Berberidaceae
through
Fumariaceae
of
New York State
Richard S. Mitchell
New York State Museum
LIBRARY
MAR 11 1983
NcW YUr\K
botanical garden
Contributions to a Flora of New York State V
Richard S. Mitchell, Editor
1983
Bulletin No. 451
New York State Museum
The University of the State of New York
THE STATE EDUCATION DEPARTMENT
Albany, New York 12230
Digitized by the Internet Archive
in 2017 with funding from
IMLS LG-70-15-0138-15
https://archive.org/details/bulletinnewyorks4511newy
Berberidaceae
through
Fumariaceae
of
New York State
Richard S. Mitchell
New York State Museum
Contributions to a Flora of New York State V
Richard S. Mitchell, Editor
1983
Bulletin No. 451
New York State Museum
The State Education Department does not discriminate on the basis of age, color, creed, disability,
marital status, veteran status, national origin, race, or sex in the educational programs and activities
which it operates. This policy is in compliance with Title IX of the Education Amendments of 1972.
Inquiries concerning this policy may be referred to the Department’s Affirmative Action Officer, Edu-
cation Building, Albany, NY 12234.
The University of the State of New York
THE STATE EDUCATION DEPARTMENT
Albany, New York 12230
THE UNIVERSITY OF THE STATE OF NEW YORK
Regents of The University (with years when terms expire)
1988 Willard A. Genrich, Chancellor, LL.B., L.H.D., LL.D.,
1988 J. Edward Meyer, Vice Chancellor, B.A., LL.B.,
L. H.D. Chappaqua
1986 Kenneth B. Clark, A.B., M.S., Ph.D., LL.D., L.H.D.,
D.Sc. Hastings
on Hudson
1989 Emlyn I. Griffith, A.B., J.D. Rome
1983 Mary Alice Kendall, B.S. Rochester
1984 Jorge L. Batista, B.A., J.D., LL.D. Bronx
1986 Laura Bradley Chodos, B.A., M.A. Vischer Ferry
1987 Martin C. Barell, B.A., I.A., LL.B. Kings Point
1984 Louise P. Matteoni, B.A., M.A., Ph.D. Bayside
1987 R. Carlos CARBALLADA, B.S., L.H.D. Rochester
1988 Floyd S. Linton, A.B., M.A., M.P.A., D.C.L Miller Place
1988 Salvatore J. Sclafani, B.S., M.D Staten Island
1989 Mimi Lieber, B.A., M.A. New York
1985 Shirley C. Brown, B.A., M.A., Ph.D. Albany
1983 Robert M. Best, B.S. Binghamton
President of The University and Commissioner of Education
Gordon M. Ambach
Executive Deputy Commissioner of Education
Robert J. Maurer
Deputy Commissioner for Cultural Education
Carole F. Huxley
Director, State Science Service
Richard H. Monheimer
Chief Scientist, Biological Survey
Norton G. Miller
State Botanist
Richard S. Mitchell
PREFACE
OUR GOAL in producing this series is to present a useful and authoritative account of the plants
of New York State. These contributions are intended to reflect the knowledge and taxonomic
opinions of specialists who prepare the manuscripts while following a generalized format for consis-
tency. Inclusion of ecological, distributional, medical and economic information on each species is
also one of our major aims. Habitat references, flowering times, pertinent synonymy, etc., apply
specifically to New York plants rather than to the entire ranges. Complete illustration should
facilitate identification of specimens for those who are not formally trained in botany. Descriptions
are original, ordered and as complete as possible to provide sequential cross-referencing.
Distribution maps accompany species of seed plants, ferns, mosses, lichens and some groups of
fungi. These are plotted by counties to eliminate pinpointing endangered habitats, while offering
an accurate visual picture of past collecting. Maps are based on the master file at the New York
State Museum, Albany, and supplemented by available data (specimens examined by the authors)
from herbaria housing significant New York collections. Specific data or literature citations for any
map may be obtained, on approval, from the Museum.
We hope that these bulletins will serve individuals with interest in the flora, as well as to
provide information for state and federal agencies, conservation organizations, industry and the
scientific community. With these works go our hopes for the preservation and wise use of a pre-
cious and lifegiving resource — our state’s plantlife.
The New York State Flora Committee
The steering council of the New York State Flora Committee met for the first time on January
19, 1976, and established as its goals the promotion of study of the state’s plant resources and the
publication of this series of Museum Bulletins. These contributions will be continually updated
after publication for possible incorporation into larger volumes at a later date.
Members of the council at the time of this publication are:
Richard S. Mitchell, Chairman, State Botanist, N.Y. State Museum, Albany (Vascular Plants)
Charles J. Sheviak, Curator of Botany, N.Y. State Museum, Albany (Vascular Plants)
Edwin H. Ketchledge, College of Environmental Science and Forestry, Syracuse (Bryophytes)
Clark T. Rogerson, New York Botanical Garden, New York (Fungi)
George J. Schumacher, Biology Department, SUNY, Binghamton (Algae)
CONTENTS
Page
Preface iii
The New York State Flora Committee iii
Acknowledgments v
Important Note v
Legend vi
Berberidaceae 1
Podophyllum 1
Jeffersonia 3
Caulophyllum 5
Berberis 6
Lardizabalaceae 10
Akebia 10
Menispermaceae 12
Menispermum 12
Papaveraceae 15
Sanguinaria 16
Chelidonium 17
Glaucium 20
Macleaya 21
Eschscholzia 23
Papaver 24
Argemone 34
Fumariaceae 38
Dicentra 39
Adlumia 44
Corydalis 45
Fumaria . 51
Appendix I. (fungi) 53
Appendix II. (insects) 57
Bibliography 59
Index to Latin Names 65
IV
ACKNOWLEDGMENTS
The illustrations in this treatment are originals by Kathryn M. Conway. The appendices on
insects and fungi are the work of J. Kenneth Dean, with reviews by Timothy McCabe, Clark
Rogerson and John Haines, to whom I offer my thanks. Special thanks go to Gerald Ownbey and
the Steering Council of the Flora Committee for review. Arthur Cronquist also offered helpful
suggestions. Sincere thanks go to the staffs and curators of the following institutions who offered
their complete cooperation and hospitality when I visited their collections: the U.S. National Her-
barium (US), the New York Botanical Garden (NY), Cornell University (CU,BH), Environmental
Science & Forestry, Syracuse (SYRF), Harvard University (GH,NEBC), Brooklyn Botanical Gar-
den (BKL), the Buffalo Museum of Science (BUF) and Colgate University (GRCH). Data from the
above collections are added to the master file at the New York State Museum (NYS). This file was
started by Homer House around 1915 and was largely produced and maintained by Stanley J.
Smith from 1948-1978. The classification system employed in organizing this flora is that of Cron-
quist (1981).
IMPORTANT NOTE
All economic uses, folklore, medical and pharmaceutical notes,
uses as foodstuffs, etc., are compiled from the literature and do
not represent an endorsement by the authors or the New York
State Museum. Some of the uses may, indeed, be dangerous if
incorrectly employed. Some are not effective and are presented
for historical interest only.
LEGEND
FOR ALL MAPS IN THIS PUBLICATION THE
FOLLOWING SYMBOLS APPLY:
Solid dot — specimen seen by author; data on file at the State Herba-
rium (NYS)
Circle — Field observation with location data and observer’s name on
file (NYS)
Hollow triangle — Literature citation on file (NYS)
FOR ALL ILLUSTRATIONS IN THIS PUBLICATION THE
FOLLOWING LETTER-DESIGNATIONS APPLY:
A.
Habit sketch
F. Flower
B.
Silhouette
G. Inflorescence
C.
Fruit or fruit cluster
H. Flower parts (breakdown)
D.
Seed
K. Rhizome
E.
Bud
M. Root
VI
Berberidaceae (Barberry Family)
The Berberidaceae: a family of shrubs and primitive herbs comprising 10 (or more) genera and up to 600 species
worldwide. Most of its taxa are species of Berberis (Barberry) including the compound-leaved segregate genus,
Mahonia (Grape Holly). Distribution of the family centers in Asia, with lesser representation in Europe, north
Africa and the Western Hemisphere. The Pacific Coast of North America has as native genera: Vancouveria,
Achlys and Berberis (Mahonia). In eastern North America Diphylleia cymosa and Berberis canadensis are primarily
southem-Appalachian, and are known only from cultivation in New York State. Our native genera ( Podophyllum ;
Caulophyllum; Jeffersonia) are herbaceous relicts of the ancient Arctotertiary Forest, each with one species in
North America and an east-Asian counterpart. Two species of Berberis escape cultivation in the state. Members of
this family are important in horticulture, the pharmaceutical trade and as alternate hosts to diseases of cereal-grain
crops. Some are also poisonous.
FAMILY DESCRIPTION
Mostly shrubs or perennial herbs from perennial rootstocks, rhizomes or tubers. Leaves are alternate, fascicled or
sub-opposite, and may be simple or pinnate ly compound (or reduced to spines), sometimes basally articulated,
entire to variously toothed or lobed or with marginal spines, estipulate or stipuled, some genera with swollen
petiole bases. Flowers are solitary or borne in racemes, cymes or panicles, usually bracteate. Flowers are radially
symmetrical, bisexual; the perianth is of two to several series of free parts which may be somewhat similar or
clearly differentiated into calyx and corolla, 3 to 5-merous; petals (4) 6-9; sepals (4) 6 to many, often petaloid.
Stamens may be equal to the petals in number or more numerous (4-18); the outer whorl is often opposite the
petals and is sometimes associated with petaloid nectaries. Anther sacs dehisce by valves (slits in Podophyllum).
The ovary is superior, 1 per flower with (1) 2 to several ovules on a sub-basal to parietal placenta. Fruits are few-
seeded berries or capsules. Seeds are often drupe-like with fleshy to homy endosperm, sometimes with an aril.
KEY TO GENERA
1. Plants woody, much-branched shrubs, usually with sharp spines 4. Berberis (p. 6)
1. Plants herbaceous, without spines (2)
2. Flowers borne in panicles, greenish-yellow to purple, less than 1 cm wide; seeds borne naked, with the
appearance of glaucous, blue berries 3. Caulophyllum (p. 4)
2. Flowers solitary, white to creamy, over 1 cm wide; seeds borne within a berry or capsule (3)
3. Leaves only basal, long-petioled with 2 terminal leaflets; fruit a capsule, scapose 2. Jeffersonia (p. 3)
3. Leaves mostly cauline, often peltate with several lobes; fruit a berry, peduncled ... .1. Podophyllum (p. 1)
1. PODOPHYLLUM
Common Names: May-apple, Mandrake
Authority: Linnaeus, Species Ph, p. 505, 1753
This is a genus of primitive plants of uncertain relationships, sometimes put in their own family, Podophyllaceae. One
species is native to eastern North America, and one or more are known from Asia. Our common, forest species, P.
peltatum !.., is sometimes grown as an ornamental or transplanted to woodlots where it thrives. The seeds and
foliage are poisonous, but the pulpy flesh of the fully ripe fruit is reportedly edible.
1
1 Podophyllum peltatum L.
Common Names: May-apple, Wild Mandrake,
Wild Jalap, Indian-apple, Ground-lemon, Hog-
apple, Puck’s-foot
Type Description: Linnaeus, Species Pi., p. 505,
1753
Origin: Arctotertiary Forest
Habitats: Rich, moist woodlands and borders,
usually in partial shade
Habit: Erect, perennial herbs
Flowering: May
Fruiting: (July) August
General Distribution: Western Quebec and New
England to Minnesota, south to Texas, Loui-
siana and the southern Appalachians (n.
Florida)
Description: Plants with bisexual flowers; stigma (2) 4-8 lobed, capitate, convoluted, fleshy, 2-5 mm broad, 2-3
mm high, persistent in fruit; style lacking; ovary 1 per flower (rarely 2-8 carpels), superior, fusiform to subspheri-
cal, 8-12 mm long, 5-8 mm wide, many-ovuled, smooth, becoming a fleshy, ovoid berry in fruit, 3. 5-5. 8 cm long,
ca 2-4 cm wide at maturity, somewhat spindle-shaped with obtuse, tapered ends, pale yellow to deep yellow often
tinged with pink (uncommonly orangish to deep red), smooth, the pulpy placenta surrounding the numerous seeds;
seeds elliptic, 3-4 mm long, almost totally enclosed in their pulpy arils; stamens usually 12-20 or more (rarely 6,
opposite the petals), 8-10 mm long; anther sacs 4-6 mm long, dehiscing along longitudinal slits (not valved) with a
broad, fleshy connective; filaments stout, 3-6 mm long, broader near the bases; petals 6 or 9, free, in 2-3 series
(sometimes with smaller petals interspersed), greenish-white to creamy-pink (rarely orange to reddish), obovate to
obspatulate with rounded tips, mostly 1-3 cm long, 1-2.5 cm broad; sepals 6, in 2 series, greenish- white, early-
deciduous, cucullate, enclosing the 9-18 mm bud; sepal-like bractlets associated with the base of the bud early-
deciduous, 2-3 in number, greenish, oval, ca 3 mm long, 2 mm wide; receptacle tough, somewhat expanded in
fruit; peduncle 3-7 cm long, sparsely to densely villous and somewhat flattened and ribbed, arching outward from
the common axil of 2 terminal leaves to bear a single flower, elongating and pendulous in fruit; leaves 1 or 2 (3)
2
per stalk, oriented horizontally, suborbicular to reniform-cordate (or peltate), usually deeply (3) 5-9 lobed, palmate
with deep sinuses and ± bifid lobe tips, with a few shallower lobes and apiculate (to blunt) serrations and denta-
tions of varying sizes along the margins, glabrescent, and darker green above, soft-villous below; solitary leaves
robust, usually suborbicular and 5-9 palmately cleft, often peltate near the margin, 15-30 (40) cm broad, each on
a ribbed stalk 15-40 cm tall which arises directly from a (bracteate) rhizome; paired leaves (very rarely 0, 1 or 3)
of the fertile stalks 3-7 (9) palmately cleft with deep sinuses and marginally attached petioles (or slightly peltate);
petioles (4) 7-16 cm long, ribbed and deeply grooved, ± villous toward the blade, attachment subopposite (to
alternate) subtending the peduncle at the stem apex; stem glabrous, grooved and minutely caniculate, 15-30 (40)
cm tall; basal bracts (scale leaves) strap-like ± sheathing, 2-5 (7) cm long, ca 1 cm wide, pale, creamy-green,
borne alternately near the point of stem-attachment to the branching rhizome, 2-8 (11) mm thick (colony-forming)
with tough, fibrous roots. (2n = 12)
Infraspecific Variation: Developmental irregularities are not uncommon in this species. Populations, as well as
individual teratological specimens, have frequently been written up in the literature and occasionally named. Plants
developing 2-8 carpels (or separate fruits) per flower have been designated forma polycarpum Clute. Plants with
dark, reddish fruits and flowers have been called forma deamii Raym. Leafless flowering stalks are known (though
rare), called forma aphyllum Plitt. , and flowers may be accompanied by 1 or 3 leaves on occasion, rather than two.
Another developmental variation is the fusion of the peduncle to one of the petioles and the production of a floral
bract.
Importance: May-apple is cultivated in temperate climates in gardens and woodlots, and may escape within and
outside its natural range. It is a poisonous plant; however, the pulp of the fully ripe fruit is widely known to be
edible. They are not tasty, and are mostly eaten by children, who have reportedly been severely poisoned by
partially ripe fruit. The pulp may be made into a good marmalade or added to fruit drinks, but the seeds may
contain measurable amounts of toxins. The foliage and roots of the plant are particularly poisonous, containing
Podophyllotoxin and related resins and lignins. Podophyllin, the crude extract, is a violent cathartic and laxative,
and has been used in folk medicine in very small doses. It has also been used as a cytotoxic substance in the
treatment of certain types of cancer and venereal warts. The practice of treating tumors in this manner was learned
from American Indians. Internal poisoning results in severe gastroenteritis, purging and possible death. Even
handling the powdered rhizome may cause skin lesions, conjunctivitis and other symptoms.
2. JEFFERSONIA
Common Names: Twinleaf, Rheumatism-root
Authority: Barton, Trans. Amer. Phil. Soc., vol. 3, p. 342, 1793
A genus of two species: Jeffersonia diphylla of the eastern part of North America and J. dubia of Manchuria. Both
species are cultivated, and ours has been used in folk medicine.
3
1. Jeffersonia diphylla (L.) Pers.
Common Names: Twinleaf, Rheumatism-root,
Ground-squirrel Pea, Helmet-pod
Type Description: Linnaeus, Species PL, p. 505,
1753
Synonyms: Podophyllum diphyllum L., Jefferso-
nia binata Bart., J. bartonis Michx.
Origin: Aretotertiary Forest
Habitats: Rich, calcareous woodlands in deep
soils
Habit: Erect, acaulescent, perennial herbs
Flowering: Early May (NY)
Fruiting: June-August
General Distribution: Western N. Y. State and
adjacent Ontario to Minnesota, Iowa and south
to Tennessee and Georgia
Rarity: Near its northern limits in western New
York, this species is recorded from fewer than
25 locations in the state.
Description: Plants with bisexual flowers; stigma 1 per ovary, truncate at tip, 2-lobed, fleshy, convoluted, 1-1.5
mm broad, persistent; style stout, 1-2 mm long, persistent in fruit; ovary 1 per flower, superior, elliptic-oblong,
5- 7 mm long, 3-5 mm wide, stipitate, glabrous, many-ovuled, becoming a capsule in fruit; capsule 2. 2-2. 8 cm
long, 1.3-1. 8 (2.1) cm broad, bilaterally symmetrical, oriented ± horizontally by a contortion of its basal stipe,
ovoid-fusiform, somewhat falcate, the terminal portion attenuated abruptly toward the persistent style beak, the
surface greenish-tan, muricate-tuberculate, the capsule dehiscing by a large, ringent, subterminal adaxial pore with
revolute, lipped margins, the upper fruit then becoming a lid; seeds numerous, oblong, 5-7 mm long, 2-2.5 mm
wide, surfaces red-brown, muricate; aril pale, wing-like, ca 3 mm long; stamens usually 8; anther sacs linear, 5-6
mm long, dehiscing by 2 long valves which are apically attached; filaments ca 2 mm long; perianth parts in 2-3
series, all white (greenish or lavender tinted), free; petals 8 (9) in 2 series or acyclic, 6-18 mm long, 3-12 mm
wide, narrowly to broadly obovate; sepals usually 4 (3-5), petaloid, narrowly spatulate, attenuated toward bases,
6- 16 mm long, 2-5 mm wide; flower (fruit) borne singly; receptacle conspicuously jointed to the stipe, ebracteate
(rarely with a linear bract); scape slender, 7-55 cm tall, glabrous, ± ribbed; leaves all basal, glabrous, borne on
long petioles from the rhizomes; leaf blade deeply divided into 2 leaflets (rarely undivided, renifbrm); leaflets
borne opposite one another in the same plane forming near mirror-images (“angel-wings”), 1-11 cm long, 0.5-6. 5
cm wide, palmately veined from their lateral points of attachment, arcuate, half-oval, the upper lobe somewhat
rounded, the lower with a pointed tip, proximal leaflet-margins usually entire, distal margins shallowly sinuate-
dentate (or lobed); petioles slender, glabrous, ± caniculate, 7-60 cm tall, often apiculate-tipped (apex between
4
leaflets); basal bracts 3-8 cm long, 4-7 mm wide (much-reduced along rhizome), lanceolate, with pseudo-parallel
veins and ± fleshy (phyllodal); rhizome tough, branched (colony-forming), 5-12 mm broad with fibrous, matted
roots. (2n = 12)
Infraspecific Variation: Plants of Twin-leaf are quite variable in size, some populations being dwarfed with dwarfed
flowers. The leaflets may occasionally be joined into a single reniform blade (as in J. dubia of Asia). In forma
lobata Clute, the leaflets are deeply marginally lobed.
Importance: This species is a very attractive ornamental oddity, grown primarily for its foliage and unusual pod,
since flowering is brief. Extract of the root is aromatic and considered to be stimulant, used in tonics and some-
times taken for rheumatism. Also, this extract has been used in folk medicine as a substitute for Snakeroot ( Aristo -
lochia serpentaria).
3. CAULOPHYLLUM
Common Names: Blue Cohosh, Papoose-root
Authority: Michx., Fl. Bor. Amer., vol. 1, p. 205, 1803
A genus of two species widely disjunct in eastern North America and eastern Asia. They are sometimes cultivated,
but are also quite poisonous.
1. Caulophyllum thalictroides (L.) Michx.
Common Names: Blue Cohosh, Papoose-root,
Squawroot, Blueberry-root, “Blue Ginseng”
Type Description: Linnaeus, Species Pi., p. 312,
1753
Synonym: Leontice thalictroides L.
Origin: Arctotertiary Forest
Habitats: Cool, moist woodlands in partial to full
shade and rich soils
Habit: Erect, perennial herbs
Flowering: April — May
Seeding: June — August
General Distribution: New Brunswick & Nova
Scotia to eastern Manitoba, southward in the
Appalachians to Georgia
5
Description: Plants with bisexual flowers; stigma 1 per ovary, minute, its unilateral surface linear-oblong; style 1
per ovary, somewhat falcate, 1. 1-1.5 mm long, tapering and enlarged toward the ovary; ovary 1 per flower,
superior, ovoid and ± gibbous, glabrous, enlarging from 2 mm at fertilization to 3-4 mm, then ruptured by the
developing ovule(s); ovules 2, one often aborting, the other enlarging greatly on a columnar stalk, breaking through
the ovary summit to become a berry-like seed (or both seeds developing); seed spheroid, 6-11 mm in diameter,
the integuement blue-purple, glaucous, the surface becoming ± rugose with age; endosperm homy; funicular stalk
pedicel-like, clavate, 5-7 mm long, stout, greenish-glaucous with a glabrous surface, attached at the receptacle
where the ruptured pericarp often persists, looking much like a bract; stamens 6, free; anther sacs 1.5 mm long,
clavate, dehiscing by hooded, subterminal pores; filaments ca 1 mm long, ± flattened; petals (5) 6, 1.3-1. 8 mm
long, clawed, spatulate, with blunt, flabellate to reniform expanded tips, somewhat fleshy, dusky purple (to yellow-
green); sepals (5) 6, petaloid (each subtending a petal), obovate to narrowly elliptic with entire to erose margins
and rounded to acute tips, yellow-green with a purple blush (to deep purple or brownish); involucre (subtending
the corolla) of 3-4 sepaloid bracts, linear-ovate, ca 2 mm long; pedicels 5-11 mm long in flower, up to 22 mm long
in fruit, shallowly ribbed, often with tiny, acute-tipped bracteoles and bracts; inflorescence and flowers opening
early with the developing leaves, paniculate or racemose, usually of 5-12 flower; leaves 2 or 3, borne at 2 levels,
usually sessile and consisting of whorls of biternate, triternate or pinnatisect compound leaflets, or the lower 2
leaves subopposite; upper leaflet whorl usually biternate, subtending the inflorescence; lower whorl of several
long-stalked, compound segments; immature leaflets dark purple or blackish-brown (also stalks) in claw-like clus-
ters of linear, toothed segments; mature leaflets green, somewhat glaucous below, with obovate, cuneate or half-
oval blades, 2-6 (10) cm long, 2-8 (11) cm broad, each blade usually with 1-3 (4) broad lobes with sharp sinuses,
lobe tips obtuse to acute, leaflet bases obtuse to truncate, cordate or half-cordate, surfaces glabrous, glaucous; leaf
stalks up to 25 cm long on lower whorl, ribbed glabrous; stipules absent; stem glabrous, ribbed, up to 1.4 cm thick
and a meter tall; basal bracts often 2-6, phyllodal, imbricated, 1-9 cm long, ± clasping the stem; rhizome tough,
knotty, often branched, bearing the plant on an upturned caudex which bears bract scars of previous years; roots
tough and thick, entangled in a mat around the rhizome. (2n = 16)
Importance: Blue Cohosh is sometimes cultivated in shady gardens, but should be avoided where it might be a
hazard to children. The bitter, berry-like seeds are rich in the alkaloid Methylcytisine and undetermined glyco-
sides. Saponin from the rhizomes may cause severe gastroenteritis. It has also been shown to be an orally active
smooth muscle stimulant and oxytoxic. Caulophyllum has also been implicated in cases of dermatitis and mucous
membrane irritation. The roasted seeds are said to be safe as a coffee substitute. Extract of dried rhizomes has
been used in folk medicine (small quantities) as an emenagogue, diuretic and antispasmodic.
4. BERBERIS
Common Name: Barberry
Authority: Linnaeus, Species Pi., p. 330, 1753
Berberis is a genus of over 200 species, primarily distributed in Asia and Africa. The segregate genus, Mahonia,
differs in having compound (often spiny-margined) leaves and in lacking stem spines. New York State has two
introduced species of the genus \vdiich occasionally become naturalized after escaping cultivation. The nearest
native species is B. canadensis Mill, of the southern Appalachians. Barberries are relatively common in cultivation
except in grain-growing areas. Due to their role as alternate hosts to stem rust (Puccinia graminis) of wheat, B.
vulgaris and B. canadensis have been largely eradicated in some states. The wood, bark and berries of some
species have folk uses.
6
Description: Plants with bisexual flowers; stigma 1, orbiculate, depressed in the center; style usually absent (or
short); ovary 1, superior with 2-9 (15) erect ovules, becoming a several-seeded, indehiscent berry, usually oblong
and fleshy; seeds lacking arils, but with crustaceous integuements, containing elongate embryos and albuminous
endosperm; stamens 6, free; anther sacs opening by hinged valves; filaments tactile; petals 6, free, in 2 whorls,
each subtending a stamen, margins inrolled, 2 glandular structures at the base, just above the short petal claw;
sepals usually 6, petaloid, yellowish; floral bracts 2-3, small, subtending the calyx directly; pedicels slender;
inflorescence of few to many flowers borne in racemes or umbels (sometimes solitary) from short, bracteate shoots;
leaves simple (excluding Mahonia) ± spatulate, articulated near the base, borne in fascicles on short shoots in the
axils of branched or unbranched spines (rarely spineless); stems shrubby with arched branches and yellow inner
bark and wood, up to 3 (6) meters tall from fibrous root systems,
KEY TO SPECIES OF BERBERIS
1. Mature leaves with toothed margins; flowers and fruits borne in elongate racemes; some spines forked
1. Berberis vulgaris (p. 7)
1. Mature leaves with entire margins; flowers and fruits borne singly or in small umbels; spines unbranched
2. Berberis thunbergii (p. 9)
1. Berberis vulgaris L.
Common Names: Common Barberry, European
Barberry, Jaundice-berry, Piprage
Type Description: Linnaeus, Species PL, p. 330,
1753
Origin: Eastern Europe
Habitats: Thickets, roadsides, waste places, fen-
ce rows and woodland borders, as an escape
from cultivation
Habit: Profusely-branching, deciduous shrubs
Flowering: May — June
Fruiting: June, persisting year-round
General Distribution: Escaping in temperate cli-
mates; a native of Eurasia
7
Description: Plants with bisexual flowers; stigma 1 per ovary, discoid, ± concave in the center, ca 1.5 mm broad,
0.3 mm thick, the surface rugose, margins overlapping the ovary tip, persistent in fruit; style absent; ovary 1 per
flower, 1.9-2. 4 mm long, ca 1 mm wide, cylindric, several-ovuled, becoming an elliptic to subglobose berry in
fruit, 7-12 mm long, 5-9 mm wide, glossy scarlet to dark red; seeds often 2-6 per fruit, ca 4 mm long, ellipsoid-
clavate and somewhat curved with wrinkled surfaces; stamens 6; anther sacs opening by conspicuous valves; fila-
ments fleshy ± clavate, ca 2 mm long; petals 6, free, each subtending a stamen, creamy yellow, 3^ mm long, 2-3
mm wide, each with 2 elongate, abaxial glands ca 1 mm long at the base, (petals) cucullate, obovate with entire to
minutely erose margins; sepals 6, free, petaloid, cream yellow to greenish, ovoid-cucullate, 4-5 mm in diameter;
bracts 2-4 (6), yellow-green, 1.5-2. 5 mm long, 0.5-1. 5 mm wide, obtuse to acute tipped with entire margins;
pedicels 5-8 mm long ± caniculate, each subtended by a minute, lanceolate bract; inflorescences dense, axillary
cymes 3-6 cm long, usually of 12-15 flowers; peduncle of the inflorescence 1-1.5 cm long borne on short-shoots,
smooth or very obscurely ribbed, flattened, occasionally with 1 or 2 naked bracts of the type that subtend pedicels;
leaves borne in fascicles, tightly spiraling (on short-shoots), 2.5-6 (8) cm long, 0.8-2. 8 cm wide, obovate, with
rounded to obtuse tips and attenuate bases, the margins profusely serrulate-toothed, each tooth tipped with a
flexuous spine ca 1 mm long, the revolute margin of the leaf contiguous with the spines, leaf surfaces green to red-
purple, glabrous, the lower with a prominent network of reticulate veins; petioles glabrous, ± winged toward the
blade, up to 1.5 cm long, conspicuously jointed near the base, the 2-3 mm below the joint expanded and
sheathing, persistent on the short-shoot; short-shoots bearing both leaves and inflorescences, varying in length with
age, usually less than 1 cm long, 4-5 mm wide, sheathed in the tightly spiraling petiole bases of dropped leaves;
spines present at the base of short-shoots (or absent) greenish to shiny red-brown at maturity, each with a tough
expanded base and 2-3 spine branches at the tip (or single), spine branches mostly 4-10 mm long; twigs and stems
furrowed with grayish-brown to chalky bark which peels away in thin, coriaceous strips; plants branching freely,
forming erect to bushy-sprawling shrubs up to 2 (3) meters tall, from dense root systems. (2n = 28)
Hybridization: Although it has not been reported from New York State, Berberis x ottawensis Schneid. is thought
to be the natural hybrid of B. vulgaris and B. thunbergii DC., and might be expected where the two species occur
together.
Importance: Common Barberry is best known as an enemy of the wheat farmer, since it may serve as alternate
host of the devastating rust, Puccinia graminis. Eradication programs have been implemented in agricultural areas
of the midwestern United States and Canada with considerable success. In New York State the plants may be
found in cultivation and naturalized, but they are less commonly grown than the Japanese Barberry (B. thunber-
gii). The berries are suitable for making jelly, wine and pies. The yellow wood has been used in furniture inlay and
extracted for a yellow die for wool and leather. The wood and bark are also sources of Berberine, a broad-
spectrum antibiotic. In folk medicine, a tonic from the bark or roots is said to be alterative, but could be
dangerous in quantity.
8
2. Berberis thunbergii DC.
Common Name: Japanese Barberry
Type Description: De Candolle, Syst. vol. 3, p.
9, 1821
Synonym: B. vulgare of Kew Index not L. (in
part)
Origin: Eastern Asia
Habitats: Fencerows, thickets, old fields and
waste places as an escape
Habit: Much-branched, deciduous shrubs
Flowering: Late April-May
Fruiting: June-October (may persist year-
around)
General Distribution: Escaping circumboreally
in temperate climates; a native of Japan
Description: Plants with bisexual flowers; stigma 1 per ovary, discoid, ± concave in the center, ca 1.5 mm broad,
0.4 mm thick, the surface rugose, margins overlapping the ovary tip, persistent in fruit; style absent: ovary 1 per
flower, 1.5-2 mm long, ca 1 mm wide, broadly cylindric with 2-several ovules, becoming a tough, ellipsoid to
subglobose, red berry in fruit, 7-10 mm long, 6-8 mm wide, glossy, smooth; seeds 1 or 2 (-4), elongate-elliptic,
5-6 mm long, 2.8-3 mm wide, often with a flattened side, surface rugose to papillose; stamens 6; aether sacs
opening by valves; filaments thick, ca 1.5 mm long; petals 6, free, each subtending a stamen, creamy-yellow ca 3
mm long and broad, ovate-cucullate with entire margins, with basal glands ca 0.7 mm long; sepals 6, free,
petaloid, ovate-cucullate yellowish with a blush of red, 3-4 mm long and broad; bracts subtending the calyx 2-4,
strap-like, reddish-tinged, ca 2 mm long; pedicels glabrous, finely ribbed, 4-10 mm long, each with a minute,
subulate bract at base; flowers borne on short-shoots, singly, paired or in umbellate inflorescences of 3 (-5)
flowers; peduncle of inflorescence slender, ribbed, glabrous, 3-8 mm long (or absent on single flowers); leaves
borne is fascicles, tightly spiraling on the short-shoots, 0.5-2. 5 (3.5) cm long, 2-18 mm wide, entire, spatulate to
obovate with rounded to obtuse tips, margins ± revolute, surfaces glabrous, green to red-purple, blade gradually
attenuated to an indistinct petiole, except in the basal 2 mm where it is unwinged and obscurely jointed; short-
shoots bearing both leaves and inflorescences, 2-5 mm long and broad, sheathed in the spiraling, expanded petiole
bases of leaves of former seasons, reddish to gray-brown; spines present at the bases of short-shoots or with their
buds in the axils, 5-8 (11) mm long, with tough, expanded bases, the spine usually unbranched, with small.
9
abortive lateral spines at the junction with its sheathing base; twigs and stems deeply furrowed, greenish when
young to red-brown, the bark stripping, dark brown to nearly black; wood yellow; plants profusely divaricately
branching, up to 2.5 meters tall, from a dense root system. (2n = 18)
Infraspecific Variation and Hybridization: This species has a number of cultivated forms. Among them are color
and growth-form mutants including the diminutive cv. Minor (Box Barberry) and the narrow, upright cv. Erecta
(True-hedge Column-berry). The hybrid of B. thunbergii and B. vulgaris (B.x ottawensis) has not been reported
from New York State.
Importance. This species is widely cultivated and can occasionally become a noxious weed of abandoned fields and
fence rows.
Lardizabalaceae (Lardizabala Family)
A family of about 6 genera and 30 species of vines (rarely shrubs) native to eastern Asia and western South
America. In the United States those cultivated are: Akebia, Lardizabala, Decaisnia, and Stauntonia. By far the
most commonly cultivated species is Akebia quinata, which is relatively hardy and may escape as far north as
eastern Canada. Fruits of members of this family are often edible.
FAMILY DESCRIPTION
The plants are usually monoecious or dioecious woody vines with alternate leaves which are palmately compound
( Decaisnia , the exception, is a shrub with bisexual flowers and pinnately compound leaves). The perianth may
consist of one or two whorls and may differ in size or coloration between male and female flowers. Perianth parts
are free, in whorls of threes, including such reduced series as petaloid nectaries. When petals are absent or
reduced, sepals are often petaloid; stamens are 6, free or basally united. There are 3-15 free ovaries, each with (1)
2 to many ovules, parietal placentation and a single sub-sessile stigma. The fruits are berries or fleshy follicles
which dehisce along a single ventral suture. The embryo is small, straight and embedded in fleshy endosperm.
1. AKEBIA
Common Names; Five-leaf, Three-leaf Akebia, Chocolate Vine
Authority: Decne., Arch. Mus. Hist. Nat., Paris, vol. 1, p. 195, 1839.
A genus of two species of vines, native to China, Japan and Korea. In addition to the following species, the Three-
leaf Akebia, A. trifoliata (Thunb.) Koidz., is cultivated, but not known to escape in New York State.
10
1. A kebia quinata (Houtt. ex Thunb.) Decne.
Common Names: Five-leaf Akebia, Five-leaf,
Chocolate Vine
Type Description: Houtt., in Thunb. Fl. Jap., p.
148, 1784
Synonyms: Akebia lobata (Houtt. ex Thunb.)
Decne., Rajania quinata Houtt. ex Thunb.
Origin: Eastern Asia
Habitats: Thickets, vacant lot borders, waste
places and clearings as an escape from cultiva-
tion
Habit: High-climbing, perennial, semi-evergreen
vines
Flowering: May-June
Fruiting: July-September
General Distribution: A sporadic escape from
cultivation in North America; native to eastern
Asia
Description: Plants monoecious; female flowers: stigma 1 per ovary, cleft and bilabiate, 1.0-1. 5 mm broad, glossy
purple-brown; style absent; ovaries usually 4-6 per flower, free, superior, cylindric, 3-5 mm long, 1-2 mm wide,
often arched and divergent, 1-loculed, 1-carpelled, 1 to 2 ovaries per flower maturing into large, fleshy, berry-like
follicles, (follicle) up to 8 cm long and 4 cm wide, compressed dorsiventrally, oblong, purse-like, dehiscing along an
adaxial suture, purple or creamy, blotched with purple, glaucous with a juicy, yellow-green fleshy endocarp,
bearing a cylindric, parietal placenta with numerous seeds; seeds purplish-black, 4-5 mm long, ca 3 mm wide,
trapezoidal to narrowly tapered toward their blunt tips; rudimentary anthers 0-6, minute, less than 1 mm long;
petals absent; perianth (female flowers) of 3 petaloid, free sepals, dusky purple, strongly cucullate, often with
slightly reflexed margins, 7-12 (14) mm long, 5-9 (12) mm broad; receptacle becoming enlarged, woody, jointed
to the fruit; male flowers: ovaries absent or represented by sterile tissue; stamens 6, free at base, 2. 5-3.5 mm
long; anther sacs comprising most of the length of the stamens, falcate, inwardly arched to form a turbinate
androecium, purplish, dehiscing first by pores, then extrorsely valved; filaments minute, flattened; petals absent;
perianth (male flowers) of 3 petaloid sepals, free, rose-purple to dusky-purple, flat to weakly cucullate, ovate to
triangular-obtuse, 3-6 mm long, 2-5 mm wide, receptacle not enlarging; inflorescence a cyme-like, few-flowered
panicle, usually with 3-8 male flowers borne terminally and 2-4 female flowers attached below; pedicels slender,
glabrous, ribbed, yellow-green, 1-5 long in male flowers, up to 22 mm long in female flowers, placing them in
11
proximity with the males; pedicels subtended by linear, pellucid bracts, ca 1 mm long; peduncles mostly 1-2 cm
long, borne singly or several per node, subtended by broad coriaceous bracts at the nodes; leaves compound, 5-
palmately divided; very young leaflets linear-lanceolate, strongly apiculate; mature leaflets broadly oval to oblan-
ceolate, emarginate or rounded, often with a tiny, reflexed apiculation persisting at the tip, obtuse to acuminate at
base, 1.5-5. 5 (7) cm long, 0.6-3. 2 (4) cm broad, glabrous; petiolules glabrous, 1-10 mm long, somewhat ribbed or
winged; petioles slender, glabrous, ribbed, up to 15 cm long, slightly expanded at bases; stipules absent; bud
scales broad, oval to triangular-acute, coriaceous; nodes enlarged, woody; internodes smooth or ribbed when
young, becoming dotted with oval, protuberant lenticels, eventually forming bark which is yellowish to red-brown,
splitting longitudinally, becoming charcoal to grayish and fibrous on older stems; stems twining and climbing up to
15 meters or more from a fibrous, perennial rootstock. (2n = 32)
Importance: This vine is attractive and easily propagated by stem or root cuttings; seeds usually are set only after
hand pollination. It is widely cultivated, but may spread aggressively and escape. The gelatinous pulp of the fruit is
eaten in countries where the plant is native. The leaves are also extracted to make a tea, and young shoots are
woven into baskets in Japan.
Menispermaceae (Moonseed Family)
This large group is primarily tropical and absent from Europe. It comprises up to 80 genera and 400 species,
depending upon the authority consulted, but probably has far fewer defensable taxa. The plants are usually dioe-
cious vines. This family has historically been included under Magnoliaceae, Berberidaceae and Annonaceae. In
North America, north of Mexico, the representatives are: Cocculus (2 spp.) in southern states, Cissampelos (1 sp.)
from Florida, Calycocarpum (1 sp.) from the south and midwest, and Menispermum (1 sp.) from the eastern U.S.
and adjacent Canada. Over 100 alkaloids have been extracted from members of the family, about % of which are
not found outside Menispermaceae. A number of species, including Menispermum canadense L. are known to be
poisonous. The root extract of a tropical genus, Chondrodendron, yields curare — used locally as a coating to poison
darts and arrowheads. In medicine curare is used in controlled doses to paralyze skeletal muscles temporarily
during surgery.
FAMILY DESCRIPTION
Members of this family are primarily dioecious, woody vines (rarely shrubs or trees). Leaves are alternate, simple
(or trifoliate), entire to deeply palmately cleft and estipulate. The unisexual flowers are small and greenish with
both calyx and corolla usually present and 6-parted. Stamens are 3 to many, often 6, free or connate at base (some-
times monodelphous). Ovaries are unfused, superior, 3-6 per flower, each a one-loculed carpel; placentation is
parietal. The stigma is capitate or discoid, often scalloped or lobed. The fruit is a drupe or achene. Seeds contain a
curved embryo, with or without endosperm.
1. MENISPERMUM
Common Names: Moonseed, Yellow Parilla
Authority: Linnaeus, Species Pi., p. 340, 1753
Menispermum is a genus with one species in eastern North America (M. canadense) and one in eastern Asia (M.
dahuricum DC.), a classic pattern of Arctotertiary disjunction. These may constitute a single species, but require
further study. Moonseed is cultivated and may be dangerous to children, since the grape-like fruits are poisonous.
12
1. Menispermum canadense L.
Common Names: Moonseed, Yellow Parilla,
“Texas Sarsaparilla”
Type Description: Linnaeus, Species Pi., p. 340,
1753
Synonyms: Menispermum angulatum Moench,
M. virginicum Willd., M. mexicanum Rose
Origin: Arctotertiary Forest
Habitats: Woods and thickets, especially moist
bottomlands and streambanks
Habit: Perennial vines
Flowering: June — July
Fruiting: July — September
General Distribution: Western Quebec and New
England to Georgia and west to Oklahoma
Description: Plants dioecious; female flowers: stigma 1 per ovary, ca 0.2 mm wide, rotate with minutely scalloped
margins; style short, reflexed outward; ovaries 2 (4), reniform, ca 1 mm long, the abaxial surface convex, each
ovary becoming a fleshy, glaucous drupe, 1. 1-1.4 cm in diameter, subglobose, but developing unevenly to leave
the stigma displaced laterally as a small protuberance 2-4 mm from the fruit base, fruit color dark purple to black;
ovules 2, one aborting, the other becoming a crescent-shaped seed, 7-9 mm in diameter, laterally compressed, ca
3 mm wide with 3 prominent ridges along the outer perimeter, their walls with spiny ribs and furrows; embryo
curved within an albuminous endosperm; staminodes usually 6, 1.0-1. 5 mm long, each composed of a thick fila-
ment and bulbous tip; petals 4-8, free, ovate, greenish-white, 1.5-1. 8 mm long, ca 0.8 mm wide; sepals 4-8, free,
ovate, greenish-white, 1.7-2. 1 mm long, ca 1 mm wide; male flowers: ovaries absent; stamens 12-24, from 2.0-
2.5 mm long; anther sacs globose; filaments fleshy, free at base; petals 2-6, greenish, lanceolate, ca 1.5 mm long,
0-4 mm wide; sepals greenish, 4-8, ovate, ca 2 mm long, 1 mm wide; inflorescences 3-9 cm long, axillary panicles;
pedicels 1-3 mm long (up to 1 cm in fruit), ribbed, glabrous or with a few pilose hairs; foliar bracts subtending
pedicels and peduncles, linear- lanceolate, 1-2 (4) mm long with ciliate margins; peduncles and rachises shallowly
ribbed, often sparsely villous; leaves oval with cuspidate tips when young, becoming shallowly (to deeply) 3-7 (9)
lobed, the lobes rounded to acute-tipped at maturity, often with a residual apiculation, leaf bases truncate to
13
cordate (peltate), upper surfaces bright green, very sparsely short-villous, lower surfaces pale to gray-green,
sparsely to moderately villous, mature leaves up to 20 (30) cm broad, obscurely peltate, the petioles attached near
the basal leaf margins, elongating with age becoming up to 20 (25) cm in length, ribbed and sparsely villous, often
twisted; stems twining, reddish-brown, ribbed, sparsely to densely arachnoid- villous, especially near the growing
tips, climbing up to 6 meters from a tough, fibrous root system. (2n = 52)
Importance: Like other members of its family, this species is occasionally cultivated, primarily for its foliage and
climbing habit, and it may escape. It is a poisonous plant, the toxic principles being isoquinoline alkaloids (includ-
ing Dauricine) with an action much like curare. There are reported cases of convulsion, paralysis and death in
children after they ingested the grape-like fruits. The fruits are, however, not apparently injurious to birds for
which they serve as food. It is a common phenomenon for birds to thrive on fruits or seeds which are poisonous to
mammals, and they should never be used as indicators of the edibility of plant materials for humans.
14
Papaveraceae (Poppy Family)
The Papaveraceae: a family of some 25 genera and over 200 species worldwide. The plants are mostly herbaceous,
often with showy flowers, and a number of the genera are important horticulturally. The species treated here are
mostly naturalized after escape from cultivation, since Bloodroot (Sanguinaria canadensis L.) is the only member of
the family native to the state. The largest genus of the family is that of true poppies, Papaver, with 60-80 species.
In addition to their usefulness as ornamentals, poppies and their relatives are valuable sources of alkaloids. The
best known of these is Opium Poppy (Papaver somniferum L.), whose seeds also have commercial uses. Members
of Papaveraceae are most common and prolific in southern, dry climates, but range to boreal and arctic habitats.
Mediterranean climatic zones of southern Europe and the west coast of the U. S. and Mexico are particularly rich
in members of the poppy family.
FAMILY DESCRIPTION
Herbs or rarely shrubs or trees with milky sap (to watery), often with latex and characteristically containing iso-
quinoline alkaloids. Plants annual, biennial or perennial with unbranched or branched stems or scapose from
rhizomes or caudices. Leaves are usually alternate, but may be paired or whorled near the branch tips; they are
often divided or lobed and cut, but may be entire. Plants are glabrous to densely pubescent, and may have setae
or spiny armature. Flowers are bisexual, usually showy and brightly colored, solitary or in paniculate or umbellate
clusters. The calyx is of 2-4 distinct sepals which enclose the bud and usually drop with the opening of the flower.
The corolla is of 4-6 or 8-12 petals (rarely absent) which are distinct or very slightly fused at base. Stamens are 4
to many, centripetal, usually numerous with 2-celled anthers. Filaments are slender, clavate or alate-petaloid.
Ovary 1, superior, of 2 to many carpels, but unilocular (some genera have septations which break down in fruit
development). Styles are very short, usually obsolete; stigmas are (1) 2-18 or more, corresponding with carpel
number. They may be fused into a persistent radiate disc which caps the fruit. The fruits are various types of
capsules (or rarely follicles), dehiscent by valves, sutures or indehiscent. Placentation is parietal, and each capsule
usually bears many seeds. Seeds are quite varied in color, shape, reticulations, caruncles, aril types, etc. They
contain small embryos and albumenous, oily or mealy endosperm.
KEY TO GENERA
1.
3.
3.
5.
5.
Perianth of 2 creamy, petaloid sepals; petals absent; flowers in a much-branched, terminal panicle
4. Macleaya (p. 21)
Perianth of 2 or more whorls of unlike parts; petals present; flowers borne singly or in small, umbellate
clusters (2)
2. Plants acaulescent, the leaves and flowering scapes borne directly from a rhizome; leaves palmately veined
1. Sanguinaria (p. 16)
2. Plants caulescent, the leaves and flowers borne on above-ground stems; leaves pinnately veined . . . . .(3)
Leaves multifid with linear segments . 5. Eschscholzia (p. 23)
Leaves not multifid, but lobed, pinnatisect or entire (4)
4. Capsules more than 10 times longer than broad (6)
4. Capsules no more than 5 times longer than broad (5)
Leaves spiny; capsule opening by 4-6 valves 7. Argemone (p. 34)
Leaves not spiny; capsule opening by pores or indehiscent 6. Papaver (p. 24)
6. Petals 1 cm long or less; flowers in umbellate clusters . .2. Chelidonium (p. 17)
6. Petals 2 cm long or more; flowers solitary 3. Glaucium (p. 20)
15
1. SANGUINARIA
Common Names: Bloodroot, Puccoon.
Authority: Linnaeus, Species Pi., p. 505, 1753
A North American genus with a single species, but very closely related to the eastern Asian genus Eomecon, which
should possibly be included in Sanguinaria. The single species of Eomecon differs from S. canadensis in having
the sepals fused. It is also pubescent with cordate leaves and more than one flower per scape; however, Sanguina-
ria also will occasionally have two or more flowers in a bracteate inflorescence, suggesting common ancestry.
1. Sanguinaria canadensis L.
Common Names: Bloodroot, Puccoon, Puccoon-
root. Red Puccoon, White Puccoon, Tet-
terwort, Indian-paint, Red-root, Corn-root,
Pauson, Sweet-slumber, Snake-bite, “Tur-
meric”
Type Description: Linnaeus, Species Pi., p. 505,
1753
Synonyms: Sanguinaria acaulis Moench., S.
vernalis Salisb., S. dilleniana Greene
Origin: Arctotertiary Forest
Habitats: Rich woods, often alluvial soils in
calcareous places
Habit: Acaulescent, scapose, perennial herbs
Flowering: April-May (NY)
Fruiting: May -June (July)
General Distribution: Nova Scotia to Manitoba,
west to Kansas and south to e. Texas and
northern Florida
Description: Plants with bisexual flowers; stigma strongly 2-lobed, 1-2 mm in diameter, persistent; style narrowly-
cylindric, contiguous with the ovary, up to 1.5 mm long in fruit; ovary fusiform, 4-7 mm long, 2-3 mm wide,
glabrous, becoming an elliptic-fusiform capsule which is brown, glabrous, glaucous, 2. 5^. 5 (6) cm long (excluding
style) 0.8-1. 5 cm wide, 2-valved, acropetally dehiscent along 2 sutures, the valves not dropping, unilocular with 2
placentae, capsule base often somewhat stipitate at maturity; seeds numerous, obovate, 3.2-3. 7 mm long, 2. 2-2. 8
mm wide (excluding aril) with a basal nipple, greenish-tan when immature, becoming rich rust-red to dark red-
brown at maturity, the seedcoat minutely and irregularly pitted-reticulate; aril creamy-golden, crested and wing-
like, often longer than the seed (up to 4.2 mm); stamens 14-30, varying in length; anther sacs oblong, 2-4 mm
long, golden; filaments 3-11 mm long, slender; petals free, usually 8 (6-12), in 2 or more series, narrowly oblong
16
to elliptic or spatulate, entire (rarely lobed) 1-2 (3) cm long, 0.4-1. 2 (1.6) cm wide, white to creamy (pink); sepals
2, membranous, glabrous, cucullate, often notched irregularly at apex, 1.0-1. 5 cm long, early deciduous; flower
borne singly, scapose (very rarely 2-3 pedicillate flowers with bracts); scape naked, slightly ribbed, glabrous, 10-20
(35) cm tall; leaves 1 per node, arising from near ground level, each associated with a stipe, enfolding the bud
when young, growing throughout the season and changing shape from cordate-ovate to broadly reniform at matu-
rity, up to 20 cm long, 30 cm wide with large, irregular dentations and crenations as well as incisions with rounded
sinuses which make the blade palmately lobed (also palmately veined), blade tip rounded, basal lobes cordate to
sagittate, leaf surfaces glabrous, the lower surface pale green to conspicuously glaucous (especially when young);
mature petioles up to 35 (45) cm tall, ribbed, glabrescent, subtended by 2-6 ovate to lanceolate bracts, often with
falcate, e margin ate or obtuse tips, partially sheathing, membranous, 0.5-10.5 cm long, 5-12 mm wide, with
pseudo-parallel veins; stems consisting only of tough, branching rhizomes 5-20 mm in diameter with fibrous roots
and orange-red sap. (2n = 18)
Infraspecific Variation: Plants of this species vary in the degree of leaf lobing and in their size, color and glaucous-
ness, as well as the length and branching of the rhizome; fruiting stipes may overtop the leaves or not. Variety
rotundifolia (Greene) Fedde combines some variations of these features with large, undulate, practically uncut
leaves, and probably deserves recognition as a clinal, southern race at best. Flower color varies from white to pink,
and the cultivar Multiplex, has numerous petals replacing the stamens and ovary. These variations have been
found on numerous occasions, and their discovery in the wild usually generates some colorful horticultural litera-
ture. Multiplex has also been called var. plena Barcl. as well as cv, “Flore Pleno ’ in the past.
Importance: Sanguinaria is potentially poisonous, but not well documented as a threat to man or livestock. The
plants contain isoquinoline alkaloids, of which Sanguinarine, named for the genus, has been investigated as a
poisoning agent from Argemone. The generic name comes from the blood-like appearance of the juice, which was
used by American Indians as a facial stain. Medicinal use has been quite varied: early North American settlers used
a tea of Bloodroot to gargle for sore throat, while a similar solution was used by the Native Americans of the
Mississippi Valley as an internal treatment of rheumatism. In quantity or undiluted solution the Sanguinarine is
emetic, causing vomiting and diarrhea and possible shock, fainting and coma. The extract of powdered rhizome
became popular in medicine during the early 19th century, especially in Europe, and it is still a common medicine
in Russia. British doctors developed a flour paste with Sanguinaria extract and zinc chloride which was used to
treat various warts, polyps and superficial cancers of the nose and skin. It was applied after erosion of the tumor
with acid. This practice fell into disrepute until more recent studies showed Sanguinarine and Chelerythrine from
the plants to be active against types of carcinoma and sarcoma in mice. A revival and refinement of 19th century
techniques is now used with considerable success against certain carcinomas in humans and pets. Protopine, known
from this and other genera of the family, shows bradycardial action. These plants are popular in cultivation, being
easily propagated by rhizome cuttings. They are prized in Europe where they are not native.
2. CHELIDONIUM
Common Names: Celandine, Swallow -wort
Authority: Linnaeus, Species Pi,, p. 505, 1753
A genus with a single species, though species from other genera have historically been included in it. Chelidonium
majus L. is a common weed in northeastern North America and in Eurasia including areas of Europe where it is
also native. The acrid, yellow-orange juice is used medicinally and may be poisonous when undiluted.
17
1. Chelidonium majus L.
Common Names: Greater Celandine, Celandine,
Swallow-wort, Rock-poppy Tetter-wort, Kill-
wort, Wart-wort
Type Description: Linnaeus, Species Pi., p. 505,
1753
Origin: Europe
Habitats: Roadsides, yards and walls, moist
waste places and disturbed woodlands as an es-
cape from cultivation; now firmly naturalized
Habit: Erect and spreading biennial or perennial
herbs
Flowering: April-August
Fruiting: May -October
General Distribution: Naturalized over most of
the eastern United States and Canada; native
and ubiquitous in Eurasia
Description: Plants with bisexual flowers; stigma 1, with 2 rough faces outwardly directed ca 45° from vertical (roof-
like), persistent in fruit, not enlarging; style 0.8-1. 1 mm long, cylindric, glabrous, persistent; ovary 1 per flower,
superior, linear-cylindric, 4-7 mm long, ca 0.5 mm wide, becoming a linear capsule, glabrous, 2-4 (5) cm long, 2-
3 (4) mm wide, straight or slightly curved, brown to tan at maturity, dehiscing from the base upward along sutures
associated with two slender ribs, the valves eventually curling and falling away; seeds borne in two rows, few to
numerous, 1.2-1. 7 mm long, 0. 7-1.0 mm wide (excluding the aril) ellipsoid, slightly flattened on one side, reticu-
late, shiny, red-brown, the aril plump, crested, wing-like, 0.5-1. 8 mm long, creamy-golden; stamens 4-15 or
more, yellow, 4-6 mm long; anther sacs elliptic, ca 1.5 mm long; filaments slender but expanded upward; petals 4
in two pairs, 7-11 (17) mm long, 5-9 (14) mm wide, elliptic to narrowly ovoid, bright yellow (creamy or orange-
yellow); sepals 2, enclosing the bud) early-deciduous, yellow-green ca 8 mm long, glabrous or with a very few
villous trichomes; inflorescences near the branch tips, stalked umbels, usually 3-6 flowered; pedicels slender, often
glabrous and somewhat ribbed, elongating up to 3 cm in fruit; bracts at the pedicel bases minute, glabrous or
ciliate; peduncle of the umbel 3-5 (7) cm long, glabrous to sparsely villous, strongly grooved; leaves glaucous,
sparsely villous to almost glabrous, deeply 5-7 (or more) pinnatifid with variously lobed and dentate (rarely lacini-
ate) margins on the broad, rounded lobes; cauline leaves reduced upward, the smaller ones subtending the pedun-
cles sometimes scarcely lobed; basal leaves formed each year in a rosette; peduncles sparsely villous, winged and
grooved; stems winged and grooved, sparsely to densely villous, up to 1.3 meters tall from a tough, biennial or
perennial taproot and fibrous roots with orange-yellow latex and sap. (2n = 12)
18
Infraspecific Variation; A large-flowered form (up to 3.5 cm in diameter) is known for this species. The variety
laciniatum (Mill.) Syme has deeply incised leaves.
Importance: This plant was introduced into North America by European settlers who used it as a medicinal herb.
The yellow-orange sap has been extracted since ancient times and used in dilute solutions for various internal and
external ailments. The teachings of Pliny and Dioscorides refer to it in treatment of “film of the eyes”. The
technique was supposedly learned from swallows who were said to rub Celandine in the eyes of their young.
Mixed with milk or made into a salve, it is still used by some nature healers and reputed to aid with problems
ranging from cataracts to impotence. The juice has been used to erode corns and warts, hence the strange common
name of Wart-wort. This juice is rich in alkaloids such as Berberine, Protopine, Chelidonine and Sanguinarine, etc.
and plants are included on poisonous plant lists. Deaths from ingestion of the bitter foliage have been reported
from Europe but not North America. Professor M. L. Fernald once reported eating the very young greens in salad
without ill effects, but it should be avoided as a foodstuff. Spartesine, found in Chelidoniurn, has been used to
restore heart arhythmia and in cases of atrial fibrillation. It is also a diuretic, and shows some experimental hy-
poglycemic activity. The juice and powder have been used for toothache and by dentists during and after tooth
removal.
19
3. GLAUCIUM
Common Names: Horned Poppy, Sea Poppy
Authority: Miller, Gard. Diet. abr. ed., p. 4, 1754
This is a genus of about 20 species native to southern Europe and central Asia. The following species occurs in
New York State as an escape from cultivation.
1. Glaucium flavutn Crantz
Common Names: Horned Poppy, Horn-poppy
Type Description: Linnaeus, Species Pi., p. 506,
1753
Synonyms: Chelidonium glaucium L., Glaucium
glaucium (L.) Karst., Mosenthinia glaucium
(L.) Kuntze, Glaucium luteum Scop.
Origin: Mediterranean Region
Habitats: Sandy to gravelly places, seashores,
waste places and borrow pits as an escape
Habit: Erect or spreading biennials or perennials
Flowering: June-August
Fruiting: July -October
General Distribution: Escaping in the northeast-
ern U. S. and eastern Canada; native to
southern Europe and northern Africa
Description: Plants with bisexual flowers; stigma 1 per flower, ca 2 mm tall, 3 mm wide, with two laterally
diverging or slightly reflexed lobes, the other 2 opposite lobes folded upward upon one another giving the stigma a
hat-like appearance, receptive surfaces glandular to minutely tuffed-puberulent, (stigma) persistent in fruit becom-
ing 4-5 mm wide, ca 3 mm high; style absent; ovary 1 per flower, superior, linear-cylindric, 1.0-1. 5 cm long, 1-2
mm wide, muricate, especially above, becoming an extremely elongate, tuberculate, pod-like capsule, greenish-tan
to brown, 15-40 (50) cm long at maturity, 4-7 (9) mm wide, conspicuously jointed to the receptacle, arching
upward or outward and dehiscing along 2 sutures from the tip to near the base, 2-chambered with corky tissue
surrounding the numerous seeds; seeds 1. 5-2.0 mm long, ca 1 mm wide, dark red-brown to chocolate with a
moderately prominent reticulum of oblong fields; stamens numerous, 5-14 mm long; anther sacs linear-oblong, ca
2 mm long, 0.4 mm wide, yellow; filaments yellow, slender but flattened, 3-12 mm long; petals usually 4,
obovate, 2. 5-3. 5 (4) cm long and broad, golden yellow (pale orange), entire; sepals 2, early-deciduous, glaucous,
20
glabrous to short-strigose, enclosing the bud, oblong-lanceolate with minutely spined tips; flowers borne singly,
terminally or in branch axils; peduncles glaucous, ribbed, glabrous, 3-15 (25) mm long, stout and elongating little
in fruit, subtended by bracteal leaves which are ovate to lanceolate and shallowly lobed with acute, apiculate tips
and clasping bases, surfaces glaucous, glabrous or with a few short, stout-based hairs; cauline leaves similar, but
larger and more deeply lobed, up to 10 cm long, 4 cm wide; basal leaves in a dense rosette, up to 35 cm long, 10
cm broad, deeply pinnatisect, lower lobes serrate and dentate, the upper more deeply lobed with apiculate tips,
surfaces sparsely to densely white-tomentose; petioles tomentose, flattened and basally strongly ribbed, 2-5 cm
long (absent on cauline leaves); stems stout, ribbed, glaucous, glabrescent, branched above, up to 9 dm tall, from a
tough first-year caudex and stout biennial or perennial taproot system with yellow latex. (2n = 12, 24)
Infraspecific Variation: In Europe G. flaviim is distinguished from G. leiocarpum Boiss. on the basis of its capsule
being tuberculate throughout its length, and lacking constrictions between the seeds. In our materials the tubercu-
late surfaces are primarily on the upper portions of the capsules (as in G. leiocarpum), but capsules lack constric-
tions as they should in G. flavam.
Importance: The Homed Poppy is widely cultivated and escapes in sandy or disturbed soils, especially along
coasts. The oil from pressed seeds has been burned in lamps and used in food preparation and soap manufacture.
In folk medicine the extract of the plant is known as a purgative, hydragogue and sedative. The juice has been
used an an adulterant of opium.
4. MACLEAYA
Common Names: Plume Poppy, Tree Celandine
Authority: R. Brown, in App. Denh. Trav., p. 218, 1826
A genus of two species of eastern Asian herbs. They are atypical of the Poppy Family, having no petals and few
seeds (one in M. microcarpa) per capsule. They are most closely related to the genus Bocconia in which they were
first described. As cultivated plants, Plume Poppies are grown for their foliage since the flowers are not showy.
The following species escapes cultivation in New York State.
21
1. Macleaya cordata (Willd.) R. Br.
Common Names: Plume Poppy, Tree Celandine
Type Description: Willdenow, Species Pi., ed.
2, p. 841, 1799
Synonyms: Bocconia cordata Willd., B. japonica
Andre, B. cordata var. thunbergii Miq.
Origin: Eastern Asia
Habitats: Disturbed ground and sandy places as
an escape from cultivation
Habit: Tall, colony-forming perennials
Flowering: Late July-August
Fruiting: August-Oetober
General Distribution: Northeastern United
States as an escape; native of Asia, also escap-
ing cultivation there
Description: Plants with bisexual flowers; stigma 1 per ovary, glandular, clavate or 2-lobed, 0.5-1. 2 cm long, about
half as wide, persistent but enlarging little in fruit; style 1, short, cylindric, 0.3-0. 8 mm long in fruit; ovary 1 per
flower, superior, slightly stipitate, 3-5 mm long, 1-2 mm wide, clavate, becoming an oblanceolate, 2-valved, 2-
loculed capsule, 8-18 (22) mm long (smaller ones aborted), 4-5 (7) mm wide, flattened with 2 slender ribs at the
margins, dehiscing as the valves break away from the ribs in the upper half of the fruit, capsule stipitate at base;
seeds (2) 4-6, plump, ellipsoidal, dark purplish-brown when mature, surface with an indistinct reticulum the aril
plump, linear-lanceolate, ca 1 mm long, creamy-golden; stamens 20-30 in number, 5-8 mm long; anther sacs
linear, 3-5 mm long ca 0.3 mm wide; filaments 2-3 mm long, delicate, filamentous; petals absent; sepals 2,
cucullate, 5-9 mm long, ca 2 mm wide, creamy-white to pale salmon-pink; inflorescences l~several, terminal and
axillary, much-branched panicles, lanceolate to lance-elliptic in outline, (5) 20-50 (65) cm in length, about Vz as
broad; pedicels (3) 4-7 mm long, slender and conspicuously jointed to the receptacle; floral bracts subtending the
pedicels coriaceous, 3-ribbed, lanceolate, 1. 5-2.0 mm long with lacerate to glandular-ciliate margins; peduncle and
rachis of the inflorescence glabrous, terete, somewhat ribbed, glaucescent, the peduncle 3-15 cm long; leaves
much reduced upward on the stem, suborbicular to ovate-cordate in outline, with 5-11 large lobes and oval to
elliptic sinuses, minor lobing and blunt dentations near the lobe tips, glaucous, densely white, short-villous below,
glabrous above, lower leaves up to 30 cm long, 25 cm wide, the uppermost around 5 cm in diameter; petioles (1)
22
2-16 cm long, glabrous, ribbed, glaucescent, slightly clasping at base; stems terete, ribbed, glaucous, especially at
the nodes; plants up to 2.8 m high from a stout, perennial rootstock. (2n - 20)
Importance: This species is grown mainly for its foliage as a border plant or to fill in areas of the garden difficult to
maintain. Its aggressive spreading and colony-forming tendencies and persistence after escape have not helped its
popularity as a cultivated plant.
5. ESCHSCHOLZIA
Common Names: California Poppy, Mexican Gold
Authority: Cham, in Nees, Horae Phys. Berol., p. 73, 1820
A genus of 10-14 species native to the west coast of North America. They are showy, widely cultivated and bred for
color forms. The most commonly cultivated species, E. californica, is known to escape and persist in New York State.
1. Eschscholzia californica Cham, ex Nees
Common Name: California Poppy (State Flower
of California)
Type Description: Chamisso in Nees, Horae
Phys. Berol., p. 73, 1820
Synonyms: Escholtzia and Eschscholtzia are
common misspellings of the generic name; Es-
chscholzia crocea Benth., E. douglasii Benth,
E. peninsularis Greene, E. maritima Greene,
E. cucullata Greene, E. glauca Greene, etc.
(a large number of additional synonyms and
varietal combinations exist which have no
bearing on our garden escapes which have tra-
ditionally been treated as E. californica )
Origin: West coast of North America
Habitats: Meadows, fields, roadsides as a garden
escape
Habit: Freely branching annual or perennial
herbs
Flowering: June-September (NY), from Febru-
ary on in CA
Fruiting: July- October
General Distribution: Primarily cismontane Cali-
fornia; escaping cultivation elsewhere
23
Description: Plants with bisexual flowers; stigmas 4 (6), linear, yellow to orange, 3-5 mm long, usually persistent
in fruit; styles short or indistinguishable; ovary 1 per flower, superior, fusiform-cylindric, 3-5 mm long, 1-2 mm
wide, becoming a tan, linear capsule 3-8 (10) cm long, 1-3 (5) mm wide, glabrous, 10-nerved, 2-valved, dehiscing
from the base; seeds numerous, borne on 2 placentae, spherical to elliptic, 1.5— 1.8 mm long, 1.2-1. 8 mm wide,
seedcoat dark ocher to black with a tan reticulum overlying it, the reticulum itself minutely pitted; aril absent,
stamens 16-many, usually golden-orange; anther sacs 4-8 mm long, linear; filaments 0. 2-4.5 mm long, filament-
ous, yellowish, free; petals 4 in two series, free, cuneate with broad, rounded tips, 2-6 cm long and broad,
overlapping upward to make the corolla a shallow cup, deep orange to bright yellow or pinkish (white); sepals (2)
fused to form a hood-like calyptra 1-4 cm long, 0.5-2. 5 cm broad, glabrous, pushed off by the expanding petals;
receptacle expanded into a collar-like torus, 2-rimmed, the inner erect and hyaline, the outer ribbed, spreading
2-4 mm wide; flowers borne singly at branch tips; peduncles 3-15 cm long, ribbed; leaves much-dissected with
linear segments, glaucous, glabrous to densely white-puberulent, up to 6 cm long, 4 cm broad; petioles 1-4 (5) cm
long, winged and ribbed; stems terete, ribbed, much-branched, 2-7 dm long, ascending or lax, from an annual or
perennial taproot and fibrous root system with pale, watery juice. (2n = 12)
Infraspecific Variation: This is an extremely polymorphic species even as it occurs in nature, with over 50 species
names applied to its members in the past. The more easily recognizeable varieties are explained in Munz (1963). In
addition to natural variation, cross-breeding by horticulturists has proliferated the number of forms, many of which
have names in the trade, such as; “Alba”, “Crocea” and “Rosea”.
Importance: These plants are popular in the seed-packet trade, grown as annuals, and may sometimes form winter
rosettes and persist after escape from cultivation. California Indians ate the greens boiled or roasted over hot
stones, and they were reported to have used the mildly narcotic juice as a toothache remedy.
PAP AVER
Common Name: Poppy
Authority: Linnaeus, Species PL, p. 506, 1753
A genus of 60-80 species of herbs distributed primarily in the northern hemisphere. Many species are concen-
trated around the Mediterranean Sea. They are widely cultivated and commercially valuable for their alkaloids.
Description: Plants with bisexual flowers; stigmas (2) 4-20, usually forming a persistent, radiate disc which is
sessile on the capsule; style absent or very short and broad; ovary 1 per flower, superior, oblong or orbicular,
glabrous or with setae, becoming a globose to oblong or obconic capsule with 4-20 carpels, unilocular or with
partial partitioning, dehiscent by pores (rectangular hatches) or indehiscent; seeds numerous, borne parietally,
often with reticulate and pitted surfaces; stamens centripetal, usually numerous; petals usually 4, free; sepals 2 (-4),
enclosing the bud, early-deciduous; peduncles elongating in flower and fruit, sometimes scapose, glabrous, setose
or pubescent, often glaucous; cauline leaves oval to lanceolate, usually dentate or pinnately divided or lobed, often
glaucous, glabrous to densely pubescent, petiolate, sessile or clasping; basal leaves often narrower, forming well-
defined rosettes in biennial species, otherwise much like cauline leaves; stems terete or unevenly ribbed, contain-
ing milk or colored sap and latex, arising from an annual or biennial taproot or perennial rootstock.
24
KEY TO SPECIES OF PAPAVER
1. Upper cauline leaves petioled or sessile, but not clasping the stem at their bases; buds finely pubescent or
setose (3)
1. Upper cauline leaves with clasping bases; buds usually glabrous, glaucous. (2)
2. Buds 2-4 cm long; capsules 2-6 (9) cm tall; flowers variously colored and spotted, but rarely red
1. Papaver somniferum (p. 25)
2. Buds 1-2 cm long; capsules 1.8 cm tall or less; flowers red with blackish basal spots
■ . .2. Papaver glaucum (p. 27)
3. Mature capsules not much longer than wide, subglobose to oblong or broadly conic (5)
3. Mature capsules about twice as long as wide, narrowly oblong or obconic (4)
4. Anthers yellow; plants perennial; capsules obconic, strongly ribbed 3. Papaver atlanticum (p. 29)
4. Anthers purple; plants annual; capsules narrowly oblong, not strongly ribbed .
• - 4. Papaver dubium (p. 30)
5. Petals 4-6 cm long; robust, perennial plants; buds mostly over 2 cm long 5. Papaver orientale (p. 31)
5. Petals 1.5-3. 5 cm long; slender to moderately robust annual plants; buds mostly less than 2 cm long
6. Papaver rhoeas (p. 33)
1. Papaver somniferum L.
Common Names: Opium Poppy, Common
Poppy, Garden Poppy, White Poppy
Type Description: Linnaeus, Species PL, p. 508,
1753
Synonyms: Papaver setigerum DC. (in part), P.
“mursellii” , P. somniferum ssp. hortense
(Huss.) Corb.
Origin: Western Mediterranean region, cultivars
probably derived from P. somniferum ssp. seti-
gerum and modified considerably by selection
Habitats: Waste places, old gardens, dumps, es-
caping (illegal) cultivation onto disturbed
£p-ound
Habit: Erect, sparsely branching annual
Flowering: June- September
Fruiting: July- October
General Distribution: Native to southern
Europe, but escaping cultivation throughout
the world in all but the most severe climates
25
Description: Plants with bisexual flowers; stigmas (5) 8-15 (20), sessile, forming a radiate stigmatic disc which is
flat or slightly concave or convex with conspicuous interstigmatic tissue, persistent becoming woody to papery in
texture in fruit and 8-40 mm in diameter, the pollen-receptive areas purplish-glandular radiating in a star-pattern
from the disc center to the free stigmatic lobes which may be irregularly truncate or somewhat rounded to obtuse;
styles absent; ovary 1, subglobose, smooth, enlarging greatly to become a fleshy capsule 2-6 (9) cm tall and 1.5-
6.0 (7) cm wide, with 5-20 inconspicuous nerves corresponding with carpels, sometimes dehiscent from above,
but often indehiscent in cultivated strains, subglobose to oblong, bluish green to tan with a glaucous sheen and
purple hue; seeds numerous, reniform, 1.0-1. 5 mm long, ca 1 mm in other dimensions, seed coat white, gray,
amber or bluish to black with compound reticulations and pits; stamens numerous, 1-2 cm long; anther sacs
linear, ca 4 mm long; filaments pale, slender; petals 4, broadly ovate to somewhat cuneate entire, (or lacerate), 3-
6 cm long and broad, with many pseudo-parallel veins which are often accompanied by pigmentation, white or
blue to rose-pink, deep mauve or crimson, often with streaks of above colors, each petal with a basal, dark spot of
color 1. 0-2.5 cm in diameter; sepals 2, early-deciduous, enclosing the bud, smooth, 2-4 cm long, 1-3 cm wide;
buds oblong; peduncles stout, ribbed, up to 30 cm long, glaucous and glabrous or with bristly setae in the upper
3-4 cm; upper cauline leaves sessile, perfoliate, irregularly dentate and shallowly toothed and/or lobed, glaucous,
glabrous and bluish-green, conspicuously reticulate-veined, the upper leaf pair often 3-5 cm long, 2-4 cm broad;
middle cauline leaves similar, but more deeply cut and pinnately lobed, much larger, up to 40 cm long, 15 cm
wide or more, sessile-auriculate; lower leaves somewhat smaller, narrower, often oblanceolate, not as clasping and
sometimes with short petioles; stems tough, shiny-glabrous, fluted and ridged, up to 1.5 cm thick near base, and
up to 1.5 meters tall, sparsely branched or unbranched, from a tough, deep taproot with fibrous secondary roots.
(2n = 22)
Infraspecific Variation: Because P. somniferum is a multi-use plant cultivated by man, it is extremely variable with
a history of selective breeding for drug production, seed production and for attractive horticultural features. It was
undoubtedly a variable species in nature also, and it is considered to have one native subspecies in Europe: ssp.
setigerum (DC.) Corb. (also a weed) and two in cultivation. Subspecies somniferum was probably originally derived
from ssp. setigerum , from which it differs in being larger in most aspects, having about twice the number of
stigmatic lobes, and in rarely being setulose. Subspecies songaricum Basil, is much like ssp. somniferum but differs
in having flat, obtusely tipped stigmatic lobes rather than sulcate, truncated ones. Both the cultivated subspecies
have escaped in New York State occasionally. In addition to these variations, there are a considerable number of
flower-size and color forms, and seeds vary widely in color patterns. Double-flowered plants are known under the
name “Carnation Poppy”.
Importance: Under U. S. federal law it is illegal for an unlicensed individual to obtain, transport, grow or permit
this species to be grown on land under his/her control. Prior to regulation this species was known as the Common
Poppy, and was one of the more frequent poppies in garden cultivation. It has now largely been replaced by
orange and red or pink and yellow flowered species, but many people are unaware or resist the law and continue to
grow it. Opium Poppy derives its epithet, somniferum, from the long-known sleep-inducing qualities of the milky
extract of the capsule. It is clear from Summerian idiograms that its narcotic properties were known at least 6,000
years ago. Plant remains have also been found in sites of dwellings some 4,000 years old of Stone Age lake dwellers
of Switzerland. The mind-altering effects of opium, its derivatives and precursors have much changed legal, medi-
cal, political and religious history over the centuries. Fortunes, governments and empires have risen and fallen as
the result of events involving opiates. Our own Hudson River was explored by Henry Hudson as he sought a
fabled Northeast Passage to Cathay in the Orient, which provided much of the opium that was one of the more
profitable and highly prized cargos of the day. Papaver somniferum and its subspecies have been extensively
studied, since they represent the only plants so far found to produce morphine. They also accumulate thebaine and
codeine which are morphine precursors, as well as various benzylisoquinolines. Eating of the unripe fruit has long
been known to cause stupor, coma and shallow, slow breathing; smoking opium (with tobacco) produces similar
effects when maintained past an initial euphoric state. Raw opium was used as a poison, even in ancient times, and
has a long history of abuse and use in murder and suicide — often disguised as medicine, for which it is also used.
Opiates have been used historically in treatment of pain and for narcotic, antispasmodic, hypnotic, anodyne,
analgesic and sedative effects, and to induce respiratory depression. Capsules have been used as fomentation in
sprains, bruises, neuralgia, toothache, etc. The two major grades of opium are soft (shipping) and druggist’s opium,
though many commercial names have been applied, usually having to do with the country of origin. Licensed drug
26
manufacturers produce for medical use: codeine, norcotine, laudenine, papaverine and the more dangerously ad-
dictive morphine and heroin (diamorphine). The latter is widely produced illegally and uscrupulously marketed
for high prices. By contrast, codeine is mild enough to have been used in Paragoric to sooth colicky infants to
sleep. For a few infants, however, the effect may be to stimulate wakefulness and activity for many hours. Poppy
seeds contain only a slight trace of opium, and are not known to produce a narcotic e fleet even when ingested in
quantity. They are commonly used in baking, sprinkled on rolls and bread. Black seeds are preferred for this
puipose. Poppy seeds have been reported to prevent scurvy, as they contain some Vitamin C. Paler seeded forms
are preferred for making emulsions, and the oil of the poppy seed is used in foods (such as salad dressings) and in
the manufacture of paint, varnish and soap, due to drying and adhesive properties.
2, Papaver glaucum Boiss. & Hausskn.
Common Name: Tulip Poppy
Type Description; In Boissier, Fl. Orientale, vol.
1, p. 116, 1867
Origin: North Africa
Habitats: Disturbed ground as a rare escape
Habit; Erect and spreading, branched, annual
herbs
Flowering: June-August
Fruiting: July-September
General Distribution: Native to Syria, Iraq, Iran,
cultivated and escaping widely in North Africa
and Asia, less commonly elsewhere
KM Con/tA/AV
27
Description: Plants with bisexual flowers; stigmas (6) 8-15 (18) on a radiate disc, persistent in fruit, becoming
papery to woody in texture, the disc flat to somewhat concave, 0.5-1. 1 cm broad, receptive surfaces purplish,
forming a narrow, star-like, radiate pattern from the disc center to the stigmatic lobes which are rounded to blunt
with entire to wavy (rarely laciniate) margins; styles absent; ovary 1 per flower, superior, subspherical to oblong,
glaucous, with 6-18 carpels, the single cavity partially divided by carpel walls, forming imperfect locules as it
develops into a capsule which is ovoid to subspherical, ca 1 cm tall and broad, dehiscent by pores at the lower
margin of the stigmatic disc; seeds numerous, borne parietally, 0.8- 1.2 mm long, ca 0.7 mm wide, with a complex,
deep-pitted reticulum, brownish when dry, reniform to distinctly comma-shaped; stamens 100 or more, 1-2 cm
long; anther sacs ca 2 mm long, linear, brownish, appearing golden with pollen at anthesis; filaments dark purple-
brown to black, filiform; petals 4 (-6), ovate to cuneate, 3-5 cm long, 3-5 (-7) cm broad, scarlet (to brick red)
each with a brownish-black spot near the base and a white line along the upper margin of the spot; sepals 2(-3),
early deciduous, 1. 2-2.1 cm long and broad, forming a subglobose bud, glabrous, slightly glaucous, darkly spotted
at the immediate base; peduncles ribbed, elongating during flower and fruit development, up to 35 cm long,
densely appressed-hirsute above, less so below; cauline leaves glabrous, glaucous, (2) 4-20 cm long, 1-4 cm wide,
lanceolate to obspatulate, clasping the stem, margins dentate, often pinnately narrow-lobed, the lobes obtusely
tipped to minutely apiculate; basal leaves forming a loose rosette, similar to cauline leaves but up to 25 cm long
and sometimes with short petioles; stems branching near the bases (or single), glabrous or puberulent, ribbed, up
to a meter tall, from a wiry annual (biennial?) taproot and fibrous lateral roots and a milky sap.
Infraspecific Variation: Like other poppies this species may be dwarfed in its dimensions by poor soil and dry
conditions; it may also vary somewhat in coloration, but it is much more stable in this than its close relative, the
Opium Poppy.
Importance: Tulip Poppy is a beautiful plant in cultivation, but it has not received the attention in North America that
it has in Africa and Asia where it more frequently escapes.
28
3. Papaver atlanticum (Ball) Cosson
Common Name: Morocco Poppy
Type Description: Ball, 111. Fl. Atl. vol. 1, p. 11,
1882
Origin: North Africa
Habitats: Waste places, old gardens, roadsides as
a rare escape
Habit: Short-stemmed perennials with tall scapes
Flowering: May-August
Fruiting: May- September
General Distribution: Native of Morocco, occa-
sionally escaping cultivation in Eurasia and
North America
K.PA. CONWAV
Description: Plants with bisexual flowers; stigmas 6 or 8 (10) on a flat to pyramidal radiate disc with purplish,
glandular receptive ridges, the star-shaped pattern terminating in the 6-10 stigmatic lobes which are rounded to
obtuse with undulate margins, the disc persistent in fruit becoming 6-12 mm broad and papery in texture; styles
absent; ovary 1 per flower, superior, clavate, 7-15 mm long, strongly ribbed, glaucous, becoming a clavate cap-
sule, greenish to brown, 1.5-2. 8 cm long, 4-11 mm broad at summit, strongly ribbed, dehiscing by pores just
below the margin of the stigmatic disc; seeds numerous, borne parietally, ca 0.7 mm long, 0.5 mm wide, plump
reniform to comma-shaped, shallowly reticulate; stamens numerous, centripetal, 4-11 mm long, the inner ones
longest; anther sacs golden, linear-ovate, ca 2 mm long; filaments golden, flattened (or filiform above); petals
usually 4, ovate-cuneate to triangular, 2.5-4. 1 cm long and broad, pale orange to dull orange-red, unspotted or
with a diffuse, creamy spot near each petal base; sepals usually 2, early deciduous, 1.0-1. 5 (1.9) cm long, 0.7-1. 2
cm wide, sparsely to densely pilose, forming an oblong bud; flowers solitary on scape-like peduncles; peduncles
15-60 cm long, terete or ridged deeply, appressed-hispid to hispid below; cauline leaves (2) 5-30 cm long,
oblanceolate to spatulate, pinnately lobed with few, widely spaced, narrow lobes on the winged rachis below, and
many dentate and shallowly incised lobes near the tip; basal leaves similar, up to 35 (40) cm long, densely hispid,
forming a loose rosette; petioles winged from near the bases, contiguous with the blades, hispid; stems hispid,
ribbed, up to 18 cm tall, usually short, the plant deriving most of its height (up to 8 dm) from the scapose
peduncles; plants arising from tough perennial taproots and fibrous root systems with slightly milky juice.
29
Infraspecific Variation: Plants may be dwarfed and almost stemless; flower color varies from deep orange to almost
white, and spots are not always present. This species is very closely related to P. rupifragum Boiss. & Reut.,
which differs in being subglabrous and in its smaller, brick-red flowers. Papaver lateritium C. Koch has larger,
brick-red flowers, an obovate-clavate capsule like P. atlanticum, and may be conspecific with it.
Importance: This species is not a common cultivar, but it escapes occasionally in Eurasia and North America.
Common Names: Blind Eyes, Grainfield Poppy,
Headache Poppy, Long-pod Poppy, Smooth-
fruited Poppy
Type Description: Linnaeus, Species Pi., p.
1196, 1753
Synonyms: Papaver obtusifolium Desf., P. mo-
destum Jord., P. nothum Steven, P. mairei
Batt., P. laevigatum Beib.?, P. albiflorum
(Bess.) Pacz.
Origin: Southern Europe
Habitats: Grainfields, waste places, vacant lots,
roadsides as an escape or adventive weed
Habit: Erect or ascending, unbranched or basally
branching annuals
Flowering: May-August
Fruiting: May- October
General Distribution: Widespread escape, espe-
cially in warmer climates
Description: Plants with bisexual flowers; stigmas usually 7-9, radiate on a persistent, convex stigmatic disc which
is obtusely pointed at the summit, with its glandular receptive areas borne along prominent ridges radiating to the
stigmatic lobes, the lobes obtuse, yellow-green to tan, somewhat leathery and undulate-margined, the stigmatic
disc becoming more flattened and up^ to 1 cm broad in fruit; styles absent; ovary 1 per flower, superior, oblong,
3-10 mm long, 4-5 mm wide, smooth, becoming an oblong-cylindric to obovoid capsule, 1.5-2. 2 cm long, 0.7-
1.0 cm wide, glabrous, yellow-green to tan and glaucous with visible ribs, opening by pores at the base of the
stigmatic disc; seeds numerous, borne parietally, 0.6-0. 8 mm long, ca 0.4 mm wide, reniform to comma-shaped,
purplish-gray to brownish, shallowly and boldly reticulate; stamens numerous, 5-12 mm long; anther sacs plump,
ca 1.5 mm long, purplish-lavender; filaments deep violet to purple-brown, filiform; petals usually 4, broadly oval to
suborbicular, 1. 5-3.0 (3.5) cm long and broad, red to rose-pink or red-orange, unspotted, or each petal with a dark
spot near the base; sepals 2 (-3), early deciduous, 1.2-1. 7 cm long, densely hispid, forming an oblong-ovate bud;
peduncles 15-30 (40) cm tall, appressed-hispid to strigose above, hispid below; cauline leaves 4-15 cm long 2-7
30
cm wide, bipinnately (or pinnately) lobed and dissected, the lobe tips setose, surfaces appressed-hispid, glauces-
cent; basal leaves less divided, pinnate or merely lobed; petioles short, winged, confluent with the blades; stems
hispid, ribbed, 2-6 dm tall, caulescent to subscapose, often branched near the base, from a tough, pale, annual
taproot system with milky juice and latex. (2n = 42)
Infraspecific Variation: This is a variable species in most of its features. Dwarfed plants which produce small or
abortive fruits may be difficult to distinguish from P. rhoeas under similar circumstances. The species complex to
which it belongs in Europe includes about seven taxa which might be better included in the broad range of
variability of a polymorphic P. dubium. They vary in flower, anther, filament and juice color as well as such
characters as leaf lobing, petal shape (size) and capsule attenuation. Such variations are sometimes seen in garden
varieties, belying past hybridization and artificial breeding. Least reliable among European segregates from this
species is perhaps P. albiflorum (Besser) Pacz. with white or pale pink, unspotted flowers.
Importance: Papaver dubium is cultivated, spread in hay and sown as a contaminant with grain seed. It can be a
troublesome weed in warmer climates, but it escapes only occasionally in New York State. Like other poppies it
contains isoquinoline alkaloids and should be considered poisonous.
5. Papaver orientale L.
Common Names: Oriental Poppy, “Old-
fashioned Poppy”
Type Description: Linnaeus, Species Pi., p. 508,
1753
Synonyms: Calomecon orientale (L.) Spach., Pa-
paver grandiflorum Moench., P. spectabile Sa-
lisb. , P. paucifoliatum (Trautv.) Fedde, P.
orientale var. paucifoliatum Trautv., P. orien-
tale var. parviflorum Busch
Origin: Southern Caucasus
Habitats: Disturbed sites, waste places, old gar-
dens, roadsides, as an escape from cultivation
(alpine in its native habitats)
Habits: Colony-forming, erect and spreading
perennials
Flowering: May-August
Fruiting: May- September
General Distribution: Escaping cultivation
rarely, but worldwide; native to Iran, Turkey
and southern Russia
31
Description: Plants with bisexual flowers; stigmas 8-15, on a slightly convex stigmatic disc which is persistent,
becoming leathery, up to 1.5 cm broad in fruit, the glandular receptive areas radiating from the center of the disc
along ridges to the stigmatic lobes which are obtuse or rounded with slightly undulate margins; styles absent;
ovary 1 per flower, superior, obovoid, 6-9 mm tall, 5-8 mm wide, becoming an oblong, obovoid or obconic
capsule, 1.2-2. 3 cm long, 1.0-1. 8 cm broad in the upper half, greenish-tan to brown when dry, glaucescent,
opening by pores below the rim of the stigmatic disc; seeds numerous, borne parietally, 0.6-0. 8 mm long, ca 0.5
nun wide, reniform to comma-shaped, grayish to brown when dry, evenly reticulate; stamens over 100, mostly
about 1 cm long; anther sacs 1. 8-3.0 mm long, oblong, pale violet to yellowish; filaments gray-brown to dark
purple, filamentous; petals 4 (6) or double with narrower inner petals, the outer ones suborbicular to broadly oval,
up to 6 cm long and broad, pale orange or orange-pink, lacking dark basal spots, but sometimes with pale, bluish or
white, rectangular markings near the petal bases; sepals 2 (3) in number, (2.0) 2.2-3. 4 cm long, cucullate, early
deciduous, with spreading-setose hairs, forming an oblong, pendulous bud; floral bracts absent; peduncles 10-36
cm long, nodding in bud, appressed-setose above, spreading-setose below; cauline leaves oblanceolate to spatulate,
strongly pinnatisect, especially below, with serrated lobes and tips bearing strong setae, surfaces spreading-setose,
the upper cauline leaf remote from the flower; basal leaves forming a rosette each year, longer than cauline leaves
reaching up to 60 cm long, 12 cm wide, otherwise similar; petioles varying greatly in length, winged and confluent
with the blades above, ribbed and strongly spreading-setose; stems caulescent ot almost acaulescent with 2-4
cauline leaves, stem bases surrounded by flat, but not sheathing, petiolar bases, (stems) terete or deeply grooved
and ribbed, spreading-setose above, to densely pilose near the bases, often 1-branched; plants up to 1.3 m tall
from thick, spreading, perennial rootstocks. (2n = 28)
Note: Papaver orientate has been confused in the literature with two related species which are also widely culti-
vated. Chemical, cytogenetic and taxonomic papers prior to 1974 are rich in mis-information on “P. orientate' as it
is treated there. See Goldblatt (1974) for clarification, in addition to the following brief summary:
Closely Related Species: Papaver bracteatum Lindl. (2n = 14) has well-developed, persistent, floral bracts; the
uppermost leaf is high on the stem, often proximal to the flower; buds are ± erect on the peduncles; petals are
blood-red with a black radial stripe near the base. The stigmatic disc is pyramidal at maturity.
Papaver pseudo-orient ale (Fedde) Medw., (2n = 42) may have one or more persistent floral bracts; the uppermost
leaf is high on the stem; buds are on ± erect peduncles; flowers are deep vermillion to orange, usually with dark
brown to black, rectangular or trapezoidal marks near the petal bases. Both P. bracteatum and P. pseudo-orientale
are commonly cultivated and often called “Oriental Poppy.’ Either or both may escape cultivation in the state,
though we presently have no records of this having occurred.
Infraspecific Variation: When separated from the two distinct species discussed above, P. orientate shows far less
variation than formerly indicated in the literature. Plants vary in the distance of the upper leaf from the flower, but
do not have leaves or bracts near the flower. Color of petals varies in shades of orange and pink (or white in
cultivar “Albiflora ”); pale spots may be present or absent from the petals; stamens vary from purple to brownish-
black but are consistently dark.
32
6. Papaver rhoeas L.
Common Names: Corn Poppy, Shirley Poppy,
Field Poppy, Red Poppy, Flanders Poppy,
African Rose, Corn Rose, Headache, Red-
weed, etc.
Type Description: Linnaeus, Species PL, p. 507,
1753
Synonyms: Papaver strigosum (Boenn.) Schur.,
P. timidulum Klokov, P. tenuissimum Fedde,
P. laciniaturn Hort., P. intermedium Hort., P.
integrifolium Vig. , P. rapiferum Fedde, P. um-
brosum Mott.
Origin: Mediterranean Region
Habitats: Waste places, roadsides, cultivated
ground as an escape from cultivation
Habit: Erect to spreading annuals (biennials)
Flowering: May-September
Fruiting: May-October
General Distribution: An aggressive weed of
cultivation in Eurasia and occasionally North
America, usually casually adventive; native to
southern Europe and northern Africa.
Description: Plants with bisexual flowers; stigmas 5-15 (18) on a radiate disc which is yellowish, leathery and
persistent in fruit, somewhat convex becoming flatter with maturity, 5-11 mm wide, the receptive stigmatic zones
glandular, purplish, along ridges radiating from the disc center to stigmatic lobes which are flared with overlapping
lobes, obtuse tips and wavy margins; styles absent; ovary 1 per flower, superior, ovoid, glabrous, 3-5 mm long and
broad, becoming a glabrous, glaucous capsule 8-15 (20) mm tall, 6-14 (16) mm wide, ovoid to subglobose, tannish-
green to brown at maturity with pale ribs, dehiscing by pores below the rim of the stigmatic disc, the tissue in that
region often further breaking down, leaving the disc suspended by internal tissue much like a parasol above a cup;
seeds numerous, borne parietally, 0. 7-1.0 mm long, ca 0.6 mm wide, strongly comma-shaped with a basal nipple,
amber with a purple sheen and a moderate to bold, simple reticulum of rectangular fields; stamens numerous, (2)
4-11 mm long; anthers subglobose to oblong, 0. 4-1.0 mm long, bluish-purple (often the pollen as well); filaments
lavendar to purple-black, filamentous; petals 4 (or more), broadly orbicular, entire (rarely lacerate), (1) 2-4 cm long
and broad, red, purple, rose, pink or white (various color combinations with white streaking in hybrid Shirley
Poppies), often with dark spots near the petal bases; sepals 2 (-4), hispid to villous, 0.5-2. 1 cm long, forming an
oblong bud; peduncles 7-40 (60) cm long, sparsely to densely hispid, especially above, shallowly to deeply ribbed.
33
nodding (or erect) in bud; cauline leaves hispid to glabrescent, 3-20 cm long, 1- cm wide, reduced upward, the
upper ones subopposite, clustered beneath peduncles, pinnately or bipinnately lobed (unlobed in dwarf plants),
often caudate, coarsely dentate with acuminate, setose tips; basal leaves similar, but more broadly (sometimes
bluntly) lobed, sometimes forming a rosette, up to 30 cm long, 12 cm wide near the sparingly lobed tips; petioles
absent or present only on the lower leaves, ribbed, hispid; stems slender to relatively stout, ribbed, hispid, simple
to 4-5 branched near the base; plants 1-9 dm tall arising from a tough, yellowish annual or biennial taproot with
fibrous roots and milky sap. (2n = 14)
Infraspecific Variation: Plants growing under severe drought conditions may be dwarfed in nearly all characters,
having narrow, non-overlapping petals, subentire leaves etc., and may be mistaken for depauperate P. duhium if
fruits are not well developed. Color variations in this species are numerous, including striping and spotting com-
binations as well. Shirley Poppies of the horticultural trade are bred for such variation, and are commonly streaked
with white and/or white blotched near the petal bases, with whitish filaments and yellow (rather than bluish)
pollen. Stems may be slender and unbranched to robust with several stout basal branches. Peduncles may have
appressed rather than spreading hispidity. A closely related species (perhaps better treated as a variety) is P.
commutatum Fisch. & Mey. The capsule is broad and slightly stipitate, foliage is villous and the black spots are
most prominent toward the centers of the petals.
Importance: Papaver rhoeas is the most commonly cultivated annual Poppy in most parts of North America. If one
orders poppy seeds from a catalog, the liklihood of receiving this species is extremely great, no matter what species
was advertised. The artificial American Legion poppy was designed with Shirley Poppy in mind. Papaver rhoeas
does not usually persist for long in New York State unless the soil is continually disturbed, as in cultivated fields or
dumps. In some parts of Eurasia it is known as a noxious weed of cultivated fields. The petals of Com Poppy have
been used in folk medicine for centuries, primarily as an expectorant and (reputed) mild pain killer, but mostly for
the rosy color imparted by flower pigments to the solution. These pigments have also been used to color wines.
Papaver rhoeas has been implicated in livestock poisoning in Europe. The plants are usually distasteful to animals,
but may enter their diet as a contaminant of hay or feed-grain.
7. ARGEMONE
Common Names; Prickly Poppy, Mexican Poppy
Authority: Linnaeus, Species Pi., pp. 508-509, 1753
A genus of about 20 species native to areas of low rainfall from southern North America to South America (one
species in Hawaii). The two species found in New York are infrequent escapes from cultivation, but are native
further south in the United States. These poppies are popular as garden ornamentals, and they also have alkoloids
which have been used medicinally.
Description: Plants with bisexual flowers; stigma 1, with 3-5 (7) lobes corresponding to carpel number, the lobes
flattened with felt-like, purplish receptive areas; style 1, very short, evident only in mature fruit; ovary 1, unilocular,
but of (3) 4-5 (-7) carpels and their many-ovuled placentae, becoming a dehiscent capsule, elliptic to oblong-ovate or
linear-oblong, glabrous to very spiny, dehiscing with valves splitting along fluted sutures, the vascular elements
remaining within as a cage-like framework; seeds brownish to black, 1-3 mm long; endosperm copious; embryo
straight, minute; stamens 20-250 or more; anther sacs linear, coiled after dehiscence; filaments filiform or rarely
clavate; petals usually 6, in 2 whorls, 1.5-7. 5 cm long, making flowers up to 15 cm broad, usually yellow, golden or
white (lavender-tinged); sepals early-deciduous, usually 3 (2-6), smooth to prickly or hispid, each with a subterminal
horn which is hollow and vegetative below and spined at the tip; flowers borne singly at branch tips and in their upper
leaf axils; peduncles stout, varying in length, subtended by single or paired foliar bracts; leaves reduced upward on the
stem with a basal rosette; cauline leaves glaucous, sessile, often clasping, dentate, each tooth terminating in a prickle,
the margins also usually lobed, oval to oblong in outline; basal leaves similar, oblong to oblanceolte; stem glaucous,
erect or ascending, 1-10 (-20) dm tall, smooth to closely prickly, from a persistent or ephemeral, annual or biennial
taproot with fibrous roots.
KEY TO SPECIES OF ARGEMONE
1. Flowers white; stamens more than 100
1. Flowers yellow; stamens 50 or fewer . .
34
.1. Argemone albiflora (p. 35)
2. Argemone mexicana (p. 37)
1. Argemone albiflora Hornem.
Common names: Prickly Poppy, White Prickly
Poppy
Type Description: Homemann, Hort. Hafn., p.
489, 1915
Synonyms: Argemone alba Lestib. non James
( nomen nudum), A. georgiana Croom, A. mex-
icana of authors non L., and white-flowered
plants included under it, A. mexicana var. B
Lamarck, A. mexicana li albiflora DC., A. in-
termedia Sweet (in part) as to 111. specimens (a
name excluded in Ownbey’s 1958 monograph)
Origin: Northern coast of the Gull of Mexico
Habitats: Roadsides, waste places, beaches and
dunes in sandy soil or gravelly clay, mostly ad-
ventive from gardens
Habit: Erect or spreading annuals or biennials
Flowering: (March) May-August
Fruiting: (April) June -October
General Distribution: Coastal Plain of the
southern United States from Mississippi to
North Carolina (possibly native only to
Northern Florida) escaping cultivation in New
York, New England, Illinois, Kentucky and
Arkansas
Description: Plants with bisexual flowers; stigma 1, 3-7 lobed, corresponding with carpel number, purplish be-
coming 3-4 mm broad and 2-3 mm tall in fruit where it is persistent; style very short, only evident in fruit; ovary
1, oblong with (3) 4-5 (7) carpels, becoming an oblong to oblong-elliptic capsule 12-15 (25) mm wide (excluding
spines) and 20-40 (45) mm long, armed with widely spaced, stout, spreading spines which are basally herbaceous,
unbranched and mostly 2-8 (10) mm long; seeds blackish-brown, 1.9-2. 1 mm long, ovoid with a bold reticulum and
concavities; stamens 7-12 mm long 100 or more; anther sacs linear; filaments slender; petals 6 (rarely 8) in two
whorls, white, obovate-cuneate, mostly 2-4 cm long, 2. 5-4. 5 cm wide; sepals usually 3, valvate in bud with
subterminal horns 1-3 mm long which are prickle-tipped, early deciduous, each sepal ca 2 cm long, 1.3 cm wide,
cucullate and sparsely spiney; spines of sepals 2-3 mm long; buds tending to be clustered in close proximity to
foliar bracts on very short peduncles which elongate during flowering and fruiting ranging from a few mm to 3 cm
in length during this process; peduncles stout, glaucous green to brown in fruit usually with a number of spines
toward the summit; bracteal leaves smaller, but like other cauline leaves; cauline leaves ovate to narrowly obspatu-
late, lobed %-% the distance to the midrib, glabrous above, spiny on the major veins below, coarsely toothed with
35
spine-tipped dentations, larger leaves usually 15-20 cm long, 8-10 cm wide, somewhat glaucous; basal leaves in a
rosette, narrowly oblanceolate, mostly 7-15 cm long, 2-4 cm wide, otherwise like cauline leaves; stems stout,
somewhat ribbed, glaucous, green to reddish-brown, up to 1.5 m tall, often branched, sparsely to moderately
prickly, from a deep taproot with yellow latex. (2n = 28)
Infraspecific Variation and Hybridization: This species is a member of a complex of white-flowered plants that
naturally range from northern Florida to Texas and Montana. Escape from cultivation in the east and midwest has
resulted in varied interpretations of the taxa involved, such as in Illinois where two subspecies of A. albiflora may
be found. Ownbey (1958) recognized A. albiflora ssp. texana Ownb. which differs from ssp. albiflora in its narro-
wer capsules which have numerous small spines interspersed between the large ones; the seeds are also considera-
bly smaller, averaging about 1.7 mm. Ownbey also recognized A. polyanthemos (Fedde) Ownb. which he renamed
as a segregate which had formerly been known as A. intermedia Eastw. (incorrectly A. intermedia Sweet). These
taxa all have a somatic chromosome number of 28 and hybridize freely both experimentally and where they come
into contact after escaping cultivation. Of Argemone polyanthemos, which ranges from Texas north to Montana,
Ownbey said: “A. polyanthemos is distinguished by the sparingly prickly stems, the succulent, usually shallowly
lobed leaves which are devoid of prickles on the upper surfaces, the upper ones usually clasping, and by the
stoutly, but distantly spinescent, elliptical capsules. The species is closely related to A. albiflora Hornem., and
treated more conservatively, A. polyanthemos would probably be made a subspecies of the latter.’ All three of the
entities discussed here are in cultivation and might possibly escape within our state borders.
Importance: A. albiflora and its close relatives have Protopine and Berberine type alkaloids but lack the Pavine
type also known in the genus. Though this species is not commonly cited in the literature, it can be poisonous
causing vomiting, and if ingested by livestock the oily principle may be transmitted to humans through the milk as
has been reported in A. mexicana. The plants are usually distasteful to cattle, but may be introduced into their
feed as a contaminant. A. albiflora and its relatives are widely cultivated worldwide in relatively dry garden sites.
36
2. Argemone mexicana L.
Common Names: Prickly Poppy, Mexican
Poppy, Mexican Thistle, Devil’s Fig, Bird-in-
the-bush, Yellow Thistle, Jamaica Thistle,
Queen Thistle
Type Description: Linnaeus, Species PL, p. 508,
1753
Synonyms: Ectrus mexicanus (L.) Nieuwl., Arge-
mone spinosa Moench, A. versicolor Salisb.,
A. sexvalvis Stokes, A. vulgaris Spach, A. mu-
cronata Dum., A. leiocarpa Greene
Origin: Probably the West Indies (may be native
to Florida)
Habitats: Waste places, roadways, weedy in
open, disturbed sites
Habit: Erect or spreading, annuals
Flowering: April-October (year-round else-
where)
Fruiting: May-November (year-round else-
where)
General Distribution: Originally from the Car-
ribbean area, this plant has become a prolific
weed through the warmer parts of the world,
but rarely escapes in New York State.
Description: Plants with bisexual flowers; stigma 1, 4-6 lobed, corresponding with carpel number, purple, persist-
ent, up to 2 mm high and 4 mm wide in fruit; style very short, usually visible only in fruit where it becomes 1-2
mm long; ovary, 1, oblong with 4-6 carpels, becoming an oblong to broadly elliptic capsule 12-20 mm wide
(excluding spines) and 25-45 mm long, armed with stout, ascending spines, widely spaced, mostly 6-8 (10) mm
long (or lacking spines in forma leiocarpa ); seeds 1. 6-2.0 mm long, brownish-black, reticulate due to small, often
narrow concavities; stamens 30-50 in number, 6-12 mm long, yellow; anther sacs linear, curling after dehiscence;
filaments slender; petals usually 6, obovate to obcuneate, 1.5-3. 5 cm long and wide, bright yellow; sepals usually
3, valvate in bud, each with a subterminal horn, cucullate, glaucous, sparingly prickly or smooth, 9-15 mm long
and wide, the horns 2-8 mm long; buds spherical or slightly oblong; flowers single, terminal or axillary on stout,
spiny or glabrous peduncles up to 3 cm long; bracteal leaves like other cauline leaves, but often smaller; cauline
leaves lobed V2 - % the distance from the margin to midrib (or less upward on smaller leaves) with broad, oval
37
sinuses, oval to linear-obovate in general outline, the tips of the lobes with spines up to 8 mm long, leaf color
glaucous green with bluish markings along the veins, lower leaf surfaces sparingly prickly or smooth, upper sur-
faces smooth, the upper leaves clasping the stems; basal leaves oblanceolate, up to 25 cm long, 13 cm wide; stems
stout up to a meter tall, single or usually branched, brownish to glaucous blue-green, sparingly prickly (or smooth)
with reflexed, slender spines 3-4 mm long; taproot slender and tough with bright yellow latex. (2n = 28)
Infraspecific Variation: Some authors do not distinguish Argemone ochroleuca Sweet from A. mexicana as is done
here, following Ownbey (1958). Argemone ochroleuca has pale lemon colored petals, and while this is known from
some central American populations of A. mexicana, there are bud and style characters separating the species as
well. Although it has only been found in the Florida Keys, Africa and Brazil, the non-prickly forma leiocarpa
(Greene) Ownb. deserves mention.
Importance: Mexican Poppy is widely known as a poisonous plant and a purgative. Seeds have poisoned human
beings when contaminating home-grown cereal grains, and numerous animal and bird poisonings are ascribed to
the species. Symptoms are vomiting, diarrhea, induced glaucoma, edema, fainting and coma. The plants are
usually distasteful to livestock, but contaminated feed has been blamed for epidemic dropsy and glaucoma in India
where Argemone oil was transmitted to humans in milk. In Latin America small doses of Argemone extract have
been administered as medicine for ophthalmia, and glaucoma and other ailments with mixed results. Argemone
mexicana contains Protopine and Berberine type alkaloids, but lacks Pavine alkaloids found in such species as A.
platyceras and A. munita. The leaves and seeds contain Sanguinarine, Dihydrosanguinarine, Protopine and Ber-
berine. The oil of Argemone has been burned in lamps in Mexico and used in the manufacture of soap. Although
A. mexicana is a noxious weed throughout most of the tropics, it is still widely grown in gardens.
Fumariaceae (Fumitory Family)
The Fumariaceae: a family of about 15 genera and up to 400 species, distributed mostly in temperate Eurasia. It
has received various interpretations including frequent subordination under Papaveraceae as a subfamily (Fuma-
rioideae). Although the two families are clearly related, the Fumariaceae is the larger and more coherent aggrega-
tion, unified by highly specialized and complex structural similarities in the flowers. Members of Fumariaceae also
have watery juice rather than latex, and their inflorescences are racemose or cymose. Genera such as Corydalis
Fumaria and Dicentra are known for their alkaloid content and used medicinally. They may also be poisonous,
having been proven toxic to livestock. The family is of minor importance in the horticultural trade, and some of its
members are widespread weeds.
FAMILY DESCRIPTION
Herbaceous plants, sometimes vining, with watery sap, containing isoquinoline alkaloids. Plants are annual, bien-
nial or perennial, usually with branching stems from taproots, rhizomes, bulblets or tubers. The stems and leaves
are often glabrous and glaucous but may be pubescent. Leaves are alternate, usually pinnately dissected and lobed,
cauline and/or in basal rosettes, reduced upward on the stem. Flowers are bisexual, bilaterally symmetrical or
irregular, erect or nodding, borne in racemes or cymes. There are 2 bract-like sepals which are small and often
deciduous. Petals are dimorphic, in outer and inner series of 2 members each. Outer petals are larger, often
saccate or spurred. Inner petals are narrower, apically united into an expanded, terminal crest which is connate
over the stigma and anthers, enclosing them, especially in bud (inner petal tips rarely free). Stamens 6, in two sets
of 3, their united filaments often expanded at the bases, spurred or adnate to other tissue below, each set consist-
ing of a central stamen with 2 anther sacs and 2 outer stamens each with a single sac. The stigma is 2-lobed or
capitate with 2-several receptive surfaces, borne on a slender style. The single ovary is superior, 2-many carpelled, 1-
locular, becoming a 2-valved capsule in fruit (single-seeded nut in Fumaria). Dehiscence is basipetal, acropetal or
rarely by 1-seeded joints (indehiscent). Placentation is parietal; seeds are one to many, often lustrous and arillate, each
with a minute embryo and fleshy to liquid endosperm.
38
KEY TO GENERA
1. Plants delicately vining, climbing by petioles, the central axis conspicuously elongate; corolla, spongy-saccate
and turgid, persistent in fruit 2. Adlumia (p. 44)
1. Plants lax to erect; corolla deciduous or withering in fruit (2)
2. Outer petals alike, saccate or spurred; flowers pendulous 1. Dicentra (p. 39)
2. Outer petals unlike, only the upper one basally spurred; flowers + laterally disposed (3)
3. Fruit small, globose, 1-seeded, indehiscent; seeds without arils; flowers reddish and less than 9 mm long . . .
4. Fumaria (p. 51)
3. Fruit elongate, several-seeded, dehiscent; seeds arillate; flowers yellow or (if pinkish) more than 10 mm long
3. Corydalis (p. 45)
1. DICENTRA
Common Names: Bleeding Heart, Dutchman’s-breeches, Squirrel-corn
Authority: Bernhardi, Linnaea, vol. 8, p. 457 (468), 1833 ( nom.cons .)
A genus of some 15-20 species of perennial herbs (rarely vines) of North America and eastern Asia. Most of the species
are known to be cultivated, and may escape within and outside their native ranges. Their alkaloids are used in medi-
cine, and several species are known to be mildly poisonous.
Description: Plants with bisexual flowers; stigma 1, flattened, 2-lobed; style 1, slender, persistent in fruit; ovary 1,
cylindric, becoming a 2-valved capsule, dehiscing basipetally or acropetally; seeds few to many, arillate, somewhat
flattened, reniform to oblong-spheroid, often lustrous; stamens 6, their filaments partially fused into 2 columns of 3
stamens, each opposite the outer petals; petals 4, in two series; outer petals alike, both being either saccate or
spurred at base with ± reflexed apical limbs; inner petals also alike, narrow, forming a terminal crest and fused at
their tips, enclosing the stigma and anthers; sepals small, ovate to lanceolate, appressed (not peltate); inflores-
cence a terminal cyme or raceme, branched or simple with nodding or pendulous flowers (rarely axillary, single-
flowered or with erect flowers); pedicels usually bracteolate and bracteate; peduncle usually a naked scape; leaves
pinnately or temately dissected in a basal rosette (rarely cauline) with slender, glabrous petioles; plants usually
acaulescent from rhizomes, bulblets, tubers or taproots.
KEY TO SPECIES OF DICENTRA
1. Inflorescence a raceme-like thyrse of pink to purple flowers; apex of the flower with two tiny, free lobes (not
contiguous with the petal wings) 1. Dicentra eximia (p. 40)
1. Inflorescence a simple raceme of white to creamy flowers*; apex of the flower unlobed or lobed by the lateral
wings of the inner petals (2)
2. Bases of the outer petals prolonged into hom-like spurs, arched and narrowed toward their tips; tubers
pinkish, congested, with many small bulblets 2. Dicentra cucullaria (p. 41)
2. Bases of the outer petals cordate-saccate; tubers yellowish, with few small bulblets
3. Dicentra canadensis (p. 43)
* Orange and purple in a very rare form not known from NY.
39
1. Dicentra eximia (Ker.) Torr.
Common Names: Wild Bleeding-heart, Turkey-
corn, Staggerweed
Type Description: Ker., Bot. Reg., vol. 1, pi.
50, 1815
Synonyms: Fumaria eximia Ker., Corydalis for-
mosa Pursh, Bicuculla eximia (Ker.) Millsp.
( Bikukula ), Capnorchis eximia (Ker.) Planch,
(Diclytra)
Origin: Eastern North America
Habitats: Rocky woodlands, cliffs and borders
(also cultivated)
Habit: Acaulescent, perennial herbs
Flowering: April-September
Fruiting: May -October
General Distribution: (New York) New Jersey
and Pennsylvania to the Southern Appala-
chians of Tennessee and Georgia
Rarity Status: Endangered or possibly extirpated
in New York State
Description: Plants with bisexual flowers; stigma 1 per ovary, persistent, 2-erested, flattened, ca 0.8 mm wide;
style 1, slightly ribbed, slender, 9-14 mm long, persistent in fruit; ovary 1, 2-carpelled, superior, 3-4 mm long,
ovoid, becoming a brownish, fusiform, 2-valved capsule 1.5-2. 1 cm long (excluding style), 5-8 mm wide, exserted
from the withering corolla; seeds numerous, 1.2-1. 5 mm long, somewhat flattened, dull brown; aril nearly as large
as the seed, crested and convoluted, creamy-golden; stamens 6, in columns, 1.2-1. 5 cm long; anther sacs golden,
ca 0.8 mm long, oblong; filaments free in their terminal 1.5 mm, sheathing the style in a tight tube for 4-5 mm,
united into 2 arching sheaths in the lower 7-9 mm and attached opposing the outer petals; inner petals 2, pale
pink to rose, linear-spatulate, 1.5-1. 9 cm long, expanded into cucullate lobes 3-5 mm broad at the tips and
subterminally fused enclosing the stigma; terminal lobes of the inner petals 2, free, pink, flat, 1.7-2. 1 mm long,
up to 1 mm wide, attached at the point of inner-petal fusion; inner-petal wings prominent near the petal tips, up
to 1.5 mm broad, abruptly narrowed above, not confluent with the terminal lobes; outer petals 2, pink to deep
rose-purple, saccate and cordate at the base, narrowed upward to a divergent, expanded cucullate limb, the lower
portion 1.2- 1.8 cm long, ca 4 mm broad at base, the limb 4-5 mm long, 2-3 mm wide, acute tipped; sepals 2,
persistent, 1.8-2. 3 mm long and broad, oval, with ragged margins, base truncate to subcordate, tip acute, texture
somewhat membranous; inflorescence a thyrse of 2-8 (11) fascicles, 3-10 cm long, erect or arching, raceme-like at
first but the flower clusters continuing to develop through the season producing up to 10 flowers per node, the axis
occasionally branched to produce a smaller inflorescence subtending the terminal one; pedicels irregularly bran-
ching from peduncles in fascicles, their combined length up to 1.5 cm; bracts and bracteoles two to several, often
40
in subopposite pairs, lanceolate, acute to subulate, 2-6 mm long with pellucid, entire or irregularly toothed
margins; leaves pinnately and temately compound and lobed with elongate segments, their tips obtuse to aeute-
apiculate, the surfaces glaucous, glabrescent, blade area roughly triangular, up to 30 cm long and broad; petioles
12-26 cm long, ribbed, glabrous, subtended by scales which are irregularly toothed, up to 2 cm long; stem ribbed,
usually naked, scapose (if the inflorescence is branched, a compound bract, intermediate between bract and leaf
may be produced at the node); scape 25-40 (45) cm tall, arising from a tough, scaly rhizome and fibrous root
system. (2n = 16)
Infraspecific Variation: Plants vary considerably in size, erectness, leaf-lobing and inflorescence density. Because
this species has been hybridized with others in cultivation, garden plants may not comply in every respect with the
preceding description of robust native plants known from our area. Flower color is especially variable in cultivars,
ranging from pale pink to dark purple. Inflorescences of some southern plants are not uncommonly open-paniculate.
Importance: These showy plants have long been garden favorites and may escape cultivation within and outside
their native ranges. Plants contain Protopine and other alkaloids, and are poisonous.
2. Dicentra cucullaria (L.) Bernh.
Common Names: Dutchmans-breeches, Soldier’s-
cap, Breeches-flower, Little-boy’s-breeches,
Little-boys-and girls, White Harts, Butterfly-
banners, Little Indians
Type Description: Linnaeus, Species PL, p. 700,
1753
Synonyms: Fumaria cucullaria L., Cucullaria
bulbosa Rafi, C. cucullaria (L.) Farw., Bicu-
culla cucullaria (L.) Millsp. Bikukulla) Cap-
norchis cucullaria (L.) Planch., Dicentra occi-
dentals Rydb. (in part)
Origin: Temperate North America
Habitats: Rich woodlands and cliffs
Habit: Scapose, perennial herbs
Flowering: April-June
Fruiting: May-July
General Distribution: Nova Scotia to Ontario
and Kansas, south to the Piedmont of Georgia
and Alabama (disjunct westward to Idaho, Ore-
gon and Washington State)
41
Description: Plants with bisexual flowers; stigma 1 per ovary, minute, persistent, flattened and obscurely 2-lobed;
style 1, slender, persistent, 3-4 mm long in flower and fruit; ovary 3-5 mm long, linear-cylindric, many ovuled,
becoming a fusiform capsule ca 1 cm long, 4 mm wide, brown, glabrous, dehiscing basipitally along one or both
sutures, the 2 valves conforming somewhat in outline to the seeds within; seeds glossy, ebony, 1.7-1. 9 mm long,
ca 1.6 mm wide, shallowly reniform, tuffed-arillate; stamens 6; anther sacs pale yellow, ca 0.6 mm long; filaments
7-10 mm long, slender above, winged below and united around the ovary apex, arching in 2 loosely adherent
columns of 3; perianth withering, but somewhat persistent; inner petals 2, white to creamy, fused to the outer
petals for 5-7 mm below, with a prominent mid-vein, the limbs 4-6 mm long, ca 2 mm wide at their blunt tips,
cucullate, enclosing the stigma and fused at their tips, lateral wings inconspicuous, their tips not protruding
beyond the floral apex; outer petals 2, white to creamy (pinkish-tinged), saccate and conspicuously spurred (1.0)
1. 4-2.1 cm long, the spurs arched outward, narrowing toward the tips which curve upward from the drooping
flower, petal limbs 2. 5-3. 8 mm long, reflexed ± 90° from the axis, cucullate, obtusely tipped, creamy to ocher-
yellow; sepals 2, at the inner petal bases, whitish to greenish-cream, 2-3 mm long, 1-2 mm wide, ovate to ovate-
lanceolate with ragged-dentate (to entire) margins; pedicel (3) 5-7 (9) mm long, conspicuously broadened upward
from the middle at the point of attachment of a pair of subopposite, pellucid, pale bracteoles which are 2-3 mm
long, toothed and linear-lanceolate; bract at the pedicel base similar but broader and sometimes lobed; inflores-
cence a simple raceme, usually of 6-10 flowers, often terminating in an abortive pedicel; scape slender, glabrous,
10-20 (28) cm tall; leaves basal, finely ternately and pinnately compound and lobed, the lobes glabrous, linear to
oblanceolate-spatulate, up to 2.5 cm long and 3 (5) mm wide; petiolules slender, sometimes slightly winged,
glabrous, the lower ones with slightly clasping bases; petiole 4-15 cm long, glabrous, often ribbed below, sub-
tended (along with scapes) by pellucid to scarious, pale bracts and stipules which vary from minute to 1.5 cm long
and 1.2 cm wide, ovate with irregularly dentate margins and obtuse to acute tips; scapes and leaves arising from
lenticular tubers 8-15 cm in diameter, pinkish and usually clustered with numerous smaller bulblets, tiny bracts
and tough, fibrous roots, (2n = 32)
Infraspecific Variation: Plants of the western United States tend to have broader leaf segments (3-5 mm) and have
been given varietal status by some authors [var. occidentalis (Rydb.) M. Peck], The very rare forma purpuritincta
E. H. Eames occurs on the Richelieu River near Montreal P.Q., and has purple sepals and pink and orange petals.
Importance: These plants are cultivated, being spread by tubers to shady garden spots. The species should be
considered toxic due to the presence of isoquinoline alkaloids such as Protopine, Aporphine and Protoberberidine.
There have not been reported cases of human poisoning in the U. S., but livestock losses have been known,
especially in Virginia, where cattle were allowed to graze woodlands. Experiments have shown that livestock may
ingest up to 4% of their body weight of Dicentra before showing trembling symptoms. Dried tubers of
Dutchman s-breeches have been extracted for a weak tea, said to be alterative (health-restoring), but this could be
a detrimental or even dangerous practice.
42
3. Dicentra canadensis (Goldie) Walp.
Common Names: Squirrel-corn, Turkey-corn,
Turkey-pea, Colic-weed, “Wild Hyacinth”
(from its odor)
Type Description: Goldie, Edinb. Phil. Jour.,
vol. 6, p. 329, 1822
Synonyms: Corydalis canadensis Goldie, Cap-
norchis canadensis (Goldie) Kuntze, Bicuculla
canadensis (Goldie) Millsp. (Bikukulla, Dicly-
tra)
Origin: Temperate North America
Habitats: Rich woodlands and cliffs
Habit: Scapose, perennial herbs
Flowering: April-May
Fruiting: May -June
General Distribution: Southwestern Quebec to
Minnesota and Missouri, south to North Caro-
lina and Tennessee in the Appalachians
Description: Plants with bisexual flowers; stigma 1 per ovary, 2-lobed, flattened, ca 1 mm broad, persistent; style
1, slender, 4-6 mm long in flower and fruit, persistent; ovary 1, slenderly cylindric, 5-7 mm long, several ovuled,
glabrous, becoming a smooth, olive-brown, 2-valved capsule, fusiform to slightly falcate, 1.4-1. 7 cm long, 4-6 mm
wide, striated by its veins and 2 sutures; seeds usually 5-10 per capsule, very glossy, ebony, 2. 2-2.6 mm long, ca
2 mm deep and 1.5 mm wide; aril pale, wing-like, tufted; stamens 6 in 2 loose groups of 3; anther sacs pale
yellow, ca 0.6 mm long; filaments 8-10 mm long with wings up to 1 mm broad toward the base; inner petals 2,
white to creamy, 1. 1-1.3 cm long, 1-1.5 mm wide below, not adherent to the outer petals, strongly winged,
cucullate toward tips and expanded up to 3 mm in width where they fuse at the tips to enclose the stigma; inner
petal wings ca 0.5 mm wide below to 2.5 mm broad near their tips which extrude beyond the floral apex as
rounded lobes; outer petals 2, white to creamy, 1.0-1. 6 cm long, saccate-cordate, making the flower 6-11 mm
wide, the bases rounded to somewhat elongate, but not conspicuously spurred, the outer petal limbs reflexed ±
90° from the axis, 4-6 mm long, cucullate, obtusely tipped, often pinkish or purplish-tinged; sepals 2, minute,
membranous, deltoid to ovate sometimes with an irregular lobe; pedicels 4-8 (11) mm long, broadening upward
from near the middle at the point of attachment of 2 subopposite, linear-lanceolate bracteoles which are 2-3 mm
long, pale, membranaceous with irregular margins; bract at base of pedicel oblong to spatulate, sheathing at base,
up to 4 mm long, 2 mm wide, pale, greenish, membranaceous with pink veins; inflorescence a simple raceme,
usually of 3-8 flowers, often ending in a sterile lobe or pedicel; scape glabrous to puberulent, often ribbed below,
6-22 (26) cm tall; leaves basal, finely temate and pinnatisect, the lobes glabrous, linear to oblanceolate, up to 2.5
43
cm long, 5 mm wide; petiolules narrowly winged, not clasping at bases; petioles 4-18 (2) cm long, glabrous,
caniculate, subtended (along with the scapes) by pellucid to scarious stipules and bracts up to 1.4 cm long, 1.1 cm
wide, ovate with irregular margins and obtuse to apiculate tips; scapes and leaves arising from spheroid to ovoid (or
flattened) tubers 6-12 mm in diameter, dull yellowish-brown to golden in color, borne on slender, wiry rhizomes
with very few associated bulblets and fibrous, tough roots. (2n = ca 64)
Importance: This species is sometimes cultivated by transfer of the tubers to shaded garden plots. Wildlife, such as
game birds and squirrels eat the tubers. Squirrel-corn has been extracted for a tea used in folk medicine, reputed
to cure various ailments including colic; however, it should be used with extreme caution, since it is known to
contain harmful isoquinoline alkaloids. It has been implicated in livestock deaths and should be considered poi-
sonous in quantity.
2. ADLUMIA
Common Names: Allegheny Vine, Mountain Fringe
Authority: Rafinesque, Med. Rep. II, vol. 5, p. 352, 1808 (nom. cons.)
A genus of a single species of delicate vines with a widely disjunct distribution in Korea and the mountains of
eastern North America. Adlumia is sometimes cultivated.
1. Adlumia fungosa (Ait.) Greene ex B.S.P.
Common Names: Allegheny Vine, Mountain
Fringe, Climbing Fumitory, Canary Vine, Cy-
press Vine, Fairy Creeper, Allegheny Fringe,
Wood Fringe
Type Description: Aiton, Hort. Kew., vol. 3, p.
1, 1789
Synonyms: Fumaria fungosa Ait., F. recta
Michx., Adlumia eirrhosa Raf. (not Raf. ex
DC.), A. asiatica Ohwi, A. komarovii Popov
Origin: Arctotertiary Forest
Habitats: Rich woods, usually montane, shaded
cliffs and moist, calcareous places
Habit: Slender vines twining by the petioles on
plants or rocks, biennial (or perennial in Asia?)
Flowering: June-October
Fruiting: Late June-October
General Distribution: Korea and North America
from New England, Quebec and Ontario to
the southern Appalachians of North Carolina
and Tennessee
44
Description: Plants with bisexual flowers; stigma 2-lobed, wedge shaped, enclosed by the tips of the inner petals
and capped by a waxy pollen mass; style 1, 3-4 mm long, slender; ovary 1, slender, cylindric, 0.9-1. 3 cm long,
glabrous, pale green, becoming a tan, 2-valved capsule which enlarges only to accomodate the breadth of develop-
ing seeds, dehiscent from the base along 2 prominent sutures; seeds usually 5 per capsule, shiny black, very
smooth, ca 1.5 mm long, 1 mm broad, oval with a small indentation; aril absent; stamens 6, in 2 columns opposite
the outer petals; anther sacs 8, the central stamen of each column with 2 sacs, the 2 flanking stamens with 1 sac
each; each staminal column of 3 filaments, free near the anthers, fused below into a tubular sheath which is adnate
to the lower corolla in a zone of spongy tissue; flower persistent in fruit, not readily wilting, retaining its color and
shape until necrosis sets in; petals dimorphic, in 2 pairs of 2, united for most of their lengths into a spongy, saccate
corolla which is white, pink or purple-tinged, narrowly flattened-ovoid (purse-shaped) and bilateral, 14-18 mm
long, 4-7 mm wide and 2-4 mm across, obtuse to subcordate at base; inner petals 2, opposite the outer ones, very
narrow, fused to the outer petals below, free above, their tips grooved, keeled and clawed, expanded to form a
crown-like hood which caps the stigma and entraps pollen beneath the fused, inner petal tips; outer petals 2,
forming most of the corolla sac, free at their divergent tips, the free limbs acute, strongly cucullate, yellowish, 2-3
mm long, enclosing the crest of the inner petals in bud; sepals 2, early deciduous, enfolding the young, clavate
bud, but dropping as it expands, (sepals) 2. 0-2. 5 mm long, 0.8-1. 1 mm wide, peltate, being minutely stalked well
above the blade base, lanceolate, acuminate tipped, the peltate base rounded to hastate, margins irregularly
toothed, pellucid, reddish tinged; inflorescences few to many, 1 or 2 in the axil of a leaf, much-branched cymose-
paniculate with up to 30 flowers each; pedicels (2) 5-14 mm long, ribbed and swollen near receptacle base,
glabrous or minutely scaly with one or more minute linear-lanceolate, pellucid bracts which vary in size; peduncles
smooth, usually glabrous, 1-4 cm long; cauline leaves compound, pinnately much-divided, the leaflet-bearing
portion (blade) mostly 5-10 cm long, 4-7 cm wide, leaflets minute when blade expands, becoming 5-25 (30) mm
long, 5-20 (23) mm wide, variously lobed and incised with obtuse to apiculate tips, bases obtuse to attenuate,
surfaces glabrous; basal leaves similar, but forming a stemless, first year rosette; petiolules slender, glabrous to
puberulent, terete or flattened and ridged (cauline ones) sometimes twining; petioles up to 15 cm long, usually
glabrous, terete to ribbed or winged, tortuously twisted where twining on other stalks (those of the basal rosette
not twining, flattened, ribbed and somewhat sheathing at bases); stem developing the second year, a single,
elongate axis climbing or trailing up to 5 meters from a slender, biennial taproot and fibrous root system.
(2n 1 32)
Importance: These vines are cultivated and may persist or escape.
3. CORYDALIS
Common Name: Corydalis
Authority: Ventenat (not Medic.), Choix de Pi., t. 19, 1803 (nom. cons.)
A genus of over 200 species with its center of diversity in Asia. Corydalis is distributed mostly in the boreal
Northern Hemisphere, but also has species in South Africa. Corydalis species are sometimes grown as garden
curiosities, and are rich in alkaloids, some of which are poisonous.
Description: Plants with bisexual flowers; stigma 1 per flower, apically flattened, often 2-lobed with 4-8 smaller,
receptive lobes; style 1, linear, persistent (rarely articulated and deciduous); ovary 1 per flower, cylindrie-falcate,
becoming a cylindric (to somewhat inflated) 2-valved capsule, often falcate-torulose, acropetally dehiscent and
transversely ribbed between the seeds; seeds orbicular to reniform, sometimes flattened, usually lustrous and
arillate; stamens in 2 groups of 3, the upper cluster with a nectariferous spur adnate within the upper petal spur;
inner petals 2, similar, apically fused; outer petals 2, unlike, the upper one saccate-spurred; sepals ovate to
lanceolate, small and sometimes peltately attached; flowers ± horizontally oriented, bilateral, borne in terminal
racemes or panicles which elongate with age; pedicels and peduncles varying in length, the larger ones with
leaflet-like bracts at the base; cauline leaves alternate, reduced upward, otherwise like basal ones; basal leaves
often in a rosette, pinnately dissected into many lobes, these and the lower cauline leaves with petioles; stems
semi-succulent, hollow, spreading or erect from annual or biennial root systems (less commonly perennial rhizomes
or tubers).
45
KEY TO SPECIES OF CORYDALIS
1. Flowers bicolored, pink or rose-purple and yellow; mature fruit mostly ascending, linear; stigma not 2-lobed; stems
usually erect 1. Corydalis sempervirens (p. 46)
1. Flowers yellow; mature fruit mostly spreading or drooping; stigma 2-lobed; stems lax or weakly ascending (2)
2. Mature flowers 7-9 mm long; upper petal with an undulate to toothed crest toward its tip
2. Corydalis flavula (p. 48)
2. Mature flowers 10-20 mm long; upper petal without a conspicuous, lobed crest (3)
3. Sepals lance-acuminate, usually entire, not lacerate; leaflets with lobes mostly 3-6 mm broad, conspicuously
glaucous beneath 3. Corydalis lutea (p. 49)
3. Sepals ovate to lanceolate, lobed and lacerate; leaflets finely divided into lobes, usually less than 2 mm broad
4. Corydalis aurea (p. 50)
1. Corydalis sempervirens (L.) Pers.
Common Names: Pink Corydalis, Rock Harle-
quin, Rock Fumitory, “Pale Corydalis
Type Description: Linnaeus, Species Pi., p. 700,
1753
Synonyms: Fumaria sempervirens L., Capnodes
sempervirens (L.) Borkh., Fumaria glauca
Curt., Capnodes glauca (Curt.) Moench, Cor-
ydalis glauca (Curt.) Pursh, Corydalis rosea
Eat. (Neckeria)
Origin: North America
Habitats: Rocky, sunny places or partial shade,
disturbed sites, limestone flatrock, clearings,
often in shallow soil
Habit: Slender, erect biennials (annuals)
Flowering: May-September
Fruiting: May -October
General Distribution: Newfoundland to Alaska,
south to British Columbia, Minnesota, Illinois
and Georgia (also reported from Siberia)
46
Description: Plants with bisexual flowers; stigma 1 per flower, flattened, 4-lobed, persistent in fruit becoming up
to 0.8 mm broad; style 1, broadest near the stigma, persistent in fruit, becoming 3-4 mm long; ovary 1 per flower,
linear-cylindric, ca 1 cm long, becoming a slender, 2-valved capsule, usually erect or ascending, somewhat falcate
near the tip, tan, leathery in texture, prominently veined, 2. 5-4. 5 (5) cm long, 2-3 mm wide, dehiscing acrope-
tally along 2 sutures, shedding about 20 seeds; seeds 0.8-1. 1 mm long, variously compressed (with 1 or more flat
sides) to spheroid, black, shiny, finely reticulate, with pale arils ca 0.5 mm long; stamens 6; anther sacs golden,
minute; filaments slender, pale 8-10 mm long, coherent in phalanges of 3 each, the upper phalange with a
blunt spur ca 1 mm long adherent within the upper petal spur; perianth bilaterally symmetrical, pink and yellow;
petals 4, free, but closely associated to give the flower a tubular appearance; inner petals 2, borne laterally, 9-12
mm long, the basal 5-8 mm attenuated into a narrow claw, the expanded limbs attached at their tips over the
stigma, each limb with a fold below and an obtuse lobe on its upwardly directed margin; outer petals 2, dissimilar:
upper petal 10-15 mm long, including a saccate basal spur 3-4 mm long, the spur blunt, pink, not recurved
toward the pedicel, the petal hood narrow, only slightly crested or uncrested, the tip flexed upward at maturity
with upfolded lateral margins in the terminal 2 mm of the corolla tip which is bright yellow to cream; lower petal
10-13 rnm long, unspurred; sepals 2, ovate-acute with unevenly toothed margins, attached peltately, 2-3 mm long,
somewhat translucent, pinkish-green; pedicels slender, erect, 5-20 mm long in fruit, the longer ones in the lower
inflorescence; inflorescences 1 to several, terminal on the erect branches, racemes or panicles; bracts lanceolate,
pale green 1-2 mm long, the lower ones sometimes lobed; upper cauline leaves leaflet-like, intergrading with
inflorescence bracts, sessile; lower cauline leaves like basal leaves with petioles up to 3 cm long; basal leaves
p innately dissected the larger with 5 major lobes, further dissected into terminal lobes which are oblong-elliptic,
obtuse-apiculate, ± glaucous due to surface texture, glabrous; basal petioles (1) 3-9 cm long; stems usually single,
divaricately branched above (may be dwarfed and unbranched in poor conditions), terete to irregular in x-section
due to ribbing, erect, 30-80 cm tall from a twisted, biennial taproot with thick lateral branches. (2n = 16)
Infraspecific Variation: This is a remarkably uniform, widespread species, varying phenotypically in its stature in
response to environmental stresses, and having a deep, rose color-phase prized by horticulturists.
Importance: These plants are sometimes grown in rocky gardens or along gravelly paths. Related species are
suspect in poisoning cases due to their alkaloid content, but this species has not been implicated.
47
2. Corydalis flavula (Raf. ex Desv.) DC.
Common Names: Yellow Harlequin, Yellow Fu-
mewort, Yellow Rock-harlequin, Yellow Cory-
dalis, Colic-weed, “Pale Corydalis
Type Description: Rafinesque, in Desv., Jour.
Bot., vol. 1, p. 224, 1808
Synonyms: Fumaria flavula Raf. ex Desv., Cap-
nodes flavulum (Raf. ex Desv.) Kuntze (not
Capnoides, as in House, 1924); Neckeria
flavula (Raf ex Desv.) Millsp., Corydalis gey-
eri Fedde, C. aurea a flavula (Raf. ex Desv.)
Wood, “C. flavidida ”
Origin: North America
Habitats: Moist, loose soil in rocky woodlands of
sandy to calcareous areas
Habit: Sprawling to ascending colonies of annuals
Flowering: April-May
Fruiting: April-June (July)
General Distribution: Sporadic, from New York
and Connecticut to Nebraska, south to Loui-
siana and North Carolina
Rarity Status: These plants are rare and threat-
ened in New York State and the species ap-
pears on the state list (Mitchell & Sheviak,
1981)
Description: Plants with bisexual flowers; stigma 1 per flower, flattened, the 2 major lobes widely divergent, ca 0.3
mm wide, usually deciduous in fruit; style 1, broadest near the base, 1-2 mm long, persistent in fruit, becoming
2-3 mm long; ovary 1 per flower, ca 4 mm long, 1 mm wide, falcate, becoming a cylindric, 2-valved capsule,
spreading or pendulous, (14) 16-20 (23) mm long, 2-3 mm wide, glabrous, tan, leathery, the walls contracted
between the enlarged seeds to give a beaded appearance to the capsule which is dehiscent along 2 sutures (acrope-
tally) to release about 6-12 seeds; seed lenticular with a distinct marginal ridge, black, glossy, minutely alveolate,
particularly near the margin, 1. 8-2.0 mm broad, ca 1 mm thick; aril ca 1 mm long, creamy, wing-like, wrinkled;
stamens 6; anther sacs golden, minute; filaments 5-6 mm long, slender, creamy, united into 2 phalanges, the
upper with a spur 0.5 mm long within the upper petal spur; perianth bilaterally symmetrical, creamy yellow
(sometimes small, cleistogamous); petals 4, free but closely associated to give the corolla a tubular appearance;
inner petals 2, borne laterally, 5-6 (7) mm long, clawed at bases, expanded at tips where they are united over the
stigma; outer petals 2, dissimilar: upper petal 7-9 mm long including the saccate, basal spur which is 1-1.5 (2)
mm long, its blunt tip curved under toward the pedicel; petal hood with upcurved, slightly undulate to toothed
margins near the tip, bearing an abrupt crest along its summit, the crest margin undulate to sharply toothed; lower
48
petal 5-8 mm long, unspurred, uncrested or with a small undulate crest; sepals minute, often deciduous, scarious,
lanceolate, sometimes toothed, ca 0.3 mm long; pedicels, 3-11 (16) mm long, slender, glabrous, usually reflexed in
fruit, each subtended by a bract; bracts ovate to lanceolate or lobed, 5-12 mm long, 3-9 mm broad, entire or
dentate at margins and with acute to acuminate tips, somewhat sheathing at bases; inflorescence a short, crowded
terminal or subterminal raceme; cauline leaves similar to basal ones, smaller but not reduced greatly upward, short
petioled to sub-sessile; basal leaves pinnately compound with 5-7 main lobes, these further dissected and lobed,
the ultimate leaf lobes elliptic to linear, apiculate, the surfaces glabrous, glaucous, valuable in size but mostly 4-8
mm long; petiolules and petioles slightly winged, glabrous; basal petioles up to 10 cm long; stems erect when
young, soon trailing and ascending, ribbed and fluted, juicy, branched mostly from near the base, 15-30 ( — 40) cm
long, from an annual taproot. (2n = 16)
Infraspeeific Variation: Cleistogamous plants are known in which the flowers are small and poorly developed; the
vegetative portion of such plants is often very lax and diffusely branched with larger terminal leaflets and lobes.
Importance: This species contains a number of alkaloids and is suspected in livestock poisoning in Virginia.
3. Corydalis lutea (L.) DC.
Common Names: Yellow Corydalis, Rock Fumi-
tory
Type Description: Linnaeus, Mant. Pi., ii, p.
258, 1771
Synonym: Fumaria lutea L.
Origin: Southern Europe
Habitats: Rocky places, especially walls, as an es-
cape (calciphile)
Habit: Sprawling perennials
Flowering: May-August
Fruiting: June-September
General Distribution: A rare escape in North
America, but widely naturalized in Europe
where it is also native in part
Description: Plants with bisexual flowers; stigma 1 per ovary, its 2 major lobes laterally divergent, ca 0.8 mm
wide, somewhat flattened, usually not persistent in fruit; style 2-3 mm long, cylindric, persistent; ovary 1 per
flower, slender, cylindric, 5-9 mm long, becoming a 2-valved capsule, usually pendulous, plump-cylindric, 8-12
mm long, 2-3 mm wide, somewhat constricted between seeds, tan, acropetally dehiscent along 2 sutures to release
the 3-7 seeds; seeds lenticular, black, shiny, somewhat alveolate, ca 2 mm broad, with a definite marginal ridge;
49
aril pale, wing-like; stamens 6; anther sacs golden, minute; filaments pale, slender, united in 2 phalanges, the
upper phalange with a spur 1.5-2 mm long within the upper petal spur; perianth bilaterally symmetrical, pale
yellow to dark yellow near the corolla tip; petals 4, free, but closely associated to give the corolla a tubular
appearance; inner petals 2, borne laterally, 9-13 mm long, clawed at bases, expanded into broader lobes toward
the apices where they are fused at the tips over the stigma; outer petals 2, dissimilar: upper petal 12-20 mm long
including a saccate basal spur 2.5-3 mm long, the spur narrowed toward its tip which is recurved toward the
pedicel, upper petal hood uncrested (or very slightly crested), its lateral margins entire to weakly undulate, up-
curved toward their tips; lower petal 7-10 mm long, unspurred, uncrested; sepals 2, pale, yellow-green, persist-
ent, 2-3 mm long, lanceolate with truncate bases and acute to acuminate tips, the margins entire or irregularly
serrate; pedicels slender, elongating up to 1.5 cm in fruit, reflexing with age; inflorescence a few-flowered raceme;
bracts like the sepals; upper cauline leaves leaflet-like, short petioled; lower cauline leaves much like the basal
ones, but reduced upward; basal leaves biternately to triternately compound, the segments further divided and
lobed; leaflets elliptic, lobed or unlobed, mostly 4-9 mm long, 3-7 mm wide, glabrous, green above, whitish-
glaucous below, apiculate tipped; petiolules and petioles not conspicuously winged; basal petioles up to 12 cm
long; stems lax, freely branching, ribbed, up to 40 cm long from a perennial base and fibrous roots. (2n = ±54?)
Importance: These plants are grown commonly as garden ornamentals, especially on walls and sunny slopes, and
they infrequently escape cultivation. They contain a variety of alkaloids and may be considered poisonous.
4. Corydalis aurea Willd.
Common Name: Golden Corydalis
Type Description: Willdenow, Enum., p. 740,
1806
Synonyms: Capnodes aureum (Willd.) Kuntze,
Neckeria aurea (Willd.) Millsp. (see Ownbey,
1947 for extensive, non-NY synonymy)
Origin: Western North America
Habitats: Rocky places, especially limestone, in
coniferous or mixed woodlands and clearings
Habit: Sprawling to ascending annuals or bien-
nials
Flowering: May-July
Fruiting: May-August
General Distribution: Alaska to Mexico and the
Rocky Mountains, scattered eastward in gla-
ciated areas to southeastern Quebec
Rarity Status: This species is sporadic and rare
in eastern North America, and appears on the
New York State rare plant list (Mitchell &
Sheviak, ibid.)
50
Description: Plants with bisexual flowers; stigma 1 per flower, its 2 major lobes laterally divergent, persistent in
fruit, up to 1 mm broad; style 1, slender, broadest toward base, persistent, becoming 3-4 mm long in fruit; ovary
1 per flower, 5-6 mm long, cylindric, slightly falcate, becoming a cylindric, often arcuate, 2-valved capsule,
pendulous at maturity (in ours), 1. 5-3.0 cm long, 2.5-3 mm wide, constricted between the seeds, leathery, tan,
usually with 2 prominent veins opposite the sutures, dehiscing acropetally to release (3) 7-12 seeds; seeds 1. 9-2.0
mm long, plump-lenticular to comma-shaped, very shiny black, minutely and inconspicuously reticulate, the
marginal ridge absent; aril wing-like, pale, ca 1.5 mm long; stamens 6; anther sacs golden, minute; filaments
slender, 7-8 mm long, coherent into 2 phalanges, the upper phalange prolonged backward as a spur 2-3 mm long
within the upper petal spur; perianth bilaterally symmetrical, yellow with darker yellow near the corolla tip; petals
4, free, but closely associated in a corolla which appears tubular; inner petals 2, borne laterally, narrowed in their
distal half into claws which bear small wings, the expanded blades attached over the stigma at their tips, total
length 8-10 mm; outer petals 2, dissimilar: upper petal 12-16 mm long including a saccate, basal spur 3-5 mm
long, the spur straight or slightly downcurved, but not strongly recurved toward the pedicel, petal hood with a
greenish swelling along its median, but not crested (extremely rarely with a small crest), lateral margins upturned,
entire to undulate or sharply toothed, with an apiculate tip; lower petal 9-11 mm long, unspurred uncrested, the
tip similar to the hood of the upper petal; sepals 2, scarious, ovate, 1-2 mm long, ca 1.5 mm wide, acute to
acuminate-tipped, truncate to cordate-hastate at bases, the margins irregularly toothed or incised; pedicels slender,
elongating and reflexing downward in fruit, up to 10 mm long; inflorescence a terminal or axillary raceme, usually
surpassed by the leaves; cauline leaves not much reduced upward, petioled, much like basal leaves; basal leaves
few to densely clustered in a rosette, pinnately compound into 5-7 major lobes, further dissected, the ultimate
leaflets usually deeply cleft into narrow segments, broadly to narrowly elliptic in outline, the surfaces glabrous, ±
glaucous, tips apiculate; petiolules and petioles glabrous, terete to slightly winged, somewhat clasping at bases, the
petioles 2-8 cm long, not much reduced upward; stems erect when young, soon becoming prostrate to ascending,
semisucculent, glabrous, 15-35 (50) cm long, arising from an annual root system or developing a biennial, basal
caudex with fibrous roots. (2n - 16)
Infraspecific Variation: Most of the variability of this species lies outside of New York State in the contiguous part
of its distribution range. Our materials correspond to ssp. aurea, while plants of the south-central states are treated
as ssp. occidentalis (Engelm.) Ownb., with larger, long-spurred flowers and more erect fruits.
Importance: These plants are sometimes grown in rock gardens. Due to their alkaloid content, they have been
cited in some cases of livestock poisoning.
4. FUMARIA
Common Names: Fumitory, Earth-smoke
Authority: Linnaeus, Species Pi., p. 699, 1753
A genus of over 40 species, many of which are cosmopolitan weeds, native to Eurasia and north Africa, mostly
around the Mediterranean area. In New York State the only established species is the infrequently introduced
Fumaria officinalis L., an adventive associated with agriculture. Some species of the genus contain medicinally
active alkaloids.
51
1. Fumaria officinalis L.
Common Names: Common Fumitory, Fume-
wort, Earth-smoke, Hedge Fumitory, Fume-
root, Wax-dolls
Type Description: Linnaeus, Species Pi,, p. 700,
1753
Origin: Mediterranean region
Habitats: Waste places, agricultural fields, bor-
ders, roadsides
Habit: Erect to spreading and sprawling annuals
Flowering: May-August
Fruiting: June-September
General Distribution: A widely escaped weed,
especially in warm-temperate North America;
native to southern Eurasia and north Africa
Description: Plants with bisexual flowers; stigma 1 per flower, its 2 lobes laterally divergent, flattened, up to 0.5
mm wide; style 1, slender, 5-6 mm long, articulate at base and deciduous; ovary 1 per flower, superior, fusiform,
1.2- 1.4 mm long, glabrous, becoming a single-seeded indehiscent fruit (nutlet), globose to lenticular-reniform,
usually with 2 indentations near the stigma scar, ca 3 mm wide 2.6 mm thick at maturity, greenish-tan to brown,
the surfaces rough, minutely muricate; seed 1 per fruit, ca 2 mm long, 1 mm wide, brown, ridged; aril absent;
stamens 6; anther sacs minute, golden; filaments united into 2 distinct phalanges (upper and lower), slender
above, expanded and sheathing toward the base, pale, 4-6 mm long; corolla bilaterally symmetrical; petals 4, pale
rose pink to deep purple-maroon near their tips, basally free, but closely associated to give the corolla a tubular
appearance; inner petals 2, similar, 3. 5-4. 5 mm long, borne laterally, clawed toward their bases, expanded and
ridged near their tips where they are fused-cucullate over the stigma; outer petals 2, dissimilar: upper petal 12-13
mm long including a saccate basal spur, 2.0-2. 8 mm long, straight, but with a small downcurved lobe below, petal
hood with a greenish, swollen area along the midline, laterally cucullate and blunt-tipped; lower petal 5-6 mm
long, narrowly clawed at base, the terminal 2 mm expanded, cucullate, green and maroon; sepals (1) 2-3 mm long,
whitish to translucent with green midribs and pink-tinged margins, peltate to truncate or hastate at bases, acute to
acuminate tipped with ragged, toothed margins; pedicels 2-4 mm long, sometimes flattened or grooved, glabrous;
inflorescence a narrow raceme, terminal or more often lateral or sub-terminal; bracts lanceolate, 1.5-3 mm long,
green with whitish (usually entire) margins, somewhat clasping at bases of the pedicels; cauline leaves up to 12 cm
52
long, variously compound, pinnately dissected and lobed, the ultimate segments mostly with 2-3 mm wide lobes,
oblanceolate, blunt or apiculate-tipped, glabrous; basal leaves similar though often smaller, in a rosette, few in
number or absent; petiolules and petioles glabrous, rarely entwining other low vegetation; lower petioles up to 5
cm long; stems ribbed, somewhat glaucous, ascending to sprawling up to 50 cm from a slender taproot and diffuse
annual root system which exudes a nitrous, smoky odor. (2n = 32, 14?, 16?)
Infraspeeific Variation: This species, like a number of its relatives, can produce small, pale cleistogamous flowers
which do not conform to the above description.
Importance: Fumitory can be an annoying weed of crop fields and gardens, especially further south. It has been
extracted for a yellow-green dye in the Mediterranean region. In folk medicine Fumaria officinalis has been
employed as a laxative and ’blood purifier’’ with questionable success; it does contain Protopine (Fumarine), how-
ever, which has been shown experimentally to induce bradycardia.
APPENDIX I
FUNGI ASSOCIATED WITH PLANT SPECIES IN THIS TREATMENT
To be included on this list, a fungus must occur on a species in this treatment somewhere in the United
States. If a fungus occurs in New York State and has not as yet been recorded on a host covered in this treatment,
but has been collected on such a host in some other state, it is marked with a single asterisk (*).
Abbreviations of states indicate a literature citation for each. Double asterisks (**) indicate that a herbarium
specimen with New York State host information has been seen.
PEHONOS POll ALPS
Peronospora arborescens (Berk.) de Bary, Downy mildew or false mildew on Argemone mexicana (Tex.) this nearly
cosmopolitan fungus has also been recorded on numerous Papaver sp. and Chelidonium majus on other conti-
nents
Peronospora corydalis de Bary, Downy mildew or false mildew on Corydalis aurea (Ind.), or Corydalis semper-
virens (Mass., Md., N.Y. **), on Dicentra canadensis (N.Y. **)
Peronospora dicentrae Syd. ex Gaum., Downy mildew or false mildew on Dicentra canadensis (N.Y. **), on Dicen-
tra cucullaria (Ind., Md., Mich., Mo., N.Y., Va., Wise.)
Pythium paroecandrum Drechs., Leaf and stem rot of Sanguinaria canadensis (Va.)
ERYSIPHALES
Erysiphe cichoracearum DC., Powdery mildew on various hosts including Papaver somniferam, Papaver sp., Cheli-
donium majus, and Eschscholzia calif ornica (cosmopolitan *)
Erysiphe poly go ni DC., Powdery mildew on Eschscholzia californica (Calif.), on Papaver somniferum (Ore.)
Microsphera alni DC., Powdery mildew on Menispermum canadense (Iowa, Mich., N.Y., Ohio)
Phyllactinia corylea Pers., Powdery mildew on Berberis vulgaris (Mass., Vt.)
SPHAE MALES
Diaporthe beckhausii Nits. (= Diaporthe menispermi Deam. & House), on Menispermum canadense (N.Y.**)
Diaporthe detrusa (Fries) Fuckel, on Berberis vulgaris (Mass.)
Glomerella cingulata (Ston.) Spauld. & Schrenk, on Podophyllum peltatum (Del., Tex.)
Valsa menispermi Ellis & Holw., on Menispermum canadense (Iowa)
53
HELOTIALES
Botryotinia fuckeliana (de Bary) Whetzel [ = Peziza fuckeliana de Bary, Sclerotinia fuckeliana (de Bary) Fuckel],
Gray-mold blight, [perfect state of Botrytis cinerea Pers.: Fries (Monilales) which see for host information]
Discohainesia oenotherae (Cooke & Ellis) Nannf., on Podophyllum peltatum (Va.)
Orbilia caulophylli Ellis & Everh. [ = Calloria caulophylli (Ellis & Everh.) Rehm; this species belongs in the
genus Laetinaevia but apparently the combination has not been made], on Caulophyllum thalictroides (N.Y. **)
Sclerotinia sclerotiorum (Libert) de Bary, on Papaver somniferum (Wash.)
Septotinia podophyllina Whetzel ex Groves & Elliott [ = Gloesporium podophyllina Ellis & Everh., Septogloeum
podophyllinum (Ellis & Everh.) Sacc.], on Podophyllum peltatum (Del., Md., Mo., N.J., N.Y., Pa., Va., W.
Va.)
PLEOSPORALES
Cucurbitaria berberidis (Pers.) S. F. Gray, on Berberis vulgaris (Maine, N.Y.**, S.C.)
Leptosphaeria berberidis Richon, on Berberis vulgaris (Iowa)
Pleospora media Niessl., on Papaver nudicaule (Alaska)
DOTHIDEALES
Dothidea berberidis (Wahlenb.) de Not., on Berberis vulgaris (Mass.)
Mycospherella berberidis (Auersw.) Lindau, on Berberis thunbergii (N.Y.)
USTILAGINALES
Entyloma eschscholtzia Harkn., on Eschscholzia californica (Calif.)
Entyloma fuscum Sehrot (= Entyloma fuscellum Rabehn., Entyloma glaucii Dang.), Leaf smut on Papaver dubium
(La.), on Papaver nudicaule (Iowa, Maine), on Papaver orientale (Ont.), on Papaver rhoeas and Papaver somni-
ferum (cosmopolitan)
Entyloma menispermi Farlow & Trel. (= Ramularia contexta Ellis & Everh.), on Menispermum canadense (N.Y.**
to Va., west to Minn, and Kans.)
UREDINALES
Aecidium plenum Arthur, on Argemone mexicana (Tex.)
Cerotelium dicentrae (Trel.) Mains & F. W. Anderson, a rust with 0, I on Dicentra cucullaria (N.Y.** to Md., 111.,
Iowa, Kans., S. Dak.) II, III on Laportea canadensis
Puccinia aristidae Tracy, a rust with 0, I on Corydalis aurea and Corydalis sp. (Colo., Kans., Nebr.) II & III on
Aristida and Distichlis
Puccinia brandegei Peck, on Corydalis aurea (Kans.)
Puccinia graminis Pers., a rust of enormous commercial importance, 0, I on Berberis vulgaris ** and Berberis
canadensis ** (generally distributed) II, III on Cereals and grasses
Puccinia podophylli Schw., 0, I, III on Podophyllum peltatum (Generally distributed **)
APHYLLOPHORALES
Poria punctata (Fries) Karst., on Berberis vulgaris (Md.)
Thanatephorus cucumeris (Fries) Donk (= the perfect state of Rhizoctonia solani Kuhn, which see for host informa-
tion)
MONILIALES
Alternaria alternata (Fries) Keissler, on Papaver rhoeas * (the fungus is both cosmopolitan and common)
Alternaria lancipes Ellis & Everh., on Argemone mexicana (Tex.)
Alternaria sp., on Eschscholzia californica (Tex.)
Botrytis cinerea Pers. ex Fries, imperfect state of Botryotinia fuckeliana (de Bary) Whetzel, on Berberis vulgaris
(Calif.), on Eschscholzia californica (Alaska), on Papaver nudicaule (Alaska), on Papaver orientale (Md., Wise.),
on Papaver rhoeas (Alaska), on Podophyllum peltatum (N. J.)
54
Botrytis streptothrix (Cooke & Ellis) Sacc., on Caulophyllum thalictroides (N.J., N.Y. **), on Sanguinaria canaden-
sis**
Botrytis sp., on Sanguinaria canadensis (N. Y.)
Cercospora caulophylli Peck, on Caulophyllum thalictroides (Vt., N.Y.** to Va., Mo., Wise.)
Cercospora menispermi Ellis & Holw., on Menispermum canadense (N.Y.** to Va., Kans., Wise.)
Cercospora podophylli Tehon & Daniels, on Podophyllum peltatum (111.)
Cercospora sanguinariae Peck, on Sanguinaria canadensis (Md., Mo., N.Y.**, Pa., Tex., Wise.)
Cercospora whetzelii Chupp, on Argemone mexicana (Puerto Rico)
Heterosporium eschscholtziae Harkn., on Eschscholzia californica (Calif.)
Oidium sp., on Papaver somniferum (“U. S. A.”)
Phy mat otrichum omnivorum (Shear) Dug., on Argemone mexicana (Tex.), on Berberis canadensis (Tex.), on Ber-
beris thunbergii (Tex.), on Berberis vulgaris (Tex.), on Chelidonium majus (Tex.)
Verticillium albo-atrum Reinke & Berth, on Berberis thunbergii (Conn, to Va., 111., Mich.), on Berberis vulgaris
(Conn.), on Eschscholzia californica (Calif.), on Papaver orientale (N. Y.)
SPHAEROPSIDALES
Coniothyrium insitivum Sacc., on Berberis vulgaris (Nebr., Wise.)
Coniothyrium hellebori Cooke & Massee, on Heleborus niger (N. C.)
Diplodia berberidina Sacc., on Berberis vulgaris (N. Y.)
Diplodia sarmentorum Fries, on Menispermum canadense (N. Y.)
Phoma berberina Sacc. & Roum., on Berberis thunbergii (Mass.), on Berberis vulgaris (Iowa)
Phomopsis menispermi (Peck) Grove, on Mensipermum canadense (Kans.)
Phyllosticta abortiva Ellis & Kellerm., on Menispermum canadense (Iowa)
Phyllosticta berberidis Ranenh., on Berberis vulgaris (Ky)
Phyllosticta japonica Thiim, on Berberis thunbergii (Alaska), ( — Pconidial state of Mycosphaerella berberidis )
Phyllosticta menispermicola Tehon & Daniels, on Menispermum canadense (111.)
Phyllosticta podophylii (Curtis) Wint., on Podophyllum peltatum (N.Y.** to Ala., Ark., Wise.)
Phyllosticta podophyllina Tehon & Stout, on Podophyllum peltatum (111.)
Phyllosticta sanguinariae Wint., on Sanguinaria canadensis (Mo., Tex., W. Va.)
Septoria abortiva (Ellis & Kellerm.) Tehon & Daniels, on Menispermum canadense (111., Kans., Ky.)
Septoria argemones Tharp, on Argemone alba (Okla.), on Argemone mexicana (Okla.)
Septoria chelidonii Desm., on Argemone mexicana (Puerto Rico), on Chelidonium majus (Tex.)
Septoria corydalis Ellis & J. J. Davis, on Corydalis aurea (Wise.)
Septoria podophyllina Peck, on Podophyllum peltatum (N.Y.** to Miss., Mo., Wise.)
Sphaeropsis menispermi Peck ( — Pconidial state of Physalorpora obtusa ), on Menispermum canadense (N.Y.** to
Va., Kans., Mich.)
Stagonospora berberidina Sacc., on Berberis vulgaris (N. Y.)
MELANCONIALES
Collet otrichum circinans (Berk.) Arx, experimental infection on Eschscholzia californica (Calif.)
Collet otrichum sordidum J. J. Davis, on Menispermum canadense (Wise.)
Cylindrosporium circinans Wint., on Sanguinaria canadensis (Md., Mo., Wise.)
Gloeosporium argemonis Ellis & Everh., on Argemone mexicana (Fla., Tex.)
Gloeosporium berberidis Cooke, on Berberis vulgaris (Mass., Minn., Ohio, Wise.)
Vermicularia hysteriiformis Peck, on Caulophyllum thalictroides (N.Y.**)
Vermicularia podophylii Ellis & Deam., on Podophyllum peltatum (Tex., Va.)
MYCELIA STERILIA
Rhizoctonia solani Kiihn., on Berberis vulgaris (Conn.), on Papaver nudicaule (Maine, N.Y.), on Papaver orientale
(Ind., N. J.), on Papaver rhoeas (Kans., N. J.), on Papaver somniferum (Idaho)
Rhizoctonia sp., on Podophyllum peltatum (Mo.)
55
APPENDIX II
A List of Some Insects Associated with Plant Species in this Treatment:
HEMIPTERA
Miridae
Horcias dislocatus (Say) on Caulophyllum thalictroides
Aphididae
Aphis fabae Scopoli on Papaver orientale and Papaver rhoeas
Aphis gossypii Glover on Papaver rhoeas
Liosomaphis berberidas (Kaltenbach) on Berberis thunbergii
Macrosiphon euphorbiae (Thomas) on Corydalis sempervirens and Dicentra sp.
Rhopalosiphum berberidis (Kaltenbach) on Berberis thunbergii and Berberis vulgaris
Coccidae
Lecanium persicae (Fabricius) on Berberis
LEPIDOPTERA
Pieridae
Artogeia napi (L.) on Berberis vulgaris
Artogeia rapae (L.) Cabbage Butterfly on Berberis vulgaris
Lycaenidae
Satyrium liparops (Boisduval & LeConte) on Podophyllum peltatum
Nymphalidae
Speyeria aphrodite (Fabricius) on Podophyllum peltatum
Euptoieta claudia (Cramer) Variegated Fritillary on Podophylum peltatum
Hesperiidae
Achalarus lyciades Abbot & Smith on Corydalis sempervirens
Saturniidae
Hyalophora cecropia (L). Cecropia Moth on Berberis vulgaris
Sarnia cynthia (Drury) on Berberis vulgaris
Lymantriidae
Orgyia leucostigma (Abbot & Smith) White-marked Tussock Moth on Berberis canadensis and Berberis vulgaris
Lymantria dispar (L.) Gypsy Moth on Berberis vulgaris
Lasiocampidae
Malacosoma americana (Fabricius) Apple Tree Tent Caterpillar on Berberis vulgaris
Agaristidae
Alypia octomaculata (Fabricius) Eight-spotted Forester on Berberis vulgaris
57
Noctuidae
Trichoplusia ni (Hubner) on Eschscholzia californica
Crocigrapha normani (Grote) on Caulophyllum thalictroid.es
Pseudaletia unipuctata (Haworth) Army Worm on Papaver somniferum
Mamestra curialis Smith on Dicentra spectabilis
Papaipema cerina (Grote) on Podophyllinn peltatum
Papaipema mereiekata Bird on Podophyllum peltatum
Prodenia eridanian (Cramer) on Sanguinaria canadensis
Arctiidae
Hyphantria cunea (Drury) Fall Webworm on Berberis canadensis
Pyralidae
Omphalocera dentosa Grote on Berberis sp.
DIPTERA
Cecidomyiidae
Bremia podophyllae Felt on Podophyllum peltatum
Mycodiplasis alternata Felt on Podophyllum peltatum
Youngomyia podophyllae Felt on Podophyllum peltatum
HYMENOPTERA
Tenthredinidae
Bivena quattrodecimpunctata (Nort) larva on Podophyllum peltatum
Hylaeidae
Hylaeus mesillae Cressoni (Cockerell) on Eschscholzia californica
Adrenidae
Andrena carlini carlini Cockerell on Sanguinaria canadensis
Andrena forbesii Robertson on Berberis vulgaris
Andrena hippotes Robertson on Berberis sp.
Andrena miranda Smith on Argemone sp.
Andrena miserabilis Cresson on Berberis sp.
Halictidae
Augochlora pura pura (Say) on Caulophyllum thalictroides
Augochlorella striata (Provancher) on Caulophyllum thalictroides
Augochloropsis metalica metalica (Frabricius) on Caulophyllum thalictroides
Megachilidae
Osmia lignaria lignaria Say on Dicentra sp.
Synhalonia hamata (Bradley) on Podophyllum peltatum
Xylocopidae
Ceratina calcarata Robertson on Dicentra cucullaria and Sanguinaria canadensis
58
BIBLIOGRAPHY
Acheson, R. M., et al. 1956. Distribution of anthocyanin pigments in poppies. Nature 178: 1283-1284.
Ahrendt, L. W. A. 1941. A survey of the genus Berberis in Asia. Jour. Bot. 79: Suppl. 1-80.
1961. Berberis and Mahonia. a taxonomic revision. Jour. Linn. Soc., vol. 57, 410 pp.
Airy-Shaw, H. K. 1948. Jeffersonia dubia. Bot. Mag. 164: pi. 9681.
Alexander, I. 1952. Entwicklungsstudien an Bliiten von Cruciferen und Papaveraceen. Planta 41: 125-144.
Ames, L. M. 1937. Barberries immune or highly resistant to black stemrust of cereals. Am. Arb. Bull. Pop. Inf.
IV. 5:57-72.
Anderson, E. 1934. The genus Akebia. Arn. Arb. Bull. Pop. Inf. IV. 2: 17-20.
Andrews, A. C. 1952. The opium poppy as a food and spice in the classic period. Agr. Hist. 26: 152-155.
Arber, A. 1932. Studies in floral morphology III. On the Fumarioideae, with special reference to the androecium.
New Phytol. 30: 317-354.
1938. Studies in flower structure IV. On the gynoecium of Papaver and related genera. Ann. Bot. 2:
649-664.
Asahina, H. et al. 1957. Studies of poppies and opium. Bull. Narcot. 20: 25.
Bailey, I. W. & B. G. L. Swarny. 1951. The conduplicate carpel of dicotyledons and its initial trends of specializa-
tion. Amer. Jour. Bot. 38: 373-379.
Bailey, L. H. 1922. Standard Cyclopedia of Horticulture MacMillan Co., N.Y. 3639 pp.
Baillon, H. 1871. Monographic des Menispermacees et des Berberidac6es. Hist, des Plantes 3: 1-76.
Bandelin, F. J. & W. Malesh. 1956. Aklaloids of Chelidonium majus L., leaves and stems, I. Jour. Amer. Pharm.
Assoc. Sci Ed. 45: 702-704.
Baker, R. O. 1919. The seasonal development, gross structure, ecology and geographical distribution of Sanguinar-
ia canadensis. Nat. Stud. Rev. 15: 62-73.
Barnhart, J. H. 1920. Jeffersonia diphylla. Addisonia 5: 31-32.
Bartek, J. et al. 1955. Isolation of some components of resina podophylli (Podophyllum peltatum) and comments on
their structure. (In Czech.) Chem. Listz. 49: 1550-1560.
Barton, B. S. 1793. A botanical description of the Podophyllum diphyllum of Linnaeus . . . etc. Trans. Amer. Phil.
Soc. 3: 334-347.
Bastin, E. S. 1895. Some further observations on the structure of Sanguinaria canadensis. Amer. Jour. Pharm. 67:
4-9.
Bentham, G. 1861. Notes on Menispermaceae. Jour. Linn. Soc. Bot. 5: 45-52.
Bemhardi, J. J. 1833. Uber den Charakter und Verwandtschaft der Papaveraceen und Fumariaceen. Linnaea 8:
401-477.
1838. Nachtragliche Bemerkiingen uber Papaveraceen und Fumariaceen Linnaea 13: 651-658.
Bersillon, G. 1955. Recherches sur les Papaveracees, . . . etc. Ann. Sci. Nat. Bot. XI. 16: 225-447.
Bose, P. K. 1937. Cytological studies of Argemone mexicana Linn. Jour. Indian Bot. 16: 197-208.
Boynton, K. R. 1926. Berberis thunbergii. Addisonia 10: 59.
Butters, F. K. 1909. The seeds and seedlings of Caulophyllum thalictroides. Minn. Bot. Stud. 4: 11-32.
Chapman, M. 1936. Carpel anatomy of the Berberidaceae Amer. Jour. Bot. 23: 340-348.
Citerne, P. E. C. 1892. Berberidees et Erythrospermees. Dupont, Paris, 161 pp.
Clark, L. 1923, The embryogeny of Podophyllum peltatum. Minn. Stud. Pi. Sci. 1: 111-138.
Clute. W. N. 1915. A mayapple with multiple fruits. Amer. Bot. 21: 92-93.
Crete, P. 1944. Apropos de la strophiole de Chelidonium majus L. Bull. Soc. Bot. Fr. 91: 88.
Cronquist, A. 1968. The Evolution and Classification of Flowering Plants. Houghton Mifflin Co., Boston, 396 pp.
1981. An Integrated System of Classification of Flowering Plants. Columbia Univ. Press, 1262 pp.
Cullen, J. 1965. Papaveraceae. In D. H. Davis (ed.) Flora of Turkey. 1: 221-223.
1968. The genus Papaver in cultivation. I. Wild species. Baileya 16: 73-91.
1975. Types of Papaveraceae in European herbaria. Int. Type Register, 66 pp.
Dannert, S. 1958. Zur Systematik von Papaver somniferum L. Kulturpflanze 6: 61-88.
Dawson, R, F. & C. James, 1956. Alkaloids of Papaver orientale L., I. Qualitative detection and occurrence.
Lloydia 19: 56-64.
59
Decaisne, J. 1837. Memoire sur la famille Lardizabalees. Extract des Arch. Mus. Hist. Nat., 213 pp.
Dermen, H. 1931. A study of chromosome number in two genera of the Berberidaceae: Mahonia and Berberis.
Jour. Arn. Arb. 12: 281-287.
Dickson, J. 1935. Studies in floral anatomy II. The floral anatomy of Glaucium flavum with reference to other
members of the Papaverareae. Jour. Linn. Soc. (London) 50: 175-224.
Diels, L. 1910. Menispermaceae. In: Engler, Das Pflanzenreich. 46: 1-345.
Dormer, K. J. 1954. The Acacian ty pe of vascular system and some of its derivatives. I. New Phytol. 53: 301-311.
Dreuilke, A. 1887. Les Menispermees et leurs produits. Montpellier, 45 pp.
Duke, J. A. 1970. Papaveraceous polyclave. Grit. Rev. Toxicology 3: 1-95.
1973. Annotated bibliography on opium and oriental poppies and related species. U.S.D.A. Agric. Res.
Serv., 349 pp.
Eckman, P. I. 1834. De Papavere somnifero: dissertatio botanico-medica: etc. Thesis, Uppsala.
Eichler, A. W. 1864. Versuch einer Charakteristik der natiirlichen Pflanzenfamilie Menispermaceen. Deut. Reg-
nesb. Bot. Gesel., vol. 5, 40 pp.
Elkan, L. 1839. Tentamen Monographie Generis Papaver. Regimonti Borussorum. E. J. Dalkowski, 36 pp.
Ernst, W. R. 1958. Chromosome numbers of some western Papaveraceae. Contr. Dudl. Herb. 5: 109-115.
1962a. The genera of Papaveraceae and Fumariaceae in the southeastern United States. Jour. Am.
Arb. 43: 315-343.
1962b. A Comparative Morphology' of the Papaveraceae. Ph.D. dissertation, Stanford Univ.
1964. The genera of Berberidaceae, Lardizabalaceae and Menispermaceae in the southeastern United
States. Jour. Arn. Arb. 45: 1-35.
Faberge, A. C. 1944. Genetics of the Scapiflora section of Papaver. III. Infraspecific hybrids and genetic homo-
logy7. Jour. Genet. 46: 125-149.
Fahmv, 1. R. et al. 1957. A pharmacognostical study of the seeds of a species of the genus Papaver grown in
Egypt. Pharm. & Pharmacol. 9: 541-548.
Fairliairn, J. W. & L. D. Kapoor. 1960. The laticiferous vessels of Papaver somniferum L. Planta Med. 8: 49-61.
Fedde, F. 1909. Papaveraceae (etc.) in Engler, Das Pflanzenreich. 40: 1-430.
1905. Die geographische Verbreitung der Papaveraceae. Bot. Jahrb. 36 (81): 28-43.
1936. Papaveraceae in Engler & Prantl, Die natiirlichen Pflanzenfam., ed. 2, Bd. 17B: 1-145.
Foerste, A. F. 1884. The May Apple. Bull. Torrey Bot. Club 2: 62
Friedel, J. 1927. Filiation des Papaveracees. Bull. Soc. Bot. Fr. 74: 673-688. 1929; ibid. 76: 285-291. 1930; ibid.
77; 36-42.
Fulling, E. H. 1943. Plant life and the law of man. Barberry, currant, gooseberry and cedar control. Bot. Rev. 9:
483-592.
Gagnepain, F. 1908. Revision des Lardizabilees asiatiques de Iherbier du museum. Bull. Mus. Hist. Nat. (Paris)
14: 64-70.
Gates, B. N. 1942. The dissemination by ants of the seeds of bloodroot, Sanguinaria canadensis. Rhodora 44: 13-
15.
1943. Carunculate seed disseminated by ants. Rhodora 45: 438-445.
Goldblatt, P. 1974. Biosystematic Studies in Papaver, Section Oxytona. Ann. Missouri Bot. Gard. 61: 264-296.
Granicher, S. 1906. Some notes on the pollination of flowers. Bull. Wise. Nat. Hist. Soc. 11. 4: 12-21.
Gray, A. 1848a. Ordo Papaveraceae. Gen. Pi. U.S. 1: 108-116.
1848b. Menispermaceae. Gen. Pi. U.S. 1: 69-76.
1886. Corydalis aurea and its allies. Bot. Gaz. 11: 188-189.
Greene, E. L. 1905. Revisions of Eschscholtzia. Pittonia 5: 139-194.
1905. Suggestions regarding Sanguinaria. Pittonia 5: 306-308.
Greger, J. 1930. Uber Kalkoxolat-Kristalle in der Samenschale von Fumaria officinalis L. Planta 12: 49-52.
Griffin, M. H. 1937. The Chromosome numbers of Berberis. Trans. Roy. Soc. So. Afr. 24: 203-206.
Hakim, S. A. E. 1957. Extraction and detection of poppy alkaloids. Jour. Physiol. 138: 8 p.
Halstead, B. D. 1894. Pistillodia of the Podophylhmi stamen. Bull. Torrey Bot. Club 21: 269.
Hammar, O. N. 1854. En Monografi ofver slagtet Fumaria. Berling, Lund. 58 pp.
Harper, J. L. & I. H. McNaughton, 1960. The inheritance of dormancy in inter- and intraspecific hybrids of
Papaver. Heredity 15: 315-320.
60
Harris, J. A. 1909. The leaves of Podophyllum peltatum Bot. Gaz. 47: 438- 444.
1910a. A quantitative study of the morphology of the fruit of the bloodroot, Sanguinaria canadensis.
Biometrika 7: 305-351.
1910b. Correlation in the inflorescence of Sanguinaria. Biol. Centralbl. 30:629-633.
Harshberger, J. W. 1903. Juvenile and adult forms of bloodroot. PI. World 6: 106-108.
Harvey-Gibson, R. & M. Bradley. 1916. Contributions towards a knowledge of the anatomy of the lower dicotyle-
dons. I. The anatomy of the stem of the Papaveraceae. Trans. Roy. Soc. Edinb. 51: 589-608.
Hegenauer, R. 1961. Die Gliederung der Rhoeadales sensu Wettstein im Lichte der Inhaltstoffe. Planta Med. 9:
37-46.
Herail, J. & R., Blottiere. 1886. Note sur les affinites des Lardizabalacees. Bull. Soc. Bot. Fr. 33: 521-524.
Heydenreich, K. and S. Pfeifer, 1966. Uber die Alkaloide der Gattung Papaver. II. Lindl. Pharmazie 21: 121-122.
Hildebrand, F. 1869-70. Ueber die Bestaubungsvorrichtungen bei den Fumariaceen. Jahrb. Wiss. Bot. 7: 423-
471.
Hills, K. L. & C. N. Rodwell. 1950. The recombination of some varietal characters in opium poppy. Austral. Jour.
Agric. Res. 1: 118-131.
Himmelbaur, W. 1914. Die Berberidaceen und ihre Stellung im System. Denk. Akad. Wiss. Math. Naturw.
(Vienna) 89: 733-796.
Holm, T. 1899. Podophyllum peltatum. a morphological study. Bot. Gaz. 27: 419-433.
1913. Medical Plants of North America. 78. Menispermum canadense L. Merck’s Rep. 22: 281-284.
Honeyman, J. M. 1956. On the occurrence of cyanogenic glycosides in the order Rhoeadales. Taxon 5: 33-34.
Hrishi, N. J. 1960. Cytogenetical studies on Papaver somniferum L. and P. setigirum DC. and their hybrid. From
“Genetica 1960.” Gron, 30 pp.
Hutchinson, J. 1920a. Bocconia and Macleaya. Kew. Bull. 1920: 275-282.
1920b. Jeffersonia and Plagiorhegma Kew. Bull. 1920: 242-245.
1920c. Caulophyllum thalictroides Grad. Chron. iii. 67: 63.
1921. The genera of Fumariaceae and their distribution. Kew Bull. 1921: 97-115.
1925. Contribution toward a phylogenetic classification of flowering plants: V. The genera of Papavera-
ceae. Kew. Bull. 1925: 161-168.
Jamieson, G. S. & W. G. Rose. 1943. Mexican prickly poppy seed oil. Oil-soap 20: 33-35.
Janchen, E. 1949. Die Systematishe Gliederung der Ranunculaceen und Berberidaceen, Springer Ost. St., 82 pp.
Jepson, W. L. 1922. Eschscholzia. In Fl. Calif. 1: 564-575.
Johnson, R.H. 1909. Aberrant societies of Sanguinaria and Trillium. Torreya 9: 5-6.
Kaezsmarck, F. & B. Malek. 1959. Zur Papier-chromatographie der Alkaloide von Chelidonium majus L. Planta
Med. 7: 171-173.
Kanta, K. 1960. Intra-ovarian pollination in Papaver rhoeas L. Nature 188: 683-684.
Karp, M. L. 1946. On the occasional absence of opium from the opium poppy. Compt. Rend. Acad. Sci. URSS II.
52: 537-540.
Kawatani, T. and H. Asahina. 1959. Characters and alkaloids of the artificial interspecific Fj hybrid between
Papaver orientale L. and P. somniferum L. Jap. Jour. Gen. 34: 353-362.
Kawatani, T. & T. Ohni. 1965. Chromosome numbers in Papaver Kok. Eis. Shik. Toyko. 83: 127-133.
Kern, F. D. Observations on the dissemination of the barberry. Ecol. 2: 211-214.
Kiger, R. W. 1973. Sectional nomenclature in Papaver. Taxon. 22: 579-582.
1975. Papaver of North America — north of Mexico. Rhodora 77: 410-422.
Kingsbury, J. M. 1964. Poisonous Plants of the United States and Canada. Prentice Hall, N. J., 626 pp.
Kitamura, T. and M. Sugamoto. 1961. Studies on the alkaloids of Berberidaceous plants XXXI. (In Japanese,
English summary) Jour. Pharm. Soc. Japan. 81: 254-261.
Kleinschmidt, G. 1961. Untersuch ungen uber die Alkaloid Verteilung in der Organen von Papaver orientale L. in
laufe der Vegetations-periode. Arch. Pharm. (Berlin) 294: 254-258.
Konovalova, R. A. et al. 1935. Uber die Alkaloide von Papaver Arten. I. Ber. Deutsch. Chem. Ges. 68: 2158-
2163.
Krukoff, B. A., A. Moldenke & H. N. Moldenke. 1938. Studies of American Menispermaceae with special refer-
ence to species used in preparation of arrow poisons. Brittonia 3: 1-74.
_ 1951. Supplementary notes on American Menispermaceae. V. Bull. Torrey Bot. Club 78: 258-265.
Kuester, H. L. 1926. A chemical study of the rhizome and roots of Podophyllum peltatum L. Jour. Amer. Pharm.
Assoc. 15: 259-263.
61
Kumazawa, M. 1936. Pollen grain morphology in Ranunculaceae, Lardizabalaceae and Berberidaceae. Jap. Jour.
Bot. 8: 19-46.
1937. Comparative studies on the vernation in the Ranunculaceae and Berberidaceae. (In Japanese)
Jour. Jap. Bot. 13: 573-586.
1938 a. Systematic and phylogenetic consideration of Ranunculaceae and Berberidaceae. Bot. Mae.
Tokyo 52; 9-15.
1938 b. On the ovular structure of Ranunculaceae and Berberidaceae. Jour. Jap. Bot. 14: 10-25.
Kunimoto, J. I. Studies in the alkaloids of Menispermaceae plants CLXXXII. Jour. Pharm. Soc. Japan 81: 1253-
1266.
Kuznetsova, G. A. et al. 1959. The study of the may-apple (Podophyllum peltatum L.) cultivated in the Leningrad
Region. (In Russian) Bot. Zhur. 44: 1337-1340.
Lange, J. 1961. Deciduous Barberries. Dansk. Vid. Forlag (Copenhagen) 72 pp.
Langlet, O. 1928. Einige Beobachtungen iiber die Zytologie der Berberidaceen. Svensk. Bot. Tidsk. 22: 169-184.
Lawalree, A. 1895. La chelidone double et la chelidone laciniee en Belgique. Bull. Jard. Bot. Brux. 25: 409- 410.
Lefort, L. 1950. Quelques groupes infra-specifiques luxemborgeois de Fumaria officinalis L. Soc. Nat. Luxem-
bourg. Bull. II. 44: 180-181.
Leger, L. 1894-5. Recherche sur l’appareil vegetatif des Papaveraceae. Juss. Mem. Soc. Linn. Normandie 18: 195-
623.
Leinfallner, W. 1956. Zur. Morphologie des Gynozeums von Berberis. Oster. Bot. Zeitsch. 103: 600-612.
Lestiboudois, T. 1823. Sur le ffuit des Papaveracees. Lille Rec. Trav. Soc. 1823: 181-194.
Levine, M. N., and R. U. Cotter. 1932. Susceptibility and resistance of Berberis and related genera to Puccinia
graminis. U.S.D.A. Tech. Bull. 300, 26 p.
Lewis, C. E. 1904. Studies on some anomalous dicotyledonous plants. Bot. Gaz. 37: 127-138.
Li, H. L. 1954. Akebia as a weed in the Philadelphia area. Morris Arb. Bull. 5: 58.
Lignier, O. 1897. La fleur des Crucifhrees comparee a celle des Fumariees. Assoc. Fr. Avanc. Sci. Compt. Rend.
25: 403-406.
1911. Notes anatomique sur l’ovaire de quelques Papaveracees. Bull. Soc. Bot. Fr. 58: 279-284: 337-
344; 429-435.
1915. Eschscholziees. Explication anatomique de la fleur. Bull. Soc. Bot. Fr. 62: 298-315.
Litardiere, R. de. 1921. Remarque au sujet de quelque processus chromosomique dans les noyaux diploidiques de
Podophyllum peltatum L. Compt. Rend. Acad. Sci. (Paris) 172: 1066-1068.
Ljungdahl, H. 1922. Zur Zytologie der Gattung Papaver Svensk. Bot. Tidsk. 16: 103-116.
Lloyd, J. U. & C. G. Lloyd. 1887. Caulophyllam thalictroides. Drugs Med. N. Am. 2: 141-162.
Maheshwari, P. & K. Kanta. 1961. Intraovarian pollination in Eschscholzia californica Cham., Argemone mexicana
L. and A. ochroleuca Sweet. Nature 191: 304.
Malhotra, S. K. 1960. Natural hybrid between Argemone mexicana and A. ochroleuca. Curr. Sci. Bangalore 29:
282.
Markgraf, F. 1953. Morphologish-phylogenetisch Studien an Bliiten der Rhoeadales Proc. VII Intern. Cong.
(Stockholm, 1950): 857-858.
Martin, F. W. 1958. Variation and morphology of Podophyllum peltatum. Diss. Abst. 19: 424-425.
Mauritzon, J. 1936. Zur Embryologie der Berberidaceen. Acta Hort. Gotoburg. 11: 1-17.
McNaughton, I. H. 1960. Internal breeding barriers in Papaver. Scott. Pi. Breed. Sta. Rep. 1960: 76-84.
McNaughton, I. H. & J. L. Harper. 1964. Papaver L. In: Biol. Fl. British Isles. Jour. Ecol. 52: 767-793.
Mellanofif, I. S. and H. L. Schaeffer. 1927. A study of the resins of Podophyllum peltatum L. Amer. Jour. Pharm.
99: 323-330.
Merriam, C. H. 1884. Dicentra punctured by humble-bees. Bull. Torrey Bot. Club 11: 66.
Miers, J. 1871. Menispermaceae. Contr. Bot. 3: 1-402.
Mika, E. S. 1955. Studies on growth and development and morphine content of opium poppy. Bot. Gaz. 116: 323-
339.
Mitchell, R. S. & C. J. Sheviak. 1981. Rare Plants of New York State. New York State Museum Bull. 445. 96 pp.
Miyaji, Y. 1930. Beitrage zur Chromosomenphylogenie der Berberidaceen. Planta 11: 650- 659.
62
Mowat, A. B. & S. M. Walters. 1964. Papaver. In Tutin et al., Flora Europaea, Cambridge, 1: 247-250.
Munz, P. A. 1963. A California Flora; Univ. Calif. Press, 1681 pp.
Murbeck, S. S. 1912. Untersuchungen iiber den Bliitenbau des Papaveraceen. Almquist & Wiksells, Uppsala,
168 p.
Nieuwland, J. A. 1910. Notes on the seedlings of bloodroot. Amer. Midi. Nat. 1: 199-203.
Nordhagen, R. 1959. Remarks on some new or little known myrmecochorous plants from North America and East
Asia. Bull. Res. Council, Israel D. 7: 184-201.
Musil. A. 1963. Identification of Crop and Weed Seeds. U.S.D.A. Agric. Handbook. 219, 171 p.
Negodi, G. 1937. Ulteriori contributi all cariologia, subfam. Fumaroiideae (etc.) Sci. Gen. 2: 1-25.
Norris, T. 1941. Torus anatomy and nectary characteristics as phylogenetic criteria in the Rhoeadales. Amer. Jour.
Bot. 28: 101-113.
Nowicke, J. and J. Skvarla. 1981. Pollen morphology and phylogenetic relationships of the Berberidaceae. Smiths.
Contr. Bot. 50. 83 pp.
Ohwi, J. 1931. Adlumia asiatica sp. nov. In: Symbolae ad Floram Asiae Orientalis III. Bot. Mag. Tokyo, 45: 387.
Ownbey, G. B. 1947. Monograph of the North American species of Corydalis. Ann. Mo. Bot. Gard. 34: 187-259.
1951. On the cytotaxonomy of the genus Corydalis section Eucorydalis Amer. Midi. Nat. 45: 184-
186.
1958. Monograph of the genus Argemone for North America and the West Indies. Torr. Bot. Club
Mem. 21: 1-159.
1961. The Genus Argemone in South America and Hawaii. Brittonia 13: 91-109.
Parlatore, P. 1844. Monografia delle Fumariee. Societa Tipographica, 110 pp.
Payne, W. W. 1970. Helicocytic and allelocytic stomata: unrecognized patterns in the Dicotoledonae. Amer. Jour.
Bot. 57: 140-147.
Popov, M. G. 1970. Papaveraceae. In: Komarov, V. L. Flora of the U.S.S.R. 7: 437-549.
Porter, T. C. 1877. Variations in Podophyllum peltatum L. Bot. Gaz. 2: 117.
Prain, D. 1895a. A revision of the genus Chelidonium. Bull. Herb. Boiss. 3: 570-587.
1895b. An account of the genus Argemone. Jour. Bot. (London) 33: 129-135.
Pugsley, H. W. 1919. A revision of the genera Fumaria and Rupicapnos. Joui. Linn. Soc. 44: 233-355.
1927. Further notes on the genera etc. . . ibid. 47: 427-468.
1932. Further notes on the genera etc. . . ibid. 48: 93-113.
1934. Further notes on the genera etc. . . ibid. 49: 517-529.
Raymond, M. 1948. A red-fruited form of Podophyllum peltatum. Rhodora 50: 18.
Reauburg, G. 1906. £tude organographique et anatomique de la famille Lardizabalees. Thesis, Paris, ficole Super-
ieure de Pharmacie no. 10., 127 pp.
Robertson, C. 1896. Flowers and Insects. XVII. Bot. Gaz. 22: 154-165.
Roder, I. 1958. Anatomische und Floureszenzoptische Untersuchungen an Samen von Papaveraceen. Osterr. Bot.
Zeitsch. 104: 370-381
Rogers, C. 1900. Trodden Glory: The Story of the California Poppy. W. Hebberd, Santa Barbara Calif. (1949),
129 pp.
Ryberg, M. 1955. A taxonomical survey of the genus Corydalis Ventenat, with reference to cultivated species. Acta
Hort. Berg. 17: 115-175.
1960. A morphological study of the Fumariaceae and taxonomic significance of characters examined.
Acta Hort. Berg. 19: 122-248.
Saito, K. 1957. Studies on the induction of polyploid plant deletion of five leaf Ahebia. (In Japanese) Jour. Hort.
Assoc. Japan. 26: 43-44.
Sargent, C. S. 1891. The fruit of Akebia quinata, Garden Forest 4: 136-137.
Sawada, T. 1928. Whence it originated, Rajania quinata, a conditional synonym Akebia quinata. (In Japanese)
Jour. Jap. Bot. 5: 153-157.
Sawyer, M. L. 1926. Carpelloid stamens of Podophyllum peltatum. Bot. Gaz. 82: 329-332.
Schijfsma, L. et al. 1960. A study of the colour and other characters of the seed of some varieties of oil seed
poppy. Euphytica 9: 127-140.
Schmidt, E. 1928. Untersuchungen iiber Berberidaceen. Beih. Bot. Centr. 45: 329-396.
Schmitt, C. G. and B. Lipscomb. 1927. Pathogens of Selected Members of the Papaveraceae. U.S.D.A. Agric.
Res. Serv. (ARS-NE-62), 186 p.
63
Schweizer, G. 1938. Zur Anatomie des Mohnsahmans ( Papaver somniferum L.) Deutsche. Bot. Gesell. (Berlin) 49:
414-423.
Scott, J. M. 1969. The White Poppy; A History of Opium. Funk & Wagnalls, N.Y., 205 pp.
Scott, W.R.M. & E. J. Petry. 1920 Correlation of variation in resin content of Podophyllum with certain habitats.
Rep. Mich. Acad. Sci: 21: 225-231.
Shaw, C. H. 1904. Note on the sexual generation and the development of the seedcoats in certain Papaveraceae.
Bull. Torrey Bot. Club 31: 429-433.
Soejarto, D., R. Faden and N. Farnsworth. 1979. Indian Podophyllum L. Is it Podophyllum errwdi or Podophyllum
hexandrum? Taxon 28: 549-551.
Spencer, W. P. 1944. Variation in petal number in the blood-root, Sanguinaria canadensis. Amer. Nat. 78: 85-89.
Stakman, E. C. et al. 1934. Relation of barberry to the origin and persistence of physiologic forms of Puccinia
gratninis Jour. Agr. Res. 48: 953-969.
Stermitz, F. R. 1968. Alkaloid chemistry and the systematics of Papaver and Argemone. Adv. Phytochem. 1: 161-
183.
Stermitz, F. R. & H. Rapaport. 1961. The biosynthesis of opium alkaloids. Alkaloid interconversions in Papaver
somniferum and P. orientale. Jour. Amer. Chem. Soc. 83 : 4045-4050.
Stern, K. R. 1961. Revision of Dicentra (Fumariaceae). Brittonia 13: 1-57.
Steyermark, J. A. 1952. Color forms of the may-apple. Rhodora 54: 131-134.
Suguira, T. 1940. Chromosome studies on Papaveraceae. . . etc. Cytologia (Tokyo) 10: 558-576.
Surface, F. M. 1905. Contributions to the life history of Sanguinaria canadensis. Ohio Nat. 6: 379-385.
Sullivan, B. H. & H. I. Wechsler. 1947. The cytological effects of podophyllin. Science 105: 433.
Swarbrick, 1. T. and J. C. Raymond. 1970. The identification of the seeds of British Papaveraceae. Ann. Bot. 34:
1115-1122.
Szemes, G. 1943. Zur Entwicklung des Elaisoms von Chelidonium majus. Wiener Bot. Zeitschr. 92: 215-219.
Terabayashi, S. 1978. Studies in morphology and systematics of Berberidaceae II. Floral anatomy of Mahonia
japonica (Thunb.) DC. and Berberis thunbergii DC. Acta Phytotax. et Geobot. 29: 106-118.
Thompson, N. E. & W. W. Robbins. 1926. Methods of eradicating the common Barberry (Berberis vulgaris).
U.S.D.A. Bull. 1451, 46 pp.
Tischler, G. 1920. Die Berberjdaceen und Podophyllaceen. Bot. Jahrb. 31: 596-727.
Torrey, J. and A. Gray. 1838. Berberidaceae. Flora North America. 1: 49-54.
True, R. H. & W. W. Stockberger. 1916. Physiological observations on alkaloids, latex and oxidoses in Papaver
somniferum. Amer. Jour. Bot. 3: 1-11.
Vaarama, A. 1949. The chromosome number of Fumaria officinalis. Heriditas: 35: 251-252.
Viguier, L.G.A. 1814. Histoire Naturelle Medicate et Economique des Pavots et des Argemones. 50 (1) pp.
Vesler, J. 1913. Zum Entwicklungsgeschichte von Akebia quinata. Diss. Bonn.
Vilmorin, M. L. de, 1915. Deux Lardizabalees a fruits comestibles. Bull. Soc. Acclim. Fr. 62: 89-93.
Wadmond, S.C. 1898. Leaf retardation in Podophyllum peltatum. Asa Gray Bull. 6: 66-67.
Wigdale, E. G. 1900. The structures of the rhizome and root of Podophyllinn. Quart. Jour. Pharm. 10: 40-51.
Wilkinson, R. E. 1953. Berberis thunbergii a host of cucumber mosaic virus (Marmor cucumeris) (Abs.) Phytopath.
43: 489.
Willamen, J. J. and B. G. Schubert. 1961. Alkaloid-bearing Plants and their Contained Alkaloids. U.S.D.A. Tech.
Bull. 1234, 287 pp.
Woodson, R. I. 1928. A new genus of Berberidaceae. Ann. Missouri Bot. Gard. 15: 335-340.
Yasui, K. 1937. Cytogenetic studies in artificially raised interspecific hybrids of Papaver VI. The trigenomic hy-
brids of Papaver. Cytologia, Jubilee vol.: 1101-1112.
Zaitseva, A. A. 1951. On the latex of the opium poppy. Argobiologia 3: 122-126. (in Russian)
64
INDEX TO LATIN NAMES
Adlurnia
asiatica. 44
cirrhosa 44
fungosa 44
komarovii 44
Akebia
lobata. 11
quinata 11
trilobata 10
Argeinone
alba 35
albiflora 35
georgiana. 35
intermedia 35
mexicana 37
mucronata 37
munita 38
platyceras 38
polyanthemos 36
sexvalvis 37
spinosa 37
vulgaris .................................... 37
versicolor. 37
Berberis
x ottowensis 8, 10
thunbergii . 9, 10
vulgaris .................................... 7
Bicuculla
canadensis 43
cucullaria. . 41
eximia 40
Bocconia
cordata. 22
japonica .................................... 22
Calomecon
orientale 31
Capnodes (Capnoides)
aureum 50
flavulum 48
glauca. ..................................... 46
Capnorchis
canadensis .................................. 40
cucullaria. 41
eximia ..................................... 40
Caulophyllum
thalictroides 5
Chelidonium
glaucum. . 20
majus 18
Corydalis
aurea 50
canadensis 43
flavula 48
Formosa 40
geyeri. 48
glauca 46
lutea. 49
rosea 46
seinpervirens 46
Cucullaria
bulbosa 41
cucullaria. 41
Dicentra
canadensis . 43
cucullaria 41
eximia 40
occidentals 41
Diclytra (Dielytra) 40, 43
Ectrus
mexicanus 37
Eomecon 16
Eschscholzia
californica 23
crocea 23
cucullata . 23
douglasii ................................... 23
glauca. 23
maritima 23
peninsularis. 23
Fumaria
cucullaria 41
eximia 40
flavula 48
glauca. 46
lutea. 49
officinalis. 51
sempervirens 46
Glaucium
flavum 20
glaucium 20
leiocarpum 20
luteum 20
Jeffersonia
bartonis 4
binata. 4
diphylla 3
dubia 3
65
Leontice
thalictroides 5
Macleaya
cordata 22
microcarpa 21
Menispermum
angulatum 13
canadense 13
mexicanum 13
Virginian um 13
Mosenthinia 20
Neckeria
au re a 50
flavula 48
sempervirens 46
Papaver
albiflorum 30
atlantieum 29
bracteatum 32
commutatum 34
dubium 30
glaucum 27
integrifolium 33
intermedium . 33
laciniatum 33
lateritium 30
modestum 30
mursellii 25
obtusifolium 30
orientale 31
paucifoliatum 31
pseudo-orientale 31
rapiferum 33
rhoeas 33
rupifragum 30
setigerum 25
somniferum 25
speetabile 31
strigosum 33
tenuissimum 31
timidulum 33
umbrosum 33
Podophyllum
diphyllum 4
peltatum 2
Rajania
quinata 11
Sanguinaria
acaulis 16
canadensis 16
dilliniana 16
vernalis 16
82-6841
66
ll1W il° k, Bo'ani=al Garden Library
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CONTRIBUTIONS TO A FLORA OF NEW YORK STATE
1. Mitchell, Richard S. & J. Kenneth Dean. 1978. Polygonaceae (Buckwheat Family) of New York State. Contri-
butions to a Flora of New York State I. N.Y. State Museum Bull. 431, 81 pp. $3.00
2. Mitchell, Richard S. & Ernest O. Beal. 1979. Magnoliaceae through Ceratophyllaceae of New York State.
Contributions to a Flora of New York State II. N.Y. State Museum Bull. 435, 62 pp. $3.00
3. Ketchledge, Edwin H. 1980. Revised Checklist of the Mosses of New York State. Contributions to a Flora of
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4. Andrus, Richard E. 1980. Sphagnaceae (Peat Moss Family) of New York State. Contributions to a Flora of
New York State III. N.Y. State Museum Bull. 442, 89 pp. $4.00
5. Mitchell, Richard S. & J. Kenneth Dean. 1982. Ranunculaceae (Crowfoot Family) of New York State. Contri-
butions to a Flora of New York State IV. N.Y. State Museum Bull. 446, 100 pp. $7.00
6. Mitchell, Richard S. 1983. Berberidaceae through Fumariaceae of New York State. Contributions to a Flora of
New York State V. N.Y. State Museum Bull. 451, 66 pp.
OTHER RECENT BOTANICAL PUBLICATIONS
Ogden, Eugene C. 1981. Field Guide to Northeastern Ferns. New York State Museum Bull. 444, 122
pp. $10.00
Mitchell, Richard S. & Charles J. Sheviak. 1981. Rare Plants of New York State. New York State
Museum Bull. 445, 96 pp., 55 illustrations. $8.00
Brooks, Karl L. A Catskill Flora and Economic Botany. 1979-80
Part I. Pteridophyta. The Ferns and Fern Allies. 276 pp. $4.00
Part II. Coniferales. The Conifers. 116 pp. $3.00
Part III. Apetalae. Including Poplars, etc. 374 pp. $5.00
Mitchell, Richard S. 1979. Preliminary Lists of Rare, Endangered and Threatened Plants in New York
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Slack, Nancy. 1977. Species Diversity and Community Structure in Bryophytes. New York State
Studies. N.Y. State Museum Bull. 428, 70 pp. $3.00
Ogden, Eugene C., et. al. 1976. Field Guide to the Aquatic Plants of Lake George. N.Y. State Museum
Bull. 426, 65 pp. $3.00
Ogden, Eugene C. 1974. Potamogeton in New York. N.Y. State Museum Bull. 423, 20 pp. $1.00
Ogden, Eugene C. 1974. Anatomical Patterns of Some Aquatic Vascular Plants in New York. N.Y. State
Museum Bull. 422, 133 pp., 208 maps, 67 pis. $3.00
Ogden, Eugene C., et. al. 1974. Manual for Sampling Airborne Pollen. Hafner Press, 182 pp. $10.00
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