,
and various morphological data are used with previously reported edaphic relation-
ships and seed morphology to establish relationships reported herein. This taxo-
nomic treatment reduces the taxa of the complex to | wide-ranging and variable
species, consisting of | subspecies containing 5 geographical races, and a second
subspecies containing 2 races.

Key Words: Vicia, North America, taxonomy, chromosomes, breeding system

INTRODUCTION

The Vicia ludoviciana complex (Leguminosae) is a group of
annual vetches native to North America. The complex includes
Vicia ludoviciana Nutt. ex T. & G., V. leavenworthti T. & G., V.
exigua Nutt. ex T. & G.. and varieties of each. Lassetter (1975) has
shown that Vicia hassei Wats. (V. exigua var. hassei) is not part of
this complex. Taxa of the complex are found from Alabama to
California and from northern Mexico into the Rocky Mountains of
Colorado. The center of diversity is Texas where the greatest num-
ber of taxa occur and where intergradation between taxa Is greatest.

The taxonomic relationships within the complex have been
problematic, and conclusions of different authors have not agreed.
Previous workers relied almost solely on field and herbarium data.

Shinners (1948) recognized Vicia ludoviciana (varieties tyvpica,
texana, laxiflora) and V. leavenworthii (varieties typica, occi-
dentalis) in Texas. Species were separated by peduncle length, and
varieties by overall size as well as flower characters. According to
Shinners, the distribution of taxa is at least partly associated with
soil texture. He stated that overlaps and intergradation between

‘Journal Paper No. J-9717 of the lowa Agriculture and Home Economics Experi-
ment Station, Ames, lowa. Project No. 1983.

475

476 Rhodora [Vol. 86

taxa exist. Shinners did not consider the western V. exigua as
occurring in Texas.

Turner (1959) also recognized two species in Texas, but distin-
guished them on the basis of flower number. Varieties were sepa-
rated by leaflet characters and by length of flowering axes. Like
Shinners, Turner believed that distribution was related in part to
soils, and stated that intergradation exists between varieties of each
species, and between the species. Turner considered Vicia ludovici-
ana var. texana to be V. leavenworthii var. occidentalis.

In addition to Vicia ludoviciana and V. leavenworthii, Hermann
(1960) also believed V. exigua extended into Texas and was in fact
V. leavenworthii var. occidentalis sensu Shinners and Turner. These
three species were separated by Hermann on the basis of number of
flowers per peduncle, peduncle length, and flower character.

Isely (personal communication) believed that only two species
exist although this scheme was not formally proposed. He viewed
Vicia exigua (as did Hermann, 1960) to encompass V. leavenworthii
var. occidentalis sensu Shinners. Isely believed all the other taxa
were best recognized as one of two varieties of V. /udoviciana.

Because the total range of the complex is widespread, the constit-
uent taxa have been included in several state and local floras. Cor-
rell and Johnston (1970) is the only treatment that covers a
geographical area from which all three species have been reported.
The treatment of Vicia by these authors is largely adapted from
Hermann (1960), and their view of the V. /udoviciana complex
differs only in minor details of ranges and descriptions.

The traditional taxon names of Shinners are predominately used
throughout this paper so that the reader can better relate to the taxa
being discussed. Name changes and shifts in taxon concepts are
largely reserved for the taxonomic treatment.

MATERIALS AND METHODS

Chromosome study included meiotic counts determined from
anther squashes and karyotype analyses from mitotic root material.
Field-collected buds were fixed in Newcomer’s (1953) fixative. Buds
and root tips from greenhouse plants were fixed in acetic-alcohol

1984] Lassetter — Vicia 477

(absolute ethanol: glacial acetic acid—3:1). Anther material was
stained with propiocarmine. Root tips were pretreated for 4 hours at
13-14° C in 0.002 M 8-hydroxyquinoline and were stained with
Schiff’s reagent according to the Feulgen method. Slides were made
permanent by the method of Bowen (1956). Meiotic figures and
karyotypes were drawn with a Zeiss camera lucida and a Leitz
drawing apparatus, respectively. Voucher specimens are deposited
in the lowa State University Herbarium (ISC).

Reciprocal artificial hybridizations were attempted between the
three traditional species. It was necessary to emasculate the self-
fertile flowers because pollen is often released in the bud stage. A
total of 216 crosses was made. An additional 15 emasculations with
no pollen transfers were performed as a check for apomixis. Obser-
vations were made on the breeding system.

One hundred six mass collections of about 1,200 total specimens
were made. Fifty-five morphological characters were observed and
recorded. The characters observed included those used by previous
workers that I also believed useful, as well as some additional ones.
For brevity, the entire list and explanation is omitted here, but is
available from Lassetter (1972).

Herbarium studies were conducted by utilizing material from the
following herbaria: ARIZ, ASC, BRY, CAS, COLO*, DS, F*,
FSU, GH, ISC, LA, LAF, LSU, MEXU*, MICH*, MIN, MISS,
MISSA, MO, NLU, NMC, NO, NY, OKL, OKLA, ORE, PH*,
POM*, RSA, SBBG, SMB, SD, SMS, SMU, TENN, TEX, TTC,
UARK, UC, UM, UNM, US, VDB, and University of Southern
Mississippi (herbarium designations follow Holmgren and Keuken
(1974); asterisks indicate selected material only).

Lassetter (1978a) discussed the taxonomic significance of the fol-
lowing seed characters: diameter, relative hilum length (hilum
length + seed circumference X 100), hilum length, hilum width at
wide end, hilum width at narrow (lens) end, base color of testa, and
color of mottling. Identification of taxa by seed characters alone
was shown to be unreliable due to overlapping variation.

Lassetter (1978b) reported on edaphic relationships and demon-
strated that some taxa are rather plastic and regularly occur in soils
of various texture. Other taxa were only found on sandy or sandy
loam soils.

478 Rhodora [Vol. 86

RESULTS AND DISCUSSION

Cytological data

The genus Vicia has chromosome base numbers of x = 5, x = 6,
and x = 7 (Senn, 1938; Darlington and Wylie, 1955; Fedorov,
1969). Workers presume that the lower numbers are derived from
an ancestral n = 7 (Stebbins, 1958; Rousi, 1961).

During the present study, more than 40 meiotic counts were
made, several for each taxon from various locations throughout the
range of the complex. All counts for all taxa were n = 71; (Figure |
comprises a sample); meiosis appeared normal, as previously
observed by Turner (1956).

Previous to this study only three chromosome counts were
reported in the literature. Turner (1956) reported three determina-
tions of n = 7 for Vicia ludoviciana var. typica and one determina-
tion of 2n = 14 for V. leavenworthii var. typica. Turner also listed
an unpublished count of m = 7 by Johnston for V. leavenworthii
var. occidentalis.

Figure |. Meiotic chromosomes of Vicia. Line scale = 10 microns
1. V. ludoviciana var. typica (race 1). Voucher grown from seed from S. Mc-
Daniel 8881, Dauphin Island, Alabama, (FSU #124825).
2. V. ludoviciana var. typica (race 1). Lassetter 1760, Walker Co., Texas.
3. V. ludoviciana var. laxiflora (race 5). Lassetter 1721, Val Verde Co., Texas.
4. V. ludoviciana var. laxiflora (race 5). Lassetter 1851, Llano Co., Texas.
§. V. ludoviciana var. texana (race 2). Lassetter 1773, Aransas Co., Texas.
6. V. leavenworthii var. typica (race 6). Lassetter 1832, Hill Co., Texas.
7. V. leavenworthii var. typica (race 6). Lassetter 1847, Concho Co., Texas.
8. V. leavenworthii var. occidentalis (race 4). Lassetter 1731, Culberson Co.,
Texas.
9. V. leavenworthii var. occidentalis (race 4). Voucher grown from seed from D.
Keil 1162, Maricopa Co., Arizona (ARIZ #38829).
10. V. exigua var. exigua (race 3). Voucher grown from seed collected by I. Marin,
Riverside Co., California.
11. V. exigua var. exigua (race 3). Voucher grown from seed from D. H. Hoover
s.n. San Diego Co., California, (CAS #514387).
12. V. ludoviciana var. texana (race 2). Lassetter 1779, Willacy Co., Texas.
13. V. ludoviciana var. texana (race 2), Lassetter 1783, Jim Wells Co., Texas.
14. V. ludoviciana var. laxiflora (race 5). Voucher grown from seed from M.
Fechu and F. Martizen 160, Chihuahua, Mexico (TEX s.n.).
15. V. exigua var. exigua (race 3). Voucher grown from seed from R. N. Philbrick
B65-1399 and M. Benedict, Santa Cruz Island, California (SBBG #29483).
16. V. leavenworthii var. occidentalis (race 4). Lassetter 1726, Brewster Co..
Texas.

1984] Lassetter — Vicia 479

Most karyotyping of Vicia species has been done with those
native to the Old World, for which much literature exists. Svesh-
nikova (1927) investigated 27 Old World species and found that
almost every species had a different karyotype.

Shrivastava (1963) studied eight Old World species and found
eight different karyotypes. Mettin and Hanelt (1968) studied 11 Old
World and one South American species and concluded that most
taxa in Vicia are characterized by different karyotypes, that chro-
mosome sizes also are somewhat different, and that these differ-
ences seem to apply to both Old and New World species.

pe 642 amass

ow? * Cd

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9 10 I] I2
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480 Rhodora [Vol. 86

Stankevicz (1970) studied morphological, karyological, and eco-
logical-geographical characteristics of 53 Russian species of Vicia.

Very little karyotype work has been done with New World spe-
cies. Turner (1956) illustrated 2” = 14 chromosomes of V. /eaven-
worthil var. typica, but the drawing does not show the satellited
chromosomes that are present. It was presumably made for a count
only and was not a karyotype preparation.

Lassetter (1975) utilized karyotypes and other data in a taxo-
nomic treatment separating Vicia hassei from V. exigua, and Las-
setter and Gunn (1979) used karyotypes in establishing relationships
between V. menziesii, V. nigricans, and V. gigantea.

Veerasethakul and Lassetter (1981) presented a preliminary sur-
vey of native New World Vicia karyotypes and reviewed previous
work with New World species.

Taxa of the V. /udoviciana complex all have the same basic
karyotype (Veerasethakul and Lassetter, 1981). I believe pairs one,
two, five, and seven (numbered from left to right) are the same in all
karyotypes. Some minor variation in these pairs was observed in
different preparations from the same taxon, and sometimes small
variation from cell to cell in one preparation was observed. These
differences are, | believe, due to slight inconsistencies in length
and/or temperature of pretreatment, and to differences in the pre-
cise time of mitosis at which cells were fixed.

The differences in the other three pairs of chromosomes are
believed to represent inherent karyological differences between taxa
and are given in Table 1.

With regard to evolution of karyotypes, Levitskly (1931) formal-
ized the concept of symmetrical karyotypes (chromosomes all
metacentric or submetacentric and nearly equal in size) as con-
trasted to asymmetrical karyotypes, and he proposed a trend of
increasing karyotype asymmetry in angiosperms.

According to Stebbins (1950), perennial vetches with n = 7 have
mostly symmetrical karyotypes, while karyotypes of annuals with a
reduced and derived n = 6 or n = S are highly asymmetrical.

Taxa in this complex are annuals with asymmetrical karyotypes
which are n = 7 and presumably have achieved asymmetry without
a reduction in chromosome numbers. Stebbins (1958, 1971) stated
that karyotypes of species or taxa of lesser rank (all with the same
chromosome number) that differ in size and form of individual

1984] Lassetter — Vicia 481

Table |. Variation in three chromosome pairs within the Vicia ludoviciana complex.

Taxon Pair three Pair four Pair six

V. ludoviciana telocentric sub-metacentric telocentric
var. typica

V. ludoviciana telocentric sub-metacentric sub-metacentric
var. laxiflora

V. ludoviciana sub-telocentric sub-metacentric telocentric
var. texana
Race 7 (has no telocentric sub-telocentric telocentric

traditional taxon name)

V. leavenworthii telocentric sub-telocentric telocentric
var. (pica
V. leavenworthtt telocentric sub-telocentric telocentric

var. occidentalis

V. exigua telocentric sub-telocentric sub-metacentric
var. exigua

chromosomes are best explained by an assumption of increasing
asymmetry. Ina group of taxa undergoing such a process, the most
symmetrical karyotypes would be the most primitive, and vice
versa.

From Table | and Veerasethakul and Lassetter (1981), the karyo-
type of Vicia ludoviciana var. laxiflora is the most symmetrical in
the complex. The most asymmetrical karyotype is the one common
to V. leavenworthii var. typica, V. leavenworthii var. occidentalis,
and race 7 of this taxonomic treatment (this race has never received
formal recognition and therefore has no traditional taxon name).
These three taxa, however, are morphologically dissimilar. A sim-
ilar situation was reported by Rousi (1961) who found small karyo-
type differences in different strains of V. renuifolia.

Most attempts at hybridization of Vicia taxa have not been suc-
cessful, and this lack of success has been attributed to karyotype
differences. Most workers have been unsuccessful even in crossing
different strains of a single species (Sveshnikova, 1927). Plitmann
(1967) also stated that most workers have been unsuccessful in
crossing different taxa of Vicia. Of the crosses evaluated by Plit-
mann (1967), most involved cultivated species of Eurasian origin,
and most were intraspecific crosses between subspecific taxa. Gunn

482 Rhodora [Vol. 86

(pers. comm.) even rejected all reputed “successful hybridizations”
in the literature and stated that the only successful ones have been
between some varieties of “Cracca” species, and between V. sativa
and V. angustifolia. These two closely related taxa are highly influ-
enced cultivars and were shown by Sveshnikova (1927) and Hol-
lings and Stace (1974) to have similar karyotypes. They have been
considered as a single species by Ball (1968), Gunn (1970, 1979), and
others.

Artificial hybridization

Reciprocal intraspecific crosses involving Vicia ludoviciana and
V. leavenworthii were successful and produced fully viable F;
plants, but those involving V. exigua were not successful. Dor-
mancy in V. exigua seed is difficult to break. Seeds from crosses
involving V. exigua were treated with 0.1 M thiourea (Ballard and
Buchwald, 1971), which forced radicles to emerge, but no seedlings
resulted. The same difficulty existed in germinating field-collected
V. exigua seeds for cytological work. | believe the lack of F, V.
exigua plants was largely because of faulty germination technique
and not to inviability of the hybrid seed.

Reciprocal interspecific crosses were mostly unsuccessful; most
seeds were shriveled, and seedlings were abnormal, with weak radi-
cles and kinked, stunted epicotyls. In cross 1 (Table 2), fruits
appeared normal but contained only shriveled, inviable seeds. No
F, plants resulted from cross 1, 3, 4, or 6. One F, plant resulted
from cross 2. The plant was very slow-growing, small, and stunted.
It eventually flowered, and several young fruits were produced, but
they quickly withered, and no seeds were set. Pollen from this plant
was 51 percent sterile. Cross 5 (Table 2) produced a progeny of four
F, plants. All were weak and lacked normal vigor. One plant grew
for two weeks, became chlorotic, and died. The other three flow-
ered, but produced no fruits or seed. Anthers were minute (ca.
0.15 mm) and did not dehisce. Almost no pollen grains stained with
cotton blue in lactophenol, and most were spherical or elliptical as
compared with the normal elongate appearance of mature grains.
Evidently, the anthers and pollen ceased development before either
matured. None of the five F; plants, therefore, produced any seed.

Anther squashes of the F, plants from cross 2 and cross 5 (Table
2) were not successful, and meiosis was not observed. Removal of

1984] Lassetter — Vicia 483

Table 2. Results of interspecific artificial hybridizations within the Vicia ludo-
viciana complex.

Number of
Number of crosses
crosses producing Number of
Taxa of cross attempted seed F, plants
1. V. ludoviciana var. (pica
x
V. leavenworthii var. tvpica 39 18 0
2. V. ludoviciana var. typica
x
V. exigua var. exigua 32 13 l
3. V. leavenworthii var. tvpica
x
V. exigua var. exigua 28 13 0
4. V. leavenworthii var. occidentalis
x
V. ludoviciana var. typica 3 l 0
5. V. leavenworthii var. occidentalis
x
V. leavenworthii var. typica 8 4 4
6. V. leavenworthii var. occidentalis
x
V. exigua var. exigua 7 3 0

stem tips for fixation of buds adversely affected the plants and they
never recovered.

From these data, outcrossing in individual populations seems
entirely possible inasmuch as intraspecific crosses were successful.
Hybridization between taxa seems much less likely because crosses
between different taxa produced few F, plants, and they were all
abnormal. Even progeny from V. /eavenworthii var. occidentalis X
V. leavenworthii var. typica (cross 5, Table 2) were sterile, indicat-
ing that at least some barriers to interbreeding exist between these
supposedly closely related taxa.

No apomixis was indicated.

Breeding system

Observations on the breeding system were made from living
plants in the field and greenhouse. All taxa are very efficient selfers;

484 Rhodora [Vol. 86

pollen usually is released within the young bud before the flower
opens. In wet mounts of styles from young buds, germinating pollen
grains were common.

Bagging did not reduce fruit set. When all buds but one were
removed from the peduncle and that single flower bagged, fruit set
still was successful. Shaking the plant did not alter fruit set; self-
fertilization seemed effective almost to the point of cleistogamy.

In all my field work, no pollinators were observed on these
vetches. Pollinators were seen in the immediate area, but clearly
preferred other flowers. The only record of actual insect manipula-
tion seems to be a herbarium sheet notation by Shinners, “Visited
by honeybees” (Shinners 9793, ARIZ, SMU).

It seems reasonable to conclude that a small proportion of flow-
ers probably reach anthesis with intact anthers and are at least
potential outcrossers. However, because pollinators would be
required for successful outcrossing, and pollinators seem infrequent,
outcrossing must be rare.

Plitman (1967), in studying 50 annual Vicia species of the Middle
East, concluded that self-pollination occurs in young buds precisely
as just described. He stated that self-pollination, cleistogamy, and
cross-pollination may occur in the same species, even the same indi-
vidual, but suggested no actual method by which cross-pollination
might occur in a predominantly self-pollinated taxon.

Allard and Workman (1963) and others have shown that, in pop-
ulations of many different self-fertilized species, some outcrossing
does occur. Allard et al. (1968) reviewed data supporting the
hypothesis that most inbreeders contain large stores of genetic vari-
ability and that genetic and morphological variation within species
and within populations can be considerable.

A different system evidently operates in Vicia leavenworthii var.
(ypica and other traditional taxa in part (V. /udoviciana subsp.
leavenworthii of the following taxonomic treatment). These popu-
lations were the most homogeneous of the complex. These plants
have a short style, the anthers at dehiscence are all positioned above
the stylar pubescence (Figure 2), and pollen is released directly onto
the stigma. In other taxa with longer styles (which represent V.
ludoviciana subsp. ludoviciana of the following taxonomic treat-
ment), anthers at dehiscence are at the level of stylar pubescence
(Figure 2), and pollen is released upon the stylar hairs. The stylar
hairs are antrorse, and in the confines of the folded keel, pollen is

1984] Lassetter — Vicia 485

Figure 2. Anther positions of Vicia..
1. V. ludoviciana subsp. leavenworthii (of this taxonomic treatment, races 6
and 7).
2. V. ludoviciana subsp. /udoviciana (of this taxonomic treatment, races I~ 5).

eventually worked apically to the pocket at the keel tip and effec-
tively placed onto the stigma.

Anthers in Vicia ludoviciana subsp. leavenworthii are shorter
(0.30-0.35 mm) than in subsp. /udoviciana (0.40-0.55 mm), this
condition may reflect a more effective self-pollination system. In
subsp. /eavenworthii, anthers were always observed to dehisce well
before flowers opened, and young fruits were most often already
developing at anthesis. In addition, flowers open before peduncles
and internodes elongate, and the flowers are inconspicuous among
the leafy stem tips. Even if some flowers were potential outcrossers,
they presumably would be inconspicuous to pollinators.

In Vicia ludoviciana subsp. leavenworthii, it seems that almost no
flowers could be potential outcrossers, thus this taxon must be very
highly self-pollinated.

TAXONOMIC TREATMENT

l am interpreting the Vicia /udoviciana complex as a single multi-
racial species because its elements are all confluent. An opposing
position might be supported on the basis of karyotype differentia-
tion and infraspecific incompatibility. I believe it better, however,
to limit formal names to only two subspecies that can be relatively
well defined. Several taxa which have previously been given formal

486 Rhodora [Vol. 86

species or varietal rank are recognized informally as geographical
races.

The characters listed in the key and descriptions are those that
have proved most useful in identification and recognition.

KEY TO SUBSPECIES AND RACES
OF VICIA LUDOVICIANA

1. Flowers opening after peduncles and internodes elongate, pinkish-white to deep
bluish-purple, young fruit usually not present when flowers first open; styles

(0.60) 0.80-1.40 (1.70) mm long; leaflets (5) 7-10 (13) ..... cece eee ee eee
Jars tw “@ gligciey6 Gn eve ete sore: & ai '929. w 46's 108 ew Salt eee A: subsp. /udoviciana 2

2. Style length (0.6) 0.8-1.2 (1.3) mm: calyx tube (1.1) 1.4-1.8 (2.2) mm long;
stems at midpoint glabrous to pubescent ........0ccc eee eeeecees 3

3. Flowers 1-19, averaging 5-9; fruiting peduncles (0.4) 0.6-0.9 (1.3) mm wide:
hilum/circumference ratio in percent (22.5) 29.0-37.4 (42.4); plants of
eastern Texas, Louisiana, Arkansas, and southern Mississippi and extreme
Southern Alabama..........0eeceeeaee subsp. /udoviciana race |

3. Flowers 1-10, averaging 1-3; fruiting peduncles (0.2) 0.3-0.6 (0.8) mm wide:
hilum / circumference ratio in percent (11.2) 17.4-23.7 (31.8) ........ 4

4. Flowers usually 3 with one terminal flower and a contiguous pair below;
plants of the Gulf Coastal and Rio Grande Plains of Texas ........

RCE LORE ORT On Te ae aa subsp. /udoviciana race 2

4. Flowers 1-3, each flower usually arising at a separate point on the pedun-
cle; plants of California and Baja California. ..........0000000s

epiwie ai enaele, Stele as SSE eee sugars eeu ewre ce, Sue subsp. /udoviciana race 3

2. Style length (0.9) 1.1-1.4 (1.7) mm; calyx tube (1.3) 1.7-2.1 (2.5) mm long;
stems at midpoint glabrous to densely pubescent ..........e00e05 5

5. Peduncles 1-5 flowered, (0.9) 8-35 (47) mm long when flowers are open,
usually about 1/3 the length of the subtending leaf; hilum/circumfer-

ence ratio in percent (7.9) 10.4-18.1 (20.7); plants of western Texas, the
Oklahoma panhandle, Colorado, Utah, New Mexico, Arizona, extreme
southern Nevada, and extreme southeastern California ...........

Slee 6.6 Gr bce ae 4 6p eM: Bere ee 4a a yiard- wee ee subsp. /udoviciana race 4

5. Peduncles 1-17 flowered, (15) 21-41 (120) mm long when flowers are open,
about as long or longer than the subtending leaf; hilum/ circumference

ratio in percent (16.7) 18.7-20.9 (24.3); plants of central Texas, southern
Oklahoma, and southern New Mexico..... subsp. /udoviciana race 5

1. Flowers opening before peduncles and internodes elongate, pinkish-white to light
lavender, often containing young fruit when flowers first open; styles (0.40)
0.60-0.80 (0.90) mm long; leaflets (7) LI-15 (17)... ec ce ee ce ee ee ew eee
<eaite 6 ga ra™a SS & fal 'W.-ehe eh 915.6 e944. 06 & WAP Rose B: subsp. /eavenworthii 6

6. Flowers 1-6; legumes mostly less than 30 mm long; length of hilum less than
the seed diameter; leaflets 2.5-7.0 times as long as wide; plants of central
Texas, Oklahoma, extreme northwestern Arkansas, and extreme south-
western MISSOUTL .... eee ee ee ee ee eee subsp. /eavenworthii race 6

1984] Lassetter — Vicia 487

6. Flowers 1-2; legumes mostly 30 mm long or longer; length of the hilum
greater than the seed diameter; leaflets 2.0-3.0 times as long as wide; plants
of Louisiana, Oklahoma (McCurtain Co.), scattered sites in Arkansas, and
Mississippi (Claiborne Co. and Sharkey Co.)..... eee ee eee eee
eee 0; ges We’ Se ea ara. 0d, 88 © a6 S84 Brea ales subsp. leavenworthii race 7

A: Vicia ludoviciana Nutt. ex T. & G. subspecies ludoviciana

Vicia ludoviciana Nutt. ex T. & G. Fl. N. Amer. 1: 271. 1838. LECTOTYPE Ny!: Dr.
Leavenworth, Near Nagodoches (sic!) and Natchitoches [Louisiana].

Cracca ludoviciana (Nutt. ex T. & G.) Alefeld. Bonplandia 9: 119. 1861.
Vicia ludoviciana var. tvpica Shinners. Field Lab. 16: 23. 1948.

Vicia caroliniana var. texana T. & G. Fl. N. Amer. 1: 271. 1838. LECTOTYPE NyY!:
Leavenworth, Texas.

Vicia texana (T. & G.) Small Fl. Se. U.S. p. 656 1903.
Vicia ludoviciana var. texana (T. & G.) Shinners. Field Lab. 16: 23. 1948.

Vicia thurberi S. Wats. Proc. Am. Acad. Sci. 25: 129. 1890. LECTOTYPE GH!;
ISOLECTOTYPES GH!, NY!: Thurber 299, Dona Ana [Co., New Mexico]. SYN-
TYPES GH!: Thurber 150. SYNTYPES GH !, US!: Wright 1350. SYNTYPE ISC!, NY!:
Parry 33, 1874, S. Utah.

Vicia producta Rydb. Bull. Torr. Bot. Club 28: 500. 1901. LECTOTYPE Ny!, ISO-
TYPE NY!: P. A. Rydberg and F. K. Vreeland 6006, Butte, 5 miles southwest of
La Veta, Colorado, May 22, 1900.

Vicia ludoviciana var. laxiflora Shinners. Field Lab. 16: 25. 1948. HOLOTYPE
sMU!, ISOTYPE MO!: Eula Whitehouse 15275, April 13, 1946. About 3 miles
north of Bridgeport on Highway 24, Wise County, Texas.

Vicia leavenworthii var. occidentalis Shinners. Field Lab. 16: 22. 1948. HOLOTYPE
SMU! ISOTYPE MO! NY!: C. H. Muller, Chisos Mts., Boot Spring. July 29, 1932.

Vicia exigua Nutt. ex T. & G. Fl. N. AM. 1: 272. 1838. HOLOTYPE BM (photo-
graph!). isoTYPE PH!: Nuttall, Columbia Plains (see discussion below).

Cracca exigua (Nutt. ex T. & G.) Alefeld, Bonplandia 9: 119. 1861.

DESCRIPTION. Plants diminutive to robust, 0.8—20.0 dm tall, gla-
brous to pubescent, erect to sprawling, or climbing if support is
available. Leaflets (5) 7-10 (13), (6) 10-19 (37) mm long, (0.9)
1.6-3.8 (11) mm wide, the length—-width ratio (2.0) 3.7-8.7 (16.4),
the apex acute to emarginate. Flowers opening after peduncles and
internodes elongate, the flowering peduncles (0.9) 6-30 (101) mm
long, as long as to shorter than the subtending leaf, the subtending
leaf (10) 24-46 (75) mm long. Fruiting peduncles (1.1) 21.7-59.7
(150) mm long, from shorter than to equaling and longer than the
subtending leaf, the subtending leaf (18) 31-60 (96) mm long. Flow-
ers 1-19, the standard varying from short, stubby and not showy to
long, broad and showy, the height of the reflexed standard (0.64)
0.94-1.40 (1.65) times the flower length from the calyx base to the

488 Rhodora [Vol. 86

tip of wings, length of flower from calyx base to tip of unreflexed
standard (3.5) 5.1-6.9 (9.5) mm. Upper calyx teeth (0.4) 0.7-1.2
(2.6) mm long, usually shorter than the lower tooth, the lower tooth
(0.6) 1.1-1.7 (3.0) mm long and usually shorter than the calyx tube,
the tube (1.1) 1.6-2.0 (2.5) mm long. Styles (0.60) 0.83-1.23 (1.70)
mm long, the apical (0.30) 0.52-0.74 (1.10) mm pubescent. Mature
legume (2.0) 17.8-23.2 (32) mm long. Ovules up to 8, seeds 3-9, the
hilum-circumference ratio in percent (8) 18-33 (42).

NOMENCLATURAL HISTORY AND DISCUSSION OF TYPE SPECIMENS,
Vicia ludoviciana was based on collections by Leavenworth and
Tainturier. The Leavenworth collection contains two specimens,
both of which are traditional V. /udoviciana (V. ludoviciana subsp.
ludoviciana race | of this treatment). I follow Shinners (1948) in
considering this collection as the type. The Tainturier (PH!) gather-
ing consists of four stems on a sheet also containing an unrelated
California collection by Gibbons. This collection is V. /udoviciana
subsp. /eavenworthii, cannot be considered a type for subspecies
ludoviciana, and is discussed under subspecies Jeavenworthii.

Vicia thurberi was based on four different gatherings. Twelve of
these specimens were seen, and all are V. thurberi. However, all the
duplicates of each gathering do not match exactly in label informa-
tion or collection sites and probably represent plants from different
locations. I have chosen one specimen of Thurber 299 at GH as the
lectotype of V. thurberi because it is typical, contains both flowers
and fruits, and the collection is distributed in two major herbaria.
Other types are indicated in the synonomy.

Only a photograph of the holotype of Vicia exigua was available
for this study. Howell, in preparation of the first edition of his
Marin Flora (1949) saw the holotype and affixed the following
note:

From the type description of Vicia exigua Nutt. (T. & G., Fl. N. A. |: 272) it is
obvious Nuttall saw plants from California as well as from the Columbia. |
believe that the two speciens on this sheet are from these two regions and that
the California label has been lost. From the type description it is easy to deter-
mine that the lefthand specimen is the one from “the Oregon”, the one on the
right from “Upper California”. 8/26/35.

John Thomas Howell.

Lassetter (1975), with only a photograph of the BM sheet availa-
ble, designated the righthand specimen as the lectotype. Since that

1984] Lassetter — Vicia 489

time C. R. Gunn has examined the sheet, and after a personal
conversation with him, I consider both specimens on the sheet to be
Vicia exigua; now the entire sheet may be considered holotype
material.

The remaining names reduced to synonomy under Vicia /udovi-
ciana subsp. /udoviciana were each based on only one element, and
no problems with interpretation of type specimens were en-
countered.

The three earliest specific epithets of the complex all
date from 1838. Vicia ludoviciana was chosen for the species name
because it has been associated with the plants of the widest geogra-
phical distribution. Also, little conceptual confusion should result
by choosing V. /udoviciana because the typical subspecies will
retain the epithet /udoviciana, and the kinds of plants traditionally
associated with this name will not be altered.

Subsp. ludoviciana race |

DescRIPTION. Plants diminutive to robust, 1.0-20.0 dm tall, gla-
brous to puberulent, erect to sprawling, climbing if support is avail-
able. Leaflets (5) 8-11 (13), (6) 12-16 (25) mm long, (1.2) 2.5—4.7
(11) mm wide, the length/ width ratio (2.0) 2.8-4.6 (8.5), the apex
acute to truncate or emarginate. Flowering peduncles (6.0) 16.0—
44.0 (101.0) mm long, usually about as long as or longer than the
subtending leaf, the subtending leaf (15) 24-50 (75) mm long. Fruit-
ing peduncles (9) 26-65 (112) mm long, usually about as long as or
longer than the subtending leaf, the subtending leaf (18) 45-69 (97)
mm long. Flowers 1-19, averaging 4-9, the flowers solitary or in
pairs at points on the peduncle, deep blue or bluish purple, the
standard broad and showy, the height of the reflexed standard
(0.81) 0.94-1.12 (1.22) times the length of the flower from calyx
base to tip of wings, the length of flowers from calyx base to tip of
unreflexed standard (4.2) 5.0-6.5 (8.2) mm. Upper calyx teeth (0.5)
0.8-1.2 (1.5) mm long, usually shorter, but varying to as long as the
lower tooth, the lower tooth (0.7) 1.2-1.8 (2.7) mm long, about as
long as the calyx tube, the tube (1.1) 1.41.7 (2.1) mm long. Styles
(0.70) 1.00-1.20 (1.30) mm long, the apical (0.30) 0.60-0.80 (1.00)
mm pubescent. Mature legumes (16) 18-24 (28) mm long. Ovules up
to 8. Seeds 4-8, usually about 5, the hilum/circumference ratio in
percent (23) 29-38 (42).

490 Rhodora [Vol. 86

RANGE AND HABITAT. Eastern Texas, Arkansas, Louisiana, ex-
treme southern Mississippi and Alabama. (Figure 3).

Dense woods and open woodland; thickets; grasslands and pas-
tures; floodplains and low areas; railroad and highway rights-of-
way; fence rows and field edges; weedy areas: shell ridges;
wasteland; lawns and vacant lots.

Mostly in sandy soil, but also in loams, limestone, silty clay, clay.

PHENOLOGY. Flowering in (early February) March-early May
(early July), fruiting in (late March) April-May (July).

Discussion. Taxa which compose race | are Vicia ludoviciana
var. /udoviciana sensu Hermann (1960) in part, and sensu Turner
(1959) in part, and V. /udoviciana var. typica sensu Shinners (1948)
in part.

Race | plants from Texas are typically many-flowered (averaging
7 or more) and usually have 10 or fewer lanceolate to elliptic or
oblong leaflets. This race has the middle-sized flowers of the com-
plex, with other races averaging shorter or longer. Race | plants in
Louisiana and eastward usually are more robust, typically fewer-
flowered (averaging 6 or less), the flowers usually are short and
stubby, and leaflets are mostly 10 or more and oblong to ovate.

The range of race | is essentially the eastern portion of the range
of the entire complex; this race extends west only as far as eastern
Texas (Figure 3). The southwestern limit of distribution of this phase
corresponds closely with the northeastern edge of the Rio Grande
Plain.

Race | is sympatric in part with races 5, 2, 6, and 7, and inter-
grades with all but race 6. In the western portion of its range, race |
tends to have longer upper calyx teeth and a slightly higher flower-
length/standard-height ratio and resembles race 5. In the southern
and southeastern portion of its range, flowers are smaller and fewer,
and plants intergrade with race 2. The lush fewer-flowered race |
plants of Louisiana and eastward resemble race 7.

Subsp. ludoviciana race 2

DESCRIPTION. Plants diminutive to robust, 2.1-19.0 dm tall,
often spindly, glabrous to puberulent, erect to sprawling, climbing
if support is available. Leaflets (7) 8-10 (13), (9) 13-21 (36) mm
long, (1.3) 2.2-4.1 (6.4) mm wide, the length /width ratio (2.4)

1984]

Lassetter — Vicia

491

Distribution of Vicia ludoviciana subsp. /udoviciana, race |.

Figure 3.

492 Rhodora [Vol. 86

3.8-7.6 (14.6), the apex acute to emarginate. Flowering peduncles
(3.5) 11.0-29.0 (63.0) mm long, usually about 1/2 as long as to as
long as the subtending leaf, the subtending leaf (12) 25-48 (64) mm
long. Fruiting peduncles (4) 27-61 (106) mm long, about as long or
longer than the subtending leaf, the subtending leaf (20) 31-53 (72)
mm long. Flowers 1-10, usually 6 or less, often 2 with both flowers
arising from the same point, and often 3 with | upper flower and 2
arising from a single point below, or flowers occurring singly at
points on the peduncle, bluish to deep bluish-purple, the standard
somewhat broad and showy, the height of the reflexed standard
(1.00) 1.10—-1.30 (1.39) times the flower length from the calyx base
to tip of wings, the length of flowers from calyx base to tip of
unreflexed standard (3.5) 4.5-5.8 (6.8) mm. Upper calyx teeth (0.4)
0.6-1.0 (1.5) mm long, usually shorter than the lower tooth, the
lower tooth (0.6) 1.1-1.6 (2.1) mm long, usually shorter than the
calyx tube, the tube (1.1) 1.4-1.8 (2.1) mm long. Styles (0.60)
0.80-1.10 (1.30) mm long, the apical (0.30) 0.45-0.65 (0.80) mm
pubescent. Mature legumes (13) 16-22 (27) mm long. Ovules up to
8. Seeds 4-7, usually 6, but often fewer, the hilum/circumference
ratio in percent (15) 18-24 (32).

RANGE AND HABITAT. The Rio Grande Plain and Gulf Coastal
Plain of southern Texas (Figure 4).

Open woodland; riverbanks and creek beds; mudflats: prairies
and fields; the Gulf shore; roadsides; wasteland: ditches.

In sand, sandy silt, sandy loam, sandy clay, calcareous clay, clay,
silt, over limestone, rocky soil.

PHENOLOGY. Flowering in (late February) March-April (early
May), fruiting in (late March) April-May.

Discussion. Plants which are the most well-known members of
race 2 are Vicia ludoviciana var. texana (Shinners, 1948; Hermann,
1960), but V. leavenworthii var. occidentalis sensu Turner (1959) in
part is also included.

Typical three-flowered race 2 plants have one upper flower and
two flowers arising from a common point below. On two-flowered
peduncles, flowers may arise from the same point or from separate
points. The stubby flowers, along with those of race 3, are the
shortest of the complex.

1984] Lassetter — Vicia 493

Figure 4. Distribution of Vicia ludoviciana subsp. ludoviciana, races 2 and 5.
Stars = race 2, circles = race 5.

In southern Texas, the distributions of races 2, 4, 1, 5 and 6
overlap, and intergradation with race 2 occurs. As one progresses
west up the Rio Grande Valley, race 2 steadily grades into race 4.
Along the Gulf Coast, race 2 and | resemble each other. Race 2 also
blends with race 6 in the Gulf Coast region. Although usually dis-
tinct from each other, race 2 (Figure 4) intergrades with the most
southern race 5 forms.

Correll and Johnston (1970) stated that Vicia ludoviciana var.
texana (my race 2) was particularly perplexing taxonomically. The
distribution of race 2 in Texas is rather limited and, within much of
that range, commonly overlaps and intergrades with other races.

494 Rhodora [Vol. 86

Consequently, areas of intermediate race 2 probably outnumber
areas of typical race 2. Typical race 2 populations are best found to
the exclusion of intergrading forms in Kenedy, Willacy, and
Cameron Counties.

Subsp. ludoviciana race 3

DESCRIPTION, Plants diminutive to robust, 1.6-11.0 dm tall,
glabrous to pubescent, erect to sprawling, climbing if support is
available. Leaflets (4) 6—9 (12), (9) 14-25 (37) mm long, (0.9) 1.6—-3.4
(6.3) mm wide, the length/ width ratio (3.6) 5.8-11.2 (16.4), the apex
acute to truncate. Flowering peduncles (3.0) 10.0-32.0 (66.0) mm
long, usually about 1/2 as long, but varying to as long as or much
shorter than the length of the subtending leaf, the subtending leaf
(18) 27-47 (65) mm long. Fruiting peduncles (3) 33-50 (71) mm
long, from much shorter to as long as or longer than the subtending
leaf, the subtending leaf (22) 32-57 (75) mm long. Flowers 1-3,
rarely 4, occurring singly at points on the peduncles, or 2 flowers
sometimes arising from the same point; bluish. The standard some-
what broad and showy, the height of the reflexed standard (0.91)
1.12-1.38 (1.47) times the flower length from the calyx base to tip of
wings, the length of flowers from calyx base to tip of unreflexed
standard (4.3) 4.4-6.5 (7.5) mm. Upper calyx teeth (0.6) 0.7-1.1
(1.4) mm long, usually slightly shorter than the lower tooth, the
lower tooth (0.9) 1.1-1.5 (1.7) mm long, usually shorter than the
calyx tube, the tube (1.3) 1.6-2.0 (2.2) mm long. Styles (0.70)
0.85-1.10 (1.20) mm long, the apical (0.30) 0.45-0.65 (0.70) mm
pubescent. Mature legumes (15) 18-24 (26) mm long. Ovules up to
7. Seeds 3-7, usually 5 or 6, the hilum/circumference ratio in per-
cent (11) 17-22 (27).

RANGE AND HABITAT. Southern California and Baja California,
one distant collection from northern California seen (Figure 5).

Wooded areas; moist slopes, ravines, and canyons; foothills;
along creeks; chapparal; beaches; dry wasteland.

Sand, rocky soil.

PHENOLOGY. Flowering in (February-March) April-May, fruit-
ing in (February) April-May (June).

Discussion, Race 3 is almost identical morphologically with
race 2. However, geographical separation between the two races is

1984] Lassetter — Vicia 495

Figure 5. Distribution of Vicia /udoviciana subsp. ludoviciana, races 3 and 4,
Triangles = race 3, circles = race 4.

great (Figures 4 and 5) and climatic conditions differ markedly
between these two geographical areas. For these reasons, the two
races are recognized separately.

Race 3 plants are west coast populations of The United States
and Baja California and traditionally have been known as Vicia
exigua var. exigua in part (see Lassetter, 1975) of various western
authors, and V. exigua var. exigua sensu Hermann (1960) in part.

Typical race 3 plants have 1-3 flowers usually arising at separate
points on the peduncle, but contiguous flowers are often found,

Race 3 is almost completely absent north of the Santa Barbara
area (Figure 5), but is sympatric with Vicia hassei (Lassetter, 1975)
in southern California. No intergradation between the two occurs.
Race 3 and race 4 intergrade in the western portion of the range of
the latter.

496 Rhodora [Vol. 86

Subsp. ludoviciana race 4

DESCRIPTION. Plants diminutive to robust, 0.8-12.0 dm tall,
glabrous to pubescent, erect to sprawling, climbing if support is
available. Leaflets (5) 7-9 (11), (7) 10-17 (39) mm long, (1.0) 1.4—2.2
(4.2) mm wide, the length/ width ratio (3.4) 5.9-9.7 (16), rarely up
to 28 times as long as wide, the apex acute to truncate. Flowering
peduncles usually short, (0.9) 4.0-11.0 (47.0) mm long, usually
about |/3 the length of the subtending leaf, the subtending leaf (10)
23-41 (60) mm long. Fruiting peduncles (1) 8-36 (47) mm long,
about 1/2 or less the length of the subtending leaf, the subtending
leaf (22) 28-67 (71) mm long. Flowers 1—-5, averaging 1-3, usually
only one flower occurring at any one point on the peduncle,
pinkish-white to light lavender or bluish, the keel sometimes with a
blue spot at apex, the standard not broad or showy, the height of
the reflexed standard (.62) .67—.78 (1.02) times the flower length
from calyx base to tip of wings, the flower length from calyx base to
tip of unreflexed standard (4.8) 6.1—7.4 (8.4) mm. Upper calyx teeth
(0.5) 0.7-1.0 (1.7) mm long, about subequal with the lower tooth,
the lower tooth (0.9) 1.3-1.9 (2.2) mm long, slightly shorter than the
calyx tube, the tube (1.4) 1.8-2.1 (2.4) mm long. Styles (0.90)
1.05—1.30 (1.70) mm long, the apical (0.40) 0.50-0.75 (0.85) mm
pubescent. Mature legumes (20) 21-24 (26) mm long. Ovules up to 8
(rarely 9). Seeds 5-9, usually 7, the hilum/circumference ratio in
percent (8) 11-18 (21).

RANGE AND HABITAT. Western Texas and the Oklahoma pan-
handle, west and north to Colorado, Utah, New Mexico, Arizona,
extreme southeastern Nevada, and extreme southeastern Califor-
nia (Figure 5).

Woodlands; rocky slopes and hillsides; canyons and dry washes;
stream and creek beds; plains and grasslands; desert chaparral;
roadsides; waste areas; fence rows; lawns; ditches.

In sand, limestone, gravel, calcareous clay loam, sandy loam,
sandstone rubble, lava rock seams.

PHENOLOGY. Flowering in (late February) March-April (early
May), fruiting in (late March) May-June (early August).

Discussion. Race 4 encompasses phenotypes which are best
known traditionally as Vicia producta. Other taxa which are also

1984] Lassetter — Vicia 497

included in race 4 are V. thurberi Watson, V. exigua var. exigua
sensu Hermann (1960) in part, V. /eavenworthii var. occidentalis
sensu Shinners (1948), and V. /eavenworthii var. occidentalis sensu
Turner (1959) in part.

Typical plants have one to four flowers on peduncles which are
rather short at fruiting. Individuals with short peduncles are typical
in xeric microhabitats. More robust individuals with longer pedun-
cles can be found throughout the range (Figure 5). Flowers usually
occur singly at points on the peduncle, and the fltower-length/
standard-height ratios are rather low.

Race 4 is fairly well separated from other races throughout most
of its range, but in southwestern Texas is sympatric with and inter-
grades into races 2, 5, and 6. Race 4 plants with long peduncles
appear in the Chisos Mountains and continue southeast down the
Rio Grande Valley where flowers progressively become contiguous;
race 4 there intergrades into race 2. Going northeast from the Chi-
sos and Davis Mountains, standards of race 4 become more showy
and flowers more numerous as intergradation into race 5 occurs. A
similar transition occurs in western Oklahoma. Some race 4 speci-
mens blend into and closely resemble five- or six-flowered race 6
plants. Kinney County, Texas is an area where a great diversity of
all these types of intergrading forms can be observed.

At the western edge of its range, race 4 intergrades with race 3.
Calyx tube length of the two is about the same, and flowers are
usually one or two per peduncle, but three-flowered plants in both
races are not uncommon.

Subsp. ludoviciana race 5

DESCRIPTION. Plants diminutive to robust, 1.2-8.3 dm tall,
glabrous to pubescent, erect to sprawling, climbing if support is
available. Leaflets (6) 7-10 (13), (6) 9-17 (26) mm long, (0.9) 1.6—3.8
(8.0) mm wide, the length/ width ratio (2.2) 4.5—7.5 (10.0), the apex
acute to truncate or emarginate. Flowering peduncles (2.5) 30.0-60.0
(91.0) mm long, usually about as long as or longer than the subtend-
ing leaf, the subtending leaf (12) 28-62 (94) mm long. Fruiting
peduncles (15) 34-96 (150) mm long, about as long or longer than
the subtending leaf, the subtending leaf (21) 35-65 (82) mm long.
Flowers 1-17, averaging 5-15, solitary or in pairs at points on the

498 Rhodora [Vol. 86

peduncle, mostly separated from each other when fruit is present,
pinkish-white to deep lavender-purplish, occasionally white, the
keel often with a blue spot at the apex, the standard broad and
showy, the height of the reflexed standard (0.65) 0.69-1.06 (1.45)
times the length of the flower from calyx base to tip of wings, the
length of flowers from the calyx base to tip of unreflexed standard
(5.3) 6.0-7.3 (9.5) mm. Upper calyx teeth (0.7) 1.1-1.6 (2.6) mm
long, slightly shorter than the lower tooth, the lower tooth (1.2)
1.5-2.1 (3.0) mm long, about as long as to longer than the calyx
tube, the tube (1.3) 1.6-2.0 (2.5) mm long. Styles (1.0) 1.2-1.4 (1.6)
mm long, the apical (0.55) 0.60-0.80 (1.10) mm pubescent. Mature
legumes (19) 20-25 (30) mm long. Ovules up to 8. Seeds 5-7, usually
6, hilum/ circumference ratio in percent (14) 19-23 (29).

RANGE AND HABITAT. Central Texas and southern Oklahoma,
plus two distant collections from southern New Mexico (Figure 4).

Open woods; river bottoms; rock seams and rocky slopes; prair-
ies; railroad and highway rights-of-way; waste areas: lawns and
vacant lots.

In granitic and limestone soils, serpentine and travertine, sand,
sandy loam,silty clay, rocky limestone soil, fine gravel.

PHENOLOGY. Flowering in (late February) March-April (early
May), fruiting in April-May.

Discussion. Typical members of race 5 are characterized by
large, showy standards and flower-length/standard-height ratios of
(0.65) 0.69-1.06 (1.49), and are typified by Vicia ludoviciana var.
laxiflora sensu Shinners (1948), Turner (1959), and Hermann
(1960). Calyx teeth are typically relatively long and nearly subequal.
The largest and showiest flowers of the entire complex are found in
race 5 individuals from Llano County and a few other scattered
locations in Texas, and from the mountainous areas of Oklahoma.
Race 4 flowers are about the same length, but the standard is not as
high as race 5, and therefore not as showy.

The major distribution of this race (Figure 4) appears to be three
discrete areas with two distinct gaps in Texas. Although it is possi-
ble that the gaps reflect an artifact of incomplete collecting, I feel
this explanation is not very probable since central Texas is a rather
well-collected area. I have not been able to correlate the breaks in
the distribution with geographical regions or with vegetation areas.

1984] Lassetter — Vicia 499

This race overlaps in range with and intergrades into races I, 4, 6,
and 2. In the eastern part of its range, race 5 intergrades into race 1,
and in the western and southwestern part, a change to race 4 occurs.
Few-flowered forms in central Texas often resemble race 6, and in
southern Texas, few-flowered forms grade into race 2 and race 2-4
intermediates.

B. Vicia ludoviciana Nutt. ex T. & G. subspecies leavenworthii (T.
& G.) Lassetter & Gunn (as stat. nov. in USDA Tech. Bull. No.
1601: 16. 1979.)

Vicia leavenworthii T.& G. Fl. N. Amer. 1: 271. 1838. HOLOTYPE Ny!: Dr.
Leavenworth. Arkansas.
Vicia leavenworthii var. typica Shinners. Field and Lab. 16: 22. 1948.
Cracca erotanthos Alefeld. Bonplandia 9: 118. 1861.

DEscRIPTION. Plants diminutive to robust, 1.0-12.0 dm or more
tall, glabrous to pubescent, erect to sprawling, or climbing if sup-
port is available. Leaflets (7) 11-14 (17), (5) 10-23 (25) mm long,
(1.2) 2.2-9.4 (11.0) mm wide, the length/ width ratio (2.1) 2.2-5.0
(6.9), the apex acute to emarginate. Flowers opening before pedun-
cles and internodes elongate, the flowering peduncles (0.8) 3.0-11.0
(28.0) mm long, usually much shorter than the subtending leaf, the
subtending leaf (9) 20-65 (69) mm long. Fruiting peduncles (12)
26-70 (91) mm long, usually shorter than the subtending leaf, the
subtending leaf (19) 30-92 (105) mm long. Flowers 1~6, pinkish
white to light lavender, the standard not broad or showy, the height
of the reflexed standard (0.57) 0.63-0.78 (1.04) times the flower
length from calyx base to the tip of wings,length of flower from
calyx base to tip of unreflexed standard (4.5) 5.1-6.8 (7.5) mm.
Upper calyx teeth (0.5) 1.1-1.6 (2.0) mm long, shorter than to
almost as long as the lowest tooth, the lowest tooth (1.0) 1.4-2.0
(2.2) mm long, and about as long as the calyx tube, the tube (1.3)
1.6-1.9 (2.2) mm long. Styles (0.40) 0.65—0.80 (0.90) mm long, the
apical (0.20) 0.30—-0.40 (50.0) mm pubescent. Mature legumes (18)
21-33 (35) mm long. Ovules up to 8, seeds 4-8, the hilum/circum-
ference ratio in percent (15) 18-49 (S55).

NOMENCLATURAL HISTORY AND DISCUSSION OF TYPE SPECIMENS.
The holotype of Vicia leavenworthii is good, typical and traditional
V. leavenworthii, and is a good example of race 6.

500 Rhodora [Vol. 86

There is little doubt that Cracca erotanthos of Alefeld (1861) is
the same as race 7 of this study. Alefeld stated that he obtained
specimens from a Dr. Hexamer in New York and that Hale had
collected the plant in Louisiana. There is an 1840 collection by Hale
at NY. This sheet bears no locality designation but is race 7, which
is centered in Louisiana. While it is possible (or probable) this
represents the gathering seen by Alefeld, evidence and necessity
scarcely warrant designating it a lectotype.

Alefeld specifically mentioned the fact that Cracca erotanthos
does not fit the description of Vicia ludoviciana (Nutt. ex T. & G..
1838), and that it should not be confused with V. /udoviciana, as
Cracca erotanthos possesses a long hilum. The hilum/circum-
ference ratio of race 7 is by far the largest of the complex (Lassetter,
1978a).

The specimen at PH gathered by Tainturier from Louisiana,
which was mentioned earlier, has been considered a type of Vicia
ludoviciana. This is race 7 of this study [V. /udoviciana Nutt. ex T.
& G. subsp. leavenworthii (T. & G.) Lassetter and Gunn], and
should not be thought to be representative of the typical subspecies.
It is typical race 7 of subsp. /eavenworthii.

Subsp. leavenworthii race 6

DESCRIPTION. Plants diminutive to robust, 1.0-10.0 dm tall.
Leaflets (7) 11-14 (17), (5) 10-15 (20) mm long, (1.2) 2.2-4.1 (6.3)
mm wide, the length/ width ratio (2.5) 3.4-5.0 (6.9), the apex acute
to retuse. Flowering peduncles (0.8) 3.0-11.0 (28.0) mm long, usu-
ally much shorter than the subtending leaf, the subtending leaf, (9)
20-41 (57) mm long. Fruiting peduncles (12) 26-56 (91) mm long,
slightly shorter than to longer than the subtending leaf, the subtend-
ing leaf (19) 30-50 (63) mm long. Flowers |-6, averaging 2-4, most
often one upper flower and 2 or 3 flowers arising from a common
point below, or when 2 flowered, the 2 flowers arising from the
same point. Upper calyx teeth (0.5) 1.1-1.6 (2.0) mm long, almost as
long as the lower tooth, the lower tooth (1.0) 1.4-1.8 (2.1) mm long,
about as long as the calyx tube. Mature legumes (18) 21-24 (29) mm
long, the seed hilum/circumference ratio in percent (15) 18-22 (27).

RANGE AND HABITAT. Central Texas, Oklahoma, extreme north-
western Arkansas, extreme southwestern Missouri. (Figure 6)

1984] Lassetter — Vicia 501

Figure 6. Distribution of Vicia ludoviciana subsp. leavenworthii, races 6 and 7.
Circles = race 6, stars = race 7.

Open woods; wooded hills; limestone hills; cedar brakes; pas-
tures; creek banks, river banks, stream beds, and flood plains; prair-
ies; railroad and highway rights-of-way; waste ground; vacant lots
and lawns.

PHENOLOGY. Flowering in April-early May, fruiting in April-
May.

Discussion. Excluding race 7, this race is the easiest to recog-
nize, in spite of intergradation with several others. Typical race 6
usually has three or four pale flowers (one upper flower and two or
three flowers from a common point below), very short styles, and
numerous leaflets. Traditional taxa included in race 6 are Vicia

502 Rhodora [Vol. 86

leavenworthii var. typica (Shinners, 1948), V. leavenworthii var.
leavenworthii (Turner, 1959), and V. leavenworthii (Hermann,
1960).

Race 6 is centered in Texas in the Blackland Prairie (Figure 6) as
previous authors have stated. It also occurs on the Edwards Pla-
teau, but is absent from an area known as the Central Basin or
Central Texas Granite. Race 5, however, is prevalent on these soils
which are derived from granitic parent material. Race 6 plants are
most often found on clay loam soils with moderate amounts of
calcium (Lassetter, 1978b).

Race 6 blends into race 2 on the Gulf Coast, into race 4 in western
and southwestern Texas, and into race 5 in central Texas. It shares
many characteristics with race 7, but because the two are sympatric
only in McCurtain Co., Oklahoma, confusion should be minimal.

Subsp. leavenworthii race 7

DESCRIPTION. Plants usually robust, 3.0-12.0 dm or more tall.
Leaflets (10) 11-13 (14), (13) 15-23 (25) mm long, (4.5) 5.8-9.4
(11.0) mm wide, the length/ width ratio (2.1) 2.2-2.8 (3.1), the apex
acute to emarginate, usually rounded or retuse. Flowering pedun-
cles (3.0) 3.0-9.0 (12.0) mm long, much shorter than the subtending
leaf, the subtending leaf (20) 23-65 (69) mm long. Fruiting pedun-
cles (28) 30-70 (85) mm long, usually shorter than the subtending
leaf, the subtending leaf (63) 63-92 (105) mm long. Flowers 1-2,
from the same point or separate points on the peduncle. Upper
calyx teeth (1.1) 1.2-1.4 (1.7) mm long, shorter than the lower
tooth, the lower tooth (1.5) 1.8-2.0 (2.2) mm long, about as long as
the calyx tube. Mature legumes (29) 29-33 (35) mm long, seed
hilum/circumference ratio in percent (37) 41-49 (55).

RANGE AND HABITAT. Louisiana, McCurtain County, Okla-
homa, scattered sites in Arkansas and Mississippi (Figure 6),

Deciduous woods and bottomland forests: along wooded streams;
prairies and glades; fields; roadsides; levee banks.

Mostly in heavy (clay) soils, but also in sandy soils.

PHENOLOGY. Flowering in (late February) March-April (late
May), fruiting in late March-May.

Discussion. Race 7 is the most distinctive. Most plants are very
robust, have large ovate leaflets, only one or two flowers per pedun-

1984] Lassetter — Vicia 503

cle, large legumes, and the greatest hilum/circumference percent-
age. Plants included in race 7 have traditionally been recognized as
Vicia ludoviciana, not V. leavenworthii. This race includes V. /udo-
viciana var. typica (Shinners, 1948, in part), and V. /udoviciana var.
ludoviciana (Turner, 1959; Hermann, 1960) in part.

The distribution appears to be very limited (Figure 6); however,
more complete collecting in the northern Louisiana area would be
desirable in establishing the distribution more precisely. One 19th
century collection from Florida (no specific location) was seen, but
no recent material from that state was obtained.

Hermann’s (1960) distribution map of Vicia ludoviciana var.
ludoviciana contains a dot in east-central Mississippi. I presume
this is based on S. M. Tracy 1627, Starkville, Mississippi (MISSA,
US). Labels of several duplicates of this collection are marked
“cultivated”, and one sheet, US #67561, bears the caption “Native
of Louisiana”. This collection is race 7, and represents an introduc-
tion outside its natural range.

The other Mississippi sites are thought to be natural sites.

The few-flowered, robust forms of race | found in Louisiana and
Mississippi resemble race 7, but no major problem should exist in
distinguishing between the two. The relationship with race 6 is pre-
viously mentioned, and both races 6 and 7 (subsp. /eavenworthii)
can easily be separated from the 5 races of subsp. /udoviciana,
because in subsp. /eavenworthii the flowers open before the pedun-
cles elongate.

ACKNOWLEDGMENTS

I express sincere gratitude to Dr. Duane Isely for his suggestion
of the initial dissertation problem, and his guidance throughout this
study, which represents a portion of a dissertation submitted as
partial fulfillment of the Degree of Doctor of Philosophy in Botany,
lowa State University, Ames, lowa. 1972. The facilities of the lowa
State Herbarium, supported by the Science and Humanities Re-
search Institute, were used in the preparation of this paper. The
lowa State University Agriculture and Home Economics Experi-
ment Station provided funds for laboratory supplies and some field
travel. My graduate work was supported by an NSF Traineeship,
and I was the recipient of NSF Grant GB-27935 (Grants for
Improving Doctoral Dissertations in the Field Sciences). Extensive

504 Rhodora [Vol. 86

field travel was made possible by this grant, which also provided
additional financial support. I express appreciation to the curators
of the Herbaria mentioned herein for their generosity in allowing
use of herbarium facilities, for arranging loans, and for permis-
sion to take seed samples. The following persons collected seed for
me: James M. Lang, John Lassetter, William F. Mahler, Ingrid L.
Marin, and Robert M. Stuart. Clare B. Hardham and the late
Ernest C, Twisselmann provided sound advice during my field work
in California. William Warde, formerly at lowa State University,
and Bruce Lewis, formerly at Eastern Kentucky University, assisted
in computer handling of data.

LITERATURE CITED

ALLARD, R. W. AND P. L. WoRKMAN. 1963. Population studies in predominately
self-pollinated species. IV. Seasonal fluctuations in estimated values of gentic
parameters in lima bean populations. Evolution 17: 470-480.

JS. K. JAIN, AND 1968. The genetics of inbreeding populations.
In: Advances in genetics. Vol. 14. pp. 55-131. Academic Press, New York,
Bart, P.W. 1968. Vicia. In: Tutin, T. G., V. H. Heywood, N. A. Burges, D. M.
Moore, D.H.Valentine, S. M. Walters, and D. A. Webb, editors. Flora Euro-

peae Vol. 2. pp. 129-136. Cambridge Univ. Press, Cambridge.

BALLARD, A. T., AND T. BUCHWALD. 1971. A viability test for seeds of townsville
stylo using thiourea. Australian J. Exp. Agric. Animal Husb. | 1: 207-210.
Bowen, C.C. 1956. Freezing by liquid carbon dioxide in making slides perma-

nent. Stain Technology 31: 87-90.

Corrett, D. S.. AND M. C. JouNston. 1970. Manual of the vascular plants of
Texas. Texas Research Foundation, Renner.

DARLINGTON, C. D., AND A. P. Wyciz. 1955. Chromosome atlas of flowering
plants. 2nd edition. George Allen and Unwin, Limited. London.

Feporov, AN. A., Ed. 1969. Chromosome numbers of flowering plants. V. L.
Komarov Botanical Institute, Leningrad.

GuNN,C.R. 1970. Seeds of the tribe Vicieae (Leguminosae) in North American
Agriculture. Proc. Assoc. Off. Seed Ana. 60: 48-70.

1979. Genus Vicia with notes about tribe Viciaea (Fabaceae) in Mexico
and Central America. USDA Tech. Bull. 1601. U. S. Govt. Print. Off., Wash.,
D:C,

HERMANN, F. J. 1960. Vetches of the United States—native, naturalized, and
cultivated. USDA Agricultural Handbook 168, U. S. Govt. Print. Off., Wash.,
D.C.

HOLMGREN, P. K. AND W. KEUKEN. 1974. Index Herbariorum. Part 1, The Her-
baria of the World. Oosthoek, Scheltema and Holkema, Utrecht.

HOLLINGS, E. AND C. A. Stace. 1974. Karyotype variation and evolution in the
Vicia sativa aggregate. New Phytologist 73: 195-208.

Howe tt, J. T. 1949. Marin Flora. Univ. California Press. Berkeley. (Ed. 2,
1970).

1984] Lassetter — Vicia 505

LASSETTER, J. S. 1972. A biosystematic study of the Vicia ludoviciana complex
(Leguminosae). Ph.D. Dissertation. lowa State University, Ames. Microfilm
73-9458. University Microfilms, Ann Arbor, Mich.

1975. Taxonomic status of Vicia hassei (Leguminosae). Madrono 23:

73-78.

1978a. Seed characters in some native American vetches. Sida 7:

255-263.

1978b. Edaphic relationships in the Vicia /udoviciana complex (Legumi-

nosae). lowa State Jour. Res. 53: 13-20.

~AND C.R.GuNN. 1979. Vicia menziesii Sprengel rediscovered: its taxo-
nomic relationships. Pacific Science 33: 85-101.

Levitzky,G. A. 1931. The karyotype in systematics. Trudy Prikl. Bot. 27-1, No.
1: 220-256.

METTIN,. D. AND P. HANELT. 1968. Bemerkungen zur Karyologie und Systematik
einiger Sippen der Gattung Vicia L. Feddes Repert. 77-1: 11-30.

Newcomer, E. H. 1953. A new cytological and histological fixing fluid. Science
118: 161.

PLITMANN, U. 1967. Biosystematical study in the annual species of Vicia of the
Middle East. Ph.D. Dissertation, The Hebrew University of Jerusalem, pub-
lished by the author.

Rous, A. 1961. Cytotaxonomical studies on Vicia cracca L. and Vicia tenuifolia
Roth. I. Chromosome numbers and karyotype evolution. Hereditas 47: 81-110.

SENN. H. A. 1938. Chromosome number relationships in the Leguminosae. Bib-
liographica Genetica 12: 175-336.

Suinners. L. H. 1948. The vetches and pea vines (Vicia and Lathyrus) of Texas.
Field Lab. 16: 18-29.

SHRIVASTAVA, L.M. 1963. Cytogenetical studies in certain species of Vicia. Cyto-
logia 28-2: 154-169.

STANKEVICZ, A. K. 1970. On clarification of the Vicia L. genus systematics.
Trudy Prikl. Bot. 43: 110-125.

STEBBINS. G. L., JR. 1950. Variation and evolution of plants. Columbia Univ.
Press, New york.

1958. Longevity, habitat, evolution, and the release of genetic variability
in the higher plants. Cold Spring Harbor Symp. Quan. Bio. 23: 365-378.

1971. Chromosomal evolution in higher plants. Addison-Wesley Pub-
lishing Company, Reading, Mass.

SVESHNIKOVA, I. N. 1927. Karyological studies on Vicia. Trudy Prikl. Bot. 17-1.
No. 3: 37-72.

TurNER. B. L. 1956. Chromosome numbers in the Leguminosae. |. Am. J. Bot.
43: 577: S81.

1959. The legumes of Texas. Univ. Texas Press, Austin.

VEERASETHAKUL, S. AND J. S. LASSeTTER. 1981. Karyotype relationships of native
New World Vicia species (Leguminosae). Rhodora 83: 595-606.

APARTADO AEREO #18512
CALI, VALLE
COLOMBIA, AMERICA DEL SUR

A REVISION OF THE GENUS KYLLINGA ROTTB.
(CYPERACEAE) IN MEXICO AND CENTRAL AMERICA

GORDON C. TUCKER

ABSTRACT

A revision based on herbarium study of the six species of Ai //inga occurring in
Mexico and Central America is presented. Each species is described and discussed.
Distribution maps, a key to the species, and a detailed discussion of taxonomically
useful characters in the genus are included.

Key Words: Cyperaceae, Ky //inga, Mexico, Central America

The genus Ky //inga consists of some 40 to 45 species (Kiikenthal,
1935-36; Lye, 1981). Nearly all are tropical, with no more than ten
occurring in temperate regions. The greatest diversity is in tropical
Africa and Madagascar, where as many as 35 species are found. In
the New World, there are eight species, two of which, K. brevifolia
and K. odorata, are pantropic. Three of the species occurring in the
New World, K. odorata, K. pumila and K. brevifolia, occur in the
United States (Delahoussaye and Thieret, 1967). One species, K.
nudiceps, is endemic to Cocos Island, Costa Rica.

Some workers include Ky//inga as a subgenus of Cyperus. How-
ever, Ky/linga differs from Cyperus in its two-scaled, one-flowered
spikelets. The dense, sessile spikes of Ai/linga give the plants a
different aspect from most species of Cyperus with their usually
open, branched inflorescences. The majority of recent specialists on
Cyperus and on the Cyperaceae have recognized Ky /linga as a dis-
tinct genus. Some proponents of Ky //inga as a distinct genus include:
Standley and Steyermark (1958); Koyama (1978) in the Flora of the
Lesser Antilles; Raynal (1973) who worked at different times on the
Cyperaceae of Africa, Madagascar and New Caledonia; Vorster
unpubl. Ph.D. dissertation, U. Pretoria, 1978 who studied the
southern African species, as well as the genus Cyperus; and Lye
(1981) who studied the East African Cyperaceae.

Raynal (1973) hypothesized that Ky//inga probably represented
an advanced group derived from Cyperus. He based this conclusion
on a similarity of habit of certain African species of Section Maris-
cus (Vahl) Bentham of Cyperus to that of some species of Ky /linga.
I know of no other evidence to support Raynal’s conjecture, but am

507

508 Rhodora [Vol. 86

willing to agree that Ky//inga is closely related to Cyperus. A satis-
factory understanding of the phylogenetic relationships of Ky/linga
must await adequate study of the African species.

Several regional treatments have been published for Mexico and
Central America. O’Neill (1940) provided a treatment of the three
species (Ky/linga pumila, K. brevifolia and K. tibialis) occurring
in the Yucatan Peninsula, which included the Mexican states of
Campeche, Yucatan and Quintana Roo, the Guatemalan depart-
ment of Petén, and all of Belize. Ayers (1946) included K. odorata,
K. pumila and K. brevifolia in his taxonomic treatment of the genus
Cyperus in Mexico. Standley and Steyermark (1958) included K.
pumila, K. brevifolia, K. vaginata and K. tibialis in their account of
the Cyperaceae of the Flora of Guatemala. However, they over-
looked K. odorata, misdetermining the three sheets of it at Fas “K.
pumila” or “K. brevifolia.” Svenson (1943) treated K. pumila, K.
brevifolia, K. odorata and K. tibialis in the Flora of Panama; the
present treatment adds one species, K. vaginata, to the flora of that
country. Standley (1938) included all six species of the present
treatment in his Flora of Costa Rica. Spellman etal. (1975) listed for
Belize only two out of the five species found there in the present
treatment. Molina R. (1975) omitted K. odorata from his enumera-
tion of the plants of Honduras. Standley (1931) recognized six spe-
cies of Ky/linga in his account of the Cyperaceae of Central
America.

Need for the present treatment of this genus is shown by lack of
any recent study covering the entire area from Mexico to Panama.
Many of the treatments made for a particular single country, while
useful, usually are based on few collections. Unless specializing in
the genus or family in question, an author usually does not see
enough collections to observe and describe adequately the variation
within each species. Further, species represented in a country by
only a very few collections may be overlooked because a non-
specialist has not had sufficient opportunity to learn to recognize
the species. About 40% of the specimens seen during the present
study had been misidentified, a state of affairs which emphasizes the
need for an up-to-date revision of the genus for the region.

The present treatment has been made in conjunction with a
taxonomic revision of Cyperus in Panama and Costa Rica (Tucker,
1983) and northern Central America and Mexico. Several compari-

1984] Tucker — Kyllinga 509

sons can be made between the two genera, Ky/linga and Cyperus. In
general, the author has found fewer taxonomically useful characters
in Ky/linga as compared with Cyperus, owing in large part to the
differences in inflorescence structure between the two genera. The
inflorescences of Ky/linga, composed of a sessile spike or spikes,
lack the rays, branches and various arrangement of spikes and spi-
kelets that are useful in recognizing species in Cyperus. Also, the
two-scaled, one-flowered spikelets of Ky//inga show comparatively
little variation throughout the genus, whereas in Cyperus the spi-
kelets exhibit considerable variation in length, width, imbrication
of scales, and presence of rachilla wings, as well as others.

The habitats of the six species of Ky/linga treated here are in
general very similar. Label data indicate they are plants of open,
often disturbed and usually moist situations. Such habitats include
roadside ditches, pastures, marshes but not wooded swamps, newly
graded roadbanks, gravel pits, river and lake shores, and thickets.
In addition, K. odorata occurs in pine and oak forests, and in thorn
woodlands. The most specific in habitat is K. tibialis. It is always
found close to the Caribbean on coastal dunes, under littoral trees
and shrubs, and in swales behind the dunes.

The present treatment includes a key designed to be useable on
specimens with or without underground parts. The descriptions are
based on examination of some 600 herbarium specimens from the
institutions listed in the Acknowledgments. A summary of the dis-
tribution and phenology follows each species description, as well as
a listing of representative specimens. For the rare species, Ky //inga
vaginata and K. nudiceps, all specimens seen are cited in detail. A
citation of all specimens examined for this study is made in the
Index of Exsiccatae.

COMMENTS ON TAXONOMICALLY USEFUL CHARACTERS

RHIZOMES. Five of the six species of the present treatment are
rhizomatous. Only Ky/linga pumila lacks rhizomes, being instead
densely cespitose. In K. nudiceps, K. tibialis and K. vaginata, the
rhizomes are always conspicuous provided that underground parts
have been gathered intact. In K. brevifolia, the internodes of the
rhizome are sometimes very short, about | mm in length, thus
producing a tuft of approximate culms that suggests the cespitose
species K. pumila. Such tufted plants of K. brevifolia will often also

510 Rhodora [Vol. 86

produce elongate rhizomes with the culms spaced 5-15 mm apart.
Tufted plants of K. brevifolia can be distinguished from K. pumila
by their longer anthers (0.8-1.1 mm vs. 0.2-0.4 mm) and the vertical
orientation of the longest inflorescence bract in K. brevifolia; the
longest inflorescence bract in K. pumila is horizontal to reflexed.

LEAVES. All species except Ky/linga nudiceps have been observed
with leaf blades. Three species, K. brevifolia, K. pumila and K.
odorata always bear leaf blades, and these leaf blades are linear-
lanceolate with acute apices. The remaining two species, K. vagi-
nata and K. tibialis, usually lack leaf blades. In these two species,
the leaf blades, when present (on about 1/3 of the collections seen in
this study), are linear, with the apex broadly rounded and usually
mucronate. The presence or absence of leaf blades has been a source
of confusion between K. tibialis and K. vaginata (see comments
under K. vaginata).

INFLORESCENCES. The inflorescence of Ky/linga consists of 1-4 ses-
sile spikes subtended by 2-5 usually leaf-like bracts. The shape and
number of the spikes is useful in dividing the Mexican and Central
American species into two major groups, as indicated in the Key to
the Species. The inflorescence bracts are an easily observed charac-
ter useful in several instances. For example, the extreme reduction
of the bracts in K. nudiceps separates this species from K. vaginata
and K. tibialis. The erect orientation of the longest inflorescence
bract in K. brevifolia separates that species from K. pumila and K.
odorata, in which all the bracts are borne horizontally to reflexed.
The length of the inflorescence bracts relative to the length of the
spike has been a source of confusion between K. tibialis and K.
vaginata (see comments under K. vaginata).

RACHIS. The rachis is that portion of the summit of the culm along
which the spikelets are borne. In most species it is cylindrical to
conical. A notable exception is Ky/linga tibialis, in which the rachis
is a low dome-shaped to nearly spherical structure that is a distinc-
tive character of this species. The shape of the rachis might seem a
difficult or tedious character to observe, but such is not the case. In
collections with several culms from a rhizome system, there is
nearly always at least one older culm from which the spikelets have
fallen, revealing the rachis. Shape of the rachis is the most reliable
character separating K. tibialis from K. vaginata (see comments
under K. vaginata).

1984] Tucker — Kyllinga 511

SPIKELETS. Spikelets show little morphological variation in the
genus Ky /l/inga, at least as it is represented in the New World. The
kind of variation that does occur is mostly in dimensions of the
spikelet. These differences have not been of much value in distin-
guishing related species or groups of species.

SCALES. Some variation in dimensions and in number of nerves
has been noted, but none of this variation has proven to be of
sufficient magnitude to distinguish species of Ky/linga. Color of
scales is useful in one instance: the whitish scales of K. odorata
distinguish it from other New World species which all have pale
greenish, brownish or hyaline scales.

Presence of spinulose teeth on keels of the scales has been a
source of confusion in descriptions of species. All the species of this
treatment, with the exception of Kvy/linga nudiceps, have been
observed to have either smooth or spinulose-scabrellate keels.
Plants of the same collection, and occasionally spikelets of the same
plant, may have both smooth and scabrellate keels. Ayers (1946)
used the character of smooth versus scabrellate keels to distinguish
K. odorata from K. brevifolia and K. pumila. All three of these
species have been observed either with smooth or with scabrellate
keels. Thus Ayer’s observation is not a reliable means of recogniz-
ing K. odorata.

STAMENS. A survey of the published descriptions of Ky //inga spe-
cies usually reveals more than one count of the number of stamens
per flower. For example, K. brevifolia has been reported to have
one stamen per flower (Delahoussaye & Thieret, 1967); 3 to I sta-
mens (Kiikenthal, 1935-36; O’Neill, 1940); and 1-2 (~3) stamens
(Kern, 1974). In the present study, about two-thirds of the flowers
of K. brevifolia checked had 2 stamens; the remainder had a single
stamen. These various reports suggest that stamen number ts prob-
ably not a useful character in delimiting species.

The length of the anthers has been found to be a genuinely help-
ful and reliable character. In a collection with several to many
culms, usually in at least one inflorescence spike the anthers will be
exserted, and can thus be readily observed and measured. The small
anthers of K. pumila provide a quick and reliable means of distin-
guishing that species from K. brevifolia (see key and comments
under K. pumila).

512 Rhodora [Vol. 86

ACHENES. Delahoussaye and Thieret (1967) stated that specimens
of Ky/linga ought to be collected with mature achenes for critical
determination. It is my observation that, in general, the achenes are
less important to the taxonomy of Ky//inga than in any genus of the
Cyperaceae with which I am familiar. O’Neill (1940) stated that K.
brevifolia and K. pumila differ in achene width, but the present
study cannot support this claim (see comments under K. brevifolia).

Delahoussaye and Thieret (1967) observed that the dark achenes
of Ky/linga odorata with their contrasting whitish stipitate bases
and apiculi are distinctive of the species. This is indeed a reliable
means of recognizing this species, but the achenes must fully mature
before the coloration pattern is apparent. This is the only instance
in which the writer has found mature achenes to be helpful in
determination. However, ripe achenes are not essential for critical
identification since the whitish color of the scales, a characteristic
unique to this species in Mexico and Central America, is evident
even before anthesis.

Variation of length and width of achenes falls within a rather
narrow range, 0.5 and.0.4 mm, respectively. The differences in
dimensions are not sufficiently great to be useful or reliable in
delimiting species (see discussion under K. brevifolia).

NOTES ON MEASUREMENTS

1. Height of the plant is given as length of the culm from its base to
insertion of the inflorescence bracts. Height does not include
length of the spike or inflorescence bracts.

2. Width of leaves, culms and spikes is measured at mid-length.
Achenes are measured at the widest point.

3. Width of scales was measured at the widest point, in abaxial
view, of a detached, rehydrated scale pressed flat against a flat
surface. The width may be estimated as twice the distance from
keel to margin of an intact scale viewed laterally.

TAXONOMIC TREATMENT

Kyllinga Rottb., Descr. Icon. Rar. Nov. Pl. 12. 1773. (nom. et
orthogr. cons.). TyPE species: K. brevifolia Rottb.

Cyperus subgenus Kvy/linga (Rottb.) Valck. Sur., Gesl. Cyp. Mal. Arch. 42.
1898.

1984] Tucker — Kyillinga 513

Herbaceous cespitose annuals or more frequently rhizomatous
perennials. Culms smooth, trigonous or roundly trigonous, (5-)
20-50 (—100) cm tall, 0.5-3 mm thick. Leaves with blades (1—) 5-25
cm long, 1-5 mm wide, the margins and keel scabrellate; or leaf
blades lacking, the base of the culm with I-5 scarious-margined
brownish sheaths. Inflorescence bracts 2-4, leaflike, up to 20 cm
long and 6 mm wide, or greenish brown to brown, reduced, shorter
than to equalling the spike. Inflorescence of 1-4 densely cylindrical
to ovoid or spherical sessile spikes 6-15 mm long, 4-10 mm wide.
Rachis slenderly cylindrical, conical or very nearly spherical. Spi-
kelets usually very numerous, not readily distinguishable to the
unaided eye, (1S—) 50-150, densely packed on the rachis, ovate to
lanceolate, (1.8) 2.2-3.8 (4.5) mm long, (0.7) I1-1.2 (1.4) mm wide,
consisting of four scales: the two basal scales minute, brownish,
sterile, 0.2-0.8 mm long; the two distal scales much longer, the
lower of these two (the third of the spikelet) subtending a bisexual
flower, the fourth scale slightly smaller, sterile, or infrequently bear-
ing | or 2 stamens only. The fertile scale ovate, mucronate or
mucronulate, (1.8) 2.4-3 (3.4) mm long, (0.8) 1.2-2 (2.6) mm wide,
laterally 2-4 nerved, hyaline, whitish or pale greenish or reddish
brown; keel greenish or whitish, smooth or spinulose-scabrellate,
1-3 nerved. Stamens |, 2 or 3 per flower; filaments ribbon-like,
reddish brown; anthers oblong-elliptic to linear, (0.2) 0.3-1.1 (1.3)
mm long. Styles (0.4) 0.8-1.2 (1.8) mm long; stigmatic branches 2,
(0.3) 0.6-1.5 mm long. Achenes lenticular, laterally flattened, nar-
rowly ovate to oblong or elliptic, 1—1.2 (1.5) mm long, (0.4) 0.6-0.8
mm wide, apiculate, base cuneate to rounded, substipitate to de-
cidedly stipitate, puncticulate, surface light brown to dark brown or
reddish or brown.

KEY TO SPECIES OF KYLLINGA IN MEXICO AND CENTRAL AMERICA

1. Culms densely cespitose, rhizome lacking; anthers 0.2-0.4 mm
ROE 6645 babe do eatewaeseen5es acne ueees 1. K. pumila.

1. Culms approximate to widely spaced on a usually conspicuous
rhizome: anthers 0.4-1,1 mm lone. éccivcveccasesenaxes (2)

2. Longest inflorescence bract 4-10 times longer than the longest
spike; spikes 1-4, ovoid to densely cylindrical......... (3)

3. Scales whitish; achenes dark with contrasting whitish stipi-
tate base and apiculus; rhizome short, knotted, culms

514 Rhodora [Vol. 86

approximate; inflorescence bracts all horizontal to re-
flexed; culms bulbous-thickened at the base ...........
(PRESIUES Hh DRE eS EC LES wae ae eee eS 2. K. odorata.
3. Scales hyaline, pale brownish to greenish; achenes uniform
light to medium brown; rhizome elongate, the culms (1-)
5-15 mm apart; the longest inflorescence bract erect,
looking like a continuation of the culm, the other bracts
ascendent to horizontal; culms slender at base .........
eC eee Tee Te er eee TT er eer eee 3. K. brevifolia.
2. Longest inflorescence bract at most 3 times longer
than the longest spike; spikes solitary (rarely a much
smaller sessile one borne at the base of the spike), spherical
to SUBPIODOSe Or HEMIsDNerICal .c4u00024s0ccussnnvns (4)
4. Culms 0.8-2 mm thick at the apex, leafless or with leaf
blades 2-5 mm wide; spikelets about 75-200; rhizomes
horizontal, (2-) 3-5 mm thick; inflorescence bracts
lanceolate, equalling or exceeding the spike ........ (5)
5. Spikes subglobose to ovoid; denuded rachis
cylindrical to conical; scales of rhizome lanceolate....
eS ee ee eee ee ete ee ee 4. K. vaginata.
5. Spikes spherical to globose; denuded rachis spherical to
hemispherical; scales of rhizome ovate-lanceolate .
ee ee Ce ee ee Cee ee ee 5. K. tibialis.
4. Culms 0.4—-0.9 mm thick at the apex; always leafless; spike-
lets 15-45; rhizomes oblique, 0.5-3 mm thick; inflores-
cence bracts broadly ovate, shorter than the spike......
ee er re ee ee re eee eee 6. K. nudiceps.

TREATMENT OF THE SPECIES

1. Kyllinga pumila Michaux, Fl. bor.-amer. 1: 28. 1803. Type:
U.S.A., southern Illinois, August 1795, Michaux (HOLOTYPE: P,
Herb. Michaux, photo, DUKE; ISOTYPE: P, Herb. E. Drake,
photo, DUKE).

K. caespitosa Nees, Fl. bras. 2: 12. 1842. Type: BRAzIL. Bahia, Martius s.n.
(SYNTYPE: M!); Minas Gerais, Martius s.n. (SYNTYPE: M!); Para, Martius s.n.
(SYNTYPE: M!); Rio Nigri, Martius s.n. (SYNTYPE: M!).

K. caespitosa var. major Nees, Fl. bras. 2: 12. 1842. Type: BRAZIL. Se/low (not
located).

K. laxa Schrad. ex Nees, Fl. bras. 2: 14. 1842. Type: BRAZIL. Rio de Janeiro,
Sebastianopolis, Martius s.n. (HOLOTYPE: M!).

1984] Tucker — Kyllinga 515

K. pumila B b. elatior Kunth, Enum. pl. 2: 132. 1837. Type: TRINIDAD. Sieber 2
in 1825 (HOLOTYPE: B!).

K. tenuifolia Steud., Syn. Glum. 69. 1854. C. tenuifolius (Steud.) Dandy, Catal.
Vasc. PI. S. Tomé 363. 1944. Type: not located; type locality, Senegal.

K. pumila var. elatior Boeck., Linn. 35: 413. 1868. Type: none specified; type
locality: “In Americae region calidior.”

K. flexuosa Boeck., Cyp. nov. 2: |. 1890. Type: JAMAICA. Bellevue, 27 Jan. 1888,
Eggers 3753 (HOLOTYPE: B!).

Cyperus densicaespitosus Mattf. & Kiikenth., Pflanzenreich 4(20): 597. 1936.
Based upon K. pumila.

Cespitose annual (or short-lived perennial), (5) 15-30 (55) cm tall.
Roots fibrous, brownish, rhizome none. Culms trigonous, smooth,
0.7-1.3 mm thick throughout. Leaves 1-3,(7), 4-20 (30) cm long,
1.5—3 (3.6) mm wide, flat to v-shaped, the margins and keel scabrel-
late, especially distally. Inflorescence bracts (3) 4 (5), (1) 3-10 (22)
cm long, (0.5) 1-2.5 (3) mm wide, the margins and keel ciliate-
scarbrellate, horizontal to ascendent at 30° at or before anthesis,
becoming horizontal to strongly reflexed in maturity. Spikes 1-3,
globose-ovoid to cylindric, 5-8 (11) mm long, 5~7 mm wide; second-
ary spikes smaller, globose-ovoid, 3-5 mm long, 2.5-4 mm wide;
rachis slenderly cylindrical, 3-5 mm long, 0.4—-0.7 mm thick; spike-
let pedicels 0.1-0.2 mm long, about 0.1 mm thick, separated by one
to two times their thickness. Spikelets 50-150, lanceolate to oblong-
lanceolate, (1.9) 2.4-2.8 (3.8) mm long, 0.6-0.9 mm wide, con-
tracted below into a stipitate base 0.2-0.4 mm long. Scales ovate
(1.8) 2.2-3.1 (3.4) mm long, the mucro 0.1—-0.3 mm long, 1-1.7 mm
wide, pale brownish to hyaline, laterally 2~3 (4) nerved; keel green,
with 3-10 spinulose teeth up to 0.2 mm long, or infrequently
smooth. Stamens 2; anthers elliptic, (0.2) 0.3-0.4 mm long, the
connective not prolonged; filaments (1.8) 2.4-3 mm long. Styles
0.5-0.8 mm long; stigmatic branches 2, 0.5-0.9 mm long. Achenes
oblong, lenticular, |-1.2 (1.4) mm long, 0.5-0.6 (0.7) mm wide,
apex subtruncate, apiculate, base cuneate to rounded, the stipitate
portion 0.1-0.2 mm long, about 0.1 mm wide, surface finely papil-
lose, light brown.

DISTRIBUTION. (Figure 1). Eastern United States from Pennsyl-
vania and Kansas to Florida and eastern Texas; the West Indies;
eastern and central Mexico south through Central America to
Argentina; also in tropical Africa. Moist disturbed open soils, wet
pastures, roadsides, croplands and thickets, from sea level to about
2000 m.

516 Rhodora [Vol. 86

K. pumila Michx.

Figure |. Distribution of K. pumila in Mexico and Central America.

PHENOLOGY. Flowers and fruits throughout the year.

Discussion. This species is distinguished by its cespitose annual
habit. Specimens of Ky/linga brevifolia collected without rhizomes
are frequently misdetermined as this species, but can be distinguished
by their much longer anthers, 0.8-1.1 mm long, versus 0.2-0.4 mm
longin K. pumila. The cespitose plants of K. pumila often have 10-20
culms, and usually several culms will have the anthers exserted and
readily visible.

ILLUSTRATIONS. Kiikenthal (1935-36), p. 596, fig. 63 E-G; Glea-
son & Cronquist (1952), v. 1, p. 250; Standley & Steyermark (1958),
p. 167, fig. 30 A-C.

REPRESENTATIVE SPECIMENS. MEXICO. Chiapas: Sta. Anita Huixtla, Boege /078
(MEXU); Mpio. Villa Corzo, Breedlove 37674 (ps); Mpio. La Independencia, 45-50
km E of Lagos de Montebello, Breedlove & Almeda 57709 (CAS); Mpio. Pueblo
Nuevo Solistahuacan, Breedlove & Thorne 21536 (DS, ENCB); Escuintla, Matuda
297 (MEXU). D. F. Pelouse du Pedregal pres Mexico, Bourgeau 662 (K, with K.
odorata). Guerrero: Dis. Adama, Temisco, Mexia 8795 (B, GH, LCU); Dis. Mina,
Petlacala, Mexia 9042 (B, GH, K, LCU). Hidalgo: Mpio. San Bartolo Tutotepec,
Santiago, Gimate 629 (ENCB, MEXU); Dis. Molango, Lake Atexca, Moore 2953
(BH). Jalisco: 4.5 mi NNE of Talpa de Allende, McVaugh 20232 (BM, DUKE,

1984] Tucker — Kyllinga $17

ENCB, TEX, MICH); Mpio. de Talpa, SE of Cuale, Gonzalez T. 459 (ENCB).
Mexico: Amatepec, Maruda 31275 (MEXU). Michoacan: 22 km S of Uruapan, King
& Soderstrom 4876 (MICH, TEX, US). Morelos: Cuernavaca, Deam 1/0 (GH).
Nayarit: between Chapalilla and Ixtlan on Hwy. 15, Kra/ 25659 (ENCB). Oaxaca: S
of Valle Nacional, King 2/33 (DUKE, MEXU, TEX); between Pochutla and
Miahuatlan, near Puente San Juan, Tucker 2252 (DUKE). Puebla: 2 km E of Villa
Juarez, Gonzalez Quintero 195 (ENCB, MSC). San Luis Potosi: 4 km ESE of
Tamazunchale, Rzedowski 9777 (ENCB), Mpio. Xilitla, 5 km W of Ahuacatlan,

zedowski 27714 (ENCB). Tabasco: Mpio. Cardenas, campo de investigacion
CSAT, Cowan 2468 (CAS, ENCB, MEXU). Tamaulipas: Mpio. Gomez Farias,
Rancho del Cielo, Puig 3245 (ENCB). Veracruz: Orizaba, Borteri 194 (BM, CGE):
Bourgeau 2737b (C, GH, M); Colipa, Liebmann s.n., March 1841 (C, with K.
brevifolia; K), Mpio. Santiago Tuxtla, Tepalapa, Martinez-Calderon 1480 (CAS,
DS, ENCB); 16 km E of Jalapa on the road to Cuauhtemoc, Tucker 2/35 (DUKE).

BELIZE. Belize Dis.: Boomtown, O'Neill 8974 (C, DS, GH, K, LCU, US). El
Cayo Dis.: near Camp 6, Gentle 2370 (F, GH, K, LCU, MEXU). Stann Creek Dis.:
Stann Creek Town, O’Nei// 8972 (LCU).

GUATEMALA. Alta Verapaz: Oliva prope Coban, von Tiirckheim 6 (BM, G, K,
US); Coban, von Tiirckheim I] 2271 (E, GH). Escuintla: San Andrés Osuna, Seler
2566 (GH). Huehuetenango: about Laguna de Ocubila, Standley 82650 (F). Izabal:
Quebradas, Pittier 8562 (GH, US). Guatemala: between Guatemala and San Rai-
mundo, Standley 63023 (F). Petén: La Libertad, Lundell 2498 (CAS, LCU). Sacatepe-
quez: NW of Antigua, Standley 64664 (F). Santa Rosa: Santa Rosa, Herde & Lux
3550 (GH, K, M, US). Zacapa: vic. of Zacapa, Standley 74272 (F).

HONDURAS. Atlantida: vic. La Fragua, Standley 55701 (F, US). El Paraiso: N of
Yuscaran, Standley 25710 (F). Gracias a Dios; orillas del Rio Dursuna, Ne/son 855
(BM); Intibuca: 9 km E of La Esperanza, Molina & Molina 13997 (F). Morazan: vic.
of El Zamorano, Molina 1591 (F, GH), Standley 21558 (F). Ocotepeque: Cordillera
Merendon, 10 km from Nueva Ocotepeque, Molina R. 22240 (F).

EL SALVADOR: Ahuachapan: vic. of Ahuachapan, Standley 19842 (GH). San
Miguel: vic. of San Miguel, Standley 2/088 (GH). San Salvador: vic. San Salvador,
Standley 23291 (BM, GH). Sonsonate: vic. Sonsonate, Standley 21791 (GH, US).

NICARAGUA. Chontales: Santo Tomas, Sevnrour 2785 (MSC). Jinotega: 10 km
NE of Jinotega, Atwood 2093 (F). Managua: El Rodeo, 12 km E of Managua, Garnier
4472(GH, LCU). Matagalpa: Sta. Maria de Ostuma, Williams et al. 27960 (F). Zelaya:
Rama, Seymour 718 (BM, F, GH).

COSTA RICA. Alajuela: waterfall of Rio Paz, Wilbur 33200 (DUKE); vic. Capulin
on Rio Grande de Tarcoles, Standley 50090 (US). Cartago: Rio Grande de Orosi,
Wilbur 32938 (DUKE); 2231/9 (DUKE); bords du Rio Tuis, Tonduz 8/80(US); vic. of
Tres Rios, Godfrey 67181 (DUKE, FLAS). Heredia: Paa Vulkanen Barba, Oersted
14500 (C); La Selva, OTS Field Sta., Folsom 10113 (DUKE), Sperry 11/4 (DUKE).
Limon: near Cahuita, Mac Dougal 1/63 (DUKE). Puntarenas: Osa Peninsula, vic.
Casa Medio Camino, Godfrey 66865 (FLAS); 19 km E of Esparza, Wilbur 29946
(DUKE); 4 km S of Las Alturas, Wilbur 22707 (DUKE). San Jose: Ochomogo,
Wilbur 33629 (DUKE); about 3 km E of San Marcos, Wilbur 32125 (DUKE); Los
Yoses, Wilbur 29439 (DUKE); in the saddle between Barba and Irazu, Wilbur 1/4701
(DUKE); El General, Kupper 1394 (M).

518 Rhodora [Vol. 86

PANAMA. Canal Zone: Near Gatun, Standley 27276 (US); Barro Colorado Is.,
Standley 41120(US); Ancon Hill, Standley 25168 (NY). Cocle: vic. El Valle de Antén,
Allen 106 (GH). Colén: Empire Sta., Haves 300 (BM); Chagres, Fendler 343 (GH,
NY). Chiriqui: 4 miles NE of El Hato del Volcan, Wi/bur et al. 15316 (DUKE); 6.6km
NNE of Boquete, Wi/bur & Luteyn 19275 (DUKE); vic. of Monte Lirio, Seibert 237
(GH, LCU, NY); Campamento Fortuna, Correa A. et al. 26/0 (DUKE); Darién: Rio
Chucunaque above Rio Tuquesa, Stern et al. 836 (GH, US); | mile N of Rio Sabana,
Tyson etal. 4794 (FLAS). Los Santos: 25 miles SW of Tonosi, Lewis et al. 2984 (NY).
Panama: Cerro Jefe, Tyson er al. 4301 (FLAS); Rio Tecumen, Standley 29434 (US);
Isla San Jose, Johnston 1188 (GH, NY, US).

2. Kyllinga odorata Vahl, Enum. pl. 2: 382. 1806. Type: America
meridionali, von Rohr (HOLOTYPE: C, not seen).

K. monocephala H. B. K., Nov. Gen. Sp. 2: 211. 1816. Type: VENEZUELA. Cara-
cas, HUMBOLDT (HoLotype: P; IDC microfiche Herb. H. B. K.!), nom. illeg.
Art. 63.1.

K. leucocephala Baldw., Trans. Amer. Philos. Soc. Philad. n. s. 2: 170. 1825.
TyPE: URUGUAY. Maldonado, Baldwin (HOLOTYPE: PH!).

K. sesquiflora Torr., Ann. Lyceum Nat. Hist. New York 3: 287. 1836. Cyperus
sesquiflorus (Torr.) Mattf. & Kiikenth., Pflanzenreich 4 (20): 591. 1936. Type:
u. 8. A. Middle Florida, Chapman 12 (HOLOTYPE: NY!).

K. martiana Schrad. ex Nees, Fl. bras. 2: 14. 1842. Type: BRAZIL. Prov. Para,
Martius (HOLOTYPE: M!).

K. odorata var. minor Boeck., Linnaea 35: 411. 1868. TyPE: MARTINIQUE. Sieber
/8 (HOLOTYPE: not located; IsoTYPE: M!).

K. odorata var. rigida Boeck., Vidensk. Meddel. 1894: 271. 1895. TyPE: BRAZIL.
Sao Paolo, Franca, Lofgren & Edwall 2105 (SyNTyYPE: C!).

Tufted perennial (5) 10-25 (45) cm tall. Rhizome (rarely collected)
horizontal, oblique just below the culms, 3-6 mm thick, internodes
5-12 mm long. Culms triquetrous, smooth, 0.5-1.2 mm thick just
above the sheaths, 0.5-1.2 (1.5) mm thick at the apex, bulbous-
thickened at the base, clothed with remnants of previous year’s leaf
sheaths. Leaves (1) 3-7 (10), (6) 10-18 (30) cm long, 2-3 (4) mm wide,
erect or arching, ciliate-scabrous along the margins and keel, espe-
cially apically. Bracts (2) 3-4, 1-7 (9) cm long, (0.5) 1-3 (4.5) mm
wide, ciliate-scabrous along the margins and keel, horizontal to
ascendent before anthesis, horizontal to reflexed downward parallel
to the culm in maturity. Spikes 1—3 (4), whitish, the central one the
largest, densely ovoid to cylindrical, 6-12 (17) mm long, (4) 5-7 (8)
mm wide, the other spikes smaller, shorter, ovoid, 3-5 (6) mm long,
3.5-5 mm wide. Rachis slenderly cylindrical, (4) 5-7 mm long, (0.6)
0.8-1 mm thick; spikelet pedicels very short, about 0.2 mm wide, 0.1
(—0.2) mm high, decurrent by a somewhat spongy thickened ridge.

1984] Tucker — Kyllinga 519

. odorata Vahl

Figure 2. Distribution of K. odorata in Mexico and Central America.

Spikelets (50) 75-150, ovate to ovate-lanceolate, whitish, (1.8)
2.3-2.8 (3.6) mm long, I-1.3 mm wide. Scales broadly ovate, 2-2.5
mm long, (1.2) 1.8-2.6 mm wide, whitish, often red-speckled, mucro-
nate, the apex often excurved, (5) 7 (9) nerved, the keel smooth or
with 1-6 spinulose prickles in the distal half, whitish to greenish.
Stamens 2; anthers (0.4) 0.6-0.8 (1.0) mm long, linear; filaments 2-2.4
mm long. Styles 0.6-1 mm long; stigmatic branches 2, 0.3-0.6 mm
long. Achenes lenticular, oblong-ovate, 1.2-1.5 mm long, 0.7-0.8
(0.9) mm wide, apex broadly rounded to subtruncate, apiculate, the
stipitate base 0.1-0.2 mm long, about 0.2 mm wide, surface papillose,
reddish brown to dark brown, strongly contrasting with the whitish
base and apiculus.

DISTRIBUTION. (Figure 2). Pantropic; in the New World in the
southern United States from North Carolina to Florida and eastern
Texas; the West Indies; Mexico and Central America to Argentina; in
Mexico from southern Tamaulipas (Tampico region), eastern San
Luis Potosi, Distrito Federal, southeastern Sonora (Yecora) to Nay-
arit and Chiapas; absent from the Yucatan Peninsula, although it
occurs in Belize: croplands, river banks and gravel bars, roadsides,
open pine, oak or thorn woodlands from sea level to about 2500 m.

$20 Rhodora [Vol. 86

PHENOLOGY. Flowers and fruits from May to September, and
sporadically as late as January.

Discussion. The whitish color of the inflorescence generally is
distinctive. As the achenes mature and darken, they become visible
through the sides of the scales; thus the inflorescence at maturity is
often darker than at anthesis. The broad dark achenes, with conspic-
uous whitish apiculus and stipitate base when fully mature distin-
guish this species from others of the genus in the New World.

The size of the anthers varies geographically in this species. In
collections from Central America (excluding Guatemala), the aver-
age length of the anthers is 0.5-0.6 mm (range: 0.4-0.8 mm), while in
Mexico the average is 0.7-0.8 mm (range: 0.6-1.0 mm). The ranges of
anther length for these two above regional populations do overlap.
Moreover, the average length for Guatemalan specimens was 0.6-0.7
mm, exactly intermediate between the averages for Mexico and the
remainder of Central America. Thus there seems to be continuous
variation in anther length from Mexico through Central America.
The recognition of either the Mexican or Central American popula-
tions at varietal rank would be arbitrary. No such formal taxonomic
rank is here offered.

ILLUSTRATIONS: Godfrey & Wooten (1979), p. 246, fig. 137.

REPRESENTATIVE SPECIMENS. MEXICO. Chiapas: Mpio. Angel Albino Corzo, above
Jaltenango, Breedlove 28562 (DS); La Florida, between Oxchuc and Ocosingo,
Tucker 2222 (DUKE). Distrito Federal: Tlalpam, Fisher s.n., 31 July 1926 (DS,
DUKE, US); Cerro Xochitepec, Rzedowski 24248 (DS, ENCB, MSC). Guerrero: Dis.
Mina, Manchon, Hinton 9429 (DS, K, MEXU, MICH, NY); about 10 miles from
Taxco on Cuernavaca road, Mac Daniels 125 (BH, F). Hidalgo: near Trinidad Iron
Works, Pringle 8959(C, CU, E, ENCB, K,GH, M, MEXU, MSC, NY, US): 6km N of
Pachuca, Garcia 103 (ENCB). Jalisco: Guadalajara, Palmer 253a in 1886(GH, MICH,
NY); 3 miles S of Mazamitla, Sierra del Tigre, Mc Vaugh 12980 (MICH); 5 road miles
SW of Santa Cruz de Las Flores, McVaugh 16290A (MICH). Mexico: Mpio. de
Texcoco, 8 km E of Coatlinchan, Cruz Cisneros 1314 (ENCB); Mpio. Zumango, | km
SE of San Juan Zitlaltepec, Cruz V. 7/ (ENCB):; 60 miles W of Toluca, near Michoa-
can border, Manning 531121 (GH); Temascaltepec, Volcan, Hinton 1176 (BM,
ENCB, F, K, NY); Cerro de Sacramonte, cerca de Amecameca, Rzedowski 24202
(ENCB); 5 km W de Progreso Nacional, Rzedowski 35296 (ENCB). Michoacan:
Mpio. Tancitaro, | mile S of Tancitaro, Leavenworth 376 (F, MICH, MO, NY); 0.8
miles W of Morelia on Hwy. 15, Kral 25583 (ENCB), Dis. Zitacuaro, Loma Larga,
Hinton 13103 (DS, ENCD, K, MICH, NY); 8-10 km SW of Jiquilpan, Feddema 4
(DS, DUKE, ENCB, MICH, TEX); 6-7 km N of Uruapan, Dieterle 4402 (ENCB,
MICH); Morelia, Arsene 5752 (B). Morelos: 1.5 miles SE of Huitzilac, Smith 55

1984] Tucker — Kyllinga 521

(MEXU); 12 km NW of Cuautla, Thomas 66 (MEXU). Nayarit: 9 miles N of Com-
postela, Mc Vaugh 16559 (MEX, MICH); Mpio. Acaponeta, near Labra W of Jesus
Maria, Norris & Taranto 14170 (MICH); 10 miles SE of Ahuacatlan, on the road to
Barranca del Oro, Feddema 254 (DS, DUKE, ENCB, MICH, TEX); 12 miles SE of
Tepic, Feddema 588 (DS, DUKE, ENCB, MICH, TEX), 1.5 miles W of Mazatan,
Feddema 1073 (DUKE, ENCB, MICH, TEX). Oaxaca: 53 km N of Puerto Escon-
dido on the road to Zimatlan, Roe et al. 560 (F); 16 km NE of Zanatepec, King SOS
(ENCB, MICH). Puebla: | km W of Villa Juarez, Galvan s.n., 7 Sept. 1963 (ENCB).
San Luis Potosi: Huichihuayon, Grant 540 (GH, MICH). Sinaloa: Sierra Surutato,
3 miles N of Los Ornos, Breedlove & Thorne 18409 (CAS); San Ignacio, Montes &
Salazar 471 (US); near Colomas, Rose 1/803 (US). Sonora: Yecora, Pennington 108
(TEX). Tamaulipas: Rancho del Cielo, 6 km NW of Gomez Farias in Sierra de
Guatemala, Sharp er al. 52029 (MSC). Veracruz: near Tampico, Berlandier 2137
(GH); near Orizaba, Bourgeau 2589 (C, GH, K); 5.4 miles N of Jalapa on road to
Misantla, Tucker 2077 (DUKE).

BELIZE. Belize Dis.: Sibun River, Graig Point, Gentle 1398 (NY); low pine ridge
near Manatee Lagoon, Peck 39 (GH, K).

GUATEMALA. Alta Verapaz: Cubilguitz, von Turckheim 7691] (K). Chimalte-
nango: on road to Chichicastenango, Dunn et al. 22960 (MO). Chiquimula: Volcan
Ipala, Srevermark 30529 (F). Guatemala: near Guatemala City, Tonduz 672 (GH,
NY, US). Huehuetenango: Cerro Victoria, Stevermark 49592 (F, NY). Izabal: vic.
Puerto Barrios, Standley 25057 (NY). Petén: La Libertad and vic., Aguilar H. 6
(MICH, in part, with Cyperus ischnos Schlecht.); | km N of Poptun, Harmon &
Dwyer 2719 (MO). Santa Rosa: Santa Rosa, Herde & Lux 3540 (GH, K, M, MICH,
US). Zacapa: Sierra de las Minas, between Rio Hondo and summit of mtn.. above
Finca Alejandra, Srevermark 29708 (F).

HONDURAS. Atlantida: between Tela and Lancetilla, Yuncker 458/ (MICH,
with K. pumila). Copan: Hac. Espiritu Santo, Blake 7438 (US). Gracias a Dios;
Puerto Lempira, Proctor 38922 (BM). Morazan: El Zamorano, Standley 12857 (F);
Las Flores, Cerro de Uyuca, Standley 21696 (F), region of Las Mesas, Standley
24069 (F). Yoro: N end of Lake Yojoa, Kamb 2049 (A).

EL SALVADOR. San Salvador: San Salvador, Fassett 28259 (F, GH, WIS).

NICARAGUA. Esteli: 10.6 km W of bridge at La Trinidad to San Nicolas,
Stevens 10299 (MO). Managua: Sierra de Managua, Garnier s.n., 1930-1940 (F).
Zelaya: Puerto Isabel, Seymour 2972 (F, GH).

COSTA RICA. Alajuela: between Rios Pilos and Zacaros, Brenes 17319 (F. NY),
about 4 km E of Naranjo on the new road to San Ramon, Wilbur 29960 (DUKE).
Cartago: south slope of Volcan Turrialba near Santa Cruz, Holm & Iltis 132 (A,
NY); Cartago, Oersted 14499 (C). Guanacaste: Bagaces, Opler 280 (F). Limon: delta
du fleuve San Juan, Pittier 2580 (US). Puntarenas: outskirts of Chomes, Davidse &
Pohl 1327 (MO); Cascajal, Holm & Iltis 239 (A, NY). San José: | km S of San Pablo
towards San Marcos, Mac Dougal 861 (DUKE); vic, El General, Skutch 2643 (GH,
K, MICH, NY, US); San José; Tonduz 434 (DS, F, GH, PH).

PANAMA. Canal Zone: Frijoles, Ebinger 79 (F); Mt. Hope Cemetery, Standley
28807 (US); Miraflores Lake, Tyson 1410 (FLAS). Chiriqui: Savanne bei David,
Wagner | 1/2(M). Coclé: El Valle de Anton, Seibert 474 (GH, NY). Colén: Colén,
Rose 22086 (GH, NY). Panama: near Matias Hernandez, Standley 28988 (US); Isla
San José, Bald Hill, Johnston 39 (GH).

a2 Rhodora [Vol. 86

3. Kyllinga brevifolia Rottbgll, Descr. Icon. Rar. Nov. Pl. 13. t. 4.
fig. 3. 1773. Cyperus brevifolius (Rottb.) Hassk., Cat. Hort,
Bogor. 24. 1844. Type: EAST INDIES, Kénig (HOLOTYPE: C!;
ISOTYPE, C!).

K. monocephala Thunberg, Fl. Japon. 35. 1784. Type: JAPAN, Thunberg (not
located).

K. elongata H. B. K., Nov. gen. sp. 2: 211. 1816. Type: PERU. Between Gonzana
and Loxa, Humboldt & Bonpland (P; 1DC microfiche Herb, H. B. K.!).

K. cruciformis Schrad. ex Schult., Mant. 2: 137. 1824. Type: VIRGIN ISLANDS. St.
Thomas, Ehrenberg 69 (HOLOTYPE: B!; 1soTyPEs: C! HAL!).

K. tenuis Baldwin, Trans. Amer. Philos. Soc. Philad. n.s. 2: 168. 1825. TyPE:
ARGENTINA. Buenos Aires, , Baldwin (HOLOTYPE: PH!).

Perennial, (5) 12-25 (55) cm tall, the culms tufted or widely
spaced along a conspicuous rhizome. Rhizome (1) 3-12 (30) cm
long, (0.5) 1-2 mm thick, clothed with lanceolate scales 6-13 mm
long, the internodes (2) 5-12 (30) mm long. Culms trigonous,
smooth, 0.6—1.2 (1.5) mm thick throughout. Leaves 1-3, (2) 6-15
(21) cm long, 1.5-3.5 mm wide, flat, scabrellate along the margins
and keel especially distally. Bracts 3-4, (1.5) 4-12 (18) cm long,
1-3.3 mm wide, margins and keel ciliate-scabrellate, the longest
bract nearly always erect, appearing as a continuation of the culm,
the remaining ones ascendent to horizontal. Spikes 1-3, globose-
ovoid to cylindric, 4-7 mm long, about 4 mm wide, greenish; rachis
slender, 1.5-3.5 (4.1) mm long, 0.4-0.9 mm thick; spikelet pedicels
about 0.1 mm long and about 0.1 mm thick. Spikelets (20) 40-60
(100), oblong-lanceolate, (2.2) 2.5-2.8 (3.2) mm long, (0.6) 0.7-0.8
(1.2) mm wide, the base stipitate, 0.1-0.3 mm long. Scales elliptic to
ovate, (1.8) 2.2-3.0 mm long, the mucro an additional 0.2-0.3 mm
long, (0.8) 1.1-1.6 mm wide, laterally dull whitish to pale brownish
or greenish, 2 (3) nerved, occasionally red spotted; keel greenish, |
(3) nerved, smooth or with 3-6 (—12) spinulose teeth up to 0.2 mm
long. Stamens 2 (rarely 1); filaments (1.7) 2-3 mm long. Styles
0.6-1.2 mm long; stigmatic branches 2, 0.5-1.5 mm long. Achenes
elliptic to oblong-elliptic, lenticular, 1-1.2 (1.3) mm long, 0.6-0.8
mm wide, apex broadly rounded to subtruncate, base substipitate
to decidedly stipitate, 0.1-0.2 mm long, 0.1 (0.2) mm wide, whitish,
surface papillose, brownish.

DISTRIBUTION. (Figure 3). Pantropical; in the New World from
North Carolina to Texas and Florida; also in California; southern

1984] Tucker — Kyllinga 523

. brevifolia Rottb.

Figure 3. Distribution of K. brevifolia in Mexico and Central America.

and western Mexico south through Central America and the West
andies to Brazil and Argentina. Roadsides, pastures, ditches,
marshes and streambanks, from sea level to about 2500 m.

PHENOLOGY. Flowering and fruiting throughout the year.

Discuss1IoN. Delahoussaye and Thieret (1967) reported this species
to have only one stamen per flower. However, the majority (about
2/3 of the specimens seen in the present study) had two stamens per
flower; the remainder had one. Occasionally plants of the same
collection had flowers with both one or two stamens.

O’Neill (1940) in addition to the difference in habit of the two
species, separated Ky/linga brevifolia from K. pumila by a difference
in achene width, 0.8 mm in the former, 0.6 in the latter. Apparently
this supposed distinction was based upon relatively few collections. I
cannot corroborate such a difference. The achenes of K. brevifolia
are 0.6-0.8 mm wide, while those of K. pumila are 0.5—-0.6 (0.7) mm
wide. The slight difference in the widths of the achenes is hardly a
useful means of distinguishing these two species.

$24 Rhodora [Vol. 86

ILLUSTRATIONS: Gleason & Cronquist (1952) v.1, p. 250; Kern
(1974) p. 658, fig. 70; Godfrey & Wooten (1979) p. 246, fig. 137.

REPRESENTATIVE SPECIMENS. MEXICO. Chiapas: Mpio. La Independencia, 6-10
km NNE of La Soledad, Breedlove 53203 (CAS); Mpio. Arriaga, 13 km N of Arriaga
on Hwy. 195, Breedlove & Davidse 54145 (CAS); Mpio. Pueblo Nuevo Solistahua-
can, Ton 2755 (DS, ENCB, LL); between San Cristébal and Ocosingo, near Aba-
solo, Tucker 2219 (DUKE). Michoacan: Morelia, Arsene s.n., April 1912 (E).
Nayarit: Mpio. de Tepic, Norris & Taranto 13255 (MICH): Mpio. de Compostela, 5
miles W of Compostela, Norris & Taranto 13921 (MICH); just E of San Blas,
Philbrick 733 (BH). Oaxaca: Cerro Pelon, 47 km N of Ixtlan de Juarez, Gonzalez
1016 (ENCB); between San Pedro Pochutla and Miahuatlan, near Puente San Juan,
Tucker 2253 (DUKE). Puebla: | km W of Villa Juarez, Rzedowski 17203 (ENCB).
Tabasco: Tenosique, Boca Cerro, Matuda 3559 (F, GH, LCU, MEXU, MICH);
Santa Unita, Rovirosa 708 (K, PH). Veracruz: near Santiago Tuxtla, Dressler &
Jones 184 (F, GH, NY, US): Colipa, Liebmann s.n., March 184] (C, K); Cordoba,
Matuda 310 (MICH).

BELIZE. Belize Dis., Belize, Lunde/! 1906 (LCU, MICH, MO, TEX). Corozal
Dis.: Corozal-Consejo Rd., Lundell 4973 (B, LCU, MICH, MO, NY).

GUATEMALA. Chimaltenango: near San Martin Jilotepeque, Standley 64513
(F, MICH). Chiquimula: Montafia Nonoja, 3-5 miles E of Comotan, Srevermark
31729 (F). Izabal: vic. of Quirigua, Standley 24066 (GH, NY); vic. Puerto Barrios,
Standley 24777 (GH, NY). Jalapa: Laguna de Ayarza, Hevde & Lux 3897 (GH, K,
M, MO, NY, US). Petén: Sayaxche, Stevermark 46279 (F, MICH). Retalhuleu: 9
km N of Champerico, Harmon 2303 (ENCB. MO). Sacatepeéquez: near Las Lajas,
Standley 58106 (F). San Marcos: W of Tajamulco, Stevermark 36724 (F, US).

HONDURAS. Atlantida: between Tela and Lancetilla, Yuncker 4581 (F, MICH),
Cortés: San Pedro Sula, Garcia 22 (MO). Distrito Central: San Juancito and vic.,
Pfeifer 2008 (US). Gracias a Dios: Puerto Lempira, Proctor 38922 (BM, with K.
odorata). Morazan: El Jicarito, Standley 20484 (F), 20788 (F). Olancho: Culmi,
Nelson & Romero 4652 (MO).

EL SALVADOR. Ahuachapan: vic. Ahuachapan, Standley 20290 (GH, NY, US).
La Libertad: vic. La Libertad, Standley 23205 (GH, MO. NY, US). San Miguel:
south side of Lake Olomega, Tucker 850 (F, GH, K, MO, NY). San Vicente: vic. San
Vicente, Standley 2/157(GH, NY, US). Santa Ana: N of Metapam, Rohweder 2176,
2185 (MO).

NICARAGUA. Bluefields: Waspam, van der Sluijs S. 791 (F). Managua: vic.
Managua, Narvaez 258 (F, GH). Matagalpa: Matagalpa, Zelaya M. 2298 (CAS, GH,
MSC). Zelaya: Puerto Isabel, Narvaez S. 2883 (DUKE).

COSTA RICA. Alajuela: Carrillos de Poas, Brenes 19330 (NY); near Rio Pefias
Blancas, Hepper 106 (BM). Cartago: Rio Grande de Orosi, Wilbur 30752 (DUKE);
Tuis, Tonduz 11388 (MO, US). Guanacaste: Liberia, Wilbur 31027 (DUKE). Here-
dia: La Selva, OTS Field Sta., Wilbur 33275, 33277 (DUKE). Limén: Talamanca,
border of river at Shirores, Tonduz 9218 (US); 28 miles on R. R. from Puerto
Limén, Cufodontis 642 (F). Puntarenas: vic. of Esparta, Godfrey 66977 (FLAS,
MO). San José: NE of San Jerénimo, Burger & Stolze 5336 (F, MO, NY, PMA); El
General, Skutch 2859 (GH, K, MICH, MO, NY, US); Los Yoses, Wilbur 26327
(DUKE). Isla del Coco: Wafer’s Creek, Gomez P. 3279 (F, MO, NY, PMA).

1984] Tucker — Kyllinga 525

PANAMA. Bocas del Toro: Changuinola, Lazor et al. 2608 (FLAS, PMA), VIC.
Laguna de Chiriqui, Hart 82 (K, US). Canal Zone: 3 miles W of Gamboa, Nee &
Mori 3601 (MEXU, PMA, US, WIS). Chiriqui: Finca Lerida to Boquete, Woodson
eral. 1155 (GH, NY); 2.5 miles S of Cerro Punta, Sawyer s.n., 2 March 1967 (DS,
WIS). Colon: Portobelo, Wilbur & Lutevn 11650 (DUKE, with K. pumila). Cocle:
between Las Margaritas and El Valle, Woodson et al. 1773 (GH, NY), 1774 (BM).
Los Santos: 25 miles SW of Tonosi, Lewis et al, 2944 (NY, US). San Blas: Marra-
ganti, Williams 1040 (NY, US). Veraguas: 5 miles W of Santa Fe, Liesner 945 (GH,
PMA, WIS).

4. Kyllinga vaginata Lam., Tabl. encycl. 1: 148. 1791. TYPE: PERU.
Dombey (LectotyPE: here designated: P-LA; photo: DUKE!;
ISOLECTOTYPE: C!).

K. peruviana Lam., Encycl. 3: 366. 1792. Cyperus peruvianus (Lam.) F. N. Willi-
ams. Bull. Herb. Boiss. 2 sér. 7: 90. 1907. Type: PERU. Dombey, the same as K.
vaginata (nom. illeg., Art. 63.1).

K. rigida Baldw., Trans. Amer. Philos. Soc. Philad. n.s. 2: 169. 1825. TYPE: BRAZIL.
Rio de Janeiro, Baldwin (HOLOTYPE: PH!).

K. pungens Link, Hort. Berol. 1: 326. 1827. Type: not located.

K. obtusata Presl, Relig. Haenk. 1: 183. 1828. Cyperus obtusatus (Presl) Mattt. &
Kiikenth.. Pflanzenreich 4(20): 585. 1936. Type: PERU. “In montanis Huanoccen-
sibus,” Haenke (PR!).

K. obtusata var. cvlindrostachyus Boeck., Linnaea 35: 419. 1868. Cyperus obtusa-
tus var. cvlindrostachyvus (Boeck.) Kiikenth., Pflanzenreich 4 (20): 586. 1936.
TYPE: BRAZIL. Prov. Bahia, Se//ow (HOLOTYPE: B!).

K. tenuis Boeck., Linnaea 35: 423. 1868. Type: BRAzIL. Se//ow, (HOLOTYPE: B,
Herb. Kunth!).

Perennial, 20-60 cm tall. Rhizome indurate, 2-3 mm thick,
clothed with arched, cucullate, ovate-lanceolate, brown to reddish-
brown scales 8-12 mm long, 4-7 mm wide, the internodes 2-12 mm
long. Culms single at each node, 0.8-2 mm thick just above the leaf
sheaths, 0.9-1.5 (1.8) mm thick at the apex, trigonous, hollow,
usually flattened in drying, glabrous. Leaves with short blades 2-10
cm long, 2-4 mm wide, scabrellate on the margins and keel; or,
lacking blades, the lower culm with 2-3 (4) bladeless sheaths 2-10
cm long, the sheaths with a conspicuous hyaline to light brown,
speckled border 0.5-1.2 mm wide. Inflorescence bracts 2-4, green-
ish, 1-3 (8) cm long, |.2-3.5 mm wide, horizontal to reflexed, usu-
ally erect and clasping the rachis after the spikelets fall, the margins
and keel scabrellate. Inflorescence a solitary subglobose to ovoid
head 7-11 mm long, 7-9 mm wide, rounded at the apex, truncate
below. Rachis cylindrical to conical, 3.2-5 mm long, I-I.7 mm

526 Rhodora [Vol. 86

wide. Spikelet pedicels rather closely spaced, separated by about or
less than their own width, 0.3-0.4 mm long, about 0.2 mm wide.
Spikelets 75-130, lanceolate, stipitate at the base, 2.5-4 mm long,
I-1.3 mm wide. Scales ovate, 2.7~3.4 mm long, |.6-2.2 mm wide,
laterally 3-4 (5) nerved, off-white to pale reddish brown; keel whit-
ish to greenish, 3-nerved, with 2-3 (4) spinulose teeth. Stamens 3
(the upper scale occasionally bearing 2 stamens without gynoe-
cium); filaments brownish, 1.5-3 mm long; anthers linear (0.9)
1.1-1.3 mm long, the connective tip reddish, usually conspicuous,
but less than 0.1 mm long. Styles |—1.8 mm long; stigmatic branches
2, |-1.5 mm long. Achenes lenticular, oblong-ellipsoid, 1-1.2 mm
long, 0.4-0.6 mm wide, the apex broadly rounded, apiculate, the
base cuneate, substipitate, about 0.1 mm long, puncticulate, light
brown.

DISTRIBUTION. The Greater Antilles; Belize to Panama; Suri-
nam to Peru, Brazil and northern Argentina; tropical Africa. Brack-
ish marshes; wet, sunny soil and riverbanks; sea level to about 200
m. This species is not yet known to occur in Mexico. It is apparently
rare in Central America; only the six collections cited below have
been seen. However, it is fairly common in South America, espe-
cially in eastern Brazil, Paraguay, Uruguay and northern Argentina.

PHENOLOGY. Flowering and fruiting in April, May and June.

Discussion. This species is separated from Kyllinga tibialis
chiefly by characters of the inflorescence. In K. vaginata, the spike
is truncate at the base, while in K. tibialis the spike is spherical,
rounded at the base. The cylindrical rachis of this species is strik-
ingly different from the spherical one of K. tibialis.

Kiikenthal (1935-36) and Standley (1938) treated this species as
Cyperus obtusatus (Presl) Mattf. & Kiikenth. Kiikenthal appar-
ently did not examine the type of Ky//inga vaginata. Lamarck cited
two collections in his description, Dombey from Peru, and Roussil-
lon from Senegal. The latter, however, is actually a plant of K.
tibialis. The Dombey collection is here selected as the lectotype of K.
vaginata. Two phrases in Lamarck’s description indicate such a
choice, “culmo inferne vaginato” and “involucro brevi triphyllo”.
The Roussillon collection lacks a lower culm and has only one
involucral bract, while the Dombey collection has a clearly visible
sheath on the lower culm, as well as three involucral bracts.

1984] Tucker — Kyllinga 527

The binomial Ky//inga peruviana belongs in the synonymy of this
species. Since it was also based on Dombey’s collection, it is an
illegitimate name, which Kiikenthal mistakenly placed in the syn-
onymy of K. tibialis Ledeb.

One might confuse Ky/linga vaginata with K. brevifolia, since
both species are rhizomatous. However, K. brevifolia has long leaf
blades (up to 21 cm long) longer inflorescence bracts (up to 18 cm
long) and has only one leafless sheath at the base of the culm, while
K. vaginata has 2-4.

Not all the characters used by Kikenthal (1935-36) to separate
Kyllinga tibialis from K. vaginata are reliable. He separated the two
species as follows:

Leaf blades 1-4; bracts longer than the spike; rachis cylindric

pti clen a ate DO e a a tah eka a ake meee K. vaginata
Leaf blades none; bracts shorter than the spike; rachis
NOt uy ade iar es ps dada ee ee emer K. tibialis

Both species may have leaf blades or lack them, often on adjacent
culms from the same rhizome. Similarly, the inflorescence bracts
may be longer or shorter than the spike, sometimes on adjacent
culms. The difference in the shape of the rachis holds up well, and
combined with the combined with the characters in the key used in
the present treatment, show that K. vaginata is a species amply
distinct from K. tibialis.

ILLUSTRATION. Pedersen (1970) p. 376, fig. 69 A-C.

SPECIMENS EXAMINED. BELIZE: Stann Creek Dis.: Stann Creek Town, 20
Feb. 1890, Robertson 103 (BM), and 19 Sept. 1936. Redmond 8969 (LCU).

GUATEMALA: Depto. Izabal: brackish marsh near Puerto Barrios, 25 April
6 May 1939, Standley 72157 (F, MICH).

NICARAGUA: Depto. Rio San Juan: Greytown (San Juan del Norte), 1867-68,
Tate 506 (BM, K).

COSTA RICA: Prov. Alajuela: Muelle de San Rafaél-San Carlos, June 1890,
Pittier 2587 (BR, US).

PANAMA: Prov. Bocas del Toro: Bocas; along runway bordering mangrove
swamp, 17 April 1969, Lazor, Tyson & Loftin 2408 (FLAS).

5. Kyllinga tibialis Ledeb., Diss. bot. pl. doming., p. 6. [May] 1805.
TYPE: DOMINICAN REPUBLIC. Poiteau (HOLOTYPE: LE!).

K. aphyvlla (Vahl) Kunth, Enum. pl. 2: 127. 1837. Mariscus aphyllus Vahl, Enum.
pl. 2: 373 [October] 1805. Type: SENEGAL. Dupuis (HOLOTYPE: C!).

528 Rhodora [Vol. 86

K. capitata P.-Beauv., Fl. Oware | (6), t. 31. 1806. Type: not located.

K. globosa P.-Beauv., Fl. Oware | (6), t. 31. 1806. Type: Palisot de Beauvois, Herb.
Fl. Oware (G!).

K. peruviana Lam. var. foliata Kiikenth., Fedde Rep. 12: 92. 1913. Cyperus
peruvianus var. foliatus (Kiikenth.) Kiikenth., Pflanzenreich 4 (20: 587. 1936.
Type: JAMAICA. Ocho Rios, near sea shore, 4 April 1908, Britton & Hollick 2705
(HoLotypr: B; 1soTyPe: NY!).

Perennial, 25-70 (85) cm tall. Rhizome indurate, 3-5 mm thick,
clothed with arched, cucullate, broadly ovate-triangular, brown to
reddish-brown scales, 3-6 mm long, the internodes (2) 5-10 (15)
mm long. Culms single at each node, (1.6) 2.5-3.5 mm thick just
above the leaf sheaths, (1.2) 1.6-2 mm thick at the apex, roundly
trigonous, hollow, flattened in drying, glabrous. Leaves usually
bladeless, the lower culm with 3-7 bladeless sheaths 1-5 (7) cm
long, the sheaths with a conspicuous cinnamon-brown, scarious
border 0.5—1.2 mm wide; leaf blade infrequently present, |—5 (8) cm
long, 3-7 mm wide, marginally scabrellate near the tip. Inflores-
cence bracts 3-4, brownish or greenish, 3-6 (20) mm long, (1.5) 3-4
(4.5) mm wide, at anthesis loosely clasping the spikelets, in maturity
often deflexed. Inflorescence a solitary, globose head 7-12 mm in
diameter; rachis spherical to hemispherical, 1.5-3 (4.2) mm high,
2-3 (4.2) mm wide. Spikelet pedicels closely spaced, separated by
less than their own width, 0.4-0.8 mm long, 0.2 (0.3) mm wide.
Spikelets about 100-200, broadly to narrowly lanceolate, 3.2-3.8
(4.5) mm long, I-1.4 mm wide. Scales ovate, 2.5-3.3 mm long,
1.6-2 mm wide, dull greenish or brownish white, (5) 7-9 nerved;
keel whitish or greenish, smooth or infrequently with 1-3 spinulose
teeth less than 0.1 mm long. Stamens 3; filaments whitish, ribbon-
like, 2.7~3.3 mm long; anthers linear, |-1.5 mm long, the connective
tip red, up to 0.1 mm long. Styles 0.4-1 mm long; stigmatic
branches 2, |.3-2.3 mm long. Achene lenticular, oblong-obovate,
1-1.5 mm long, 0.7-0.8 mm wide, apiculate, the apex broadly
rounded to subtruncate, the base cuneate to substipitate, the surface
puncticulate, brown to reddish brown.

DISTRIBUTION. (Figure 4). Caribbean Islands from Jamaica and
Hispaniola to Trinidad; Belize to Panama and Colombia on the
Carribean coast only; not recorded from Mexico. Also in tropical
West Africa. Sandy beaches, among littoral vegetation of the upper
beach, dunes and swales; margins of mangrove swamps and river-
banks up to about 70 m.

1984] Tucker — Kyllinga 529

. tibialis Ledeb.

Figure 4. Distribution of K. tibialis in Mexico and Central America.

PHENOLOGY. Flowering and fruiting throughout the year.

Discussion. A discussion of the differences between this species
and Kyllinga vaginata is given under the latter species.

This species is usually easily recognized by its littoral habitat, and
its dense globose inflorescence. In collections with several culms on
one rhizome, there is usually at least one old culm from which the
spikelets have fallen. The spherical rachis thus revealed is a unique
character allowing positive identification of Ky/linga tibialis.

Kiikenthal (1935-36) treated this species as Cyperus peruvianus
(Lam.) F. N. Williams, based on the illegitimate name Ky //linga
peruviana Lam. (see discussion under K. vaginata). Thus, K. tibialis
is the oldest available epithet for this species, antedating Mariscus
aphyllus Vahl by a few months.

REPRESENTATIVE SPECIMENS. BELIZE. Belize Dis.: Pine ridge near Manatee
Lagoon, Peck 40 (GH, K). Stann Creek Dis.: Middlesex, Schipp 382 (F, GH, K,
MICH, MO, NY); Dangriga, Proctor 36595 (MO); Hatchet Cay, Fosberg 54376 (F.
NY); Stann Creek Town, Croat 24111 (MO). Toledo Dis.: Nicolas Cay, Spellman &
Stoddart 2343 (MO, US); Punta Gorda, Hedger 271] (BM).

GUATEMALA. Izabal: Livingston, Deam 72 (GH, MICH, MO, NY); Puerto
Barrios, Kellerman 5133 (MEXU, US), Standley 25139 (NY, US); NE of Livingston,
Stevermark 39710 (F).

530 Rhodora [Vol. 86

HONDURAS. Atlantida: Tela, Davidse & Poh! 2176 (MO, US), Molina R. 25727
(F, MO); vic. of Ceiba, Yuncker et al. 8247 (F, GH, MICH, MO, NY). Colon:
Trujillo Beach, Saunders 142 (BM, MO). Cortés: Puerto Cortés, O'Neill 8970 (GH,
K); Gracias a Dios; alrededores de Puerto Lempira, Castro T. 191 (MO). Islas de la
Bahia: Roatan Is., Flower Bight Beach, Molina R. 20671 (ENCB, F, US).

NICARAGUA. Bluefields: El Bluff. Sermour 647 (BM, F, GH). San Juan del
Norte: vic. of San Juan del Norte, Smith 56 (ENCB). Zelaya: Puerto Cabezas,
Molina R. 1485] (GH, US); Puerto Isabel, Narvaez S. & Atwood 2944 (F, MSC).

COSTA RICA. Limon: Cahuita, Baker & Burger 22 (F); between Limén & Rio
Banano, Davidse & Pohl 1236 (F, MO); Isla La Uvita, Echeverria 45 (F); 15 km S of
Puerto Limén, Mac Dougal 1190 (DUKE), Wilbur 30599 (DUKE); 5 km S of Puerto
Limon, Wilhur 30676 (DUKE).

PANAMA. Bocas del Toro: Santa Catalina, Blackwell et al. 2697 (NY); Changui-
nola Valley, bar mouth, Dunlap 520 (F, US); Bocas, Godfrey 66317 (FLAS); Laguna
de Chiriqui and vic., Hart 79 (US); Canal Zone: Col6nto Empire, Crawford 579 (PH,
NY); Caladonia Harbor, E/more L36 (MICH, US); 1.5 km SW of mouth of Chagres
River, Nee 8927 (GH, PMA); France Field, Standley 28585 (US); vic. Fort Sherman,
Standley 31208 (US); Colon: Colon, Asplund 15157 (NY); vic. Rio Piedras near Porto
Belo, Blum et al. 2537 (FLAS); Colon, Debeaux 44 (PRC); Maria Chiquita, Ebinger
451 (F); vic. Santa Isabel, Pittier 4175 (US).

6. Kyllinga nudiceps C. B. Clarke ex Standley, Field Mus. Nat. Hist.
Ser. Bot. 4: 199. 1929. Cyperus nudiceps (Standl.) O’Neill, Lflt.
West. Bot. 4: 38. 1944. Type: CosTA RICA, Isla del Coco, Pittier
16272 (US!; Isotypes F!, GH!, LCU!, NY!).

Tufted perennial, (10) 25-40 (55) cm tall. Rhizome short, oblique,
1-3 cm long, (0.5) 2-3 mm thick, closely covered with overlapping,
reddish-brown, broadly ovate scales |-3 mm long. Roots brownish,
finely pubescent especially near the rhizome. Leaves bladeless, the
base of the culms with about 3 reddish-brown, scarious sheaths (0.6)
2-8 cm long, densely reddish spotted along the apical margins. Culms
0.4—-0.8 mm thick just above the apex of the longest sheath, 0.4-0.6
(0.9) mm thick just below the inflorescence. Inflorescence bracts 2 or
3, rather broadly ovate, 1.3-2.8 (3.5) mm long, the cusp an additional
1-2 mm long, |.5-2.5 mm wide, light reddish-brown to sordid whit-
ish, smooth or often sparsely scabrous along the keel distally, erect to
ascendent at anthesis, stiffly erect and clasping the rachis after the
spikelets fall. Spike loosely hemispherical, 3.5-4.5 mm high, 5-6 mm
wide. Rachis cylindric, (0.9) 1.3—1.8 mm long, (0.4) 0.6—0.8 mm thick:
spikelet pedicels rather densely packed, separated by less than their
own width, 0.2-0.35 mm long, 0.15—0.2 mm wide, abaxially with a
conspicuous toothlike scar from the lowest sterile scale. Spikelets
1S—45, elliptic, 2~2.8 mm long, 0.7~1.2 mm wide, dull white to light,

1984] Tucker — Kyllinga 531

greenish-brown. Scales 2-2.4 mm long, 1.4-2.1 mm wide, 7-11
nerved, keel green to dull whitish, smooth, the apiculate apex up to0.1
mm long. Stamens |, 2 or 3; filaments dirty white to light brown, 2-3
mm long; anthers linear-oblong, 0.8-1.1 mm long, the connective tip
reddish, up to 0.1 mm long. Styles 0.4-1.4 mm long; stigmatic
branches 2, 1.4-2 mm long. Achenes lenticular, broadly ovate,
1.1-1.2 mm long, 0.75-0.8 mm wide, the apex obtuse, the style
persistent, the base cuneate to substipitate, surface essentially
smooth, light brown.

DISTRIBUTION. Endemic to Isla del Coco (Cocos Island), Costa
Rica (5°31'N lat., 87°03’W long.), in the Pacific Ocean. No collection
data on habitat or elevation are available, but the highest point on the
island, Cerro Yglesia, is 634 m.

PHENOLOGY. Collections have been made in January, July and
December.

Discussion. Standley inthe publication of the species, suggested
it was most closely related to K. tibialis, as indicated by the leafless
culms. In the material cited, it was noted that K. nudiceps and K.
tibialis both also have pedicellate spikelets. All other New World
species have spikelets that are nearly sessile on the rachis. Standley
stated also that K. nudiceps could be distinguished from K. tibialis by
the former’s lack of inflorescence bracts. This is inaccurate, for
although the inflorescence bracts in K. nudiceps are small, these parts
are no smaller in proportion to the height of the inflorescence than in
K. tibialis. The bracts of K. nudiceps do differ in their cuspidate,
rather than blunt, apices, smooth margins and brownish color from
those of K. tibialis. Also, the bracts of K. nudiceps are not “perfectly
smooth,” as O'Neill and Ayers (1944) state, since the keel is usually
distally scabrellate.

SPECIMENS EXAMINED. Only three collections have been seen. Costa Rica: Isla del
Coco: July 1889, Snodgrass & Heller 944 (GH, K, US); Jan. 1902, Pittier 16272
(Hototype: US; isotypes: F, GH, LCU, NY); 5 Dec. 1959, Klawe 1501 (US).

ACKNOWLEDGMENTS

I thank Prof. Robert L. Wilbur for his advice and support in the
research and writing of this study, and for his continuing help in my
studies of the Cyperaceae in Mexico and Central America. He,

532 Rhodora [Vol. 86

along with Charlotte M. Taylor and John M. MacDougal, made
numerous collections in Costa Rica in 1980 and 1981, which have
been of much importance. Sue Dickerson and Debbie Gooch typed
the manuscript. Dr. P. J. Vorster, University of Stellenbosch, South
Africa, provided a microfiche copy of his thesis. Prof. H. Merx-
muller, Director of Botanische Staatssammlung, Miinchen (M),
kindly sent a copy of Ledebour’s “Dissertatio botanica...”. Mr. N.
Hallé, sous-Directeur, Laboratoire de Phanérogamie, Paris, sent
photographs of the types of Michaux and Lamarck. Specimens of
Kyllinga were borrowed from the following herbaria: A, B, BH,
BM, BR, C, CAS, CGE, CR, CU, DS, DUKE, E, ENCB, F, FLAS.
G, GH, HAL, K, LCU, LE, LL, M, MEXU, MICH, MO, MSC.
NY, PENN, PH, PMA, PR, PRC, S, TEX, US, WIS, and Z. My
thanks go to the curators of these institutions for making the speci-
mens available on loan.

LITERATURE CITED

Ayers, B. 1946. The Genus Ciperus in Mexico. Cath. Univ. Amer. Biol. Stud. I.

DeLanoussaye, A. J. & J.W. THierer. 1967. Cyperus Subgenus Ky /linga (Cype-
raceae) in the Continental United States. Sida 3 (3): 128-136.

GLeason, H. A. & A. CRonguist. 1952. The New Britton and Brown Illustrated
Flora. New York, van Nostrand.

Goprrey, R. K. & J. W. Wooten. 1979. Aquatic Vascular Plants of Southeast-
ern United States. Univ. Georgia Press, Athens.

Kern, J.H. 1974. Cyperaceae. /n: Flora Malesiana 7 (3).

Koyama, T. 1978. Cyperaceae. /n: R. A. Howard, ed. Flora of the Lesser Antilles
3: 220-320.

KOUKENTHAL, G. 1935-36. Cyperaceae: Scirpoideae: Cyperus. In: A. Engler, ed.
Das Pflanzenreich 4 (20).

Lye, K. A. 1981. New Taxa and New Combinations in Kyvllinga. Nord. J. Bot. 1:
741-747.

MOoLINAR., A. 1975. Enumeracion de las plantas de Honduras. Ceiba 19: 1-118,

O'NEILL, H.T. 1940. The Sedges of the Yucatan Peninsula. Carnegie Inst. Wash.
Misc. Papers 19.

.& B. Ayers. 1944. Some New World Cyperi. Lflt. West. Bot. 4: 33-38.

PEDERSEN, T. M. 1970. Cyperaceae. /n: A. Cabrera, ed. Flora de la Provincia de
Buenos Aires. Buenos Aires, I.N.T.A.

Raynat, J. 1973. Contributions a la classification de la sous-famille des Cype-
roideae. Adansonia, sér. 2, 13: 145-171.

SPELLMAN, D. L., J. D. Dwyer & G. Davipse. 1975. A List of the Monocotyle-
doneae of Belize including a historical introduction to plant collecting in Belize.
Rhodora 77: 105-140.

STANDLEY, P.C. 1931. The Cyperaceae of Central America. Fieldiana Bot. 8 (4):
239-292,

1984] Tucker — Kyllinga 533

ss: 1938. Flora of Costa Rica (Cyperus). Fieldiana Bot. 18: 96-100.

_& J. A. STEYERMARK. 1958. Flora of Guatemala (Cyperaceae). Fieldiana
Bot. 24: 90-196.

Svenson, H. K. 1943. Flora of Panama (Cyperaceae). Ann. Mo. Bot. Gard. 30:
281-324.

Tucker. G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and
Panama. Syst. Bot. Monogr. 2: 1-85.

DEPARTMENT OF BOTANY
DUKE UNIVERSITY
DURHAM, NORTH CAROLINA 27706

APPENDIX I
INDEX OF EXSICCATAE

Abbreviations of Ky/linga species

b....K. brevifolia tava Ky tibiaws
n....K. nudiceps p....K. pumila
o....K. odorata v....K. vaginata

Acosta, M. & J. Dorantes. 185 b. Aguilar H., M. 6 0. Arsene, Fr.
5646 0: 5752 0; 10110 0; 11815 b; 11896 b; 12045 b; 7-8-1910 (E) 0;
Apr. 1912 (E) b. Abbott, R. Q. 242 0. Allen, P. H. 106 p; 887 p; 1379
(GH) p, (NY) b, p; 6114 p. Asplund, E. 15157 t. Atwood, J. 2093 p.

Baker, R. & Burger. 22 t. Balls, E. K. B4303 p; B4805 o. Barkely,
F. et al. 2533 0; 7560 o. Berlandier, O. 2130 0. Bernoulli, G. 86 p.
Blackwell, Correa A. & Ridgway. 2697 t. Blake, S. F. 7327 p; 7438
o. Blum, K. E. et al. 2537 t. Boege, W. 1075 p. Botteri, M. 194 p; 195
0; 765 0, p; 773 p; s.n. July 1856 (US) b. Bourgeau, M. 662 0, p;
2737B p; 2989 o. Breedlove, D. E. 6973C 0; 14813 0; 29149 b; 37674
p; 37792 b; 38262 0; 38562 o; 53203 b. Breedlove & Almeda 57709 p.
Breedlove & Davidse 54145 b. Breedlove & Raven. 19913 p. Breed-
love & Thorne 18409 0; 21536 p. Brenckle, J. F. 47-122 b; 47-139 p.
Brenes, A. 14436 0; 15637 p; 17319 0; 19330 b; 19335 p. Burger, W.
7531 p. Burger & Liesner. 6822 p. Burger & Stolze. 5336 b. Burger
et al. 10360 t.

Calderon, S. 35 p. Castillo, G. & L. Tapia 7480. Castro T., N.
191 t. Chickering, A.M. 52 t. Clare, T. 143 t. Clarke, O. F. 501 p.
Conzatti, C. & Gonzalez, J. 648 p. Correa A., M. & B. L. Haines.
245 o. Correa A. et al. 2610 p. Cowan, C. 2440 p; 2468 p. Crawford,

534 Rhodora [Vol. 86

J.579 t. Croat, T. B. 11867 b; 24111 t. Cruz Cisneros, R. 162 b; 1020
0; 1314 0; 2511 0. Cruz V., M. 71 0. Cufodontis, G. 642 b.

Davidse, G. 875 p; 883 p. Davidse & Pohl. 1236 t; 1257 p; 1327 0;
2176 t. Davidson, M. E. 453 p. Davila V., O. 28 July 1963 (ENCB)
o. Deam, C. 10 p; 72 t; 434 p. Debeaux, G. 44 t. Deppe & Schiede
851 0. Detling, L. E. 8491 0. Dieterle, J. V. A. 3912 0; 4402 o.
Donnell Smith, J. 1843 t. Dorantes, B. 2594 p. Dressler, R. 1675 0.
Dressler & Jones. 184 b. Duke, J. A. 4442 0; 12031 p. Duke &
Mussell. 6660 t. Dunlap, V. C. 520 t. Dunn, D. et al. 22960 o.

Ebinger, J. E. 79 0; 451 t. Echeverria, J. A. 45 t; 542 p. Edwards,
M. T. 791 p. Elias, J. 476 0. Elmore, F. H. L36 t.

Fassett, N. C. 28259 0. Feddema, C. 4 0; 254 0; 588 0; 622 0; 1073
o. Fendler, A. 343 p; 349 p. Fisher, G. L. 75 0. Folsom, J. 10113 p.
Fosberg, F. R. 54376 t.

Galeotti, H. 5865 p. Galvan, M. T. 7 Sept. 1963 (ENCB) o.
Garcia, E. R. 103 0. Garcia, M. 22 b. Garnier, A. 40 b; 785 b; 4401
p; 4455 b; 4472 p; s.n. ca. 1930-1940 (F) 0. Gentle, P. 126b; 1398 o:
2370 p; 4973 b. Gentry, A. et al. 7427 t. Gilbert, L. E. 49 o. Glass-
man, S. F. 1657 p; 1744 p. Godfrey, R. K. 66016 p; 66317 t; 67181 p:;
66865 p; 66977 b; 66988 p. Gomez P., L. D. 3279 b. Gonzalez, S.
980 0; 1016 b. Gonzalez Quintero, L. 195 p; 286 0; 1453 b. Grant, V.
540 0. Grayum, M. 1219 p; 2035 p; 2198 b. Greenman, J. M. & M.
T. 5271 t. Gutierrez H., C. 92 0. Guzman C., C. 64 0.

Hammel, B. 8318 p; 9797A b. Harmon, W. E. 2303 b. Harmon &
Dwyer. 2719 o. Hart, J. 79 t; 82 b. Harvey, D. H. 5031 0. Hayes, S.
299 b; 300 p; 841 t; Hedger, J. N. 271 t. Heithaus, E. R. 287 o.
Hepper, D. N. 106 b. Heyde & Lux. 3540 0; 3550 p; 3897 b. Hilerio
A., L.97 0. Hinton, G. B. 1176 0; 9429 0; 13103 0. Hitchcock, A. S.
8443 p; 8506 p. Holm & Iltis. 132 0; 239 o. Howell, J. T. 10175 b.

Johnston, 1. M. 39 0; 1188 p. Jiménez, O. 1112 p. Jiménez M., A.
2623 b; 2838 t: 3856 b.

Kamb, P. 2049 o. Kellerman, W. A. 5133 t; 7407 p. Kerber, E. 52
p. Killip, E. P. 4057 0; 4103 b; 4532 b; 4539 p. King, R. M. 505 o;:
1876 0; 2133 p. King & Soderstrom. 4876 p. Klawe, W. L. 1501 n.
Kral, R. 25518 0; 25583 0; 25659 0; 25599 0; 25669B o. Kuntze, O.
1831 t; 1992 b. Kupper, W. 1394 p.

Lachica, L. & F. Sanchez. 2101 p. Langlassé, E. 326 0. Lankester,
C. H. 231 0. Lazier, T. 10-14 b. Lazor, R. L. 2889 b. Lazor &
Correa A. 2754 p; 2843 p. Lazor et al. 240 t; 2408 v; 2608 b:; 2790 p.
Leavenworth, W. C. 376 0. Leavenworth & Hoogstraal. 1812 o.

1984] Tucker — Kyllinga 535

Lent, R. W. 342 t; 1061 p. Lewis, W. H. et al. 570 p; 2944 b; 2984 p;
3194 t; 5517 p. Liesner, R. 945 b. Lundell, C. L. 1906 b; 2498 p.
4973 b.

MacDaniels, L. H. 125 0; 357 0. Magafia A., M. A. & R. Curiel.
391 p. Manning, W. E. & M.S. 531121 0. Martinez-Calderon, G.
1480 p. Matuda, E. 297 p; 310 b; 3559 b; 26014 p; 31275 p. Maxon,
W. & Harvey. 7896 p. MacDaniels, L. H. 125 0. MacDougal, J. M.
695 b: 755 p; b; 756 0; 861 0; 915 p; 1163 p; 1190 t; 1265 p; 1416 p.
McCorckle, J. S. C-252 b. McDaniel, S. 5083 0; 8120 p. McPher-
son, G. 1129 0. McVaugh, R. 12980 0; 16290A 0; 16559 0; 19298 0;
20232 p. Mexia, Y. 8795 p; 9042 p. Mille Pagaza, S. 68 0. Mohr, C.
& Botteri. 2 b. Molina R., A. 1591 p; 14788 t; 20671 t, 22240 p,
25727 t. Molina R. & Molina. 13997 p. Montes & Salazar. 471 o.
Moore, H. E., Jr. 2953 p. Mori, S. & Bolten. 7394 p. Muller, F.
1995 p. Murry, R. E., Jr. 431 t.

Narvaez, E. 258 b; 2883 b; 2944 t. Nee, M. 7287 b; 8927 t. Nee &
Mori. 3601 b. Nelson, C. 855 p. Nelson, C. & M. Hernandez M. 131
t. Nelson & Romero. 4524 t; 4652 b. Nichols, C. E. 915 b. Norris &
Taranto 13255 b; 13760 0; 13921 b; 14170 0; 14970 o.

Ochoa, A. 75 p. Oersted, A. 14499 p; 14499a p. O'Neill, H. T.
8970 t: 8972 p; 8974 (DS) 0, p (GH, LCU, US) p; 8975 p, 8977 p.
Opler, P. A. 280 0. Ortega, J. G. 4472 o.

Palmer, E. 6 p; 82 p: 192 p; 253a 0; 444 p. Peck, M. E. 39 0; 40 t.
Pefialosa, J. 828 o. Pennington, C. W. 108 o. Pfeifer, H. W. 1352 0:
2008 b. Philbrick, P. N. 733 b. Piper, C. V. 5684 b. Pittier, H. 9218
b: 2403 p; 2445 b, 0; 2580 0; 2583 p (?); 2587 v; 4175 t; 4215 t, 4433
b: 4465 0; 4636 0; 8562 p; 16272 n. Polakowsky, H. s.n. 24 June
1875 (BM) p. Pringle, C. G. 3438 p; 8959 0. Proctor, G. R. 38922 b,
0; 36595 t. Puig, H. 3245 p.

Redmond, R. 8969 v. Rivera, E. 7 p; s.n. 11 Aug. 1974 (PMA) p.
Robertson, J. 103 v. Rodriguez, J. V. 29 p; 274 b. Roe, K. et al. 560 o.
Rohweder, O. 2176 b; 2185 b. Rose, J. N. 1803 0; 22086 0; 23996 b.
Ross, G. B. 8-57 p. Rossbach, G. B. 3407 0; 3656 p. Rovirosa, J. 426
p; 708 b. Rzedowski, J. 1642 0; 17203 b; 18940 p; 19057 p; 24202 0;
24248 0; 27640 0; 27714 p; 31257 0; 35296 o.

Santos, J. V. 3316 o. Saunders, J. 142 t. Sawyer, J. L. s.n. 2
March 1967 (DS, WIS) b. Schaffner, J. G. 137 p. Schiede, C. J. W.
s.n., July 1829 (HAL) b. Schiede & Deppe. s.n. Oct. 1829 (BM) b.
Schipp, W. A. 382 t. Schmidt, M. s.n. in 1936 (B) p. Schmitz, J. 809
p. Seaton, H. E. 292 (F, US) p (GH) o. Seibert, R. J. 237 p; 474 o.

536 Rhodora [Vol. 86

Seler, E. 801 0; 1887 p; 2566 p. Seymour, F. C. 641 t; 718 p; 2785 p;
2972 0. Sharp, A. J. et al. 44255 0; 45935 p; 52029 0. Skutch, A. F.
2643 0; 2859 b. Smith, W. T. 55 0. Snodgrass, R. E. & E. Heller. 944
n. Spellman, D. L. & Stoddart. 2171 t; 2343 t; 2381 t; 2400 t; 2469 t.
Sperry, J. 1114 p. Standley, P. C. 1682 p; 12048 b; 12857 0; 19348 p;
19842 p; 20290 b; 20484 b; 20788 b; 21088 p; 21157 b; 21558 p;
21696 0; 21773 b; 21696 0; 21773 b; 21791 p; 21810 0; 22420 p;
22511 0; 22907 p; 23205 b; 23291 p; 23467 p; 24066 b; 24069 o:
25154 (GH) b, p (NY) b; 24356 p; 24777 b; 25057 0; 25139 t; 25168
p; 25710 p; 27276 p; 28585 t; 28807 0; 28988 0; 29434 p; 30885 b:
31208 t; 32240 p; 32742 (US) b, p; 36730 p; 40090 p; 41120 p; 41240
p; 43508 p; 53001 t; 53739 (F, NY) b (US) p; 53800 t; 55701 b, 58106
b; 63023 p; 64513 b; 64664 p; 72157 v; 72160 b; 74272 p; 72487 b:
82140 p; 82650 p. Stern, W. L. et al. 836 p. Stevens, W. D. 10299 o.
Steyermark, J. A. 29708 0; 30529 0; 31729 b; 36724 b; 39710 t:
46279 b; 49592 o. Stork, H. E. 2727 p; 3189 t. Svenson, H. K. 4495
oO.

Tate, R. 506 v; 515 t. Taylor, J. 17348 p. Taylor & Taylor. 11344
b. Thomas, C. H. 66 0. Thorne, R. F. & E. Lathrop. 41035 p. Ton,
A.S. 1685 p; 2755 b. Tonduz, A. 434 0; 672 0; 4793 p; 8180 p; 8264
p; 8849 0; 9122 t; 9218 b; 11388 b; 17906 p; 17907 p. Tucker, G. C.
2077 0; 2088 p; 2135 p; 2160 p; 2219 b; 2222 0; 2252 p; 2253 b.
Tucker, J. M. 850 b; 1326 p. Tyson, E. L. 1410 0; 1600 o. Tyson &
Smith. 4163 p. Tyson et al. 4301 p; 4794 p.

Valerio, M. 294 p. van der Sluijs, D. H. S. 791 b: S.799 t.
Vaughan, J. et al. 1086 0. von Tiirckheim, H. 6 p; 663 p; 3586 b;
7691 0; 7691B (US) p, (K) 0; 8843 t; 11 2271 p.

Wagner, M. 1-1/2 0. Webster, G. L. 12654 t. Wercklé, C. 664 p.
Whitefoord, C. 1077 p; 2390 p. Wilbur, R. L. 14701 p; 21311 p;
21614 p; 22319 p; 22455 p; 22707 p; 24899 p; 26327 b; 27511 o:
28020 (DUKE) b, p; 28941 p; 28977 p; 29439 p; 29755 p; 29946 p;
29960 0; 29970 p; 30599 t; 30676 p; 30743 p; 30752 b: 30876 p; 31027
b; 31028 p; 32125 p; 32735 p; 33005 p; 33200 p; 33275 b; 33277 b;
33313 p; 33338 p; 33629 p. Wilbur & Luteyn. 11650 (DUKE) b, p;
18888 p; 19275 p. Wilbur et al. 11998 b; 13493 p; 15316 p. Williams,
L. O. et al. 16893 0; 24325 b; 27960 (F) b, p. Williams, R.S. 1040 b.
Wilson, P. 199 0; s.n. 16 Jan. 1903 (NY) t. Woodson, R. E.& R. W.
Schery. 557 p; 770 b. Woodson et al. 1155 b; 1773 (GH) b, 0; 1774 b.
Worth, C. R. 8906 b.

1984] Tucker — Kyllinga 537

Yong, G. s.n. 18 May 1977 (PMA) p. Yuncker, T. G. 4581 (F,
MICH) o, p; 4674 t; 8247 t.
Zelaya M., H. 2298 b. Zuill, H. 575 p.

APPENDIX II
INDEX TO NAMES IN THIS TREATMENT

The number in parentheses following each binomial tells to which
species it is referred in the present treatment.

(C = Cyperus; K = Kyllinga; M = Mariscus)

C. brevifolius (Rottb.) Hassk. (3)

C. densicaespitosus Mattf. & Kiikenth. (1)
C. nudiceps (Standl.) O’Neill (6)

C. obtusatus (Presl) Mattf. & Kiikenth. (4)
C. obtusatus var. cylindrostachyus (Boeck.) Kiikenth. (4)
C. peruvianus (Lam.) F. N. Williams (4)

C. peruvianus var. foliatus (Kiikenth.) Kiikenth. (5)
C. tenuifolius (Steud.) Dandy (1)

K. aphylla (Vahl) Kunth (5)

K. brevifolia Rottb. (3)

K. caespitosa Nees (1)

K. caespitosa var. major Nees (1)

K. capitata P.-Beauv. (5)

K. cruciformis Schult. (3)

K. elongata H. B. K. (3)

K. flexuosa Boeck. (1)

K. globosa P.-Beauv. (5)

K. laxa Schrad. ex Nees (1)

K. leucocephala Baldw. (2)

K. martiana Schrad. ex Nees (2)

K. monocephala Thunb. (3)

K. monocephala H. B. K. (2)

K. nudiceps Standl. (6)

K. obtusata Presl. (4)

K. obtusata var. cylindrostachyus Boeck. (4)
K. odorata Vahl (2)

K. odorata var. minor Boeck. (2)

K. odorata var. rigida Boeck. (2)

538 Rhodora [Vol. 86

K. peruviana Lam. (4)

K. peruviana var. foliata Kiikenth. (5)
K. pumila Michx. (1)

K. pumila B. b. elatior Kunth (1)

K. pumila var. elatior Boeck. (1)

K. pungens Link (4)

. rigida Baldw. (4)

K. sesquiflora Torr. (2)
K. sororia Kunth (3)
K. tenuifolia Steud. (1)
K. tenuis Baldw. (3)

K. tenuis Boeck. (4)
K
K
M.

~

. tibialis Ledeb. (5)
. vaginata Lam. (4)
aphyllus Vahl (5)

NEW ENGLAND NOTE

THE RE-DISCOVERY OF SOLIDAGO ODORA AIT.
(ASTERACEAE) IN VERMONT

PETER F. ZIKA

Solidago odora Ait. is at the northern limit of its range in south-
eastern Vermont and is on the Vermont rare and endangered
species list (Countryman, 1978). Oakes (1842) mentioned the spe-
cies in his catalogue of Vermont plants, but by the time Brainerd et
al. (1900) compiled the Vermont flora, the vouchers could not be
located. The only specimens that have survived are those W. H.
Blanchard collected in September 1901 from Vernon, Windham
Co. (Eggleston et al., 1915). Blanchard found “one large clump of
plants”, in the vicinity of Lespedeza Xnuttalii Darl. and Aureolaria
pedicularia (L.) Raf., at the border of Line’s Woods. In October of
1982 I rediscovered a colony of 30 S. odora at the edge of open, dry
oak woods about one mile north of Lily Pond, in the central part of
Vernon [Zika 6758 (VT)]. Nearby were Aureolaria pedicularia,
Lespedeza hirta (L.) Hornemann and a small colony of Bartonia
virginica (L.) BSP. All of these species are rare plants in Vermont.

Blanchard also collected Solidago odora in 1901 from the east
side of the Connecticut River, on the Drewsville Plains and on Fall
Mountain, in Walpole, Cheshire Co., New Hampshire [ Blanchard
s.n. (VT)]. In view of the species persistence in Vernon, it seems
likely it will be rediscovered in nearby Walpole as well.

LITERATURE CITED

BRAINERD, E., L. R. Jones, & W. W. EGGLeston. 1900. Flora of Vermont. A List
of the Fern and Seed Plants Growing Without Cultivation. Free Press Associa-
tion, Burlinton, Vt. 113 pp.

CouNTRYMAN, W. D. 1978. Rare and Endangered Vascular Plant Species in
Vermont. The New England Botanical Club in cooperation with the U. S. Fish
and Wildlife Service, Newton Corner, Mass. 68 pp.

EGGLESTON, W. W.,G.L. KIRK AND J.G. UNDERWOOD. 1915. Flora of Vermont.
List of Fern and Seed Plants Growing Without Cultivation. Vermont Agricul-
tural Experiment Station Bulletin 187: 139-258.

Oakes, W. 1842. Catalogue of Vermont plants. /n: Thompson, Z. History of
Vermont, Natural, Civil, & Statistical. Published by the author, Burlington, Vt.
pp. 173-208.

PRINGLE HERBARIUM
BOTANY DEPT.
UNIVERSITY OF VERMONT
BURLINGTON, VT 05405
539

NOTICE OF PUBLICATION

THE CONNECTICUT BOTANICAL SOCIETY, INC.
Memoirs of the Connecticut Botanical Society
No. |. April, 1984

A CONVERSATION WITH HARRY AHLES
(1924-1981)

Harry Ahles was well-known in the world of botanists as an in-
defatigable collector and a person of unusually broad knowledge.
As the curator for the University of Massachusetts Herbarium,
Amherst, he was composing a flora of New England at the time of
his death. In August of 1980, three Connecticut Botanical Society
members met with Ahles and recorded his comments on the flora of
Connecticut. As the species were discussed (in phylogenetic order)
he revealed many clues to aid in field identification and to the
separation of confusing look-alikes. He described key characters,
habitat requirements, and ranges and also gave his opinions on the
status of various taxa in Connecticut and Massachusetts. A tran-
script was made of that lengthy conversation and is presented in this
Memoir in edited and annotated form. It is the first publication of a
series devoted to providing information to all who study and appre-
ciate plants. To order: send check or money order for $3.50 to:

MEMOIR

The Connecticut Botanical Society, Inc.

c/o Yale University Herbarium

Osborn Memorial Laboratory

167 Prospect Street

New Haven, CT 06511

540

AN AWARD FOR THE SUPPORT OF
BOTANICAL RESEARCH
IN NEW ENGLAND, U.S.A.

The New England Botanical Club is offering an award of $1,000
in support of botanical research to be conducted in the New
England region during 1985. It is being made to stimulate and
encourage botanical research on the New England flora and to
make possible visits to the New England region by those who would
not otherwise be able to do so. The award will be given to the
graduate student submitting the best research proposals dealing
with field studies in systematic botany and plant ecology, but
proposals for research in other areas of botany will also be
considered. This award is not limited to graduate students at New
England institutions. The NEBC’s support must be acknowledged in
any publications resulting from this study. It is encouraged that
papers based on this research be submitted to RHODORA, the
Club’s journal, for possible publication—subject to standard review
processes. The New England Botanical Club hopes to be able to
make this award on an annual basis.

Applicants should submit a proposal of no more than three
double spaced pages, including a budget (the budget will not affect
the amount of the award), and their Curriculum Vitae. Two letters,
one from the student’s major professor, in support of the proposed
research are also required. Proposals and supporting letters should
be sent before 28 February 1985 to: Awards Committee, The New
England Botanical Club, 22 Divinity Avenue, Cambridge, MA
02138. The recipient of the award will be notified by 30 April 1985.

54]

Joint Meeting: June 23-27, 1985

The annual Joint Field Meeting of
The Northeastern Section of the Botanical Society of America
The Torrey Botanical Club, and
The Philadelphia Botanical Club
will be held in East Stroudsburg, PA. Accommodations will be at
East Stroudsburg University.

There will be field trips to boreal bogs, limestone dells, upland
forests and swamps at various sites in the Pocono Mountains and
Delaware Water Gap area of Pennsylvania, as well as adjacent New
Jersey.

Space is limited and PRIOR registration is required. Full details
will be available after February |, 1985, by writing the chairman:

James K. McGrath, Vice President
Delaware Valley Conservation Society
Box 393

Lansdowne, PA. 19050

542

MEETING ANNOUNCEMENT

EIGHTH ANNUAL CONFERENCE ON THE
CONNECTICUT RIVER ECOSYSTEM

The Eighth Annual Research Conference on the Connecticut
River Ecosystem will be held on Saturday, 27 April 1985. Anyone
currently conducting research on the geology, botany, zoology, or
ecology of the Connecticut River, its floodplain, or associated
wetlands is encouraged to present a paper at the meetings.

The conference will be held at the Mountain School in Vershire,
Vermont. The school is easily accessible from Route 91 in rural
eastern Vermont, and is located.on a tributary of the Connecticut
River. The Mountain School has had a long-standing commitment
to environmental education, and presently serves as focal point for
the environmental issues and farm programs of Milton Academy.

For additional information contact:

Dr. Kevin Mattingly or Dr. Marjorie Holland

Environmental Sciences Dept. of Biology

The Mountain School Program — School of Arts and Sciences
of Milton Academy College of New Rochelle

Vershire, Vermont 05079 New Rochelle, N.Y. 10801

543

INSTRUCTIONS TO CONTRIBUTORS TO RHODORA

Submission of a manuscript implies it is not being considered for
publication simultaneously elsewhere, either in whole or in part.

Manuscripts should be submitted in triplicate (an original and
two xerox copies) and must be double-spaced (at least 3/8 of an
inch) throughout including footnotes, figure legends, and refer-
ences. Please do not use corrasable bond. The list of legends for
figures and maps should be provided on a separate page. Footnotes
should be used sparingly. Do not indicate the style of type through
the use of capitals or underscoring, particularly in the citation of
specimens. Names of genera and species may be underlined to indi-
cate italics in discussions. Specimens citations should be selected
critically, especially for common species of broad distributions. Sys-
tematic revisions and similar papers should be prepared in the for-
mat of “A Monograph of the Genus Malvastruin”, S.R. Hill,
Rhodora 84: 1-83, 159-264, 317-409, 1982, particularly with refer-
ence to indentation of keys and synonyms. Papers of a floristic
nature should follow, as far as possible, the format of “Annotated
list of the ferns and fern allies of Arkansas”, W. Carl Taylor and
Delzie Demaree, Rhodora 81: 503-548, 1979. For bibliographic cit-
ations, refer to the Botanico-Periodicum-Huntianum (B-P-H,
1968), which provides standardized abbreviations for journals
originating before 1966. All abbreviations in the text should be
followed by a period, except those for standard units of measure
and direction (compass points). For standard abbreviations and for
guidance in other matters of biological writing style, consult the
CBE Style Manual, Sth ed. (original title: Style Manual for
Biological Journals). In preparing figures (maps, charts, drawings,
photos, etc.) please remember that the printed plate will be 4 x 6
inches; be sure that your illustrations are proportioned to reduce
correctly, and indicate by blue pencil the intended limits of the
figures. (Some “turn-page” figures with brief legends will be 3 1/2 x
6 in.) Magnification/ reduction values given in text or figure legends
should be calculated to reflect the actual printed size. An Abstract
and a list of Key Words should be supplied at the beginning of each
paper submitted, except for a very short article or note.

544

LIST OF REVIEWERS OF MANUSCRIPTS
NOV. 1, 1983-NOV. 1, 1984

The editors of RHODORA are grateful to each of the following
specialists listed below for their participation in the reviewing

process.
Ralph M. Adams
Kelly W. Allred

David S. Barrington
Donovan R. Bowley
Errol Briggs

Ralph E. Brooks
William C. Burger
George L. Church
Thomas C. Dent
Glenn Dreyer
Lesley M. Eastman
George S. Ellmore
Leslie A. Garay
Robert R. Haynes
C. Barre Hellquist
Walter M. Hewitson
Harold R. Hinds
Shiu Ying Hu
Robert Kral
Thomas Lee
Michael W. Lefor
Walter H. Lewis

Leslie J. Mehroff
Richard S. Mitchell
John K. Morton
William H. Murdy
Richard W. Pohl
Anton A. Reznicek
G. K. Rogers
Ruben P. Sauleda
Alfred E. Schuyler
John C. Semple
Bruce A. Sorrie
Robin South
Herman R. Sweet
John W. Thieret
B. L. Turner
Charles H. Uhl
Lowell E. Urbatsch
Russell R. Walton
Daniel B. Ward
Gerald A. Wheeler
Carroll E. Wood

Hovora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

NORTON H. NICKERSON, Editor-in-Chief
JOAN Y. NICKERSON, Managing Editor

A. LINN BOGLE RICHARD A. FRALICK
WILLIAM D. COUNTRYMAN GERALD J. GASTONY
GARRETT E. CROW NORTON G. MILLER

ROBERT T. WILCE

VOLUME 86

1984

Che New Lugland Rotanical Club, Ine.
Harvard University Herbaria, 22 Divinity Ave., Cambridge, Mass. 02138

INDEX TO VOLUME 86

New scientific names are in bold face.

Adams, Ralph M. see Sauleda, Ruben P.

Additions to the flora of Cape Breton
Highlands National Park, Nova Sco-
tia 67-71

Additions to the flora of Ulster County,
New York 89-94

Adirondack mountain summit. A 24-
year comparison of the vegetation of,
439

Allred, Kelly W. Studies in the Aristida
(Gramineae) of the southeastern Unit-
ed States I. Spikelet variation in A.
purpurescens, A. tenuispica and A.
virgata 73-77

Angelo, Ray and Bruce A. Sorrie. Nan-
tucket field trip of the New England
Botanical Club 117

Annotated list of Minnesota Carices,
with phytogeographical and ecologi-
cal notes 151-231. Floristics 152;
annotated list 169; doubtful and ex-
cluded taxa 200; distribution maps
203-226

Aristida (Gramineae) of the southeastern
United States I. Spikelet variation in
A. purpurescens, A. tenuispica and A.
virgata. Studies in the, 73-77

Aristolochia tomentosa Sims established
at two western Massachusetts sites
115

Arnica (Compositae) subgenus Austro-
montana. Taxonomy of, 239-309

Bates, Vernon and Paul Lewis. Redis-
covery of Stylisma humistrata (Con-
volvulaceae) in Tennessee 393-394

Book review: Where have all the wild-
flowers gone? 237

Broughtonia R. Br., Cattleyopsis Lem.
and Laeliopsis Lind]. A reappraisal of
the orchid genera, 445

Broughtonia <jamaicensis Sauleda and
Adams, hybr. nov. 453

Burk, C. John. Aristolochia tomentosa
Sims established at two western Mas-
sachusetts sites 115

Burk, C. John. Geranium nepalense var.
Thunbergii and G. sibiricum natural-
ized in western Massachusetts 113

Burk, C. John see Holland, Marjorie
M.

Campbell, Christopher S. see Famous,
Norman C.

Cape Breton Highlands National Park,
Nova Scotia. Additions to the flora
of, 67-71

Catling, Paul M. A re-evaluation of Spi-
ranthes Xsteigeri Correll 469

Cat-tail (Typha latifolia L.) marsh. The
effect of power utility right-of-way
construction on, 389-39]

Cattleyopsis cubensis (Lind].) Sauleda &
Adams, comb. nov. 455

Cattleyopsis Lem. and Lailiopsis Lindl.
A reappraisal of the orchid genera
Broughtonia R. Br., 445

Cayman Islands. A new species of Phyl-
lanthus (Euphorbiaceae) from the,
121-125

Central America. A revision of the genus
Kyllinga Rottb. (Cyperaceae) in Mex-
ico and, 507

Central American species of Cerastium
(Caryophyllaceae). A revision of the
Mexican and, 339-379

Cerastium (Caryophyllaceae). A revision
of the Mexican and Central American
species of, 339-379

Cerastium cuchumatanense D. A. Good,

349: C. sinaloense D. A.
Good, sp. nov, 367; C. tolucense
D. A. Good, sp. nov. 371; fig. 350

Chaptalia (Asteraceae: Mutisieae) from
Mexico. A new widespread species
of, 127-130

sp. nov.

549

550

Chaptalia transiliens Nesom, sp. nov.
127

Chromosome number in the genus Stel-
laria (Caryophyllaceae). A new basic,
417

Connecticut River oxbow swamp forests.
The herb strata of three, 397

Croomia pauciflora (Stemonaceae).
Notes on, 131-137

Denford, Keith E. see Wolf, Steven J.

Dentaria laciniata Muhl. (Brassicaceae)
reported from Maine. Note on the first
station of, 233

Distribution and ecological characteris-
tics of Ironwood, Ostrya virginiana
(Miller) K. Koch, in northeastern Nova
Scotia 139-149

Distributions. Lomatogonium rotatum
(Gentianaceae) and Primula laurenti-
ana (Primulaceae) in Maine: new local-
ities and general, 425

Domville, Mary. Additions to the flora
of Ulster County, New York 89-94

Ecological notes. Annotated list of Min-
nesota Carices, with phytogeographi-
caland, 151-231

Effect of power utility right-of-way con-
struction on cat-tail (Typha latifolia
L.) marsh. The, 389-39]

Environmental corollaries of Panax quin-
quefolium (Araliaceae) in Delaware
County, New York. Population struc-
ture and, 431

Erratum, October 1983 issue (Vol 85, No.
844), Dent and Adams’ 119

Famous, Norman C. and Christopher S.
Campbell. Lomatogonium rotatum
(Gentianaceae) and Primula laurenti-
ana(Primulaceae) in Maine: new local-
ities and general distributions 425

Farrar, Donald R. see Parks, James
C;

First report of the fern genus Vittaria in
New York. A, 421

Rhodora

[Vol. 86

Flora boreali-americana of André
Michaux. The type localities of the,
1-66

Florida. A new species of Ziziphus
(Rhamnaceae) from, 381-387

Geranium nepalense var. Thunbergii and
G. sibiricum naturalized in western
Massachusetts 113

Geranium sibiricum naturalized in west-
ern Massachusetts. Geranium nepa-
lense var. Thunbergii and, 113

Good, David A. A revision of the Mexi-
can and Central American species of
Cerastium (Caryophyllaceae) 339-
379

Gratiola viscidula Pennell (Scrophulari-
aceae). Infraspecific variation in,
79-87

Greenidge, K. N. H. Distribution and
ecological characteristics of Ironwood,
Ostrya virginiana (Miller) K. Koch, in
northeastern Nova Scotia 139-149

Halenia alleniana Standl. ex Wilbur, sp.
nov. 326; H. crumiana Wilbur, sp.
nov. 321

Halenia (Gentianaceae) in Mexico, A
synopsis of the genus, 311~—337

Hall, David W. see Judd, Walter S.

Herb strata of three Connecticut River
oxbow swamp forests. The, 397. Flo-
ristic trends and comparisons 409:
Study sites: Hatfield oxbow 405,
Ned’s Ditch 406, Whately oxbow
409; Table: Composition of herb strata:
herbs and vines 402, woody seed-

lings 404
Hellquist, C. Barre. Observations on Po-
tamogeton hillii Morong in North

America 101-111

Hinds, Harold L. Additions to the flora
of Cape Breton Highlands National
Park, Nova Scotia 67-71

Holland, Marjorie M. and C. John Burk,
The herb strata of three Connecticut
River oxbow swamp forests 397

1984]

Infraspecific variation in Gratiola viscid-
ula Pennell (Scrophulariaceae) 79-87

Ironwood, Ostrya virginiana (Miller) K.
Koch, in northeastern Nova Scotia.
Distribution and ecological character-
istics of, 139-149

Judd, Walter S. and David W. Hall. A
new species of Ziziphus (Rhamnaceae)
from Florida 381-387

Ketchledge, E. H. and R. E. Leonard. A
24-year comparison of the vegetation
of an Adirondack mountain summit
439

Kyllinga Rottb. (Cyperaceae) in Mexico
and Central America. A revision of the
genus, 507

Laeliopsis Lindl. A reappraisal of the
orchid genera Broughtonia R. Br.,
Cattleyopsis Lem, and, 445

Lassetter, J. Stuart. Taxonomy of the
Vicia ludoviciana complex 475

Leonard, R. E. see Ketchledge, E. H.

Lewis, Paul see Bates, Vernon

Lewis, Walter H. Population structure
and environmental corollaries of Pan-
ax quinquefolium (Araliaceae) in Del-
aware County, New York 434

Lomatogonium rotatum (Gentianaceae)
and Primula laurentiana(Primulaceae)
in Maine: new localities and general
distributions 425

Maine: new localities and general distri-
butions. Lomatogonium rotatum
(Gentianaceae) and Primula laurenti-
ana (Primulaceae) in, 425

Maine. Note on the first station of Den-
taria laciniata Muhl. (Brassicaceae)
reported from, 233

Massachusetts. Geranium nepalense var.
Thunbergii and G. sibiricum natural-
ized in western, 113

Massachusetts sites. Aristolichia tomen-
tosa Sims established at two western,

115

Index to Volume 85 551

Medeolaria farlowii in New England.
Two new localities for, 235

Mexican and Central American species
of Cerastium (Caryophyllaceae). A re-
vision of the, 339-379

Mexico. A new, widespread species of
Chaptalia (Asteraceae: Mutisieae) from,
127-130

Mexico. A synopsis of the genus Halenia
(Gentianaceae) in, 311-337

Mexico and Central America. A revision
of the genus Kyllinga Rottb. (Cypera-
ceae) in, 507

Michaux, André. The type localities of
the Flora boreali-americana of, 1-66

Minnesota Carices, with phytogeograph-
ical and ecological notes. Annotated
list of, 151-231

Morton, J.K. A new basic chromosome
number in the genus Stellaria (Caryo-
phyllaceae) 417

Nantucket field trip of the New England

Botanical Club 117

NEBC Research Award Notice 126,

395, 541

Nesom, Guy L. A new, widespread spe-
cies of Chaptalia (Asteraceae: Muti-

sieae) from Mexico 127-130

New basic chromosome number in the
genus Stellaria (Caryophyllaceae). A,
417

New England. Two new localities for
Medeolaria farlowi in, 235

New England Botanical Club. Nantucket
field trip of the, 117

New England Notes:

Aristolochia tomentosa Sims estab-
lished at two western Massachu-
setts sites 115

Geranium nepalense var. Thunbergii
and G. sibiricum naturalized in west-
ern Massachusetts 113

Nantucket field trip of the New Eng-
land Botanical Club 117

Note on the first station of Dentaria
laciniata Muhl. (Brassicaceae) re-
ported from Maine 233

D232

The re-discovery of Solidago odora
Ait. (Asteraceae) in Vermont 539
Two new localities for Medeolaria far-
lowil in New England 235

New localities and general distributions.
Lomatogonium rotatum (Gentiana-
ceae) and Primula laurentiana (Primu-
laceae) in Maine: 425

New species of Phyllanthus (Euphorbia-
ceae) from the Cayman Islands. A,
121-125

New species of Ziziphus (Rhamnaceae)
from Florida. A, 381-387

New, widespread species of Chaptalia
(Asteraceae: Mutisieae) from Mexico.
A, 127-130

New York. A first report of the fern genus
Vittaria in, 421

New York. Additions to the flora of UI-
ster County, 89-94

New York. Population structure and
environmental corollaries of Panax
quinquefolium (Araliaceae) in Dela-
ware County, 431

Nickerson, N. H. see Thibodeau,
F.R.
North America. Observations on Pota-
mogeton hillii Morong in, 101-111
Note on the first station of Dentaria laci-
niata Muhl. (Brassicaceae) reported
from Maine 233

Notes on Croomia pauciflora (Stem-
onaceae) 131-137

Nova Scotia. Additions to the flora of
Cape Breton Highlands National Park,
67-71

Nova Scotia. Distribution and ecological
characteristics of Ironwood, Ostrya
virginiana (Miller) K. Koch, in north-
eastern, 139 149

Observations on Potamogeton hillii Mo-
rong in North America’ 101-111

Ostrya virginiana (Miller) K. Koch, in
northeastern Nova Scotia. Distribu-
tion and ecological characteristics of
Ironwood, 139-149

Rhodora

[Vol. 86

Ownbey, Gerald B. see Wheeler, Gerald
A.

Panax quinquefolium (Araliaceae) in Del-
aware County, New York. Population
structure and environmental corolla-
ries of, 431

Parks, James C. and Donald R. Farrar.
A first report of the fern genus Vittaria
in New York 421

Pfister, Donald H. Two new localities for
Medeolaria farlowii in New Eng-
land 235

Phyllanthus caymanensis Webster & Proc-
tor, sp. nov. 121; figs. 122, 124

Phytogeographical and ecological notes.
Annotated list of Minnesota Carices,
with, 151-231

Population structure and environmental
corollaries of Panax quinquefolium
(Araliaceae) in Delaware County, New
York 431

Potamogeton hillii Morong in North
America. Observations on, 101-111

Primula laurentiana (Primulaceae) in
Maine: new localities and general dis-
tributions. Lomatogonium rotatum
(Gentianaceae) and, 425

Proctor, George R. see Webster, Grady
L.

Reappraisal of the orchid genera
Broughtonia R. Br., Cattleyopsis Lem.
and Laeliopsis Lindl. A, 445. Key to
the genera 447; B. sanguinea 449;
negrilensis 451, Xjamaicensis Saul-
eda & Adams hybr. nov. 453; C.
cubensis Sauleda & Adams comb.
nov. 455, lindenii 458, ortgiesiana
461; L. domingensis 464

Rediscovery of Stylisma humistrata (Con-
volvulaceae) in Tennessee 393-394

Re-discovery of Solidago odora Ait. (As-
teraceae) in Vermont. The, 539

Re-evaluation of Spiranthes Xsteigeri
Correll. A, 469

1984]

Reinartz, James A. Verbascum densiflo-
rum in southeast Wisconsin 95-99
Revision of the genus Kyllinga Rottb.
(Cyperaceae) in Mexico and Central
America. A, 507. Key to the spe-
cies 513; brevifolia 522; nudiceps
530; odorata 518; pumila 514; tibi-
alis 527; vaginata 525

Revision of the Mexican and Central
American species of Cerastium (Caryo-
phyllaceae). A, 339-379. C. axillare
346: barberi 346; brachypodum
348; cuchumatanense D. A. Good, sp.
nov. 349; glomeratum 352; guate-
malense 355;juniperorum 357; ma-
drense 358; nutans 359; orithales
362; purpusii 364; ramigerum 365,
sinaloense D.,A. Good, sp.nov. 367;
sordidum 368: texanum 369; tolu-
cense D. A. Good, sp.nov. 371; trivi-
ale 372; vulcanicum 374; excluded
species 375; key tothe species 341

Right-of-way construction on cat-tail
(Typha latifolia L.) marsh. The effect
of power utility, 389-391

Rooney, Sally C. Note on the first station
of Dentaria laciniata Muhl. (Brassica-
ceae) reported from Maine 233

Sauleda, Ruben P. and Ralph M. Adams.
A reappraisal of the orchid genera
Broughtonia R. Br., Cattleyopsis Lem.
and Laeliopsis Lindl. 445

Solidago odora Ait. (Asteraceae) in Ver-
mont. The re-discovery of, 539

Sorrie, Bruce A. see Angelo, Ray

Spiranthes Xsteigeri Correll. A reevalu-
ation of, 469

Spooner, David M. Infraspecific varia-
tion in Gratiola viscidula Pennell
(Scrophulariaceae) 79-87

Statement of ownership, inside back
cover, No. 848

Stellaria (Caryophyllaceae). A new basic
chromosome number in the genus,
417

Studies in the Aristida (Gramineae) of
the southeastern United States I.

Index to Volume 85

eee

Spikelet variation in A. purpurescens,
A. tenuispica, and A. virgata 73-77

Stylisma humistrata (Convolvulaceae) in
Tennessee. Rediscovery of, 393 394

Swamp forests. The herb strata of three
Connecticut River oxbow, 397

Synopsis of the genus Halenia (Gen-
tianaceae)in Mexico.A, 311-337.H.
alata 327; alleniana Wilbur sp. nov.
326; brevicornia 315;conzattii 324;
crumiana Wilbur sp. nov. 321; de-
cumbens 332; hintonu 331; pal-
meri 323; plantaginea 334; pringlei
328: recurva 320; schiedeana 325;
key to the species 313

Taxonomy of Arnica (Compositae) sub-
genus Austromontana 239-309. A.
cernua 258; cordifolia 262; discoi-
dea 272; gracilis 277; latifolia
282: nevadensis 290; spathulata
295: venosa 300; viscosa 303; chro-
mosomenumbers 241; excluded taxa
306; key to the species 257; mor-
phology and taxonomic criteria 242;
phylogeny and phytogeography 248

Taxonomy of the Vicia ludoviciana
complex (Leguminosae) 475. Key to
subspecies and races 486; § leaven-
worthii 499; § ludoviciana 487

Tennessee. Rediscovery of Stylisma hum-
istrata(Convolvulaceae)in, 393-394

Thibodeau, F. R. and N. H. Nickerson.
The effect of power utility right-of-
way construction on cat-tail (Typha
latifolia L.) marsh 389-391

Tucker, Gordon C. A revision of the
genus Kyllinga Rottb. (Cyperaceae) in
Mexico and Central America 507

Twenty-four year comparison of the veg-
etation of an Adirondack mountain
summit. A, 439

Two new localities for Medeolaria far-
lowil in New England 235

Type localities of the Flora boreali-
americana of André Michaux 1-66.
Michaux Vol. I 6; Michaux Vol.
Il 40

554 Rhodora

Ulster County, New York. Additions to
the flora of, 89-94

United States. Studies in the Aristida
(Gramineae) of the southeastern,
73-77

Uttal, Leonard J. The type localities of
the Flora boreali-americana of André
Michaux 1-66

Verbascum densiflorum in southeast Wis-
consin 95-99

Vermont. The re-discovery of Solidago
odora Ait. (Asteraceae) in, 539

Vicia ludoviciana complex (Legumino-
sae). Taxonomy of the, 475

Vittaria in New York. A first report of the
fern genus, 421

Webster, Grady L. and George R. Proc-
tor. A new species of Phyllanthus
(Euphorbiaceae) from the Cayman Is-
lands 121-125

[Vol. 86

Wheeler, Gerald A. and Gerald B. Own-
bey. Annotated list of Minnesota Car-
ices, with phytogeographical and eco-
logical notes 151-231

Whetstone, R. David. Notes on Croomia
pauciflora (Stemonaceae) 131-137

Wilbur, Robert L. A synopsis of the
genus Halenia (Gentianaceae) in Mex-
ico. 311-337

Wisconsin. Verbascum densiflorum in
southeast, 95-99

Wolf, Steven J. and Keith E. Denford.
Taxonomy of Arnica (Compositae)
subgenus Austromontana. 239-309

Zika, Peter F. The re-discovery of Soli-
dago odora Ait. (Asteraceae) in Ver-
mont 539

Ziziphus celata Judd and Hall, sp. nov.
382, fig. 384

Ziziphus (Rhamnaceae) from Florida. A
new species of, 381-387

Vol 86, No. 847, including pages 239-395, was issued September 26, 1984

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3526 (See instruction on reverse) f

RHODORA October 1984 Vol. 86, No. 848

CONTENTS

The herb strata of three Connecticut River oxbow swamp forests
Marjorie M. Holland and C. John Burk
A new basic chromosome number in the genus Stellaria (nrstahitincese)
J. K. Morton F
A first report of the fern rat Vittaria in ie You
James C. Parks and Donald R. Farrar *
Lomatogonium rotatum (Gentianaceae) and Fens hie:
(Primulaceae) in Maine: new localities and general distributions
Norman C. Famous and Christopher S. Campbell
Population structure and environmental corollaries of Panax
quinquefolium (Araliaceae) in Delaware — New York
Walter H. Lewis :
A 24-year comparison of the isan of a an iilneniieds innsiiiala es
E. H. Ketchledge and R. E. Leonard “ ,
A reappraisal of the orchid genera Broughtonia R. Br., Calitavenale es
and Laeliopsis Lindl.
Ruben P. Sauleda and Ralph M. Adams
A re-evaluation of Spiranthes <steigeri Correll
Paul M. Catling
Taxonomy of the Vicia neitniintn ousiebind ee a
J. Stuart Lassetter :
A revision of the genus Kyllinga Rottb. apecaiaiis in shesnin an
Central America
Gordon C. Tucker
NEW ENGLAND NOTE
The re-discovery of rant odora Ait. a in Vermont
Peter F. Zika ; ‘
Notice of Publication ?
NEBC Research Award Notice
Announcement of Joint Meeting
Connecticut River Ecosystem Conference
Instructions to Contributors
List of Reviewers
Index to Volume 86

397

417

421

425

Statement of Ownership ca a eee Sidhe bade cover