Taxonomic Studies of the Genus Polymnia L
This dissertation has been 64—6976 microfilmed exactly as received
WELLS, James Ray, 1932— TAXONOMIC STUDIES OF THE GENUS POLYMNIA L.
The Ohio State University, Ph.D., 1963 Botany
University Microfilms, Inc., Ann Arbor, Michigan TAXONOMIC STUDIES OF THE GENUS POLYMNIA L.
DISSERTATION
Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University
By
JAMES RAY WELLS, B.S., M.S.
The Ohio State University
1963
Approved by
Adviser Department of Botany and Plant Pathology ACKNOWLEDGMENTS
I wish to express my sincere gratitude to Dr. T. v Richard Fisher, my adviser, who suggested this problem and under whose guidance the work wan carried out. I am also
grateful to Drs. Clara Weishaupt, J.W.A. Burley, C. E. Taft,
and Glenn W. Blaydes for reading this dissertation and for
offering helpful suggestions and criticisms.
I am indebted to the curators of the several herbaria
for making their material available.
Permission to use Goode Base Maps (Copyright by the
University of Chicago) for plotting plant distributions is
gratefully acknowledged. This permission was granted through
Goode Base Map Series, Department of Geography, The University
of Chicago.
My wife Jan, is due special recognition for her loyal
support and cooperation throughout this endeavor. TABLE OF CONTENTS Page
ACKNOWLEDGMENTS...... ii
LIST OF T A B L E S ...... iv
LIST OF ILLUSTRATIONS...... v
INTRODUCTION ...... 1
MORPHOLOGY ...... 3
GEOGRAPHIC DISTRIBUTION ...... 5
ECONOMIC IMPORTANCE ...... 7
TAXONOMY ...... 8
KEY TO SPECIES AND VARIETIES...... 11
DESCRIPTION AND DISCUSSION OF T A X A ...... 15
EXPERIMENTAL TAXONOMIC INVESTIGATIONS ...... 131
LIST OF TAXA OF UNCERTAIN S T A T U S ...... 156
LIST OF EXCLUDED SPECIES...... 160
DISTRIBUTION M A P S ...... 163
SUMMARY ...... 17^
BIBLIOGRAPHY ...... 175
AUTOBIOGRAPHY ...... 179
iii LIST OF TABLES Table Page
1 Polymnia uvedalia soil habitat data ...... 79
2 List of chromosome numbers of certain taxa of Polymnia...... 13^
3 Pollen stainability of taxa which occur naturally in the United States ...... 1^1
k Summary of interspecific hybridization attempts . 1^3
5 Summary of intravarietal hybridization attempts, selfings, and tests for ovulary abortion . . . 1^6
6 Crosses involving two plants of P. uvedalia var. uvedalia (collections 236 and 239^ from different ends of a large p o p u l a t i o n ...... 1^9
7 Summary of intraspecific hybridization attempts . 150
8 Crosses involving P. laevigata collection number 25^ ...... 153
iv LIST OF ILLUSTRATIONS Figure Page
1 Polymnia uvedalia (Holotype) ...... 10
2 Polymnia oaxacana (Isotype) ...... • 16
3 Polymnia laevigata (Isotype) ...... 21
k Polymnia canadensis ...... 2k
5 Polymnia connata (Holotype) ...... k$
6 Polymnia connata...... ^-6
7 Polymnia .jelskii ( H o l o t y p e )...... ^9
8 Polymnia .jelskii (Holotype F r a g m e n t s ) ...... 50
9 Polymnia siegesbeckia (Holotype) ...... 52
10 Polymnia siegesbeckia ...... 53
11 Polymnia riparia (Holotype) ...... 55
12 Polymnia riparia ...... 56
13 Polymnia maculata var. maculata ...... 62
l^f Polymnia uvedalia var. densipilis (Peduncle of Holotype) I T"...... 8l
15 Polymnia uvedalia var. floridana (Peduncle of Holotype) I 7~...... 97
16 Polymnia uvedalia var. floridana ...... 98
17 Polymnia quichensis (Holotype) ...... 10^
18 Polymnia meridensis (Holotype) ...... 107
19 Polymnia sonchifolia ...... 110
20 Polymnia apus (Holotype)...... 113
21 Polymnia apus (TIsotype) ...... 11 ^
v LIST OF ILLUSTRATIONS— (Continued) Figure Page
22 Polymnia suffruticosa ...... 116
23 Polymnia pyramidalis ( H o l o t y p e ) ...... 119
24 Polymnia glabrata (Holotype) ...... 123
25 Polymnia parviceps (Holotype) ...... 125
26 Polymnia microcephala (Holotype) ...... 127
27 Polymnia fruticosa ...... 129
28 Polymnia laevigata pollen mother cell at diakinesis (Wells 255) ...... 135
29 Polymnia laevigata pollen mother cell at early metaphase (Wells 2 5 5 ) ...... 136
30 Polymnia uvedalia var. uvedalia pollen mother cell at diakinesis (Wells 211) ...... 138
31 Polymnia uvedalia var. uvedalia pollen mother cell at diakinesis (Wells 2 1 1 ) ...... 139
32 Polymnia canadensis pollen mother cell at diakinesis (Wells 218) ...... 140
33 Polymnia canadensis var. radiata (pollen parent) X Polymnia canadensis var. canadensis (ovule parent) hybrid ...... 151
34 Polymnia laevigata (pollen parent) X Polymnia canadensis var. canadensis (ovule parent) hybrid . 154
35 Polymnia bakeriana (Isotype)...... 157
36 Known distribution of Polymnia canadensis excluding Canada ...... 163
37 Known distribution of Polymnia uvedalia excluding Bermuda 164
38 Known distribution of Polymnia laevigata ...... 165
39 Known distribution of Polymnia oaxacana and Polymnia quichensis ...... 166
40 Known distribution of Polymnia apus and Polymnia maculata var. adenotricha ...... 167
vi LIST OF ILLUSTRATIONS— (Continued) Figure Page
41 Known distribution of Polymnia maculata vars, maculata and glabricaulis ...... 168
42 Known distribution of Polymnia riparia ...... 169
43 Known distribution of Polymnia glabrata. Polymnia parviceps, and Polymnia microcephala . 170
44 Known distribution of Polymnia pyramidalis and Polymnia connata ...... 171
45 Known distribution of Polymnia siegesbeckia, Polymnia fruticosa, and Polymnia suffruticosa . 172
46 Known distribution of Polymnia sonchifolia and Polymnia meridensis...... 173
vii INTRODUCTION
The generic name Polymnia is derived from the Muse
"Polyhymnia" who was one of the Greek goddesses of song and poetry. This Linnaean genus dates back to 1751*
Heretofore, no known comprehensive study of the genus
Polymnia has ever been carried out. This is an attempt directed toward producing a semblance of order and understanding in a poorly-understood genus. Earlier workers were not sufficiently aware of the limits of the genus Polymnia and partly for this reason it was often confused with other genera among which were:
Adenostemma, Axiniphyllum, Berlandiera, Didelta, Guizotia,
Melampodium, Montanoa, Rumfordia, Tetragonotheca, and Wedelia.
E. L. Core (1955) and others have pointed out that a
"revision" of a genus may be based only on studies of herbarium
specimens from one continent or a smaller area whereas a "mono
graph" is world-wide in scope and takes into account other
disciplines such as: cytogenetics, anatomy, physiology, mor
phology, paleobotany, and ecology. This study of the genus
Polymnia does not readily fit into either of these two categories
but has the characteristics of being world-wide in scope of
coverage and based largely on studies of herbarium specimens.
Also, where practicable, considerations are given to geography,
cytology, ecology, and genetics. More thorough evolutionary considerations must await extensive travel and collections in the western hemisphere, particularly South America. Hybridization experiments along with cytological investigations will have to be carried out with all taxa.
This, then, is an effort directed toward delimiting the taxa as they are now understood and at the same time laying the foundation for more experimental investigations and refinement of present concepts. A cytotaxonomist must know the currently valid name of the plants with which he is working. Although these names may not be truly representative of phylogenetic relationships, they will, at least, serve as a starting point
from which various taxa may be combined or separated as experi mental evidence becomes known.
Citations to herbarium specimens examined are made in accordance with abbreviations cited by Lanjouw and Stafleu
(1959); Field Museum of Natural History (F), Gray Herbarium
(GH), Indiana University (IND), Louisiana State University (LSU),
Missouri Botanical Garden (MO), New York Botanical Garden (NY),
University of North Carolina (NCU), Ohio State University (OS),
Ohio University (OU), Philadelphia Academy of Science (PH),
University of Tennessee (TENN), University of Texas (TEX), United
States National Herbarium (US). MORPHOLOGY
Polymnia species are for the most part perennials. Their root system is generally thought to be fibrous although no refer ence is known to me of the type of root system of the tree forms which grow in South America. In P. uvedalia, P. sonchifolia, and P. connata the underground parts are tuberous.
The stems reach diameters up to 15-20 cm and attain heights of up to 12 meters. Stems may be terete, angular, or grooved. The stems may be glabrous or with glands and/or hairs.
Sometimes the stems are purple-spotted or have purple stripes.
Leaves in all cases known to me are opposite. In the species which attain tree size the leaves are unlobed. In herbaceous or shrubby species they may be lobed or unlobed.
The ray flowers are monosporangiate and ovulate. The
disc flowers produce pollen but no fruits or seeds. The United
States species are for the most part self-sterile. The pre
dominant color of corollas is yellow although white, yellowish- white, orange or reddish can be found. A pappus has not been
found on flowers of any species.
The leaves and peduncles vary from glabrous through
puberulent, pilose, glandular-pilose, or densely stipitate-
glandular. The type of vesture has been given considerable im
portance by Blake (1917a), (1917b), and (1935).
3 k
Previous authors have treated the phyllaries as being in two series. I have interpreted the lower set of about four to 6i£ broadened leaf-like structures associated with each head as phyllaries. The upper set is of the same number as the ray flowers and one of these "phyllaries" subtends each of the ray flowers in much the same manner as the chartaceous disc pales subtend the disc flowers. It seems to me to be morphologically sound to treat this upper set of structures as pales of the ray flowers and not an upper series of phyllaries. Polymnia laevigata and P. canadensis have biseriate phyllaries; other species have uniseriate phyllaries. GEOGRAPHIC DISTRIBUTION
The genus Polymnia is, for the most part, a temperate genus. In regions near the equator species are found as high as 3,000 meters or even higher. This genus is apparently indigenous and endemic to the western hemisphere. Its species are believed to occur in every country on the two continents although no collections were examined from British, Dutch, or
French Guiana.
Outside the continental limits of North and South
America P. uvedalia has been collected in Bermuda. Reports of occasional transplants to Europe can be found.
When all the specimens examined are compared to the total area of known ranges of distribution the number of plants collected per square mile is indeed small. More extensive col lections have been made in the United States than any other country. In this country no species have been reported west of central Texas and Oklahoma.
Collection data support the idea of a disjunct distri bution between the United States and Mexico. Any formerly continuous distribution between these two countries probably preceded the occurrence of arid lands in the southwest.
Alternate raising and lowering of the ocean levels probably did not destroy the mountainous refugia in Central and
5 South America. In Guatemala taxa are found which closely re semble other Guatemalan species and at the same time bear a marked resemblance to species in northern South America.
The species of Polymnia most distinct morphologically,
P. suffruticosa, occurs at low altitudes in Venezuela. This
species is known only from the Amazonas Territory in Venezuela and has been found at altitudes of 130 to 500 meters. ECONOMIC IMPORTANCE
In the United States no species of Polymnia is recognized as being of significant economic value. However, the importance
of plant species as soil builders and agents in abating erosion necessarily includes the several species of Polymnia.
Various species of Polymnia have been used as food by
cattle and man. In Venezuela cattle have been known to eat
leaves of P. riparia. P. sonchifolia (P. edulis) is cultivated
by natives in Bolivia and Ecuador for the sweet napiform tubers
or roots— the exact organ being unknown to me. The tubers or
fleshy roots of P. connata are eaten in Argentina where plants of
the species are apparently cultivated to some extent.
Some medicinal value seems to be derived from three
other species. W. H. Camp noted on his collection No. E 1980
that in Ecuador leaves of P. fruticosa are heated and tied onto
the legs as a cure for rheumatism. Leaves of P. pyramidalis
are used in Venezuela as a treatment for headache. The leaves
are heated and placed on the forehead. From P. glabrata parts
of the plant are used in treating the common cold.
Old wood of P. glabrata is used by natives in Peru—
presumably as fuel.
7 TAXONOMY
Polymnia L., 1751• Nova Plantarum Genera. Amoenitates Academicae. Vol. 3s22.
Alymnia Neck.. Elem. I (1790) 31.
Polymniastrum Lam., Illustr. (1798) t. 712; III. (1823) 287.
Uvedalia DC., Prodr. V. (1 8 3 6) 516.
Mostly if not all erect perennial herbs or trees ex
ceeding 12 meters in height, stems terete, angular or caniculate,
up to 20 cm thick at base, bark of large species becoming rough
on older stems; at least the lower leaves opposite, entire to
deeply lobed, the lobes and/or main veins pinnate or palmate,
sessile or petiole winged to the sometimes auriculate base or,
distinctly petiolate, the leaf shape often quite variable on the
same plant; heads few to many'; the phyllaries usually **— 6, in 1
or 2 series; ligulate flowers in 1 or infrequently 2 series,
monosporangiate, carpellate, corollas 2-3-dentate or without
teeth, yellow, white, or orange-red; pales subtending ray flowers
usually concave, the tip sometimes projecting to an elongate
point; disc flowers apparently monosporangiate, staminate, of
about the same number as the ray flowers or usually many more
than the ray flowers, their corollas tubular, usually yellow or
sometimes purplish at least when dried; pappus wanting; pales
subtending disc flowers usually chartaceous or infrequently of firmer texture and subulate, with or without prominent veins; receptacle flattened, somewhat convex to decidedly convex; fruits forming only from ovularies of ray flowers, the brown, purplish- black or black fruits obovoid or spherical and slightly flat tened laterally, or 3-5-angled, the angles infrequently winged,
7 mm or usually less in the largest dimension.
Type species: Polymnia uvedalia L. Probably collected by Frederick Gronovius in Virginia prior to 1751* A copy of the photograph of the holotype appears below. tllrflpiZT fi+3. fy b f fit- JJtffJ/fuA*Y
Fig. 1.— Polymnia uvedalia (Holotype) S cale: .63X KEY TO SPECIES AND VARIETIES
1. At least the middle or lower leaves pinnately lobed, sinuses extending at least halfway to midvein; ray corollas orange to orange-red, white, or very pale y e l l o w ...... 2
1. Leaves entire or palmately lobed, sinuses sometimes broad and shallow, ray corollas distinctly yellow or ray flowers biligulatea ...... 4
2. Leaves sessile or petiole winged to base, ray corollas orange to orange-red (Guatemala, Honduras, Mexico) . . * ...... (A) P. oaxacana
2. Leaves distinctly petiolate, ray corollas white to very pale y e l l o w ...... 3
3. Stems glabrous, fruits 5-angled, ray corollas white (Ala., Fla., Ga., Mo., T e n n . ) ...... (B) P. laevigata
3. Stems stipitate-glandular or glandular and pilose especially upward, fruit 3-angled, ray corollas white or pale yellow (eastern Canada, eastern to midwestern United States) ...... (C) P. canadensis
4. Ray flowers biseriate or biligulate ...... 5
4. Ray flowers in one series, the corolla single and unaccompanied by a second corolla or bractlike structure attached at top of fruit ...... 6
5. Ray flowers biseriate, fruit biconvex, sometimes tuberculate, without wings, herbs (Argentina, Brazil, Paraguay, Uruguay)...... (D) P. connata
5. Ray flowers biligulate, the second corolla or bractlike structure attached at top of the 4-angled, winged fruit, small trees or shrubs (Peru)...... (E) P. jeiskii
3.The term "biligulate" is taken from the original description and is intended to mean two complete and separate ray corollas.
11 6. Margin of pales subtending ray flowers involute, prolonged to an elongate point, stipitate- glandular below, the glands up to 1 mm long . . • 7
6. Margin of pales subtending ray flowers not in volute, apex acute or acuminate but not prolonged into an aristate point, if glandular then glands sessile or nearly so ...... 8
Disc flowers about 15 or fewer, ray corollas about 5-6 mm long, phyllaries 1 cm long or usually less (Bolivia, Brazil, Paraguay, Peru) ...... (F) P. siegesbeckia
Disc flowers about 30 or more, ray corollas about 10 mm long, phyllaries usually longer than 1 cm (Bolivia, Colombia, Ecuador, Guatemala, Peru, Venezuela) ...... (G) P. riparia
8. Herbs; leaves, at least the lower, lobed, the lobes sometimes separated by shallow and broad sinuses; fruits ovoid or obovoid ...... 9
8. Trees, woody shrubs or herbs; leaves entire, dentate, or crenulate...... * ...... 1 ^
Mature flowering stems purple-spotted; middle and lower leaves palmately veined and lobed; fruit laterally flattened, ovoid, multistriate, k mm or more in length . 10
Mature flowering stems not purple-spotted; middle and lower leaves if palmately veined and lobed then fruit asymmetrically obovoid and/or angular, spherical or less than k mm in l e n g t h ...... 1 1
10. Fruit 5 nun long or less, about k mm wide (Costa Rica, Guatemala, Honduras, Mexico). .(H) P. maculata
a) peduncles hispid-pilose . . .(I) var. maculata b) peduncles densely stipitate- glandular ...... (J) var. adenotricha c) peduncles and stems glabrous (K) var. glabricaulis
10. Fruit larger, more than 5 nun long and 4 mm wide (United States, Bermuda) ...... (L) P. uvedalia
a) peduncles denselypilose. .(M) var. densipilis b) peduncles densely stipitate- glandular ...... (N) var. uvedalia c) peduncles somewhat glandular and sparsely pilose or pubescent ...... (0) var. floridana 13
11. Stems and upper surface of leaves glabrous; leaves lanceolate, distinctly petiolate, the petiole not winged (Costa Rica, Guatemala) .... (P) P. quichensis
11. Stems and upper surface of leaves hairy, leaves broadly ovate or triangular; petioles winged to base or leaves s e s s i l e ...... 12
12. Plants of coarse texture; stems scabrous or hispid; leaves hispid above, peduncles stout; phyllaries ovate, 12 -l^f mm long, 1 0 -1 2 mm wide (Colombia, Venezuela) . . . (Q) P. meridensis
12, Plants of finer texture; stems and upper surface of leaves pilose or strigose; peduncles of smaller diameter; phyllaries lanceolate, up to 2 .5 cm long, 8 -9 mm wide or l e s s ...... 13
15. Winged portion of leaf longer than broad; peduncles pilose and glandular; phyllaries 15 mm long or less, up to 8 mm wide; pales of ray flowers 7 mm long or less, about 2 mm wide (Bolivia, Colombia, P e r u ) ...... (R) P. sonchifolia
13. Winged portion of leaf about as broad as long; peduncles stipitate-glandular; phyllaries up to 2 .5 cm long, about 9 mm wide; pales of ray flowers about 10 mm long, 3 mm wide ( M e x i c o ) ...... (S) P. apus
1 ^. Leaves lanceolate, small, less than 10 cm long and 3 cm wide, entire or only' remotely crenulate, densely pilose-glandular above and below, fruit oblong, equally quadrangular (Venezuela) ...... (T) P. suffruticosa
l^f. Leaves larger, distinctly dentate or lobed, if entire then fruit not equally quadrangular nor leaves densely pilose above ...... 1 5
15. Herbs; stems, upper surface of leaves, and peduncles glabrous; leaves lanceolate, distinctly petiolate; fruits spherical 5-6 mm in diameter (Costa Rica, Guatemala) ...... (P) P. quichensis
15. Trees or shrubs; hairs and/or glands present on leaves, stems, or peduncles; leaves ovate to broadly lanceolate, with or without winged petioles; fruit less than 5 mm in diameter, obpyramidal or flattened-obovoid, often angular...... 16 Ik
16. Upper parts of stems and peduncles densely pilose; leaves broadly triangular-ovate, the base attenuate and sometimes distinctly petiolate, margins entire or with small teeth about 1 mm high, densely tomentose below (Ecuador, Venezuela) ...... (U) P. pyramidalis
16, Upper parts of stems and peduncles glabrous, glandular and/or with hairs; leaves oblanceolate to broadly ovate, entire or deniate, the teeth up to k mm long, sessile or petioles winged to base, the lower winged portion sometimes auriculate, lower surface glabrous to tomentose ...... P. glabrata complex
a) Leaves without auriculate bases, oblance olate, long-acuminate, margins often entire; heads usually more than 5 om in diameter (Peru)...... (V) P. glabrata
b) Leaf bases sometimes auriculate, lanceolate to broadly ovate, margins usually with rounded teeth; heads about 5-6 mm in diameter (Bolivia, Peru) . . (W) P. parviceps
c) Leaves apparently all entire, heads about 5 mm in diameter (Ecuador) ...... (X) P. microcephala
d) Leaves shining-glabrous above, below densely tomentose, peduncles and phyl laries densely stipitate-glandular (Colombia, Ecuador, Peru). . (Y) P. fruticosa DESCRIPTION AND DISCUSSION OF TAXA
(A) Polymnia oaxacana Schultz Bip. Leopoldina. 23:89. 1 8 8 7.
Polymnia nelsonii Greenm. Proc. Amer. Acad. 39:99* 1903* (Isotype GHJ, US!).
Herbs to 2 meters in height; stems round, erect, sparingly hirsute to densely pilose, often purple-striped; leaves opposite, sessile or petioles winged to base, the bases auriculate, 20 cm or more in length, up to 16 cm wide, pinnate-pinnatifid, the lobes 5-7> acuminate, dentate, dark green above, pale green beneath, both surfaces sparingly pubescent or upper surface scabrous, upper leaves with small lobes separated by shallow sinuses or entire; peduncles puberulous to glandular-pilose; phyllaries 5» uniseriate, basally united, ovate to ovate- lanceolate, sparsely or densely pilose below, glabrous above; ray flowers about 10, their corollas yellow to reddish-purple, up to 7 mm wide, 15 mm long; disc flowers numerous, the corollas about 6 mm in length, their styles becoming long-exserted at maturity and exceeding by about 2 mm the corolla; pales of ray flowers ovate-lanceolate, about 5 mm long; disc pales about k mm long or less, their midvein prominent; fruit dark brown or black,
asymmetrically obovoid, somewhat flattened, about 3 mm long,
2.5 mm broad (Fig. 2).
Type: MEXICO: Trapiche de la Concepcion, Liebmann,
387 (Isotype FI, US!). 15 16
/f:
Fig. 2.— Polymnia oaxacana (Isotype) Scale: .28x
The principal character used in separating P. nelsonii from P. oaxacana is the color of ray corollas; those of the latter are violet, those of the former are yellow.
Examination of two isotypes of P. oaxacana as well as other specimens indicates that the ray corolla color changes with age and/or drying. The labels of certain herbarium specimens bear a note referring to the color of the ray corollas as "orange- yellow" or "orange.” However, their color when dried is as violet-purple as that of the P. oaxacana isotypes. One col- 17 lection, R. M. King 3338, had variegated corollas, portions of which were orange and the remainder distinctly violet-purple.
P. C. Standley noted on a collection label that P. oaxacana was "planted with maize" but the purpose for such planting was not listed.
Haploid chromosome numbers for P. oaxacana are reported as being "about 20" by R. M. King and "16" by B. L. Turner et al.
References to these counts are found in Table 2.
A copy of the photograph of the isotype appears above.
Specimens Examined: GUATEMALA: Alta Verapaz: Coban,
**300 ft. alt., H. von Turckheim, 187, May, 1886 (F, GH, US);
Coban, H. von Turckheim, II 130**, July, 1906 (US); Coban, 1350 m. alt., H. von Turckheim, II 1**9^» Aug., 1907 (F, NY); Coban,
H. von Turckheim, II l**9*f, Dec., 1907 (GH, US); Coban, **000 ft. alt., H. Johnson, 20, in 1919 (GH, NY, US); Coban, open hill sides and in high forest, **000 ft. alt., H. Johnson, 58 8, Sept. 5,
1920 (US); Rt. 7 2 mi. W of Santa Cruz Verapaz, R. M. King, 3338,
July 13, I960 (TEX); Ba.ja Verapaz: Rt. 5 about 1** mi. N of
Salama, R. M. King, 3288, July 11, I960 (US); 16 mi. from Salama on road to Coban, 1700 m. alt., J. G. Hawkes et al., 1932,
Nov. 11, 1958 (F). Huehuetenango: Trail between Democracia and
Santa Ana Huista, 800-1000 m. alt., J. A. Steyermark, 51306,
Aug. 25, 19^2 (F, GH, US); between Democracia and Santa Ana
Huista, 800-1000 m. alt., J. A. Steyermark, 51309, Aug. 2 5, 19^2
(F). Quiche: J. I. Aguilar, 1193, in 19*^2 (F 11 9 7857).
HONDURAS: Comayagua: Near Siguatepeque, 1050 m. alt., T. G. Yuncker et al., 5779> July 11, 1936 (US); near El Achote above plains of Siguatepeque, 13.50 m. alt., T. G. Yuncker et al., 6000
July 21, 1936 (GH, MO); Barranco de Trincheras, 1150 m. alt.,
L. 0. Williams and A. Molina, 14665, Aug. 8, 1948 (F, GH);
Barranco de Trincheras, A. Molina, 5815, Aug. 29, 1955 (F).
El Paraiso: Drainage of Rio Yeguare, 1500 m. alt., P. Herrada and L. 0. Williams, 15973, Sept. 11, 1949 (F). Intibuca: 13 nii from Esperanza on road to Siguatepeque, J. G. Hawkes et al.,
2083, Nov. 26, 1958 (F). Morazan: Drainage of Rio Yeguare,
S. F. Glassman, 1 8 5 7, July 5, 1948 (F, NY); Uyuca, 1238 m. alt.,
J. V. Rodriguez, 804, Sept. 9, 1943 (F); 3 mi. from Zamorano in pine-oak woods, 1060 m. alt., A. Molina, 10151, July 26, 1946
(F,US); slopes of La Montanita, 1530 m. alt., L. 0. Williams and
A. Molina, 10160, July 28, 1946 (F, US); lower slopes of Cerro de Uyuca, 1530-1600 m. alt., P. C. Standley and A. Molina, 4217,
Feb. 22, 1947 (F); Piedra Herrada, I65O m. alt., A. Molina, 423,
July 26, 1947 (F, US); Piedra Herrada, lower slopes of Cerro de
Uyuca, 1500-1650 m. alt., P. C. Standley, 12001, Aug. 7, 1947
(F); near Rio Agua Amarilla above El Zamorano, 1000-1200 m. alt.
P. C. Standley, 12753, Oct.-Nov., 1948 (F); La Montanita, 1400-
1750 m. alt., P. C. Standley, 14781, Nov. l8, 1948 (F); Trail between Las Flores and La Labranza, 1500-1575 alt., P. C.
Standley, 20568, June 30, 1949 (F); Cerro de Uyuca, near La
Labranza, 1600-1800 m. alt., P. C. Standley, 20707, July 2, 1949
(F); between Quebrada Grenadillo and Las Flores, 1600 m. silt.,
A. Molina, a749, July 27, 1950 (F, US); near Rio Agua Amarilla above El Zamorano, 1000-1200 m. alt., P. C. Standley, 2 6 5 8 6,
Sept. 8, 1950 (F); between Santa Lucia and Valle de Angeles,
1800 m. alt., A. Molina, 3325, Sept. 26, 1950 (F); drainage of
Rio Yeguare, A. Molina, **093, Aug. 28, 1951 (F, US). MEXICO:
Chiapas: Near San Cristobal, 7000-8000 ft. alt., E. W. Nelson,
3221a, Sept. 18, 1895 (US); between Teneapa and Yajalon, 3000-
5000 ft. alt., E. W. Nelson, 325^, Oct. 13, 1895 (US); Hacienda
Monserrata, C. A. Purpus, 9102, Sept., 1923 (GH, MO, NY); Fenis,
C. A. Purpus, 10527, May, 1925 (NY, US). Guerrero: Tixtlancingo,
Liebmann, 3 8 6, (US 1265093)* Michoacan: Dos Tetecos, Arsene, sn., Nov., 1907 (US 1159009)* Oaxaca: Totontepec, E. W. Nelson,
823, July 21-27, 189^ (NY, US).
(B) Polymnia laevigata Beadle Bot. Gaz. 25:278. 1898.
Perennial herbs up to 2.5 meters in height; root system fibrous, shallow; stems glabrous or somewhat puberulent, ir regularly purple-striped, hollow, up to 2 cm in diameter at soil level; nodes often purple; leaves light green, spicy-aromatic, thin, glabrous or puberulent, petiolate, the adaxial surface of the petiole and veins puberulent, lower leaves 5-H-pinnately- lobed, the lobes serrate, acuminate, up to 27 cm wide and ^1 cm long, upper leaves with few lobes or lobes none; peduncles pubescent and glandular, heads nodding; phyllaries of 2 types, the lower lanceolate, 1 mm wide, 2-3 mm long, the upper broadly ovate, 3 mm wide and long with minutely-bristled margins; ray flowers ^f-6, the corolla yellowish-white, tridentate, about 3 mm 20 broad and long, the tube densely pilose outside; disc flowers about 1 5 , pale yellow, the acute lobes with fimbriate margins; ray pales broadly ovate; disc pales obovate with large veins; fruit brown, 5-ribbed, about 3 long and wide (Fig. 3)»
Type: Tennessee: Franklin Co.: Cowan, C. D. Beadle,
5837, Aug. 21, 1897 (Isotype NY!).
This is the rarest of the three species of Polymnia which occur in the United States. The several specimens examined represent only seven counties in five states. I am not prepared to interpret the true geographic distribution from the apparent distribution mapped for this species. Although there are proba bly populations at low altitudes the station I visited near
Monteagle, Tennessee, is on the Highland Rim at an altitude of
1 ,0 0 0 feet.
Polymnia laevigata, like its closest relative P. cana densis, can be found growing in shade in calcareous soil.
Representative genera associated with P. laevigata about 4.8 miles southeast of Monteagle include Fraxinus, Salix, Cercis,
Solidago. Hydrangea, and Actinomeris. At this location P. laevigata is quite abundant on both sides of U. S. route 4l.
Several transplants were taken from the population referred to above and placed in the Ohio State University green house. One of these, no. 254, possessed very unusual pollen of various shapes and sizes. The term "heteromorphic pollen" was
first coined by Fisher and Wells (1 9 6 2) to describe this condition 21 \ which we believe to be the first such pollen record for flowering plants.
I have never observed any abnormal meiosis in smeared preparations of pollen mother cells of this species. A haploid chromosome number of 15 was counted in the course of this in vestigation. Experimental, studies involving this species are discussed later.
J
EarcuKnuKci,r^ .... -■ -'.-/•.-’i-R;.'i■ •
Fig. 3.— Polymnia laevigata (Isotype) Scale: .28X 22
Specimens Examined: Alabama: Tuscaloosa Co.: Ravine near Warrior River about 8 mi. above Tuscaloosa, R. M. Harper,
3091, July 30, 1933 (MO, NY, PH, US); below Lock 13, along
Warrior River about 8 mi. from Tuscaloosa, R. M. Harper et al., kkj>, (TEX). Florida: Jackson Co.: Marianna Caverns State Park,
R. K. Godfrey, 54255, Oct. 21, 1955 (US); Caverns State Park,
A. F. and R. Godfrey, sn., Sept. 8, 1961 (OS). Georgia; Walker
Co.: E slope of Pigeon Mt., P. Wilson, 170, Aug. 1, 1900 (NY,
US); E slope of Pigeon Mt., R. M. Harper, 333, Aug. 1, 1900 (NY).
Missouri: Pemiscot Co.: 3 mi* SSW of Portageville, J. A.
Steyermark, 82261, July 29, 1956 (F, MO). Tennessee: Franklin
Co.: Cumberland Mt. at Sewanee, H. Eggert, sn., Sept. 12, 1898
(MO 1767352 and MO 1767412); near Sewanee, E. J. Alexander et al., sn., Sept. 24, 1933 (NY). Marion Co.: on bluff 5 Mi* SE of
Monteagle by U. S. rt. kl, E. S. Ford and N. H. Russell, 2195,
Sept. 18, 1946 (TENN); on roadside of U. S. rt. kl 4.8 mi. SE of
Monteagle, J. R. and J. M. Wells, 254 and 255, Aug. 1, 1961 (OS).
Rhea Co.: Valley of Little Piney Creek, R. E. Shanks et al.,
14060, Oct. k, 19^9 (TENN).
(C) Polymnia canadensis L. Sp. PI. 9 2 6. 1753*
Polymnia variabilis Poiret. Encyc. Meth. Bot. Suppl. 5:505-50^T l8o4.
Polymnia canadensis L. var. discoidea Gray. Gray's Lessons in Botany and Vegetable Physiology. 248. l88l.
Polymnia canadensis L. var. radiata Gray. Syn. FI. N. Am. 1 : part 2 , 2 3 8. 1884.
Polymnia radiata (Gray) Small. Flora of the SE U. S. 13467 1 9 0 3 . 23
Osteospermum canadense House. Bull. K. Y. State Mus. 63:2^3-244. 1923.
Polymnia canadensis L. f. radiata (Gray) Fassett. Rhodoreu 34: 9 6. 1 9 3 2 .
Perennial herbs up to or exceeding 1 meter in height, root system fibrous, shallow; stems terete or angular, yellowish- green and often purple-spotted, densely glandular-pubescent, especially upward; leaves petiolate, thin, dark green above, pale green below, both surfaces glabrous to puberulent, the shape variable, lower leaves deeply pinnatifid, up to 26 cm wide, 39 cm long, lobes usually 5-7» upper leaves with fewer lobes or lobes none then leaves ovate or ovate-lanceolate, dentate to nearly
entire; phyllaries about 5» densely stipitate-glandular and sticky, broadly or narrowly ovate; ray flowers about 5j their
corollas white or pale yellow, 3 -1 0 mm wide and up to 15 mm long;
disc flowers about 3 0, 7 urn long or nearly so, their corollas
yellowish; pales of ray flowers ovate to ovate-lanceolate; pales
of disc flowers elliptic to oblanceolate, about as long as disc
flowers, parallel main veins 8 or fewer, the upper margin
ciliate; fruit dark brown to black, sometimes mottled with light brown spots, trigonous, about 2 mm wide and 4 mm long (Fig. 4).
Type: CANADA: Collected by Peter Kalm.
Most of the confusion in regard to this species has
centered around the length of ray corollas. I have treated all
forms under one taxon. This is discussed further in the section
on experimental investigations. Fig. k»—-Polymnia canadensis Scale: .3X
Polymnia canadensis is usually found in shaded and alkaline situations such as talus slopes near a stream. In Knox
County, Tennessee, I observed it growing on moist limestone almost completely devoid of soil.
The haploid chromosome number of the type variety as well as var. radiata is 15. In addition to counts which I made,
Dr. L. C. Anderson of Michigan State University sent me a camera lucida drawing with a haploid count of 15 clearly indicated. A voucher specimen also accompanied the drawing. 25
In addition to the distribution as shown on Figure 3 6,
P. canadensis has been collected in Ontario, Canada.
Specimens Examined: CANADA: Ontario: Niagara Falls
Queen Victoria Park, C-. C. Mellor and J. A. Shafer, sn., Aug.,
1889 (PH); Niagara Falls Glen, H. L. Fisher, sn., Aug. 7, 1901
(PH); Queenston Heights, Niagara, J. Macoun, 26668, July 7, 1901
(MO, NY, US). Alabama: Bibb Co.: C. Mohr, sn., June 2, 1883
(US). Warnock, C. Mohr, sn., June 10, 1895 (US). Clay Co.:
Talladega Mt., F. S. Earle, 1023, Aug. 26, 1897 (F, NY). Colbert
Co.: S of Florence, H. Eggert, sn., July 21, 1899 (MO). Jackson
Co.: North Sauty, D. Isely, 2869, July 2, 19^3 (PH); 2 mi. W of
Scottsboro, G. R. Cooley and F. Harper, ^500, Mar. 21, 1956 (GH).
Lauderdale Co.: Greenhill, Tennepec Valley, C. Mohr, sn.,
July 28, 1896 (US). Madison Co.: 7 mi» NW Huntsville, J. D.
Smith, sn., Sept. 5, 188^ (F, GH, PH,US); Monte Sano, C. F. Baker,
1355, Sept. 13, 1897 (MO, NCU, NY, OS); Monte Sano near Hunts ville, W. M. Canby et al., 113, Oct. 7, 1898 (MO, NY, US);
Huntsville, J. D. Smith, sn., Sept. 5, 19?? (GH); Huntsville,
W. Rhoades, sn., Aug., 1931 (GH); N side Monte Sano, R. M.
Harper, 3^07, June 18, 1935 (GH, MO,NY, US). Arkansas: Baxter
Co.: above Norfolk Dam, D. Demaree, 235^8, July l^f, 19^2 (MO,
NY). Benton Co.: near War Eagle, Boston Mts., D. Demaree, 6 7 8 6,
June 1, 1929 (F, TENN, TEX); D. M. Moore and W. A. Lee, 102,
Aug. 21, 1931 (TEX). Carroll Co.: Eureka Springs, N. M. Glat- felter, July 17, 1898 (MO). Conway Co.: Petit Jean State Park,
D. Demaree, 25277, Sept. 2^, 1 9 ^ (MO); Petit Jean State Park, J. E. Moore, 1123, July 28, 1955 (TENN). Franklin Co.: White
Rock Mts. near Mountainsburg, N. C. Fassett and V. M. Watts,
19803, June 12, 1938 (F). Garland Co.: Hot Springs, F. L.
Harvey, 7, July, ? (GH). Logan Co.: Magazine Mtn., H. R. Pyle,
59^1 Oct. 9, 1936 (TEX); Magazine Mtn., D. Demaree, 17715,
June 18, 1938 (F); Magazine Mts., C. Heiser and D. Demaree,
25170, July 30, 19kk (MO). Newton Co.: W. H. Emig, 73, Aug. 2,
1913 (MO). Pope Co.: Nogo, G. M. Merrill, 58 2, July 9, 19^1
(TEX). Stone Co.: Onia, De Demaree, 278 3 8, June l8 , 19^9 (TEX).
Yell Co.: Mt. Nebo State Park, D. Demaree, 20558, Aug. 30, 1939
(NY), Connecticut: New Haven Co.: Wallingford, E. B. Harger,
^89^, Sept. 15, 1905 (PH); Northford, G. H. Bartlett, sn.,
Sept. 2, 1906 (GH). Middlesex Co.: Pistapang Mt., C. A.
Weatherby, 3382, Sept. 9, 1913 (US); Pistapaus pond, Durham,
R. W. Woodward, sn., Sept. 19, 1913 (GH). District of Columbia:
L. F. Ward, sn., Oct. 22, 187O (US); High Island, L. F. Ward, sn., April 2, 1 8 7? (US 1^2^01); L. F. Ward, sn., July 27, 1879
(GH); E. S. Burgess, sn., Oct. 2, 1888 (US); E. S. Steele, sn.,
July 16, 1897 (GH); Glen Echo, E. S. Shute, sn., July 16, 1897
(US); High Island, F. J. Braendle, sn., Aug. 1 5 , 1898 (US);
E. L. Morris, 2^0, Aug. 13, 1899 (US); Va. sideof Potomac River,
E. L. Morris, 282, Aug. 25, 1899 (F). Georgia: Walker Co.:
Pigeon Mt., R. M. Harper, 361, Aug. 3, 1900 (F, GH, MO, NY, US).
Illinois: Calhoun Co.: Kampsville, L. M. Turner, 27^, June, 1930
(F). Carroll Co.:3 mi* N of Palisades Park, F. J. Hermann, 8866,
June 17, 1937 (NY); G. D. Parker, 17^5, Aug. 3, 1939 (F). Champaign Co.: Urbana, F. C. Gates, 1950, Sept. 21, 190? (US);
Brownfield Woods, A. S. Pease, sn., Sept. 23, 1909 (OH); near
Urbana, E, S. Steele, sn., Aug. 11, 1910 (US); Brownfield Woods,
G. N. Jones, 12511, July 20, 1940 (MO); Univ. woods, G. N. Jones,
14065, July 2, 1941 (MO); Brownfield Woods, G. N. Jones, l44l6,
Aug. 3t 1941 (MO). Cook Co.: A. Chase, sn., July 27, 1896 (PH);
Cook or Kane Co.: Elgin, C. Gronemann, sn., (F 55962*0; Cook or
Kane Co.: Vasey, sn., (F 467396); Cook or Kane Co.: H. C.
Benke, 1800, July 12, 1916 (F 663396). Hancock Co.: Augusta:
B. Mead, sn., Aug., 1845 (GH, MO). Henderson Co.: Oquawka,
H. N. Patterson, sn. (F 209338, F 97605, US 593719); Oquawka,
H. N. Patterson, sn., Sept., 1876 (MO); Oquawka, H. N. Patterson,
187 (GH); H. N. Patterson, sn., (MO 1767398, F 103994, F 209334,
F 257*f5*f, F 209337); Oquawka, H. N. Patterson, 6 6, (US 783870);
Woods near Oquawka, H. N. Patterson, sn., Oct., 1876 (F 46*1-207);
Oquawka, H. N. Patterson, sn., Oct., 1880 (US). Jackson Co.:
G. H. French, sn., Sept. 24, 1878 (US); Grand Tower, H. A. Gleason,
1800, Aug. 25, 1900 (GH). Jasper Co.: Near Newton, R. Ridgway,
1584, Oct. 24, 1^21 (MO). Jersey Co.: Elsah, R. Hoffman, sn.,
June 19, 1918 (MO). Jo Daviess Co.: Warren, L. M. Umbach, sn.,
July 8, 1896 (F, PH); S Galena, M. C. Lodewyks, 407» Aug. 2, 1937
(MO); Apple River Canyon State Park, J. A. Steyermark, 40768,
July 6, 1941 (F); Johnson Co.: E. J. Palmer, 16672, Oct. 3> 1919
(MO). La Salle Co.: La. Salle, sn., Sept., 1882 (MO 1767397);
Starved Rock State Park, 0. E. Lansing, Jr., 3871» July 8, 1914
(F ); Starved Rock Park, F. E. McDonald, sn., Sept., 1919 (F); Starved Rock Park, F. H. Thone, 99* June-Sept., 1921 (F, MO).
McLean Co.; Bloomington, B. L. Robinson, sn., Aug., 1886 (GH);
1 mi. W of Funk's Grove, L. M. Rohrbaugh, 123, Aug. 9j 19^9
(TEX). Monroe Co.; W. M. Bailey and J. R. Swayne, 5097» July
1^» 1952 (NCU). Ogle Co.; above Oregon, M. S. Bebb, sn., in
1872 (F). Peoria Co.: J. T. Stewart, sn., in 1869 (F ); Chil- licothe, M. Fisher, sn.,July 3» 1880 (F); Kickapoo Valley,
V. H. Chase, IOO8 3, Aug. 30, 19^8 (NY, TEX). Randolph Co.:
Prairie du Rocher, G. Engelmann, sn., Nov., 18^5 (MO). St. Clair
Co.: H. Eggert, sn., July 5» 1877 (MO); Carondolet, E. Douglas, sn., Sept. ^f, 1891 (MO); Sugar Loaf and Falling Spring, J. M.
Greenman, 3935i Oct. 5» 1918 (MO); 1 mi. Wof Falling Spring,
J. A. Steyermark, 1230, (MO). Saline Co.: Harrisburg, H. A.
Gleason, 8 6 5, July 21, 1899 (GH). Stark Co.: 3 mi. NW of Wady
Petra, V. H. Chase, sn., Aug. 15, 1895 (F); 3 mi. NW of Wady
Petra, V. H. Chase, 1536, Aug. k t 1907 (OS). Tazwell Co.:
Marshland border N of E Peoria, V. H. Chase, 11377» Aug. 2,
1950 (F). Union Co.: Wolf Lake, H. A. Gleason, 9026, June 29,
19^0 (NY); G. D. Fuller and R. Fisher, 17^, Aug. 1, 19^0 (F);
G. D. Fuller, 6 8 8, July 17, 19^1 (F). Vermillion Co.: Salt
Fork near Fairmount, G. N. Jones, 1 3 2 3 6, Oct. 6, 19^-0 (GH, MO,
TEX). Wabash Co.: Mt. Carmel, H. Shimer, sn., in 1866 (GH).
Woodford Co.: F. E. McDonald sn., Sept., 1890 (GH); F. E.
McDonald, sn., Aug., 1891 (F). Indiana: Carroll Co.: Delphi,
J. N. Rose, sn., July, 1892 (US). Howard Co.: Melfalfa Park,
C. M. Ek, 91, July 3, 1938 (US). Jefferson Co.: Hanover, A. H. Young, sn., July, 1875 (NY); Hanover, H. M. Coulter, sn., July 7,
1877 (PH); Hanover, J. M. Coulter, sn., July 11, 1877 (F).
Lawrence Co.: Tunnelton, J. Wynn, 9 6, July 30, 19^5 (TENN).
Union Co.: Liberty, J. N. Rose, sn., July 8, 1886 (F). Porter
Co.: Chesterton, J. R. Churchill, sn., Aug. 13, 1925 (MO).
Vermillion Co.: Montezuma, F. J. Herman, 8^3 6, Oct. 2, 1936
(GH, NY); across Wabash River from Montezuma, R. C. Friesner,
19^30, Aug. 11, 19^5 (NY). Warren Co.: J4 mi. W of Kramer, C. C.
Deam, sn., July 8, 1918 (PH). Wells Co.: 34 mi. NW of Bluffton,
C. C. Deam, sn., June 22, 1905 (MO, US). Iowa: Allamakee Co.:
B. Shimek, sn., Aug. 11, 1917 (MO, NY). Clayton Co.: Bixby,
L. H. Pammel, sn., Sept. 18, 1918 (MO); McGregor, L. H. Pammel,
1030, Aug. 12, 1925 (GH, MO); McGregor, L. H. Pammel, 4l, Aug.
26, 1926 (GH). Dubuque Co.: H. C. Benke, 37^1, Aug. 18, 1923
(F). NW of Luxemburg, B. Shimek, sn., July 3» 1929 (MO, NY).
Fayette Co.: B. Fink, sn., June, 1 89 ^ (GH);.B. Fink, 331» Aug. 1,
189^ (US); J. R. Garaner, 75^, July 27, 1899 (NY). Madison Co.:
Pammel State Park, M. J. Fay, ^97^, Aug. 22, 1952 (US). Van
Buren Co.: 2 mi. ESE of Bonaparte, R. A. Davidson and R. F.
Thorne, 1803, Sept. 17, 1953 (NY, TEX, US). Winneshiek Co.:
Decorah, E. W. Holway, sn., July 16, 1888 (MO); B. Shimek, sn.,
July 20, 1927. Kentucky: Adair Co.: Canoe Creek, D. Gibson,
112, July 5j 1958 (F); D. Gibson and J. Watson, Oct. 23,
1959 (F). Allen Co.: SE Scottsville, II. A. Gleason, 8882,
June 25, 19^0 (NY). Anderson Co.: Salt River SW of Lawrenceburg,
E. T. Wherry and F. W. Pennell, 1369^, Sept. 7* 1927 (PH). Bell Co.: T. H. Kearney,. Jr., ^7 8, Sept., 1893 (F, GH, MO, NCU, OS,
US). Edmonson Co.: Mammoth Cave, H. W. Lix, ^35, July 11,
19^8 (US); Brownsville near Green River, S. F. Price, sn.,
July 26, 1896 (MO). Estill Co.: W. A. Anderson, sn., Aug. 28,
19^1 (MO). Fayette Co.: Lexington, C. W. Short, sn. (NY).
Fulton Co.: Hickman, C. W. Short, sn., (PH 782V 7O ) . Harlan Co.:
Big Black Mtn., T. H. Kearney, 288, Aug., 1893 (NY). Jessamine
Co.: High Bridge, F. T. McFarland, 3227, May 1^, 1939 (MO).
Madison Co.: near Bighill, L. B. Smith and A. R. Hodgdon, 3768,
June 10, 1937 (GH, US). Mercer Co.: Bluffs of Dicks River, A.
King, 6^, June 5, 1916 (F). Pike Co.: Elkhorn Cliff on Frank fort Pike, J. W. Singer, 309, July 1, 1930 (US). Kansas:
Anderson Co.: 2 mi. S of Garnett, W. H. Horr and R. L. McGregor,
3229, June 22, 19^9 (TEX, US). Maryland: Montgomery Co.: above
Cabin John, P. Dowell, 17^9, Sept. 28, 1902 (GH); above Cabin
John, J. H. Painter, 102^, Aug. 21, 1904 (MO). Washington Co.: base of Round Top, F. Shreve and W. R. Jones, 752, July 13, 1906
(US). Michigan: Berrien Co.: F. W. Johnson, 1299, Sept. 10,
191^ (F). Ingham Co.: G. Thurber, sn., in i860 (GH); SE corner of Baker woodlot M.S.U. campus, L. C. Anderson, 2180, Sept. 17,
1962 (MSU). Jackson Co.: S. H. and D. R. Camp, sn., July
1897 (F). Minnesota: Fillmore Co.: C. 0. Rosendahl, 129^,
July 26, 1903 (NY). Goodhue Co.: Cannon Falls, J. H. Sandberg, sn., Aug., 1891 (F); Cannon Falls, J. L. Scofield, sn., July,
1893 (F). Houston Co.: along Root River, C. 0. Rosendahl, 5378,
Aug. l^f, 1927 (GH); 1.9 mi. N of Iowa-Minn. state line, J. W. Moore, 15978, Aug. 23, 1943 (GH). Nicollet Co.: C. A. Ballard, sn., July, 1892 (F, GH). Waseca Co.: Waseca, E. P. Sheldon, sn., June, 1891 (NY, US). Winona Co.: Winona, J. M. Holzinger, sn., June, 1890 (F); J. M. Holzinger, sn., Sept., 1901 (NY).
Missouri: Barry Co.: Seligman, Dewart, sn., Aug. 21, 1892 (MO).
Barry Co.: Eagle Rock, B. F. Bush, 92, Sept. 17, 1896 (MO, NY);
Eagle Rock, B. F. Bush, 155, June 22, 1897 (MO, US); Eagle Rock,
B. F. Bush, 3224, Aug. 12, 1905 (GH, MO, NY, US); Eagle Rock,
E. J. Palmer, 6291, July 17, 1914 (MO); along river, B. F. Bush,
15192, Sept. 10, 1935 (MO). Benton Co.: Big Buffalo, yfz mi. N
Zora, J. A. Steyermark, 7310, Sept. 24, 1938 (F). Callaway Co.:
SE of Fulton, J. A. Steyermark, 26130, Sept. 10, 1957 (F, MO).
Camden Co.: J. A. Steyermark, 4836, Aug. 9, 1932 (F). Carter
Co.: Current River, W. Trelease, 557, Sept. 10, 1897 (MO); 2 mi*
E of Big Spring State Park, J. A. Steyermark, 68230, June 19,
1949 (F). Christian Co.: 4 mi. SE of Chadwick, J. A. Steyer mark, 23059, July 6, 1937 (F, MO). Clark Co.: B. F. Bush, sn.,
Aug. 26, 1892 (MO); along Des Moines River SE of Dumas, J. A.
Steyermark, 68922, Aug. 13, 1949 (F); NE-facing bluffs, Des
Moines River, J. A. Steyermark, 6 8 923, Aug. 13, 1949 (F). Craw
ford Co.: on S side of Meramec River, J. A. Steyermark, 41342,
June 15, 1941 (F); 2}& mi. N of Steelville on Meramec River,
J. A. Steyermark, 4l4o8, June 16, 1941 (F). Dallas Co.: 2 mi.
SW of Windyville, J. A. Steyermark, 13733, July 18, 1934 (MO).
Daviess Co.: about 2 mi. SW of Jameson, J. A. Steyermark, 69551,
Apr. 30, 1950 (F). Dent Co.: SW of Jodwin near Shannon Co. line, J. A. Steyermark, 19405, Aug. 3, 1935 (MO); Montauk State Park,
J. A. Steyermark, 12755, Aug. 9, 1936 (MO). Douglas Co.: 2>& mi.
SE of Ann, J. A. Steyermark, 23625, July 25, 1937 (MO). Franklin
Co.: SE of Gay Summit, J. A. Steyermark, 1295, July 17, 1927
(MO). Gasconade Co.: near Mt. Sterling, Gasconade River, J. A.
Steyermark, 9151, Aug. 11, 1933 (MO); 3 mi. NW of Bay at Brown
Shanty Ferry, J. A. Steyermark, 27873, July 4, 1939 (F); 5 ml*
S of Pershing, J. A. Steyermark, 72700, Sept. 4, 1951 (F).
Greene Co.: E. M. Shepard, sn., in 1880 (GH); near Turner, P. C.
Standley, 88ll, Sept. 4, 1911 (US); near Springfield, P. C.
Standley, 9559, Aug. 28, 1912 (US). Jackson Co.: Kansas City,
K. Mackenzie, 300, July 22, 1895 (MO); Kansas City, K. K.
Mackenzie, 848, July 5, 1896 (MO); Kansas City, B. F. Bush, 848,
July 5, 1896 (US); Kansas City, K. K. Mackenzie, sn., July 11,
1900 (NY). Jefferson Co.: Kimmswick, F. Wislizenus, 182, Aug. 9,
1885 (MO); along Big River, J. K. Kellogg, 1289, Aug. 21, 1927
(MO). Laclede Co.: along Ningua River 4j£ mi. SW Eldridge, J. A.
Steyermark, 27110, June 23, 1939 (F). Lincoln Co.: 4 mi. NE
Troy, J. A. Steyermark, 26035, Sept. 9, 1937 (F, MO); near Natural
Bridge, J. A. Steyermark, 72325, Aug. 6, 1951 (F). McDonald Co.:
Noel, E. J. Palmer, sn., Aug. 8, 1908 (MO); B. F. Bush, 5030,
Aug. 8, 1908 (MO); Noel, E. J. Palmer, 4133, Sept. 5, 1913 (MO);
SE of SW City, J. A. Steyermark, 5574, May 31, 1938 (F. MO).
Maries Co.: 6-7 mi. SE of Vienna, J. A. Steyermark, 27630,
June 30, 1939 (F). Marion Co.: Hannibal, J. Davis, 1217, May,
1906 (MO); Hannibal, J. Davis, sn., July 9, 1913 (MO); Riverview Park, Hannibal, J. Davis, sn., April 26, 1915 (TEX); Hannibal,
J. Davis, 4584, July 3, 1915 (MO); Hannibal, J. Davis, 4626,
July 26, 1915 (MO); Hannibal, J. Davis, 4628, July 26, 1915 (MO,
TEX); Hannibal, J. Davis, 6l8 6, Aug. 26, 1915 (MO, TEX); Han nibal, J. Davis, 6454, Nov. 8, 11, 1915 (MO); Hannibal, J. Davis,
1583, July 6, 1916 (MO, TEX); Hannibal, J. Davis, 8912, Sept. 8,
1916 (MO). Miller Co.: along Osage River W of Capps, J. A.
Steyermark, 6 8 3 3, Sept. 20, 1938 (F). Oregon Co.: between Fidler
Spring and Eleven Points River, J. A. Steyermark, 5381, May 24,
1938 (F). Ozark Co.: J. A. Steyermark, 15569, Sept. 28, 1934
(MO). Phelps Co.: Jerome, J. H. Kellogg, 449, June 22, 1913
(MO), Pike Co.: along Miss. River, J. A. Steyermark, 25926,
Sept. 6, 1937 (F, MO); 1)6 mi. NW of La., J. A. Steyermark, 25928,
Sept. 6, 1937 (F, MO). Polk Co.: along Pomme de Terre River,
4)6 mi. NE of Bolivar, J. A. Steyermark, 27303, June 25, 1939 (F) 0
Pulaski Co.: Big Piney Bluffs, W. Trelease, 556, Aug. 12, 1897
(MO); Dixon, M. Child, sn., Feb. 23, 1934 (MO); on Big Piney River
SW of Hooker, J. A. Steyermark, 1934la, July 28, 1935 (MO); 2 mi.
NE of Waynesville, J. A. Steyermark, 19339, July 28, 1935 (MO);
near Prewitt Spring, J. A. Steyermark, 20092, Sept. 19, 1936 (MO);
NW of Hooker on Gasconade River, J. A. Steyermark, 25396, Aug. 26,
1937 (MO). Ralls Co.: 5/6 mi. V/ of Center, J. A. Steyermark,
25874, Sept. 5, 1937 (F, MO, NY, TENN). Reynolds Co.: Black
River W of Gads Hill, 10 mi. SE of Redford, J. A. Steyermark,
14219, Aug. 3, 1934 (MO). Ripley Co.: 5 mi. NE of Bennett,
J. A. Steyermark, 11914, July 17, 1936 (MO). St. Charles Co.: Mo, River E and W of hwy. 6l bridge, B. Bauer, V?9» Aug, 5, 1939
(F); ^ mi, NE Schluersburg, 3 mi* NW of Defiance, J, A, Steyer mark, 72¥f3 and 72*+51» Aug, 13, 1951 (F)* St, Louis Co,: Creve
Coeur Lake, G. Engelmann, sn., Sept., 1838 (MO); St. Louis, H.
Eggert, sn., in 1877 (GH); H. Eggert, sn., July 5* 1877 (MO).
Letterman, sn., Aug. 27, 1878 (US); Allenton, G. W. Letterman, sn., in 1880 (MO); Allenton, J. H. Kellogg, sn., Aug., 1880 (MO)
Allenton, G. W. Letterman, sn., July, 1882 (GH); Allenton, G. W.
Letterman, sn., Aug., 1883 (MO); Allenton, J. H. Kellogg, sn.,
Aug. 15, 1885 (MO); Allenton, G. W. Letterman, sn., July 30,
1890 (MO); Creve Coeur Lake, Hersford, sn., July 9* 1891 (MO);
H. Eggert, sn., Sept. 1^, 1892 (GH, MO); F. Hugel, sn., Sept. l^f
1892 (MO); Allenton, G. W. Letterman, sn., Aug., 189^ (MO, PH,
TEX); Creve Coeur, Pollard, sn., Aug. 31, 1895 (MO); Creve Coeur
Lake, Norton, sn., Sept. 17, 1898 (MO); Creve Coeur Lake, Norton sn., Nov. 23, 1900 (MO); Creve Coeur Lake, A. G. Johnson, sn.,
July *f, 1906 (MO); H. W. Edmonds, sn., Sept. 12, 1928 (MO); on
Meramec River at Mt. Ridge, J. A. Steyermark, 9^39* Sept. 12,
1928 (MO). Saline Co.: 1 mi. S of Nelson, J. A. Steyermark,
20Vf3, Oct. 11, 1936 (MO); mi. SE of Herndon, J. A. Steyer mark, 73966, July 2^, 1952 (MO). Shannon Co.: B.F. Bush, sn.,
June 28, 1888 (MO); B. F. Bush, sn., July 17, 1891 (MO); along
Jack's Fork of Current River, 6 mi. NW of Monteer, J. A. Steyer mark, 26910, June 1939 (F). Stone Co.: Galena, E. J.
Palmer, 4-659, Oct. 14-, 1913 (MO); near Baxter, J. A. Steyermark,
19533, Aug. 22, 1935 (MO); B. F. Bush, 15202, Sept. 11, 35
1935 (MO). Taney Co.: 1 mi. S of Swan, W. Trelease, 5^2, in
1897 (MO); Swan, W. W. Eggleston, 12258, Oct. 7-9, 1915 (US);
3 mi. S of Kissee Mills, J. A. Steyermark, 22733* June 28, 1937
(F, MO); 2 mi. N of Mincy, J. A. Steyermark, 5545, May 29, 1938
(F, MO). Texas Co.: along bluffs of Jack's Fork of Current
River (MO). Washington Co.: Big River near Bliss, J. A.
Steyermark, 1 8 3 8, July 31, 1932 (F). Wright Co.: Bryant Creek
SE of Cedar Gap, J. A. Steyermark, 23668, July 26, 1937 (F, MO).
New York: Broome Co.: Binghampton, C. F. Millspaugh, 885
(F 18707)* Cattaraugus Co.: Collins, A. E. Perkins, sn., Aug.
29, 1926 (NY); Collins, A. E. Perkins, sn., Aug. 10, 1930 (US).
Cayuga Co.: Lower Paine's Creek Ravine, A. J. Eames et al.,
13155, Aug. 3, 1.919 (GH). Chemung Co.: T. F. Lucy, 1379,
Aug. 13, 1882 (MO); Elmira, T. F. Lucy, 5977, Sept. 25, 1894
(NY); T. F. Lucy, 13764, Aug. 9, 1898 (F, NY). Erie Co.:
Buffalo, G. W. Clinton, sn. (NY); Niagara Falls, W. H. Seaman,
sn. (US 78778^); Buffalo, I. C. Martindale, sn., in 1866 (PH
583925)* Genesee Co.: Bergen swamp, E. D. House, sn., Aug. 15,
1917 (GH); Bergen swamp, K. M. Wiegand, IO98O, Aug. 17, 1918
(GH); W. C. Muenscher, 15998, Aug. 30, 1924 (GH). Madison Co.:
Chittenango ravine below falls, K. M. and M. C. Wiegand, 1281,
Aug. 10, 1913 (GH); Chittenango Falls, F. E. and A. K. Harrison,
sn., June 12, 1936 (GH). Monroe Co.: Riga Swamp, W. A. Matthews,
4l83, July 13, 1940 (NCU). Niagara Co.: T. F. Allen, sn., Aug.,
1807 (NY); Niagara, W. Cooper, 1 8 2 8, (NY). Onondaga Co.:
Syracuse, Middlesex Institute, sn. (GH); F. C. Straub, sn., in 1891 (US 329910); Quarry in Onondaga, N. Hotchkiss, 1824, Sept.
29, 1924 (US). Ontario Co.: J. Hall, sn., 1828-1834, (F
476528). Seneca Co.: Sheldrake Point, N. W. Folwell, 619,
(GH, US 1802474); Ovid, Brewer and Chickering, sn., Aug. 23,
1858 (MO). Tompkins Co.: Ithaca, E. Schuyler, sn. (PH 715289); sn., Aug. 7, 1875 (US 266782); F. V. Coville, Sept. 25, 1884
(US); Enfield, A. J. Eames, 5262, Sept. 5, 1915 (GH); Enfield
Gorge, W. C. Muenscher and A. R. Bechtel, 713, Sept. 4, 1920
(US).’ Yates Co.: H. P. Sartwell, sn., (F 476010); Pennyan,
Buckley, sn., (MO 1767427). North Carolina: Near Hot Springs,
57l4d, Aug. 11, 1898 (NY); 6.4 mi. NW of Hot Springs on French
Broad River, H. E. Ahles and J. A. Duke, 46426, July l4, 1958
(NCU); 3.1 mi. SE of Trust on NC rt. 63, H. E. Ahles and J. A.
Duke, 50540, Oct. 8, 1958 (NCU). Mitchell Co.: 2 mi. from Red
Hill on NC rt. 26, J. A. Duke and H. E. Ahles, 47228, July 23,
1958 (NCU). Yancey Co.: 1.4 mi. ENE of Sioux road to Huntdale,
H. E. Ahles and J. A. Duke, 50800, Oct. 9, 1958 (NCU). Ohio:
Adams Co.: edge of Woodland near Serpent's Mound, G. W. and
H. L. Hall, 2092, Sept. 10, 1956 (OU). Ashtabula Co.: L. E.
Hicks, sn., Sept. 5» 1931 (OS). Athens Co.: W. A. Kellerman, sn., Aug. 11, 1892 (OS). Auglaize Co.: St. Marys, A. Wetzstein, sn., Sept. 23, 1896 (OS); Wapokoneta, W. Kayser, sn., in 1929
(OS). Belmont Co.: Barnesville, E. E. Laughlin, 976, July 30,
1907 (GH); Barnesville, E. E. Laughlin, sn., Aug. 19, 1910 (OS);
Rt. 147, about 4 mi. W of Bellaire, C. Weishaupt, sn., Sept. 7,
1955 (OS). Champaign Co.: V/. C. Werner, sn., Sept. 10, 1892 (OS); Cedar Swamp, E. C. Leonard, l877» Oct. 7, 1917 (US); 5 mi.
NE of Tremont, G. M. Leonard, 5571, July 24, 1925 (US); 5 mi. SE of Thackery, E. C. Leonard, 16091, Sept. 7, 1931 (US). Clermont
Co.: C. Mohr, sn. (US 783869); Loveland, J. F. James, sn. (OS
389OO). Clinton Co.: V/z mi. NE of Clarksville, E. E. Terrell,
2508, Sept. 12, 1952 (OS). Cuyahoga Co.: Chippewa Creek near
Cuyahoga River, E. M. Herrick, sn., July 5, 1957 (OS). Delaware
Co.: Lewis Center, R. Crane, 3083, Aug. 25, 1928 (NY). Liberty
Twp., J. Beatley, sn., Jan. 11, 1947 (OS). Erie Co.: Cedar
Point, R. F. Griggs, sn., Aug. 4, 1900 (OS). Franklin Co.:
Sullivant, sn. (GH); E. E. Bogue, sn., July 9, 1892 (OS); W. A.
Kellerman, sn., Aug. 11, 1895 (OS); Hayden Falls, E. C. Leonard,
1877, Oct. 12, 1915 (US); Hayden Rd. ravine, J. R. Wells, 229,
Feb. 26, 1961 (OS). Geauga Co.: SE of Bainbridge Race Track,
E. M. Herrick, sn., July 6, 1957 (OS). Greene Co.: John Bryan
State Park, D. Demaree, 11524, Aug. 18, 1935 (GH, MO, OS. US).
Hamilton Co.: C. G. Lloyd, sn., July 18, l88l (GH, PH); Cin
cinnati, C. G. Lloyd, 5977, Aug. 12, 1890 (F, MO); W. H. Aiken,
sn., Aug. 1, 1897 (OS). Hancock Co.: mi. W of rt. 698 and
1 mi. N of rt. 30 in Van Buren Twp., C. Weishaupt, sn., June 16,
1961 (OS). Hardin Co.: L. C. Riddle, sn., June 7, 1899 (OS).
Highland Co.: V/. A. Kellerman, sn., July 13, 1899 (OS); Paint
Creek, A. Cronquist, 3997, Aug. 8, 1945 (NY). Huron Co.:
Greenfield Twp., J. A. Sanford, 1379, Sept. 1, 1877 (OS); New
London, J. R. Watson, sn., Aug. 25, 1895 (F); Olena, 0. E.
Jennings, sn., Aug. 16, 1901 (OS). Jefferson Co.: Mt. Pleasant, W. A. Kellerman, sn., June 21, 1900 (OS). Licking Co.: St.
Albans Twp. on bank of Lobdell Creek 1 mi. N mi. E of Alexand
ria, C. Weishaupt, sn., July 30, 1957 (OS). Lorain Co.: Oberlin,
E. S. Steele, sn., July, 1888 (OS); Oberlin, A. E. Ricksecker,
sn., July 25, 1895 (OS, US). Lucas Co.: Toledo, J. A. Sanford,
sn., in 1877 (OS). Miami Co.: Ludlow Falls, S. E. Horlacher,
sn., July l8, 1903 (OS). Monroe Co.: Clarington, W. A. Keller
man, sn., June 26, 1900 (OS). Ottawa Co.: Catawba Island, E.
Classen, sn., in 1891 (OS); Catav/ba Island, W. A. Kellerman, sn.,
July 3, 1895 (OS); Port Clinton, E. L. Moseley, sn., Aug. 1^,
1895 (US); Green Island, D. J. Pinkava, 95, June 28, 1958 (OS);
Green Island, K. Leon, sn., June 28, i960 (OS); Middle Bass
Island, T. E. Fisher, 2071, June 10, 1961 (OS). Portage Co.:
D. M. Brown, sn., July 7, 19^0 (OS); Rt. 303 2 mi. W of Streets-
boro, E. M. Herrick, sn., June 25, 1957 (OS). Ross Co.: Bain-
bridge Caves, R. R. Deisbach, IO8 7, Aug. l*f, 1922 (F); Huntington
Twp., Bartley and Pontius, 79, July 10, 1936 (NY). Summit Co.:
Rt. ik Vfi mi. SE of Twinsburg, E. M. Herrick, sn., June 27, 1957
(OS). Warren Co.: 5 mi. E of Lebanon, C. A. Dambach, sn., Aug.
5, 1937 (OS). Oklahoma: Adair Co.: 23 mi. NE of Tahlequah on
rt. 10, C. S. Wallis, 7586, July 13, 1958 (TEX). Cherokee Co.:
Hanging Rock Camp 16 mi. NE of Tahlequah, V. Gibson, 3 6, June 26,
19^2 (NCU); 9 mi. NE of Tahlequah, J. F. Logan, 111, Oct. 5,
19^6 (TEX); 3.5 mi. NE of Tahlequah on rt. 10, C. S. Wallis, 176,
Aug. 30, 1950 (TEX); 3 mi. NE of Tahlequah, C. S. Wallis, 7559,
July 13, 1958 (TEX). Mayes Co.: 2.8 mi. NE of Locust Grove on rt. 82, C. S. Wallis, 765^, July 20, 1958 (TEX). Leflore Co.: near Page, G. W. Stevens, 278^, Sept. 10, 1913 (GH). Pennsylvania
Bradford Co.: Towanda, Brown, sn., June 10, 1897 (PH). Lan caster Co.: Safe Harbor on bank of Susquehana, A. P. Garber, sn«, Sept. 19, 1868 (NY, US); Pond Hill, J. K. Small, sn., Aug.
16, 1889 (F); Banks of Susquehana at Safe Harbor, A. E. Urban, sn., Aug. 20, 1922 (GH); Ravine at Shenk's Ferry, F. A. Tanger, sn., Aug. 15, 1938 (GH). Luzerne Co.: A. A. Heller, sn., Aug.
16, 1889 (GH, PH); Lily Lake, J. K. Small, sn., Aug. 15-16, 1889
(GH). Lycoming Co.: Ralston, B. Meridith, sn., July 1, 1898
(PH). York Co.: near Marietta, T. C. Porter, sn., (F 3 18276);
T. C. Porter, sn., Oct. 6, i860 (GH); sn., Feb. 20, 1895 (PH).
Tennessee: Campbell Co.: J. K. Underwood and A. J. Sharp, 151j
July k, 1933 (TENN); along Powell River, Kelly, 2^50, July l8,
193^ (TENN). Cannon Co.: near Woodbury, H. K. Svenson, 9280,
July ^f, 1938 (GH). Carter Co.: Forge Hill, A. B. Grindstaff, sn., July 2, 1956 (TENN); Sinking Creek, J. Pearman, sn., June
23, 1956 (TENN). Cheatham Co.: Kingston Springs, II. K. Svenson,
51, June 2kt 1922 (GH); Ashland City, D. Demaree, ^5799» June 30,
1962 (OS). Cocke Co.: Newport, Rugel, sn., Aug., 18^-2 (NY).
Davidson Co.: Nashville, A. Gattinger, sn., (F 88370); Nashville,
C. Williamson, sn., July, 1897 (PH); Banks of Cumberland River,
? W. W. Ashe, 571^, Aug. 11, 1897 (GH, NCU, NY, OS,US); near
Donelson, H. K. Svenson, 9^5^ 1 July 2 5, 1938 (GH). Franklin Co.:
I-I. Eggert, sn., July 8, 1897 (M0); Cowan, 571^c, Aug. 21, 1897
(NY). Giles Co.: between Bethel and Minor Hill on limestone bluff, R. E. Shanks et al., 9177, Sept. 12, 1949 (TENN). Grainger
Co.: Buffalo Springs preserve, Tampico, A. W. Morrison and Brown,
70, July 2, 1936 (TENN). Hancock Co.: between Sneedville and
Mooresburg, M. B. Wilson, 3972, July 14, 1935 (TENN). Hawkins
Co.: Chestnut Ridge, J. Wolfe, 19126, Aug. 9, 1955 (TENN). Knox
Co.: T. H. Kearney, Jr., sn., Aug., 1891 (OS); Knoxville, A.
Ruth, 4016, July, 1897 (NY); A. Ruth, sn., Sept. 10, 1897 (GH);
Knoxville, A. Ruth, 640, Sept., 1898 (NY); Knoxville, A. Ruth,
6 6 5, Sept., 1898 (MO, US); H. M. Jennison et al., 1310, June l4,
1929 (GH, MO); Univ. of Tenn., A. J. Sharp, 1735, June 30, 193^
(TENN); near Mascot and American Limestone Co., A. J. Sharp, et al., 24879, June 30, 1958 (TENN); Cherokee bluffs .2 mi. from
Alcoa hwy. bridge, J. R. and J. M. Wells, 249, July 30, 1961 (OS).
Lawrence Co.: near Fall River, A. J. Sharp et al., 9699, July 12,
1948 (TENN). Lincoln Co.: near Elora, 5714a, July 31, 1899 (GH).
Marion Co.: Hales Bar Dam,' A. J. Sharp et al., 48-71* June 16,
1948 (TENN). Maury Co.: SE of Culleoka, R. E. Shanks et al.,
5 8 1 3, Aug. 8, 1947 (TENN). Monroe Co.: Little Tenn. River below
Skona Lodge, A. J. Sharp and F. Galyon, 21613, Oct. 13, 1956
(TENN). Montgomery Co.: Clarksville, A. H. Howell, 624, July 20,
1910 (US). Rutherford Co.: Fosterville, 571^, Aug. 19, 1897
(GH, MO, NCU, NY, OS, US). Rutherford Co.: A. Ruth, 53, Aug. 19,
1898 (GH); 10 mi. SE of Murfreesboro, H. R. DeSelm, 7^6, July 16,
1955 (TENN); on rt. 99 between Murfreesboro and Rockvale, A. J.
Sharp and R. A. Pursell, 25485, Oct. 11, 1958 (TENN); Eagleville,
D. Demaree, 45793, June 29, 1962 (OS). Sevier Co.: Gatlinburg, 7 mi. trail, G. 3. Miller, 2252, Oct. 15, 1 9 ^ (US). Sumner Co.:
Bledsoe Creek E of Gallatin, A. J. Sharp and E. Clebsch, V7-*f69,
July 12, 19^7 (TENN). Unicoi Co.: Nolichucky River, Erwin, A. J
Sharp et a l ., 3157, Aug. 19, 193^ (TENN); Nolichucky River,
Erwin, A. J. Sharp and J. K. Underwood, 3219, Aug. 18, 193^
(TENN); Nolichucky River near Erwin, A. J. Sharp and J. K. Under wood, 3220, Aug. 19, 193^ (TENN); Unaka Springs between Long and
Short Branch, C. Lyle, 19636, July 7, 1955 (TENN). Van Buren-
Bledsoe Co.: H. H. Iltis, 339^, May 2, 19^7 (TENN). White Co.:
Below Great Falls Dam near Webb's Camp, A. J. Sharp et al., 5053,
July 31, 19^7 (TENN). Wilson Co.: Lebanon, D. Demaree, V7731,
June 22, 1962 (OS). Vermont: Rutland Co.: W. W. Eggleston,
1375, Sept. k, 1899 (GH, NY, US); W. W. Eggleston, 2 0 3 6, Sept. 3,
1900 (MO, PH); N. F. Flynn and E. B. Chamberlain, sn., July 17,
1901 (US); E. M. Kittredge, sn., Sept. 10, 192*f (PH). Virginia:
Arlington Co.: F. Blanchard, sn., Aug. 23, 1891 (MO); F.
Blanchard, sn., July 17, 1892 (MO). Carroll Co.: on Farmer Mtn.
on New River, J. K. Small, sn., July 12, 1892 (F, GH, MO).
Fairfax Co.: above Chain Bridge, P. Dowell, 3070, Aug. 6, 190^
(GH); opposite Plummer Island, P. Dowell, 5397, Aug. 1, 1908
(GH). Frederick Co.: Cedar Creek, Middleton, F. W. Hunnewell,
11233, Aug. 5, 1929 (GH). Giles Co.: below Cascades of Little
Stony, H. P. Totten, 1, July 7, 1936 (NCU); Cliffs along New
River, J. M. Fogg, Jr., 13027, Aug. 5, 1937 (MO); along New River
J. M. Fogg, Jr., 1^642, June 29, 1938 (GH). Greene or Page Co.:
Summit of Bear Fence Mtn., E. H. Walker, 2952, July 28, 19^0 (US) Lee Co.: on the pinnacle, J. K. Small, sn., July 27» 1892
(F, GH). Loudoun Co.: R. McVaugh, 6617, Sept. 6, 1942 (NY);
River opposite Brunswick, L. B. Smith, 5510, Aug. 4, 1951 (US).
Madison Co.: Shenandoah Nat'l. Park, E. H. Walker, 2^99, May,
1934 (US); Whiteoak Canyon, F. J. Hermann, 14843, Aug. 22, 1958
(US). Montgomery Co.: A. H. Curtiss, sn., Aug., 1872 (F, MO); sn., Aug. 1887 (MO 1767^25). Nelson Co.: Wallace, 2475, (F
376260). Page Co.: Stony Man Mtn., W. Palmer and W. H. King,
123, Aug. 24, 1901 (NY, US). Page or Rappahannock Co.: Marys
Rock Trail, E. H. Walker, 2347, May 9, 1938 (US). Rappahannock
Co.: Marys Rock Trail, E. H. Walker, 2432, June 19, 1938 (US);
F. R. Fosherg, 33229, Oct. 22, 1950 (NY). Rockbridge Co.:
Natural Bridge, E. B. Bartram, sn., Sept. 12, 1907 (PH); J. R.
Churchill, sn., Aug. 11, 1924 (GH, MO); near Lexington, J. R.
Churchill, sn., Aug. 21, 1924 (MO). Warren Co.: near Front
Royal, G. S. Miller, Jr., sn., in 1897 (US). West Virginia:
Fayette Co.: Gauley Bridge, W. W. Eggleston, 5544, Oct. 17-18,
1909 (GH, MO, NY); W. V a . Agr. Exp. Stn., C. F. Millspaugh, 603
(NY); Fayetteville, L. F. and F. R. Randolph, 1304, Aug. 14, 1922
(GH). Greenbrier Co.: White Sulphur Springs, A. Brown, sn.,
July 16, 1892 (NY). Hardy Co.: Wardensville, W. Va. Univ.
Biol. Expedition, sn., Aug. 13, 1930 (GH). Jefferson Co.:
Harpers Ferry, J. D. Smith, sn., Oct. 1, i860 (US). Mason Co.:
Rt. 33 near New Haven, J. R. and J. M. Wells, 234, July 22, 1961.
Ohio Co.: Wheeling, H. N. Mertz, sn., Oct. 5, 1878 (NY); A.
Eliver, sn., in 1909 (PH 584019); H. N. Mertz, sn., Oct. 5, 1878 (NY); Wheeling, A. MacElwee, sn., July 25, 1909 (PH). Pendleton
Co.: below Hermit Island, E. E. Berkley, 1524, Aug. 8, 1930 (MO)
Circleville, Nelson Gap, H. A. Allard, 2156, Aug. 29, 1936 (US);
H. A. Allard, 3 5 8 9, Aug. 21, 1937 (GH, US). Raleigh Co.: 1 mi. below Richmond Farm near Hinton, J. P. Tash, 6 6 3, Aug. 23, 1910
(GH). Wisconsin: Brown Co.: Ashwaukenon, J. H. Schuette, sn.,
July 26, 1881 (GH); Flintville, J. H. Fauette, 70, June 22, 1887
(F). Calumet Co.: Brillion, H. C. Benke, 1263, in 1914 (US).
Crawford Co.: Prairie du Chien, N. C. Fassett, 4498, Aug. 19,
1927 (GH). Dane Co.: Madison, T. I. Hale, sn., (GH). Dodge
Co.: Neda, N. C. Fassett et al., sn., July 4, 1932 (GH). Grant
Co.: Glenhaven, N. C. Fassett, 13037, Sept. 10, 1930 (F, GH);
Glenhaven, N. C. Fassett, 12641, Sept. 6, 1930 (MO). Milwaukee
Co.: H. E. Hasse, sn., July 31,i 1880 (NY). Sauk Co.: Baraboo,
R. H. True, sn., Aug. 1, 1892 (GH); Devils Lake, N. C. Fassett,
2753, Sept. 17, 1925 (GH); J. M. Greenman, 5037, June 23, 1939
(MO).
(D) Polymnia connata (Sprengel) Blake. Cont. U. S. Nat. Kerb. 26:238. 1930.
Gymnolomia connata Sprengel. Syst. Veg. 3s6l0. 1826.
Polymnia silphioides DC. Prodr. 5s516. 1 8 3 6.
Polymnia macroscypha Baker ex Martius. FI. Bras. 6, part 3, 1 5 8. l884. (Syntype FI).
Polymnia silphioides DC. var. perennis Bettfreund. FI. Arg. 2:116-117. 1900.
Polymnia andrei Arechavaleta. Ann. Mus. Nac. Montev. 1 : series 2, 35-38. 1905. hk
Annual or perennial herbs up to 3 meters in height, aromatic; stems angled, hispid or pilose; leaves sessile or petiole winged to base, the bases connate by their auriculate bases, up to ^0 cm long and 3^ cm wide, above scabrous or strigose, below puberulent to pilose, often resin-dotted; upper leaves ovate-lanceolate, serrate or dentate; lower leaves tri angular, acuminate, the bases hastate or truncate, abruptly tapering to the lower winged portion, the lobes coarsely and finely dentate, peduncles densely glandular-pilose, glandular, or pilose; phyllaries about 5» ovate to lanceolate, basally connate, glandular or pilose below; ray flowers in 2 series of about 20 flowers each, corollas yellow, 8 mm long or less, about
2 mm wide, the pilose tube about 2 mm long; disc flowers 3 mm long, about 30 or more, upper portion of corolla often dark- purple when dried, pilose outside; pales of ray flowers concave, lanceolate, pilose below, glabrous above, about 7 mm long, acumi nate to long-acuminate; disc pales membranceous, 7 mm long or less, lanceolate, the margins pilose; fruit dark brown, com- pressed-obovoid, striate, up to 7 mm long and 5 mm wide (Figs.
5 and 6).
Type: BRAZIL: Sello, Sprengel Herb. No*. 1791*
A copy of tfyfe photograph of the type specimen appears below.
The type is irk,the Schultz Bipontius herbarium. Blake
(1930) points out that the type is labelled "Monte Video" although the type was described as being from Brazil, Sello. Fig. 5»— Polymnia connata (Holotype) Scale: .50X
Because of the poor quality of a small, original photo graph of the type specimen I have included a photograph of another herbarium specimen (Fig. 6) of P. connata.
As the specific epithet implies, the leaves are basally
connate by their auriculate bases. Other species appear to have basally connate leaves but close inspection will reveal that the auricled bases are separate but often overlapping. A character istic of this species, unique in Polymnia, is the biseriate
condition of the ray flowers. b6 A
Fig. 6.— Polymnia connata Scale: .28x
The tuberous underground portions of this species as in
P. sonchifolia are used as a source of human food. If hybrids between these two species are possible perhaps an improved food source could be obtained by extensive breeding programs.
Specimens Examined: ARGENTINA: Buenos Aires: Partido de La Plata, Palo Blanco, A. Krapovickos, 2700, Nov. 30, 19^5
(NY, US). Corrientes: Cept. San Martin, Yapeya, A. M. Huidobro,
3726, Dec. 18, 19^6 (F); Dept. San Martin, Tapebicua, A. M.
Huidobro, 3788, Dec. 18, 19^6 (F). Eva Peron: Alrededores de La Plata, Los Talas, A. L. Cabrera, 2784, Mar. 26, 1933*
Mislones: Dept. Candelaria, Bertoni, 2184, Oct. 13, 1945 (F);
Dept. Cainguas, Puerto Leoni, G. J.Schwarz, 1572, Nov. 27,
194-5 (F); Dept. Iguazu, Puerto Experanza, J. E. Montes, 94-25,
Oct. 15, 1950 (NY). Tucuman: S. Venturi, 152 (US); Dept.
Graucas, Bid de la Cuesta, alt. 1700 m., S. Venturi, 4-121,
Apr. 20, 1926 (US); Cerro Nogalito, alt. 1200 m., S. Venturi,
8798, Apr. 12, 1929 (US). BBA3IL: Santa Catarina: 33 km SE
of Campos Novos on road to Sao Jose do Cerrito, alt. 700-900 m.,
L. B. Smith and R. Klein, 11204, Feb. 10, 1957 (US). Rio de
Janeiro; J. Weir, 505, 1861-2 (F); Sao Paulo: J. Weir, 505,
1861-2 (F). PARAGUAY: Alto Parana: K. Fiebrig, 5728, in 1909
(US). Guaira: Villarca, P. Jorgensen and E. Hassler, 4444-,
Jan., 1932 (NY). URUGUAY: Colonia: Bank of San Juan, H. H.
Bartlett, 21260, Dec., 1943 (US). Rocha: 25 km N of Castillos,
H. H. Bartlett, 21385, Jan. 22, 1944 (US). Treinta y Tres:
Vergara, W. G. Herter, 94100a, Dec. 1933 (F, NY, US).
(E) Polymnia jelskii Hieronymus. Engl. Bot. Jahrb. 36:484. 1905*
Since the only material examined was a small fragment
of the type specimen the species description is that taken from
the original description by Hieronymus.
A shrubby tree about 2 meters in height; branches sulcate-
angled, villous, hairs jointed, rusty-colored, about 4 mm long;
lower leaves cordate-ovate, some cuneate, petiole winged and
narrowly decurrent, acuminate, membranaceous, triplinerved, thickly pubescent below chiefly on veins; petiole winged (wing about 3 nini wide on both sides) about 8 cm long, leaves up to 47 cm long and 25 cm wide, upper leaves smaller, almost sessile, ovate, long-acuminate; inflorescence corymbose, many-headed, the heads small; peduncles 1-3 cm long, about 1 mm thick; bracts ovate, up to 8 mm long and 3 mm wide, short-acuminate or almost blunt, involucral scales ovate, 5-6 mm long, 3^-4 mm wide, 11-13- nerved, margins ciliate, inner several boat-shaped; ray flowers
10-12, corollas biligulate, outer ligules quadrate-ovate, about
2j£ mm long, 1)4 mm wide, apex tridentate, 7-nerved; inner ligule
1)4 mm long, J4 mm wide, 1-nerved, tube about 1 mm long, villous; disc flowers about 3 0, corollas about 3 mm long; fruit glabrous, dark brown compressed obovoid, 4-angled, outer angles narrowly winged (Figs. 7 and 8).
Type: PERU: near Callacate, Const, de Jelski, 687, May,
1879 (only fragments of Holotype TJSl).
The type specimen pictured below is in the Berlin Her barium. Figure 7 is a copy of a photograph of the type in the
Field Museum Herbarium. Figure 8 is a photograph of the fragments of the holotype which I examined.
I have noted that the peduncles are densely villous and that the hairs are light brown. Fruit size is 3 mm long and 2 mm wide. The outer angles of the fruit are winged. Disc pales are 3-lobed and have a single main vein.
Hieronymus in describing P. .jelskii used the term
"biligulate" in referring to the ray corollas. I interpret 49 I
jy l«» 8»y»tr‘'
UNITED STATES NATIONAL MUSEUM
Fig. 7»— Polymnia .jelskii (Holotype) Scale: ,24x biligulate to mean two separate and complete corollas attached at the top of a single fruit.
This interpretation is based on observations of a fruit and associated flower parts examined from fragments of the type material. At the top of the fruit appeared two widely separated ligule-shaped structures about 2 mm long. The inner (adaxial) of these two was obviously corolla-like in structure and texture.
The outer "ligule" was about the same size and only slightly more coarse in texture. I suggest that this outer "ligule" is morphologically an extension of the pericarp, i.e., of the same 50
Fig. 8. — Polymnia jelskii (Holotype fragments) Scale: 2X layers of cells as those which make up the wings of the fruit.
If this is true then we are not really observing two separate corollas (i.e., two sets of petals, as Hieronymus probably thought) but a single corolla and a ligule-shaped extension of the pericarp.
This species appears to be well-defined although it bears
certain affinities to P. pyramidalis or some species in the P.
glabrata complex. 51
(F) Polymnia siegesbeckia DC. Prodr. 5:516. 1 8 3 6.
Annual herbs to 2 meters in height; stems often orange- yellow when dried, glabrous below, stipitate-glandular above or glandular and pilose; leaves sessile or petioles winged to base, the lower basally connate, deltoid, acuminate, the base cuneate, puberulent or strigose above, sparsely puberulent below; peduncles stipitate-glandular or glandular and pilose; phyllaries usually 5, ovate to ovate-lanceolate, about 7 mm long, up to 5 mm wide, below glandular, glandular-puberulent or glabrous; ray flowers about 1 5 , in 1 series, corollas yellow, 5 -7 mm long, 2 mm wide or less; disc flowers about the same number as the ray flowers or fewer than the ray flowers, 5 -6 mm long; pales of ray flowers ovate, long-acuminate, concave, about 5 mm long, margins of long-acuminate tip inrolled, stipitate-glandular below, glabrous above; pales of disc flowers lanceolate, about 5 mm. long; fruit dark brown to black, obovoid, longitudinally striate, up to 3 mm wide and ^ mm long (Figs. 9 and 10).
Type: BRAZIL: Rio Janeiro, A. Gaudichaud, 6 9 2, 183 ^
(type not seen, Holotype fragments Fl).
The type specimen is in the Delessert Herbarium. Figure
9 is a copy of a photograph of the type specimen.
Polymnia siegesbecki& is a relative of P. riparia.
However, there are several distinct; differences between the two.
In P. riparia the overall size of the plant is greater and there are more disc flowers than in P. siegesbeckia. The pales of ray
flowers are attenuate in both species but the attenuate portion 52
Fig. 9»— Polymnia siegesbeckia (Holotype) Scale: .36 of the pales in P. riparia commonly exceeds the entire length of the ray pale in P. siegesbeckia. Another among the many dif ferences is that P. riparia is a perennial and P. siegesbeckia is an annual.
In Peru "yarita" is a common name of P. siegesbeckia.
The outline of some lower leaves of P. siegesbeckia is shown in Figure 10.
Specimens Examined: BOLIVIA: Santa Cruz: La Cruz, 0.
Kuntze, June, 1892 (NY, US). BRAZIL: Minas Gerais: Caldas, 53
uttixa t’tiu aattouimvmot
n.»U f StatllUata* ■*wt. W- M / ‘ /,/ ? *,/...<
Fig. 10.— Polymnia siegesbeckia Scale: .31X
A. F. Regnell, I 195 (US 1^02098). Parani: Ponta Grossa, P.
Dusin, sn., Jan. 3» 1910 (US); Ponta Grossa, P. Dusln, 9503*
June 3, 1910 (NY, US). Rio de Janeiro: A. Glaziou, 15071*
Aug. 6 , 1885 (US); J. Miers, sn., (US 1^20709); G. Gardner, 253,
Aug.-Nov., 1836 (GH, NY). PARAGUAY: Guaira: Villarrica, P.
Jorgensen, (F 698OO9 , US 15^0955)• PERU: Cuzco: San
Miguel, Urubamba Valley, l800 m. alt., 0. F. Cook and G. B.
Gilbert, 892, May 25, 1915 (US); Junin: Rio Pinedo N of La
Merced, 700-900 m. alt., 2359^, May 30, 1929 (NY, US). San Martin: Zepelacio, near Moyabamba, 1200-1800 m. alt., G. Klug,
3399, Dec., 1933 (NY, US).
(G) Polymnia riparia H. B. K. Nov. Gen. et Sp. PI. *f:282-283. 1820.
Perennial herbs or shrubs to k meters in height; stems glabrous to densely villous; leaves up to 3^ cm wide, 50 cm long, sessile or petioles winged to base, the winged portion gradually becoming narrower toward base, lower leaves deltoid to broadly ovate, lobes often small then sinuses broad and shallow, margin dentate, the teeth callous-tipped, upper surface scabrous to sparsely pilose, lower surface densely pilose to sparsely veluti- hous, upper leaves ovate to ovate-lanceolate or rhombate; peduncles glandular or pilose with glands; phyllaries about 5, broadly ovate or ovate-lanceolate, 12 mm long or less, up to 10 mm wide, glabrous or sometimes puberulous to glandular-pilose; corollas of ray flowers yellow, about 10 mm long, up to 6 mm wide, the tube pilose especially on lower portion or pilose with
glands; disc flowers about 6 mm long, corollas yellow (sometimes purple, at least when dried), puberulous outside; pales sub
tending ray flowers ovate-lanceolate, the upper margins involute
resulting in a spear-shaped point up to 1 cm long above the
broadened basal portion, lower surface of ray pales with large
stalked glands up to 1 mm long, pilose hairs sometimes occurring
with the glands; pales of disc flowers about as long as disc
flowers, lanceolate; fruit, flattened-obovoid, dark brown, striate,
about 3«5 nun long, 3 mm wide (Figs. 11 and 12). 55
Fig. 11.— Polymnia riparia (Holotype) Scale: *30X
Type: COLOMBIA: On banks of Magdalena River between
Cascajo and Angostura de Carare, Humboldt, l6*f0.
Figure 11 is a copy of a photograph of the holotype in the Berlin Botanical Museum.
Figure 12 is a photograph showing typical leaves at upper nodes as well as the elongate, filiform pales of the ray flowers.
The attenuate pales of the ray flowers are uniquely
characteristic of P. riparia and P. siegesbeckia. These two
species seem to be closely related. However, P. riparia has two or three times as many disc flowers as P. siegesbeckia. Their 56
Fig. 12.— Polymnia riparia Scale: ,J2X range of geographic distribution overlaps in Bolivia where variants occur.
In Panama P. riparia also seems to be less well-defined.
These variants which are found in Bolivia and Panama may be the result of hybridization or they may be ecological variants.
The closest relative of P. riparia in Central America is thought to be P. maculata. The ranges of these two species overlap in Guatemala.
Because of the dissimilarities in gross morphology between P. quichensis and P. oaxacana these two species are not considered as likely to hybridize with P. riparia as is P. maculata.
The chromosome numbers of P. maculata and P. riparia are likely the same. A haploid count of "about 15" is reported for
P. riparia (Table 2) while haploid eounts of 16 and 17 are listed for P. maculata.
Specimens Examined: BOLIVIA: La Paz: near Coroico,
1100 m. alt., 0. Buchtien, 2438, in 1912 (US). Oruro: Los
Yungas, 0. Buchtien, 2438, Dec., 1908 (US). COLOMBIA: Antioqui&
Daniel, 903, Sept. 14, 1936 (US). Boyacli.: Honda Valley, A.
Joseph', 10l4a, July, 1923 (US); NW of Bogota near Mt. Chapon,
A. E. Lawrence, 225, in 1932 (NY). Caldas: Salento, F. W.
Pennell, 8 9 2 8, July 25-31, 1922 (US); Salento, 2000-2200 m. alt.,
F. W. Pennell, 8931, July 25-31, 1922 (NY, US); Rio Quindio above
Armenia, F. W. Pennell et_al., 8 7 3 1, July 2 5, 1922 (NY, US); Rio
Quindio above Armenia, Cordillera Central, F. W. Pennell et al.,
8702, July 25, 1922 (US). Cauca: 1000-2200 m. alt., Lehman,
4549 (F 578368); near Popayan, 1700-2500 m. alt., F. C. Lehmann,
5528 (F 578357, US 1402126); Micay Valley near Rio San Joaquin,
1400-1500 m. alt., E. P. Killip, 7842 and 7 8 6 6, June 29-30, 1922
(US); Valley of Rio Palo between Tacueyo and La Tolda, 1780-1900,
Dec. 19, 1944 (F); Quebrada de Bermejal, I65O-I6OO m. alt., J.
Cuatrecasas, 23762, Mar. 10, 1947 (F ); Popayan, 1800 m. alt.,
S. Y. Agredo, 170 and 250, Mar., 1948 (F). Cundinamarca: Bogota
1321 (NY, US 1480755); Monte Redondo to Quetame, 1400-1500 m. alt
F. W. Pennell, 1844, Sept. 7, 1917 (NY); Guaduas, Ariste-Joseph, 1015, July, 1923 (US); Sasaima, Apolinar-Maria, sn., July, 1931
(F); La Vega, 1300 m. alt., E. P. Arbelaez and J. Cuatrecasas,
53^2, June 1939 (F); near San Bernardo, 1600-1800 m. alt.,
J. Cuatrecasas, 9636 and 9602, June 23, 19^+0 (F); between San
Francisco and La Vega, 1650-1730 m. alt., H. Garcia-Barriga,
10950, Jan. k, 1 9 ^ (US); Estacifin Santana above Sasaima, 1600-
1700 m. alt., A. Dugand and R. Jaramillo, 3822, July 25-29,
19^5 (US); between Guaduas and El Alto de Aguaclara, H. Garcia-
Barriga, 123^9, July 2k, 19^7 (US). Magdalena; Santa Marta,
H. H. Smith, 913, 1898-1899 (NY, US). Meta; Villavicencis, l4^f, T. A. Sprague, Jan. 25, 1899 (US). Norte de Santander: near Chinacota, l^fOO m. alt., E. P. Killip and A. C. Smith,
20799, Mar. 18, 1927 (US); Valley of Rio Margua between Miranda and Alto de la Aurora, 950-1087 m. alt., J. Cuatrecasas, 13^12,
Nov. 23, 19^1 (F, US). Narino; Cerro de las Pinas on trail to
Delicias, 1640 m. alt., J. A. Ewan, 15932, July 26-30, 1 9 ^ (NY,
US). Santander; between El Jaboncillo and Surata, 1500-1800 m. alt., E. P. Killip and A. C. Smith, 16^37, Jan. 3, 1927 (NY).
Tolima; road from Fresno to Falan in canon of Rio Guali, 1120-
1700 m. alt., H. Garcia-Barriga, 8323, Dec. 5, 1939 (US). Valle del Cauca; Valley of Rio Sanquinini, 1250-1^-00 m. alt., J.
Cuatrecasas, 15539, Dec. 10-20, 19^3 (F). ECUADOR: Cotapaxi;
8 km W of La Esperanza, E of Pilalo, 500 m. alt., C. B. Reiser,
Jr., 5050, Apr. 10, 1962 (IND). Pichincha; between Cansacoto and Tandapi, I. Holmgren, 804, July 15, 1920 (US). GUATEMALA:
Quezaltenango; S-facing slopes of Volcan Santa Maria between Santa Maria de Jesus and Calamach6, J. A. Steyermark, 3319^,
Dec. 31i 1939 (F); S-facing slopes of Volcan Santa Maria between
Santa Maria de Jesus and Calahuache, 1300-2000 m. alt., J. A.
Steyermark, 33^86, Jan. 1-2, 19^0 (F); along Rio Samala near
Santa Maria de Jesus, 1500-16^0 m. alt., P. C. Standley, 8462^,
Jan. 25, 19^1 (F); between Finca Pirineos and Patzulin, 1200- ikOO m. alt., P. C. Standley, 86756, Feb. 9, 19^1 (F). San
Marcos: 9 mi. NW of El Porvenir on SW-facing slopes of Volcan
Tajumulco, 1300-2000 m. alt., J. A. Steyermark, 37758, Melt. 1^,
19^0 (F, GH). PERU: Ayacucho: between Huanta and Rio Apurimac,
750-1000 m. alt., E. P. Killip and A. C. Smith, 22691, May 7, 17»
1929 (US). VENEZUELA: Distrito Federal: near Caracas, 800-
1000 m. alt., H. Pittier, 9787, Sept. 11, 1921 (NY); Cerros del
Avila, 1600 m. alt., E. Pittier, 3 6, Oct. 7, 1921 (US); Los
Venados, Elias, 52, Aug., 1939 (F); Los Venados, 1500 m. alt.,
C. Vogl, V 70, Oct. 1, 19^0 (F). Miranda: Los Teques in German
plantations, 1^00-1500 m. alt., H. Pittier, 7510, Oct. 19, 1917
(US); Petare, 800 m. alt., E. Pittier, 9787, Sept. 11, 1921 (US);
Ocumare del Tuy, 950 m. alt., L. Williams, 12^43, Oct. 15, 1939
(F); Ocumare del Tuy, 1500 m. alt., H. Gines, 41^0, in 1951 (US).
(H) Polymnia maculata Cav. Icones et Descr. PI. 3!l^» 179^•
Coarse herbs to 5 meters tall, stems furrowed, purple-
spotted, glabrous to hispid-pilose or scabrous; leaves sessile or
petiole winged to base, as large as ^5 cm long and 30 cm broad,
basally connate, shape variable, deltoid to broadly ovate, main
veins and lobes of lower leaves palmate or sometimes pinnate, lobes serrate and dentate, acute or acuminate, sometimes hastate, the base truncate or cuneate, below densely canescent, hispid- pilose or sparsely puberulent, sometimes with yellowish sessile glands and hairs, upper leaves unlobed, ovate-lanceolate; peduncles glabrous, densely stipitate-glandular, hispid-pilose or pilose and glandular; pjiyllaries 5-6, in 1 series, basally united, ovate to lanceolate, up to 16 mm long and 10 mm wide, vesture as variable as that of peduncles, margin hairy; ray flowers up to about 20, corolla yellow, tridentate, 1 cm wide or less, up to 2.5 cm long, disc flowers 8 mm long or less, corolla yellow, puberulent on the lower surface of lobes; fruit black, flattened-ovoid, longitudinally striate, about 5 nim long or less, up to 4 mm wide.
Type: "Habitat in Nova-Hispania. Floruit in R. horto
Matritense mense Decembri 1793•"
Polymnia maculata is morphologically quite different from most other species of Polymnia; but it is difficult to find good characters by which one can separate it from P. uvedalia. The fruits of P. uvedalia appear to be consistently larger than those of P. maculata. The leaves of P. maculata most frequently have truncate to attenuate bases and acute lobes but in some cases they have hastate bases and long-acuminate lobes. In both species peduncles are glabrous or nearly so, pilose, glandular, or pilose- glandular. Blake (1917a, 1917b, 1935) has used these variations as criteria for recognition of several varieties within each of the two species. Haploid chromosome counts of 16 have been re ported for both species (Table 2). 61
In the southern part of its range P. maculata might be confused with P. riparia with which it occurs sympatrically.
Polymnia maculata, however, has purple-spotted stems, ray pales which are not inrolled along the margins and certain differences in vesture of the ray pales. Also, the leaves of P. riparia are less deeply divided.
Habitats in which P. maculata grows include damp thickets, moist slopes where oak trees grow, tropical evergreen forests, sandy loam in open sun, coffee plantations, coconut groves, and heavy red clay loam of pine forests.
Common names used for plants of this species are "furca"
in Costa Rica and "mirasol" in Guatemala.
(I) Polymnia maculata Cav. var. maculata comb. nov.
Polymnia maculata Cav. var. vulgaris Blake, Contr. Gray Herb. 52:33. 1917. (Isotype FI, GH!, NY!, US!).
Polymnia maculata Cav. var. hypomalaca Blake. Contr. Gray Herb. 52:33. 1917* (Paratype F!, GH!, MO!, NY!, US!).
Branches and peduncles hispid-pilose and sparsely glandu
lar; leaves densely or sparsely canescent (Fig. 13).
Blake's varieties vulgaris and hypomalaca were originally
separated on the basis of type of vesture of the lower leaf sur
face. These two varieties are separable when about the upper 30
cm of flowering stems are examined. This is the approximate
length of most of the stems on herbarium sheets. However, some
tall specimens can be found for which at least two herbarium sheets
have been used in mounting the single specimen. In such cases
the leaves at lower nodes would key out to var. vulgaris while 62
fs?&,
Fig. 13*— Polymnia maculata var. maculata Scale: .28X the upper would key out to Blake's var. hypomalaca. In view of this consideration I have combined the two varieties under a single variety.
Blake's epithet "vulgaris" is replaced by "maculata" in conformation with Section 5» Article 26 of the International
Code of Botanical Nomenclature (1961).
Haploid chromosome counts of 16 and 17 have been reported for var. maculata (Table 2).
Specimens Examined: BRITISH HONDURAS: Stann Creek:
Kendal, 150 ft. gilt., W. A. Schipp, 80*f, June 26, 28, 1931 (F, 63
GH, MO, NY). El Cayo: Vaca, P. H. Gentle, 2215, Feb. 22, 1958
(F, GH, MO); El Cayo, H. H. Bartlett, 12096, Mar. 3 0, 1931 (GH,
US); El Cayo, C. L. Lundell, 6 6 1 9, June-Aug., 1936 (US). COSTA
RICA: Ala.juela: in San Pedro de Montes, J. M. Orozco, 3 6 0,
Oct. 21, 1938 (F). Cartago: S of Orosi, 1800 m. alt., R. M.
King, 5383, Aug. 22, 1962 (SS). San Jose: Tucurrique, 635 m. alt., A. Tonduz, 13270, Feb., 1899 (US); Puntarenas: near Buenos
Aires, Pittier, 4923, in 1892 (GH). near San Josl, H. Pittier,
427, Aug., 1888 (US); Tucurrique, 800 m. alt., J. Leon, 743,
May 20, 1941 (F). EL SALVADOR: Santa Ana: NW flank of Volcan
de Santa Ana, 1710 m. alt., J. M. Tucker, 128l, Apr. 23, 1942
(F, NY, US). San Vicente: Volcan de San Vicente, 1200-1500 m.
alt., P. C. Standley, 21496, Mar. 7-8, 1922 (GH, NY, US).
GUATEMALA: Alta Verapaz: Coban, 4300 ft. alt., H. von Turckheim,
1102, Jan., 1887 (F, GH, NY, US); Coban, H. von Turckheim, II
6 5 8, Nov., 1902 (US); Coban, H. von Turckheim, 8427, Nov., 1902
(F, GH, NY, US); Coban, H. von Turckheim, II 1334, July-Aug.,
1906 (F, GH, MO, NY); H. von Turckheim, II 65 8, Aug., 1907 (US);
H. von Turckheim, 4015, Dec., 1912 (F); near Coban, 1260-1440 m.
alt., P. C. Standley, 69485, Mar. 26, Apr. 15, 1939 (F); along
rt. 5 about 1 mi. S of Coban, R. M. King, 3306, July 2, i960
(TEX); along rt. 5 about 2 mi. NE of San Pedro Carcha, R. M. King,
3326, July 12, i960 (TEX, US); Fields along rt. 5 about 1 mi. S
of Coban, R. M. King, 3306, July 12, i960 (US); on rt. 7 about
17 mi. W of San Cristobal, R. M. King, 3355, July 13, I960 (TEX,
US). Amatitlan: Villa Nueva, Heyde and Lux, 4509, Nov., 1889 64
(F, US). Baja Verapaz: between Purula and Cucuni, 0. F. Cook,
305, June 6, 1904 (US); between Purula and Tuceru, 0. F. Cook and C. B. Doyle, 305, June 5» 1904 (US); 17 mi. from Salama on road to Coban, Patal, J. G. Hawkes et al., 1936, Nov. 11, 1958
(F). Chimaltenango: Finca Santa Emilia near Pacayal, 3,200 ft. alt., J. Bequaert, 23, Feb. 19, 1931 (F, GH); near Tecpam, 7,200
ft. alt., A. F. Skutch, 6 0 8, Sept. 12, 1933 (GH, NY, US); Alameda,
J. R. Johnston, 3 6 9, Nov. 20, 1936 (F, GH); Volcan de Agua, 7,500
ft. alt., July 22, 1937 (F); Calderas, J. R. Johnston, 1103,
Oct. 25, 1937 (F); at Cuilapa, J. R. Johnston, 1088, Oct. 31,
1937 (F); near Finca La Alameda, I83O m. alt., P. C. Standley,
59065, Dec. 7, 1938 (F). Chiquimula: Volcan Ipala near Amatillo,
900-1510 m. alt., J. A. Steyermark, 30499, Oct. 25, 1939 (F);
Cerro Tixixi 3-5 N of Jocotin, J. A. Steyermark, 31553,
Nov. 10, 1939 (F). El Progreso: Finca Piamonte and summit of
Volcan Santa Luisa, 2400-3333 m. alt., J. A. Steyermark, 43592,
Feb. 5, 1942 (F, NY). Escuintla: 1100 ft. alt., J. D. Smith,
2384, Mar., 1890 (F, GH,US). Guatemala: San Rafael, 7000 ft.
alt., E.W.D. Holway, 62, Jan. 10, 1915 (GH); 10 Ion S of San
Raimundo, 1800 m. alt., P. C. Standley, 6 2 889, Jan. 18, 1939 (F,
NY); I. Aguilar, 3 6 6, in 1939 (F 100094-1) „ Huehuetenango: Todos
los Santos, 2727, Sept., 1896 (NY). Izabal: Quebradas, H.
Pittier, 8617, May 19-22, 1919 (GH, NY, US). Jalapa: Inuaz,
W. A. Kellerman, 7701, Jan. 8, 1908 (F, NY); near Jalapa, I36O
m. alt., P. C. Standley, 77151, Nov. 7-18, 194-0 (F). Jutiapa:
near San Jose Acatempa, 1300 m. a lt., P. C. Standley, 77567, Nov. 19, 19*1-0 (F). Quezaltenango: ravine below Fuentes
Georginas above Zumil, 25OO m. alt., J. A. Steyermark, 3^ 8 3,
Jan. 20, 19*1-0 (F). Quiche: Nebaj, 7100 ft. alt., A. F. Skutch,
1753. Nov. 21, 193*1- (GH). Retalhuleu: San Felipe, 2055 ft. alt.
J. D. Smith, 2863, April, 1892 (F, GH, US); San Felipe, W. A.
Kellerman, 5*t*t7, Feb. k, 1906 (F, OS); Las Victorias, 660 m. alt.
Rojas, 276, Mar. 21, 1921 (US); P. C. Standley, 6 6 676, Feb. 27,
1939 (F); near Las Delicias S of Retalhuleu, 200 m. alt., P0 C.
Standley, 88113, Feb. 22, 19*1-1 (F); near Retalhuleu, 2*f0 m. alt.,
P. C. Standley, 88673. Feb. 17, Mar. 1, 19*H (F). Sacatepequez:
Antigua, W. A. Kellerman, *f809, Feb. 15, 1905 (OS, US); slopes of Volcan de Agua, S of Santa Maria de Jesus, 1800-2100 m. alt.,
P. C. Standley, 59505. Dec. 10, 1938 (F); above Pastores, P. C.
Standley, 60771, Dec. 23, 1938 (F, NY). San Marcos; Finca
Cauqutz, El Quetzal, 1150 m. alt., G. Salas, 3 8 7, May 31, 1923
(US); Rio Cabus near Malacatan, 300 m. alt., P. C. Standley,
68880, Mar. 15, 1939 (F); Mtns. along rt. 1 about 12 mi. W of
San Marcos, R. M. King, 3151, July, i960 (TEX, US); about 3 mi.
W of El Carmen on rt. 1, R. M. King, 3128, July 1, i960 (TEX, US)
Santa Rosa: Las Cruces, 6000 ft, alt., Heyde and Lux, 6l6l,
Oct., l89*f (F, GH, MO, NY, US); near Cuilapa, 895 m. alt., P. C.
Standley, 77993, Nov. 20-27, 19*K) (F); near Cuilapilla, 900 m.
alt., P. C. Standley, 7 80 2 3, Nov. 23, 19**-0 (F). Solala; Volcan
Atitlan, 6 5OO ft. alt., W. A. Kellerman, 63IO, Feb. 6, 1907 (F,
OS). Suchitepequez: Mazatenango, G. Bernoulli, 576, May, 187O
(NY); Mazatenango, 1250 ft. alt., W. A. Kellerman, *<-965, Feb. 28, 1905 (OS, US); Mazatenango, W. A. Kellerman, 5305, Mar. 2, 1905
(F, OS); Mazatenango, W. A. Kellerman, 5337j Feb. 12, 1906 (OS);
Mazatenango, W. A. Kellerman, 538^, Feb. 15, 1906 (F); near
Patulul, 330-600 m. alt., P. C. Standley, 62171, Jan. 5, 1939
(F). Zacapa: on trail between Bio Hondo and Finca Alejandria,
1000-1500 m. alt., J. A. Steyermark, 297^8, Oct. 11, 1939 (F).
HONDURAS: Atlantida: near La Ceiba in coconut grove, T. G.
Yuncker et al., 8*f22, July l^f, 1938 (F, GH, MO, NY). Cortes:
Lake Yojoa along Rio Lindo, V)0 m. alt., L. 0. Williams and A.
Molina, 1^593, Aug. 8, 19^8 (GH, MO). Olancho: near Cqtacamos along river, ^50-500 m. alt., P. C. Standley, 13^16, Mar. 18-26,
19^9 (F, MO); 1 km from Canaverales along Rio Lindo, 600 m. alt.,
A. Molina, 5511, Aug. 25-26, 1955* MEXICO: Chiapas: Ghiesbreght,
577, 1864-1870 (GH); near San Cristobal, 7000-8000 ft. alt., E. W.
Nelson, 3221, Sept. l8, 1895 (US); Silsepec, E. Matuda, 76^,
Dec. 31, 1936 (GH, MO, US); Acacoyagua, E. Matuda, 17527, Feb.
29, 19^8 (F, NY); Sultepec, 2300 m. alt., E. Matuda, 29262, Sept.
21, 1953 (NY). Coahuila: Arteaga, 2000 m. alt., G. B. Hinton, l68l6, July 2b, 19^9 (GH). Distrito Federal: Barranca de Santa
Fe, M. Bourgeau, 59°, July 22, 1865 (GH, NY, US); near Santa Fe,
J. N. Rose and J. H. Painter, 7996, Oct. 18, 1905 (US); El
Rosario, L. II. MacDaniels, 70^-, Sept. 1, 1936 (F); Santa Fe, 2^50 m. alt., E. Matuda, 21^07, Aug. 12, 1951 (NY); Cuajiraalpa, Rio
Hondo, 2^00 m. alt., E. Matuda, 262 9 8, June 29, 1952 (NY).
Guerrero: Plains near Tecpam, 2100 m. alt., A. F. Skutch, 6 0 8,
Sept. 12, 1933 (US). Hidalgo: near Ixmiqulpan, J. N. Rose, 91^8, in 1905 (US); Chapulhuacan, L. A. Kenoyer, 756, Nov. 27, 1937
(F). Jalisco: Sierra del Tigre, 3 mi* S of Mazamitla, 2100-
2200 m. alt., R. McVaugh, 13, 1 6 9 , Sept. 22, 1952 (US). Mexico:
Rio Hondo Canon, E.W.D. Holway, 3562, Oct. 4, 1889 (GH); Bar ranca above Santa Fe, 85OO ft. alt., C. G. Pringle, 1356^,
Sept. 1, 1905 (GH, US); Nanchititla, G. B. Hinton, 5227, Nov. 18,
1933 (F. GH, MO, NY, US); near Rio Hondo, 2500 m. alt., E. Matuda,
21837, Sept. 9 , 1951 (NY). Michoacan: Patzcuaro, E.W.D. Holway,
3598, Oct. 10, 1899 (GH); near Patzcuaro in fields, C. G.
Pringle, 3326, Oct. 23, I89O (F, GH, MO, US); Morelia, Arsene, sn., in 1909 (F 387306); near Morelia Loma Sta. Maria, 1900 m. alt., G. Arsene, 5336, in 1910 (MO, US); near Morelia, 1950 m. alt., G. Arsene, 5 8 8 2, Sept. 4, 1910 (F, GH, MO, NY, US); near
Morelia, 2000 m. alt., G. Arsene, sn., Sept. k, 1910 (GH); near
Morelia Rincon Carrieres, 2000 m. alt., G. Arsene, 828^, Oct. 17,
1912 (F, MO, NY). Nayarit: along rt. 15 about 1 mi. N of Tepix,
R. M. King, 3 6 8 8, Aug. 11, i960 (TEX,US). Nuevo Leon, 6600 ft. alt., A. J. Sharp, ^5689, July 12, 19^5 (NY). Oaxaca: W slope of Mt. Zempoaltepec, 7700-8000 ft. alt., E. W. Nelson, 596,
July 5 to 13, 18 9 ^ (US). Puebla: Huejatzingo, G. Arsene, 1790,
Nov. 17, 1908 (MO, US); Puente del Emperador, La Venta, 83OO ft. alt., A. J. Sharp, ^523, Aug. 25, 19 ^ (NY). Quer£taro:
Tequisquiapan, G. B. Hinton, 13^7, Aug. 20, 1932 (GH). San Luis
Potosi: Alvarez, 2200-2^00 m. alt., F. W. Pennell, 17888, July
30-31, 193^ (US). Vera Cruz: San Rafael, C. H. Townsend, 20
(US 66A-525); Orizaba, Botteri, 237 (GH); Orizaba, Muller, sn., in 68
1855 (NY); Wartenberg near Tantoyuca, L. C. Ervendberg, 286, in 1858 (Gil); E. Iverber, 8, July 21, 1882 (US); Kt. Orizaba,
Cordoba, 2700 ft. alt., H. E. Seaton, 420, Aug. 20, 1891 (F, GH,
US); Jalapa, 4-300 ft. alt., F. W. Johnston, sn., Sept. 22, 1906
(NY); C. A. Purpus, 2 8 5 3, Dec., 1907 (NY); Boca del I-ionte, C. A.
Purpus, 3024, Sept., 1908 (F, GH, MO, NY, US); Orizaba, 4700 ft. alt., G. L. Fisher, 191, Aug. 9, 1924 (F, MO, US); Cordoba, 0. A.
Plunkett, 191, Aug. 27, 1932 (F); Cordoba, 2700 ft. alt., G. L.
Fisher, 35356, July 27, 1935 (F, MO, NY,US); near Jalapa, 4600 ft. alt., L. II. MacDaniels, 356, Aug. 30, 1935 (F); Orizaba,
L. A. Kenoyer, A 166, Aug. 12, 1938 (F); between Tecamaluca and
Ojo Zarco above Orizaba, 4300 ft. alt., A. J. Sharp, 44770,
Sept. 12, 1944 (NY); roadside bank below El Puerto and above
Acultzingo, 7450 ft. alt., A. J. Sharp, 44785, Sept. 13, 1944
(NY); in gorge below Ixolo Falls near Jalapa, 3300 ft. alt.,
A. J. Sharp, 4 5 8 0 3, Aug. 2, 1945 (NY); near Tapalapan, NW of
Santiago, R. L. Dressier and Quentin Jones, 48, Aug. 11, 1953
(GH, NY, US); Cordoba along rt. 150, R. M. King, 2 6 6 5, June 6, i960 (TEX, US). Zacatecas: near Plateado, J. H. Rose, 2749,
Sept. 2, 1897 (US). NICARAGUA: Zelaya: A. Molina, 2338, Apr.
23, 1949 (F). PANAMA: Chiriqui: near Monte Lirio valley of upper Rio Chiriqui Viejo, 1300-1900 m. alt., R. J. Seibert, 292,
June 27, July 13, 1935 (US); Volcan de Chiriqui, 85OO ft. alt.,
M. E. Davidson, 978, July 15, 1938 (GH); 1 mi. SW of Boquete,
4000 ft. alt., P. H. Allen, 4726, July 20, 1947 (MO); about 8 mi. 69
W of David on interamerican hwy., R. M. King, 5 2 8 5, July 29,
1962 (US); along road to David about 1 mi. S of Boquete, R. 'M,
King, 5 3 1 7, July 30, 1962 (US).
(J) Polymnia maculata Cav. var. adenotricha Blake. Contr. Gray Herb. 52:3^. 1917*
Peduncles and branches densely stipitate-glandular, hispid-pilose hairs sometimes present; leaves below sparsely pubescent, often glandular-dotted.
Type: GUATEMALA: Santa Rosa: Chupadero, 5^00 ft. alt.,
Heyde and Lux, 3307, Aug., 1892 (Holotype GH!, Isotype Ft, US!).
COSTA RICA: Cartago: San Rafael de Cartago, H. Pittier, 6993,
Aug. 28, 1892 (Paratype GH!); Cartago, A-250 ft. alt., Cooper,
5827, Dec., 1887 (Paratype F! , GII! , US I ). MEXICO: San Luis
Potosi: Tancanhuitz, Nelson, A401, May 2, 1898 (Paratype F!,
GH!, US!). Vera Cruz: Zacuapan, Purpus, 2853, Dec., 1907
(Paratype GH!); Mirador, Sartorius, sn., (Paratype GH!).
Specimens have been examined which are intermediate between varieties adenotricha and vulgaris.
R. M. King reports via personal communication (1962) a haploid chromosome count of 33 plus one fragment for var. adenotricha. This is indeed unexpected in view of two previous haploid counts of 16 and one count of 17 for the var. vulgaris
(Table 2). If the chromosome count of 33 plus one fragment is consistent for the variety adenotricha and this variety is truly reproductively isolated from var. vulgaris— which indeed may be
the case— then perhaps var. adenotricha should be elevated to specific rank. Specimens Examined: COSTA RICA: Cartago: Dulce Nombre, ikOO m. alt., P. C. Standley, 35376, Feb. 27, 192^ (US); R. T.
Rojas, 130, Dec. 30, 1924- (US); in vicinity of Finca Las Concavas,
P. C. Standley, ^+1530, Dec. 7-8, 1925 (US); k km E of Cartago,
R. M. King, 5^06, Aug. 28, 1962 (US). San JosS: 1500 m. alt.,
J. D. Smith, 6609, Mar., 1896 (US); A. Alfaro, 32^23, Jan., 192*+
(US); near San Jos£, 1150 m. alt., P. C. Standley, 33300, Feb.,
192*+ (GH, US); N. Valerio, 13*+*+, July, 1935 (F). GUATEMALA:
Amatitlan: Villa Nueva, 3500 ft. alt., Heyde and Lux, *+509,
Nov., 1889 (F). Jalapa: Cerro Alcoba, E of Jalapa, 1300-1700 m. alt., J. A. Steyermark, 325*^, Dec. 2, 1939 (F). Quezalten- ango: Montana Chicharro on SE-facing slopes of Volcan Santa
Maria, 2-*+ mi. S of Santa Maria de Jesus, 1*+00-1500 m. alt.,
J. A. Steyermark, 3^270, Jan. 17, 19^0 (F). HONDURAS:
Comayagua: 6 km N of Siguatepeque, 1200 m. alt., T. G. Yuncker et al., 6311, Aug. 4, 1936 (GH, MO, US). Morazan: Zamorano,
800 m. alt., J. V. Rodriguez, l*+35» Nov. 1, 19*+3 (F); near Hoya
Grande, L. 0. Williams and A. Molina, 10*+*+2, Aug. 25, 19^6 (F);
Mt. Uyuca in guamil, S. F. Glassman, 2173, Aug. 7, 19^8 (F, NY); drainage of Rio Yeguare, 1700 m. alt., A. Molina, 1096, Sept. *+,
19^8 (F); V/ slope of Cerro de Uyuca on trail toward Tatumbia,
1500-1600 m. alt., P. C. Standley, 22733, Aug. 17, 19^9 (F);
Region of El Jicarito above El Zamorano, 900-1000 m. alt., P. C.
Standley, 2*+857, Dec. 1, 19^9 (US); between Santa Lucia and Valle de Angeles, 1800 m. alt., A. Molina, 3317, Sept. 26, 1950 (F); drainage of Rio Yeguare, 800 m. alt., A. Molina, 5368, Dec. 15, 71
195^ (F). MEXICO: Chiapas: Coapilla, S. Juzepczuk, 1?00, in
1926 (US 1 ^9295^)* Distrito Federal: Mexico City, E.W.D. Holway,
305^, Oct. Ik, 1898 (GH),. Guanajuato: L. A. Kenoyer, 2226,
Aug. 20, 19^7 (GH). Hidalgo: Chapulhuacan, L. A. Kenoyer, sn.,
Nov. 27, 1937 (MO); at border of Hidalgo-San Luis Potosi on hwy. below Chapulhuacan, 2100 ft. alt., H. E. Moore, 50^1, Sept. 22,
19^9 (GH). Michoacan: near Patzcuaro, C. G. Pringle, 3326,
Oct. 23, 1890 (GH). Nuevo Leon: near Monterrey, C. K. and M. T.
Mueller, 250, July, 1933 (F, GH, TEX); Dulces Nombres, 1850 m.
alt., F. G. Meyer and D. J. Rogers, 2689, June 28, 19^8 (MO, US);
Cerro del Viejo, 15 mi. W of Dulces Nombres in Zaragoza, F. G.
Meyer and D. J. Rogers, 3031> Aug. 19, 19^8 (MO, US). San Luis
Potosi: Cerro Grande 5 km NW of Guadalcazar, Rzedowski, 5G6O,
Oct. 3» 195^ (US); 27 mi. E of San Luis Potosi, J. Crutchfield
and M. C. Johnston, 5640, Sept. 19, i960 (TEX); 7 mi. NE of
Xilitla on road to Jalpan, R. M. King, M+27, Apr. 1, 1961 (NY,
TEX). Vera Cruz: along rt. 150 about V/z mi. W of Cuitlahuac,
R. M. King, 2368, Jan. 27, I960 (TEX, US).
(K) Polymnia maculata Cav. var. glabricaulis Blake. Wash. Acad. Sci. 25:313-31^. 1935.
Peduncles glabrous; stems glabrous except for villous
hairs at the node; leaves scabrous above, sparsely puberulent
below.
Type: MEXICO: Tamaulipas: Cerro Carrizo, near San
Jose, ^800 ft. alt., Bartlett, 10508, July 19, 1930 (Holotype
US! ). 72
The peduncles and stems of Blake's var. glabricaulis are distinctly more glabrous than those of any other specimen of P. maculata I have examined. The holotype examined is the only specimen of this variety known to me. Whether the lower portion of the stem is glabrous or not is subject to conjecture.
The stem portion of the holotype is about 15 cm long. Polymnia maculata may grow to heights of 5 meters. I presume that this glabrous condition extends throughout the remainder of the stem.
In all probability this is likewise an assumption on Blake's part. Nevertheless, there does not yet seem to be sufficient evidence for the rejection of this taxon as a separate entity.
(L) Polymnia uvedalia L. Sp. PI. Second Edition. Vol. 2, 1503. 17^57
Osteospermum uvedalia L. Sp. PI. First Edition, Vol. 2, 923. 1753^ CHolotype).
Polymnia macrophylla Raf. Florula Ludov. 70-71. l8l7.
Polymniastrum uvedalia (L.) Small in Small and Carter. FI. Lane. Co. 302. 1913.
Smallanthus uvedalia Mackenzie in Small. Manual of Southeast. FI. l4o6. 1933.
Perennial herbs up to or exceeding 3 meters in height; root system fibrous but some enlarged roots or underground stems are often present and resemble tubers; stems furrowed, purple- spotted, glabrous to densely pilose, about 3 cm in diameter at base or usually less; leaves up to 39 cm wide, 68 cm long, sessile or winged to base, palmately 3-5-lobed and veined, ovate to del toid, the base abruptly cuneate to the winged portion, scabrous 73 to nearly glabrous above, below glabrous or resin-dotted and puberulous, hairs larger on main veins, upper leaves sparsely dentate, often unlobed; peduncles glabrous, densely pilose, stipitate-glandular, or glandular-pilose; phyllaries *f-6, up to l2 mm wide and 20 mm long, broadly ovate to ovate-lanceolate, glabrous above, below glabrous or with sparse pubescence or greenish-yellow sessile glands; ray flowers 7-1 3 i corollas yel low, 3 cm long or less, about 1 cm wide, the tube 2 mm long, pilose or glandular-pilose outside; disc flowers ^5-8 0, yellow, about 1 cm long, the corolla somewhat pilose outside; ray pales broadly ovate, acuminate; disc pales lanceolate, chartaceous;
frixits 6 mm long, b mm wide, longitudinally multistriate, ovoid and somewhat laterally compressed, color first white then red then purple then black.
Type: Probably collected in Virginia by Frederick
Gronovius.
The three varieties of P. uvedalia as recognized by
Blake (1917b) are var. genuina, var. densipilis, and var.
floridana. Infraspecific epithets such as "genuina" and "vera"
are illegitimate except where they repeat the specific epithet.
This is in accordance with Section 5» Article 2b of the Inter
national Code of Botanical Nomenclature (1961). The correct
epithet for the type variety becomes uvedalia.
The three varieties of P. uvedalia which I now recognize
are var. uvedalia, var. floridana, and var. densipilis. As Blake
pointed out (1917b) these varieties differ in the vesture of peduncles. The var. uvedalia has densely stipitate-glandular peduncles while those of var. densipilis are densely pilose.
The peduncle vesture of var. floridana is in some respects inter mediate between that of vars. uvedalia and densipilis. However, the degree of pilosity and size of glands and hairs is greatly reduced in var. floridana from that of vars. uvedalia and densi pilis . Intermediates between these three varieties have been found.
Although each of the three varieties in general conforms to a specific geographical distribution, specimens have been col lected outside the usual range (Fig. 37) • Th-e var. densipilis seems to occur in the western part of the species range while
Blake's var. genuina is mainly limited to the north and north eastern portion of the range. Variety floridana occurs in the southern part of the range; i.e., in the South Atlantic and central gulf states. More than one variety may be found at the same station. Variety densipilis was collected with variety genuina. W. H. Duncan collected a glabrous plant along with var. floridana in Georgia. The paucity of records for Mississippi and Alabama is probably due to unavailable information, not to rarity of the plants.
I believe that P. uvedalia and P. maculata are the same
species. This belief is based on their similarity in chromosome
number as well as gross external morphology. The size of fruit
which I have used to separate them seems to hold true but ad
ditional characters should exist if the separateness of the two
entities is to continue to be recognized. 75
Transplants of the two species to a common research garden followed by a series of hybridization experiments would probably shed appreciable light on how distinct the two entities really are. Herein lies one of the most interesting experimental taxonomic problems in the genus. A haploid chromosome number of
16 has been counted for both species and since their morphology is so strikingly similar I would expect hybrids between the two to be obtainable.
Let us assume that the genus originated in Central or
South America. With the spread of maculata-like forms northward in Mexico then northeastward into the eastern United States, a continuous range of distribution could have existed. Subsequent occurrence of hot and dry climates in the southwestern part of the United States destroyed most or all of the moist shaded habitats of the species, consequently a disjunct distribution resulted.
From the distribution map (Fig. 37)» it appears that a disjunct distribution exists betv/een the United States and Mexico.
If the variation in peduncle vesture was not present at the time of maximum species spread then there would have resulted with
this disjunct distribution, an example of parallel evolution occurring in regions now known as the United States and Mexico or some country south thereof.
Polymnia uvedalia has been collected in the islands of
Bermuda. All three varieties are known to occur there. Blake
(1917b) makes the following statement concerning the species 76 occurrence on the islands:
On the Bermuda Islands, where it is certainly introduced and seems to be rapidly naturalizing itself, this form is the only representative of the species. The plant is not noticed in Reade's list of Bermuda plants (1 8 8 3), and as it is very unlikely that so conspicuous a plant could have escaped his notice, it seems very probable that it is a recent introduction.
Support for a theory that highly mutable genes control peduncle vesture is attempted belpw:
1) Something is known about the geographic range of distri
bution of the three varieties of P. uvedalia in the
United States.
2) Each variety in the United States has its own range of
distribution with some overlap present.
3) All three varieties occur on Bermuda.
4) The plants were probably introduced into Bermuda since
1883.
5) The plants transplanted to Bermuda must have come from
the United States because the species does not occur
elsewhere.
6) Owing to the rather distinct geographic distribution for
each variety it is possible but not probable that 2 or
all of the three varieties would occur in a single popu
lation some of which was transplanted to Bermuda.
7) This transplant has undergone changes in Bermuda; :Le.,
mutations have occurred in an isolated environment far
removed from other plants of the same species. 77
Therefore, rather highly mutable gene(s) could be pre sumed to control peduncle vesture. The fact that distinct ranges of distribution exist in the United States for each variety may be due to controlling factors in the environment other than heat and/or moisture. It is not likely that such extremes as we know in the United States would be likewise present in this sub tropical climate.
Mature specimens were collected in Bermuda as early as
1905. This means that the date of transplanting probably occurred between 1883 and 190^ or 1905 (Blake, 1917b).
A theory alternate to that outlined above would be that hybrids were transplanted to Bermuda and outbreeding has resulted in the appearance of all three forms.
I have succeeded in achieving reciprocal crosses between all three varieties (see chapter on Experimental Investigations for further detail) but as yet have not found the proper environ ment in which hybrid seeds germinate. When this obstacle has been overcome more can be said about the inheritance of the vesture of peduncles as well as about which of the two theories is more credible. On the other hand there may be genetic incom patibility between these varieties which results in failure of hybrid seeds to germinate under any set of environmental conditions.
Seven soil samples were analysed from seven different populations in which P. uvedalia plants were growing. The soil samples were taken with a tubular Hoffer Soil Sampler made by the Elano Corporation in Xenia, Ohio. Each sample, 13/16 inch 78 in diameter and 6 inches long, was kept in a sealed plastic bag for approximately 10 days prior to testing. All soil tests were performed by The Ohio State University Soil Testing Laboratory,
Agricultural Extension Service, on the Columbus campus#
Table 1 shows the results of the soil test data. The purpose of this table is not to show the extremes of certain environmental factors but rather to give some indication of ranges which exist.
Soils associated with P. uvedalia are in all probability usually more acidic than those in which P. canadensis and P. laevigata grow. The pH of two soil samples from two stations of
P. canadensis populations and of a single sample from a P. laevigata station were determined by the soil testing service referred to above. All three samples were more basic than any of- the seven samples shown on the Table 1.
There appears to be no significant difference in soil acidity from one variety of P. uvedalia to another. I believe that additional soil samples would indicate that soil pH is not the controlling factor in peduncle vesture.
The largest population of P. uvedalia which I have en countered occurred along U. S. highway 33 in Calhoun County,
West Virginia, 3«1 miles from the Calhoun-Gilmer County line.
This is my collection site no. 213. 1 was informed of this popu lation via personal, communication ivith R. W. Cruden and its measurements are approximately 150 feet long and 20 feet wide.
It is a dense stand, part of which is under the canopy of trees. TABLE 1 POLYMNIA UVEDALIA SOIL HABITAT DATA
Collection Per Cent Number Location Variety Soil Type PH Organic Matter
213 Calhoun Co* uvedalia light reddish 6.7 3.0 W. Va. brown silty clay loam
248 Gaston Co* floridana grayish brown 6.4 ^•5 N. C. sandy loam
244 Stanly Co. floridana light brown 5.3 1.5 N. C. sdllt loam
243 Chataham Co. floridana dark brown 5.8 2.0 N. C. silt loam
247 Mecklenberg Co. densipilis light reddish 6.2 2.0 M. C. brown sandy loam
246 Union Co. densipilis light reddish 5.7 2.0 N. C. brown silt loam
242 Lee Co. densipilis brown silt 5.3 1.5 N. C. loam 80
Plants found in association with P. uvedalia (here all apparently the var. uvedalia) include: Robinia, Ulmus, Malus,
Phleum, Vitis, Rubus, Heliopsis, Daucus. Actinomeris, Erigeron,
Carex, and Monarda.
Plants of P. uvedalia growing under tree species tend to flower later than those in more exposed sites. However, Polymnia uvedalia, like P. canadensis and P. laevigata, are all generally favored by at least some shade.
After checking this population for two growing seasons
I have yet to find a single seed in the mature fruit. This leads one to conclude that this large population is a single clone which is self-sterile. Plants from each end of the population were crossed with no seed set (Table 6). Large seeds are formed when crosses are made with plants from geographically widely separated areas. If the number of new stems added annually is determined it might be possible to estimate the age of this popu lation.
A haploid chromosome number of 16 was counted for all three varieties of P. uvedalia (Table 2).
(M) Polymnia uvedalia L. var. densipilis Blake. Rhodora. 19:^8. 1917.
Peduncles and branches of the inflorescence densely pilose, glands not easily seen or else absent (Fig. 1*0.
Type: Texas: Comal Co.: Ten feet high, rare on the
shady banks of the Guadeloupe near New Braunfels, Lindheimer,
38l, Aug., 1850 (Holotype GHI). BERMUDA: along roads, Botanic
Garden, Paget, J. W. Harshberger, sn., June, 1905 (Paratype GHI), Fig. !*».— Polymnia uvedalia rar. densipilis (Peduncle of Holotype) ScaleI 6.71
U3l)( near Bailey*8 Bay* S* Brown* ^9^* Feb. 10-Mar. 9* 1908
(Paratype GHI* USI). Louisianat no date or location given*
Hale* an. (Paratype GHI)*
Variety densipilia has the largest range of distribution of any of the three varieties. It covers the western* southern* and southeastern portion of the species range.
There is some evidence that the shortest peduncles have the greatest amount of piloslty. It would appear that as peduncle elongation ensues the pilose hairs, already,formed, 82 are scattered by intercalary growth of the peduncle. The degree of pilosity, then, would be an environmental expression.
Specimens Examined: BERMUDA: Tucker's Town, S. Brown,
4?0, Feb. 10, Mar. 9, 1908 (F, GH, NY); near Tucker's Town, S.
Brown and N. L. Britton, 321, Aug. 31, Sept. 20, 1905 (F, GH,
NY, US); near Middle Road, Smith's Parish, A. H. Moore, 306l,
July 21, 1905 (US); Abbot's Cliff, S. Brown and N. L. Britton,
879, Aug. 27, Sept. 21, 1912 (NY); Paget, F. S. Collins, 348,
Aug. 21, 1913 (NY, US)j Arkansas: Benton Co.: E. K. Plant, sn., in 1899 (NY); Plank, sn., (MO 1767439)* Carroll Co.: Eureka
Springs, E. J. Palmer, 4448, Sept. 22, 1913 (MO). Garland Co.:
Magnet Cove, D. Demaree, 16155, Sept. 4, 1937 (NY); Cedar Shades
Road, Hot Springs, F. J. Scully, 6 7, Sept. 22, 1937 (MO). Hemp stead Co.: Fulton, E. J. Palmer, 12681, Sept. 8, 1917 (MO, NY).
Hot Spring Co.: Malvern, D. Demaree, 16292, Sept. 19, 1937 (F,
GH, MO, NY); Caddo River, De Gray Dam, D. Demaree, 46494, Sept.
27, 1962 (OS). Howard Co.: Mineral Springs, D. Demaree, 9743,
Oct. 14, 1932 (NY, MO, TENN). Marion Co.: Buffalo Rock State
Park, D. M. Moore, 480747, Aug. 19, 1948 (TEX). Montgomery Co.:
Bluffs in Blakely Dam reservoir, D. Demaree, 34313, Sept. 19,
1953 (NY). Pulaski Co.: Little Rock, H. E. Hasse, sn., Aug.
30, 1885 (PH); Little Rock, H. E. Hasse, sn., Sept., 1886 (MO,
NY). Delaware: New Castle Co.: Wilmington, highlands of the
Brandywine, Morong, sn., Sept. 1, 1873 (NY); Wilmington, opposite
Bancroft factory, A. Cummins, sn., July 30, 1890 (MO, NY).
Sussex Co.: Coast of Delaware, J. H. Churchill, sn., Sept. 1 3 , 1908 (MO). District of Columbia: G. McCarthy, sn., (F 348749);
Island opposite Georgetown, A. Schott, sn., Aug. 25, 1859 (F);
H. Brummel, sn., in 1869 (US 71549); Low island, L. F. Ward, sn.,
Sept. 10, 1876 (US); L. F. Ward, sn., Sept. 21, 1879 (GH);
Windy Gap, E. S. Burgess, sn., Oct. 1888 (US); Eiver Flats, E. S.
Steele, 47, Sept. 1, Aug. 17, 1896 (MO); Canal Hoad, E. S.
Steele, sn., Aug. 3* 1899 (US); near Chain Bridge along river,
H. D. House, 282, Aug. 19, 1904 (NY); Great Falls, Va., E. C.
Leonard, 846, Aug. 17, 1919 (US). Florida: Lake Co.: Leesburg,
H. O'Neill, sn., Sept. 26, 1929 (US). Volusia Co.: Hammock S
of Daytona, J. K. Small, 869O, May 8, 1918 (GH, NY). Georgia:
Baldwin Co.: on Indian mound Indian Island plantation, F.
Harper, 1123, June 10, 1939 (PH). Illinois: Jackson Co.: G. H,
French, sn., Sept. 21, 1878 (OS); Pope Co.: 1 mi. S of Herod
'Williams Hill, G. S. Winterringer, 1500, Aug. 17, 1948 (TEX, US).
St. Clair Co.: Bluffs Lake, H. Eggert, sn., July, 1882 (MO).
Union Co.: E of Wolf Lake, W. M. Bailey and J. R. Swayne, 5102,
Aug. 27, 1951 (NCU). Kentucky: Clark Co.: 3 mi. S of Indian
Fields, M. E. Wharton, 5105, Aug. 7, 1939 (TENN). Louisiana:
Natchitoches Co.: E. J. Palmer, 8717, Sept. 28, 1915 (MO).
Maryland: Montgomery Co.: Potomac River Flats, Cabin John,
J. H. Painter, 9 6 7, Aug. 14, 1904 (MO). Worcester Co.: Stockton
K. H. Rusby, sn., Aug., 1889 (NY). Missouri: Barry Co.: B. F.
Bush, 15193, Sept. 11, 1935 (MO). Boone Co.: Robey's Spring,
L. Jeffrey and F. Dronet, 1110, Sept. 8, 1933 (TENN). Butler Co.
Poplar Bluff, sn., Aug., 1879 (MO). Cape Girardeau Co.: between Mocassin Springs and beyond Sheppard Point, J. A. Steyermark,
64092, Sept. 22, 1946 (F). Douglas Co.: 2-2)6 mi. NW of Sweden,
J. A. Steyermark, 65057, Sept. 22, 1947 (F). Dunklin Co.:
Campbell, B. F. Bush, sn., Oct. 24, 1892 (MO); Campbell, B. F.
Bush, 6229, Sept. 7, 1910 (MO); Crowley's Ridge along St.
Francis River, J. A. Steyermark, 6655, Sept. 18, 1938 (F).
Jasper Co.: Webb City, Carterville, E. J. Palmer, 327, Sept.
28, 1902 (MC); Carterville, E. J. Palmer, 866, Aug. 26, 1906
(MO); 5 mi. SE of Carthage, E. J. Palmer, 3159, Sept. 11, 1910
(MO); Oronogo, E. J. Palmer, 18775, Aug. 22, 1920 (GH); Oronogo,
E. J. Palmer, 32652, Aug. 28, 1927 (NY). Jefferson Co.: Kim- mswick, F. Wislizenus, 1 8 3 , Aug. 23, 1885 (MO). McDonald Co.:
Noel, E. J. Palmer, 4138, Sept. 5, 1913 (MO, US). Madison Co.: along Castor River, J. A. Steyermark, 66100, Aug. 27, 1948 (F).
Oregon Co.: 1 mi. N of Greer, J. A. Steyermark, 27986, Aug. 27,
1939 (F); along Eleven Point River, J. A. Steyermark, 72495,
Aug. 20, 1951 (F). Ozark Co.: Norfolk Dam along N fork of White
River, J. A. Steyermark, 66230, Sept. 5, 1948 (F); between
Martin's Ford and Brixey Spring, J. A. Steyermark, 69OII, Aug.
22, 1949 (F). Reynolds Co.: 3 mi» S of Oates on Black River,
J. A. Steyermark, 19729, Sept, 21, 1935 (MO). Shannon Co.: 12 mi. E of Eminence, J. A. Steyermark, 28007, Aug. 26, 1939 (F).
St. "Charles Co.: yfz mi. S of Femme Osage, J. A. Steyermark,
64247, Oct. 13, 1946 (F, MO). St. Louis Co.: H. Eggert, sn.,
Aug., 1886 (MO); near Allenton, G. W. Letterman, sn., Sept.,
1890 (F, NY, PH, TEX, US); Allenton, G. W. Letterman, sn., Aug., 1895 (MO); Creve Coeur, N. M. Glatfelter, sn., Sept. 22, 1900
(MO); Creve Coeur Lake, J. H. Kellogg, sn., Sept. 22, 1901 (MO);
Rocky Hill Meramec hds., R. Hoffmann, sn., Sept. 2, 1917 (MO).
Stoddard Co.: bordering Mingo Swamp NW of Egypt School, J. A.
Steyermark, 66120, Aug. 28, 19^8 (F). Taney Co.: b mi. SW of
Protem, J. A. Steyermark, 66372, Sept. 6, 19^8 (F); 53^-6 mi. SE of Protem, J. A. Steyermark, 69^6 8, Sept. 29, 19^9 (F); 6 mi. SE
Forsyth, J. A. Steyermark, 69503* Sept. 30, 19^9 (F). New
Jersey: Hudson Co.: foot of Cliffs, Weehawken, W. M. Van
Sickle, sn., Aug. 20, 1895 (US). North Carolina: 2 mi. S of
Rowan Co. line and 3 mi. E of Watts X Road, H. E. Ahles and R. S.
Leisner, 19653* Sept. 23, 1956 (NCU). Chatham Co.: E of Car- bonton power plant, P. Kessler, 5 8 5, Sept. 8, 1955 (NCU); E of
Carbonton power plant, J. R. and J. M. Wells, 2^3, July 26,
1961 (OS). Cumberland Co.: Cape Fear River E of Fayetteville,
H. E. Ahles, 36656, Oct. 11, 1957 (NCU). Granville Co.: 2.7 mi.
E of Dickerson and 5 mi. W of Granville-Vance Co. line, H. E.
Ahles and R. S. Leisner, 1763^, July 2^f, 1956 (NCU). Halifax
Co.: near Winton, R. K. Godfrey, 5185* July l8, 1938 (GH).
Lee Co.: 1 mi. SW Moncure, L. S. Beard, 1073* July 28, 1955
(NCU). Mecklenburg Co.: 1 mi. N Matthews, J. R. and J. M.
Wells, 2^7, July 27, 1961 (OS). Orange Co.: Chapel Hill, W. C.
Corres, sn., Sept. 26, 1909 (NCU); Battle's branch S of II. V.
Wilson's, H. R. Totten, sn., July 22, 1915 (NCU); between Neville
and Morgan Creeks, Radford and Stewart, 226a, Oct. 1^, 1939 (NCU)
Richmond Co.: NW of Rockingham near Pee Dee River, A. E. Radford 86 l448l, July 24, 1956 (NCU). Stanly Co.: near bridge SW of
Albermarle, J. R. and J. M. Wells, 245, July 26, 1961 (OS).
Union Co.: Rocky River on N. C. 205 N of New Salem, H. E.
Ahles and J. Haesloop, 51477, July l4, 1957 (NCU). 'Wake Co.: near Crabtree Creek N of Raleigh, R. K. Godfrey, 2053, Aug. 18,
1937 (NY). Warren Co.: Jordan's Creek at Roanoak River, C. R.
Bell, 4317, Oct. 24, 1956 (NCU). Oklahoma: Adair Co.: 1 mi.
W of Watts, C. S. Wallis, 7788, Sept. 6, 1958 (TEX). Cherokee
Co.: 7.8 mi. NE of Tahlequah, C. S. Wallis, 7751, Sept. 6, 1958
(TEX). Delaware Co.: 11 mi. W of Stateline hwy. 33, D. Bo
Fowler, 34, Oct. 5, 1946 (TEX). Johnston Co.: near Tishomingo,
H. W. Houghton, 3518, Oct., 1915 (NY). Murray Co.: Arbuckle
Mtns., E. J. Palmer, 44058, Oct. 10, 1937 (MO). Oklahoma Co.:
N Canadian River, 1 mi. S of Harrah, U. T. Waterfall, 3098,
July 26, 1941 (GH). Pushmataha Co.: 5 mi. W and .7 mi. S of
Albion, U. T. Waterfall, 191, Aug. 20, 1932 (NCU). Rogers Co.:
Verdigris, B. F. Bush, 280, Oct. 5, 1894 (NY). Tulsa Co.: New
Block Park, H. A. Hawk, 17, Sept. 25, 1941 (MO). Pennsylvania:
Lancaster Co.: along railroad N of station, A. MacElwee, 1228,
Aug. 31, 1899 (PH). South Carolina: Charleston Co.: South
Santee River on U. S. 17, H. E. Ahles and R. 3. Leisner, 32171,
July 21, 1957 (NCU). Colleton Co.: 1 mi. S of Ritter along
Ashepoo River, H. E. Ahles and C. R. Bell, 21049, Oct. 12, 1956
(NCU). Greenwood Co.: near Troy, J. Davis, sn,, Aug. 15, 1921
(TEX); Saluda River E of Dyson, A. E. Radford, 26612, July 7,
1957 (NCU). Darlington Co.: along Lauther's Lake, B. E. Smith, V?2, Aug. 3* 19^0 (NCU). Williamsburg Co.: 8 mi. SE of Gourdin,
R. K. Godfrey and R. M. Tryon, Jr., k jk , July 10, 1939 (GH, MO); k mi. W of Gourdin near U. S. hwy. 52, A. E. Radford, 28360,
Aug. 22, 1957 (NCU). Tennessee: Lake Co.: Reelfoot Lake, 0. E0 and G. K. Jennings, 68 3* in 192^ (TENN). Texas: Comal Co.:
New Braunfels Comanche Spring, F. Lindheimer, 95^* Aug., I85O
(F, GH, MO, NY, PH, TEX,US). Dallas Co.: Dallas, J. Reverchon,
^69 (F 832995» MO 1767297). Gonzales Co.: edge of ash bog,
Ottine, E. R. Bogusch, 2^-68, Sept. 18, 1927 (TEX); Hillside above
Karnstodt's marsh, E. C. Tharp, 7380, Aug. 5* 1929 (TEX'). Gregg
Co.: C. L. York, sn., Sept. 1, 19^1 (TEX). Harris Co.: Crosby,
G. L. Fisher, 13* Sept. 25, 1921 (US). McLennan Co.: below
Lake Waco, L. D. Smith, 1037* Sept. 30, 19^7 (F). Tarrant Co.: near Trinity River, A. Ruth, 331* Oct. 1, 1920 (F, GH). Travis
Co.: Austin, B. C. Tharp, 900, July, 1920 (US); Austin, E. D.
Schulz, 351* July, 1920 (US). Virginia: Brunswick Co.: Seward
Forest near Triplett, K. L. Fernald and J. B. Lewis, 1^75^*
Sept. 12, 1 9 ^ (GH, PH). Fairfax Co.: Curtis, sn., (F 1 3 6 6 5).
Gloucester Co.: Dutton, E. T. 'Wherry and F. V/. Pennell, 12758,
Sept. 1, 1925 (MO, PH); S of White Marsh, M. L. Fernald and B.
Long, 8892, July 22, 1938 (GH). Henrico Co.: Westhampton, M. L.
Fernald et al., 6722, Aug., 1936 (GH). Isle of Wight Co.:
Burwell's Bay, James River below Fort Boykin, M. L. Fernald and
B. Long, 1 2 8 7 6, Aug. 30, 19^0 (GH). King George Co.: Potomac
River near Farview Beach, H. H. Iltis, 2258, Sept. 1-2, 19^3 (US).
Norfolk Co.: Ocean view near Norfolk, J. R. Churchill, sn., 88
Aug. 9, 1927 (MO). Princess Anne Co.: N of Blackwater River,
M. L. Fernald and B. Long, *f253> Aug. 7> 193^ (GH); Great Heck,
M. L. Fernald and B. Long, 5118, Sept. 5» 1935 (GH). Prince
William Co.: Quantico, Tidestrom, 76 2 8, Aug. 21, 1915 (US).
Shenandoah Co.: River E of Woodstock, H. Allard, 93&3j Aug. 10,
19^1 (US). Sussex Co.: Nottoway River at double bridge 6 mi.
NW of Jarratt (GH).
(N) Polymnia uvedalia L. var. uvedalia comb. nov.
Polymnia uvedalia L. var. genuina Blake. Rhodora. 19:^7* 1917. (Syntype FI, GHI, M01, NCU!, NYI, US!).
Peduncles and branches of the inflorescence densely stipitate-glandular, pilose hairs few or usually none.
Type: Probably collected in Virginia by Frederick
Gronovius.
Specimens Examined: BERMUDA: Paynters vale, D. W.
Marble, 7^9,. Mar. 29, May 3, 1909 (NY). Alabama: Etowah Co.:
Gadsden, G. R. Vasey, sn., 1878 (NY). Lee Co.: F. S. Earle,
2120, Aug. 7, 1899 (F)« Madison Co.: Monte Sano, C. F. Baker, sn., Sept. 15, 1897 (OS). Delaware: New Castle Co.: Alapokas woods on Brandyv/ine Creek, R. R. Tatuall, 350^, Aug. 31» 1937
(GH). Sussex Co.: Rehoboth, J. R. Churchill, sn., Sept. 13,
1908 (GH). District of Columbia: opposite Analostan Island,
E. L. Morris, 311, Sept. 7, 1899 (F); C. S. Williamson, sn.,
1905 (PH). Georgia: Bartow Co.: 3 S of Allatoona Dam,
W. H. Duncan, 8*fl6, June 19, 19^8 (NY). Fannin Co.: H. H.
Smith, 2 5 8 5, Aug. 5» 1909 (F)o Habersham Co.: J. IC. Small, sn., Sept. 1-3, 1 8 9 ^ (NY). Lincoln Co.: Price's Island in Savannah,
W. H. Duncan, 9 8 2 8, June 29, 19^9 (TENN). Rabun Co.: road to
Dillard, A. J. Sharp et al., 29^7, July 28, 193^ (TENN).
Stephens Co.: NNW of Currahee Mt., W. H. Duncan, 1174-1, Aug. 10,
1950 (MO). Whitfield Co.: E. M. Harper, 3 1 6, July 27, 1900
(NY, US). Indiana: Jefferson Co.: Hanover, J. M. Coulter, sn.,
Aug. 11, 187^ (F). Lawrence Co.: N of Huron, R. M. Kriebel,
3592, Aug. 11, 1935 (GH, MO). Orange Co.: 3 mi. W of Paoli,
C. C. Dean, 1734-7, July 14, 1915 (GH). Illinois: Union Co.:
Anna, A. B. Seymour, 1209, Aug. 16, i860 (GH). Kentucky: Bell
Co.: Mtns. near Pineville, K. K. Mackenzie, 935, Aug., Sept.,
1904 (MO). Edmondson Co.: Mammoth Cave, A. H. Howell, 5 6 9,
July 4, 1909 (US). Estill Co.: 1 mi. W of Harg., M. E. Wharton,
3136, July 20, 1938 (NY). Harlan Co.: along Poor Fork of Cum berland River, T. H. Kearney, Jr., 251, Aug., 1893 (F, GH, MO,
NCU, NY, OS, US). Kenton Co.: Banklick Creek, J. M. Lea, sn.,
July, 1870 (PH). Mercer Co.: Burgin, A. King, 2 8 7, Aug. 24,
1916 (F). Ohio Co.: mi. W of Deamville on rt. 54, T. R.
Fisher, 2023, Aug. 9, 3.960 (OS). Powell Co.: V/2 mi. S of West
Bend, M. E. Wharton, 2343, July 30, 1938 (TENN). Warren Co.:
Bowling Green, S. F. Price, sn., Aug., 1892 (MO). 'Wayne Co.:
Beaver Creek SW of Monticello, L. B. Smith and A. R. Hodgdon,
3913, July 12-14-, 1937 (GH); NW of Monticello, L. B. Smith and
A. R. Hodgdon, 40l8, July 17, 1937 (GH). Baltimore Co.: W. R.
Jones, sn., Aug. 30, 1906 (F). Montgomery Co.: near Plummer
Island above Cabin John, P. H. Dowell, 1?4S, Sept. 28, 1902 (GH); NW of Cabin John, A. Chase, 2627, Sept. 10, 1904 (F). Washington
Co.: Antietam, C. D. Lippincott, sn., Sept. 17, 1896 (PH).
Mississippi: Attala Co.: Natchez Trace Parkway, W. B. McDougall
1397, July 14, 19^7 (US). Missouri: Barry Co.: W. Trelease, sn in 1898 (MO). New Jersey: Hudson Co.: Weehawken, W. M. Van
Sickle, sn., Aug. 20, 1895 (OS); Weehawken, T. F. Allen, sn.,
(NY); 'Weehawken, T. F. Allen, sn., Aug. 8, 3.864 (GH). Sussex Co.
2 mi. SW of Wallpack Center, J. L. Edwards, sn., Aug. 12, 1951
(NY). New York: Cayuga Co.: above falls Springport, A. J.
Eames et al., 1 3 1 5 6 , Sept. 14, 1919 (GH). Erie Co.: Cattaraugus
Indian Reservation, A. E. Perkins, sn., Aug. 6, 1950 (US).
Ontario Co.: W. E. Aikin, sn., (PH); J. Hail, 2 8 9, 1828-1834
(F). Yates Co.: H. P. Sartwell, sn., (F 476011). North Caro lina: Alexander Co.: roadbank near Wilkes Co. line, Radford
and Stewart, 1 6 0 6, July 23, 1940 (NCU); 5 mi. N of Taylorsville,
A. E. Radford, 1 8 1 8 0 , Sept. 15, 1956 (NCU); 5J£ mi. N of Tajiors-
ville, A. E. Radford, 18179, Sept. 15, 1956 (TENN). Ashe Co.:
3.2 mi. E of Bena, A. E. Radford, 3 8 6 8 3, Aug. 1, 1958 (NCU).
Caldwell Co.: eastern slopes of Blowing Rock Mtn., J. K. Small
and A. A. Heller, 423, July 31, 1891 (F, GH, MO, NY). Gaston
Co.: .6 mi. SW of Gastonia, J. R. and J. M. Wells, 248, July 27,
1961 (OS). Graham Co.: 3 mi. NW of Pocahontas, A. E. Radford,
16018, Aug. 19, 1956 (NCU). Haywood Co.: Waynesville, C. Hall,
501, July 15, 1882 (US); Waynesville, C. Hall, 522 and 523, in
1882 (US); near Eagles Nest, P. C. Standley, 5581, Sept. 7, 1910
(US); Twin Spring Mt., Coker et al., sn., Aug. 28, 1926 (NCU). Iredell Co.: .2 mi. E of Monbo, H. E. Ahles and J. A. Duke,
48858, Aug. 7, 1958 (NCU). Jackson Co.: Nantehala, L. N. John son, 1380, July 21, 1888 (F); E of Sylva near railroad crossing,
E. Quarterman, 112, July 18, 1938 (PH). McDowell Co.: 1 mi. E of Little Switzerland, K. K. Wiegand and W. E. Manning, 3315,
Aug. 23, 1927 (GH); .4 mi. SE of jet. N. C. 26 and U. S. 221 on
N. C. 26, H. E. Ahles and C. R. Bell, 17763, Sept. 1, 1956 (NCU).
Madison Co.: Warm Springs, J. H. Redfield, 5612, Aug. 23, 1876
(MO); .9 mi. NW of jet. U. S. 19 and 23 on U. S. 23, H. E. Ahles and J A. Duke, 46305, July 1**, 1958 (NCU). Mecklenburg Co.:
Irvins Creek 3.1 mi. N of Matthews, H. E. Ahles and J. A. Duke,
4 8 583, Aug. 6, 1958 (NCU). Mitchell Co.: 3.9 mi. SW of Bakers- ville, H. E. Ahles and J. A. Duke, 47199, July 23, 1958 (NCU).
Polk Co.: near Columbus, E. C. Townsend, sn., Sept. 2, 1897
(US); Melrose, D. C. Peattie, 1104, Aug. 8, 1921 (F, NCU). Swain
Co.: near Swayney, J. Mooney, sn., Sept. 9, 1913 (US)j Ridge near Bryson City, L. Barksdale, sn., July 1, 1935 (NCU); V/z mi.
S of Lauda near N. C. 28, A. E. Radford, 17277, Aug. 24, 1956
(NCU). Watauga Co.: .6 mi. W of Ashe-Watauga Co. line, H. E.
Ahles and J. A. Duke, 47724, July 25, 1958 (NCU). Wake Co.: near Raleigh, R. K. Godfrey, sn., Aug. l8, 1937 (GH). Yancey Co.
Daybrook-Burnsville, A. E. and L. M. Radford, 2 5 8 7, July 9, 1946
(NCU). Ohio: Adams Co.: between Serpent Mound and locust grove
A. R. Harper, sn., July 23, 1923 (OS). Athens Co.: Waterloo
Game Preserve, W. B. Price, sn., Aug. 1, 1954 (OU). Belmont Co.:
Steubenville, H. N. Mertz, sn., Aug. 23, 1878 (US). Clarke Co.: Little Honey Creek, G. H. Shull, sn., Aug. 27, 1898 (OS).
Clermont Co.: Loveland, D. L. James, sn., (OS 38890). Coshocton
Co.: H. N. Moldenke, 13054, July 8, 1942 (OS). Cuyahoga Co.:
W. Krebs, sn., 1896 (OS). Gallia Co.: Vinton, W. A. Kellerman, sn., July 10, 1901 (OS). Highland Co.: Bartley and Pontius, sn.,
July 26, 1931 (OS); Brush Creek Twp., sn., Sept. 8, 1931 (OS);
Concord Twp., K. M. Roads, sn., June 24, 1938 (OS); above Baker's
Fork near Fort Hill, A. Cronquist, 4044, Aug. 12, 1945 (NY).
Jackson Co.: bluff along Salt Creek, F. Bartley and L. L.
Pontius, sn., July 14, 1935 (OS). Lawrence Co.: Symmes Creek
Valley, W. A. Kellerman, sn., Oct. 28, 1892 (OS). Meigs Co.:
Salem Twp., C. H. Jones, sn., Sept. 1, 1935 (OS). Noble Co.:
Summerfield, H. Eeymer, sn., June 28, 1896 (OS). Scioto Co.:
McDermott, B. Wigginton, sn., Aug. 15, 1935 (PH); McDermott,
D. Demaree, 11511, Aug. 15, 1935 (i'O, OS, US); C. H. Jones, sn.,
July 31, 1937 (OS). Tuscarawas Co.: 2/4 mi. N of New Phila delphia, J. Sterki, sn., Sept. 25, 1925 (OS). Vinton Co.: Moon- ville, V/. P. Porter and R. S. Wamsley, sn., July 22, 1939 (OS);
Vinton Furnace Experimental Forest, G. W. Hall, 19^3, July 29,
1956 (OU). Oklahoma: Johnston Co.: near Tishomingo, H. W.
Houghton, 3518, Oct., 1915 (GH). Pennsylvania; Allegheny Co.: near Natrone, J. A. Shafer, 782, July 21, 1900 (GH, PH). Bedford
Co.: 2)4 mi. S of Hyndman, D. Berkheimer, 4028, July 31, 1943
(TEX,US). Delaware Co.: Wawa, F. W. Pennell, sn., in 1904 (US).
Fayette Co.: J. Bright, 284, Aug. 2, 1936 (TEX). Franklin Co.:
Mercersburg, T. C. Porter, sn., Aug. 19, 1948 (F, NY). Greene Co.: Perry Twp., 3. S. Dickey, 129, Aug. 20, 1922 (GH).
Huntington Co.: T. C. Porter, sn., Aug. 18, 1882 (NY). Lan caster Co.: A. P. Garber, sn., in 1868 (F); % mi. SE of Millers- ville, F. A. Tanger, sn., Aug. 1, 1938 (GH). Northampton Co.:
Easton, T. C. Porter, sn., Aug. 30, 1869 (GK); Easton, T. C.
Porter, 3k2k, Sept., 1870 (MO, NY); Easton, T. C. Porter, sn.,
Aug. 10, 1886 (F); Chestnut Hill, T. C. Porter, sn., July 12,
1890 (US); Easton, T. C. Porter, sn., July 30, 1893 (US). York
Co.: J. Crawford, 91092 (PH 5036^). South Carolina: Abbeville
Co.: 5.6 mi. ESE of Abbeville, A. E. Radford, 25887, June 29,
1957 (NCU). Anderson Co.: J. Davis, 78^+3, Aug. 29, 1917 (M0);
J. Davis, 7989, Aug. 23, 1917 (PH); Concord Church, J. Davis, sn.,
Aug. , 1920 (TEX). Jasper Co.: 1.7 mi. S of jet. rts. J>k and
31 on rt. 3k, C. E. Bell, 3969, June 29, 1956 (NCU). Kershaw
Co.: 5 “I* WNW of Shamokin, A. E. Radford, 2770k, July 28, 1957
(NCU). Oconee Co.: 3 mi. N of Tamasee, A. E. Radford, 17837,
Sept. 2, 1956 (NCU). Pickens Co.: W of Pickens, A. E. Radford,
1 6 5 8 7, Aug. 22, 1956 (NCU). Williamsburg Co.: 8 mi. SE Gourdin,
R. K. Godfrey and R. M. Tryon, Jr., ^5^, July 10, 1939 (F, TENN).
Texas: Tarrant Co.: A. Ruth, 331, Sept. 2k, 1925 (US). Tennes see: Blount Co.: near Weare's Cove Road, A. J. Sharp and H. M.
Jennison, 53^, Aug. 1, 1935 (TENN); Chilhowee Mt., A. J. Sharp and H. M. Jennison, 876, Aug. 22, 1935 (TENN); Bacon’s Ferry,
H. N. Moldenke, 19309, Aug. 29, 19^7 (NY); above Chilhowee Dam,
A. J. Sharp and N. Veloira, 20966, Oct. 7, 1956 (TENN); Cades
Cove, A. J. Sharp, 29728, Oct. 10, 1961 (OS). Campbell Co.: 94
Stiner Bridge along Powell River, Kelly, 2452, July l8, 193^
(TENN). Cheatham Co.: Craggie Hope, H. K. Svenson, 4216, Aug.
22, 1930 (GH). Claiborne Co.: Powell River Valley, A. J. Sharp et al., 3 0 8 5 6, Sept. 30, 1962 (TENN). Coffee Co.: H. R. DeSelm,
604, June 28, 1955 (TENN). Davidson Co.: Baker's Station, H.
Eggert, sn., July 13, 1897 (MO, NY); Joelton, II. K. Svenson,
120, July 16, 1922 (GH, PH); H. K. Svenson, 7331, July 23, 1935
(GH, TENN). Franklin Co.: Cowan, 788b, Aug. 21, 1897 (GH, NY,
US). Grainger Co.: Tampico, Morrison, sn., Aug. 21, 1936
(TENN). Knox Co.: T. H. Kearney, Jr., sn., Aug. 1891 (OS);
Knoxville, A. Ruth, sn., Aug., 1893 (US); A. Ruth, 52, Aug. 15,
1894 (GH); A. Ruth, sn., Sept., 1894 (NY); A. Ruth, 270, Aug.,
1895 (F, MO, PH); A. Ruth, 40l8, July, 1897 (GH); Knoxville City
Park, E. Norton, sn., June 10, 1940 (TENN); bank of Tenn. River,
Knoxville, N. Russell, NR-3026, Sept. 9, 1946 (TENN); near East
Tenn. State Hosp., A. J. Sharp, 22638, Sept. 22, 1957 (TENN); between Neyland Drive and Loudon Lake, Knoxville, A. J. Sharp,
24946, Oct. 5, 1958 (TENN). Lawrence Co .: near West Point,
A. J. Sharp et al., 9756, July 12, 1948 (TENN). Marion Co.:
5 mi. SE of Monteagle by U. S. rt. 4l, E. S. Ford and N. H.
Russell, 2215, Sept. 1 8 , 1946 (TENN). 6 mi. SE Monteagle, J. R. and J. M. Wells, 253> Aug. 1, 1961 (OS). Monroe Co.: near Union
Grove, A. J. Sharp and W. H. Ellis, 29699* Oct. 8 , 1961 (OS).
Polk Co.: E. T. Wherry and F. W. Pennell, 14025, Sept. 14, 1927
(PE). Sumner Co.: Old Hickory Reservoir near Old Indian Lake,
J. R. and J. M. Wells, 286-291, Sept. 19, 1962 (OS). Unicoi Co.: Nolichucky River near Erwin, A. J. Sharp and J. K. Underwood,
31971 Aug. 19, 193^ (TENN). Wayne Co.: k mi. W of 'Waynesboro,
H. H. Welch, sn., Aug. 10, 193^ (TENN). White Co.: A. J. Sharp et al., 5587, July 31, 19^+7 (TENN). Virginia: Amherst Co.:
James River Gorge between Snowden and Balcony Falls, R. S. Freer
2257, Aug. 16, 195^ (GH). Augusta Co.: near Fordwick and
Craigsville, E. S. Steele, 90, Aug. 30, 1913 (US). Botetourt
Co.: on rt. 622 near Arcadia, R. S. Freer, 2 1 8 9 , July 28, 195^
(GH). Fairfax Co.: Arlington, Tidestrom, 8131, Oct. 11, 1916
(GH). Loudon Co.: near Harper's Ferry, R. McVaugh, 6613,
Sept. 6, 19^2 (NY). Princess Anne Co.: Great Neck, M. L.
Fernala and B. Long, 5118, Sept. 5» 1935 (US). Rockbridge Co.:
J. R. Churchill, kO, Aug. 10, IS 2k (MO); near Lexington, J. R.
Churchill, sn., Aug. 28, 192^ (NO); near Lexington, J. R. Church ill, sn., Aug. 21, 192^ (GH). Shenandoah Co.: Strasburg, H. A.
Allard, 7305, Aug. 2k, 1939 (GH, US). Smyth Co.: S fork of
Holston River, St. Clair's Bottom, J. K. Small, sn., July 30,
1892 (F, GH, LSU, NCU, NY, PH,.US); Middle Fork of Holston River
J. K. Small, sn., Aug. 6, 1892 (F, MO, US). 'West Virginia:
Cabell Co.: Pleasant Valley, F. A. Gilbert, 7^3» Aug. 1, 1938
(F, GH, 110, NY, PH, TENN, US). Calhoun Co.: 3.1 mi. from
Gilmer-Calhoun Co. line on rt. 33> J« R. Wells, 235, July 23,
I960 (OS). Marion Co.: kmi. N of Fairview, S. S. Rickey, 101,
Aug. 15, 1922 (GH). Mingo Co.: near Varney School, E. E.
Berkley, 973, July 7, 1930 (GH, MO). Ohio Co.: Wheeling, H. N.
Mertz, sn., Aug. 23, 1878 (NY); Wheeling, H. W. Mertz, 1380, (F 96
208l*f*f and F 208l*f3)» Fendleton Co.: Hermit Island Smoke Hole,
E. E. Berkley, 1^76, Aug. 7* 1930 (GH, MO). Randolph Co.:
Tygard Valley, C. F. Millspaugh, ^77* July 11, 1890 (NY); Nut- tall, C. F. Millspaugh, 1111, Aug. 11, 1891 (NY). Summers Co.:
Camp Brookside, M. C. Beals, sn., Aug. 13-15* 1958. Tucker Co.:
Canoan Valley, H. A. Allard, 1 1 5 2 8 , Aug. 3* 19^5 (US).
(0) Polymnia uvedalia L. var. floridana Blake. Rhodora. 19;^8. 1917.
Peduncles and inflorescence branches sparsely pilose and glandular (Figs. 15 and 16).
Type: Florida: Brevard Co.: Thicket, Indian River region, A. Fredholm, 5 6 2 6, Dec. 2, 1902 (Holotype GHl, Isotype
US!); Chapman, 788c, without date or locality in Florida (Para- type GHl, NYt, US!). Leon Co.: at Tallahassee, G. V. Nash,
2327, Aug. 7-9, 1895 (Paratype FI, GHl, MOi, NYI, US I).
The var. floridana seems to be found mainly in the southern and southeastern portion of the species range. An oc casional record appears in the northern portion of the species range, however, as in southern Indiana and Delaware. Of the specimens examined there were fewer of this variety than of any of the other two.
Examples of var. floridana can be found which lead one to think that this variety is intermediate between var. uvedalia and var. densipilis. The peduncles of var. floridana are on oc casion nearly glabrous except for the sparse puberulence and a few small glands. 97
Fig. 1 5 .— Polymnia uvedalia var. floridana (Peduncle of Holotype) Scale: 5X
The photograph of P. uvedalia var. floridana (Fig. 16) shows typical leaf outlines of the species.
Specimens Examined; BERMUDA: near Bailey's Bay, S.
Brown, *f9^i Feb. 10, Mar. 9? 1908 (F. NY); Paget, F. S. Collins,
3^8, Aug. 21, 1913 (GH). Alabama: Clay Co.: T. G. Harbison, sn., July, 1902 (GH). Cullman Co.: 3 mi. N of routes 65 and 31 near Hanceville, T. R. Fisher and J. M. Speer, 21^7* Sept. 11,
1962 (OS). Madison Co.: Monte Sano, C. F. Baker, 1358, Sept. 13»
1897 (MO, NY). Delaware: New Castle Co.: Wilmington, E. 98
* -
Fig. 16.— Polymnia uvedalia var. floridana Scale: .29X
Tatuall, sn., Aug., 18^2 (GH); near Wilmington, W. M. Canby, sn.,
Aug., 1873 (F); Woods on Brandywine Creek N of Wilmington, R. R.
Tatuall, 350^, Aug. 31» 1937 (GH). Florida: Citrus Co.:
Komosassa, Combo, sn., Sept. 10, 1898 (US). Duval Co.: Ft0
George Is., L. H. Lighthipe, 531* Oct. 26, 1897 (NCU, NY).
Franklin Co.: Apalachicola, sn. (MO 783801); Apalachicola,
Chapman, 788c, July-Aug., 1897 (GH, NY, US). Levy Co.: W. A.
Murrill, sn., June 18, 1939 (MO). Liberty Co.: Rock Bluff
Landing, Apalachicola River, K. M. Wiegand and W. E. Manning,
331^, Aug. 6, 1927 (GE); Apalachicola River at Aspalaga, R. Itral, 3090, July 25, 1956 (GH). Manatee Co.: 1 mi. N of Manatee
River, J. H. Simpson, 108, Sept. 12, 189 O (F, US). Putnam Co.:
Palatka, A. K. Curtiss, sn., June, 188? (NCU 7 8 6 8 0). Volusia
Co.: S of Daytona in Hammock, J. K. Small, 8 6 9 0, May 8, 1918
(TENN, US); Daytona, E. H. Butts, sn., Sept. 6, 19^3 (GH).
Georgia: Bartow Co.: Allatoona Creek 3 mi. S of Allatoona Dam,
W, H. Duncan, 84l6, June 19, 1948 (GH, MO, US). Berkeley Co.:
Bank of Santee Canal 1 mi. N of Moncks Corners, K. M. Wiegand and W. E. Manning, 3311* July 13, 1927 (GH). Catoosa Co.: Little
Sand Mt. 3 ml* NE of Ringgold, A. Cronquist, 5 6 0 8, Aug. 8, 1948
(F). DeKalb Co.: base of Stone Mtn., J. K. Small, sn., July 4,
1893 (F); Stone Mtn., H. Eggert, sn., July 24, 1897 (MO).
Douglas Co.: 2 mi. E of Villa Rica, A. Cronquist, 5565* July 27,
1948 (GH, MO, NY, PH, US). Early Co.: Chattahoochee River,
R. M. Harper, 1225, Aug. 14, 1901 (GH, MO, NY, US). McIntosh
Co.: S end of Sapelo Island, W. H. Duncan, 20324, July 19, 1956
(GH, NCU, TEX, US). Upson Co.: along Double Branch SE of Wood bury, A. Cronquist, 5504, July 17, 1948 (GH, MO, NY, PH, US).
Wilkes Co.: Kettle Creek 6 mi. SW of 'Washington, W. H. Duncan,
11409, July 1 8 , 1950 (GH, MO). Illinois: Alexander Co.: McClure,
E. J. Palmer, l66ll, Sept. 29, 1919 (MO). Jackson Co.: G. H.
French, sn., Sept. 21, 1878 (F,US). Kentucky: Edmondson Co.:
Mammoth Cave, H. W. Lix, 307, July 25, 1947 (US). Lyon Co.:
Kuttawa, W. W. Eggleston, 5278, Sept. 27, Oct. 9, 1909 (NY).
Louisiana: Assumption Co.: Paincourtville, E. Marionneaux, sn.,
July, 1917 (LSU). Lafayette Co.: Girard Woods, Lafayette, G. B. 100
Claycomb, sn., Aug. 10, 1942 (LSU). Lincoln Co.: Sugar Creek,
J. A. Moore, 5444, Sept. 12, 1953 (LSU). Terrebonne Co.: 12 mi, below Houma (LSU). Mississippi: Harrison Co.: Biloxi, S. M.
Tracy, 4329, Aug. 12, 1898 (F, MO, NY, OS, US); Biloxi, F. E.
Lloyd and S. M. Tracy, 526, Sept. 12, 1900 (NY); Biloxi, S. M.
Tracy, 6972, July 4, 1900 (NY). Missouri: Barry Co.: Roaring
River, W. Trelease, sn., Sept. 7, 1898 (MO). Ozark Co.: 4 mi.
NE of Dormis and 4 mi. SE of Dora, J. A. Steyermark, 15578,
Sept. 28, 1934 (MO). Shannon Co.: B. F. Bush, sn., Sept. 15,
1888 (MO). St. Louis Co.: G. Engelmann, sn., Sept., l84l (MO).
North Carolina: Alexander Co.: 5# mi. N of Taylorsville, A. E.
Radford, 18179, Sept. 15, 1956 (TENN). Caldwell Co.: 2 mi. NW of Collettsville near Johns River, A. E. Radford, 14953, Aug. 4,
1956 (NCU). Currituck Co.: 3 mi. N of Waterlily, H. E. Ahles and J. A. Duke, 4812, July 31, 1958 (NCU). Forsyth Co.: 1.9 mi.
NW of Vienna toward Dozier, H. E. Ahles and J. A. Duke, 49083,
Aug. 8, 1958 (NCU). Gaston Co.: 1 mi. E of Mt. View near
Crowders Mt., H. E. Ahles, 18920, Sept. 17, 1956 (NCU). Guil
ford Co.: Lower Richland Creek, L. Melvin, sn., Sept. 4, 1955
(NCU). Lee Co.: 1 mi. W of Moncure below Lockville Dam, L. S.
Beard, 1192, Aug. 2, 1955 (NCU). Lincoln Co.: 3 .8mi. S of
county line on U. 3. route 321, C. R. Bell, 15239, Sept. 10,
1958 (NCU). Martin Co.: 4.2 mi. NW of Williamstown, A. E. Rad
ford, 39502, Aug. 9, 1958 (NCU). Mecklenburg Co.: Irvins Creek
3.1 mi. N of Matthews, H. E. Ahles, 4 8 5 8 3, Aug. 6, 1958 (NCU);
1 mi. N of Matthews, J. R. and J. M. Wells, 247, July 27, 1961 101
(OS). Macon Co.: Highlands, C. Boynton, sn., Aug., 1888 (US).
Orange Co.: II. L. Blomquist, 6997j Aug. 22, 1932 (PH). Pasquo tank Co.: 1.2 mi. E of Nixonton toward Weeksville, H. E. Ahles and J. A. Duke, ^8115, July 31, 1958 (NCU). Pitt Co.: 1.1 mi.
N of Grimesland, A. E. Radford, 3963^, Aug. 10, 1958 (NCU).
Randolph Co.: Uwharrie River V/ of Asheboro, C. R. Bell, 1^192,
Aug. 1, 1958 (NCU, TEX). Stanly Co.: Falls of Yadkin River,
J. K. Small, sn., Aug. l8, 1892 (F)j Long Creek near Albermarle on N. C. rt. 27, H. E. Ahles and R. S. Leisner, 19817, Sept. 2^f,
1956 (NCU). Wake Co.: Raleigh, R. K. Godfrey, 5927, Aug. 8,
1938 (GH, NCU). Oklahoma: Johnston Co.: H. W. Houghton, 3518,
Oct., 1915 (MO). Pennsylvania: Lancaster Co.: near mouth of
Tucquan, J. K. Small, sn., Aug. 7, 1890 (F); A. MacElwee, 1228,
Aug. 31, 1899 (NCU). South Carolina: Anderson Co.: near Ander son, J. Davis, 285, Aug. 4, 1920 (MO). Beaufort Co.: 3 mi* N of Beaufort on U. S. rt. 21, H. E. Ahles and C. R. Bell, 18030,
Sept. 6, 1956 (NCU). Berkeley Co.: 4 mi. S of Moncks Corner,
H. E. Ahles and J. G. Haesloop, 30730, July 5, 1957 (NCU).
Cherokee Co.: 1.5 mi. S of jet. Co. 99 and U. S. 29A, H. E.
Ahles and J. G. Haesloop, 308 9 0, July 11, 1957 (NCU). Darling ton Co.: back of Darlington Country Club, B. E. Smith, 602,
July 21, 19^1 (NCU). Dorchester Co.: Cattle Creek S3W of
Reevesville, R. S. Leisner and H. E. Ahles, 31937, July 20, 1957
(NCU). Edgefield Co.: Beaverdam Creek, 3.5 mi. SW of Cleora,
A. E. Radford, 2 6 3 6 2, July 1, 1957 (NCU). Fairfield Co.: W
fork of Little River between Shelton and Woodward, C. R. Bell, 9337, June 26, 1957 (NCU). Georgetown Co.: Pee Dee River, 2.6 mi. NE of Yauhannah, A. E. Radford, 28637, Aug. 23, 1957 (NCU).
Jasper Co.: 4 mi. E of Coosawhatchie, K. M. Weigand and W. E.
Manning, 3312, July 20, 1927 (GH). Lexington Co.: Congaree
River, Cayce, A. E. Radford, 27018, July 9, 1957 (NCU). Saluda
Co.: 4.5 mi. NW of Batesburg, A. E. Radford, 26985, July 8,
1957 (NCU). Spartanburg Co.: Pacolet River below Rainbow Lake dam, C. R. Bell, 10393, Oct. 5, 1957 (NCU). Williamsburg Co.:
8 mi. SE of Gourdin, R. K. Godfrey and R. M. Tryon, Jr., 454,
July 10, 1939 (NY, PII, US). York Co.: Dam Creek E of Clover,
H. E. Ahles, 34653, Sept. 19, 1957 (NCU). Tennessee: Cocke Co.:
English Mtn., J. A. Chapman, sn., Aug. 4, 1956 (TENN). Franklin
Co.: H. Eggert, sn., Sept. 8, 1898 (MO). Grundy Co.: 2 mi. NW
Monteagle on U. S. rt. 4l, J. R. and J. M. Wells, 2 5 6, Aug. 1,
1961 (OS). Hardin Co.: NE of Gillises Mills, A. J. Sharp et al.
9472, July 10, 1948 (TENN, TEX). Knox Co.: Roaring Springs,
A. J. Sharp, 3939, July 8, 1935 (TENN). Polk Co.: E of Ocoee by U. S. rt. 64, E. S. Ford and N. II. Russell, 2337, Sept. l8,
1946 (TENN). Virginia: Bedford Co.: A. H. Curtiss, sn., Aug.
25, 1871 (NY). Fairfax Co.: Mt. Vernon, J. W. Chickering, sn.,
(F 205112); 7 mi. S of Alexandria Hunting Creek, F. R. Fosberg,
23957, Sept. 12, 1945 (NCU). James City Co.: Williamsburg, E. J
Grimes, 4219, Aug. 11, 3.921 (GH). Princess Anne Co.: Munden,
K. K. Mackenzie, 1669, Mar. 19,1905 (MO); Great Neck, M. L.
Fernald and B. Long, 5116, Sept. 5, 1935 (GH). Shenandoah Co.:
E of Maurertown on Shenandoah River, H. A. Allard, 9371, Aug. 10, 103
1941 (US). Surry Co.: James River below Sunken Meadow Beach,
M. L. Fernald and B. Long, 9197, Aug. 23, 1938 (Gil); NW of
Chippokes Beach of Cobham Bay, M. L. Fernald and B. Long, 12877,
Aug. 25, 1940 (GH, US).
(P) Polymnia quichensis Coulter. Bot. Gaz. 20:48. 1895*
Polymnia latisquama Blake. Wash. Acad. Sci. l6:421<> 1926. (Holotype US!, Isotype GHl).
Stems up to 3 meters tall, terete, smooth or sparingly puberulent; leaves lanceolate or ovate-lanceolate, 30 cm long or less, up to 8 mm wide, serrate, some teeth about 1 mm long and callous-tipped, others smaller, some lower leaves with short lobes separated by shallow sinuses, above glabrous to scabrous, beneath sordid-pilose in axils of main veins or puberulent along the veins, bases oblique or attenuate, apex acuminate to very long-acuminate; peduncles glabrous or sparsely puberulent; heads
few or solitary; phyllaries broadly lanceolate to round-ovate, up to 1 .7 cm wide, 2 cm long; ray flowers 7-1 5 , corollas yellow,
3 cm long or less, up to 9 mm wide; disc flowers about 1 cm long,
corolla yellow; ray pales broadly lanceolate, as large as 13 mm long, 8 mm wide; pales of disc flowers linear, about 7-8 mm long and 1.5 mm wide; fruit black, spherical, 5-6 mm in diameter (Fig.
17).
Type: GUATEMALA: Quiche: Chiul, 8000 ft. alt., Heyde and Lux, 3375, Apr., 1892 (Holotype F!, Isotype GHl, MOI, NYI,
US!). 104
Fig. 17.— Folymnia quichensis (Holotype) Scale: .30
I examined one unusual isotype which is deposited in the
Gray Herbarium. This specimen, Heyde and Lux 3375 had two or three smaller and separate petaloid structures associated with each ray flower. These petaloid parts were about 7 mm long and lo5 mm wide. I am unable to interpret their morphology. They might be petaloid stamens or may represent a second corolla present on each ray flower— hence biligulate ray corollas.
There is obvious overlap in each of the characteristics used by Blake in separating P. latisquama from P. quichensis. In 105 treating these two as one species I suggest that Blake's P. latisquama which is from Costa Rica may be a clinal variant of
P. quichensis otherwise recorded only from Guatemala. The fact that no specimens of P. quichensis were observed from countries between Costa Rica and Guatemala is probably due to lack of col lection rather than absence of the plants in those interlying regions. Specimens of Polymnia collected from Nicaragua were almost completely absent in some of the larger United States herbaria. I have examined only one specimen of Polymnia from that countryl
Specimens Examined; COSTA RICA; Cartago; S slope of
Volcan de Turrialba, near Finca del Volcan de Turrialba, 2000-
2400 m. alt., P. C. Standley, 35340, Feb. 22, 1924 (GH, US);
Rio Birris, S slope of Volcan de Irazti, P. C. Standley, 35^12,
Feb. 23, 1924 (GH, US); on road to Volcan Irazfi, M. A. Chrysler,
5544, Aug. 2 3, 1940 (F). GUATEMALA: Huehuetenango; 3Yz m i. E of San Mateo Ixtatan, 2900 m. alt., J. A. Steyermarlc, 48502,
July 10, 1942 (F); at Cruz de Lim6n, between San Mateo Ixtatan and Nuca, 2600-3000 m. alt., J. A. Steyermark, 49833, July 31»
1942 (F). Jalapa; near Soledad, between Jalapa and Mataquescu- intla, 2000-2500 m. alt., J. A. Steyermark, 32621, Dec. 4, 1939
(F). Quich.6; Nebaj, 85OO ft. alt., A. F. Skutch, 1704, Nov. 17,
1934 (GH). San Marcos; 2700 m. alt., P. C. Standley, 68575*
Mar. 14, 1939 (F); Barranco Eminencia, 2700 m. alt., P. C. Stand ley, 68555, Mar. 14, 1939 (F); 2100-2400 m. alt., P. C. Standley,
68472, Mar. 14-15, 1939 (F); between San Sebastian and Todos 106
Santos on slopes of Volcan Tajumulco, 3000-3800 m. alt., J. A.
Steyermark, 36952, Mar. 1, 19^0 (F); Barranco Eminencia road between San Marcos and San Rafael Pie de la. Cuesta, P. C. Stand ley, 8 6 2 7 8, Feb. 6, 19^1 (F); Barranco Eminencia between Finca
La Lucha and Buena Vista, 2500-2700 m. alt., P. C. Standley,
86225, Feb. 6, 1941 (F, US); Barranco Eminencia, road between San
Marcos and San Rafael Pie de la Cuesta, P. C. Standley, 86364 and 864l8, Feb. 6,1941 (F);
(Q) Polymnia meridensis Steyermark. Fieldiana, Bot. 28:667* 1953.
Polymnia cocuyensis Cuatrecasas. Rev. de la Acad. Colomb. 9:2^7. 1954. CHolotype FI).
Stems stout, rough, glandular-hispid, up to 3 meters tall; leaves sessile or the petiole winged to base, scabrous above, below densely puberulent, broadly ovate, lobes about 5 and ser rate-dentate, sinuses shallow, bases truncate to somewhat sagit tate; upper leaves without lobes, ovate-lanceolate; peduncles stout, densely hirsute to glandular-hispid; phyllaries about 5* broadly ovate, obtuse, about 12-14 mm long, 10-12 mm wide, densely glandular or the glands among hairs; ray flowers up to about 2 0, corolla yellow, 8-9-nerved, pilose below, the tube densely pilose, about 10 mm long, 4 mm wide; disc flowers 5 -8 mm long, ray pales lanceolate, about 7 mm long, pales of disc flowers 5 -7 mm long;
fruit black, obovate, 3 mm wide, 4.5-6 mm long (Fig. 18).
Type: VENEZUELA: Merida: Rocky thickets on slopes below
Chachopo 3045 m. alt., J. A. Steyermark, 57519* July 22, 1944
(Holotype FI). 107
Fig. l8.— Polymnia meridensis (Holotype) Scale"! .2&X
From the specimens examined I infer that P. meridensis is a polymorphic species. When one reads the type descriptions
of Steyermark's P. meridensis and Cuatrecasas's P. coccuyensis he finds that those two species overlap. The type specimens of
these two species were examined. After such examination along
with another specimen of P. coccuyensis annotated as such by
Cuatrecasas, I am unable to separate the two species and there
fore have placed P. coccuyensis in synonomy with P. meridensis.
Still another collection— Aristeguieta No. 2^8^— had leaves which
were more rugose but yet not sufficiently different to be worthy 108 of taxonomic segregation. It is probable that the few specimens examined do not represent the full range of variation which exists within this species.
steyermark's P. meridensis, collected in 1944, was not described until 1953* This description antedated that of P. coccuyensis by one year hence P. meridensis is retained on the basis of priority of publication.
P. meridensis resembles P. riparia but differs in the more robust and heavily-vestured stems and peduncles. The ray pales are noticeably unlike in the two. In P. riparia they are elongate and concave with large stipitate glands below which are up to 1 mm long. In P. meridensis the ray pales are oblong- lanceolate and not noticeably pointed and glands are absent or small•
Specimens Examined: COLOMBIA: Boyaca: Cocuy, 275^ m. alt., J. Cuatrecasas, 1700, Sept. 14, 1938 (F); Pauna, on hwy.
to Muzo, 2850 m. alt., K. Garcia-Barriga, 13237, Nov. 12, 1948
(US). Cundinamarca: Bogota Ciudad Universitaria Jardin Botanico,
2600 m. alt., K. Garcia-Barriga, 1084l, Mar. 19, 19^3 (US);
Guavio River valley, 4 km N of Ubala, 1725 m. alt., M. L. Grant and F. R. Rosberg, 9375, May 29, 1944 (F). VENEZUELA: Merida:
La Carbonera, 2500 m. alt., L. Aristeguieta, 2484, Sept., 1956
(F, NY).
(R) Polymnia sonchifolia Poepp. and Endl. Nov. Gen. ac Sp. PI. 3:47 -41H 1845.
Polymnia edulis Wedd. Ann. Sci. Nat. Bot. Series 4, 7:114. 1857. 109
Herbs to about 2 meters tall, stems stout, angular, furrowed, densely pilose above, often orange-yellow when dry; the tuberous underground portions are a source of human food; lower leaves broadly ovate and hastate or nearly so, as large as
33 cm long and 22 cm wide, the short lobes separated by broad shallow sinuses, lobes and sinuses dentate, the teeth callous- tipped, bases of expanded portion deltoid or abruptly cuneate, the lower part winged, wings up to k cm wide, 1 ^ cm long, auricu- late at base, main veins 3 » upper surface pilose, the lower pubescent, upper leaves ovate-lanceolate, without lobes and hastate bases; peduncles densely glandular-pilose; phyllaries 5> uniseriate, ovate-lanceolate, as large as 15 mm long, 10 mm wide, glandular-pilose below, nearly glabrous above, margins pilose; ray flowers about 15 or fewer, corollas yellow, 12 mm long or less, up to 7 mm broad, the tube densely pilose outside; disc
flowers about 7 mm long, corollas yellow, slightly pubescent on
outside; ray pales ovate-lanceolate, about 6 mm long, 2 mm wide; pales of disc flowers obovate, upper margin denticulate, about
3 mm long and 2 mm wide; fruit purple (mature ones not seen)
(Fig. 19).
Type: "Crescit in montibus calcareis herbidis Peruviae
orientalis ad Cassapi." (Type not seen).
Although P. edulis may be a commonly used scientific name
for this taxon it is antedated 12 years by P. sonchifolia. The
original description of P. sonchifolia is quite extensive and is
accompanied by detailed drawings of the various reproductive and 110
Fig. 19.— Polymnia sonchifolia Scale: .27X vegetative parts. There is, therefore, little doubt as to the taxon being described.
Poeppig and Endlicher describe the fruit as being J- angled, obovate, striate and purple.
Heiser has noted on the herbarium label of his collection
6068 a haploid chromosome number of JO for this species.
Polymnia sonchifolia along with P. connata have under ground tuberous parts which are eaten by natives in South America.
Although these two species superficially resemble each other the biseriate ray flowers and connate leaf bases of P. connata make it readily distinguishable from P. sonchifolia whose leaf bases Ill are separate and whose ray flowers are uniseriate. These two species are geographically separated. Polymnia sonchifolia grows in western South America while P. connata is found in southeastern South America.
"Llacon" is a common name for P. sonchifolia in Bolivia and in Ecuador the common name "jicama" is used.
Specimens Examined: BOLIVIA: Cochabamba: M. Bang,
1229, in 1829 (NY, US). La Paz: near Sorata, 2700 m. alt., G.
Mandon, 28, Apr., 1959 (F. NY); near Sorata toward Iminapi,
H. C. Cutler and M. Cardenas, 10003, Mar. 29, 1947 (F). Tari.ja:
K. Fiebrig, 2 8 5 2, Apr. 24, 1904 (US). COLOMBIA: Cauca: S. Y.
Agredo, 216, Mar., 1948 (F). ECUADOR: Chimborazo: Riobamba,
2800 m. alt., A. Rimbach, 727 (NY, F 863 6 8 8). Imbabura: T.
Guerro and C. B. Heiser, Jr., 5073A, Apr. 25, 1962 (IND).
Pichincha: Quito, I. Holmgren, 130, Dec. 29, 1919 (US).
Tungurahua: 2400 m. alt., C. B. Heiser, Jr., 6 0 6 8, May 25, 1962
(IND). PERU: Lima: Wawra, 2604, 1868-1871 (NY).
(S) Polymnia apus Blake. Contrib. U. S. Nat. Herb. 22:604. 1924.
Probably an erect perennial herb, the lower part not observed; stem sulcate with spreading pilose hairs, upward be coming stipitate glandular; leaves sessile or winged to base, triplinerved, the lower broadly ovate, the upper ovate, up to 35 cm long and 32 cm wide, lobes short and sinuses shallow and broad, the lobes gradually becoming obsolete near the apex, margins ir regularly dentate, the teeth callous-tipped, upper surface short- 112 pilose, the pilosity especially prominent along the main veins, lower surface densely pubescent; peduncles stipitate-glandular; phyllaries about 5» basally united, lanceolate, callous-tipped, up to 9 mm wide and 2 .5 cm long, below stipitate-glandular and pilose, margins pilose, above sparsely glandular to glabrous; ray flowers about 1 0 , corollas yellow, tridentate, about 2 cm. long, 5 mm wide, short-pilose on veins below; disc flowers about
6 mm long; pales of ray flowers ovate-lanceolate, acuminate, about 11 mm long, h mm wide; disc pales lanceolate-ovate, about
7 mm long and 2 mm wide; fruit black, not noticeably striate,
(only immature forms seen) (Figs. 20 and 21),
Type: MEXICO: Nayarit: Territorio de Tepic, between
Santa Gertrudis and Santa Teresa, J. N. Rose, 2079* Aug. 8 ,
1897 (Holotype USI).
The only specimen of P. apus examined other than the type is Rose 2077 which is in the herbarium of the New York
Botanical Garden. This specimen may be an isotype on which the collection number was mistakenly listed as 2077 instead of 2079*
All other information of the herbarium label of Rose 2077 is identical to that of the holotype.
The holotype was originally identified as P. edulis
(P. sonchifolia) which is not known to occur north of Colombia.
From P. edulis it differs in morphology of leaves, ray corollas, and phyllaries as well as vesture of peduncles.
Polymnia apus appears to be sufficiently different from other Polymnia species to warrant its acceptance as a separate 113
i. v Tt-*rU. -«/ ■' ...
....'.■■vvi.vjfj'."''' - : Is-- -
Fig. 20.— Polymnia apus (Holotype) Scale: .27X species. Admittedly, only two specimens do not present much of a range of variation whereas a larger sample might result in a modification of this viewpoint. --- f it*., i— * >■ 4_
Fig. 21.— Polymnia apus (?Isotype) Scale: .28X~
(T) Polymnia suffruticosa (Baker) Badillo. Boletin de la Sociedad Venezolana de Ciencias. 10:310. 19^6.
Melampodium suffruticosum Baker ex Martius FI. Bras. 6: part 3» 1 ^2 . IHB¥~(Holotype).
Polymnia stenocarpha Blake. Bull. Torr. Bot. Club. 5 8:b&7. 1 9 3 1 .
Shrubs or herbs to 2 meters in height, branching often dichotomous, stems terete or nearly so, densely pilose upward, less so below; leaves small, ovate-lanceolate, up to 10 cm long and 3 cm wide, their margins crenulate, densely pilose and glandular above and below, petioles up to 1 cm long on larger 115 leaves, the smaller leaves sessile or nearly so; peduncles about
1 cm long, densely pilose; phyllaries in 2 series, the lower 2
(infrequently 5) ovate-lanceolate, pilose, opposite, and at right angles with the upper pair which are broadly ovate, puberulent below and on tip above, shining-coriaceous, with a texture similar to that of ray pales; ray flowers about 7» corollas yellow, up to 3 nun wide and mm long, the tube 1 .8 mm long or less and pilose outside, persistent annular disc at base of corolla; disc flowers 15 -2 0, 5 mm long, lobes of the corolla bordered by a prominent vein; ray pales coriaceous, petaloid, yellowish— at least when dried— broadly ovate, 3 nun wide, 5 mm long, several prominent longitudinal parallel veins persistent almost to edge of pale; pales of disc flowers narrowly lanceolate, about 3 mm long, the outer disc pales somewhat wider than the inner ones, a prominent midvein persisting to the tip; fruit brown, oblong- quadrangular, about 3 mm long and 1 .5 mm wide, annular disc about
.3 mm high persistent on upper end of fruit (Fig. 22).
Type: VENEZUELA: Amazonas: on rocky, dry ground near
Esmeralda, on ? tributary of the Orinoco River, Spruce, 3225
(not seen).
Polymnia suffruticosa is easily separable from all other species of the genus. This species might be worthy of subgeneric status on the basis of its very distinctive morphology; however, no such category has yet been erected for this genus.
I know of no record for the occurrence of P. suffruticosa
outside the Amazonas province of south-central Venezuela. It 116
Fig. 22.— Polymnia suffruticosa Scale; .2^X would not be particularly surprising to learn of its presence in neighboring regions of Brazil and Colombia.
This species probably occurs at the lowest altitudes
of all South American species of Polymnia. All collections
examined were made between 130 and 500 meters altitude.
This species along with P. canadensis and P. laevigata
has more than a single series of phyllaries associated with
each head.
Specimens Examined: VENEZUELA: Amazonas: Esmeralda,
Upper Orinoco, 130 m. alt., L. Williams, 15 382, May 15, 19^2 117
(F, US); Upper Orinoco, 1JO m. alt., L. Williams, 16150, June,
19^2 (F, US); between Esmeralda and base of Cerro Duida, 150 m. alt., J. A. Steyermark, 57736, Aug. 21, 1 9 ^ (F, US); Cerro
Moriche, Rio Ventuari, 200-500 m. alt., B. Maguire et ail., 3 0 8 3 8,
Jan. 13, 1951 (NY); Rio Ventuari, 200 m. alt., B. Maguire et al.,
3100^, Jan. 17, 1951 (NY); between mouth of Rio Paru and Las
Carmelitas, 150 m. alt., B. maguire et al., 31575» Feb. 19» 1951
(NY); Esmeralda Ridge, Alto Rio Orinoco, 150-200 m. alt., B.
Maguire and J. J. Wurdack, 3^666 (F, NY, US).
(U) Polymnia pyramidalis Triana. Ann. Sci. Natur. Bot. Ser. k . 9: 4o-4l* 155^
Polymnia eurylepis Blake.. Bet. Gaz. 7^5^15* 1922. (Holotype USJ, Paratype US!).
Trees exceeding 12 meters in height, stems up to 20 cm in diameter, bark at least sometimes very dark, becoming rough on older stems; leaves ovate to broadly ovate, sessile or petiole winged to base or distinctly petiolate, up to 25 cm wide and *f2 cm long, lower surface tomentose especially along veins, upper surface glabrous or sparingly strigose, dentate or denticulate; peduncles villous to hispid; phyllaries about 6, 1 cm long or less, up to 7 nun wide, ovate, uniseriate, glabrous or with few pilose hairs, margins pilose; ray flowers about 20, corollas yellow, 1 cm long or less, about k- mm wide; disc flowers about 3 mm long, corollas yellow, puberulent outside; ray pales concave, obovate, 5 mm long or less; pales of disc flowers oblanceolate,
about 3 mm long, margin denticulate, midvein prominent; fruit black, flattened-obovoid, up to 4 mm long and 3 mm wide (Fig. 23). 118
Type: COLOMBIA: "Prov." de Bogota, 2700 m. alt.,
Triana, 1323 (type not seen).
Plant size of this species is probably greater than that of any other species of Polymnia. In his original description
Triana (1 8 5 8) pointed out that the plants are trees up to *f0
feet tall.
The epithet "pyramidalis" is derived from the pyramidal shape of the crown. Triana (1 8 5 8) noted that natives in
Colombia use them around their houses as a shelter from the wind.
He further points out that the plants with their large size, pyramidal crown, and almost constant garnish of yellow flowers
serve as ornament.
In 1923 Blake annotated certain specimens from the United
States National Herbarium as P. eurylepis but later (in 1932)
annotated those same specimens as P. pyramidalis.
Common names for plants of the species include: "polaco"
in Ecuador, "anime" in Colombia and Venezuela, "anime manso" in
Venezuela, and "arbolaco" in Colombia. These three countries are
the only ones from which I have seen specimens.
Heiser's collection of P. pyramidalis ^917 was reported
by him to have a haploid chromosome number of "about 3 0."
Figure 23 is a copy of a photograph of the holotype. 119
TVFES OF THE FANIS HCNSANIUM
Fig. 23.— Polymnia pyramidalis (Holotype) Scale: .32X
Specimens Examined: COLOMBIA: Antioquia: Copacabana,
Daniel, 280, Mar. 28, 193^ (US); near Santa Elena between Medel lin and Rio Negro, 2300-2500 m. alt., G. Gutierrez, V. and A. L.
Delisle, 1119* Oct. 20, 19^6 (US). Boyacli: 6 km E of Socha in valley of Rio Arobispo, 3050-3070 m. alt., F. R.Fosberg, 22308,
Nov. 11, 1 9 ^ (F, US). Caldasi Old Quindio trail, 2500-2800 m. alt., E. P. Killip and T. E. Hazen, 9099, Aug. 1, 1922 (NY, US).
Cauca: open land in Cordillera Central, 2300-2500 m. alt., E. P.
Killip, 6859, June 17-18, 1922 (NY, US). Cundinamarca: Rio San
Francisco, above BogotH., 2700-2800 m. alt., F. W. Pennell, 2275, Sept. 27-29, 1917 (NY); Bogota, on road to Choachi, S. Juzepczuk,
5 1 8 9, June 7, 1926 (US); Sabana de Bogota, 2600 m. alt., J.
Cuatrecasas, 135^8, Jan. 8, 19^2 (F, US); between Cachipay and
La Florida, 1700 m. alt., J. Cuatrecasas, 136l*t, Mar. 22, 19^-2
(F, US); SE of BogotS. on trail toward UbanquS, 8900 ft. alt.,
E. L. Little, Jr. and R. R. Little, 9 2 3 8, Jan. 7, 19^5 (F, US);
NE of Bogota, 9000 ft. alt., E. L. Little, Jr. and R. R. Little,
9792, Mar. 19, 19^5 (F, NY, US); 2700 m. alt., 0. Haught, 5773,
May 28, 19^7 (US). Narino: between El Encano and Pasto, 2700-
2900 m. alt., J. Cuatrecasas, 1196^, Jan. 11, 19^1 (F, US). Norte de Santander: between Mutiscua and Pamplona, 2700-3^00 m. alt.,
E. P. Killip and A. C. Smith, 19693, Feb. 23, 1927 (NY, US);
Culga Valley near Tapata, 1500-2100 m. silt., E. P. Killip and
A. C. Smith, 20W+, Mar. 3-8, 1927 (NY, US); Pamplona, J.
Cuatrecasas and H. Garcia-Barriga, 1 0 2 k k t July 26, 19^0 (US).
Santander: near La Baja, 2700-3500 m.'alt., E. P. Kippip and
A. C. Smith, 180^3, Jan. 1^-31, 1927 (NY, US). Valle del Cauca:
Valley of Rio Bugalagrande, 2300 m. alt., J. Cuatrecasas, 20^55,
Mar. 27, 19^6 (F); Valley of Rio Bugalagrande, 2900 m. alt., J.
Cuatrecasas, 20886, Apr. 17, 19^6 (F). ECUADOR: Azuay: Cuenca,
E. W. D. and M. M. Holway, 97^, Sept. 10, 1920 (US). Carchi: between Paja Blanca and El Cucho, 2900-3200 m. alt., M. A. Solis,
10^75, July 17, 19^5 (F). Imbabura: near Ibarra, 2350 m. alt*,
Y. Mexia, 7^15, July 3, 1935 (US). Tungurahua: Ficoa, near
Ambato, A. Pachano, 135, Feb., 1919 (US); Rio Pastaza between
Banos and Patate, 1900 m. alt., C. B. Heiser, Jr., ^891, Feb. 10, 121
1962 (IND); Pelaquin, near Ambato, 3320 m. alt., C. B. Heiser,
Jr., 4917, Feb. 22,1962 (IND). VENEZUELA: Merida: Santo
Domingo, 2200 m. alt., A. Jahn, 128, Dec. 2, 1910 (US); San
Rafael, 3150 m. alt., A. Jahn, 7 8 7, Jan. 22, 1922 (US); San
Rafael de Mucuchies, H. Pittier, 12916, Feb. 6, 1928 (NY, US);
Timotes, 2000 m. alt., H. Pittier, 12942, Feb. 9» 1928 (NY, US);
San Rafael, 3185 m. alt., W. Gehriger, 10, Apr. 4, 1930 (F, NY,
US); along stream bed W of Timotes, 2315 alt., J. A. Steyer- mark, 55684, Mar. 22, 1944 (F, US); Sto. Domingo, 2600 m. alt.,
L. Aristeguieta, 2421, Sept., 1956 (F, NY, US). Tachira: above
Betania at base of Paramo de TamH., 2285 m. alt., J. A. Steyer- mark, 55684, July 14, 1944 (F).
Polymnia glabrata Complex
The largest single taxonomic problem in the genus Polymnia is the disposition of certain taxa which I have chosen to term the "Polymnia glabrata complex."
Members of this complex are all trees or shrubs which are found in western South America— specifically in south western Venezuela, Ecuador, Peru, and Colombia. The extremes of this complex are distinct but many of the specimens intermediate between these extremes are not distinct from other specimens.
Although only about 60 herbarium specimens were examined from species in this complex, the sample is large enough to bring to attention the size of the problem. I do not foresee any solution to this problem until thorough studies of these South
American plants are made in the field. The species considered to constitute this complex are:
P. glabrata, P. fruticosa, P. parviceps, and P. microcephala.
(V) Polymnia glabrata DC. Prodr. 5:515* 1 8 3 6.
Shrubs or small trees up to 8 meters tall; stems puberulent or glabrous; leaves ovate to rhombic-ovate, long- acuminate, sessile or petiole winged to base, base of expanded portion oblique or attenuate, margin crenate or sinuate to entire, glabrous above, below puberulent or glabrous; peduncles puberulent phyllaries ovate to round-ovate, puberulent or minutely glandular, up to 7 mm long and 5 mm wide; ray flowers about 1 0 , their corol las yellow, 6 mm long, 2-3 mm wide; disc flowers about 4 mm long,
corolla yellow; pales of ray flowers broadly ovate, 2 mm wide,
3 mm long; disc pales about 2 mm long, upper margin denticulate, midvein prominent, fruit black, obovoid, about 3 mm long and 2 mm
wide (Fig. 24).
Type: CHILE: Haenke, 8192 (Holotype not seen).
At the same time that De Candolle (1 8 3 6) described P.
glabrata he also recognized a (3 angustifolia. About the ^
angustifolia he said "foliis oblongo-lanceolatis subintegerrimis."
All of the specimens I observed had leaves the same shape as
those of the holotype photographed (Fig. 24). The type specimen
is in the Geneva Herbarium. Since this photograph is not that of
@ angustifolia there must be growth forms with leaves narrower
than any I have observed. Therefore, the status of the taxon
angustifolia should not be altered until there is sufficient
evidence for making a change. In Peru P. glabrata is called "poque" and is used in treating the common cold*
Fig. 2 k t— Polymnia glabrata (Holotype) Scale: .35X
Specimens Examined: PERU: Huancavelica: near Cordova,
R. D. Metcalf, 30279, Mar. 27-28, 19^2 (US). La Libertad:
Santiago de Chuco, 3150 m. alt., A. Lopez, 982, June 8, 1953
(US). Lima: Matucana, C. H. Townsend, 150^, Jan. 30, 1913 (HS)
San Buenaventura, 2800-2900 m. alt., F. W. Pennell,.145^2, June
17, 1925 (NY, US); Rio Blanco, 3000-3500 m. sit., E. P. Killip
and A. C. Smith, 216^7, Apr. 15-17, 1929 (NY, US). 124
(W) Polymnia parviceps Blake. Contrib. U. S. Nat. Herb. 22: 6o4. 1924.
Trees or shrubs to 5 meters in height; stems puberulent above (the lower part not seen); leaves sessile or petiole winged to base, the basal portion of the expanded blade cuneate to at tenuate, acuminate, 35 cm long or less, up to 18 cm wide, dentate, the teeth blunt and rounded, lower surface puberulent to tomentose especially on main veins; peduncles puberulent; phyllaries obovate to ovate, glabrous, about 3 mm long and 2 mm wide; ray flowers 10 -1 5 , corolla yellow, about 3 mm wide, up to 6 mm long, disc flowers about 2 5, the length 4 mm or less, the corolla glabrous; ray pales obovate, glabrous, about 4 mm long and 2 mm wide; pales of disc flowers about 3 mm long, lanceolate, chartaceous, their upper margins denticulate, midvein prominent; fruits black, about 3 mm long and 2 mm wide, obovoid and flat tened or obpyramidal (Fig. 25).
Type: PERU: Cuzco: Torontoy, 2400 m. alt., 0. F. Cook and G. B. Gilbert, 8l8 , May 20, 1915 (Holotype US!).
Polymnia parviceps appears to be intermediate between
P. glabrata and P. microcephala. From the former it differs in having slightly smaller phyllaries. From the latter it differs in having a noticeably dentate leaf margin. Leaf margins of P. glabrata are denticulate or entire. The leaves of P. parviceps axe on the average broader than those of P. glabrata. However, intermediate forms can be found which have the effect of obscuring the definite limits of the two taxa. 125
. ' . f ..- .. V '--.
Fig. 25.— Polymnia parviceps (Holotype) Scale: .33X
In Peru P. parviceps is called "yarita."
Specimens Examined: BOLIVIA: La Paz: Unduavi, 8000 ft. alt., Ex Herb. Parke Davis and Co., sn., Oct., 1885 (F);
Larecaja, near Sorata, 2600-2700 ft. alt., G. Mandon, 50, Apr.,
1859 (F, NY); Sorata, 8000 ft. alt., H. H. Rusby, 1669, Feb.,
1886 (NY, US); Sorata, E.W.D. and M. M. Holway, 558, Apr. 21,
1926 (US). PERU: Ayacucho: Prov. De La Mar, A. Weberbauer,
5557* May, 1910 (US); Pampalaca, between Huanta and Rio Apurimac,
3200 m. alt., E. P. Killip and A. C. Smith, 22268, May *f, 5» 18,
1929 (NY, US). Cuzco: Prov. Urubamba, 2000 m. alt., C. Vargas, 126
2802 (NY); Cuzco, 3350 m. alt., K. Krause, 596, Mar., 1925 (NY);
F. L. Herrera, 1480, Mar., 192? (US).
(X) Polymnia microcephala Hieron. Engl. Bot. Jahrb. 29:33* 1900. Polymnia lehmannii Hieron. Engl. Bot. Jahrb. 28:599* 1901. (isotype F!, USl).
Shrubs or small trees to 8 meters tall; branches glabrous
to pubescent; leaves sessile or petiole winged to base, rhombic-
ovate, acuminate, the broadened portion of blade at base at
tenuate, entire, below sparingly puberulent to tomentose, resin-
dotted, above glabrous except along the puberulent veins;
peduncles slender, about 3 cm long or less; phyllaries k - 5,
ovate-lanceolate, 2 -5 mm long; ray flowers about 1 0 , corollas
yellow, about 7 mm long or less, up to 3 mm wide; disc flowers
about 2 5, 2 -3 mm long; pales of ray flowers ovate, up to 3 mm
long, 2 mm wide, glabrous; pales of disc flowers about 2 mm
long; fruit black, smooth, obovate, laterally flattened, about
2.5 mm long or less (Fig. 26).
Type: ECUADOR: Chimborazo: In valle Pollatanga, 2000
m. alt., A. Sodiro, 2^/2 (Holotype fragments US!).
The identity of P. lehmannii with P. microcephala is
noted here. The type specimen of each of the two species was
taken at about 2000 m. alt. in Ecuador. After examining types
of P. lehmannii and fragments of the type of P. microcephala
along with a photograph of the type of P. microcephala I conclude
that no significant difference exists between them. They in fact Fig. 26.— Polymnia microcephala (Holotype) Scale: .29X are quite similar in many respects to P. parviceps but differ in having an entire leaf margin.
Figure 26 is a copy of a photograph of the holotype in
the Berlin Herbarium.
Specimens Examined: ECUADOR: Cuenca: 2000-2600 m. alt.
F. C. Lehmann, *f895 (F, US).
Polymnia fruticosa Benth. PI. Hartw. 209* 18^5.
Polymnia arborea Hieron. Engl. Bot. Jahrb. 29:33* 1900. 128
Aromatic trees or shrubs up to 12 meters in height; leaves lanceolate to ovate, acuminate, glutinous, coriaceous, up to 25 cm long and 11 cm wide, sessile or petiole winged to base, base of widest portion of blade attenuate, the bases some
times auriculate but not connate, entire or denticulate; margins, at least when dried, revolute; densely tomentose below, above shining glabrous; peduncles and lower surface of phyllaries
densely stipitate-glandular, the glands sometimes accompanied by hairs; phyllaries 4-5 in 1 series, ovate-lanceolate, about 1 cm long, 7 mm wide or less; ray flowers about 1 2 , corollas yellow,
9 mm long or less, up to 4 mm wide; disc flowers about 4 mm
long, corollas yellow; ray pales ovate, about 5 mm long, 2 mm
wide, minutely glandular below; disc pales denticulate along
upper margin, midvein prominent; fruit obovoid black, 4 mm long
or less, about 2 mm broad (Fig. 27)•
Type: "In montibus Quimia, prope Rumibamba" (? Colombia)
Hartweg, 1158 (Isotype NY!, fragments of Isotype F!, US!).
Riobamba, a city in the province of Chimborazo is thought to be
the intended name for "rumibamba."
The closest relative of P. fruticosa is thought to be
P. glabrata. The principal distinguishing characteristic between
these two is the densely glandular condition of peduncles and
phyllaries in P. fruticosa. Leaf shape is often similar in the
two species but the leaves in P. glabrata are less coriaceous
than those of P. fruticosa. Also, in P. glabrata the lower 129
flAKIS Of ECUADOR
Fig. 27.-—Polymnia fruticosa Scale: ,28X surface of the leaves is glabrous or nearly so whereas in P. fruticosa it is densely tomentose.
"Colla" and "polaco" sire common names for P. fruticosa in Ecuador. The haploid chromosome number of this species is
"over 25" according to Heiser (Table 2).
Specimens Examined: COLOMBIA: Narino: 5 km from
Tuquerres, headwaters of Rio Pasqual, 3000 m. alt., J. A. Ewan,
16893, Feb. 7, 19^5 (F, US). ECUADOR: Azuay: along Rio
Matadero, 9600-10,000 ft. alt., W of Cuenca, W. H. Camp, E-I98O, 130
Mar. 3> 19^5 (NY); along Rio Tarqui, 18 km S of Cuenca, 8300-
9000 ft. alt. (NY, US). Bolivar: Western Cordillera valley of
Chimbo River near Guaranda, 2800 m. alt., A. Rimbach, 629 (F
839586); Casubagua, 2860 m. alt., M. Acosta-Solis, 5995» Sept.
29, 19^3 (F); between El Sato and Illubin, 2200 m. alt., M.
Acosta-Solis, 6557» Oct. 25, 19^3 (F); Pucara de Telunbela,
2600-3000 m. alt., M. Acosta-Solis, 68l4, Nov. 18, 19^3 (F).
Imbabura: Cotocachi, I. Holmgren, 921, Aug. 30, 1920 (US).
Pichincha: near Quito, Jameson, 10^9 (NY); Cotocollao, L. Mille,
^90, in 1897, (NY); Cotocallao, A. M i l e , sn., Nov. 12, 1920
(US); Borde del camino de La Magdalena al Cinto, C. Firmin, 676,
Oct. 25, 1928 (US); E. Ilalo, E. K. Balls, 579^, Dec. 12, 1938
(US); Ciudadela Universetario, Quito, 28l8 m. C. B. Heiser, Jr.,
^851, Feb. 3i 1962 (IND); near Quito, 3000 m. alt., C. B. Heiser,
Jr., 501^, Mar. 31> 1962 (IND). Tungurahua: W of Cotalo on road
to Banos, 2600 m. alt., C. B. Heiser, Jr., 6016, May 11, 1962
(IND). PERU: Lima: Chosica, Macbride and Featherstone, 532,
May 2, 1922 (US). EXPERIMENTAL TAXONOMIC INVESTIGATIONS
Three species of Polymnia are found in the United States.
These are P. canadensis, P. laevigata, and P. uvedalia. The last species is composed of three morphologically distinct varieties; namely, P. uvedalia var. uvedalia, P. uvedalia var. densipilis, and P. uvedalia var. floridana. These taxa in the
United States have been the subject of experimental investi gations for the past three years.
Experimental plant taxonomy employs a wide variety of approaches. Among those used during this study were pollen stainability and morphology, chromosome number and morphology, and hybridization studies. Several more years will be required before such a study can be completed. Nevertheless, some useful information has been gained as a result of these efforts; con sequently, the phylogenetic relationship of these United States
taxa is now somewhat more than pure conjecture.
Several trips were made to the southern and southeastern
United States in order to obtain living specimens of all the
above-mentioned taxa. The transplants were kept at conditions
prevailing in the Ohio State University greenhouses except for
certain plants which were permitted to overwinter outside the
greenhouse.
131 132
All chromosome material was fixed in a solution of three parts of 95 per cent ethanol and one part acetic acid and main tained at about 3°C for 2 k hours. The acetic acid-alcohol mixture was then replaced by 70 per cent ethanol and the material returned to a temperature of 3 °0 until examinations were made.
The preserved young anthers and root tips were stained with aceto-carmine. Examinations and photographs of the chromo somes were made after the stained material had been smeared between a slide and cover slip.
Pollen stainability was used as an indicator of pollen viability. The stain used for pollen grains was aniline blue which was prepared by combining one gram of aniline blue crystals,
25 grams of phenol crystals, and 25 milliliters each of glycerin, lactic acid, and distilled water.
The pollination technique used was suggested by R. W.
Cruden. For each pollination I removed the monosporangiate, staminate disc flower with a pair of forceps and used this
flower itself, on which mature pollen was located apically, as a brush. Pollination was considered effected when pollen could be seen with the unaided eye on the stigma of the ovule parent.
Using the technique as outlined by Comar and Zacheile
(19^2) I wished to see if two morphologically distinct species
such as P. uvedalia and P. canadensis could be separated on the
basis of content of chlorophylls a and b. The chlorophylls were
extracted from 10 gram wet weight samples of leaf tissue from
each species. 133
The chlorophyll a content in both species was about 5 mg/liter of solution made according to the method referred to above. The chlorophyll b content was slightly in excess of k mg/liter of solution. Dr. Eugene Carr informs me, via personal communication, that in obtaining the 1 0 -gram sample of leaf tissue from the plant there is as much as 10 per cent sampling error. Results indicated less than 10 per cent difference between these two species for amounts of chlorophylls a or b.
Prior to hybridization experiments, counts of chromosome numbers were made of P. canadensis vars. canadensis and radiata,
P. laevigata, and P. uvedalia vars. uvedalia, floridana, and densipilis. These are the only taxa of Polymnia which are known to grow naturally in the United States. The counts which I made and those made by others are listed in Table 2. Photomicrographs of some of the smears used in making these counts follow. Voucher specimens are deposited in the herbarium of The Ohio State Uni versity.
Figure 28 is a photomicrograph of a microsporocyte smear of P. laevigata at diakinesis. Fifteen pairs of chromosomes are clearly visible. The darkest object in the figure labelled S is a stain crystal. The chromosome pair labelled H is thought to be heteromorphic with a satellite extending out from but still con nected to the homologue farthest to the right.
Figure 29 is a photomicrograph of a pollen mother cell from the same P. laevigata plant at metaphase. The satellite is not now apparent. Its absence is probably due to greater TABLE 2 LIST OF CHROMOSOME NUMBERS OF CERTAIN TAXA OF POLYMNIAa
Haploid Collection Taxon Number Ascertained by Number Year
P. canadensis var. canadensis 15 J. R. Wells 218 I960 p. canadensis var. canadensis 15 L. C. Anderson 2180 1962 p. canadensis var. radiata 15 J. R. Wells 249 1961 p. fruticosa over 25 C. B. Heiser, Jr. 4851 1962 p. laevigata 15 J. R. Wells 255 1961 p. maculata var. adenotricha 33 + 1 R. M. King 5046 1962 fragment p. maculata var. adenotricha 16 M. Powell King 4427 1961 p. maculata var. maculata 16 B. L. Turner, W. L. Ellison, R. M. King 2665 I960 p. maculata var. maculata 17 B. L. Turner, W. L. Ellison, R. M. King 3306 I960 p. oaxacana about 20 R. M. King 5288 I960 p. oaxacana 16 B. L. Turner, W. L. Ellison, R. M. King 3338 I960 p. pyramidalis about 50 C. B. Heiser, Jr. 4917 1962 p. riparia about 15 C. B. Heiser, Jr. 5050 1962 p. sonchifolia 30 C. B. Heiser, Jr. 6068 1962 p. uvedalia var. densipilis 16 J. R. Wells 247 1961 p. uvedalia var. floridana 16 J. R. Wells 248 1961 p. uvedalia var. uvedalia 16 J. R. Weils 211 1961
Any chromosome counts attributed to J. R. Wells were made during this investigation. Records of all other counts were obtained from herbarium labels except King No. 5046 which was via personal communication. H -P- Fig. 28.— Polymnia laevigata pollen mother cell at diakinesis (Wells 255). Fig. 29 • — Polymnia laevigata pollen mother cell at early metaphase (Wells 255)• 137 contraction of the chromosomes. It was observed that the chromosomes at metaphase are only about half as large as those at diakinesis.
Figures 30 and 31 show the approximate size of the 16
chromosome pairs of P. uvedalia at diakinesis. Wo differences
were observed in the chromosome number or morphology among the
three varieties of P. uvedalia.
The 13 chromosome pairs of P. canadensis may be counted
in Figure 32, which is a photomicrograph of a pollen mother cell
at diakinesis.
The percentage of pollen stainability was high in all
taxa (Table 3)* Since this stainability test was used as an
indicator of pollen viability it is inferred that no selective
mechanism such as gametophytic lethality is operating in the
representatives of the taxa which were examined during this study.
Ranges of diameters of pollen grains can be useful in
separating P. uvedalia, P. canadensis, and P. laevigata taxonomi-
cally. These ranges of diameters are P. laevigata P.
canadensis 33/i“^0/* and *f2/«-52j*. for P. uvedalia. All pollen, with
the exception of heteromorphic pollen (Fisher and Wells, 1962),
was spherical with many spines which projected outward in all
directions. All ranges are based on a random sample of 10 pollen
grains. Measurements were taken under oil immersion lens. Each
of the measurements is from the tip of one spine to the tip of
another spine which projected outward in an opposite direction
but on the same plane. The average diameter of pollen grains 138
Fig. 30.— Polymnia uvedalia var. uvedalia pollen mother cell at diakinesis (Wells zuT. 139
Fig. 31.— Polymnia uvedalia var. uvedalia pollen mother cell at diakinesis (Wells 2TTT. 140
Fig. 32.— Polymnia canadensis pollen mother cell at diakinesis (Wells 218)'. TABLE 3 POLLEN STAINABILITY OF TAXA WHICH OCCUR NATURALLY IN THE UNITED STATES. ALL COUNTS ARE BASED ON A RANDOM SAMPLE OF 500 POLLEN GRAINS. COLLECTION 255 WAS COUNTED FROM HERBARIUM MATERIAL. ALL OTHER COUNTS WERE MADE FROM LIVING PLANTS
Collection No. Stained No. Stained Per Cent Taxon Number Positively Negatively Stainable
P. canadensis var. canadensis 279 481 19 96.2 p. canadensis var. canadensis 282 491 9 9 8 .2 p. canadensis var. radiata 265 487 13 97.4 p. laevigata 25*fa 464 36 9 2 .8 p. laevigata 255 499 1 99.8 p. uvedalia var. densipilis 246 498 2 99.6 p. uvedalia var. densipilis 242 495 5 99.0 p. uvedalia var. floridana 244 492 8 98.4 p. uvedalia var. floridana 243 491 9 9 8 .2 p. uvedalia var. genuina 211 495 5 99.0 p. uvedalia var. genuina 226 497 3 99.^ p. uvedalia var. genuina 256 484 16 9 6 .8 p. uvedalia var. genuina 239 498 2 99.6 p. uvedalia var. genuina 231 496 4 99.2 p. canadensis var. radiata 265 X 279 495 5 99.0 (pollen parent) X P. canadensis var. canadensis (ovule parent)
aCollection 254 is the plant from which heteromorphic pollen was described (Fisher and Wells 1962), ikz mounted in distilled water is not significantly different from
that of those mounted in acetocarmine.
Species which are cross-fertile obviously have many
genes in common. Therefore, fertility of hybrids between
species may be used as an indicator of phylogenetic relation
ships.
In Table k is presented a summary of attempts at
achieving hybrids between different species of Polymnia. In all
hybridization experiments the number of crosses means the number
of actual flowers (not heads) cross pollinated. A total, of two
seedless fruits were formed from 151 attempts at crossing P.
uvedalia and P. canadensis. These results are predictable on
the basis of differences in chromosome number of the two species,
their morphology, and their different ecological habitats. I
consider these two species to be cross-sterile.
When P. laevigata was crossed with P. canadensis 105
fruits were formed from 205 crosses attempted, i.e., fruit set
was about 50 per cent.
If all 38 matings are considered, the per cent of fruit
set was usually low. Two exceptions were matings 10 and 12 which
had 100 per cent and 92 per cent fruit-set respectively. Mating
10 involved a sample of only eight attempts at hybridization
while in mating 12 the heads had been emasculated. I attribute
a partial reduction in yield of fruits to a loss of some of the
ovulate flowers as a result of inadvertent breaking of conducting
elements between ovulary and receptable during emasculation. TABLE 4 SUMMARY OF INTERSPECIFIC HYBRIDIZATION ATTEMPTS
Number of Number of Per Cent Mating Crosses Fruits Fruits Number Pollen Parent Ovule Parent Attempted Formed Formed
1 P. canadensis var. P, laevigata 35 21 60,0 canadensis
2 P. laevigata P. canadensis var, 42 22 52.4 canadensis
3 P. canadensis var, P, laevigata 85 47 55.3 radiata
k P, laevigata P, canadensis var, 43 15 34.9 radiata
5 P. canadensis var, P, uvedalia var, 82 0 0.0 canadensis uvedalia
6 P, uvedalia var, P, canadensis var. CO 1 2.6 uvedalia canadensis (seedless)
7 P. canadensis var, P, uvedalia var, 31 1 3.2 canadensis floridana (seedless) Evidence for the theory that through emasculation a reduction in yield is effected is thought to be found in the results of matings 29 and 30. A higher per cent of fruit-set was achieved in mating 30 than in mating 29. There was no emasculation in mating 30 yet var. uvedalia is nearly self- sterile according to the results of mating 11. Therefore, the higher per cent yield of fruits in mating 30 is probably due not to self-fertilization but to the maintenance of continuity in vascular elements between ovularies of ray flowers and the receptacle.
In Tables *t-8 I have entitled one column "Number of
Fruits Formed." Parthenocarpic fruits sometimes occur in
Polymnia uvedalia; therefore, there is not a 100 per cent cor relation between fruit formation and seed-set. Random samples of fruits from all matings revealed seeds. Consequently, a high correlation is indicated between fruit formation and seed-set.
Obviously, not all the fruits were examined for seed content since it was desirable to have the hybrid seeds germinate and be useful in further tests.
Although no crosses were made between P. laevigata and
P. uvedalia they are assumed to be cross-sterile. Reasons for this assumption are l) there is a basic difference in chromosome number— the haploid being 16 for P. uvedalia and 15 for P. laevigata— and, 2) P. canadensis is cross-sterile with P. uvedalia although P. canadensis is cross-fertile with P. laevigata. In 145 other words, P. canadensis is more closely related genetically to P. laevigata than to P. uvedalia.
In addition to knowing something about which crosses could be made at the species level I wished to check for per cent cross-fertility and self-fertility within the same varieties of P. canadensis, P. uvedalia, and in P. laevigata. The results of such tests and of tests for ovulary abortion appear in Table 5»
The results from "matings" 19-22 indicate that ovularies do not develop into fruits parthenogenetically. let, some fruits are formed from selfings in all taxa except P. uvedalia var. densipilis (mating 16) of which only 15 were attempted. In this case, I believe a larger sample would have resulted in formation of a few fruits. Based on the results obtained in mating 11, the expectation in mating 16 would be less than one fruit formed.
Other conclusions drawn from Table 5 are l) the two varieties of P. canadensis are only about 10 per cent self-
fertile, 2) the three varieties of P. uvedalia are also low in self-fertility; i.e., less than 20 per cent, 3) uvedalia var.
floridana appears to be more self-fertile than the var. uvedalia,
and 4) self-fertility in P. laevigata is unexpectedly high— in
excess of 50 per cent.
Self-sterility is thought to be rather common in species
of Compositae. If all of the few known populations of P.
laevigata are highly self-fertile this might help account for the
limited distribution of the species; iew, in only seven counties
of the United States. Species which are self-fertile tend to be TABLE 5 SUMMARY OF INTRAVARIETAL HYBRIDIZATION ATTEMPTS, SELFINGS, AND TESTS FOR OVULARY ABORTION
Number of Number of Per Cent Mating Crosses Fruits Fruits Number Pollen Parent Ovule Parent Attempted Formed Formed
8 P. canadensis var. P. canadensis var. canadensis canadensis (selfed) 148 14 9.5 9 P. canadensis var. P. canadensis var. 45 5 11.1 radiata radiata (selfed) 10 P. canadensis var. P. canadensis var. 8 8 100.0 canadensis canadensis (not selfed) 11 P. uvedalia var. P. uvedalia var. 139 9 6 .5 uvedalia uvedalia (selfed) 12 P. uvedalia var. P. uvedalia var. 25 23 9 2 .0 uvedalia uvedalia (not selfed) 13 P. uvedalia var. P. uvedalia var. 18 10 55.6 uvedalia uvedalia (not selfed) 14 P. uvedalia var. P. uvedalia var. 36 7 19.4 floridana floridana (selfed) 15 P. uvedalia var. P. uvedalia var. 68 28 41.2 floridana floridana (not selfed) 16 P. uvedalia var. P. uvedalia var. 15 0 0.0 densipilis densipilis (selfed) 17 P. uvedalia var. P. uvedalia var. 55 25 45.5 densipilis densipilis (not selfed) TABLE 5— (Continued)
Number of Number of Per Cent Mating Crosses Fruits Fruits Number Pollen Parent Ovule Parent Attempted Formed Formed
l8 P. laevigata P. laevigata (selfed) b9 21 55.3
19b P. canadensis var. 5 k 0 0.0 canadensis 0 0 20b P. uvedalia var. 8*t 0 . uvedalia
21b P. uvedalia var. 51 0 0.0 floridana
22b P. uvedalia var. 30 0 0.0 densipilis
emasculation was carried out in these crosses* The flowers of heads involved in these tests were emasculated; no pollination of any kind followed. ik $ ~ more homozygous than those which are self-sterile. Consequently,
fewer new genetic recombinations would result where selfing
occurs. This would mean a greater chance of survival of plants
that were genotypically similar to the parents, which themselves
are already presumed to have limited ecological amplitude.
The non-selfed matings 10, 12, 13, 15, and 17 each in
volved parents from widely separated populations such as from
different states or different counties within the same state.
The higher percentage of fruit-set from these matings than from
the selfings serves as a check on the efficacy of the pollination
technique. In other words, the lower per cent yield of fruits
from selfings than from matings 10, 12, 13, 15i and 17 is prob
ably not the result of pollination techniques.
The data in Table 6 are offered as further evidence that
P. uvedalia is self-sterile or nearly so. However, this was
not the main objective of matings 23-26.
My collections 236 and 239 came from an almost solid
stand of P. uvedalia var. uvedalia. This population is located
in Calhoun County, West Virginia, and measures about 150 feet long
and 20 feet wide. Plant no. 236 came from one end of this popu
lation and no. 239 came from the other end, i.e., they had been
growing about 150 feet apart before being transplanted to The
Ohio State University greenhouses.
Including reciprocal crosses, 112 cross pollinations were
attempted between these two plants. No fruits were formed hence
self-sterility is indicated, at least within this population. ]A9
TABLE 6 CROSSES INVOLVING TWO PLANTS OF P. UVEDALIA VAR. UVEDALIA (COLLECTIONS 2?6 AND 239) FROM DIFFERENT ENDS OF A LARGE POPULATION
Number of Number of Mating Crosses Fruits Number Pollen Parent Ovule Parent3, Attempted Formed
23 236 239 31 0 2b 239 236 25 0 25 236 236 (selfed) 26 0 26 239 239 (selfed) 12 0
All pollen-producing flowers in the heads of ovule parents were removed except those which were selfed.
Then, this largest of all P. uvedalia populations known to me is
thought to be a single clone. If the annual increase in area of
this population could be ascertained, it would be possible to
estimate the age of this population.
Matings 27-3^» whose results appear on Table 7, are crosses
within the same species but between different varieties of that
species. All varieties of each species are cross-fertile with
all other varieties of that same species. Since P. uvedalia is
self-sterile or nearly so, the data of matings 30 and 31 need not
be treated differently from any other in the same table. Crosses
between different varieties of P. uvedalia result in almost 50
per cent fruit set while those between the two varieties of P.
canadensis result in almost 60 per cent fruit set.
A photograph of a hybrid plant (P. canadensis var. radiata
X P. canadensis var. canadensis) in flower from mating 28 is
Figure 33 below. Flowers on this plant are larger than on either TABLE 7 SUMMARY OF INTRASPECIFIC HYBRIDIZATION ATTEMPTS
Number of Number of Per Cent Mating Crosses Fruits Fruits Number Pollen Parent Ovule Parent Attempted Formed Formed
27 P. canadensis var. P. canadensis var. 58 4l 70.7 canadensis radiata
28 P. canadensis var. P. canadensis var. 71 34 V?.9 radiata canadensis
29 P. uvedalia var. P. uvedalia var. 62 36 58.1 densipilis uvedalia
30a P. uvedalia var. P. uvedalia var. 71 52 73.2 densipilis uvedalia
3la P. uvedalia var. P. uvedalia var. 8 4 5 0 .0 uvedalia densipilis
32 P. uvedalia var. P. uvedalia var. 48 21 43.8 densipilis floridana
33 P. uvedalia var. P. uvedalia var. 8 3 37.5 floridana densipilis
3^ P. uvedalia var. P. uvedalia var. 4o 24 6 0 .0 floridana uvedalia
^ 0 emasculation was carried out in these crosses. 150 151
Fig. 35*— P» canadensis var. radiata (pollen parent) X P. canadensis var. canadensis (ovule parent) hybrid. Scale: .70X. of the parents. Flower size is thought to be a manifestation of hybrid vigor. The ray corollas on the hybrid measure about 1 cm
long and 1 cm wide, those on the var. radiata parent were 6 mm
long and 6 mm wide, and those of the canadensis parent were 3 nun
long and wide. The possibility that the large ray corollas result
from the selfing of var. radiata, which normally has large ray
flowers, is ruled out since the var. radiata was used as the
pollen parent. The head photographed in Figure 33 measured 3 cm
in diameter; i.e., from tip of ray flower to tip of ray flower.
The receptacle was 1 cm wide, ray corollas were 1 cm wide and
long, and teeth of the ray corollas were about k mm long. Corol
las of disc flowers were about 5 ®m wide and were translucent. Meiosis in this hybrid appeared normal. I regularly observed 15
pairs of chromosomes in smeared pollen mother cells. 152
Having examined flowers of P. canadensis from several herbaria, I observed that ray flower size varies all the way from about 3 ma up to 1.5 cm in length and from about 5 ejb. to 10 mm in width. Although the extreme conditions were recognized for the experimental portion of this study the ray corolla length is a highly variable character. Neither var. radiata nor var. canadensis can be associated with a specific geographic region within the species range. Therefore, in the light of all evidence it becomes impossible for me to draw a line which separates the two varieties and I have thus placed var. radiata in synonymy with var. canadensis. However, the decision regarding these varieties will be subject to change upon cytological analysis of hybrids.
Soon after heteromorphic pollen (Fisher and Wells, 1962) was discovered in P. laevigata (Wells 25^0« crosses were made in which this plant was used both as the pollen and ovule parent.
Since pollen grains from collection 25^ possessed from one to four nuclei it seemed that there could be several possible chromosome numbers among any hybrids. Of the four fruits formed
from the crosses listed in Table 8 only the single fruit of mating
37 had a seed which germinated. This hybrid has leaves only
slightly different in morphology from all other P. laevigata X
P. canadensis var. radiata hybrids. However, this hybrid has not
yet been examined cytologically.
Figure 3^ is a photograph of another morphological hybrid
variant resulting from the cross P. laevigata X P. canadensis var. TABLE 8 CROSSES INVOLVING P. LAEVIGATA COLLECTION NUMBER 25k
Number of Number of Per Cent Mating Crosses Fruits Fruits Number Pollen Parent Ovule Parent Attempted Formed Formed
35 P. laevigata P. canadensis var, 14 1 7.1 canadensis
36 P, canadensis var. P. laevigata 11 0 0.0 canadensis
37 P, laevigata P. canadensis var, 5 lb 20.0 radiata
38 P, laevigata P. laevigata (selfed) 17 2 11.8
£ All pollen-producing flowers in the heads of ovule parents were removed except those which were selfed. Fruit contained a viable seed which germinated. 153 15^ /
Fig. — £• laevigata (pollen parent) X P. canadensis var. canadensis (ovule parent) hybrid. Scale; .28X. canadensis. The leaves of this plant are smaller and considerably thicker than the usual Polymnia hybrid and, the plant is dwarfed and has a rosette habit of growth. I have examined root tip smears from this hybrid; preliminary counts indicate approximately
30 chromosomes. An exact count of the number of chromosomes has not yet been satisfactorily made.
Two kinds of problems have arisen with respect to hybrid seeds and seedlings. These are: l) in matings 1, 2, 3» 8, 9»
10, and 18 there was for the most part a failure of hybrids to flower, and 2 ) seeds were produced but failed to germinate in matings 1 1 , 1 2 , 1 3 , 14, 1 5 , and 1 7 .
Attempts to induce flowering have included exposure to photoperiods of 8 and 12 hours in a 2A--hour regime, overwintering outside the greenhouse at Columbus, and cutting away all parts 155 above the soil level. Although few hybrids flowered, confir mation of hybridization was possible on the basis of the dominant leaf shape of P. laevigata over P. canadensis. Reciprocal crosses
did not produce contrary results.
I was not able to obtain any germination of hybrid seed
from P. uvedalia crosses. Treatments of fruits included scarifi
cation, placing fruits on moist filter paper for as long as two months, cold treatment of one month at -15°C, cold treatment of
one month at 3°C, placing the fruits in concentrated sulfuric
acid for 1 -3 minutes, and allowing^jsome to overwinter in pots of moistened soil outside the greenhouse. It seemed desirable at
the outset in crosses between varieties of P. uvedalia to know
something about the inheritance of peduncle vesture, since this
is of primary importance at the varietal level. Each type of
vesture retains its integrity when allowed to grow in the same
set of greenhouse conditions. I suggest that this character
may be controlled by a single gene or possibly only a few genes.
Future experimental studies with plants of this genus
will include efforts directed toward obtaining and analyzing
hybrid plants in flower. The scope of coverage will at least
include those taxa which occur naturally in the United States. LIST OF TAXA OF UNCERTAIN STATUS
P. bakeriana Glaziou Soc. Bot. Erance. Mem. 3» 393-^88. 1910.
£• bi^entis Linn, ex Jackson. Index Linn. Herb. 119 (1912) nomen. Quid?
P. dentata Moench. Supplementum ad Methodum Plantas. 2^2. 1802.
P. depressa Sessl and Mocino. Flora Mexicans. 213. 1887.
P. graved ens Rottboell. Acta Literar. Universitatis hafniensis. 299. 1787.
P. odoratissima Sessl and Mocino. La Naturaleza. 1^8-1^9» I89O.
P. scandens Willdenow.
P. uribea Jaramillo. Man. Bosq. y Maderas Tropicales. 2:221. Fig. 134. 1931.
When more is known about the eight taxa listed above most of them will probably be placed among the list of excluded species while some may be placed in synonymy with species now recognized.
The reason for this list, then, is that in my opinion insufficient information is available to justify a more exact treatment.
Polymnia bakeriana Glaziou was first collected in central
Brazil. On the label of this specimen there is a question mark apparently in Glaziou's writing. Perhaps this means that he was
doubtful that this was a new species of Polymnia. No Latin
description accompanies the name but the "original description"
consists of the following: "Polymnia bakeriana Glaz. n. sp.? in
herb. Paris, Berol., Kew., Genev., etc.— Fazenda de Santa Cruz et
156 157
Sao Christovao, Rio Jan., 6029» 7691* Herb., FI. roses.
Octubre-Novembre." A photograph of Glaziou's collection 6029 follows.
Fig. 3 5.— Polymnia bakeriana (Isotype) Scale: .28X
The above photograph was taken from a specimen obtained from the Field Museum in Chicago. This species does not closely resemble known species of Polymnia. Polymnia bakeriana has alternate leaves and fertile disc flowers. These two character istics are sufficient to cause its placement in some genus other than Polymnia. 158
The only reference to P. bidentis known to me is that following its name in the above list of species, Polymnia bidentis is mentioned only by name among a list of other species of Polymnia in the Index to the Linnaean Herbarium. It could be that the specific epithet was inadvertently placed in the wrong genus. Otherwise, it is doubtful that P. bidentis ever existed.
The original description of P. dentata by Konrad Moench in 1802 follows:
Polymnia dentata pericarpiis apice dentatis, foliis connatis ellipticis, subserratis, pedunculis terminalibus ternis: intermedio nudo: lateraJLibus foliosis, calycibus, pentaphyllis.
Caulis herbaceus, subangulatus, ramosus, subscaber. Folia obtusa trinervia, utrinque retrorsum scabra. Rami oppositi, apice trifidi, triflori. Calyx flosculis longior: foliolis ovatis integris, aequalibus, glaberrimis. Flores feminei 8-10 plani emarginati, disco longiores: disci plures quinquefidi. Pericarpia in orbem posita glaberrima, compressa.
This short description suggests a plant resembling P. maculata or possibly P. meridensis but I do not feel prepared to proceed beyond this suggestion.
The original description of P. depressa is also a short one and is quoted from Sesse and Mocino (1 8 8 7).
Radix fibrosa. Caules procumbentes, herbacei, spithamam longp., hirsuti. Folia alterna, pinnatifida, laciniis dentato-lobatis, terminalis saeqius rotundata, indivisa. Petioli vaginantes. Pedunculi oppositifolii, raro interfoliacei, longissimi, erecti, uniflori. Corollae lutae. Semina obovata, plana, margine crenata, Paleae receptaculi lineares, apice latiore, subspath- ulatae, villosae Habitat in pratis de Atotonilco el Grande. Floret Septembri. Perenniis. 159
Several of the above characters seem typical of Polymnia but alternate leaves are not known to be characteristic of any presently recognized species.
The "original description" of P. graveolens is quoted from page 299 in a work by Christian F. Eotboell (1787).
"Polymnia graveolens, venenata. Indis et Negris cara planta, cujus decoctum parva copia summunt, ut efficax antisiphyliticum."
This description is insufficient to delineate this taxon described in 1 7 8 7.
Polymnia odoratissima was originally described by Sessl and Mocino in 1 8 9 0 . This description is a comparatively long one in Latin. After studying the description I infer that P. riparia is the plant being described. The odoriferous quality of foliage is characteristic of many if not all Polymnia species.
No citations to specific collections accompany the description.
P. scandens was apparently described by Carl Willdenow.
Since this is the only reference known to me, all of Willdenow's works will have to be examined for further information before a definite decision can be made.
Attempts to obtain the original description of P. uribea
from some of the largest botanical libraries in the United States and from libraries in Chile and Colombia have been fruitless.
Search for such descriptions will be continued in an effort to ascertain the correct status of these two species. LIST OF EXCLUDED SPECIES
P. abyssinica Linn. f. Suppl. 383. = Guizotia abyssinica Cassini, in Diet. Sc. Nat. lix. 248. 1826-1834.
P. africana Poiret, Encyc. Meth. Bot. Suppl. IV. 48l-483« l8l6. = Wedelia africana Beauv. FI. Owar. II. 19. t. 6 9•
P. aspera DeCandolle, Prodr. Syst. Natur. Regni Vegetabilis, Part 5, 515. 1836. = Axiniphyllum scabrum Blake. Contrib. U. S. Nat. Herb. 26: 2487 1930.
£• australis Spreng. ex DC. Prod. V. 5^2. 1 8 3 6. = Wedelia latifolia DC. Prod. V. 541. 1 8 3 6.
P. benghalensis Poiret ex Steudel Nom. ed. II. 2, 377* = Wedelia calendulacea Rich in Pers. Syn. II. 490.
P. calendulacea Poiret Ency. Suppl. IV. 482. l8l6. - Wedelia calendulacea Rich in Pers. Syn. II. 490.
P. calicina Poiret Encyc. Supp. IV. 482. l8l6. = Wedelia calycina Rich in Pers. Syn. II. 490.
P. carnosa Linn. f. Suppl. 384. = Didelta carnosa Aiton. Hort. Kew. ed. I. III. 2 5 6. 1789*
P. caroliniana Poiret. Ency. Suppl. V. p. 505» 1804. = Berlandiera tomentosa Nuttall. Trans. Am. Phil. Soc. VII. 343^ l W I
P. corcovadensis Glaz. Soc. Bot. France Mem. 393-488. 1910. = Adenostemma viscosum var. triangulare (DC.) (incorrectly attributed to Benth. by) Baker. Mart. FI. Bras. 6, pt. 2: 186. 1876.
P. crenata Poiret Ency. Suppl. IV. 482. l8l6. = Wedelia crenata Rich in Pers. Syn. II. 490.
P. cruciana Poiret Ency. Suppl. IV. 482. l8l 6. = Wedelia cruciana Rich, in Pers. Syn. II. 490.
160 l6l
P. frondosa Bruce. Travels VII. t. 52. 1813. = Guizotia abyssinica Cass, in Diet. Sc. Nat. lix 248. 1826- 18 FU
P. gracilis Poiret. Ency. Suppl. IV. 483. l8l6. = Wedelia gracilis Rich in Pers. Syn. II. 490.
P. grandis Hort. Berol. ex Kunth, Ind. Sem. Hort. Berol. 13. 1 M 7T = Montanoa bipinnatifida C. Koch. Wochenschr. VII: 406. 1864.
P. liebmannii Schultz Bipontius. Leopoldina. 23: 8 9. 1887. = Montanoa liebmannii Blake. Contr. Gray Herb. 52: 37. 1917.
P. mollis Poiret. Ency. Suppl. IV. 482. l8l6. = Wedelia mollis Pers. Syn. II. 490.
P. nervata Jones. Contr. West. Bot. 12: 44. 1908. = Montanoa patens Gray. Proc. Amer. Acad. 21: 3 8 8. 1886.
P. parviflora Poiret Ency. Suppl. IV: 482. 1816 . = Wedelia parviflora L. C. Rich. Pers. Syn. II. 490.
P. perfoliata Poiret Ency. Suppl. V. 5 0 6. l8o4. = Melampodium perfoliatum H. B. and K. Nov. Gen. et Sp. IV. 27^1 1820.
P. revoluta Auct. ex Steud. Nom. ed. II 2: 377. = Melampoditun perfoliatum H. B. and K. Nov. Gen. et Sp. IV. 27^. 18201
P. revoluta Auct. ex Steud. Nom. ed. II 2: 377. = Melampodium perfoliatum H. B. and K. Nov. Gen. et Sp. IV. 27^U 1820.
P. scabra Zucc. Abh. Akad. Wiss. Miinchen. 1: 313. 1832. = Axiniphyllum scabrum Blake Contrib. U. S. Nat. Herb. 26: 243: 1950:
P. serrata Poiret Ency. Suppl. IV. 482. l8l6. = Wedelia serrata Rich, in Pers. Syn. II. 490.
P. spinosa Linn. f. Suppl. 384. = Didelta spinosa Aiton. Hort. Kew. ed. I. 3: 2 5 6. 1 7 8 9.
P. standleyi Steyermark. Field Mus. Pub. Bot. 23: 106. 1944. e Rumfordia standleyi (Steyerm.) Standi, and Steyerm. ?ield Mus. Pub. Bot. 23: 26>5* 1947. tarapacana Phil. Index Kewensis Suppl. Is 3^1 . 1886-1895. (mentioned by name only). = Polylepis tarapacana Philippi. Anales del Museo Nacional Chile. 2l. ”1891“ --- tetragonotheca Linn. Syst. ed. XII. 576. = Tetragonotheca helianthoides Linn. Sp. PI. 905* 1753* verapazensis Standley and Steyermark. Field Mus. Pub. Bot. 23:l4*f. 1 9 ^ . = Bumfordia guatemalensis (Coult.) Blake. Journ. Wash. Acad. Sci. 1 8 : 25. 1928. wedelia Linn. Mant. 1: 118. = Wedelia fructescens Jacq. Select. Am. 217. t. 130, or in sensu Hook., f. ex Benth. in Journ. Linn. Soc. 13s ^39 • 1873* • • •«
• • •• • • •• • ••
P. conaden si s
2 0 0 MILES
l-'ig. 36 .— Known distribution of Polymnia canoiensis excluding Canada. P. u v e d a I i a D z var. dens ipi I is F : va r. floridana 9: var. uvedalia
200 MILES
Fig. 37»— Known distribution of Folymnia uvedalia excluding Bermuda. 165
i 4
P. laevigata
200 MILES rig. 3S.— Known distribution of Polymnia laevigata. 20
<6
P. oaxaca P. quiche
SCALE lyOMILCS
CONIC PROJECTION
1(0 105 100 95 90 65 80 166 Fig. 39.— Known distribution of Polymnia oaxacana and P. quichensis. 0 P. macuiata ad e nof r i cha
SCALE
CONIC PRO JEC TIO N
o^ -^3 Fig, — Known distribution of Polymnia apus and P. macuiata var, adenotricha 25
20
acuiato var. maculata
A P- moculata var. glabricaulis
SCALE
CONIC PROJECTION
Fig, *fl.— Xnown distribution of Polymnia maculata vars. maculata and glabricaulis. 169
SOUTH AMERICA
SCALE
CONIC PROJECTION
Fig. k2%— Known distribution of Polymnia riparla. 170 SOUTH AMERICA
10 «
Fig, k^>,— Known distribution of Polymnla glabrate., £• parviceps, and P, microcephala. 171
SOUTH AMERICA
A P. c on n o to
£ P. py.ramidalis
SCALE
SINUSOIDAL PROJECTION
Fig. hk,— Known distribution of Polymnia pyramidalis and P. connata. 172
SOUTH AMERICA
£ P. fruticosa £ P. >i Fig, ^5,— Known distribution of Polymnia siegesbeckia, P, fruticosa, and P. suffruticosa, 173 SOUTH AMERICA A P . in • f i d « n a i » 0 . P. aoftehifalin Fig. ^6»— Known distribution of Polymnia sonchifolia and P« meridensis. SUMMARY A comprehensive study of the genus Polymnia has been undertaken from the approach of classical and of experimental taxonomy. The experimental investigations involve only the taxa which grow naturally in the United States. Nineteen species and six varieties are presently recognized. These totals are subject to change as future experimental evidence is gained. During the course of this study I have examined ap proximately 2 ,0 0 0 herbarium specimens from 13 herbaria located in the United States. Many type specimens were examined; where types themselves were not available photographs of types were examined when possible. Both the types and the photographs of other types were of inestimable value in helping ascertain the known limits of each taxon. The distribution map for each taxon is based on herbarium specimens examined. 174 BIBLIOGRAPHY Arechavaleta, J. 1905* Nueva contribuci6n al conocimiento de la flora del Uruguay. Anales del Museo Nacional de Montevideo. l:ser. 2, 35-38. _____ . 1908. Flora Uruguay III. Anales del Museo Nacional de Montevideo. 6:295* Baker, J. G. in K.F.P. von Martius. 1 8 7 6. Flora Brasiliensis. Vol. 6:part 2. . l88*f. Flora Brasiliensis. Vol. 6:part 3* Badillo, V. M. 19^6. Contribucion al conocimiento de la sistematica y distribuciSn geografida de las compuestas en Venezuela. Boletin de la Sociedad Venezolana de Ciencias Naturales. 10:279-320. Beadle, C. D. 1 8 9 8. Notes on the botany of the southeastern states. I. Botanical Gazette. 255 2 7. 8 Bentham, George. 18^5* Plantae Hartwegianae. Bentham, George, and J. D, Hooker. 1 8 7 6. Genera Plantarum. Vol. 2, Reeve and Co., London. Bettfreund, Carlos. 1900. Flora Argentina. Vol. 2. Bipontius, Schultz in F. W. Klatt. 1 8 8 7. Beitrage zur Kenntniss der Compositen. Leopoldina. 23:89. Blake, S. F. 1917a. New and noteworthy Compositae, chiefly Mexican. Contributions from the Gray Herbarium. 52:3^. ______. 1917b. Polymnia uvedalia and its varieties. Rhodora. 19:219. ______. 1922. New South American Asteraceae collected by W. D. Holway. Botanical Gazette. 7^:^* ______. 192^t. New American Asteraceae. Contributions from the National Herbarium. 22:60^-605. ______. 1926. Five new American Melampodiinae. Journal of the Washington Academy of Sciences. 16:421. 175 176 Blake, S. F. 1930. Notes on certain type specimens of American Asteraceae in European herbaria. Contributions from the United States National Herbarium. 26:227-263. Blake, S. F. in Gleason. 1931. Botanical results of the Tyler- Duida expedition. Bulletin of the Torrey Botanical Club. 58:487-488. Blake, S. F. 1935* New Asteraceae from the United States, Mexico, and South America. Journal of the Washington Academy of Sciences. 25:313-314. ______. 19^5* Asteraceae described from Mexico and the south western United States by M. E. Jones, 1908-1935* Contri butions from the United States National Herbarium. 29: 132. Cavanilles, A. J. 1794. leones et descriptiones plantarum, quae aut in hortis hospitantur. Vol. 3* Comar, C. L., and F. P. Zscheile. 1942. Analysis of plant extracts for chlorophylls a and b by a photoelectric spectrophotometric method. Plant Physiology. 17:198-209. Core, E. L. 1955. Plant Taxonomy. Prentice-Hall, Inc., Englewood Cliffs, N. J. Coulter, John M. 1895* New or noteworthy Compositae from Guatemala. Botanical Gazette. 20:48-4-9. Cuatrecasas, Josl. 1954. Notas a la flora de Colombia XIII. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales. 9535* DeCandolle, A. P. 1 8 3 6. Prodromus Systematis Naturalis Regni Vegetabilis. Vol. 5. Fassett, N. C. 1932. Notes from the herbarium of the University of Wisconsin— VII. Rhodora. 34:96. Fernald, M. L. 1950. Gray1s Manual of Botany. American Book Company, New York. Fisher, T. Richard, and J. R. Wells. 1962. Heteromorphic pollen grains in Polymnia. Rhodora. 64:336-340. Glaziou, A.F.M. 1910. Plantae Brasiliae centralis a Glaziou lectae. Societe Botanique de France. Memoires. 3! 393-488. Gleason, H. A. 1952. The New Britton end Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. Vol. 3* The New York Botanical Garden, New York. I. 177 Gray, Asa. l88l. Gray's Lessons in Botany and Vegetable Physiology. Ivison, Blakeman, Taylor and Co., New York. 1884. Synoptical Flora of North America. Vol. 1, part 2. Ivison, Blakeman, Taylor and Co., New York. Greenman, J. M. 1903. New and otherwise noteworthy Angiosperms from Mexico and Central America. Proceedings of the American Academy of Arts and Sciences. 39s99» Hieronymus, G. 1900. A. Sodiro, S. J.: Plantae Ecuadorenses II, Botanische Jahrbucher fur Systematik, Pflanzen- geschichte und Pflanzengeographie. 29:33* ______. 1901. Plantae Lehmannianae praesertim in Colombia et Ecuador collectae, additis qxiisbusdam ab aliis collector- ibus ex iisdem regionibus allatis determinatae et descriptae. Compositae II. Botanische Jahrbuchner fur Systematik, Pflanzengeschichte und Pflanzengeographie. 28:599-600. ______. 1905. Plantae Peruvianae a claro Constantino de Jelski collectae. Compositae. Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie. 36:484-485. Humboldt, F.H.A., J. A. Bonpland, and Carolus Kunth. 1820. Nova Genera et Species Plantarum quas in Peregrinatione Oribus Novi Collegerunt, Descripserunt, partim adumbraverunt. 4:282-283. Jackson, B. Daydon. 1 8 8 5. Index Kewensis. Vol. 2 and supple ments. Lanjouw, J., and F. A. Stafleu. 1959* Index Herbariorum. Fourth Edition. 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Steyermark. 19*+^a. Studies of Central American Plants— IV. Botanical Series, Field Museum of Natural History. 23:106. _. 19^b. Studies of Central American plants— V. Botanical Series, Field Museum of Natural History. 23:1*14. ______. 19^7. Studies of American plants— VII. Botanical Series, Field Museum of Natural History. 23:265. Steyermark, Julian A. et al. 1953* Contributions to the flora of Venezuela. Botanical Exploration in Venezuela— III. Fieldiana: Botany. 28:667-668. Triana, J. M. 1 8 5 8. Choix de plantes de la Nouvelle-Grenade. Annales des Sciences Naturelles. Botanique, Series *f. 9:*K)-4l. Turner, B. L., W. L. Ellison, and R, M. King. 1961. Chromosome numbers in the Compositae. IV. North American species with phyletic interpretations. Am. Jour. Bot. ^8:216-228. Weddell, H. A. 1857. Notice sur L'Ahipa et L'Aricoma, Plantes Alimentaires Du Haut— Perou. Annales des Sciences Naturelles. Botanique, Series k, 7:11^. Weishaupt, Clara, i9 6 0. Vascular Plants of Ohio. Published by Harold L. Hedrick, Columbus, Ohio. AUTOBIOGRAPHY I, James Ray Wells, was born in Delaware, Ohio, on May 28, 1932. My grade school attendance was in south and north Alabama, South Carolina, and Florida. I attended high schools in Ohio and in east and middle Tennessee. The high school diploma was obtained at Isaac Litton High School in Nashville, Tennessee, in 1950* Colleges attended were: Trevecca College and Peabody College in Nashville, University of Tennessee in Knoxville, University of Michigan, and The Ohio State Uni versity. One year was spent at Trevecca College and another at Peabody College. I then transferred to the University of Ten nessee at Knoxville where I received the B.S. degree in 195^ and the M.S. in 1956. I was enrolled at the University of Michigan for the summer session of 1955* Military service was in the U. S. Army and I spent most of a 2-year enlistment at the Chemical Corps Research and Development Command in Frederick, Maryland. I then taught for one year in the Miamisburg, Ohio, public school system. My first quarter at The Ohio State Uni versity was in the summer of 1959 and I have been in regular attendance since. Beginning in the fall of 1963 I have accepted a position as Assistant Professor in the Department of Biology at East Carolina College located in Greenville, North Carolina. 179