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LOWDEN, Richard Max, 1943- REV I SI ONARY AND POPULATION STUDIES IN THE AMERICAN AQUATIC PLANT GENUS PONTEDERIA L.

The Ohio State University, Ph.D., 1971

University Microfilms, A XEROX Company , Ann Arbor, Michigan REVISIONARY AND POPULATION STUDIES IN THE AMERICAN

AQUATIC PLANT GENUS PONTEDERIA 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

Richard Max Lowden* B,A., M.Sc.

The Ohio State University

1971

Approved by

Adviser Graduate Program in Botany To my Parents for their continued interest in Education ACKNOWLEDGMENTS

I have sincerest admiration for my industrious assistant,

Rafaela Joaquin, my wife, for the countless hours she rendered to this

research in the field and laboratory. The receipt of a Grant-In-Aid of

Research from The Society of the Sigma Xi is acknowledged, which en­

abled me, while carrying out other research, to study populations in

British Honduras during December 1969. I am most indebted to the

Organization for Tropical Studies, Inc. (OTS) for an introduction to

tropical Biology (Costa Rica, summer of 1968) and for granting an OTS

Pilot Research Grant F 70-23 (The Ohio State University Research

Foundation Project No. 3031-Al) to R. L. Stuckey (Faculty Adviser,

Department of Botany, The Ohio State University) and myself (Principal

Investigator) for aid in performing the necessary field research in

Mexico and Central America during the summer of 1970.

To the Graduate Committee of the Department of Botany and the

Graduate School Fellowship Committee (The Ohio State University), I am

most grateful for the University Dissertation Year Fellowship awarded

to me for the completion of this investigation.

Appreciation is expressed to Curators of the following herbaria

for specimens loaned: Brooklyn Botanic Garden (BKL), Brooklyn,

National Museum of Canada, National Herbarium (CAN), Ottawa, Ontario;

Duke University Herbarium (DUKE), Durham, North Carolina; Field Museum

of Natural History, Department of Botany (F), Chicago, Illinois; The Florida State University Herbarium (FSU), Tallahassee, Florida; Arnold

Arboretum and Gray Herbarium of (A, GH), Cambridge,

Massachusetts; The University of Michigan Herbarium (MICH), Ann Arbor,

Michigan; New England Botanical Club Herbarium (NEBC), Cambridge,

Massachusetts; Missouri Botanical Garden (MO), St. Louis, Missouri;

New York Botanical Garden (NY), Bronx, New York; The Ohio State

University Herbarium (OS), Columbus and The Franz Theodore Stone

Laboratory Herbarium (FTSL), Put-In-Bay, Ohio; The Academy of Natural

Sciences of Philadelphia (PH), Philadelphia, Pennsylvania; Smithsonian

Institution, Department of Botany (US), Washington, D. C.: and

University of Wisconsin Herbarium (WIS), Madison, Wisconsin. Voucher specimens for investigations in Pontederia have been deposited in The

Ohio State University Herbarium, Columbus, Ohio.

Constructive criticism of the manuscript by J. A. Schmitt,

E. D. Rudolph, R. L. Stuckey, and T. F. Stuessy (Dissertation Reading

Committee, Department of Botany, The Ohio State University) is grate­ fully appreciated. Thanks are extended to R. M. Giesy, T. J* Johnson

(Department of Botany, The Ohio State University) and D* W. Stevenson

(Graduate Program in Botany, The Ohio State University), for assistance in photography of plates.

iii VITA

September 27, 1943 . . . Born, Columbus, Ohio

June 1964 ...... B.A., Botany Department, The Ohio State University, Columbus, Ohio

June 1964-June 1966 . . Peace Corps Volunteer, United States Peace Corps, taught Biology and Mathematics at Austin High School, Stann Creek Town, British Honduras, Central America

December 1967 ..... M.Sc., thesis entitled "A Vascular Flora of Winous Point, Ottawa and Sandusky Counties, Ohio,’' Botany Program, The Ohio State Uni­ versity, Columbus, Ohio

September 1966-June 1968 Research Assistant to Professor T. R. Fisher, National Science Foundation Grant, The Ohio State University, Columbus, Ohio

July-August 1968 .... Student in the course, Tropical Biology--An Ecological Approach, sponsored by the Organization for Tropical Studies (OTS) in Costa Rica, Central America

September 1968-June 1970 Teaching Assistantshlp, Graduate Program in Botany, The Ohio State University, taught Biology 100 during Autumn quarter 1968 and Winter quarter 1969; Botany 410 during Spring and Summer quarters 1969; and Botany 102 during Spring quarter 1970

January 1970 ...... Awarded a Grant-In-Aid of Research from The Society of The Sigma Xi for study of "William A. Schipp's Botanical Explorations in the Stann Creek and Toledo Districts, British Honduras 1929-1935"

May 1970 ...... A Pilot Study research grant was awarded by The Organization for Tropical Studies to Associate Professor Ronald L. Stuckey (Faculty Adviser to Mr. Lowden, Academic Faculty of Botany, The Ohio State University) and Mr. Richard M. Lowden (Principal Investigator)

iv aiding field research carried out in Central America for this Dissertation

June 27, 1970 Married Rafaela Andrea Joaquin of the Dominican Republic

June 1970-June 1971 • . Awarded a University Dissertation Fellowship, Graduate School, The Ohio State University, Columbus, Ohio

4 PUBLICATIONS

Lowden, Richard M. 1969. A Vascular Flora of Winous Point, Ottawa and Sandusky Counties, Ohio. Ohio J. Sci. 69 (5):257-284.

______. 1970. William Ashbrook Kellerman’s Botanical Expeditions to Guatemala (1905-1908). Taxon 19 (l):l9-35.

1970. William A. Schipp's Botanical Explorations in the Stann Creek and Toledo Districts, British Honduras (1929-1935), Taxon 19 (6):831-861.

FIELDS OF STUDY

Major Field: Botany

Studies in Vascular Plant Taxonomy (Plant Systematics). Associate Professor Ronald L. Stuckey

Studies of Botanical Collectors and Explorers in Central America. Professor Emanuel D. Rudolph and Associate Professor Ronald L. Stuckey

v CONTENTS Page

ACKNOWLEDGMENTS...... 11

VITA ...... lv

TABLES ...... vll

FIGURES ...... vlil

PLATES ...... xii

PONTEDERIA L. THE GENERIC NAME!: NOMENCLATURE SINCE 1737 .. . 1

REVISION OF PONTEDERIA L.: TAXONOMY ...... 5 Disposition of Excluded Names ...... 197

A GENERIC BASIS: CYTOLOGY ...... 199

GEOGRAPHICAL DISTRIBUTION OF PONTEDERIA L.: THE AMERICAS . . . 204

A TRISTYLOUS GENUS WITH AN ISOLATED FLORAL FORM: POPULATION REPRODUCTIVE BIOLOGY ...... 211

STATISTICAL ANALYSIS OF PHENOLIC QUALITATIVE FEATURES: CHROMATOGRAPHIC D A T A ...... 228

B10SYSTEMATIC ASPECTS: EVOLUTIONARY IMPLICATIONS ...... 245

APPENDIX (TABLE 7 ) ...... 251

LITERATURE CITED ...... 286

vl TABLES Table Page

1 Representative length measurements of perigones, stamens and styles in Pontederia L...... 195

2 Seed germinations in growth chamber conditions, with and without hardened perigone bases ...... 210

3 Indication of floral form frequency in Pontederia L. based on population and herbarium studies . . . • 222

4 Spot colors of phenolic compounds in Pontederia L. and related genera ...... 237

5 Summary of spot occurrence of phenolic compounds in Pontederia L. and related genera (Table 4) • . ■ . 238

6 A comparison of two dimensional paper chromatographic profiles of Pontederia L. and related genera based on summary of phenolic spot occurrence (Table 7) , 240

7 Spot occurrence of phenolic compounds in Pontederia L. and related genera as exemplified by population study and herbarium specimens examined ...... 251

vii FIGURES Figure Page

1 Numerous blue-purple flowered inflorescence) clasping subtending spathe and flowering stalk leaf of Pontederia cordata L. var. cordata, U.S., Ohio, Ottawa County, V/inous Point"! 3% miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay ...... 22

2 Pontederia cordata L. var. cordata in marsh habitat, U.S., Ohio, Ottawa County, Winous Point, 3% miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay 22

3 Fruiting specimen of Pontederia cordata L. var. cordata, showing cordate flowering stalk leaves and clasping sheath at base of flowering stalk stem shoot, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS) ...... 23

A Fruiting and flowering specimen of Pontederia cordata L. var. cordata, showing sagittate-hastate leaves, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS) ...... 25

5 Geographical distribution of Pontederia cordata L. var. cordata in Ontario, Quebec, New Brunswick, Prince Edward Island and Nova Scotia Provinces of Canada . 27

6 Geographical distribution of Pontederia cordata L. var. cordata (dots), var. lancifolia (Muhl.) Torr. (triangles), and var. ovalis (Hart.) Solms (stars) in the United States 29

7 Numerous white flowered inflorescence, clasping sub­ tending spathe and flowering stalk leaf of Pontederia cordata L. var. cordata in British Honduras, C.A., 9 miles from Belize City, along the Northern Highway ...... 31

Pontederia cordata 1.. var. cordata in drainage ditch habitat, British Honduras, C.A., 9 miles from Belize City, along the Northern Highway ...... 31

vili FIGURES (continued) Figure Page

9 Provisional lectotype of Pontederia lancifolla Muhl.( in Muhlenberg's Herbarium (PH), showing a narrow lanceolate flowering stalk leaf ...... 110

10 Holotype of Pontederia lanceolata Nuttall (specimen on the left of the herbarium sheet) collected at Savannah, Georgia (PH), showing a narrow lanceolate flowering stalk l e a f ...... 112

11 Geographical distribution of Pontederia cordata L. var. cordata (dots) in British Honduras (solid lines represent highways), C.A., and var. lancifolla (Muhl.) Torr. (triangles) in Cuba, West Indies .... 114

12 Pontederia cordata L. var. lanceolata (Nutt.) Grisebach, Cuba, Plantae Cubenses Wrightianae, C. Wright 3260 (US 33961), showing narrow lanceolate leaves ...... 116

13 Holotype of Pontederia oval is Mart, in Roemer & Schultes, (M), photograph (US), showing ovate leaves 137

14 Specimen cited as Pontederia ovalis Mart, in Martius, Flora Brasiliensis, collected by George Gardner 3483 ( N Y ) ...... 139

15 Geographical distribution of Pontederia cordata L. var. cordata (dots), var. lancifolia (Muhl.) Torr. (triangles) and var. ovalis (Mart.) Solms (stars) in ...... 141

16 Holotype of Pontederia lanceolata Nutt. var. vichadensls Hermann, 21 Jan. 1944, , Comisaria del Vichada, San Jose de Ocune, F. J . Hermann 11045 (US 2169640) 143

17 Numerous-flowered inflorescences of Pontederia sagittata Presl growing in association with Sagittaria, Honduras, Dept. Cortes, Puerto Cortes, 3 Ave.-5 Calle 0 ...... 148

18 Pontederia sagittata Presl with drooping fruit clusters and sagittate leaves in a woody habitat, Mexico, Laguna Catemaco, Arroyo Agrio, 6 kms. from Catemaco town, between Coyame and San Andres Tuxtla .... 148

ix FIGURES (continued) Figure Page

19 Pontederia sagittata Presl, sagittate flowering stalk leaves, Mexico, State of Vera Cruz, along the Trans Isthmian highway (route 185), 17 kilometers northeast of Minatitlan, R M. King 933 (US 2397514) 149

20 Geographical distribution of Pontederia sagittata Presl (Presl) Schery in Mexico and Central America .... 151

21 Young plant specimen of Pontederia sagittata Presl, showing short petioles of flowering stalk leaf (arrow) and long petioles of leaves attached to rhizome, Mexico, State of Chiapas, fields along route 225, ca. 1 mile north of Suchiate, R. M. King 3430 (MICH) ...... 153

22 Numerous-white-flowered inflorescences of Pontederia parvlflora Alex., shallow cordate based leaves, Panama, Province of Herrera, Los Llanos de Santa Maria 163

23 Pontederia parviflora Alex, in wet field habitat, Panama, Province of Herrera, Los Llanos de Santa M a r i a ...... 163

24 Flowering specimen of Pontederia parviflora Alex., showing clasping subtending spathe and flowering stalk leaf, Panama, Province of Herrera, Los Llanos de Santa Maria, Lowden 20 (OS)...... 164

25 Geographical distribution of Pontederia parviflora Alex, in Panama and Colombia, stars represent sighted records 166

26 Fruiting specimen of Pontederia parviflora Alex. Schery, Panama, Province of Herrera, Los Llanos de Santa Maria, Lowden 20 (OS) 168

27 Fewer-flowered inflorescence of Pontederia rotundifol1a L. f. showing two yellow-orange spots on the largest perianth lobe, El Salvador, Dept. La Libertad, Laguna de Zapotitan ...... * . ..173

28 Pontederia rotundifolia L. f. in a swampy habitat at Pito Solo, south end of Lago Yojoa, Dept. Comayagua, Honduras ...... 173

x FIGURES (continued) Figure Page

29 Reniform leaves and short inflorescence of Pontederia rotundifolia L. f., Brazil, Brazilian Amazonia, Territory of Rondonia, Prance, Forero, VJrigley, Ramos & Farias 5914 (NY^ • * • '. . '. I I . . 7 • . • 174

30 Geographical distribution of Pontederia rotundifolia L. f. in Central America, arrow points to insert of canal zone ...... *.*..,.176

31 Sagittate leaves of Pontederia rotundifolia L. f. attached to trailing prostrate shoot, Nicaragua, Dept. Granada, Lago Nicaragua, Lowden 18 (OS) . . . 178

32 Fewer-flowered inflorescence and subovate loaves of Pontederia subovata (Seub. in Mart.) Lowden, arrow points to a ligule of a petiole sheath, Paraguay, in regione lacus Ypacaray, E. Hassler 12503 (MO 8 4 7 6 2 8 ) ...... 188

33 Geographical distribution of Fontederia rotundifolia L. f . (dots) and Pontederia subovata (Seub. in Mart.) Lowden (triangles) in South America ...... 190

34 Narrow subovate leaf variant of Pontederia subovata (Seub. in Mart.) Lowden, Brazil, Minas Gerais, Municipio Lagoa Santa, Lagoa Santa, L. B. Smith 6702 (US 2173371) ...... 192

35 Legitimate and illegitimate pollinations in Pontederia L. floral forms, arrows represent the transfer of pollen from anthers to stigmas, code to floral forms (Sml) SHORT STYLE, 3 medium stamens, 3 long stamens; (sMl) 3 short stamens, MEDIUM STYLE, 3 long stamens; (smL) 3 short stamens, 3 medium stamens, LONG STYLE; (slL) 3 short stamens, 3 long stamens, LONG S T Y L E ...... 218

36 Summary two-dimensional paper chromatographic profile of phenolic compounds found in Pontederia L. and related genera 235

37 Graphic representation comparing two dimensional paper chromatographic profiles of Pontederia L. taxa based on coefficients of similarity (Table 6) . . . 243

38 Speculative scheme of evolution. Arrows represent divergent evolution from a hypothetical ancestral stock ...... 249 xi PLATES

Plate Page

I Inflorescence, pendulous ovule and fruits...... 8

II Seeds, endosperm and embryo 10

III Hardened perigone bases 12

IV Hardened perigone bases ...... 14

V Chromosome configurations in microsporocytes . . . • . 202

VI Floral forms 220

xii PONTEDERIA L. THE GENERIC NAME!:

NOMENCLATURE SINCE 1737

Conspectus of Pontederia L.

Carl Linnaeus (1737), In communication with Gronovius, described

Pontederia In the first edition of Genera Plantarum as having ovate trllocular capsules with many seeds, flowers with six stamens and two­ lipped perlgonea (perianth). In the Hortus Cllffortianus (1737),

Linnaeus dedicated the genus to Julio Pontedera (1688-1757), Professor of Botany, Padua, at which time Linnaeus had chiefly in mind the

American Pickerelweed of Maryland and Virginia. Specific reference was also made to the Virginian plant, with a spike inflorescence and auricled leaves, cited by Petiver, Morlson and Plukenet. In the Species

Plantarum of 1753, Linnaeus enumerated three species of Pontederia In the class Hexandria: (1) ovata, habitat in Malabar [a plant with one stamen, Incorrectly placed by Linnaeus in the class Hexandria, and now belonging to the Marantaceae], (2) P. cordata, habitat in Virginia, with reference to Hortus Cl iffortianus (1737) and Gronovius Cthe

American Pickerelweed with indehiscent one-seeded utricles and spike

Inflorescences] and (3) P^. hastata, habitat in India; Cthe Monochoria

Presl of today, with dehiscent many-seeded capsules and umbelliform

Inflorescences]. Linnaeus repeats the same fruit description of the genus in the fifth edition of Genera Plantarum (1754): capsules tri- locular and many-seeded. Adanson (1763) made the generic name Naruktla synonymous with the Linnaean Potederia. It is clear In Species Plantarum (1753) and the first and fifth editions of Genera Plantarum (1737, 1754) that to Linnaeus

(Fernald, 1925), "Pontederia vas a group of superfically similar but structurally quite dissimilar plants." However, Rafinesque (1830) stated that the Linnaean generic description of Pontederia called for capsules trilocular and many-seeded, therefore Monochoria Presl

(Pontederia hastata L.) must be Pontederia and the one-seeded utricle plant, _P. cordata L., was placed in Rafinesque's new genus Unlsema

(1808, Umsema). This was perhaps the first occasion (Rafinesque, 1808) that the nomenclature as applied to Pontederia by Linnaeus was questioned and brought to the attention of American botanists. Ten new species and numerous varieties of Unlsema for North America were described in "characteristic style" by Rafinesque (1830, 1833, 1836,

1837). Unlsema was later revived hy Farwell (1924, 1928) who also restricted the genus Pontederia to P_. hastata, the only species named by Linnaeus (1753), that had a many-seeded, three-celled capsule.

According to the International Code of Botanical Nomenclature

(1966, Article 52), "when a genus is divided into two or more genera, the generic name must be retained for one of them." Pursh (1814),

Nuttall (1818), Torrey (1824), Solms-Laubach (1883), Sprague (1924),

Fernald (1925), Schwartz (1927), and Castellanos (1951) have retained and reinstated Pontederia cordata L. as the type of the American

Pontederia L. Hitchcock and Green (1929) suggested _P. cordata L. as the "standard-species" of Pontederia L ., since it Is "the best known of the three original species and the only one now retained in the genus." Established custom which has prevailed over the last two centuries has recognized P. cordata (the American Pickerelweed with unilocular one-seeded indehiscent fruits) as the type species of

Pontederia, even though Linnaeus described the genus as having tri- locular many-seeded capsules.

In the present treatment, I also follow this precedent and propose that Pontederia cordata be conserved as the nomenclatural type of the Linnaean Pontederia in accordance with established custom which has prevailed over the last two centuries. If cordata L. is not conserved as the nomenclatural type of Pontederia L., then taxono­ mists must adopt another generic name for the American Pickerelweed and probably accept P. hastata L. (nomenclatural type of Monochoria

Presl) as the nomenclatural type of Pontederia L. This latter course of action would result in unnecessary nomenclatural changes and con­ fusion for those taxonomists who associate Pontederia L. with the

American plant and Monochoria Presl with the Asian plant.

Conspectus of Reussla Endl.

In Solms-Laubach1s (1883) treatment of the Pontederiaceae,

Pontederia L. was recognized as being generally distinct from Reussla

Endl. Both have one-seeded indehiscent fruits and two-lipped perigones

(perianth). Pontederia was characterized by having 3 lobes in each lip, whereas, Reussla was characterized by having 5 lobes in the upper lip and 1 lobe in the lower lip. This classification scheme was followed by

Schwartz (1927, 1930), Schulz (1942), and Castellanos (1951, 1958). In accordance with this distinction, Castellanos (1951) transferred

Pontederia rotundifolia L. f• to Reussla based on his reported obser­ vation of 5 lobes in the upper lip of the perigone. From my own observations In nature, there Is no doubt that

Pontederia species (Figs. 1, 7, 22; P. subovata was not observed In 3 nature) Including P. rotundifolia (Fig. 27) have the y arrangement, even though the deepest incisions of the perigone occur on both sides of the smallest lobe of the lower lip. However, the character of the 3 5 number of lobes In each lip, either y or y, appears to be unsuitable for distinguishing Pontederia from Reussla. It Is next to impossible to discover from dried specimens the number of lobes in each lip.

Additional evidence obtained in the present investigation indicates that Pontederia and Reussla are so morphologically, cytologically and chemically Blmilar as to warrant the treatment of Reussla as a subgenus of Pontederia. In the International Code (1966, Appendix III, p. 226 and Article 14), Reussla is a conserved name, therefore it competes with Pontederia, each based on different nomenclatural types and against which Reussla is not explicitly conserved. Pontederia is the earlier name, thus it has priority over Reussla. REVISION OF PONTEDERIA L.: TAXONOMY

Pontederia Linnaeus, Sp. PI. 288. 1753

Narukila Adanson. Families Des Plantes 54, 581. 1763.

Umsema Raf. [=Unlsema Raf.] Med. Repos. II 5:352. 1808.

Pontederas Corth. mut.] Hoffmannsegg, Verz. Pfl. 137. 1824.

Unlsema Raf., Medical Flora 2:105. 1830.

Reussla Endl., Genera Plantarum 1:139, 1836.

UnlBemma Corth. mut.3 D'Orbigny, Dictionnaire Universal D ’Histoire Naturelle 12:761. 1843.

Fontederaea (ia) Corth. mut.3 O. Kuntze non L., Rev. Gen. 2:719. 1891.

Hirschtia K. Schum. ex Schwartz in Engler, Bot. Jahrb. 61 (no. 139):41. 1927.

Aquatic perennial herbs, emergent to floating; rhizomes branched, stems erect or prostrate; leaves (phyllodes) lanceolate, ovate, cordate, sagittate, hastate or reniform; leaf of floral shoot with short, other­ wise leaves with long petioles (Fig. 21); petiole bases sheathed and clasping stems, sheath of petiole with a ligule (Fig. 32); leaves obtuse, entire margined with parallel to outwardly curved venation; foliage ranging from dark green-pale green-grayish; vegetative parts often containing anthocyanin; peduncles longest with maturation of infruc- tescences (Fig. 18) and subtended by a spathe (the terminal phyliode of the floral shoot; very early in floral development the inflorescence Is enclosed by the spathe, Plate I-a, and the petiole base of the flower

5 stalk leaf from which it emerges); sheaths (expanded phyllode bases) in the axil of floral shoots (Fig. 3); inflorescence a few to many- flowered spike (Fig. 1), flowering throughout the year in the tropics; flowers (Figs. 1, 27) bisporangiate, zygomorphic (bilaterally sym­ metrical), bilabiate, blue-purple, white or white-green, largest lobe of perigone with a bilobed yellow Bpot; young perigones (Plate VI) of

6 basally connate parte; stamens 6, unequal, in two groups of 3 each, adnate to the perigone tube; anthers introrse, dorsifixed, versatile, dehiscing longitudinally; young perigone tube and filaments covered with hairs terminated with pinheadlike glands; species heterostylous

(tristylous) or homoatylous (Plate VI); stigmas 3-lobed with each lobe bifid; carpels originally 3, united, with 2 aborting and 1 remaining, unilocular and 1-seeded; ovules pendulous (Plate I-b), anatropous; placentation parietal; pericarp high in glutanous content; seeds ovoid or reniform (Plate Il-a, d), enclosed by hardened perigone bases,

toothed, smooth or spinulose ridged (Plates III, IV); endosperm white mealy surrounding a long cylindrical embryo (Plate Il-b, c); fruit a utricle (Plate I-c, d; 1-seeded and indehiscent).

Type species: Pontederia cordata L., U.S., Virginia, communicated to

Linnaeus by Gronovius. (Type not seen, the possibility

of a type in the Linnaean Herbarium is being investi­

gated).

Distribution: The Americas. 7

KEY

Hardened perigones toothed or smooth ridged (Plate III; Plate IV-a-b); floral bearing shoots erect (subgenus Pontederia):

Style length unequal to stamen length (Plate Vl-a-c); leaves cordate, sagittate, reniform, ovate or lanceolate.

Hardened perigones tooth ridged (Plate III);

leaves cordate, sagittate (unauricled), hastate or reniform ...... la. F_. cordata var, cordata

leaves narrow, lanceolate (leaf base not lobed) ...... lb. P^. cordata var. lancifolla

leaves ovate to ovate-lanceolate (leaf base not lobed) ...... lc. JP. cordata var. ovalis

Hardened perigones smooth ridged (Plate IV-a);

leaves sagittate (usually deeply auricled) ...... 2* P.- sagittata

Style as long as the three longest stamens (Plate Vl-d);

leaves subcordate ...... 3. P_. parviflora

Hardened perigones spinulose ridged (Plate IV-c,d); floral bearing shoots prostrate (subgenus Reussia) :

flowers long lasting (more than 8 flowers per in­ florescence) ; leaves reniform, sagittate or cordate ...... A. P. rotundifolia

flowers ephemeral (usually fewer than 12 flowers per inflorescence); leaves subovate to sublanceolate ...... ,..5.P. subovata PLATE I

Inflorescence, Pendulous Ovule and Fruits

A* Young inflorescence with subtending spathe of Pontederia sagittata Presl from Puerto Barrios, Guatemala (Lowden 11), length 15 mm.

B. Pendulous ovule attached to upper portion of ovulary and long style of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21) , length of ovule 1 mm.

C. Utricle of Pontederia rotundifolia L. f. from Granada, Lago Nicaragua, Nicaragua (Lowden 17, 18), length 8.5 mm.

D. Utricle of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21) , length 8.5 mm.

8 9

4 PLATE II

Seeds, Endosperm and Embryo

A. Reniform seed of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21), length 3.5 mm.

B. Portion of copious mealy endosperm surrounding embryo of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21), length A mm.

C. Linear cylindrical embryo of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21), length 2 mm.

D. Ovoid seed of Pontederia rotundifolia L. f. from Granada, Lago Nicaragua, Nicaragua (Lowden 17, 18), length 3 mm.

10 11 PLATE III

Hardened Perigone Bases

A. Tooth-rldged hardened perigone base of Pontederia cordata L. var. cordata from Winous Point, western end of Sandusky Bay, Ohio, U.S. (Lowden 36, 37), length 8 mm.

B. Tooth-ridged hardened perigone base of Pontederia cordata L. var. cordata from 32 miles north of Belize City, British Honduras (Lowden 27), length 9.5 mm.

C* Tooth-ridged hardened perigone base of Pontederia cordata var. lancifolla (Muhl.) Torr. from vicinity of Colama, Cuba (Britton and Cowell 9693), length 6 mm.

D* Tooth-ridged hardened perigone base of Pontederia cordata L. var. oval is (Mart.) Solms from State of Minas Gerais, Brazil (Irwin 8904), length 6 mm.

12 13 PLATE IV

Hardened Perigone Bases

A. Smooth-ridged hardened perigone base of Pontederia sagittate Presl from Vera Cruz, Mexico (Lowden 6 ), length 8 mm.

B. Tooth-ridged hardened perigone base of Pontederia parviflora Alex, from between Pacora and Chepo, Panama (Lowden 21), length 8 mm.

C. Spinulose-ridged hardened perigone base of Pontederia rotund!foIla L. f. from Granada, Lago Nicaragua, Nicaragua (Lowden 17, 18), length 10 mm.

D. Spinulose-ridged hardened perigone base of Pontederia subovata (Seub. in Mart.) Lowden from Logoa Santa, Minas Gerais, Brazil (Smith 6702), length 10 mm.

14 15 Pontederia subg. Pontederia (Type of subg.: Pontederia cordata L . , Sp. PI. 288. 1753)

Hardened perigone bases toothed or smooth ridged (Plate III;

Plate IV-a, b); floral bearing shoots erect; main steins with short internodes; underground rhizomes long (mature plants).

1. Pontederia cordata L . , Sp. PI. 288. 1753.

Tristylous (Table 1); plant maximum height 9-13 dm; inflo­ rescence length 2.0-16.0 cm; perigone blue to white; hardened perigone bases toothed ridged (Plate Ill-a-d); anthers blue; peduncle length (inflorescence base to floral shoot leaf base)

5.0-33.0 cm; leaves cordate, sagittate (unauricled), ovate or lanceolate, leaf width 0.4-21.0 cm; spathe (terminal phyllode of floral shoot) length 2.0-7,4 cm; floral shoot leaf petiole (in­ cludes sheath base of petiole) length 4.0-31.0 cm; petiole length of other leaves 28.7-59.5 cm, maximum ligule length of petiole sheath 4.0-10.5 cm; sheath (axil of floral shoot) maximum length

22.3-28.0 cm; fruits and seeds reniform.

Comments, enclosed in brackets under the synonymy (nomencla­

ture) of each variety, indicate (unless otherwise stated) the part(s) of the taxon which is(are) thought, in accordance with the author's treatment of the new combination, to be synonymous with

the taxa here described. Those comments found in parentheses refer

to the type of a new taxon.

la. Pontederia cordata L. var. cordata, Sp. Pi. 288. 1753,

Umsema obtusifolia Raf., Med. Repos. II 5:352. 1808. (=P. cordata L. Type not seen.) 17 Urosema mucronata Raf., Med. Repos. II 5:352. 1808. (Raf. 1808* =P. cordata L. Habitat, Raf. 1837, cited ’’found by Mr. Kingston in Virginia 1800, seen in his herbal." Type not seen.)

Pontederia angustlfolla Pursh, Fl. Am. Sept. 224. 1814. ("P. foliis elongato-triangularibus sensim acutis basi truncato-subcordatis, corollae laclniis lineari- lanceolatis." "In high mountain-lakes: New York to Carolina." Holotype OXF?)

? Unlsema sagittate Raf., Fl. Ludov. 18. 1817. (Saglttaria? cerulea, "Flbchiere k feuilles en fleche, a fleurs en fepi. Rob. p. 340." Type not seen.)

Pontederia cordata L. var. angustlfolia (Pursh) Torrey, Fl. U.S. 1:343. 1824.

Unlsema deltlfolia Raf., Medical Flora 2:105, 107. 1830. (Habitat, Raf. 1830, "In west Kentucky, Tennessee, Alabama, &c." Raf. 1837, ". . . and Florida." Type not seen.)

Unlsema purshiana Raf., Medical Flora 2:107. 1830. (Habitat, "In the Southern States." =Pontcderla angustifolla Pursh. Type not seen.)

Unlsema media Raf., Medical Flora 2:107. 1830. (Habitat, Raf. 1830, "From New York to Carolina." Raf. 1837, ". . . called P. angustif. by many, ..." Holotype NY!)

Unisema media Raf. var. albiflora Raf., Medical Flora 2: 107. 1830.

Unisema media Raf. var. angustlfolla Raf., Medical Flora 2:107. 1830.

Unlsema obliquata Raf., Medical Flora 2:107. 1830* (Habitat, "In New Jersey and Virginia." Type not seen.)

Unlsema latlfolia Raf., Medical Flora 2:107. 1830. (Habitat, Raf. 1830, "very common, chiefly in the Southern States." Raf. 1837, "the most common kind from New Jersey to Florida, in marshes and streams, . . •" Type not seen.)

Unisema latifolia Raf. var. elatior Raf., Medical Flora 2:107. 1830.

Unlsema latifolia Raf. var. undulata Raf., Medical Flora 2:107. 1830. 18

Unlsema latifolia Raf. var. albiflora Raf., Medical Flora 2:107. 1830.

Unlsema latifolia Raf. var. pallida Raf., Medical Flora 2:107. 1830.

Unlsema acutlfolla Raf., Medical Flora 2:107. 1830. (Habitat, "Found by Bose in Carolina; I have never seen it." Type not seen.)

Unlsema heterophylla Raf., Medical Flora 2:108. 1830. (Habitat, Raf. 1830, "From New York to Louisiana." Raf. 1837, "... found at Trenton Falls of Delaware and on the R. Ohio, . . . , therefore the P. angustif. -—*• of some botanists." Type not seen.)

Unlsema heterophylla Raf. var. lanceolata Raf., Medical Flora 2:108. 1830.

Unlsema heterophylla Raf. var. stenocardia Raf., Medical Flora 2:108. 1830.

Unisema rotundlfolia Raf., Medical Flora 2:108. 1830. (Raf. 1830, "... perhaps a variety of the last [U, heteropity 11a Raf.3." Habitat, "In the Western States, rare, . . ." Raf. 1837, "Kentucky and Illinois." Type not seen.)

Pontederia cordata L. var. albiflora Short, Transylvania J. Med. Assoc. Sci. 32:3, 7. 1835. (Habitat, ". . .Mr. Griswold, pointed out this variety to us in a pond near Louisville." Marshes around Louisville, Sept. 1835, C. W. Short, Lexington, Kentucky, holotype PH 7838831, isotype NY1)

Pontederia caerulea Maund, Bot. Gard. 6:No. 551. 1836. (Habitat, plant under cultivation in Great Britain, introduced 1830 and cited as "From the unexpected loss of the specimen from which our drawing was taken, we are unable to determine whether or not it be more than a variety of cordata." Type not seen.)

Unisema peduncular!s Raf., New Fl. 2:75. 1837. (Habitat, "Carolina to Florida," cited as "purshlana? Raf. m. fl. (angustlfolla? Pursh)." Type not seen.)

Unlsema peduncularis Raf. var. parviflora Raf., New Fl. 2:75. 1837.

Pontederia nymphaelfolia Kunth, Enum. PI. 4:126. 1843. (Habitat, Brazil, collected by Sello 235, holotype in Museo Botanico Berolinensi, isotype PH! Photograph of type NY!) 19

Pontederia sagittata Seub. non Presl in Mart., Fl. Braa. 3:95. 184 7^ [name of Solms in DC. Monog. Phan. 4:533. 1863, believed to be Incorrectly attributed by Solms to Seubert, Instead of to Presl]

Pontederia cordata L. var. typlca, Solms in DC. Monog. Phan. 4:532. [that part synonymous with Pontederia angustlfolla Pursh, Pontederia cordata L. var. angustlfolla (Pursh) Torr.; Pontederia cordata L. forma brasillansis Solms in DC. and Pontederia sagittata Seub. in Mart, (name of Solms In DC.)3

Pontederia cordata L. forma angustlfolla (Pursh) Solms in DC., Monog. Phan. 4:532. 1883. [that part synonymous with Pontederia angustlfolla Pursh and Pontederia cordata L. var. angustlfolla (Pursh) Torr.]

Pontederia cordata L. forma braslllensls Solms in DC., Monog. Phan. 4:533. 1883. (Habitat, "In America meridional! Brasilia: Villa nova de Almeida, Princ. M. Wied; Minas Gerais (Clausen); Port Alegre (Isabelle); Buenos-Ayres (Bade); Paraguay (Bolansa, n. 607); Uruguay (Lorentz, n. 131)." Specimens not seen.)

Pontederia rotundlfolia L. f. var. nymphaelfolia (Kunth) Solms In DC. Monog. Phan. 4:534. 1883.

Pontederia cordata L. [var.] lanclfolla (Muhl.) Morong. Mem. Torrey Bot. Club 5:105. 1894. [that part synonymous with Umsema obtuslfolla Raf. and Umsema mucronata Raf.; Pontederia angustlfolla Pursh; and Pontederia cordata L. var. angustlfolla (Pursh) Torr.)

Narukila cordata (L.) Nieuwland, Amer. Midi. Naturalist 3:101. 1913.

Narukila cordata (L.) Nieuwland var. lanclfolla (Muhl.) Nieuwland, Amer. Midi. Naturalist 3:101. 1913. [that part of Pontederia cordata L. var. lanclfolla (Muhl.) Morong synonymous with Umsema obtuslfolla Raf. and Umsema mucronata Raf.; Pontederia angustlfolla Pursh; and Pontederia cordata L. var. angustifolla (Pursh) TorrTl

Pontederia cordata L. forma angustlfolla (Pursh) House, New York State Mus. Bull. 243-244:62. 1923.

Pontederia cordata L. forma latifolia (Raf.) House, New York State Mus. Bull. 243-244:62. 1923. 20

Pontederia cordata L. forma albiflora (Raf.) House, New York State Mus. Bull. 243-244:62. 1923.

Unisema cordata (L.) Farvell, Pap. Michigan Acad. Sci. 3:91. 1924.

Unisema cordata CL.) Farwell forma angustlfolia (Pursh) Farwell, Pap. Michigan Acad. Sci. 3:92. 1924.

Unlsema cordata (L.) Farwell forma latifolia Farwell, Pap. Michigan Acad. Sci. 3:92. 1924. (Habitat, New Jersey, Secancus, 13 Sept. 1890, H. II. Rusby, lectotype MICH1 Habitat, New Jersey, Franklin, Aug. 1873, H. H. Rusby, paratype MICH!)

Pontederia lanceolata Nutt, forma trullifolia Fernald, Rhodora 27:81. 1925. (Habitat, Florida, Okeechobee region, Brevard Co., 3 August 1903, A. Fredholm 5927, holotype GHl)

Pontederia lanceolata Nutt, forma brasiliensls (Solms) Fernald, Rhodora 27:81. 1925.

Unlsema lacifolja (Muhl.) Farwell forma trullifolia (Fernald) Farwell, Amer. Midi. Naturalist 11:73. 1928.

Pontederia cordata L. forma taenia Fassett, Rhodora 39: 274. 1937. (Habitat, Maine, Lincoln Co., Damariscotta Lake, Jefferson, shallow mucky cove, 28 August 1936, N. C. Fassett 16067, holotype WISI, isotypes GHI and MO 11498911)

Pontederia cordata L. forma bernardl Lepage, Naturallste Canad. 82:101. 1955. (Habitat, Canada, Quebec, Nominingue, zone tourbeuse autour du lac Violon, 12 aout 1951, Jean-Paul Bernard 386, holotype CAN 234514!)

Plant maximum height 11 dm (Figs. 2, 8); inflorescence (Figs.

1, 7) length 2.0-16.0 cm; perigone blue, purple or white; peduncle length (inflorescence base to floral shoot leaf base) 5.0-33.0 cm; leaves cordate (Fig. 3) or sagittate (Fig. 4, unauricled), leaf width

2.2-21.0 cm; spathe (Fig. 7, terminal phyllode of floral shoot) length

2.0-7.4 cm; floral shoot leaf petiole (includes sheath base of petiole) length 4.0-31.0 cm; petiole length of other leaves 28.7-59.5 cm, 21 maximum ligule length of petiole sheath 10*5 cm; sheath (Fig* 3, axil of floral shoot) maximum length 28.0 cm.

Type locality: United States, Virginia, communicated to Linnaeus by

GronovluB.

(Type not seen, the possibility of a type in the Linnaean

Herbarium is being investigated)

Distribution: Canada, United States, British Honduras and South

America (Figs. 5, 6, 11, 15). Fig. 1. Numerous blue-purple flowered in­ florescence, clasping subtending spathe and flowering stalk leaf of Pontederia cordata L. var. cordata, U.S., Ohio, Ottawa County, Winous Point, 3^ miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay (photographed, July 1966, R. M. Lowden).

Fig. 2. Pontederia cordata L. var. cordata in marsh habitat, U.S., Ohio, Ottawa County, Winous Point, miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay (photographed, July 1966, R. M. Lowden). Fig. 3. Fruiting specimen of Pontederia cordata L. var. cordata, showing cordate flowering stalk leaves and clasping sheath at base of flowering stalk stem shoot, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS).

23 21+

Pontenp'tii1* L 0*'ftwni v USnr'u* T i Trum ihlp, l ] /? 1 1 * 9 §w For* ClTr;- on «rr 9*v ^ F* \, C * r r *TF 1 *<• R * * '. iV Fig, 4. Fruiting and flowering specimen of Pontederia cordata L. var, cordata, showing sagittate- hastate leaves, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS).

25

I»*w f

( s

v * Fig. 5. Geographical distribution of Fontederla cordata L. var. cordata in Ontario, Quebec, New Brunswick, Prince Edward Island and Nova Scotia Provinces of Canada (map used, Department of Mines and Technical Surveys, Surveys and Mapping Branch, Canada, south of latitude 75 , The Map Distribution Office, Ottawa, Canada).

27

Fig. 6. Geographical distribution of Pontederia cordata L. var. cordata (dots), var. lancifolia (Muhl.) Torr. (triangles)! and var. ovalls (MartY)Solms (stars) in the United States (map used. Bureau of the Census, U.S. County Outline Map, 1 April I960, Department of Commerce, Superintendent of Documents, U.S. Government Printing Office, Washington 25, D.C.).

29 30 Fig* 7. Numerous white flowered inflorescence, clasping subtending spathe and flowering stalk leaf of Pontederia cor­ data 1.. var. cordata in British Honduras, C.A., 9 miles from Belize City, along the Northern Highway (photographed, December 1969, R. M. Lowden).

Fig- 8* Pontederia cordata L. var. cordata In drainage ditch habitat, British Honduras, C.A., 9 miles from Belize City, along the Northern Highway (photographed, December 1969, R. M. Lowden). 32

SPECIMENS EXAMINED*:

CANADA: PROVINCE OF NEW BRUNSWICK: 20 August 1876, shallow water,

Norton, Geo, N. Hay (CAN 29999), (no day or month! 1879, Fredericton, ex herb, J. Fowler (US 261092), 8 August 1927, Bhallow water of Third

Pond, Grand Manan, Charlotte County, C, A. and Una F, Weatherby 5769

(CAN 29998, GH). 20 July 1929, shores and woods around Lower & Upper

Chamcook Lakes near St. Andrews, M. 0. Malte (CAN 29997), 14 July

1955, Rothesay, near St. John, shallow pond, H. J. Scoggan 12440 (CAN

237932). 21 July 1955, shallow water at edge of lake 8 miles north of

St. Stephen, H. J. Scoggan 12589 (CAN 237934). 16 August 1956, Grand

Lake, east of Fredericton, shallow water, H. J. Scoggan 13455 (CAN

243854). PROVINCE OF NOVA SCOTIA: ANNAPOLIS COUNTY— +19 July 1921, muddy or wet sandy margin of Lamb's Lake, M. L. Fernald, E. B. Bartram,

B. Long, and N. C. Fassett 23578 (GH). +10 August 1955, edge of runnel among typical variety, west of Lequille River, E. C. Smith,

W. B. Schofield, J. C. Taylor, D. H. Webster, L. B. Sllpp, and P. A .

Bentley 11871 (CAN 235789). COLCHESTER COUNTY— 29 July 1954, common

in marginal water of Earltown Lake, E. C. Smith, D. H. Webster, and

P. A. Bentley 11742 (CAN 250168). DIGBY CO.— 9 August 1921, shallow rocky and peaty cove at head of Wentworth Lake, M. L. Fernald and

B. Long 23579 (GH, PH 587759). GUYSBOROUGH COUNTY— +5 Sept. 1925,

Horton's Pond, West Cook's Cove, H. G. Perry, R. H. Wetmore, G. C .

Hicks, and A. R. Prince 10013 (GH). HALIFAX COUNTY— +6 August 1927,

sandy lake bottom, Quoddy, A. R. Prince and C. E. Atwood 508 (WIS).

A plus sign (4) before a citation indicates there is no dot for that specimen(s) represented on distribution maps. PICTOU COUNTY— +25 August 1906, Millstream, C. B. Robinson 509 (NY).

30 July 1913, Valley of the , margin of Taylor's Lake, Sunny

Baae CSunnybrae], Harold St. John 1404 (GH). SHELBURNE COUNTY— 9 Sept.

1921, peaty shore, Lake John, M. L. Fernald and B. Long 23581 (GH, PH

587760). 20 August 1928, shallow water, Clyde River, A. R. Prince and

C. E. Atwood 1323 (WIS). YARMOUTH COUNTY— +27 July 1920, shallow water at sandy margin of Clearwater Lake, Belleville, B. Long and

D. H. Linder 20624 (GH). +4 August 1920, shallow water at rocky margin of Goose Lake, Argyle, M. L. Fernald and P. White 20625 (CAN

45015, Gil, PH 589644, US 1104233). 13 August 1921, shallow water,

Cobbly border of Vaughan (Tusket) Lake, Gavelton, M. L. Fernald and

B. Long 23580 (GH, PH 587962). CAPE BRETON ISLAND— 19 August 1898,

in a creek, Louisburg, Macoun (CAN 29996). 21-25 July 1901, edge of

pond, marshy soil, North Sydney, Cape Breton, C. D. Howe and W. F.

Lang 649 (GH, NY). NOVA SCOTIA Y— +27 July 1910, Italy Cross, N. S.,

John Macoun (CAN 29995). PROVINCE OF ONTARIO: [no date], Lake

Timagami, Northern Ontario, Edgar E. and Dorothy Anderson (MO 1003519).

21 June 1866, Clinton, Lincoln Co., C. A. Gross Herbarium 2988 (PH

586496). 31 July 1874, in shallow water, near Belleville, Macoun

(CAN 45010). 26 July 1877, Toronto, Macoun 1905 (GH). [no day or month) 1879, Sarnia Bay, Dodge (MO 1983377). +21 August 1884, in water,

Carrelman, Macoun (CAN 116593). 15 Sept. 1884, in water, St. Clair

Flats, J. M. Macoun (CAN 45016). 13 August 1885, Kingston, Round Lake, ex coll. J. Fowler (MO 1983331). 1 Sept. 1889, Lake Muskoka, Port

Sandfield, Dr. and Mrs. N. C. Britton and Miss Millie Timmerman (NY).

10 July 1892, ponds and lake borders, Muskoka, Yneadborough [?] (CAN 45009). 17 July 1892, In a pond at Grimsley CGrimsby], only one patch of It seen, Armstrong (US 33953). 9 August 1894, Rideau River, John

Macoun (MO 1983332, US 231992 and 232004). +12 July 1897, pond at mouth of "The Eight", Lincoln Co., W. C. McCalla 548 (CAN 45008).

22 July 1897, near Toronto, Blltmore Herbarium (GH). 11 July 1898, near Toronto, Blltmore Herbarium (US 980486). 13 July 1898, Smith's

Falls, Kingston, ex coll. J. Fowler (F 83713). +22 July 1899, Lake,

Bomie Chese Mts., L. M . Umbach (F 86661). +31 July 1900, in Cache

Lake, flora of Algonquin Park, Macoun (CAN 45012). 20 August 1900, in the discharge of Whites Lake, flora of Algonquin Park, Macoun (CAN

45011). 10 July 1901, Niagara Falls, John Macoun (CAN 45013). 23 July

1902, Sarnia, Herb, of G. L. Fisher (US 481790). 31 July 1902, Plevna, ex coll. J. Fowler (GH). 22 August 1905, Rideau River, Huramans Bridge,

W. H. Harrington 2355 (CAN 116590). 7 July 1911, Cumming's Bridge,

Ottawa, John Macoun (CAN 116592). 8 August 1922, shallow water,

Temagami Forest Reserve, C. E. Hindson 431 (CAN 45006). 28 July 1923, shallow muddy creeks, flora of Temagami Forest Reserve, Ingall Lake,

W. R. Watson 1041 (Gil). 2 August 1928, Beausoleil Isl. Georgia B.,

H. Lloyd (CAN 45005). +20 August 1928, Knophle bog, Chikopi, Cedric

L. Porter 235 (MICH). 10 Sept. 1932, plants of the islands and shore of northern Lake Huron, in one foot of water of Rock Lake near 45

56* N, 80° 54' W, French River Harbor, Northwest part of Parry Sound,

Carl 0. Grassl 2301 (MICH). 12 Sept. 1932, plants of the islands and shore of northern Lake Huron, 45° 56' N, 80° 55' W, waters edge, French

River mouth, Walter Koelz 4810 (MICH). 31 August 1933, Flora of Bruce

Peninsula, Stokes Bay, F. V. Krotkov 7230 (GH). 22 July 1934, flora of 35

Bruce Peninsula, marsh, Stokes Bay, P. V. Krotkov 8825 (NY, WIS).

4-11 August 1935, west of White Fish, margin of White Fish Lake,

E. C. and T. G. Yuncker 5444 (F 800225, FSU 43682). +5 Sept. 1935,

in muck near shore, Carp Lake, flora of Algoma District, Lat. 47° 00’ N,

Long. 84° 45' W, T. M« C. Taylor, R. C. Hosle, R. E. Fitzpatrick, S. T .

Losee, and A. Leslie 2368 (CAN 45004, GH). 13 August 1936, plants of

the region between Lake Superior and Lake Nlplsslng, bank of Vermilion

River near Whitefish, 46° 24' N Lat., 81° 14' W Long., Carl 0. Grassl

7494 (MICH). 16 August 1936, plants of the region between Lake Superior

and Lake Nipissing, frequent clumps in 1 to 2 feet of water in Blind

River, 46° 11' N Lat., 82° 59' W Long., Carl 0. Grassl 7493 (MICH).

19 Sept. 1936, plants of the region between Lake Superior and Lake

Nipissing, with Bricaulon in 3 to 10 inches of water in Clear Lake near

Novar, 45° 28' N Lat., 79° 15' W Long., Carl 0. Grassl 7495 (MICH).

18 July 1938, Hurdman Bridge, riviere Rideau, fls. bleues, Adrien

Dubois 543 (NY). +24 August 1938, In water N.E. Shore, Lang [?] Lake

Erie, H. A. Senn and J. H. Soper 511 (NY). +18 July 1942, flora of

the Georgian Bay Islands opposite Shawanaga Township, Parry Sound

District, off 500A (govt. no. of island), Hemloch Island, 3 feet of water near E9, E. D. McDonald, Jr. 152 (CAN 196950, US 1924507). 2/7

1946, Chalk River, Dominion Forest Exp. Station, in the river near stat.,

I. Hustlch (CAN 151941). +25 July 1948, Kent County, in 3 feet of water along marshy shore, Lot 1, conc. VIII, Dover Twp. St. Luke's Bay,

L. St. Clair, common, J. H. Soper and H. M. Dale 4068A (GH, US 2066603) and 4068B (GH, M0 1635054, US 2066604). 10 August 1950, growing in 3-4 feet of water, Dover Tp., Kent Co., #24-St. Patricks Cove, ca. 1 mile S of Mitchell Bay, J. K. Shields 276 (CAN 228347, MO 1648213). 23 July 36

1952, Frontenac County, Loughborough Township, Upper Rock Lake, 7 miles east of Perth Road, rocky lakeshore in one foot of water, ex­ tensive patches, flowers blue, J. M. Glllett 6579 (NY). 6-8 August

1952, flora of Georgian Bay, Byng Inlet, Hopewell Bay, common in water,

A. E. Porsild 18547 (CAN 271915). 25 July 1956, Nipissing District, in shallow water at edge of lake, north shore of L. Nipissing 2 miles south of Sturgeon Falls (Loc. 472), J. H. Soper and G. Fleischmann

6593 (CAN 257907, NO 1730458). +13 August 1959, Haliburton County, open swamp near lake edge (Loc. 6), point on west shore of Paudash

Lake % mile N of Ross Is., C. E. Heidenreich 333 (CAN 299123). 5 Sept.

1960, Frontenac County, in one foot of water, Black Lake Prov. Park, just S of Hy #7, between Maberley and Arden, M. Dutnais (MICH). 8 Sept.

1964, Muskoka District, B-grows at the edge of lake, leaves and in­ florescence aerial, rooting in mud, Margaret Lake, 8 mi SSE of Dorset,

T. W. Chow 648 (CAN 300508). 14/8 1966, Long Lake, Waters Twp., Sud­ bury District, shallow water, K. Winterhalder KW 66191 (CAN 323222).

15 August 1968, Muskoka County, 45° 54* N, 79° 20* W, in one foot of water with mucky soil, Gravenhurst, Silver narrows near Gull Lake ca.

2 miles E of Gravenhurst, S. J. Ives and J. McCarron 101 (CAN 319511).

PELEE ISLAND, ESSEX CO.— 17 August 1967, common in shallow water of

North Lagoon, north end of Pelee Island, Ronald L. Stuckey 5161 (OS).

July 1968, growing in North Lagoon in about 2 ft of water, R. Kalinsky

209 (FTSL). 11 July 1968, locally common in shallow water of North

Lagoon, NW corner of Pelee Island, Ronald L. Stuckey 7065 (CAN 320716,

OS). GEORGIAN BAY AREA— 20 August 1904, shallow water, loc. Go Home,

O.G.U. [?] (CAN 45007). PROVINCE OF PRINCE EDWARD ISLAND: QUEENS 37

C0UNTY--+9 August 1912, shallow water, sandy margin of Lake Verde,

M. L. Fernald, Bayard Long, and Harold St. John 7131 (CAN 45014, GH,

NY, PH 738854, US 1202564). PROVINCE OF QUEBEC: 24 July 1903, by

Bell’s Lake, near Wakefield, Macoun (CAN 116589). 30 August 1905, in pools of water, Ste. Anne de Beaufue [Beaupre], Macoun (CAN 45003).

+25 July 1908, shallow water, river margins, brooks, etc., Fairy Lake,

W. H. Harrington 2355M (CAN 116591). 25 August 1922, Templeton, M. 0 .

Malte (CAN 232019). 10-11 August 1923, swampy shore of Lake Champlain,

PhiltlDipsburg, Missisquoi County, Clarence H. Knowlton (GH). 31 July

1924, St. Cesaire, Bro. Majoni [73 (CAN 45002). 29 Sept. 1925, Saint-

Jerome, lac a la Barbotte, Plantes des Laurentides, F. Maric-Victorin

20316 (F 568569, MO 944284). 19 August 1926, Magog, riviere,

J. Rousseau 24223 (GH). 13 August 1929, Nominingue, ccmte de Labelle,

Fr. Edmond Roy, c.s.v. 1363 (CAN 229412, US 2096414, WIS). 15 July

1930, Nicolet, comte de Nicolet, marais, F. Regis-Stanislas 192 (NY).

August and Sept. 1930, Mascouche et Lachenaie, comte de L'Assomption,

P. Louis-Marie et Alii 1420 (CAN 45001). 25 August 1934, lie Sainte-

Therese, Pres de Saint-Jean, Vallee du Richelieu, Fr. Rolland-Germaln

43521 (GH). 21 August 1943, Baie de la Valliere, comte de Richelieu, le long du ruisseau Pot-au-Beurre, F. Marle-Victorin, F. Rolland-

Germain, and Ernest Rouleau 2296 (CAN 229411, NY). 25/7 1946, Granby,

Bord de riviere, F. Fabius 403 (CAN 207004), +2 August 1947, Saint-

Adolphe, comte d ’Argenteuil, anses du lac, Frerc Rolland-Germaln 401

(FSU 46361). +7 August 1952, St.-Adolphe, comte d ’Argenteuil, dans

1 m. d ’eau, lac Ste-Marie, Frere Rolland-Germain 7602 (CAN 265885, FSU

70952, US 2421634). 30 July 1965, Lac Memphremagog, comte de 38

Stanstead, Bord boueux du lac, extremite Nord, cote Ouest, Philippe

Forest 290 (CAN 322890). 13 August 1968, Lac Vaseux, Weedon, canton de

Weedon, comte de Wolfe, bord marecageux du lac (tres abondant), Claude

Hamel et Samuel Brlsson 15210 (CAN 321148).

* UNITED STATES: ALABAMA: BALDWIN CO.— 6/12 [19]09, Magnolia Springe, borders of ponds and streams, showy native aquatic plant, hardy per. h . ,

1-4 ft, flowers 4-5M, In dense long spikes, June-Oct., l-seeded utricle, leaves ovate cordate-sagitate, root-horizontal rootstock, P. 0 .

Schallert 757 (DUKE). 30 July 1947, entrance to Bay, Minette Basin,

Mobile Delta, emergent, common in low marsh, Francis X. Lueth L70

(DUKE). 15 July 1970, 4 miles east of Mobile, Battle Ship Parkway,

Routes 31-90-98, Mobile Bay, Mobile Bay Causeway, brackish water,

R. M. Lowden 5 (OS-4). MOBILE CO.— [probably Mobile Co.— +[no date],

Estate of Adams Jewett (MICH). 10 May 1842, Mobile, Estate of Adams

Jewett (MICH-3). 4 July 186 7, margins of ponds and ditches, Mobile,

Car. Mohr (MICH). 13 May 1878, Mobile, swamps, shallow water, C. Mohr

(US 784455). March 1879, Mobile (OS). CO. ?— +16 May 1896, Hanvana

Glen, L. M. Underwood (NY). +August, 1919, Bay shore, or river front,

Magazine Pt., E. W. Graves 653 (MO 977690). ARKANSAS: CRAIGHEAD CO.—

6 August 1929, Lake City, near water, very rare, D. Demaree 7075 (US

1438477); Lake City, St. Francis River, D. Demaree 7076 (F 722544, GH).

CO. 7--+21 June 1920, Big Luke, Hornersville, F. P. Metcalf 626 (US

1134709). CONNECTICUT: FAIRFIELD CO.— 4 Sept. 1916, Green Pond, town of Sherman, Francis W. Pennell 8592 (PH 732440). 16 August 1962,

Trumbull, sand and alluvium, strand of Poquonock R. with Scirpus

Purshlanus F., Frank C. Seymour 20245 (NEBC). HARTFORD CO.— h3 August 39

1882, sand-bar of Connecticut River, Hartford, C. Wright (GH). 28 June

1889, Bristol, water, John Hendley Barnhart 317 (NY 2535). +7 August

1892, Glastonbury, Frances Wilson (NEBC). 31 July 1898, Southington, swampy ground, C. H. Blssell 614 (NEBC). 3 August 1898, Southington, ditches and also streams, L. Andrews 685 (GH). 8/10 1899, town of

Southington, in shallow water, C. H. Blssell 420 (NY). +15 August

1902, Boardman's Pond, East Hartford, C. A. Weatherby 145 (NEBC).

LITCHFIELD CO.— July 1891, West Goshen, Ct., L. M. Underwood 3l25a (NY) and 3125 (NY). MIDDLESEX CO.— 20 August 1892, waterhole in meadow,

Portland, Francis Wilson (NEBC). NEW HAVEN CO.— +31 July 1887, Oxford,

E. B. Harger 2825 (PH 561260). +15 August 1896, plants of Tranquillity

Farm, Mlddlebury, W. M. Shepardson (NEBC). 8-9-(19358, marshy meadow bordering Mill River, North Haven, aquatic herb, M. R. North 105 (NY).

CO. ?— +[no date!, Charles Wright (MO 1983336). 15 August 1873, Rock­ ville, Herb. F. N. Pease (NY). +18 July 1883, Easton, pond at Perry's

Mill, Herbarium of L. N. Johnson 2988 (F 1430029). +10 July 1884,

Greenville, G. R. Lumsden 44248 (CAN 165676). +18 July 1921, Wash­ ington, borders of pond, A. W. Evans (MO 935313). +30 June 1941,

Mt. Carmel Pond, W. T. Edmondson (WIS). DELAWARE; NEW CASTLE CO.—

6 July 1843, Wilmington, marsh, Rev. J. H. Brakely (?). July 1860,

Wilmington, ponds and ditches, common, Herb. Wm. M. Canby (NY). 23 July

1890, Wilmington, ditches, Herb. A. Commons (PH 540654). +14 August

1908, Smyrna, N. in Smyrna River, Bayard Long (PH). +3 August 1916,

Augustine Beach, mud, Francis W. Fennell 7785 (NY). 8-2 1923, in water, along Delaware River, south of New Castle, plants of Chesapeake

Bay Region, Ivar Tldestrom 11574 (GH). +19 July 1925, New Castle, tide swamp, R. R. Drelsbach 3576 (PH). + 29 August 1925, Christiana, pond,

R. R. Dreisbach 4044 (PH). +1 July 1934, marsh near road, 1.25 miles

W of Delaware City, Esther L. Larsen 573 (PH). +18 July 1934, Delaware

City, coal wharf, Esther L. Larsen 643 (GH). +20 July 1934, NE side of

Pea Patch Island, Esther L. Larsen 671 (PH). SUSSEX CO.— +5-6 July

1908, Milton Creek, N. L. Britton 76 (NY). +21 July 1941, Blades, swampy field near Nanticoke River, Edward L. Owens (PH 799903),

25 June 1942, muddy bank, outlet of Burton's Pond, about 5 miles SW of Rehoboth Beach, Rogers McVaugh 6532 (F 1411200, GH, NY 2380).

CO. 7— +28 June 1898, Hollyoak, tidal shore of Delaware River, Albert

Commons (PH 540655). +13 August 1911, Drawyers Creek, Delaware, J. R .

Churchill (GH, M0 105376 7). DISTRICT OF COLUMBIA: [no date], near

Washington, Herbarium of Wm. H. Seaman (US 787756). 8-11 1846,

Georgetown, Detwiller Herbarium (PH). 19 June 1873 flowers, 27 July

1877 fruit, in vicinis Washington, Lester F. Ward (MO 773740). 21 June

1885 [?], Washington, Schott (F 44042). 12 Oct. 1887, near Analostan,

Washington, E. S. Burgess (NY). 28 June 1891, Bank of Potomac,

Washington, Herb. Ferdinand Blanchard (MO 749785 and 1983355). 19 July

1893, Anacostia, F. L. J. Boettcher 161 (MO 1983354). 9 August 1897,

C. & 0. Canal, above Georgetown, Thos. H. Kearney, Jr. (FSU 17918, OS).

30 Sept. 1897, Pot. flats D.C., A. J. Pieters (MICH). FLORIDA:

ALACHUA CO.— 17 May 1940, In shallow water, margin of lake, 2 miles S of Gainesville, flowers blue-purple, Robert F. Martin 1457 (DUKE 112823).

CALHOUN CO.— 6 June 1956, abundant in a small stream flowing into

Chlpola River, about 200 yards north of US 20, Paul L. Redfearn, Jr .

2205 (FSU 32552, GH). CITRUS CO.— 12 July 1958, marshy borders of Tso[a]la Apopka Lake, R. K. Godfrey 57183 (FSU 42084). COLLIFR CO.—

5 April 1952, Deep Lake, 75 miles from Miami, Henry Field and Yusuf

Lazar (F 1577348, US 2097116). COLUMBIA CO.— +21 July 1893, shallow pond water, Lake City, Alligator Lake, A. L. Quaintance (F 228621).

+June-July 1898, Lake City, A. S. Hitchcock (F 234452). +20 May 1964, marsh in drainage canal bordering swamp forest, N outskirts of Lake city, R. K. Godfrey 63760 (FSU 87274). DADE CO.— +1 Jan. 1916, Praires about Royal Palm Hammock, John K. Small 7242 (NY, US 1738173). DIXIE

CO.— 3-V-1958, along Florida 349, 3 miles S of US 19, W side of road, alt. 25 feet, edge of roadside pond, ca. 50 yards across, 2 inches water over highly organic, sandy muck, cypcraceous and juncaceous species, Plnus sp,, Nyssa sp., Quercus sp., Myrica, ring the pond, perennial herb, ca. 30-40 inches, upright in large clumps, total stand

20 by 3 yards, violet-blue coralios w. yellow mark on upper perianth part, roots pinkish, Alfred Traverse 723 (GH). DUVAL CO.— May [no day or year], tidal creeks, NE Florida, A. H. Curtiss 2988 (BKL, GH, NY-2,

US 937962). 1893, St. Johns River, A. Fredholm (US 214864). 2 July

1894, near Jacksonville, A. H. Curtiss 4941 (F 147405, MO 1983361, US

224475). July-August 1900, Jacksonville, A. S. Hitchcock (F 231897).

26 May 1902, River swamp, A. Fredholm 5237 (GH). FLAGLER CO.— 26 July

1961, edges of cypress-gum swamp, 3 miles W of Bunnell, this number,

RKG 61162, each portion of plant from a separate stem, gathered more or less randomly in '’population", R. K. Godfrey and Grady W. Relnert

61162 (FSU 66 748-6 6 754). FRANKLIN CO. — 1898 [?], Apalachicola, A. W .

Chapman (MO 760556 and 1983411). 30 Oct. 1910, St. Vincent Island,

W. L. McAtee 1737 (US 514747). 29 June 1956, common on wet sandy- 42

clay-alluvium of Apalachicola River bank, or in shallow standing water,

NW of Apalachicola and 1-3 miles N of RR bridge, R. Krai and P. L .

Redfearn 2734 (FSU 29154). 2 April 1960, edge of cypress swamp, % mile

W of Palmdale and US 27, 6 miles S of Highlands-Glades County line,

S4, T41S, R30E, flowers purple-blue, D. B. Ward 1-6 with BTY 421 (FSU

94955). HENDRY CO.— 28 III 1959, Hendry County, in water, flowers

purple, roadside canal, road between Big Cypress Reservation and

Clewiston, seminole use— useless, mikasukl seminole name - hicakna: blo:ci, William C. Sturtevant 12 (US 2524641). HILLSBOROUGH CO.—

5/21 1900, Tampa, Herb. S. M. Tracy (NY). 11 April 1923, Tampa,

Hillsboro County, Hillsboro River, J. R. Churchill (MO 1049448).

24 August 1960, SE of Tampa, one mile N of US 301 & Fla. 674 inter­

section, wet grassy border of Cypress pond with Myrica, Magnolia virginiana, Persea Borbonia, Smilax, and pond cypress, Olga Lakela,

Jackie Fatman, and James P. Ray, Jr. 10063 (GH). +19 Nov. 1964,

swamp forest, 8 miles S of Zephyr Hills, R. K. Godfrey 65144 (FSU

91244). JACKSON CO.— +19 May 1960, open marshy area bordering cypress- gum pond, 1 3/4 miles E of Grand Ridge, pieces of plant on separate

sheets, my number 59541, collected from separate stalks more or less at random in population11, what is a plant and what is a clone was not determined, R. K. Godfrey 59541 (FSU 56388-56399, 56402-56405). 21 Sept.

1963, 2 miles S of Alford on US 231, modified cypress-sawgrass habitat, exposed, Plants 1-8 (two 8's), 9-17 (two 17’s), 19, R. Christensen and

Rex Kerstetter 5 (FSU 121442-121458). +26 April 1967, Hays Spring Run, ca. 5.5 miles W of Greenwood, S32, T5N, R10W, Taxodium-Nyssa anuatlca-

Fraxlnus tomentosa, local colony in overflow area, flowers purple, Sidney McDaniel 8826 (FSU 1X3730). JEFFERSON CO.— +Jan.-Mar. 1890,

head of Wacissa River, Rev. L. H. Llghthlpe (BKL). +15 April 1956,

rare in shallow still water or muddy banks, headwaters of Wacissa

River, R. Krai and J. P. Gillespie 2289 (FSU 35558). 25 Sept. 1963,

S end of hake Miccosukee at US 90 bridge, floating clones in partial

shade, Plants 1-13 (two 11’s), Cone without a Plant no.3, Christensen,

Kerstetter, and McDaniel 7a (FSU 121541-121553)• 25 Sept. 1963, S end of Lake Miccosukee at US 90 bridge, floating clones in exposed habitat,

Plants 1-3 (two 3'a), 5 (two 5’s)-8» Christensen, Kerstetter and

McDaniels 7B (FSU 121554-121561). 2-25 1967, abundant in ditches along

logging road in swamp, 3 miles east SR 59 and US 98, Jean Wooten 1072

(FSU 110044). LAFAYETTE CO.— 13 June 1956, common on muck of bottomland bay, 8 miles SSW of Mayo, R. Krai and P. L. Redfearn 2645 (FSU 26959).

LAKE CO.— 16-30 April 1894, collected in vicinity of Eustis, margins of lakes, Geo. V. Nash 449 (GH, MICH, MO 1983409, NY, US 227931 and

937956). LEE CO.— July-August 1900, Myers, swamps, A. S. Hitchcock 354

(F 101421, GH, MO 1983358, NY, US 387386). 1 March 1916, vicinity of

Fort Myers, edge of Whisky Creek, Paul C. Standley 12959 (US 896396).

29 March 1916, vicinity of Fort Myers, edges of lakes, Jeanette P .

Standley 104 (F 457297, GH, MO 800741, NY, US 569560). March 1938, pond in cypress swamp, near Hendry Creek, 10 miles S of Fort Myers,

Paul C. Standley 576 78 (F 918741). LEON C0.--+Summer, 1895, near

Tallahassee, N. K. Berg (F 159955 and 159956, NY). +15 May 1956, abundant in marshes along the shore of Lake Jackson, R. K. Godfrey

54830 (FSU 26817, GH), +8 June 1956, very abundant in a small lake just N of Lake Munson, Paul L. Redfearn, Jr. 2262 (DUKE 136296, GH, 44

NY). 29 Sept. 1958, In shallow water at upper end of Lake Miccosukee,

Richard S. Mitchell (FSU 44692). 17 May 1960, abundant in shallow stream, open area, 3.5 miles W of Tallahassee, each piece of plant on other sheets, my number 59518, was collected from a separate stalk gathering them more or less randomly in one "population", thought what a plant is here I know not, for whatever this is worth if any­ thing, R. K. Godfrey 59518 (FSU 56365-56387), +18 Sept. 1963,

Tallahassee, pond on T.S. and E. Road, 1 mile E of S. Monroe St., large floating clones in exposed habitat, plants growing here are large and robust, the vast majority of the plants are floating and only those next to the shore are attached, all of the plants seen here had cordate leaves, the habitat is fully exposed and several plants were in flower, there seems to be a great deal of asexual reproduction here by rhizomes as the plants are found in large clumps, Plants 1-4, 7-23,

25, R. Christensen and Rex Kerstetter 2 (FSU 121483-121498). LEVY-

CITRUS COS.— 21 May 1948, Withlacoochee River, below Yankeetown, growing in water, Carl and Jeanne Janish 351 (MO 1734721). LEVY CO.— 4 June

1958, in about one foot of water, cypress pond, 1.5 miles W of Bronson,

R. K. Godfrey and William Lindsey 56987 (FSU 39831, GH). 8 June 1961, marshy shores of Long Pond, N of Chiefland, each piece of leaf from a different stalk collected more or less randomly in "population", my number 60879, R. K. Godfrey and Grady W. Reinert 60879 (FSU 65104-

65119). 10 May 1963, floodplain woodland, swamp, along the Suwannee

River at Manatee Springs, R. K. Godfrey and Richard D. Houk 62824 (FSU

79660-79661, 79665). MADISON CO.— +[no date3, standing water of road­ side ditch and adjacent cypress pond, colony appearing rather uniform, bumblebees seen visiting flowers, ca, 10 miles NE of Monticello on

Fla. 146, William P. Adams 175 (FSU 48852). +June-July 1898, A. S.

Hitchcock (MO 1983359). 1 Oct. 1955, in shallow water, sandy-peaty bottom, Mystic Lake, near Madison, R. K. Godfrey, R. Krai, and J. P .

Gillespie 53979 (FSU 15566, GH). 8 April 1961, flowers bluish-lavender, abundant in shallow water of muddy pond, ca. 4 miles S of Madison, S12,

T1S, R9E, Barbara Moore 21 (FSU 94951, US 2604328). 14 June 1961, found on the border of a pond that was drying up, 4 miles W of Green­ ville on Hwy. 90, Gwynn W. Ramsey and Richard Mitchell 54 (FSU 67875).

NASSAU CO.— 2 June 1958, in marshy borders of stream, drainage canal,

Fernandina, R. K. Godfrey and William Lindsey 56829 (DUKE 140593, FSU

40334, GH). 28 July 1961, marshy borders of swamp along the St. Marys

River, NW of Yulee, each portion of plant of this gathering, my number

61227, was taken from a separate stalk more or less randomly scattered about the "population1', I know not what is a plant or a clone, R. K .

Godfrey and Grady W. Reinert 61227 (FSU 66891, 66900-66903, 66909-66917).

23 May 1964, marshy clearing of swamp, vicinity of O'Neil, R. K .

Godfrey 64115 (FSU 31524). OKALOOSA CO.— 25 June 1964, swamp astride

Rocky Bayou, ca. 2 miles E of Niceville, R. K. Godfrey 64356 (FSU

89842). OSCEOLA CO.— 5 April 1936, pond, Loughman-Campbell's Highway,

Mary L. Singeltary 58 (DUKE 46219). PINELLAS CO.— 28 Oct. 1907, in fresh water near St. Petersbury, Mrs. Chas. C. Deam 2895 (BKL-2, NY).

26 April 1908, edge of pond, near St. Petersburg, Mrs. Chas. C. Deam

5024 (BKL, F 243606, US 513474). 18 April 1926, creek side Gulfport,

St. Petersburg, Mary H. Williams (MO 1072997, PH). +21-28 Dec. 1949, pond margin, Pasadena, Robert F. Thorne 9587 (GH). POLK CO.— 29 March

1925, Lake Alfred, E. M. Gilbert (WIS). +6/12 1931, Bartow Swamp, James B . McFarlin 5768 (MICH). +5/9 1931, Kossuthville, James B .

McFarlln 5179 (MICH). PUTNAM CO.— +28 April-3 May 1897, shallow water

Johnson, John Hendley Barnhart 2093 (NY 2535). ST. JOHNS CO.--22 May

1964, swamp 4.5 miles W of St. Augustine, R. K. Godfrey 63986 (FSU

37057). SANTA ROSA CO.— 27 June 1941, sandy shore of Escambia Bay,

Pensacola, R. BrInker 142 (MO 1225256). 18 August 1941, growing in water along edge of stream flowing through swamp at Chimney Point on

Hwy. 90, at Escambia Bay, R. BrInker 48 7 (MO 1225291). SARASOTA CO.—

25 March 1943, Sarasota, Anne E. Perkins (GH). SEMINOLE CO.--2 May

1960, in small stream near Wagner, Richard J. Eaton, James P. Ray, Jr. and George R. Cooley 7511 (GH). SUMTER CO.— 1-17 March 1879, Esperanza

John Donnell Smith (US 937964). 13 June 1958, one-half mile west of

Tarrytown, abundant on peat-muck of cypress hardwood swamp, flowers deep blue, Robert Krai 6857 (FSU 44929, GH, NY). TAYLOR CO.— 13 June

1956, shallow water and muck of pond border, 10 miles E of Perry,

R. Krai and P. L. Redfearn 2582 (FSU 31494, GH, NY). +13 June 1956, common on muck soil of cypress swamp 4 miles S of Salem, R. Krai and

P. L. Redfearn 2697 (FSU 27271, GH, NY). VOLUSIA CO.--+21 May 1885,

Hardwood Grove, G. C. Whitlock (GH). 2 May 1895, Port-Orange, F. C .

Straub 134 (GH). 5-16 1959, Highland Park Dead River, NW of DeLand,

Elmer C. Prichard 1170 (GH). 5-27 1964, margins of pools, Daytonia

Beach, Delzie Demaree 50160 (FSU 93463). WAKULLA CO.— +12 August 1957 in ca. 15 Inches of water, drainage canal, St. Marks Wildlife Refuge, near Oak Island, R. K. Godfrey 55894 (FSU 35106). 29 May 1960, open pools in gum swamp, 1.8 miles E of Newport, each piece of plant on separate sheets, my number 59565, was from a separate stalk the gathering being more or less random in the "population", I know not what a plant may be, or a clone, for whatever' value this may be, if any,

R. K. Godfrey 59565 (FSU 57095-57120). +5-17 1966, S.R. 375, 5 miles

W Panacea, Jean L. Wooten 287 (FSU 107366). FLORIDA: CO. ?--+Cno date), Southern Flora, Herb. Chapman (NY, US 964062). + [no date),

Chapman (NY). +Cno date), Delzie Demaree 10257 (MO 1047381). + [no date), F. Cozzens (NY). + [no day or month), F. Rugcl (F 119381, MO

1983408 and 1983410, US 512724). +July-Sept. [no year), miry margins of ponds and rivers, distribution of the duplicates of the Chapman

Herbarium 752e (GH, NY, US 335049). +3 April 1892, B. F. Leeds (F

403969). +26 Dec. 1895-11 Jan. 1896, New River, A. S. Hitchcock (F

233684). +[no day or month) 1897, from Chapman (MO 1983412). +12 May

1916, Lake Jamonia, W. D. Pierce (US 424676). +12 March 1930, in swamp, Wekiwa Springs, F. S. Blanton 6536 (NY). GEORGIA: CAMDEN CO.--

+12 May 1930, in wet ditch, Colesburg, plants about 2 feet tall, flowers blue, Harold N. Moldenke 1163 (DUKE 09720, MO 1000716, NY). CHATHAM

C0.--+May 1826 [?), common about Savannah in bogs and ditches (PH).

+16 June 1923, Savannah, Herbarium of Lee Sowden (PH 839935). LONG

CO.— 10 June 1950, about 30 inches tall, corolla light purple, at edge of pool of water in swamp woods in Altamaha River Swamp, 4% miles SW of

Ludowici, Wilbur H. Duncan 11089 (GH). McINTOSH CO.— 26 May 1909,

Darien, Swamp, Huron H. Smith 2095 (F 247744, WIS). 19 Sept. 1956, corolla blue, the upper lobe with a yellow area ringed with white, leaves, except one, basal, two-ranked at right angles to horizontal, fleshy rhizome, south-central part of Sapelo Island, Wilbur H. Duncan

20564 (DUKE 142603, F 1578181, GH, US 2262699, WIS). RICHMOND CO.— 48

+ 27 July 1923, edge of pond no. 2, Augusta, S. F. Hildebrand (DUKE

09730). SUMTER CO.— +21 July 1897, shallow water in pine-barren pond

E of H[?Juntington, Roland M. Harper (BKL, NY). CO. ?— +Cno date],

S. Boykin (Ga.), (PH 783935). + [no date], Ga., Rev. T. C. Porter (PH

704190), ILLINOIS: COOK CO.— Cno date], Calumet Rio, near Chicago,

tHerbarium John J. Brown M.D.] (WIS). July 1859, Chicago, from F.

Scammon, M.D. and Rinywood [?], Vasey (F 32836). 12 August 1880, South

Chicago, (F 1352930). 16 August 1941, on 131st St. between Wolf Rd. and Will-Cook Rd. W of Palos Park Pond, in wettest portions, Julian A.

Steyermark 40917 (F 1328583). FRANKLIN CO.— 5 July 1940, near

Christopher, George Neville Jones 12337 (GH). HENDERSON CO. — tno date], ponds, in low grounds near Oquawka, Harry N. Patterson (F

135235 and 135236, MO 910840). LAKE CO.— Cno date], Fox Lake, Herb. of Charles S. Raddin (F 1211108). +28 July 1907, in water 2-3 M deep,

Grass Lake, abundant, collected on a 360 K'm walk through Eastern

Wisconsin & NE Illinois, Frank C. Gates and R. R. Sleeper 2241, col. no. 1754.1 (MICH). 6 August 1926, Antioch, fresh water lake, Nellie

V. Haynle 181 (F 1514618). MARSHALL CO.— 12-16 Sept. 1906, near Henry,

Dr. Seth E. Meek (F 193380-3). McHENRY CO.— August 1881, McHenry,

A. B. Seymour and 5 August 1880, Big Spring, Mahomet, water, fls. blue,

T. J. Burr ill and A. B. Seymour (DUKE 66221). OGLE CO.— 9 August 1885,

Oregon, Herbarium of Merton B. Waite (DUKE 66206). PEORIA CO.— +[no date], Dr. J. T. Stewart (F 114809). ROCK ISLAND CO. — 7-20 1887,

Port Byron, R. A. Harper (WIS-2). ST. CLAIR CO.— +9 August 1877, "in sun”, H. Eggert (M0 739480-739481); Bluffs Lake, H. Eggert (MO 1983389-

1983393). UNION CO.— 30 August 1952, muddy shores of Wolf Lake, Franklin B. Buser 1895 (F 1472292). WILL CO.--22 August 1904, ponds,

Joliet, H. C. Skeels 461 (F 177347). CO. 7— +[no date], morais in

IiriJinois, Juillet (MO 1983385). +16 June 1891, Lake areas, Centre-

ville, E. Douglass (MO 1983387-1983388). +6 July 1897, DuPage river,

Naperville, L. M. Umbach (F 434558). +20 July 1898, Stony Creek,

Blue Island, 0. E. Lansing, Jr. 368 (F 57704). +July 1901, Des Plaines

River, Summit, L. M. Umbach 12989 (WIS). +Cno day or month] 1904,

M. Gandoger C?] (MO 713769). +7 July 1906, River, Romeo, L. M. Umbach

1107 (WIS). 24 July 1909, Naperville, pond, L. M. Umbach 4095 (WIS).

+13 August 1932, Beach, along Dead River, Nellie V. Haynie 2538-A (F

1525409). INDIANA: DeKALB CO.--30 July 1933, Diamond Lake, 35 miles

N of Auburn, Cora Shoop (F 907946). ELK. I!ELKART] CO.— August 1890,

small lake near Mich., C. H. Kauffman (MICH). KOSCIUSKO CO.— +7-31

1897, Eagle Lake, Chas. C* Deam (F 123136, NY). 5 August 1908, ponds,

Syracuse, M. W. Strahler 28484 (WIS). 30 August 1941, margin of

Dewart Lake, T. G. Yuncker and Winona H. Welch 10687 (GH). LAKE CO.—

+July 1870, "Miller's", ex Herb. Henry H. Babcock (MO 1983380),

+21 July 1898, sloughs, Miller's, L. M. Umbach (US 351283, WIS).

+3 Sept. 1900, border of Wolf Lake, Agnes Chase 1457 (MO 19833 76, PH

515311). +27 July 1909, River, Millers, L. M. Umbach 4160 (WIS).

+14 August 1912, Lake margin, Wolf Lake, L. M. Umbach 6047 (WIS).

+20 August 1913, margin of Calumet River, Miller, Huron H. Smith 5770

(F 408143). 18 July 1914, Bog, Millers, L. M. Umbach 6 725 (NY-2).

+9 July 1915, Miller, G. Ejfrlg (F 1562067). +24 July 1915, Miller,

H. C. Benke 1701 (F 555378, US 1521315). +1 July 1934, marshes N of

Grand Calumet Millers, Carl Buhl 474 (PH 726097). +6 August 1935, ditch on the S side of US 12, E of Miller, R. M. Tryon, Jr. 1751

{DUKE 103087). +27 August 1942, Miller (Gary), H. C. Benke 6179 (F

1126363). LAPORTE CO.— 20 July 1878, Clear Lake, Laporte, J. M .

Coulter (F 363418). +VIII/16 1909, Sagnnay [?], J. A. Nieuwland (MO

741951). VIII, 10 1912, Hudson Lake, J. A. Nieuwland (BKL, MO 742869).

+24 August 1925, The Dunes of Lake Michigan, Pine Lake, J. R. Churchill

(GH, MO 1053768). MARSHALL CO.— 1900, near Lake Maxlnkuckee, J. T .

Scovell and H. Walton Clark 1254 (F 131630, US 407234). PORTER CO.—

16 August 1920, Wa[u]hob Lake, near Valparaiso, Donald C. Feattle (GH).

+5 VIII 1923, Dunn at Tamarack, In 6 inches of water in open bog,

Herb, of Marcus W. Lyon, Jr. (MICH). +22 VIII 1924, Dunes at Mineral

Springs, in about h m water in Little Lake, Herbarium of Marcus W .

Lyon, Jr. (MICH). +4 VIII 1929, dunes at Wycliffe, E end Long Lake,

Herbarium of Marcus VJ. Lyon, J r . (MICH). 21 July 1930, Valparaiso,

Lake Wauhob, H. C. Benke 5304 (F 652820). +8 August 1953, In shallow water of Goose Lake NW of Baileytown, flowers bluish-purple, Munsell

Color No. PB-P 6/8, Floyd A. Swink 2530 (F 1496908). +12 August 1956,

in shallow water along shore of Mud (Goose) Lake, Baileytown, H. R .

Bennett (NY, US 2261306). ST. JOSEPH CO.--July 1891, Mishawaka,

E.B.U. (F 260514). STEUBEN CO.— +24 July 1906, low border of Bass

Lake, Chas. C. Deam 1245 (US 607253). 20 August 1933, shallow water,

James Lake, Angola, Clarance Clark (OS). 9 July 1961, in shallow water of a small lake along Ind. #120, just E of Orland, N. C. Henderson

61-572 (FSU 70891). VIGO CO. — [no date], 5-mile Pond, Terre Haute, private collection of Dr. Barton W. Evermann (US 645862). CO. ?—

+[no date], J. L. Darlington (PH). +2 August 1876, gathered at Pine Station, Geassly [?] (F 289262). +29 June 1878, Pine Sta., Herbarium of E. S. Bastin (F 1344424). +29 June 1895, sloughs, Pine, L. M .

Umbach 2986 (WIS). +4 August 1900, Slough, Clarke, 0. E. Lansing, Jr.

1078 (F 85773). +6 Oct. 1900, ditch, Dune Park, L. M. Umbach 11309

(WIS). +7 August 1905, Clark Junction, edge of pond, south Chicago

Area, Herb, of Frank C. Gates 121 (MICH). +20 June 1909, sloughs,

Dune Park, L. M. Umbach 3980 (WIS). +13 August 1910, bed of drying

Blough, Gibson, 0. E. Lansing, Jr. 2833 (F 282249, GH, WIS). + [no day or month] 1926, Dune Park, D. C. Peattle 2302 (F 565055). IOWA:

APPANOOSE CO.— +7-19 1896, Herb, of T. J. and M. F. L. Fitzpatrick

(MO 1983394). BENTON CO.--[no day or month] 1879, Vinton, J. J .

Davis (WIS-2). BOONE CO.— 20 August 1898, Ogden, L. H. Pammel (GH).

DECATUR CO.— +17 August 1898, borders of ponds, J. P. Anderson (MO

1983396). +8-10 1899, ponds, swamps, infrequent, Herb, of T. J. and

M. F. L. Fitzpatrick (F 123443, NY). HAMILTON CO.— 4 August 1936,

Goose Lake, lh miles E of Jewell, George J. Goodman 2961 (M0 1230019,

NY). HENRY CO.— August 1897, Mt. Pleasant, J. H. Mills 528 (GH, M0

1983395, US 324291). LINN CO. — [no date] 18__, Cedar Rapids, Albert

S. Hitchcock (MO 1983397). LOUISA CO.— +August 1897, cut off ponds on alluvial flat, near Parker's landing, Muscatine Isl., B. Shlmek and

P. C. Myers (MO 1245456). +17 July 1927, south side of Dewey's swamp

SW of Cone, B. Shimek (WIS). MUSCATINE CO.— 20 July 1922, Bwampy border of pond among old dunes, 8 miles NW of Muscatine, medium styles,

B. Shlmek (NY). 29 Sept. 1951, shallow water of sluggish tributary running E into Muscatine Slough, Sec. 12, Seventy-six Twp., Robert F .

Thorne and E. 0. Beal 13917 (US 2230265). KANSAS: CHEROKEE CO.— 52

+17 August 1897, ponds, Geo. L. Clothier 1098 (GH, MICH, MO 1983420,

NY, US 353900). +Summer 1898, L. V. Harvey (PH 531898). KENTUCKY:

BALLARD CO.— 18 August 1923, Wickllffe, in water Swan Pond, Frank T.

McFarland and W. A. Anderson, Jr. 16 7 (MO 903743). DAVIESS CO.— July

1851, Owensboro, Herb, of GeorRe Scarborough (BKL). JEFFERSON CO.—

14 July 1939, ponds, 525 ft., growing in old quarry, Louisville,

M. Seargent [?] ^2^ (GH). CO. ?— +Cno date], ponds in the Barrens of

Ky. fl. June, C. W. Short Herbarium (PH 783881). +[no date], C . W .

Short, M.P., Lexington, Kentucky (GH, NY). LOUISIANA: BEAUREGARD

PARISH— 29 April 1955, roadside ditch near Longville, flowers purple,

Cooley & Brass 4072 (GH). CALCASIEU PARISH — [no day or month] 1904, vicinity of Lake Charles, Andrew Allison 106 (F 286523, MO 1983364,

NY, US 514042) and 220 (F 286528, MO 1983365, NY, US 514058). 18 May

1915, Lake Charles, swamps, E. J. Palmer 7689 (MO 793920, NY, US

1531680). IBERIA PARISH— 15 Oct. 1963, one clump noted in shallow water along edge of upland lake, Avery Island, John W. Thieret 16504

(DUKE 158871, FSU 85370, US 2433308). JEFFERSON PARISH— 2 May 1899, very abundant in ditches, collected at Gretna, opposite New Orleans,

Carleton R. Ball 329 (CAN 165675, F 93385, GH, MO 1983367, NY, US

382721). LAFAYETTE PARISH— +7 July 1942, Anse La Butte pond, also roadside ditches, G. B. Claycomb (GH). 5-16 1967, small pond, 5 miles

S of Rayne, Jean Wooten 1027 (FSU 109978). ORLEANS PARISH— 30 April

1846, New Orleans, in swamp-[?], A. Fendler (MO 1983373). 21 Oct. 1885, swamp, New Orleans, J. F. Joor (MO 1983369). 5 July 1886, swamp north of New Orleans, J. F. Joor (MO 1983371). [no day or month] 1896, New

Orleans, fron R. Dodson (MO 1983366). 2 Sept. 1923, New Orleans, sunny marsh, Otto Degener 564 (NY-2). 23 August 1936, drainage ditch,

New Orleans, Saint BernardC?], Delzle Demaree 13505 (MO 1178796, NY,

US 1699228)* OUACHITA PARISH— 31 July 1955, roadside swamp, Calhoun

S27 T18N R1E, John A. and Jack Moore 6270 (GH). ST. BERNARD PARISH—

+5/23 1960, edge of cypress swamp along Lake Borgne Canal about % mile

E of Violet, fairly common, Robert J. Lemaire 1037 (US 2348560).

ST. CHARLES PARISH— 26 April 1955, in shallow water, very highly organic soil with much undecomposed vegetable matter, near Sarpy

Brothers well #5 Sarpy Wildlife Refuge, approx. 15 miles W of New

Orleans on Hwy 61, E. Lowe 98 (GH). ST. MARTIN PARISH— July 1878, in ditches and ponds, Pointe a la Hache, St. Martinville, A. B. Langlols

(M0 981882). ST. MARY PARISH— 19 June 1893, near Morgan City, E. 0 .

Wooton (&S 735171). +20 April 1933, ditch along roadway and bayous,

Bayou Sale, Bynum, Ingram, Joynes (BKL). ST. TAMMANY PARISH--13 July

1919, vicinity of Covington, Bro. Anect 92 (F 505010). TANGIPAHOA

PARISH— 23 August 1938, in a canal along road, near the Tangipahoa

River, 1 mile W of Robert, D. S. and H. B. Correll 10541 (DUKE 72059,

GH). 5-6 1963, 2 miles S of Ponchatoula, swamp, Roomie Wilson 396

(FSU 81704). PARISH ?— +[no date}, "Louisiana", Sainturier C?], (PH).

+August 1847, Croton; Vide Synopsis Flor West, No. 1458, Louisiana,

Herb, of J. L. Riddell. M.D. (NY). +16 April 1887, West End, La.,

J. F. Joor (MO 1983372). +1 Nov. 1888, west End Swamp, La., J. F. Joor

(MO 1983370). +20 June 1917, swamp near Sabine River, J. Bequaert

(GH). +1 August 1925, Delta Duck Club, Inarantine C?], C. C. Spury

418 (US 1467125). MAINE: ANDROSCOGGIN CO. — [no day or month] 1893,

1894, South Poland, Kate Furbish (NEBC). Cno day or month] 1894, East

Livermore, Kate Furbish (NEBC). 16 July 1896, in shallow water Ea. Auburn, E. D, Merrill 1043 (NEBC), 23 July 1915, pond, Leeds,

Clarence H. Knowlton (NEBC). CUMBERLAND CO,--9 1897, Westbrook,

A.H,N,, Herb. F, L. Ricker 1043 (US 914665). Sept, 1900, West Baldwin,

Kate Furbish (NEBC). 18 August 1903, Mountfort’s Mill Pond, Town of

Cumberland, Edward B. Chamberlain 623 (NEBC). 26 June 1913, The Park,

Brunswick, Kate Furbish (NEBC). 2 Oct. 1969, Pleasant River at Route

95, 1 mile S of Route 202, Gray In Cumberland County, J. N. M. Howells

J9 (OS). FRANKLIN CO.— Sept. 1894, Pease Pond, E. Wilton, Kate Furbish

(NEBC). +3 August 1937, Webb Lake, in lake, Dale W. Jenkins (OS).

13 August 1941, Swale, New Vineyard, Clarence H. Knowlton (NEBC).

HANCOCK CO.— Cno date], Orland, H. M. Atkins (NEBC). 16 July 1880,

Flora of Mount Desert I., Somesville, Wm. H. Dunbar and 28 July 1892, shore of Northwest area, Great Pond, Flora of Mount Desert Island,

Edward L. Rand (NEBC). 18 August 1888, NC?]adlock’s Upper Pond, Flora of Mount Desert I., Edward L. Rand (NEBC). 27 August 1888, Seal Cove

Meadow, Flora of Mount Desert I., Edward L. Rand and 29 August 1890,

Breakneck Pond, Flora of Mount Desert Island, Edward L. Rand (NEBC).

14 June 1889, N. E. branch, Flora of Mount Desert Island, Rand and

Redfield (NEBC). 25 August 1890, Somes Brook, Somesville, Flora of

Mount Desert Island, Edward L. Rand and 16 August 1889, Brooks, Somes­ ville, Flora of Mount Desert Island, Rand and Red field (NEBC). 30 July

1890, Mill-dam, Somesville, Flora of Mount Desert Island, John Red- field (MO 126184, PH). 6 August 1910, Lower HaSlock Pond, Mt. Desert,

Emile F. Williams (GH). 15 Sept. 1969, outlet, Alamoosooc Lake, Route 1,

Orland, Hancock County, J. N. M. Howells 4 (OS). KENNEBEC CO.--Cno date], pond margin, Readfield (PH 608102). 31 August 1873, Manchester, 55

F. Lamson Scribner (NEBC), 16 August 1904, River, Flora of Clinton,

Ralph Carleton Bean (NEBC). 13 July 1933, Kennebec Co., shallow water of a stream, Readfield, N. C. Fassett 15893 (WIS), 2 August 1933, wet shore of Cobbossee Stream, Gardiner, N. C. Fassett 15734 (F 1430028),

12 Sept. 1969, Cobbosseecoutee Stream, Town Line, Litchfield and

Gardiner, Kennebec County, Route 95, J. N. M. Howells 1 (OS). KNOX

CO.— 12 August 1913, Thomaston, D. W. Fellows 5599a (US 785921). July

1928, Camp Medomak, near Washington, Julian A. Steyermark (F 887461).

2 July 1929, Otter Island S of Friendship in Penobscot Bay off coast of

Maine, Julian A. Steyermark (F 887404). 20 July 1929, peaty margins of south end of Washington pond, near Washington, Julian A. Steyermark

1080 (M0 1026287). 15 Sept. 1969, North Pond, N side of Route 1, 1 mile E of Lincoln County line, Warren, Knox County, J. N. M. Howells 7

(OS). LINCOLN CO.— 2 August 1898, in shallow water Penaquid C?1 river, Bristol, E. B. Chamberlain 682 (NEBC). +7 August 1930, at high-tlde level, estuary of Eastern River, Dresden Mills, N. C .

Fassett 10523 (WIS). 11 August 1937, Campbell Pond, Boothbay, N. C .

Fassett 18812 (MO 1149720) and 18813 (WIS). 28 August 1937, Lincoln

Co., shallow water, Damarlscotta Pond, Bunker Hill, N. C. Fassett

18814 (NEBC, WIS-2) and 18820 (F 917294, NEBC, WIS). 3 Sept. 1969,

Lincoln County, in pond 3 miles S of Newcastle center off Route 1

(the Newcastle Rest Area), abundant, associated with Nymphaea, Typha, and Sparganlum, R. M. Lowden 31 (OS-5). OXFORD CO.--July 1884, Hart­ ford, J. C. Parlin (GH). 14 Sept* 1893, Buckfield, J. A. Allen (WIS).

11 August 1893, Buckfield, J. A. Allen (DUKE 66213). 1 Sept. 1894,

Bryant's Pond, Ex Herb. J. W. Blankinshlp (NEBC). Cno day or month] 1897, Gilead, Kate Furbish (NEBC), +8/5 1929, Albany, Crooked River,

large clumps of great beauty, Herb. Leston A. Wheeler (NEBC). 22 July

1942, Brook, Sumner, Ralph Carleton Bean (NEBC). PENOBSCOT CO.— July

1869, streams, Hudson, F. P. Briggs (NEBC). 10 July 1890, River

intervale, Orono, M. L. Fernald (NEBC). 25 August 1916, shallow waters by Otter Chain Ponds, Milford, M. L. Fernald and Bayard Long

13171 (NEBC, PH 575297). 12 Sept. 1969, Dolby Flowage, East Millln-

ocket, Penobscot County, Route 157-11, J. N. M. Howells 2 (OS).

14 Sept. 1969, South Branch Penobscot River at Route 116, Howland, a

distributary of the Penobscot running immediately into the Piscataquis

River, freshwater, Penobscot County, J. N. M. Howells 3 (OS).

PISCATAQUIS CO.— 22 August 1867 [?], Spender Pond, [East Middlesex],

C. E. Smltte [?] (NEBC), SAGADAHOC CO.— 24 August 1920, very abundant on tidal mud-flats, estuary of the Cathance River, Bowdoinham, N. C .

Fassett 36 (F 501770). 24 August 1921, Tldal-flats about Merrymeeting

Bay, Bowdoinham, sandy shore at high-tide level, W side of the Kennebec

River, opposite the foot of Swan Isl. [7], N. C. Fassett 159 (NY).

August 1921, Tidal-flats about Merrymeeting Bay, Bowdoinham, N. C .

Fassett 223 (WIS). 3 Sept. 1969, Sagadahoc County, Cathance River at

the Old Feldspar Mill off Cathance River where the Cathance River

empties into the estuary by the Cutler home, 1 mile from Topsham, l^s miles from Bowboinham, fresh water, abundant, R. M* Lowden 29 (OS-4).

3 Sept. 1969, Sagadahoc County, muddy River (tidal river), Foreside

Road near Topsham (off route 24), abundant, associated with Nuphar advena and Saglttaria latlfolia, R. M. Lowden 30 (OS-9). SCMERSET

CO.— [no day or month] 1873-8, Cambridge, Valley of Sebasticook River, F. S. Bunker (GH, NEBC). +21 July 1894, Wood's Dam, Mercer,

E. Edwards, Thos. C. Porter (7 703735). WALDO CO.— 19 July 1912,

St. Georges Lake, Liberty, J. A. Cushman (NEBC). 13 August 1930,

swampy lake shore, Pitcher Pond, near Rocky Point, Lincolnville,

G. B. Rossbach 112 (NEBC). +8-13 1936, shallow water on margin of

Megunticook Lake, Ray C. Friesner 9764 (DUKE 41003, F 885741, NY-2).

+8-16 1952, boggy margin of Mullen's Pond, N side of Megunticook Lake,

Ray C. Friesner 25210 (DUKE 147954, WIS). 15 Sept. 1969, small

stream, S side of Route 3, adjacent to cemetery, 1.4 miles W of

Marrill-Searsarout line, in Marrill Waldo County, J. N. M. Howells 5

(OS). 15 Sept. 1969, Ruffwghau Meadows Pond north Searsarout, Waldo

County, J. N. M. Howells 6 (OS). WASHINGTON CO.--+2 August 1916,

Pond, Marshfield, Clarence H. Knowlton (PH 562994). 14 August 1928,

River Machias, Clarence H. Knowlton (NEBC). 13 August 1931, Brook,

Northfield, Clarence H. Knowlton (NEBC). +9 August 1932, shallow water, Love Lake, Crawford, Clarence H. Knowlton (NEBC). +14 August

1939, Pond, shallow water, Marion, Clarence H. Knowlton (NEBC).

8 August 1940, shallow water, outlet of Lake, Lambert Lake, Clarence

H. Knowlton (NEBC). YORK CO.— 26-7 1873, River margin, Limington,

J. H. Pillsbury (PH 608082). 27 July 1894, in shallow water, common,

North Berwick, John C. Pari in (NEBC). 27 August 1919, in shallow water, ponds and streams, Kennebunkport, Mrs. Amy M. Pier 196 (DUKE

114389). 12 Sept. 1919, Kennebunkport, Amy M. Pier 3 (NEBC). 2 Oct.

1969, Kennebunk River on Town line and Arundel, York County, Rt. 95,

J. N. M. Howells 8 (OS). 2 July 1970, York Co., Bonnie Bay Pond,

North Berwick, W of Route 4, R. M. Lowden 1 (OS-3). July 1970, York Co., Kennebunk River, boundary between Kennebunk and Arundel Towns,

Route 95, freshwater, R. M. Lowden 4 (OS-2). MARYLAND: CECIL CO.—

7-4 1903, Northeast, Stewardson Brown (PH 508194). 7-11 1923, forming

colonies in water, Chesapeake City, Ivar Tldestrom 11402 (GH).

15 July 1933, tidal marsh, along branch of Back Creek, N of Chesapeake

City, Bayard Long 41115 (PH). +30 June 1946, swamp, 1 mile SE of

Hance's Point, C. Irvin Stiteler (GH, PH). HARFORD CO.— +17 August

1865, Port De-Posite, J. J. Carter (PH). 2 August 1902, in bog at

foot of Havre de Grace Light, George H. Shull 158 (MO 1983352, NY).

KENT CO. — 25 June 1895, Betterton, A.M.G. (PH). 18 August 1900,

Chestertown, E. G. Vawatta [7] (PH). MIDDLESEX CO.--+5 July 1893,

Middlesex Co., Concord River, Walter Deane (GH-5, NEBC-2). WORCHESTER

CO.— +24 August 1934, edge of stream, near old Furnace, G. F. Beaven

80 (DUKE 32101). CO. ?--+24 July 1890, Kings Creek, J. H. Holmes 87

(US 33947). +3 Sept, 1902, Little Gunpowder River, between tides,

young stages, George H. Skull 312 (GH, MO 1983353, NY). MASSACHUSETTS

BARNSTABLE CO. — [Date ?], 28 July 19L?, Welfleet [Wellfleet], F. S.

Collins (NEBC). 22 August 1888, Hyannisport, J. R. Churchill (MO

1053765 and 1053766). 22 August 1888, Hyannisport, stock sectioned,

Walter Deane and 23 August 1888, fruit, Half-Way Pond, Walter Deane

(NEBC). 4 July 1896, Wequenoket [Wequaquet] pond, Centreville,

Emile F. Williams (GH). 4 July 1896, Great Pond, Centreville, Cape

Cod, Emile F. Williams (PH 633849). 12 Sept. 1897, Nine Mile Pond,

Centreville, Emile F. Williams (NEBC). 4 Sept. 1898, Nine Mile Pond,

Cape Cod, J. M. Greenman 358 (MO 737907). 4 Se [Sept.3 1901, Harper’s

Pond, East Orleans, T. W. Edmondson 2249 (NY 2569). 30 July 1902, Orleans, John Murdock, Jr. (F 469568). 11 August 1906, So. Orleans,

Slnnellfe Pond, John Murdoch, Jr. 2075 (F 470352, NEBC) and 2083 (F

470360, NEBC). 5 July 1907, Dennis Pond, Yarmouthport, E. W. Slnnott

(NEBC). 11 July 1907, Aunt Jemlnias Pond, Eastham, Frank S. Collins

278 (NEBC). 9 Sept. 1913, damp sandy borders of small ponds W of

White Pond, not In water, Chatham, M. L. Fernald and Bayard Long 9139

(GH-2, NEBC-2, PH 551153). 7 August 1914, Pocasset, Bourne, Flax

Pond, F. S. Collins 2833 and 25 August 1914, BrewBter, F. S. Collins

3131 (NEBC). 16 Sept. 1916, Barnstable, pond, Clarence H. Knowlton

(PH 562462). 16-17 Sept. 1916, Southwestern section, Barnstable, edge of water, R. C. Bean, F. W. Bird, and C. H. Knowlton (NEBC). 28 July

1918, wet, peaty quagmire-margin of pond S of Hawk's Nest Pond,

Harwich, M. L. Fernald 16515 (NEBC). 22 August 1918, sandy and peaty margin, Lower Simmons Pond, Dennis, M. L. Fernald and Bayard Long

16516 (NEBC, PH 575298). 25 July 1919, borders of streams, Yarmouth­ port, I. V. and M. A. Freeman (BKL). 9 July 1924, Nobska Point, Woods

Hole, Small lake N of light house, Otto Degener 566 (NY). 8 August

1928, Boggy swale, along Mashpee R., Mashpee, John M. Fogg, Jr. 3602

(MO 951845, NEBC). BERKSHIRE C0.--1 August 1894, vicinity of Great

Barrington, C. L. Pollard (US 309048). 25 August 1902, Pittsfield,

R. Hoffman (NEBC). 24 July 1912, Lake Buell [Buel], R. Hoffman (NEBC).

20 July 1916, ice pond, Housatonic, R. Hoffman (NEBC). 29 July 1920,

Lake Buel, New Marlboro, J. R. Churchill (MO 1053770, NEBC-2). 31 July

1951, inlet of Stockbridge Bowl, Thomas S. Githens 134 (PH 845207).

BRISTOL CO.— [no date}, New Bedford, Herb. E. Williams Hervey (NEBC).

Cno dateJ, New Bedford, Herbarium of Thomas A. Greene (WIS). July 1890, North Easton, Carl Blomberg (NEBC). 19 July 1911, Swansea, in

stream, Herbarium of S. N. F. Sanford (NEBC). 16 July 1932, Westport,

shallow water, S. N. F. Sanford (NEBC). 17 Sept. 1935, Seuddlng Pond,

Taunton, F. C. Seymour (DUKE 66209). 16 Sept. 1957, bases swollen

apparently to resist drought, with Bidens laevls, in marsh along

Segreganset River, region of conglomerate rock, Dighton, Frank C .

Seymour 17735 (WIS). DUKES CO.— 4 Sept. 1898, Chilmack, common,

Sydney Harris (NEBC-2). 22 June 1917, ditch in cranberry bog, Edgar-

town, F. C. Seymour 1634 (GH, NEBC). 12 Sept. 1917, cranberry bog,

Edgartown, Flora of Martha's Vineyard, F. C. Seymour 1142 (DUKE

66211, GH, NEBC, NY, PH 738599, US 1103889). [no day or month} 1924,

Gay Head, edge of pond near town, [Gay Head is on Martha's Vineyard],

Otto PeRener 562 (NY). 4 August 1927, Nashawena, E end of island,

swampy margin of large pond, John M. Fogg, Jr. 2368 (NEBC). ESSEX

CO.— Cno date], Salem, Mass., C. Pickering (PH). +[no date], Magnolia,

Mass., '98, Miss A. F. Rogers (NEBC). 28 July 1868, Ipswich River,

Ipswich, very common, Morong (NY). 10 July 1877, Chebacco Pond,

Beverly, J. C. Phillips (NEBC). 20 July 1877, Manchester, Herbarium of William Coolidge Lane (GH). 13 July 1879, Lynnfield, Herbert A .

Young (NEBC-2). 7-8 1885, North Andover, Herbarium of C. H. Morse and Georgetown, mill pond, Herb. Charlotte N. S. Horner (NEBC).

13 Sept. 1903, Salem, edge of ponds, F. F. Forbes 915 (NEBC). 14 July

1909, Amesbury, Herbarium of Edwin B. Bertram, L. P . (PH 648755).

+21 August 1913, Byfield, an evidently fresh marsh, very wet, Donald

White 393 (NEBC). 30 July 1933, Peabody, R. B. Machlntosh (NEBC).

14 July 1950, edge of Ipswich River, Middleton, Ralph Carleton Bean (NEBC). HAMPDEN CO.— July 1884, Monson, water, E. L. Morris 194

(BKL) and 3233 (BKL). 10 Sept. 1915, Flora of Springfield, frequent

In ponds and pools, Island Pond, L. Andrews (NEBC). 9 Sept. 1922, pond shore, Ludlow, F. W. Hunnewell and H. K. Svenson (NEBC). 11 July

1925, in quiet water along the shore, Muddy Brook, Chicopee, F. C .

Seymour 460 (DUKE 66218, GH, MO 927742, WIS). HAMPSHIRE CO.— +31

August 1927, Brookfield, Quabog Lake, Alfred S. Goodale 52482d (DUKE

66210). 24 July 1932, bank of Connecticut River, Hadley, F. W .

Hunnewell 12.503 (NEBC). MIDDLESEX CO.--[no date], Middlesex Flora,

G. E. Davenport (NEBC). [no date], Jamaica Plain, Herb. C. E. Faxon

(GH). [no day or month] 1849, West Newton, Anna P. Jones (NEBC).

29 July 1854, W. Cambridge, from Herb. William Boott (GH). 10 August

1857, Concord, river bank, Edward S . Hoar (NEBC). July 1878, in water edges of ponds, Ashland, Morong (F 134393). July 1878, Ashland, flowers July, fruits September, Herb. Thomas Morons (NY). 26 Sept.

1878, Cambridge, C. E. Perkins (CAN 165677). July 1880, Plants of

Westford, Emily F. Fletcher (GH, NEBC). 29-VII 1883, 21 VIII 1884,

Bedford, Herb. Charles W. Jenks (NEBC). 15 August 1885, Weston,

F. S. Collins (NEBC). 3 July 1890, Waushakim [?] Pond, So. Framingham

E. L. Sturtevantii (MO 1983334). 26 July 1893, Concord, young plant is shallow water of Concord River, Walter Deane (GH-4, NEBC-2).

18 July 1901, North Reading, Arthur S. Pease 281 (NEBC). 1 August

1902, Wilmington, Silver Lake, Geo. H. Tilton (NEBC). +23 August

1904, East Lexington Meadow and vicinity, Lexington, in water, G. C.

Worthen (US 646065). +7 July 1906, shallow water, Hardy's Pond,

Waltham, Clarence H. Knowlton (NEBC). 14 July 1907, Concord, Edwin B. Bertram (PH 532623). 25 July 1911, Flora of Sherborn, Brook, Martha

Louise Loomis 361 (NEBC). 7 Sept. 1913, Cambridge, Clear River Marsh

back of Stellman Infirmary, flowering plant, extreme south end, Walter

Deane (NEBC). +7-24 1915, Flora of Wakefield, Clapp's Island, water,

Winfield S. Ripley, Jr. (NEBC). +24 July 1919, Newton Centre, Austin

H. Clark 26 (F 505379, MO 867108). 11 July 1920, Charles River,

Natick, ex Herbarium of F. W. Hunnewell, 2nd and Westford, Miss E. F .

Fletcher (NEBC). +16 August 1924, edge-of-pond, Melrose, Mass. (NEBC).

11 July 1934, Ayer, D. Leighton Ordway and Charles Bullard (NEBC).

20 July 1957, Concord, shallow water, edged of river, specimen shows

the three flower forms, R. J. Eaton (NEBC). 20 Sept, 1961, In very

shallow water on muddy sand, Concord, edge of Bateman's C?] Pond,

R. J. Eaton 5145 2/3 (NEBC). NANTUCKET CO.— 5 Sept. 1904, Polpis,

plants collected on Nantucket Island, Eugene F. Bicknell (NY). 8 Sept.

1904, Maxcy's Pond, Nantucket Island, Eugene P. Bicknell 255 (NY).

19 Sept. 1907, Polpis Pond, Nantucket Island, Eugene P. Bicknell 256

(NEBC). 19 August 1921, Hummock Pond, Francis W. Pennell 11059 (PH

729511), 25 August 1960, Slasconset, Nantucket Is., Swamp, Frank C .

MacKeever N492 (NY). NORFOLK CO.--+19 July 1870, Punkapog Pond,

Canton, J. R. Churchill (MO 1053769). +20 July 1862, Dedham, J. R .

Churchill (MO 1053764). +22 August 1882, Lily Pond, Cobasset,

W. Deane (NEBC). 12 July 1883, Needham, Herb, of T. Otis Fuller (NEBC).

+29 Sept. 1883, Chas. River, Dedham, Walter Deane (NEBC). +3 Sept.

1888, Ponkapog, Canton, fruit, Geo. G. Kennedy (GH). +26 June 1890,

Paul's Bridge, Milton, Geo. G. Kennedy (PH 633848). +5 July 1890,

Milton, Neponset River, Walter Deane (NEBCO. +5 July 1890, Paul's Bridge, Milton, Geo. G. Kennedy (WIS). +5 July 1890, Neponset River,

Paul's Bridge, Milton, Geo. G. Kennedy (PH 633847). +12 July 1890, white, near Paul's Bridge, Milton, Geo. G. Kennedy (NEBC). +19 July

1919, Milton [?], Pine Tree Grown, at Broon [?] road, Herbarium

Nathaniel Thayer Kidder (NEBC). +1 Sept. 1923, Great Pond, weymouth,

Herb, of Clarence A. and Austin W. Cheever (NEBC). PLYMOUTH CO.—

[no day or year! July, S. Scituate, Herb, of Wm. P. Brooks (NEBC).

26 August 1923, Cushing pond, Hingham, Clarence H. Knowlton (NEBC).

SUFFOLK CO.— +[no day or month] 1816, Boston, N.A., ex Herbario William

Mitten, Dr. Boot (US 937958). +22 August 1882, shallow water, Read- ville, Herb. Jas. Lund. (NEBC). +3 Sept. 1888, W. Roxbury, Herb. C. E .

Faxon (GH). +17 July 1919, Stony Broon, N. T. Ridder [?] (NEBC).

WORCHESTER CO.— +[no date], Ltmbridge [?], Herb. S. F. Tower 14790

(NEBC). +[no day or month] ca. 1854, Lvnenburg [?], Mrs. W. B. Rogers

(NEBC). 24 July 1879, Beaver St., Worcester, Hattie Merrificld (NEBC).

+25 July 1880, Westminster, C. W. Mlnott (US 789263). +17 August 1929,

S. Spectacle Pond, Athol (S. Spectacle Pond is near South Athol, but ia located in the township of New Salem), Alfred S. Goodale, Walter C .

Markert, and Randall H. Piper (NEBC). +20 August 1931, Dana, Pottapang

Lake, A. S. Goodale, S. F. Potsubay, and F. R. St. John (NEBC).

+18 July 1935, Hardwick, edge of small river, Charles Foster Batchelder

(NEBC). +3 August 1935, Harvard, shallow margin of mill pond, Charles

Foster Batchelder (NEBC). +4 Sept. 1944, in clumps near shore, Clam­ shell Pond, Clinton, F. C. Seymour 7108 (DUKE 114443, NEBC). CO. 7--

+16 July 1888, just north of Nonquit [?], E. L. Sturtevanti (MO

1983333). +Sept. 1895, Wellerery [?], Ex. Herbarium of Wellesley College (US 270864). MICHIGAN: ALGER CO.--+29 July 1966, colonies frequent at N end of Sand Lake, ca. 3^ miles NW of Deerton, flowers blue, E. G. Voss 12263 (MICll). ALLEGAN CO.— +August 1898, East Lake,

N. A. Dohmen 48 (F 58091). +19 August 1902, W end of Swan Lake, In lh feet of water, W. F. Wight 7 (NY, US 430059 and 430060). +7/20

1937, T2N R13W Sec. 4-5, Dumont Lake, sta. no. 1, 1 foot of water,

42° 35’ N Lat., 85° 51’ W Long., marl bottom, D. C. Chandler 125

(MICH). +7/26 1937, T2N R15W Sec. 13, Round Lake, sta. no. 1, 8 inches of water, 42° 33' N Lat., 86° 01* W Long., bottom of sand and some fibrous mat, D. C. Chandler 177 (MICH). BARRY CO.— 14 August

1936, Long Lake, 1 foot of water, bottom of fibrous and pulpy peal,

L. A. Woodbury 317 (MICH). 8/14 1936, T3N $10W Sec. 28-29, Long Lake,

6 Inches of water, 42° 38' N, 85° 30' W, bottom pulpy and fibrous peat, L. A. Woodbury 326a (MICH). 16 August 1940, Fine Lake, water 2

(0-2) ft., mare bottom type, J. Funk (MICH). +5 July 1958, occasional, mucky shore at SW end of Crooked Lake, NE 1/4 Sec. 22, Pralrieville

Tp., ca. 3 miles SW of Pralrieville, E. G. Voss 7231 (MICH). BERRIEN

CO.--3 July 1911, ditch, in meadow, 1 mile W of Stevensville, 0. E .

Lansing, Jr. 3235 (F 293088, GH, US 753376, WIS). BRANCH CO.— 4 July

1959, uncommon in shallow water of outlet near N end of Brown's Lake,

N edge Sec. 5, California Tp., ca. 2 miles NW of California, E. G.

Voss 8792 (MICH). CALHOUN CO.— +31 July 1938, Wabascow Lake, 3 (1-3) ft. of water, mare fibrous peat bottom, P. Eschmeyer 503 (MICH).

CASS CO.— +8 July 1941, Hemlock Lake, 0-1 foot of water, fibrous peat bottom, E. W. Roelofs (MICH). CASS-VAN BUREN COS.— 9 August 1904,

Lake, Magician Lake, L. M. Umbach (WIS). 20 July 1915, Bogs, Magician Lake, L. M. Umbach 7109 (WIS). CHIPPEWA CO.— 13 July 1963,

frequent, wet shore of Little Duck. Lake, Sugar Island, Sec. 26, ca.

1% miles NW of Homestead, E. C. Voss 11241 (MICH). CLARE CO.—

+12 August 1916, Border of Lake, Lake Station Mich., R. R. Drelsbach

(MICH). +12 August 1916, edge of Cranberry lake, Lake Station, Mich.,

R. R. Drelsbach (F 531611, PH 585276). CHEBOYGAN CO.— 31 July 1932, outlet of Black Lake, into Black River, Flora of Douglas Lake Region,

J. H. Ehlers 5076 (F 1566656, MICH). 8-9 1967, isolated colonies within shallow water, upper Black River, Jean Wooten 1766 (FSU 115911).

EMMETT CO.— 4 August 1930, aquatic herb growing in a floating mat on

French Farm Lake, collected in the vicinity of the University of

Michigan Biological Station, Frank C. Gates 16440 (GH). 18 July 1935, in shallow water, French Farm Lake, Flora of the Douglas Lake Region.

J. H. Ehlers 5859 (MICH). GOGEBIC CO.— 8-11 1933, Cisco Lake, shallow water-sand, Nellie V. Haynie 1867 (F 1525407). GRATIOT CO.— 11 July

1891, Alma, Charles A. PavlB (MICH). HILLSDALE CO.--6/25 1937,

Hillsdale, T5S R3W Sec. 15, Big Cobb Lake, sta. no. 1, 18 inches of water, 42° 03f N Lat., 84° 38' W Long., marl bottom, D. C. Chandler 4

(MICH). HOUGHTON CO.— 27 July 1948, common in shallow water and on mucky shores, Sturgeon Slough E of Chassell, Charles D. Richards 1141

(MICH-2). INGHAM CO.— 11 August 1938, Lake Lansing, 3 (1-3) ft. of water, fibrous peat bottom, P. Eschmeyer 699 (MICH). IOSCO CO.— 21

July 1935, Hale, in water in Loon Lake, R. R. Drelsbach 8533 (MICH).

JACKSON CO.--9 July 1838, Jackson, in small lakes (the original label is in the hand of John Wright), collected by John Wright, Botanist, with the aid of George Bull, assistant botanist (MICH). +10 July 1838, Jackson Co., small lakes, In water (the pencil label Is thought to be the work of an herbarium assistant, ca. 1870), collected by

John Wright, botanist with the aid of George H. Bull, assistant botanist (MICH). KENT CO.--28 August 1891, Reed*s Lake, Grand Rapids,

H. M. Bailey (MICH). 7-7 1941, Lake shore, Crawford Lake, 30 miles NE of Grand Rapids, C. W. Bazuin 3341 (MICH). LIVINGSTON CO.— +5 Sept.

1936, In shallow water, Strawberry Lake, J. H. Ehlers 6163 (MICH).

29 July 1938, stream from Lime Kiln to Sandy Bottom Lake, 1 foot of water, sand mare bottom, M. R. Raymond 394 (MICH). +4 August 1938,

Carpenter Creek, %-l^ feet of water, silt detritus gravel bottom,

M. R. Raymond 437 (MICH). +22 July 1942, Grid P-22, pond in tamarack swamp, as socle s-Nuphar, F. N . Hamerstrom, Jr. 532 (MICH). MASON CO.—

12 July 1910, Hamlin Lake, Bog, Ralph W. Chaney 150 (F 296891, GH, NY,

US 752988). 18 August 1956, occasional at edge of water, sandy E shore of Hopkins Lake, ca. 5 miles SE of Ludington, E. G. Voss 3266

(MICH). MECOSTA CO. — 7 August 1927, shore of Lake, Chippewa Lake,

R. R. Drelsbach 5258 (PH 655474). +20 July 1940, Tubbs Lake, 2 (1-3) feet of water, muck bottom, J . Funk (WIS). +31 July 1940, Bullhead

Lake, 1 (0-2) feet of water, peat bottom, J. Funk (MICH). MISSAUKEE

CO.— 12 August 1941, Missaukee Lake, 1 foot of water, pulpy peat bottom, J. Funk 484 (MICH). MONROE CO.— +5 August 1949, Pte. Moulllee

State Game Area, Silver Creek, N of Pte. Moulllee State Game Area,

Malcolm McDonald 5315 (MICH). MONTCALM CO.— 28 July 1941, Crystal

Lake, 1 (0-1) foot of water, mare pulpy peat bottom, E. W. Roelofs 369

(MICH). MUSKEGON CO.--7-7 1941, 1 mile E of Meva [Mona] Lake growing in wet sand, C. W. Bazuin 3216 (MICH). 27 August 1955, locally 67

abundant in water at edge of Half Moon Lake, NE 1/4 Sec. 14, Casnovia

Tp., ca. 2.5 miles NW of Casnovia, flowers blue, Edward G. Voss 2833

(MICH). NEWAYGO CO.— +8-17 1941, Crystal Lake Wooster, Lake shore and

in water, C. W. Bazuin 3904 (MICH). OAKLAND CO.— 20 July 1916,

Lakeville, John M. Sutton 266 (MICH). 4 Sept. 1938, Milford Pond, 3

(1-3) feet of water, clay muck bottom, P. Eschmeyer 995 (MICH).

OCEANA CO.— 6 July 1963, occasional, sandy mucky shore of Jakes Lake,

Sec. 10, Claybanks Tp., ca. 5^ miles NW of Rothbury, E. G. Voss 11208

(MICH). ONTONAGON AND GOGEBIC COS.— 20 June 1938, Gogebic Lake, 3

feet of water, muck bottom, H. Kilpela (MICH). OSCEOLA CO.--+22 Sept.

1936, T19N R8W Sec. 19, Little Long Lake, 44° 2' N, 85° 20* W, D. S.

Shetter (MICH). OTTAWA CO.— 22 July 1887, Spring Lake, E. A. Fopenoe

(PH 608400). PRESQUE ISLE CO.— 31 July 1940, in water, Black Lake,

H. A. Gleason 9976 (NY). 25 July 1949, marsh along N shore of Swan

Lake, 4 miles E of Rogers City, abundant, flowers blue, with yellow

spot on upper lobe, Rogers McVaugh 11011 (MICH). 9 August 1953, on a marshy point at the mouth of Rainy River, E end of Black Lake, associ­

ated with Acorus calamus, Elzada U. Clover and C. Wood 26261 (MICH).

ROSCOMMON CO.— 28 August 1939, Muskegon River, 0-1 foot of water, muck bottom, D. Shetter (MICH). SHIAWASSEE CO.— +22 July 1941, Hopkins

Lake, 1 (%~1) foot of water, fibrous peat bottom, E. W. Roelofs 320

(MICH). 23 July 1941, Shiwassee River, Corunna, Wendell A. Bush 2

(MICH). ST. CLAIR CO.--24 July 1899, in shallow water about Lake St.

Clair, abundant there in spots, C. K. Dodge (MICH). ST. JOSEPH CO.—

+13 July 1938, Minnewaukon Lake, 2 (2-4) feet of water, fibrous and pulpy peat and rubble bottom, P. Eschmeyer 243 (MICH). 30 July 1953, Portage Lake, C. R. Hanes, S. J. 78 (MICH). VAN BUREN CO.— 25 July

1910, South Haven (Little Bear Lake), very common, C. H. Kauffman

(MICH). +14 August 1913, Sister Lakes, flowers purple, Judge Arthur

W. DeSelm 58 (F 408579). 9 Sept. 1928, Lake Bronson, Lawton, Edgar

Anderson (MO 942491). +7/14 1937, T1-2S R13W, North Lake, sta. no. 1,

1-2 feet of water, 42° 19 f N Lat., 85° 49' W Long., fibrous peat bottom, D. C. Chandler 101 (MICH). +3 July 1941, Lake of the Woods,

1 (0-1) foot of water, marl fibrous peat bottom, E. W. Roelofs (MICH).

WASHTENAW CO.— 13 July 1861, Ann Arbor, purple (MICH). +20 July 1867,

Whitmore Lake, M. W. Harrington (MICH). 6 July 1884, A.A. [Ann Arbor],

Huron river, G. B. Sudworth (MICH). 30 Sept. 1901, Ann Arbor, Huron R. , shallow water, organic soil, F.B.H.B. (OS). June 1917, Huron River,

Ann Arbor, J. H. Ehlers (MO 852245). 11 August 1917, Huron River Ann

Arbor, Beulah Wells (MICH), +4 August 1920, shallow water, Whitmore

Lake, Francis L* Stearns [?] 1^ (MICH). +22 July 1923, shore of

Arnold Lake, Lodi Township, F. G. Gustafson (MICH). 29 August 1926, vicinity of Ypsilanti, Frain's Lake, Nathan A. Harvey (MICH). 6 August

1927, in water at edge of Huron River at Scio, E. W. Erlanson 5288

(MICH, NY). 8 August 1928, banks of Huron River, Ann Arbor, Duke and

Heater 26 (WIS). +3 July 1936, 42° 20' N, 84° 5' W, T2S R3E [?]

Sec. 4, Mill Lake 1-2 feet of water, numerous large patches, C. 0.

Grassi 7262 (MICH). +3 July 1936, 42° 22' N, 84° 5» W, T1S R3E Sec.

22, Green Lake, 2 feet of water, frequent colonies, C. 0. Grassl 7263

(MICH). +15 Oct. 1937, in 12 inches of water at NE corner of Cassidy

Lake, 42° 21' N Lat., 84° 4 f W Long., Carl 0. Grassl 7726 (MICH).

WAYNE CO.— [no date], Detroit, Bigelow (M0 1983379). CO. 7— +[no date], [presumably Ionia Co.— if Wheeler and Smith's Catalog], C. F.

Wheeler (MICH). +July 1891, Eagle L . , E.B.U. (F 260075). +13 July

1916, Bankson [?] Lake, G« Elfrig (F 1562374). +[no day or month]

late summer 1931 [?], northern end of the Lower Peninsula, H. A.

Gleason, Jr. (DUKE 123274). MINNESOTA: AITKEN CO.— August 1921,

Rice Lake, W. F. Kubichek 143 (MO 921189). CARLTON CO.— 23 August

1934, Moose Lake, John B. Mayle and Etlar L. Nielsen 1961 (US 1725244).

CHISAGO CO.— +July 1892, Green Lake, B. C. Taylor (NY, US 33952, WIS).

CROW WING CO.— 5 Sept. 1930, in 1-2 inches of water, mucky bottom, at

edge of marsh, more abundant further out, on N side of Borden Lake,

Garrison Twp., N. Hotchkiss and Percy Jones 4114 (US 1568779).

RAMSEY CO.— August 1890, Ponas, Ex Herb. J. H. Sandberg (F 433286).

WASBASHA CO.— 23 August 1926, shores of Half Moon Lake, Weaver, N. C.

Fassett and N. Hotchkiss 3304 (GH). WASHINGTON CO.— +17 Sept. 1929,

Echo Lake, H. J. Posting (DUKE 09728). MISSISSIPPI: HANCOCK CO.—

29 April 196 7, sandy soil, low mixed hardwoods, along Jordan River S

of Kiln, coastal flatwoods, S. B. Jones 11846 (FSU 113007). HARRISON

CO. — 7/3 1896, Biloxi, S. M. Tracy 3437 (NY). 8-16 1949, Biloxi, low

wet areas or shallow water, E. J. Alexander 28171 (NY). JACKSON CO.—

15 May 1895, Ocean Springs, Miss J. Skehan (MO 1983363, US 308066).

+8-9 1953, shallow water, Orange Grove, P. 0. Kreole, Delzie Demaree

33626 (US 2133797). CO. 7— +August-Sept. 1900, South Pass, the

Mississippi Delta and Adjacent Islands, F. E. Lloyd and S. M. Tracy 248

(NY). +16 July 1932, E. Pascagoula R. [probably Jackson or George Co.],

River bank, F. A. and W. W. Barkley 254 (MO 1099391). MISSOURI: BATES

CO,——1 Oct. 1938, one clump bordering wet mud along branch from artesian well to Prairie Lake, sect. 20, in bottoms of Mariais des

Cygnes River, 1 1/4 miles SW of Rapinsville, Julian A. Steyermark 9975

(F 965281, MO 1169067). BUTLER CO.— 25 June 1934, marsh near Popular

Bluff, James W. Cunningham (MO 1083328). CLAY CO.--+20 June 1897, swamps-local, quite abundant in the swamp at Randolph, Kenneth K .

Makenzje (MO 1983413, NY, US 318333). CRAWFORD CO.— 12 August 1939, margin of lake from Blue Spring branch, 2 miles SE of Bourbon, Julian

A. Steyermark 27912 (F 1065287). GRUNDY CO.— 10 August 1952, common,

"Gooseneck Lake" (natural lake), in bottoms of Thompson River, T60N,

R24W, northwest sect. 34, 6% miles S of Trenton, Julian A. Steyermark

74216 (F 1438758, M0 1983414). JACKSON CO.— +7-8 1884, shallow lakes, a very rare species not known here now, B. F. Bush (MO 1983400).

+9 August 1896, Lake City, common, B. F. Bush 417 (MO 1983415, US

281182). JASPER CO.— 23 August 1873, Joplin, in water, C. Broadhead

(MO 1983416). +1 August 1907, local in slough, near Waco, E. J. Palmer

1087 (MO 762206). +11 August 1907, Waco, slough, E. J. Palmer 1087

(MO 1983399). +15 July 1920, in shallow water, muddy sloughs, Forest

Mill, E. J. Palmer 18330 (NY). LINCOLN CO.— +7 August 1927, Kings

Lake, John H. Kellogg 824 (MO 957101). PIKE CO.— +6 Oct. 1956, natural sink-hole pond surrounded by Quercus palustris, T55N R3W Sect.

28, 5 miles SW of Ashburn, common and dominant with Nelumbo, Julian A .

Steyermark 82860 (MO 1755195). ST. CHARLES CO.— +29 August 1933,

Dardenne Lake near Peruque, Julian A. Steyermark 8887 (MO 1136183).

+7 June 1941, marsh, Douglas E. and Dorothy Wade 2177 (WIS).

ST. LOUIS CO.— 13 Sept. 1885, Natural Bridge Road, St. Louis, Herb, of

Frederick Wlslizenns (MO 1983398). 17 June 1886, Allenton, H. Eggert (MO 758945, US 754847). 17 June 1887, Allenton, H. Eggert (BKL, CAN

165679, F 410593 and 410594, MO 758946). 6-8 1892, St. Louis, Calvary

rd. woods pond and 6-18 1892, Hickory Grove, St. Louis, Herb, of N. M .

Glatfelter (MO 1983401). 10 June 1896, Jefferson Barracks, ponds,

John H. Kellogg (MICH, MO 910841, NY). 7-24 1907, North Amer. Tract,

Chas. Fullgraf (MO 1983419). 28 June 1910, St. Louis, Earl

E. Sherff 114 (GH). August 1911, Lily Pond M.B.G., Moses Craig (MO

1983417). STODDARD CO.— +12 July 1933, sloughs, low woods, E of Fisk,

Ernest J. Palmer and Julian A. Steyermark 41534 (GH, MO 1071214, NY).

CO. ?— +Oct. 1867, dried up swamps, Amenbar C?3 Bottom, St. Louis,

George Engelmann. M.D. (MO 1983384). NEW HAMPSHIRE: BELKNAP CO.—

24 July 1907, shallow water, border of Hill’s Pond, Alton, J. A .

Cushman and S. N. F. Sanford 1341 (NEBC). 11 July 1964, deadwater at end of Lake Waukewan boundary of Center Harbor, New Hampton, G. B .

Rossbach (NEBC). CARROLL CO.— +August 1917, in Lake, Chocorua, Herb.

W. G. Farlow (NEBC). +7-20 1938, Lake Ossipee, Osslpee, W. V. Brown

(DUKE 63379). CHESHIRE CO.— 9 August 1890, Jeffrey, Brook, Walter

Peane (7 703734). 7 Sept. 1890, shore of Gilmore Pond, Jaffrey,

Walter Deane (7 703733). 24 July 1896, Jaffrey, Chriesonhild C?3

Robinson 75 (GH). 20 July 1898, sandy shores of Gilmore Pond,

Jaffrey, B. L. Robinson (NEBC). 8 August 1918, Rlndge, wet ground at edge of mill-stream, Charles Foster Batchelder (NEBC, PH 577885).

4 Sept. 1918, Richmond, shallow water, edge of Wheeler Pond, Charles

Foster Batchelder (NEBC). 6 Sept. 1931, Nelson, shallow water, shore

Rye Pond, Charles Foster Batchelder (NEBC). 25 July 1940, Swanzey

Lake, R. E. Schultes 356 (NEBC). COOS CO.— 27 August 1918, Baker’s Pond, Lancaster, Arthur Stanley Pease 17264 (NEBC). 27 August 1918,

Horseshoe Pond, Northumberland, Arthur Stanley Pease 17309 (NEBC).

+6 August 1952, muddy shore, Pontook Res., Dummer, Arthur Stanley Pease

36817 (NEBC). 25 July 1955, Long Pond, Errol, Arthur Stanley Pease

37985 (NEBC). GRAFTON CO.— 21 July 1923, Squaw Cove, Squam Lake,

Holderness, H. K. Svenson (BKL). 2 Oct. 1931, Mink Brook, Hanover,

John F. Reen (DUKE 24798). 10 July 1933, Foster Pond, Alexandria,

Charles Bullard (NEBC). HILLSBOROUGH CO.— +14 July 1909, Peterboro, shallow water, Nubanusitt River, C. F. Batchelder 2007 (NEBC). 4 August

1917, Merrimack, shallow sandy-bottomed pond, Charles Foster Batch- elder (NEBC). +28 July 1920, Hillsborough, shallow water, North Branch

River, Charles Foster Batchelder (MO 1077710, NEBC). 12 August 1930, muddy t?3 shore of Gumpuz Pond, Pelham, Fred'kS. Beattie (NY).

19 August 1944, ditch by pond, Weare, Clarence H. Knowlton (NEBC).

MERRIMACK CO.— 26 August 1903, edge of pond, East Andover, M. A. Day

(GH). 26 July 1921, Hooksett, shallow pond, Charles Foster Batchelder

(NEBC). 15 August 1932, Tannery brook meadow, Danbury, Charles Bullard

(NEBC). 3 Sept. 1933, Eagle Pond, Wilmot, Charles Bullard (NEBC).

ROCKINGHAM CO.— August Cno day or year}, Greenland, [J. Blake3 484

(US 2590253). 10 Sept. 1916, Hampton Falls, shallow water, edge of pond, Charles Foster Batchelder (NEBC). 19 July 1919, Derry, boggy shore of pond, Charles Foster Batchelder (NEBC). +13 July 1927, Fremont, shallow water, Loon Pond, Charles Foster Batchelder (NEBC). 8 Sept.

1969, Exeter River behind the gym at the Phillips Exeter Academy,

Exeter, R. M. Lowden 32 (OS-6). 8-16 1936, Coffin's Millpond, Hampton, edge of water, Albert Chandler 226 (MO 1882051). 8-16 1936, Hampton, Albert Chandler 1265 (MO 1126962). +21 July 1938, Rye, P. L. Wright

(WIS). STRAFFORD CO.— 8 August 1888, Rochester, presented to NEBC by

Mrs. Edward Bartow (NEBC). 28 August 1938, Strafford, muddy shore of

Wild Goose Pond, A. R. Hodgdon and Edson Chamberlain 2893 (NEBC).

27 June 1946, Strafford Co., Durham, shallow water, oyster River above dam, A. R. Hodgdon, Robert Harrington, and Win. Jahoda 6898 (NEBC).

5 July 1950, Barrington, along Isinglass River near bridge N of East

Barrington, A. R. Hodgdon, Alan and Anthony Hodgdon 6735 (NEBC).

4 July 1970, Bellamy River, Exit 7-Spaulding Turnpike, Strafford Co., freshwater, R. M. Lowden 2 (OS-5). SULLIVAN CO.— 30 August 1962,

Washington, Rte. 31 shallow water at edge of May Pond, M. C. Cowden

(NEBC). NEW JERSEY: ATLANTIC CO.— August 1879, Hammonton, F. L .

Bassett (PH). *20 August 1893, M a y ’s (Mays! Landing, flora of vicinity of Philadelphia, within a radius of 50 miles, Arthur N. Leeds (PH).

4 August 1907, Pleasant Mills, Edwin B. Bartram (PH 648669). 21 August

1910, Pleasant Mills, near mouth of Hammonton Creek, Bayard LonR 4656

(PH). +24 August 1912, bogs and boggy thickets, Sleeper Branch of

Atsion River, near Parkdale, Bayard Long and Francis Pennell 7354 (PH).

+24 June 1922, Cranberry Bog W of Station, H. B. Meredith

(PH). +6 August 1937, alluvial marsh, within 1 mile S along Great Egg

Harbor River, Catawba, Bayard Long 51190 (PH). 22 June 1946, in mud along stream, near May's Landing, Ellys T. and Harold N. Moldenke 18315

(NY 2830). BERGEN CO.— 27 July 1879, Ridgewood, from Dr. W. M . (NY

2577). 17 July 1884, Carlstadt, Albert Niederer (NY). 12 July 1886,

Norwood, J. L. Zabriskle (BKL). 4 July 1893, Harrington Park, C . L.

Pollard (US 309050). 4 August 1905, Tenafly, H. Dautun (BKL). +21 July 1907, Woodridge, Henry Dautun (BKL-2). 22 July 1932, Head of

Overpeck Cr., swamps, Englewood, George Lelderman (WIS). 13 August

1932, Head of Overpeck Cr., swamps, Englewood, George Lelderman 23

(WIS). BURLINGTON CO.— 29 June 1879, marsh, Riverton, H. Russell

Bassler (OS). 21 July 1881, Atsion, Ex. Herb. C. D. Fretz (PH).

6 June 1888, Atsion, Herb. M. E. Leeds (MO 1073628). 3-6 July 1899,

between Tuckerton and Atsion [probably Burlington Co.), C. F. Saunders

and W. N. Clute (PH 504716). 4 July 1910, Medford, Wltmer Stone (PH).

5 Sept. 1910, Hartford, Parker Creek just north of P.R.R., Bayard Long

4922 (PH). 15 July 1911, Bordentown, along Crosswicks Creek, Bayard

Long 6040 (PH). 29 July 1911, Moorestown, North Branch of Pensauken

Creek, Bayard Long 6384 (PH). +30 Jun^ 1912, shallow water, margin of

Batsto River, near site of old furnace, Hampton Furnace, Bayard Long

7783 (PH). 10 August 1914, sandy and muddy tidal shore, W along

Delaware River, Kinkora, Bayard Long 10725 (PH). 27 June 1922, swamp,

Rococos Creek, Mount Holly, H. B. Meredith (PH). 3 July 1922, Mullica

River, near RR bridge, Atsion, Geo. Wm. Bassett (GH, WIS). +14 July

1923, mud along Batsto River, Quaker Bridge, R. R. Drelsbach 1637

(PH). 29 July 1932, edge stream, Cranberry bog, 1% miles SE Atsion,

John M. Fogg, Jr. 4594 (PH). 11 July 1946, border of small marshy pond nearby Mullica River, Crowleytown, 2 miles WNW of Green Bank,

Hollis Koster C9-5-2 (PH). CAMDEN CO.— August 1851, near Camden,

T. F. Seal (?). [no day or month] 1885, Camden Co., presented by

S. S. Garrlgnes (MICH). July 1870, Camden, Ex Herb. Isaac C. Martin- dale (PH). 3 Sept. 1892, Kalghn's pt, Camden, flora of vicinity of

Philadelphia, within a radius of 50 miles, Alexander MacElwee, Jr. (PH 515308 and 515309). 10 July 1901, between Camden and N. Cramer

Hill, C. C. Stewart 2052 (NY 2576). 16 June 1906, Pen[nDsauken, Witmer

Stone (PH). 17 August 1910, Fish House, shore of Delaware River just below P.R.R. Sta., Bayard Long 4525 (PH). 26 July 1911, Delair, shores of Delaware River above P.R.R. Sta., Bayard Long 6299 (PH). +20 July

1922, tidal marsh, YorkBhip, John M. Fogg, Jr. 364 (NY 2503, WIS).

20 August 1922, marshes of Newton Creek, Fairview, Camden, Geo. Wm.

Bassett (GH). 20 Sept* 1922, marshes of Newton Cr. N of Collings Rd., height 6° 2*, Fairview, Camden, Geo. Vfln. Bassett (PH). 9 July 1926, shallow water of tidal marsh, N branch of Newton Creek at A.C.R.R.,

West Collingswood, J. W. Adams 371 (PH). 10 July 1926, shallow water of tidal mud flats, S branch of Pen(n]sauken Creek, just S of Park

Avenue bridge, PentnDsauken, J. W. Adams and Harry W. Trudell 381

(GH, PH). +15 July 1928, roadside bog between Atsion and Parkdale,

William H. Witte (NY). +2 August 1940, peaty bog along Mullica River, ca. 1.5 miles N of Parkdale, Bayard Long 54927 (PH). CAY MAY CO.—

12 June 1899, Race Course Pond, West Cape May, 0. H. Brown (PH).

12 August 1910, West Cape May, around nponds,r N of main road to Cape

May Point, Bayard Long 4439 (PH). 15 July 1922, Cape May Point, salt marsh, along tidal stream, J. M. Fogg, Jr. (PH). 23 July 1940, shagnous and peaty bog, Old Bogs, ca. 1 mile SSW of South Seaville,

Bayard Long 54773 (PH). 6 August 1940, shallow water, Deans Branch of

East Creek, 1.5 miles S of Belleplain, Flora S. Fender 4921 (PH).

CUMBERLAND CO.--19 July 1906, Manumuskin, Pine Barren Region, southern

New Jersey, Witmer Stone 8068 (PH). 12 August 1923, boggy swale, E side of Maurice River at Landis Ave., 2 (7!] m. w. of Vineland, Geo. Vftn. Bassett and B. Long (PH), +19 July 1925, Millville, in pond just over bank of Maurice River, in water about 4-6 inches, black, rich, sandy mud, flowers violet blue, between Silver Run and Milleville,

J. W. Adams (PH), 14 August 1927, peaty margin, Maurice River at

C»R,R. of N.J., 2,5 miles W of Vineland, J. W. Adams 993 (MO 954908),

+6 August 1929, marsh, Dorchester, 0. H, Brown (PH), +13 August 1931, along Muskee Creek, about 1 mile SE of Port Elizabeth, Walter M, Benner

4349 (PH). 18 Sept. 1932, wet, peaty margin of Cumberland Lake,

Manumuskin R., Cumberland, Bayard Long 38515 (PH), ESSEX CO.--+14 Sept.

1895, below Washington Park, flora of vicinity of Philadelphia, within a radius of 50 miles, Albrecht Jahn C?3 (PH). 25 August 1896, Newark, in mud and water, Herb, of Marcus W. Lyon, Jr. (US 1101386). GLOUCESTER-

CAMDEN COS.--8-31 1878, Westville, J. B. Brinton (PH). 29 July 1888,

Westville, J. Bernard Brinton (OS). GLOUCESTER CO.— 2 Sept. 1884,

Bridgeport (MICH, PH). 8-7 1892, Swedesboro, flora of vicinity of

Philadelphia, within a radius of 50 miles, Charles D. Lippencott (GH,

PH). 17 August 1922, border of Mathews Brook, W Woodbury, Bayard Long

26589 (PH). 5 August 1923, on dam of Riders bog h mile S of pond,

Malaga, Geo. Wm. Bassett (PH).25 July 1940, shallow water, margin of pond on Little Ease Run, Franklinville Lake, Franklinville, Bayard Long

54824 (PH). HUNTERDON CO.— 7-24 1897, Stockton, river border, Herb, of

H. L. Fisher (MO 1074001, PH). +28 August 1932, shallow back-water pool along Delaware River, Treasure Island, Bayard Long 38234 (PH).

MERCER CO.— [no date), Princeton (NY). June 1876, Princeton, G.

Maclaskie C?3 (NY). 20 July 1885, Princeton, John E. Peters (PH).

August 1887, near Trenton, L. Sigafoos (PH). MIDDLESEX CO.— 27 August 1885, Woodridge, L. H. Llghthipe (BKL)* 14 July 1892, Milltown, A* P .

Selby (OS). 6/30 1931, Spotsvood, J. A. Drushel 7583 (MO 1031027).

7/19 [?] 1931, Spotswood, J. A. Drushel 7616 (BKL). MONMOUTH CO.—

[no date], Freehold, 0. R. Willis (MICH). MORRIS CO.— (-[no date],

Budds Lake, Black River, R. C. Perry (MO 977002). +8 July 1884,

Hanover, E . J . Fond (US 619915). +18 July 1909, Budd Lake, near outlet,

Harold W. Pretz (PH 555257). OCEAN CO.--24 June 1895, in water with a sandy bottom, Tom's River, Alexander MacElwee, Jr. (PH 515314). 25-31

July 1898, Tom’s River, abundant in slow streams and ponds, Willard N .

Clute 233 (NY). 14 July 1905, New , flora of vicinity of Phila­ delphia, J. Grove (PH). 7 August 1907, Tom’s River, Witroer Stone (PH).

8 June 1946, sphagnoua bog along Oswego River ca. 1 mile N of Warren

Grove, F. C. Schmid, Jr. (PH). SALEM C0.--1 Sept. 1908, Salem, Witmer

Stone 10562 (PH). 30 June 1910, Husted to Vineland, vicinity of

Centerton, Witmer Stone 12546 (PH). 26 July 1922, margin of streamlet,

S tributary to Alloway Creek, Penton, Bayard Long 26305 (PH). 17 August

1922, border of dam SE of Elmer, H. B. Meredith (PH). 25 August 1923, wet, sandy, tidal shore, NW along Oldmans Creek, Auburn, Bayard Long

28933 (PH). 5 August 1924, in swamp, Penn’s Grove, flowers blue,

Harold N. Moldenke 2160 (NY). +3 Oct. 1934, flowers purple-blue, in shallow water, along Game Creek N of Biddles Landing, J. W. and Myrtle

T. Adams 1739 (PH). 23 August 1935, tidal swale along Delaware River, about 0.75 mile S of Penns Grove, John M. Fogg, Jr. 9475 (PH).

21 August 1941, shallow water, Centerton Lake, Centerton, Dorothy and

Douglas Wade (WIS). SUSSEX CO.— =July 1878, Franklin, Henry H. Rusby

2988 (MICH). +12-14 August 1890, Budd’s Lake, John K. Small (F 433868, PH 568263, US 298335). +4 July 1910, Hewitt Pond along L.&H.R.R. about % mile NE of Andover Junction, Harold W. Pretz 2641 (PH 555258).

UNION CO.— +28 July 1902, Springfield, Katharine Meigs (BKL). WARREN

CO.— +15 August 1889, Cedar Lake, Uselma C. Smith 1463 (PH 527497).

+10 August 1910, Foul Rift, mouth of spring E shore of Delaware River,

Charles C. Bachman (PH). 26 August 1949, shore of Pequest Creek, just

E of Belvldere, R. L. Schaeffer, Jr. 32249 (PH). +23 August 1951, swamp % mile SE of Pleasant Valley, R. L. Schaeffer, Jr. 37682 (PH).

+13 August 1957, alluvium %-l ml NW of Columbia, R. L. Schaeffer, Jr.

(PH). CO. ?— +25 August 1877, Squaw Marsh, Miss G. D. Clements (MO

126183). +10 August 1882, 12th st., Ex. Herb. C. D. Fretz (PH).

+13 July 1887, Hospitality Bridge, 8th St., presented through Witmer

Stone (PH). +5 July 1893, ditches, Granton, Wm. M. Van Sickle (BKL).

+4 July 1904, Speedwell, Pine Barren Region, southern New Jersey,

Witmer Stone (PH). +8/16 1925, Goodman, J. A. Drushel 5700 (MO

955609). NEW YORK: ESSEX CO.— 1 August 1921, Lake Harris, Newcomb,

1550 ft. alt., H. D. House 8431 (CAN 165678). 13 August 1927, Lake

Harris, Newcomb, H. D. House 15380 (MO 942658). HAMILTON CO.--3 August

1930, in Brown's Tract at Raquette Lake Station, Indian Lake, Bertha

B. Lambert 5 (PH 701096). LIVINGSTON CO.— 2 Sept. 1926, in narrow lagoon, SW corner of Conesus Lake, P. R. Burkholder and W. C. Muenscher

16485 (GH). MADISON CO.--15 August 1895, South Bay, in shallow water near shore of Oneida Lake, in about 1 foot of water, William R. Maxon

(NY, US 6 74531). CO.— 18 July 1905, New SpringviUe,

Staten Island (Richmond, New York City), Philip Dowell 3900 (MO 703291).

5 August 1905, SW of Jamaica (Queens New York City], Long Island, E. P. Bicknell 252 (PH 736023). ONONDAGA CO.— 21 August 1916, South

Bay, Cicero, K. M. Wlegand 6133 (GH), 18 August 1927, Marcellus, Mill

Dam, Nine Mile Creek, Alfred S. Goodale 52134d (DUKE 66216). ORANGE

CO.— 2 August 1891, showing rootstock, radical leaves and stipules on the basal sheaths, Greenwood Lake, shallow water, Thos. Morong (NY).

+15 Sept. 1895, muddy lake shore, Orange Lake, John Hendley Barnhart

1183 (NY). +2 August 1936, margin of Sucherland Pond, plants of the

Black Rock Forest, Hugh M. Raup 7811 (GH). OSWEGO CO.— August 1869,

Oswego, Wm. A. Buchurt [?] (F 310912). RENSSELAER CO.--[no day or month] 1828-1834, Troy, James Hall (F 476861). ROCKLAND CO.— +30 July

1957, shore Cranberry Pond, Ramapo, Ramapo Township, J. Harry Lehr 20

(NY). ST. LAWRENCE CO.— 14 July 1914, slow streams and ponds,

Gouverneur, Orra Parker Phelps 200 (BKL). 14 July 1914, still water,

Gouverneur, Orra Parker Phelps 295 (GH, NY). SARATOGA CO.— 30 June

1932, in cove along across from Moses Kill, W. C. Muenscher and A. A. Lindsey 3159 (MO 1088204). 22 Sept. 1939, shallow water,

Saratoga lake, 11. D. House 27080 (NY). SENECA CO.— Sept. 1858, ponds,

Ovid, Brewer et Chickerlng (PH 752308). SUFFOLK CO.— 5[?3 July 1873, low grounds, Riverhead, Herb. H. W. Young (F 136821). 26 July 1920, wet shore of Sweezy Pond, Southampton, Harold St. John 2623 (GH, PH

635475). +30 June 1936, shallow water, Peconlc River, Calverton, Long

Island, S. A. Cain (MO 1199360). TOMPKINS CO.— 13 August 1879, Ithaca,

Rutherford P. Hayes 545 (OS). 10 August 1885, Ithaca, Frederick V .

Covllle (US 295553). 11 July 1893, Fall Creek, Cayuga Marshes near

Ithaca, H. von Schrenk (MO 1983340). 15 July 1893, Fall Creek, near the lake, near Ithaca, H. von Schrenk (MO 1983338). 15 August 1893, 80

Fall Creek (near mouth), near Ithaca, Herman von Schrenk (MO 1983341).

21 August 1916, in water bayou near Field Station, bottom land woods along Fall Creek at head of Cayuga Lake, A. J. Eames 6135 (GH). July

1919, Renwick, near inlet, Ithaca, R. Hitchcock and A. R. Bechtel 11725

(MO 979706). ULSTER CO.— 15 July 1910, vicinity of Highland Falls, in the Hudson Highlands, altitude about 50 meters, Edgar A. Mearns 38

(US 647914). WARREN CO.— 20 August 1936, about edge of Lake Kinnear,

Pine Log Camp, NE of Luzerne, Grace M. Tees (PH 737752). WASHINGTON

CO. — Cno day or month] 186 7, East Greenwich, Dr. Asa Fitch (MICH).

+13 August 1914, Tripoli pond, southern W. Fort Ann, Flora of the Lake

George Region, Stewart H. Burnham (GH). 2 August 1919, Middle Gran­ ville, L. M. Dougan (MO 1032869). WAYNE CO.— 16 August 1929, marsh,

Sodus Point, J. L. Lowe (MICH). WESTCHESTER CO.--[no date], White

Plains, 0. R. Willis (NY). 24-7 1874, Scarsdale, (NY). +15 July

1895, water, Ardsley, John Hendley Barnhart 1065 (NY 2535). +2 Sept.

1916, ’’Shadow River" Lake Waccabuc, Francis W. Pennell 8404 (PH 732438).

+15 July 1932, shallow water, mouth of Frouplesome Brook, St. Elenors

Lake, Yonkers, H. A. Gleason, Jr. 1172 (DUKE 123341). CO. ?«=+[no date],

Niagara River, Susan M. Williamson (WIS). +[no day or month] 1845,

New York ?, J. F. Joor (MO 1983368). +11 July 1877, Tuckahoe, E. S .

Burgess (NY). +[no day or month] 1887, Perch Lake, Ex. Herb. H. H .

Keyes (BKL). +July 1887, Onondaga Lake, Ex. Herb. M. L. Overacker

(MO 773123). +July 1887, West Park, C. H. Hall (BKL). +1 July 1887,

Jones Point, Hudson River, Ex. Herb. M. 0. Steele (GH). +August 1887,

Mohigan, J. W. Martens (BKL). +30 June 1889, Lake Mohigan, J. W. M .

[J. W. Martens] (BKL). +July 1889, Mohigan, J. W. M . [J. W. Martens] (BKL). +20 July 1891, Busti, E. M. Wilcox (OS). +26 July 1891,

Rockland Lake, T. M . (NY). +August 1895, Peconic River, Long Island,

H. von Schrenk (MO 1983339). +1 August 1895, Davenport, Herb, of

Wm. L. Fisher (PH 817442). + Cno day or month] 1896, Tusedo Park,

W. H. Lewis, Jr . (NY-2). +July 1899, Renwick, Lake Border, Herb, of

Elizabeth Seelman (BKL). +17 Sept. 1905, Valley Stream, Long Island,

E. P. Bicknell 253 (PH 734944). +27 July 1920, near Yankeetown, edge of swampy lake, Otto Degener 563 (NY). +Summer 1924, Fourth Lake,

Webb Township, Chester H. Myron (DUKE 66217). +22 August 1931,

Oscawana Lake, seed no. 143, P. Wilson (NY-2). NORTH CAROLINA:

BEAUFORT CO.— 6/25 1935, in marsh along Cockold Creek, near Belhaven,

Don Correll 1703 (DUKE 32309). 9 August 1938, marsh at Belhaven,

R. K. Godfrey 4387 (GH, US 176 7456). BERTIE CO.— 7/2 1935, low moist weedy soil, Edenhouse Point, Don Correll 1940 (DUKE 33124). BLADEN

CO.— 21 May 1939, growing in edge of clear lake, sandy bottom, White _ - LJ' V Lake, Francis Harper 949 (PH 846097). BRUNSWICK CO.— 14 July 1926, marsh land just W of Wilmington in the Coastal Plain, A. A. Heller 14067

(MO 932528). 14 August 1930, Southport, H. L. B . (DUKE 09722). 31 May

1938, marsh near Wilmington, R. K. Godfrey and I. V. Shunk 4150 (GH, US

1767267). +29 August 1938, marsh at Lake Waccamaw, R. K. Godfrey 6311

(GH). 2 Sept. 1955, abundant in brackish ditch at Brunswick-New

Hanover Co. line, plants about 5 ft. tall above the water level, corollas a clear sky-blue, flowering spikes erect, fruiting ones re­ curving, Rte. 17, W of Wilmington, R. B. Channel1 and H. F. L. Rock 648

(DUKE 160833, FSU 18539, GH). CAMDEN CO.— 7/2 1935, open marshy soil near Elizabeth City, Don Correll 2078 (DUKE 31991). 7 July 1941, swamp land N of Elizabeth City, Madeline Hill 119 {DUKE 72320). CARTERET CO.—

10 June 1938, marsh, Newport, R. K. Godfrey 4442 (GH, US 1767491).

+11 August 1946, Ellis lake, Coll. Dave Ulmer, Hollis J. Rogers 3413

(DUKE 120599, FSU 23011). 12 August 1946, swamp by Prison Camp 202 at New Port, Hollis J. Rogers 3418 {DUKE 120514). CHATHAM CO. — 28 July

1955, N.C. 1 mile W Moncure behind lockvllle Dam on Deep River, in shallow soil on rock in riverbed, wet, 1. S. Beard 1124 (GH0. +7/2

1935, in water on edge of river, near Edenton, Don Correll 2043 (DUKE

31951). COLUMBUS CO.--8 Oct. 1933, swamp, W of Bolton, H. J. Posting and H. L. Blomqulst 33732 (DUKE 18939). 29 August 1938, marsh, Lake

Waccamaw, R. K. Godfrey 6311 (US 1768411). 18 May 1957, occasional in drainage ditch about 4 miles S of Hallsboro, perianth blue-violet,

R. L. Wilbur 5846 (DUKE 141905). CRAVEN CO.— 23 August 1967, Lake

Ellis at Camp Bryan on the eastern shore of the shallow lake in the southern part of the county just across the Carteret County line, common along shallow, perianth violet, R. L. Wilbur 9536 (DUKE 186126).

DARE CO.— 7-13, 1957, shallow margin of ditch beside US 64, 7.5 miles

E of East Lake, ditch, water 0-3*', pH 6.3, chloride 735 ppm, organic matter .127., measured resistance at 30°C— 2.5 X 10, Ernest 0. Beal 3255

(FSU 60745). FORSYTH CO.--[no date], Salem (PH). [no day or month]

1824, W-S [Winston-Salem], N.C., P. O. Schallert 5144 (DUKE 09724).

GATES CO.--+7/6 1935, wet rich low soil near Raduco, Don Correll 2280

(DUKE 32311). GRANDVILLE CO.— 23 June 1956, infrequent in patches along banks of Tar R. about 6 miles S of Oxford and just below Provi­ dence on Butler Hwy., growing in water in moderate sun, James P.

Gillespie and J. David Gillespie 376 (FSU 29647, GH). 4 July 1956, 83 wet sand bar in Tar River about 10 miles S of Oxford on Hwy. 96, growing In water, James P. Gillespie 403 (FSU 29363). GREENE CO.—

6/21 1935, in water of swamp near Snow Hill, Don Correll 1366 (DUKE

32310). HALIFAX CO.— +7/18 1938, meadow, Winton, R. K. Godfrey 5244

(DUKE 49722). HENDERSON CO.— 8 June 1898, near Hendersonville, A. M .

Huger (NY). HERTFORD CO.— 7/6 1935, in water at river edge near

Winton, Don Correll 2302 (DUKE 32264). 18 July 1938, meadow near

Winton, R. K. Godfrey 5244 (GH). HOKE-SCOTLAND CO. LINE— 6/11 1938,

shallow water, Lumber R., R. K« Godfrey 4591 (DUKE 49575). MARTIN

CO.— 23 June 1967, 4.6 miles NW of Williamston on N.C. #147, near

Conno Creek, scattered plants in wet soil at edge of Conno Creek, petals purple, marked inside with yellow, "thrum"-type flower, R. E.

Weaver 823 (DUKE 183969). NEW HANOVER CO.— August 1925, near Wil­ mington, H. L. B . (DUKE 09723). PASQUOTANK CO.— 3 July 1885, swamps,

Elizabeth City, B. Shimek (US 518191). 24 July 1935, open water,

shallow, swamp about 4 miles N of Elizabeth City, H. J. Posting 35512

(DUKE 31512, NY). PERQUIMANS CO.--+18 June 1927, edge of swamp by road, Winfall, K. M. Wiegand and W. E. Manning 727 (GH). 27 May 1933,

River edge, Hertford, M. Glasson (DUKE 19073). 20 July 1938, marsh, near Hertford, R. K. Godfrey 5332 (GH, US 1768025). ROBESON CO.—

+7/4 1940, swamp, Herb, of P. 0. Schallert (WIS). SCOTLAND CO,--+11

June 1938, shallow water, Lumber River, R. K. Godfrey 4591 (GH). +7/4

1940, swamp, Herb, of P. 0. Schallert (MO 1062964). ROWAN CO. 7— 7/5

1921, border of pond along roadside, Spencer, P. 0. Schallert (GH).

STANLY CO.— 6 July 1956, lake margin, branch of Lake Tillery about 1.5 miles N of Norwood, R. S. Leisner and Harry E. Ahles 16287 (FSU 54145). TRANSYLVANIA CO.— +10 June 1961, Horsepasture Gorge, beside stream,

1/4 mile S of river crossing, C. L. Rodgers 6175b (DUKE 150749).

TYRRELL CO.— 6/29 1935, In water of marsh near Columbia, Don Correll

1886 (DUKE 32236). 9 June 1938, marsh, 15 miles S of Columbia, R. K .

Godfrey 4339 (US 176 7417). VANCE CO.— 1 Oct. 1956, pond shore. Mill

Pond on Weaver Creek, 3.7 miles NNE of Epsom, R. S. Lelsner and Harry

E. Ahles 20418 (FSU 53291). WASHINGTON CO. ?— 28 July 1899, in and along the Roanoke River above Plymouth, Maryland C. Hemmick 3 (BKL,

F 430758, US 343858 and 641366). 26 July 1955, very abundant along mucky margin of Phelps Lake at Pettigrew State Park, perianth dark purple with the two largest lobes with a greenish yellow spot, R. L .

Wilbur 3950 (GH). CO. ?— +[no date), Gates [Gates Co.) and Montgom.

[Montgomery Co.) (PH). + [no day or month) 1885, G. McCarthy (US

221554). +5 June 1896, swamps, Biltmore, Biltmore Herbarium 752 (MO

1983405, US 284287). +22 June 1897, swamps, Biltmore, Biltmore

Herbarium 752b (F 412301, GH, MO 1983404, NY, US 331160, WIS). +3

July 1908, Lake Ellis, [probably Carteret Co.), W. H. Brown 70 (US

512908). +9 July 1919, Albemarle Sound, species examined in osmotic pressure studies by J. Arthur Harris C19360 (GH). OHIO: ASHLAND CO.—

+ 7/4 1911, Round Lake, L. S. Hopkins (OS 45967). ASHTABULA CO.— 8-18

1932, Geneva-on-the-Lake, Lawrence E. Hicks (OS 842290). 7/17 1936,

Geneva-on-the-Lake, Mrs. E. Kennaweg (PH 778734). 27 August 1944,

F. J. Tyler and H. Crofoot (OS 43669). CARROLL CO.— 30 June 1963,

Section 18, Perry Twp., shallow water in SE extension of Leesville

Lake, Erwin M. Herrick (OS 72562). +19 August 1963, Washington Twp., abundant in Still Fork Creek at Specht Marsh, Allison W. Cuslck (OS 73318-2). CUYAHOGA CO.— August 1897, shallow water, Berea, J. R .

Watson (OS 9890). DEFIANCE CO.--=5 Sept. 1898, E. L. Fullmer (OS 9889).

+1 Sept. 1898, E. L. Fullmer (OS 9892). 22 Sept. 1969, Little Pond,

Defiance County, off route 49, R. M. Lowden 34 (OS-4). 22 Sept. 1969,

Big Pond, Defiance County, off route 49, R. M. Lowden 35 (OS-4).

ERIE CO.— 13 August 1894, Vermilion, A. E. Ricksecker and W. M. Dick

(MICH). 13 August 1894, Vermil , A. E. Ricksecker (OS 78314). 8/2

1895, Cedar Point, E. L. Moseley (F 239859). 7/9 probably 1901, Cedar

Point, Mead (OS 45436). 15 July 1913, Cd. Pt. (Cedar Point), E. L .

Fullman (OS 99130). 17 July 1970, Erie Co., one small plant seen in

sand along beach adjacent to water of Carp Pond at E end of pond,

Kelleys Island, Lake Erie, Ronald L. Stuckey 8980 (OS). FRANKLIN CO.—

+31 May 1922, Jackson Twp., John H. Schaffner (OS 9888). FULTON CO.—

+29 August 1893, S.E., J. S. Hlne (OS 9887). GEAUGA CO.--+July 1895,

Newberry, J. R. Watson (OS 9886). +1 July 1895, Berea, J. R. Watson

(WIS). HAMILTON CO.— +9 August 1901, Cedar Pt., Chas. S. Mead (OS

9885). HURON CO.— +3 August 1894, Vermilion, Alfred E. Ricksecker

(US 217577). LAKE CO.--11 August 1889, Fainesville, Ex Herb. Wm. C.

Werner 3074 (OS 9884). 18 August 1924, Mentor Marsh, Roscoe J. Webb

5464 (GH). LICKING CO.— 11 July 1895, Licking Reservoir, [Buckeye

Lake, Probably Licking Co.3, E. E. Bogue 1690 (OS 87174). 22 July

1897, Licking Reservoir, W. A. Kellerman, K. F. Kellerman, and W. R .

Beatty (OS 9883). 25 July 1897, Licking Reservoir, W. A. Kellerman

and K. F. Kellerman (OS 9882). July 1934, Buckeye Lake, K . Boyd (NY

2264). 12 Oct. 1969, Cranberry Bog, Buckeye Lake, R. M. Lowden 33

(OS-4). LORAIN CO.— +23 July 1900, swamps, Oak Point, A. Monosmlth 86

(OS 9881, WIS). +31 August 1965, at edge of marsh Oak Point, Black

River Twp., George T. Jones (OS 78329). LUCAS CO.— July 1878, Toledo,

J. F. James (OS 9880). 27 August 1968, common in shallow water,

Magee Marsh, Crane Creek State Park, Sec. 12, SE corner of Jerusalem

Twp., ca. 10 miles E of the city of Oregon, Ronald L. Stuckey 7531

(OS 90569). MEDINA CO.— 19 July 1903, Chippewa Lake, V. Sterks (OS

46054). 28 June 1911, Chippewa Lake, L. S. Hopkins (OS 77062).

OTTAWA CO.— [no day or month] 1890, Lakeside, A. P. Selby (OS 87172).

July 1897, Lakeside, J. D. Simklns by W. G. Sanger (OS 75938). 28 June

1937, East Harbor, Ellis Ray Shilling, Jr. (FTSL). 5 July 1939, East

Harbor, Earl L. Core 7303 (FTSL). 17 July 1946, Haunck's Pond, Middle

Bass, H. E. Simmons (FTSL-2). 7/4 1950, in water (8”) of a pond,

Catawba Island, D. Seaman (FTSL). 6-22 1957, Haunck's Pond, Middle

Bass Is., watery muddy bottom, R. Cruden 31 (FTSL). 7/10 1959, IV water l^s feet deep, Haunck's Pond, Middle Bass, T. R. Fisher 1823

(FTSL). 27 July 1961, dense spike subtended by spathe-like bract,

Haunck's pond, Middle Bass Island, Thelma D'Almalne 20 (OS 101758).

6/25 1963, Middle Bass Island, Haunck's Pond, growing on the edge of the mud flat, quite abundant, group IV, HP6IV (FTSL). 27 June 1963,

Middle Bass Island, Haunck's Pond, Yard one of trax., mudflat, group

III, H.P.l (FTSL). 30 July 1963, Ottawa County, Middle Bass Island,

Haunck's Pond, SW corner on the mud flat, R. M. Lowden 46 (OS). 27

July 1965, Haunck's Pond, Middle Bass Island, in 12 inches of water in center of pond, abundant, Duke W. Wood 255 (FTSL). 27 June 1966,

Winous Point, 3h miles SW of Port Clinton, off St. Rt. 53, SE h

Sec. 22, Bay Twp., aquatic-swampy area, Richard M. Lowden 564 (OS). 29 July 1968, a few plants on mat floating between open water zone and shrub zone at East Harbor State Park, Ronald L. Stuckey 7179 (OS 90946),

27 June 1969,' Ottawa County, Vinous Pt., 3% miles SW of Port Clinton off State Route 53, Cent, SE \ sec, 22, Bay Township, R. M, Lowden 37

(OS-16), 10 August 1969, Ottawa County, Winous Pt., 3% miles SW of

Port Clinton off state route 53, cent. SE \ sec. 22, Bay Township,

R. M. Lowden 36 (OS-16). 13 July 1970, Haunck's Pond, Middle Bass

Island, about 20 plants along N gravel bank in about 12" of water, mixed with Nuphar advena, Marvin L. Roberts 553 (OS). PERRY*C0.—

+26 July 1895, Reservoir Park, W. A. Kellerman (OS 9879). PORTAGE

CO. — [no date] Brady's Lake, Wm. Krebs (OS 9878). 24 July 1905,

Cuyahoga R. at Hiram Rapids, Roscoe J. Webb 528 (OS 74447) and 779

(GH). 8 Sept. 1970, Portage Co., flowers light violet, common in . shallow water, muddy bottom along shore of Twin l.akes in the village of Twin Lakes, Franklin Twp,, Robert R. Haynes 3447 (OS). STARK CO.—

[no date], dans les fosses et lacs pres Canton, 69, Nicholas Riehl

Herbarium (MO 1983375). July 1835, Lakes, Canton, Riehl (MO 1983374),

+28 June 1898, Congress Lake, W. A. and K. F. Kellerman (OS 9877).

+4 July 1899, Congress Lake, A. D. Selby 1691 (OS 87171). SUMMIT CO.—

+26 June 1955, Twinsbury Twp., Tinker's Creek Valley flood plain and higher ground adjacent, 3/4 to 1^ miles SE of Twinsburg, Ervin M .

Herrick (OS 54720). WAYNE CO.--+5 July 1899, Brown’s Lake, A. D .

Selby and J. W. T. Duvel 399 (OS 87177). +21 July 1899, Doner's Lake,

A. D. Selby 401 (OS 9874 and 87178). +7 Sept. 1899, Brown's Lake,

J. W. T. Duvet 400 (OS 87176) +29 August 1901, Fox Lake, J. R. Watson

2648 (OS 87173). WILLIAMS CO.— 8 August 1899, Nettle Lake, A, D . Selby 1692 (OS 87175). July 1932* Leslie L. Pontius and Floyd Bartley

(US 1630451). 21 July 1932, Nettle Lake, Bartley and Pontius (OS 9873)

+18 Sept. 1967, NE h Sec. 33, T9S, R4W, Northwest Twp., occasional In

shallow water at SE edge of Mud Lake Bog, ca. 1% miles SW of Cooney,

Ronald L. Stuckey 6100 (OS 87407). 16 Sept. 1969, occasional in clumps in shallow water along shore of Nettle Lake, NE % Sec. 23, T9S, R1E,

Northwest Twp., ca. 9 miles W of Pioneer, Ronald L. Stuckey 8556 (OS).

CO, ?— +8/1 1891, Put-in-Bay and Detroit, The N. M. Glafelter Her­ barium (MO 1983378). OKLAHOMA: CHEROKEE CO.--21 June 1936, water 2 feet deep, fish hatchery no. 3, Talequah CTahlequah], James de Gruchy

135 (GH, NY). McCURTAIN CO.--4 June 1937, in water, sandy clay,

Yashaw Creek, Broken Bow, U. T. Waterfall (NY). 4 June 1937, Yashaw

Creek, 3% miles S of Broken Bow, U. T. Waterfall 669 (NY 2603). 23

June 1948, swamp, Grassy Lake, 4 miles E and 4 S of Tom, U. T. Water­ fall 8024 (PH 852519). 6 June 1949, mud flats, edge of Grassy Lake,

4 miles S and 4 miles E of Tom, U. T. Waterfall 8883 (GH). OTTAWA CO.-

+27 August 1913, at lower margin of erect vegetation, border of pond, near Ottawa, G. W. Stevens 2448 (GH). PENNSYLVANIA: BERKS CO.— +24

Sept. 1939, along stream in pasture 3/4 mile E of Kenneys, Hans

Wilkens 6064 (PH). BUCKS CO.— 22 July 1876, Neshaminy Creek, Bridge

Point [*=Edison], H. B. Meredith (PH). August 1882, Rockhill, Ex. Herb.

C. D. Fretz (PH). August 1882, near New Hope, Sarah E. Paxon (PH).

+[no day or month] 1886, Tohickon Creek, Mrs. 0. U. Melcher (PH),

July 1893, Neshaminy, Agnes Woodman (MO 1983350). +30 July 1901,

Delaware River near Kintnersville, J. A. and H. F. Ruth (PH). +6

August 1910, shores of Tohickon Creek, Weisel, Walter M. Benner 280 (PH). 19 August 1910, shallow water, Little Neshaminy Creek, Bayard

Long 4589 (PH). 26 August 1921, Delaware River, near upper bridge,

Morrisville, Walter M. Benner 1748 (PH). +1 July 1922, Tohickon Park,

in Tolickon Creek, Ex. Herb. C. D. Fretz (PH). +8 July 1922, in swamp

along road leading N from Diehl’s Pottery, N.W. of Rockhill Station,

East Rockhill Township, Walter M. Benner 1954 (PH). +14 August 1923,

shore of Delaware River, Point Pleasant, Walter M. Benner 2311 (PH).

+25 July 1924, water in creek, Bristol, R. R. Drelsbach 3065 (PH).

+24 Sept. 1932, sandy muddy tidal shore, s.e. along Delaware River,

Eddington, Bayard Long 38531 (Pll). CRAWFORD CO. — [no day or month]

1891, Oil Creek, Thos. C. Porter (? 703739). DELAWARE CO. — [no date],

Read, Chipa, [Chester, Pa.] (PH). +11 July [no year], T [?] lnicum,

in water, Benjamin H. Smith (PH). «7-29 1865, Delaware River Bank,

near Del. State Line, C*Mareus Hook], Herb, of William Trimble (PH).

8 August 1891, Ridley, marshes near Darby Creek Sta., Benjamin H. Smith

(PH). +5 July 1946, muddy border of Henry's Pond by Plush Mills Road,

Swarthmore, Samuel C. Palmer (PH). ERIE CO.— 26 June, July 9 1879,

Presque Isl., [G. Guttenberg, Coll.] (? 704706). 7-12 1928, Cranberry

Pond, Presque Isle, W. R. Witz (NY). 6-26 1941, Pond on Presque Isle,

W. R. Witz 912 (PH 844517). +18 August 1948, In shallow water along

shore, Pleasant Lake, E. F. Westlake, B. F. Donley and H. A. Wahl 6101

(GH). LANCASTER CO.— 6 June 1859, Susquehanna, Safe Harbor, T. C .

Porter (PH). August 1890, Peach Bottom, A. F . Eby (MO 1983345).

August 1891, Peach Bottom, A. F. Eby (MO 1983344). 22 July 1894,

Peach Bottom, A. F. Eby (M0 1983346). +22 August 1894, Fites Eddy,

A. F. Eby (MO 1983343). +5 August 1903, M Falls Ferry, J. J. Carter (PH). LEHIGH CO.— +6 August i860, Lehigh [probably in the Lehigh River

above Easton], Herb, of Thomas C. Porter (?). +13 July 1868, Chain Dam

in Lehigh, Thos. C. Porter 490 (?). +18 July 1899, Island Park, in

Lehigh, above Easton, Thos. C. Porter (?). +15 August 1908, below

Rockdale, along Lehigh River, Harold W. Pretz 1459 (PH). +25 July

1909, Rockdale, Backwater's of Lehigh R., above, Bayard Long (PH).

+28 July 1912, Lagoon, opposite Lehigh Gaf Station, along Lehigh River,

Harold W. Fretz 4898 (PH). 19 July 1913, open marsh 7/8 mile NW of

Rockdale, along Lehigh River, Harold W. Pretz 5869 (PH). LUZERNE CO.—

+15 August 1889, Lily Lake, A. A. Heller (F 396268). 16 August 1889,

Lily Lake, John K. Small (F 177891). +9 August 1921, North Pond,

Sweet Valley, Ross Twp., H. B. MeredLth (PH 585927). MONROE CO.—

[no date], Long Pond, Pocono (PH), [no day or month] 1860, Long Pond,

Pocono, (? 704704). 31 July 1860, Pocono Mountain, Traill Green (F

63427). 25 July 1904, Long Pond, plants of the Pocono Plateau, John

W. Hershberger (F 189255, GH, MO 1983347, Ph 525013). MONTGOMERY CO.—

+12 Sept. 1943, shallow water of old canal by Schuylkill River below

Ford Street, Bridgeport, Bayard Long 60346 (PH). NORTHAMPTON CO.—

[no day or month] 1829 [?], Hellerst. [HellertownJ, Detwlller Her­

barium (PH). +1 August 1953, alluvium 1 mile NW of Riverton, R. L .

Schaeffer, Jr. 44545 (PH). PHILADELPHIA CO. — [no date], Philada.,

The Bernhardt Herbarium (MO 1983349). [no day or month] 1832, in Lecha

fluvio prope Philadelphiam, Unio itiner, C. J. Moser (MO 1983348).

8 July 1869, shores of Schuylkill R* above Phila., J.H.R. 8465 (MO

1983351). 1 July 1888, in water west bank of Schuylkill river, below

south St. Bridge Philadelphia, Herb, of Alex. McElwee 302 (PH 515524). 20 July 1889, near Philadelphia, J. M. Greenman 1268 (MO 738348).

15 August 1894, in ditches, in the "Neck", Gibson's Pt., Phila., flora of vicinity of Philadelphia, within a radius of 50 miles, Alexander

MacElwee, Jr. (PH 515310). 26 August 1936, shore of Delaware River near mouth of Pennypack Creek, Holmesburg, Walter M. Benner 7333 (PH).

SUSQUEHANNA CO,— 13 Sept. 1939, edge of Kenny's Pond 2 miles N of West

Auburn, H. A. Wahl 460 (GH). WAYNE CO.— 17-19 June 1921, South Sterling,

Alexander MacElwee (PH 636285). YORK CO.— 7 July 1890, 2 miles N of

Wrightsville, John K. Small (F 178331). +7-30 1893, York Furnace,

Herb. M. E. Leeds (PH 700963). +28 June 1899, Kevinski Lake, York

Furnace, Alex. MacElwee 758 (GH, NY, PH). 21 July 1906, Peach Bottom,

J. J. Carter (PH). CO. ?— +8 July 1902, Darby Creek [probably

Delaware Co.3, C. C. Stewart 2488 (NY 2576). +7 Sept. 1910, border of Mountain Lake, Burlington, Herb, of S. Stetson (NY). +7-10 1914,

Essington, William Randolph Taylor (PH). +22 June 1933, pools, Morris

Farm, Bristol [probably Bucks Co.], J. M. Greenman 4822 (MO 1063977).

RHODE ISLAND: NEWPORT CO.— 21 August 1913, shallow water, Fresh Pond,

Block Island, M. L. Fernald, F. W. Hunnewell, 2d, and Bayard Long 9138

(PH 551152). 1 0 [Oct.] 1916, Block Island, Gravatt (US 1118368).

16 July 1932, in shallow water, Tiverton, Sawdy Pond, S. N. F. Sanford

(NEBC). PROVIDENCE CO.--+[no day or month] 1840, Blackstone R.,

(NEBC). +July 1884, Berkeley, John F. Collins (NEBC). 7 July 1892,

Spectacle Pond, Cranston, C. A. Regester and J. F. Collins (GH, NEBC).

+25 August 1893, Mashapaug Pond, Elmwood, J. F. Collins (NEBC-2).

+31 July 1908, Rafer Williams Park, T. Hope (NEBC). WASHINGTON CO.—

+30 August 1919, seedlings, Moscow Pond, Hopkinton, Washington Co., R. W. Woodward (NEBC). +31 July 1947, at edge of small stream, Exeter, corolla blue, Harold N» Moldenke 19046 (CAN 198766, NY). SOUTH

CAROLINA: ANDERSON CO.— 17 July 1919, Piedmont, river banks, Rev. John

Davis (BKL, MO 900034 and 1025123). BERKELEY CO.— 28 May 1957, Crawl

Creek on SC. 45, east of Pineville, John G. Haesloop and Harry E. Ahlea

26651 (NY). CHARLESTON CO.— +2 August 1915, Blake Marsh, C. P .

Alexander 56 (US 910388). CLARENDON CO.— +11 May 1917, Santee River

Bottom, W of St. Paul, Wltmer Stone 607 (PI1 571462). COLLETON CO.—

+8 Sept. 1915, Ashepoo River, C. P. Alexander 253 (US 910454). GEORGE­

TOWN CO.--23 August 1915, Hobcaw Plantation, Georgetown, C. P.

Alexander 170 (US 910429). 6/14 1936, open, wet woods in savannah about 3 miles N of Plantersville, D. S. Correll 5305 (DUKE 135374).

9 July 1939, marsh, Waccamaw River, Georgetown, R. K. Godfrey and

R. M. Tryon, Jr. 348 (DUKE 58161, F 1602502, GH, MO 1186111,. US

1837248). GREENVILLE CO.— 17 July 1919, Piedmont, Rev. John Davis (MO

1023332 and 1073742). HORRY CO.--25 May 1957, wet ditch by U.S. Rt.

501, 10 miles NW of Myrtle Beach, C. Ritchie Bell 7792 (NY). ORANGE­

BURG CO.— 8 May 1907, swamp along Edisto Riv., Orangeburg, H. D. House

3332 (NY). CO. ?— +[no date], Rev. J. Backman, Charleston, S. Carolina,

C. W. Short Herbarium (PH 783879). [no day or month] 1889, Miss

Plckney (NY). TENNESSEE: GRUNDY CO.— 27 April 1936, shallow water of

Goose Pond, near Pelham, alt. 1000 ft., H. K. Svenaon 7610 (BKL).

23 June 1938, shallow water of Goose Pond, Pelham, H. K. Svenaon 9112

(BKL). CO. ?— +June 1896, marsh at Wolf Creek, E. Tenn., A. Ruth 1048

(NY). +June 1896, swamp near Wolf Creek, E. Tenn., A. Ruth (MO 1983381).

+June 1932, Reelfoot Lake at Walnut Log Lodge, Clarence E. Moore B4 (MO 1042636), TEXAS: BEXAR CO. — Cno date], "Plentiful", San Antonio,

E. H, Wilkinson 127 (MO 1983425), Cno day or month] 1897, San Antonio

E. H. Wilkinson 16 (MO 1983426), 3 Oct, 1900, San Antonio, common along the river, B, F. Bush 1258 (MO 1983427), Cno day or month]

1900-1902, near San Antonio, E. H, Wilkinson (MO 1983424). Cno day or month] 1904, Gust. Jermy (NY). 1 June 1911, tropical life zone, San

Antonio, Mr. & Mrs. J. Clemens 130 (MO 808 724). 12 April 1938, in shallow water, I. W. C. campus, San Antonio, Sister Mary Clare Metz

(NY). HARRIS CO.--Oct. 1842, Houston, F. Llndhelmer (GH-2). +Oct.

1842, in swamps, Houston, etc. (MO 1983433). ROBERTSON CO.— +16 June

1943, in stream on bog near Newbaden, Fred A. Barkley 13029 (MO

1271889, US 1870412). WALLER CO.--8 June 1872, ponds, Hempstead,

Elihu Hall 668 (F 454597, MO 1983462, NY-2, US 33955). CO. 7— +Cno date], San Antonio River, Gust. Jermy (MO 1983422, NY). +Cno date],

Mr. Charles Wright (GH). +Cno date 7], in swamps (MO 1983432). +4

July Cno year], pond near College, J. Reverchon (MO 1983437). +Cno date], S. Antonio River, G. Jermy bought 1904 (MO 1983434). [no date]

Berlandier 2516 Herb. Berlandierum Texano-Mexicanum (MO 1983429).

+Cno date], Herbarium Berlandierianum Texano-Mexicanum, Berlandier

1086 (MO 1983428). +April 1834, fl. violacei, De S. Patricis a Goliar

2516=1086, Herbarium Berlandierianum Texano-Mexicanum 1086 (GH).

+Aprll 1834, fl. violacei, De S. Patricis a Goliar (PH 783936). +July

1842, Lynchburg, F. Lindheimer (MO 1983431). +[no day or month] 1890,

Hockley, W. F. Thurrow (F 375194). +9-6 1907, swamp, Sheldon,

J. Reverchon (MO 1983436). +1 August 1920, San Antonio River, Miss

Ellen D. Schulz 191 (US 1103400). VERMONT: ESSEX CO.— 2-12 August

1899, Brunswick, Willard W. Eggleston 1647 (GH). 30 August 1940, Bow Fond, Guildhall, Arthur Stanley Pease 28364 (NEBC). RUTLAND CO.— 7

August 1909, Rutland, Herb. Elroy C. Kent, G. L. K . (NEBC). Sept. 1920,

Lake St. Catharine Wells, P. L. Dutton and C. W. Dodge (NEBC). 4 Sept.

1920, shallow water Lake St. Catherine Wells, D. Lewis Dutton (GH, MO

865499). 11 July 1941, Hortonia Lake, shallow water, Hubbardton,

Clarence H. Knowlton (NEBC). TRAVIS CO. ?— 19 August 1943, small pond,

M. Harpin 44h (MO 1273546). WINDHAM CO.— +[no day or month] 1848,

Woodstock, bequest of Thomas Hope (NEBC). 26 July 1910, Jamaica, flora of West River Valley and tributaries, Cold Pond, Leston A .

Wheeler (NEBC). 30 July 1921, flora of West River Valley Vt. and tributaries, Athens, Lily Pond, F. T. Randall (NEBC). WINDSOR CO.—

29 August 1935, in still water at edge of pond, Tyson, flowers blue,

Harold N. Moldenke 8779 (MO 1202830, NY). CO. ?— +25 August 1880,

Colchester Pond, Herb. F. H. Hosford (F 261869). +30 July 1899, North

Hero, plants of Western Vermont, Ezra Bralnerd (GH). VIRGINIA: FAIR­

FAX CO.— 7 August 1910, swamp, near Great Falls, Philip Dowell 6435 (GH).

GLOUCESTER CO.— 5 July 1924, swamp-Beaver Swamp Creek, 1 mile N

Gloucester, K. B. Meredith (PH 624096 and 624141). GREENSVILLE CO.—

+12-13 June 1939, wooded bottomland of Fontaine Creek SE of Taylor's

Millpond, M. L. Fernald and Bayard Long 10183 (GH, PH). HANOVER CO.—

Cno date], Richmond, DeChalmot (US 367286). 17 June 1922, wet mucky soil, Chickahomlny Swamp, 6 miles N of Richmond, L. F. Randolph and P.

Merriman 249 (GH). ISLE OF WIGHT CO. — 7-28 June 1893, near Franklin, plants of southeastern Virginia, A. Arthur Heller 957 (F 352611 and

430418, GH, MO 1983356, NY-3, PH, US 33950 and 937957). JAMES CITY

CO.— +24 June 1939, marsh by Powhatan Creek, J. X. Baldwin, Jr. 309 (GH, MICH). NANSIMONI) CO.— 19 June 1938, margin of Creek, wooded bottomland of Somerton Creek, near Factory Hill, M. L. Fernald and

Bayard Long 8184 (GH, PH). 20 July 1938, margin of Somerton Creek, near Factory Hill, M. L. Fernald and Bayard Long 8646 (BKL, F 1075875,

GH, MO 1253027, NY, PH, US 1810315). 11 June 1944, swamp, 4 miles W of Suffold, Leslie Hubricht B2486 (MO 1281717). NORFOLK CO.— 2 July

1892, near Suffolk, in the vicinity of Norfolk, N. L. and Elizabeth G.

Britton and Anna M. Vail (NY). 9 July 1898, near Northwest, Thos. H.

Kearney, Jr. 1551 (US 356241). 14 Sept. 1945, fresh tidal reed-marsh along Northwest River below Northwest, M. L. Fernald and Bayard Long

14925 (PH, US 2003552). +22 Sept. 1933, edge of brackish marsh by

Blackwater R . , Pungo [?] Ferry, M. L. Fernald and Ludlow Griscom 2806

(GH). SOUTHHAMPTON CO.— 9-29 June 1893, about Franklin, A. Arthur

Heller 957 (OS). 22-23 June 1936, sandy alluvial bottomlands of Three

Creek, Drewryville, M. L. Fernald, Bayard Long and R. F. Smart 5702

(GH, NY, PH). +19 June 1938, sandy shore of Blackwater River, NE of

Smith's Ferry, at Bailey's Seine Beach, M. L. Fernald and Bayard Long

8645 (F 1075881, GH, NY, PH, US 1810316) and 19 July (MO 1253026).

CO. 7--+19 July 1904, Slough, Four-Mile Run, flora of the Potomac

Valley, Agnes Chase 2497 (F 167509). +28 August 1904, shallow water,

Hunting Creek, near New Alexandria, H. D. House 420 (NY). +20 August

1915, Chopa Wausic Creek, between the Nelumbo and Sclrpus colonies,

I. Tidestrom 7588 (GH, US 1736237). +26 August 1915, Point Micon

Reach, between Zlzanla and Nelumbo colonies, I. Tidestrom 7682 (GH).

+25 July 1940, F-4, in swamp near White Hill Lake, with red maple, gum, et al., almost fall light, occasional here, S. C. Ryan and Frank E. 96

Egler AO-189 (NY 2258). WISCONSIN: ADAMS CO.— +26 July 1939, Strongs

Prairie, T18N R4E Sec. 3, the soil Is sandy, growing In Wisconsin River

bottom land, which is subject to occasional inundation, generally In

the spring, to a depth of 4 ’ to 8', growing in low sloughs in 1* to

1%' of water, extremely mucky bottoms, from 3' to 3 V long, Robert p .

McIntosh C-633 (F 1564398, MICH). ASHLAND CO.— +17 July 1896, Lake

Superior region, near Kakagen River, L. S. Cheney (WIS). 16 July 1936,

Ashland Co., Torrey L., Morse, Louis Knowlton 97 (WIS). 23 Sept. 1952,

Lincoln Co., in water about 3 dm deep, sandy bottom Muskalunge

CMuskellungeD Lake, T35N, R6E, sec. 12, tn Bradley, Frank C. Seymour

15019 (WIS). BAYFIELD CO. — 3 Sept. 1959, shore of East portion of Lake

Namekagon by Hy. D, 2 miles NW of Namekagon, NW corner of S. 11, T43N

R6W Sect. 11, P. Weber, D. Ugent and J. Zimmerman (WIS-2). P'JFFALO . . -

6 August 1926, shallow water, pool in river bottoms, Nelson, N. C .

Fassett and N. Hotchkiss 3209 (GH, NY, WIS). BURNETT CO.— 16 August

1929, near shore, Oak Lake, Sect. 20 T40N R14W, Gaslyn, W. T »

McLaughlin 642 (WIS). 19 August 1929, sandy shore Corner's Lake,

Sect. 35 T40N R16W, Yellow Lake, W. T. McLaughlin 647 (WIS). 20 August

1929, marahy shore, Goose Lake, Sect. 11 T40N R14W, Birch Island,

W. T. McLaughlin 645 (WIS). 21 August 1929, grass invaded sand shore,

Bass Lake, Sect. 25 T38N R15W, Coomer, W. T. McLaughlin 646 (WIS). 24

August 1929, off sandy shore, Oak Lake, Sect. 20 T40N R14W, Gaslyn,

W. T. McLaughlin 641 (WIS). +30 August 1930, Mud Hen Lake, H. J.

Posting 3066 (DUKE 09729). 6 August 1945, in shallow water in Lang

Lake, NE of Webster, T40N R15W Sec. 16, J. W. Thomson (WIS).

CHIPPEWA CO.— 1 Oct. 1949, 15 miles N of Bloomer, shallow water, Turk Lake, T32N R9W, N. C. Fassett and J. C. Budd (WIS). 24 July 1950,

Chick Lake, T32N R9W near New Auburn, William T. Helm (WIS). +27 June

1959, Lake Larabee, large rich roggy lake shore, Sphagnum, Serracenia, orchids, many Ericaceae, a few Tamaracks, water with Saglttaria,

Nymphea, Nuphar, T32N R9W Sect. 24, NW %, J. Patraan, R« Long and G .

Strulk (WIS). +2 August 1959, occasional, none flowering, Cather Lake,

T32N R9W Sect. 3, SW T. R. Swan 89 (WIS). +3 August 1959, abundant, much in flower, Skaw Lake, T32N R9W Sect. 11, SW%, F ■ R . Swan 84 (WIS).

+9 July 1960, emergent, 1* of water, thick silt bottom, T32N R9W Sect.

11, SW i, Skow Lake, T. A. Ebert (WIS). 24 June 1965, in water along the shore of Long Lake, purple flowers, Mollie Harker and Betsy Paddock

(WIS). DANE CO.— [no date], Madison, T. J. Hale (WIS). July [no day or year], Madison, L. Wingra, Herb. W. H. Chase (WIS). 15 August 1876,

Madison, L, Wingra, W. H. Chase (WIS). 1 August 1888, margins of shallow ponds, Madison, L. S. Cherrey (WIS). 24 July 1945, Arboretum

University of Wisconsin, section 41 (arboretum map), R. A. McCabe (WIS).

FOREST CO.--6 July 1930, Htles, Katherine P. Smith (WIS). +2 Sept.

1956, along creek road through Spruce-Fir-Hemlock-Thuja woods, near shores of Julia Lake, T38N R12E Sect. 6 , E. S. Bradford's property,

Katharine Snell (WIS). 8/8 1963, Pine Lake 10 yds. of NE shore 3-4 feet of water, approachable from federal boat landing forest road 2185,

Nicolet National Forest, T37N R12E Sect. 16, Gerald F . Levy (WIS).

GRANT CO.— 5 Sept. 1930, in mud, Wisconsin River bottoms, Sect. 14

T6N R6W, opposite Bridgeport, N. C. Fassett 13477 (WIS). IRON CO.—

10 August 1964, in a few inches of water, mainly Band, silted over with thin muck layer, in Channel between 1st Black Lake and Lake of The Falls off cty. trunk FF, 6 miles W of Mercer, T43N R2E Sect. 36, Stephen R .

Meyer 45 (WIS). JEFFERSON CO.— +15 August 1969, emergent aquatic in

warm shallow water of Goose Lake, with Utricularla vulgaris, T7N R16E

Sect. 22, Bill Tans 149 (WIS). JUNEAU CO.— 3 August 1891, Camp

Douglas, banks of Lemonweir River, Edgar A. Mearns (GH, US 670830).

+8/17 1938, Miller Ditch, T18N R3E Sect. 14, John Catenhusen (WIS).

LaCROSSE CO.— 10 July 1956, marshy shore of Lake Onalaska, T16N R7

Sect. 6, Onalaska Twp., Thomas G. Hartley 1345 (WIS). +July 1957,

near North Bend, Twon Farmington, Alvin M. Peterson 324 (WIS).

LANGLADE CO.— +10 July 1904, P. H. Dunehl [?] (WIS). LINCOLN CO.—

+16 July 1950, Crystal Lake, T35N R6E Sec. 11, twp. Bradley, Albert

Lerch (WIS). 19 August 1956, mudbanks of drained river bed and river,

Prairie River just outside of Merrill, N of city limits on Hy. 51,

T32N R7E S. 6, Margaret M. Iwen 322 (WIS). +5 August 1962, Red maple,

poplar, birch stand on the E bank of the Wisconsin River, along H'wy

107, 5-6 miles S of Gilbert and junct. with H'wy 51, T33N R6E Sect. 3,

G. Levy and F. Iltls (WIS). MARQUETTE CO.— +4 Sept. 1962, Black

Spruce and Tamarack Swamp, flora of the old bed of glacial Lake

Wisconsin and the adjacent terminal moraine, Sec. 33 T16N R8E, Paul D .

Sorensen 4744 (WIS). ONEIDA CO.— 18 July 1914, shallow water of

Norway Pt., Pelican Lake, Roland E. Kremers (WIS). 13 Sept. 1927, in

0.5 m water, sphagnous shore of Hill Lake, Minocqua, Sect. 9 T39N R6E,

N. C. Fassett 5359 (WIS). +8-12 1930, Garth Lake, still water, Nellie

V. Haynie 1899 (F 1525289). 15 August 1930, Isle-o'Dreams, Round Lake,

Three Lakes, Nellie G. Smith 153 (WIS). 8-13 1942, Birch Lake, near

Minocqua, E. Sella (F 1412214, 1412181, 1520948 and 1520960). +20 July 1947, water 6-18’* deep at edge of Clear Lake, Sec. 17 T39N R7E, Philip

B. Whitford 1910 (WIS). +29 August 1957, Lake Thompson along shore,

mostly In 2-3' water, T36N R9E Sect. 2, SESE, R. Melville 492 (WIS).

+5 July 1958, open Carex-Sphagnum floating mat, edge of A. with a little

Chamaedaphne, "Wisconsin Gardens" Nature Preserve on S short of Lake

Shlsehogama, T39N R5E Sect. 2, Hugh H. Iltis and W. Buckmann 11458 (WIS).

+26 August 1964, margin of Carol Lake, plant rooted to the lake bottom,

depth of water 2-3 inches, the leaves and flowering axis erect and

above the water surface T39N R7E Sec. 9, J. G. Iyer 205 (WIS).

OZAUKEE CO.--+30 July 1934, pond, Cedarburg bog, H. Cutler 241 (WIS).

POLK CO.--25 August 1929, narrow sand shore. Twin Lakes, Sect. 29

T33N R16W, Amery, W. T. McLaughlin 638 (WIS). +23 August 1960, very

common in shallow lakes and swamps, growing in open water usually less

than 2 feet deep, Sagittaria and cattail common in same area, this

plant from Grass Lake, T37N R17W Sect. 5, West Sweden Twp., Miles

Johnson (WIS). 26 Sept. 1965, in shallow water about 2 feet deep, in

Balsam Lake a medium hard water lake, 4 miles E of Balsam Lake

Wisconsin, T34N R16W Sect. 6, SW k, C. T. Lind 44 (WIS). PRICE CO.—

+Cno date], T37, NR1, W Sect 13 (WIS). +August 1909, color blue,

Charles Goessl [7] (WIS). +18 July 1941, Lac du Flambeau, 7% miles

SW at Turner L., growing in shallow water and muck on lake margin sun­

light, common in locality, all plants blooming at date of collection,

[John Neesel 643 (WIS). RACINE CO.— +July 1843, I. A. Lapham (MO

1983383). ROCK CO.— +8-23 1930, now wet sandy shore, Nellie V. Haynie

2038 (F 1514482). SAUK CO.— +8/14 1942, Sec. 34 T7N R13E, Lake

Chapman, Nina Leopold (WIS). +5 Oct. 1945, Section 18 T12N R9E, R. A .

McCabe (WIS). SAWYER CO.--17 August 1929, sandy shore Whitefish Lake, abund. in shallow water with peaty bottom, flowers blue, T37N R11W

Sect. 4, J. G. Mauritz 1688 (WIS). WAUKESHA CO.— Cno day or month}

1928, Lake Deuoon [?], Muskego, Elizabeth Ehrler (WIS). WAUPACO CO.—

13 Sept. 1931, muddy shore of White L., Weyauwega, N. C. Fassett and

J. W. Rhodes 13985 (WIS). WOOD CO.— +1 August 1961, growing in water,

Gottschalk cranberry Co., T21N RAE Sect. 9, Walter A. Skroch 175 (WIS).

CO. 7— +July 1881, Green Bay Sloughs, J. H. Schuette (F 350070, GH,

NY, US 752395, WIS). +25 July 1884, Muskego Lake (NY-2). STATE ? —

+[no date], August flowers, Fresh Pond, Map C?], J. T. Rothrock

(F 321995). +12 July flowers, South Hadley, J. T. Rothrock (F 321996).

+Baltimore (GH). Phld., C. C. G. Mon (MO 1983342). 3960 (MO 1983360).

+Ex Museo botanico Berolinensi, Ex Herb. A. Braun (MO 1983435). +The

Bernhardi Herbarium (MO 1983438, 1983439, 1983440, 1983441). +N. A.

Beyrich [?] (MO 1983442). +Ex coll. D. N. Dean (M0 1983443, 1983444).

+The Henry Eggert Herbarium, bought 1905 (MO 1983445). +Herb. J. Q. A.

Fritchey, bequeathed 1897 (MO 1983446). +(M0 1983447). +8-17 Mystic,

(MO 1983449). +(M0 1983450). +Herb. of the College of New Jersey,

Scientific Dept., Wm. H. Leaman (NY). +Herb. Pursh propr., Herbarium

Hookerianum 1867 (NY). +July, Catskile, Herb. W. DeF. Day (NY).

+Torrey Herbarium (NY). +Herbarium Columbia University New York (NY).

+Muhlenberg Herbarium, [457: Sheets 1 and 2], (PH-2). +(PH-2).

+Essex Co., Mafs [?], Oakes (PH). +Pontederia canadenfis. Not,

Mrs. Parcival, [Schweinitz ined.3, (PH). +Boston Little, 162 (PH).

+Au. [Aug.] 10, New Haven, MaSheum [?], Herbm . of H. H. Eatan, C. W.

Short Herb., (PH 783880). +Benjamin Smith Barton Herbarium, collected by Dr. Gray of Southhampton, Va. (PH). +Benjamin Smith Barton Her- Sect. 22 T39 R9W, 10 miles S of Hayward, N. C. Fassett 10311 (WIS),

18 August 1931, T42N R6W, Teal Lake, Clam Lake, J. H. Steenls 782 (WIS).

26 July 1934, muddy shore of Holly Lake, Hayward, Aldo Leopold and

N. C. FaBsett (WIS), +2 Sept. 1962, shallow, 5 dm deep water, vicinity of camp site and boat landing, Pickerel Lake (Lake of the Pines), T38N

R3W Sect. 14, NW %, Hugh H. Iltls 20618 (WIS). SHAWANO CO.--[no date],

Showano, I. A. Lapham (WIS). TAYLOR CO.— 23 August 1966, blue flowers, in shallow water on N. Spirit Lake, about 6 miles N of Rib Lake, frequent, T33N R2E Sect. 6, Wm. J. Barnes 381 (WIS). VILAS CO.—

2 Sept. 1929, locally abundant, Big Wildcat L., Boulder Junction,

N. C. Fassett 9045 (WIS). + [no day or month] 1936, R. A. Elder, Jr.

(WIS). 5 August 1941, mud 1 foot, High L., Boulder Jet., L. A. Fraser

(WIS). +8-25 1941, White Sand Lake, edge of shallow water, J. E .

Potzger 9108 (WIS). +17 July 1953, Irving Lake, T41N R8E Sect. 2,

Delle N. Swindale 2600 (WIS). +2 Sept. 1955, Thunder Lake Marsh, three lakes, Rita Horner (WIS). +5 Sept. 1959, White Birch Bog, common in 1%* deep water, edge of lake, T42N R7E Sect. 35, NE % of NE %,

H. H. Iltis 15398 (WIS). 2 7 June 1962, Vilas Co., shallow calcareous waters ca. 3-5 dm deep, with Lobelia dortmanna, Gratiola lutea,

Sparganium angustifollus, Eriocaulon pellucidum, Pontederia cordata,

Brasenia schreberi, N end of Middle Ellerson Lake (T41N, R6E, NE %

Sect. 29), ca. 9 miles NNW of Woodruff, Hugh H. Iltis and Shoichi

Kawano 20371 (WIS). +14 August 1966, Bog, edge of Spring Lake, T39N

R10W Sil, K. W. Wood 127 (WIS). WASHBURN CO.— 23 August 1929, narrow sand strip, marshy shore Big Casey L., Sect, 21 T40N R13W, Laurel P.O.,

W. T. McLaughlin 643 (WIS). +11 July 1966, Hunt Hill Audubon Camp, abund. in shallow water with peaty bottom, flowers blue, T37N RllW

Sect. 4, J. G. Mauritz 1686 (WIS). WAUKESHA CO. — Cno day or month]

1928, Lake Deuoon C?D, Muskego, Elizabeth Ehrler (WIS). WAUPACO CO.—

13 Sept. 1931, muddy shore of White L., Weyauwega, N. C. Fassett and

J. W. Rhodes 13983 (WIS). WOOD CO,— +1 August 1961, growing in water,

Gottschalk cranberry Co., T21N R4E Sect. 9, Walter A. Skroch 175 (WIS).

CO. ?— +July 1881, Green Bay Sloughs, J. H. Schuette (F 350070, GH,

NY, US 752395, WIS). +25 July 1884, Muskego Lake (NY-2). STATE ?—

+[no date], August flowerB, Fresh Pond, Map [7], J. T. Rothrock

(F 321995). +12 July flowers, South Hadley, J. T. Rothrock (F 321996).

♦Baltimore (GH). Phld., C. C. G. Mon (MO 1983342). 3960 (MO 1983360).

+Ex Museo botanico Berolinensi, Ex Herb. A. Braun (MO 1983435). +The

Bernhardt Herbarium (MO 1983438, 1983439, 1983440, 1983441). +N. A.

Beyrich [?] (MO 1983442). +Ex coll. D. N. Dean (MO 1983443, 1983444).

+The Henry Eggert Herbarium, bought 1905 (MO 1983445). +Herb. J. Q. A.

Fritchey, bequeathed 1897 (MO 1983446). +(MO 1983447). +8-17 Mystic,

(MO 1983449). +(MO 1983450). +Herb. of the College of New Jersey,

Scientific Dept., Wm. H. Learoan (NY). +Herb. Pursh propr., Herbarium

Hookerlanum 1867 (NY). +July, Catskile, Herb. M. DeF. Day (NY).

+Torrey Herbarium (NY). +Herbarium Columbia University New York (NY).

♦Muhlenberg Herbarium, C457: Sheets 1 and 2], (PH-2). +(PH-2).

♦Essex Co., Mafs C?]» Oakes (PH). +Fontederia canadenfis. Not,

Mrs. Parcival, CSchweinitz ined,], (PH). +Boston Little, 162 (PH).

+Au» CAug.] 10, New Haven, MaSheum C?3, Herb1”, of H. H. Eatan, C. W.

Short Herb., (PH 783880). +Benjamin Smith Barton Herbarium, collected by Dr. Gray of Southhampton, Va. (PH). +Benjamin Smith Barton Her­ barium, collected by Benjamin Smith Barton (PH). +Aug., ca. 1860,

Del ab Richmond, Wm. Wynne Wlster Herbarium, presented by Estate In

1899 (PH). +CJuly}, Blue, D. Reece, Leconte, £4 In water with

Saggitarla, one pistil, 6 stamens, 3 long and 3 very short but furnished with anthers and apparently perfect, Aug. 1st (PH). +Plant Endog.

Carol. Mer., L. R. Glbbes (US 33949). +Herb. of I. A. Lapham (WIS).

+N. America, from the Herbarium of Dr. F. J. Young, Germany (WIS).

STATE ? [date]: +1833 [?3, Amen'r, bottom, [probably Missouri}, (MO

1983386 . +Aug. 1834, Hartford [?], (NY). +Aprll 1838, collection of

Prof. Louis R. Glbbes (NY). +20 August 1853, Medford, W. Boott, coll.

Henry Glllman, (NY). +Sept. 1860 (MO 1983448). +13 June 1861 [?],

Prescott C. W. , B. Billings 1. [?], (CAN 227187). +11 June 1868, Hort

Huber B. 237 (NY). +18 August 1869, M. Gandoger, Flora Galllea ex-

slccata, £318, Hab. in paludls Galliae orient, prope St. Georges-de-

Reneneln (Rhone), locadicto "Laye", abr [?} subspont, (MO 1983469).

+20 July 1877, Brauford, Ct. C?3, William Trelease (MO 1983337). +15

July 1883, Norwich, Ct. [?], W. A. Setchell (NY). +12 July 1886,

Closter, Herb, of S. L. Zabrlskie (BKL). +8 August 1891, Water Lily

Creek, Lake George, Geo. D. Hulst (BKL). +6/18 1892, Pic-nic Grove,

Calv. Rd., 536, The N.M. Glatfelter Herbarium bought 1909 (MO 1983402).

+8-4 1893, Pond, Wellesley, Ex Herbarium of F. W. Hunnewell, 2nd (NEBC).

+6 July 1894, Charles River, Herb, of Grace Gilbert, (GH). 14 July

1894, DuPage River S of Naperville, (WIS). +5 Oct. 1894, Verdigris, common, plants of Indian Territory, B. F. Bush 579 (MO 1983421). +2

July 1895, Phoenix, George T. Hastings (NY 256 7). +1 Sept. 1896,

Creek head of Harris Bay, Lake Geo., Herb, of Geo. D. Hulst (BKL). 103

+August 1897, in Lake, Good Luck Lake, Hamilton Co. [probably New York],

H. J. B. 162 (NY 2251). +July 1900, Wilson Ct., pool near Norwalk

River, A. E. Carpenter (OS). +9-29 1901, Herb, of Ohio Wesleyan Uni­

versity (NY). +7/28 1904, S. C. J . (MO 1983418). +August 1907 [?],

R. L. Mann (WIS). +July 1928, backwater, bays, etc., Beardstorm, (F

847679). +12 Sept. 1928, iles a l'embou chuce de la Riviere de

Prairies, Adrien Dubois (NY). +25/7 1946, Granby, Bord de riviere,

Frere Fabius 403 (NY 3134). CENTRAL AMERICA: BRITISH HONDURA :

BELIZE DISTRICT— Nov. 1933, Northern River, Percy Gentle 944 (GH,

MICH, M0 1064062, NY). 24 Dec. 1969, 9 miles north of Belize City,

along Northern Highway, Belize District, R. M. Lowden 24 (OS-12).

30 Dec. 1969, Belize District, 32 miles north of Belize City along

Northern Highway, R. M. Lowden 27 (0S-5). 30 Dec. 1969, Belize

District, 29 miles north of Belize City along Northern Highway, Cowhead

Creek, R. M. Lowden 26 (OS-5). 30 Dec. 1969, Belize District, Hattie-

ville-Burrell Boom Road, 5% miles from Burrell Boom, R. M. Lowden 28

(OS-6), COROZAL DISTRICT— 1931-1932, Percy H. Gentle 388 (F 713016,

MICH, US 1587455). EL CAYO DISTRICT— 1 April 1958, "Water Lily",

flowers white, in high ridge, 41 miles Section, Belize-Cayo Road,

Percy H. Gentle 9708 (DUKE 163229, F 1600873, MICH, NY, US 2572823).

ORANGE WALK DISTRICT--Feb. 1933, Hillbank [Hill Bank] Lagoon, herb,

occasional, (herb, flowers very pale mauve-white, occasional in shallow water), R. S. Pelly 43 (F 679802) and 44 (F 679803). STANN CREEK

DISTRICT— 18 June [no year], vie. Cockscomb, 400 ft. altitude, 2h ft. height, diam. k in., flower white, slight perfume, rare, William A .

Schipp S119 (F 659211). 11 March 1932, very showy aquatic growing in small pools in shade, flowers white with faint mauve spot, "rare",

23 mile, Stann Creek Valley, 250 ft. alt., 2 ft., W. A. Schipp 955 (F

657813, GH, MICH, Mo 1029553, NY). TOLEDO DISTRICT— 28 Nov. 1951,

**Water Lily", flowers white with yellow, in swampy damp area, 15 miles,

San Antonio-Punta Gorda Road, Percy H. Gentle 7531 (F 1600922, NY, US

2572821). SOUTH AMERICA: ARGENTINA: PROVINCIA OF BUENAS AIRES — 27

Nov* 1938, Depto. Campana, Campana, muck and shallow water of river- bank, full sun, area grazed by cattle, alt. 10 m . , perennial herb to

1 m., fl. blue, common, associated with Scirpus, W. J. Eyerdam and A. A .

Beetle 23070 (GH). PROVINCIA OF CHACO— +[date ?], coman [?3 en el agua, Los Pilsor [?], P. Jorgensen 2529 (US 1055202). 12 Dec. 1928,

Dept. Resistencia, Colonia Benitez, 150 m. alt., color de la flor azul, en un estero, S. Venturi 7907 (GH, US 1545332). +XI 1940, Enrique

Urien, Campo Bonazzola, lote 9, Dept. Tapenaga, Banados, flor olerte palido, A* P. Rodrigo 2381 (NY-2). +21 1 1947, Margarita Belen, acuatica, R. M. Aguilar 1035 (MO 1634309). PROVINCIA OF CORRIENTES —

XI 1936, Mercedes, 12 leguas al N., banados del Azul, "0[C3amalote",

A. P. Rodrigo 750 (NY). +26 X 1967, Prov. Corrientes, Dept. Mburucuya,

Paso Ceario, en el agua, fl. celestes, A. Lourtelg 2111 (US 2547107).

PROVINCIA OF MISIONES— +6 II 1947, Dept. Apostoles, Tres Galpones, flora Lila, planta alt 0.60 mts. [73, A. M. R. Kuldobro 4817 (MO

1634335). +I-II 1936, camlno de Reconquista a Vicanor Molinas, flor

Ilia, M. M. Job 866 (US 1691680). TERRITORY OF FORMOSA— 5 [13918, abundate en lagunas Formosa, P. Jorgensen 3005 (MO 866779). 8 [13918, abundate en [?3 Aeros Formosa, P. Jorgensen 3005 (GH). 2 [13919, abundate en banadas [?3 Formosa, P. Jorgensen 3005 (US 1065639). PROVINCIA 7— 23 Dec. 1888, Bord de l'Yl (Durazaro) [?], M. M. Alvares,

Frommel, Lavenir, Lefebvre 8 (F 814320). 18 XI 1907, plantae In

clvitate argentine Misiones collectae, Posadas, ad flum, Alto-Parana

pr. praed, "La Granja" [?], E. L. Ekman 1344 (NY). BRAZIL: PROV.

MINAS GERAIS— +[no date], Brasilia, I. s35 Sele [?] (PH 618815).

PARANA— XI 1960, plantas Do Brasil, estado do Parana, Rio Pequeno,

auto estude Curitiba-Paranagua, aquatica, flores azues, R. Brage and

E. Vesretra [?] 320 (US 2446929). RIO DE JANEIRO--+June 1832, Brasilia,

Prof* Rio de Janeiro, L. Rledel, iter Brasiliensis 1821-36, (NY).

Feb. 1833, Brasilia, Rio de Janeiro, pr. Lorena, L. Rledel, iter

Brasiliensis 1821-36 (NY). RIO GRANDE DO SUL— +1828 [?], aquatica

paludosa, L. Leopoldo 360 (GH). (no day or month] 1940, flowers cherry

red, in watery places, vicinity of Sao Leopoldo, Jose Eugenio 99 (NY).

+8 XI 1949, Rio dos Sinos, B. Rambo 44322 (US 2026910). 24 XI 1949,

Lagoa dos Barros, pr. Osorio, B. Rambo 44574 (F 1508888). 30/10 1961,

Km. 19 da BR-37, entre Guaiba e Arroio dos Ratos, em brejos, flores

azuls com labelo amarelo, G. Pabst 6417 and E. Pereira 6590 (F 1656303,

NY). SANTA CATARINA--5 Dec. 1956, 14.5 km. from C, Mun. Curitibanos

and Campos Novos, west of Curitibanos toward Campos Novos, alt. 850-

950 m., L. B. Smith and R. Klein 8304 (US 2338189). 26-28 Dec. 1956,

Mun. Chapeco, Fazenda Campo Sao Vicente, 24 km. west of Campo Ere, alt.

900-1000 m., L. B. Smith, Pe. R. Reitz and 0. Sufrldlnl 9468 (US

2338215) and 9482 (US 2338216). 20-21 Feb. 1957, south banhado with

corduroy causeway, Mun. Chapeco, Fazenda Campo Sao Vicente 24 km. west

of Campo Ere, alt. 900-1000 m., L. B. Smith and R. Klein 11624 (US

2338259). SAO PAULO— XII 1900, Campinas, Jose De Campos Novaes 1231 (US 390003). 17/10 1917, n. vulgar "Aguape", Butantan, S. Paulo, planta do brejo, fl. coerulea, F. C. Hoehne 726 (NY). COLOMBIA:

C0MMISSAR1A MAGALENA— +12 Oct. 1944, succulent herb growing In shallow

standing water, about 50 cm. high, flowers white with yellow spot,

spike very showy (shoot Is erect), much visited by butterflies, near

Valencia, alt. about 200 m., Oscar Haught 4406 (F 1324271, US 1709345).

COMMISSARIA VALLE [DEL CAUCAD--+16 May 1944, acuatica, moja corlacea,

flexible verde clara, perianto bianco o bianco rosado, costa del

Paclfico, rio Cajambre, Quebrada de Guapecito, 0-5 met. alt., J.

Cuatrecasas 17703 (F 1354697 and 1354698). PARAGUAY: +1853-6, ex­ ploration of the River La Plata and adjacent countries. Dr. Edward

Palmer (US 33962). +1888-1890, Central Paraguay, Thos. Morons, Luque

(18 Dec. 1888) 301 (F 164911, NY, MO 1983468, PH, US 245052 and 937959) and Thos. Morong, Villa Rica 490 (GH, NY). 1885-1895, Laguna Ypacarai,

Dr. E. Hassler 306 (NY-2) and 306a (NY). Nov. 1913, Paraguarla

Centralis, in reglone lacus Ypacaray, Dr. E. Hassler 12683 (GH, MO

847475, NY, US 1057500). +[date ?3, petals blue, one with yellow spot, common in water, Prof. Pedro Jorgensen 3891 (F 705003, GH, MO 971944,

NY, PH 660102, US 1483689). 2 X 1892, Villa Florida, Otto Kuntze (NY).

+[date ?], [place 73, Dr. E. Hassler 3803 (NY). [date 73, In altoplanitie et decllvils, Sierra de Maracayu, Dr. E. Hassler 5363

(GH, NY). +1888-1890, Pllcomayo River, Thos. Morong 1040 (NY-2).

+Sept. 1892, Sued Paraguay, am Rio Febienary [?3, Otto Kuntze (NY).

25 XI 1950, Dep. Paraguarl, Paraguarl, Cerro Hu, Sparre Y. Vervoorst

564 (MO 1769422). +31 8 1954, herb 30-40 cm. with short rhizome, fls. blue, with yellow circle on upper petal, swampy campo, A. L. Woolston 107

314 (NY), URUGUAY: DEPARTMENT CANELONES — 6 III 1910, Toledo, Pando,

Santa Lucia, 20-30 M.S.M., Santa Lucia, Orilla del Rio, W. Herter (US

1233908), Nov. 1926, n.v. Carnalote, plant 1 m * , fl. coerul., In aquls,

Loc. Diet. Toledo, Herter fll. 522 (F 568273, GH, MO 933846, NY, WIS).

+Nov. 1926, n.v. Camalote, plant 1 m., fl. coerul., in aquis, Loc.

Diet. Independencia, Herter 522a (US 1341936). 18 II 1937, Rincon de

Pando, en el agus, Herb. C. Piego Legrand 949 (F 1004191). DEPARTMENT

MALDONADO— +M. 1 Cl9]41, Lau [San] Carloa, Descole [?] 104 (GH).

DEPARTMENT 7-1892-93, Sta. Elena, SorianaCo], 0. V. Aplln (MO

1688406, NY). 8 Feb. 1921, "Camalote", Laz[s]cano, District of Rocha,

Alex. Wetmore 882 (US 1066031). LATIN AMERICA 7-1787-1795-1804,

Plantae Novae Hispaniae, a Sesse, Moclno, Castillo et Maldonado lectae

4979 (F 848212).

lb. Pontederla cordata L. var. lancifolia (Muhl.) Torrey, Fl. U.S. 1:343. 1824.

Pontederla lancifolia Muhl., Cat. 34. 1813. Type locality cited below.

Pontederla lanceolata Nuttall, Gen. 1:216. 1818. ("Hab. near Savannah in Georgia - Dr. W. Baldwyn. Also in South Carolina." Savannah, Georgia, holotype PHI, Figure 10.)

Pontederla lancifolia Muhl. Cat., Elliott, Botany 1:382. 1821.

Pontederla cordata L. var. lancifolia (Muhl. & Elliott) Torrey, Fl. U.S. 1:343. 1824.

Unlsema lancifolia Raf., Alt. Jour. 178. 1833. (Cited as "The Pontederla lancifolia Mg. and Elliot." Habitat, Raf. 1837, "Florida to Texas," "Pont, lanceo­ lata Mg. Elliot." Type not seen.) 108

Pontederla cordata L. var. lanceolata (Nutt.) Grisebach, Cat. PI. Cub. 252. 1866. (Wr. [Wright] 3260 cited only. Figure 12.)

1 Pontederla cordata L. var. typica, Solms In DC. Monog. Phan. 4:532. 1883. [that part synonymous with Pontederla lanceolata Nutt.]

Pontederla cordata L. forma angustlfolia (Pursh) Solms In DC. Monog. Phan. 4:533. 1883. [that part synonymous with Pontederla lanceolata Nutt.]

Pontederla cordata L. [var.] lancifolia (Muhl). Morong, Mem. Torrey Bot. Club 5:105. 1894. [that part synonymous with Pontederla lancifolia Muhl.]

Naruklla cordata (L.) Nleuwland var. lancifolia (Muhl.) Nieuwland, Amer. Midi. Naturalist 3:101. 1913. [that part synonymous with Pontederla lancifolia Muhl.]

Hlrschtla heterantherlmorpha K. Schum. ex Schwartz in Engler, Bot. Jahrb. 61 (no. 139):41. 1927. (Habitat, "Brasilien: Im Staate Goyaz, leg. A. Glazion n. 22228." Type not seen.)

Pontederla. heterantherimorpha (K. Schum.) 0. Schwartz in Engler, Bot. Jahrb. 61 (no. 139):41. 1927.

Unlsema lancifolia (Muhl.) Farwell, Am. Midi. Nat. 11:73. 1928.

? Pontederla oblonga Larranaga, Pub. Inst. Hist. Geog. Uruguay 2:134. 1930. (Cited as "Foliis oblongis, florlbus splcatls.*1 Type not seen.)

Plant maximum height 13 dm; inflorescence length 2.0-15.0 cm; perlgone blue to blue purple; peduncle length (inflorescence base to floral shoot leaf base) 5.0-33.0 cm; leaves narrow lanceolate to ovate lanceolate (Figs. 9, 10), leaf width 0.4-8.3 cm; spathe (terminal phyllode of floral shoot) length 2.0-6.5 cm; floral shoot leaf petiole

(Fig. 12, includes sheath base of petiole) length 4.0-21.0 cm; petiole

(Fig. 12) length of other leaves 29.0-59.5 cm, maximum ligule length of petiole sheath 10.5 cm; sheath (axil of floral shoot) maximum length

28 ■ 0 cm ■

Type locality: United States, Muhlenberg cited "Habitat. Car.

Carolina." Plant "spear-leaved." Lectotype (PH!).

Figure 9.

Distribution: United States, Cuba and South America (Figs. 6, 11, 15). Fig. 9. Provisional lectotype of Pontederla lanclfolia Muhl., in Muhlenberg1s Herbarium (PH), showing a narrow lanceolate flowering stalk leaf.

110

Fig. 10, Holotype of Pontederla lanceolata Nuttall (specimen on the left of the herbarium sheet) collected at Savannah, Georgia (PH), showing a narrow lanceolate flowering stalk leaf.

112 113

f/fnrtmftfa

niini Mi''"' Fig. 11. Geographical distribution of Fontederla cordata L. var. cordata (dots) in British Honduras (solid lines represent highways), C.A., and var. lancifolia (Muhl.) Torr. (triangles) in Cuba, West Indies (maps used, British Honduras Survey Department, Paslow Building, Belize City and National Geographic1s, Atlas of the World, enlarged second edition, map of Cuba).

114 115

r *

no - MILCO

r w Fig. 12. Pontederia cordata 1,. var. lanceolata (Nutt.) Grisebach, Cuba, Plantae Cubenaea Wrightianae, C. Wright 3260 (US 33961), showing narrow lanceolate leaves.

116

118

SPECIMENS EXAMINED*:

ALABAMA: GENEVA CO.--9 May 1967, 8 miles south of Samson, common in shallow water in Taxodium ascendens-Nyssa biflora-Cyrilla swamp, corolla blue with yellow spot on upper median lobe, roots rose-purple, Sidney

McDaniel 8916 (FSU 114729). CALIFORNIA: +29 June 1941, Adelante, The

Garden of Anson and Anita Blake, Berkeley, five feet high, flower color violet-blue, two yellow spots on upper lobe, N. Floy Bracelln 1434 (NY).

+22 August 1943, Adelante, The Garden of Anson and Anita Blake,

Berkeley, 4-5 ft. high, cultivated, N. Floy Bracelln 2581 (F 1503364).

CONNECTICUT: NEW LONDON CO.— 19 August 1913, border of Dodge's Pond

Niautic, east Lyme, E. B. Harger 6328 (NEBC). WINDHAM CO.— 19 August

1918, Thompson, shallow water of Long Pond, C. A. Weatherby 4373 (NEBC).

DISTRICT OF COLCMBIA: ? June 1896, Potomac River, Herb. E. L. Morris

3046 (BKL). FLORIDA: BAY CO.— 1 May 1959, marsh, vicinity of Mine

Defense Lab, Panama City, Meredith Jones (FSU 48290). BREVARD CO.— *

10-20 March 1916, Indian River near Rockledge, Edwin B. Bertram (PH

648754). 30 July 1957, SW shore Lake Poinsette % mi. N of St. Johns

River inlet, frequent on muck of broad marsh, R. Krai 5331 (FSU 34272).

30 Sept. 1963, specimens taken randomly from 3 widely separated strands in the marshes along the St. Johns River, below Lake Poinsett, Brevard-

Orange Co., R. K. Godfrey and Richard D. Houk 63089 (FSU 83275-83280).

BROWARD CO.— 31 Jan. 1940, 11 mi. west of Davie, flB. pale blue, growing in water in open glades, R. J. Seibert 1191 (MO 1182424, PH 792681).

18 April 1952, 25 mi. n. of Miami, on U.S. 27, Henry Field and Yusuf

Lazar (F 1577353, US 2097117 and 2097118). 30 May 1963, Old Glory

The use of the asterisk (*) within a specimen citation indi­ cates that part of the specimen(s) which is(are) var. l a n d folia (Muhl.) Torr. as contrasted with var. cordata. 119

Island, Pool 2A Conservation Area, in the everglades about 10 miles NW of Fort Lauderdale, H. E. Hackett 23 (DUKE 161786). CALHOUN CO.— i-Cin part] wl Sept. 1963, % mile west of Ochlocknee River on SR 20, de­ pression near bridge, nearly fully shaded, Plants 1-13, R. Christensen and R. Kerstetter 6 (FSU 121466). CHARLOTTE CO.— +7/21 1965, Caloosa

Forest/Expt. Area, headquarters pasture, deep sand, fresh water marsh, perennial, common, flower color blue, R. T. Parrott 112 (DUKE 175340).

COLLIER CO.— 15 April 1952, Pinecrest, s. of Tamiaml Trail between mile 40 and Ochopee, Henry Field and Yusuf Lazar (US 2097120). 15

August 1960, 20 miles south of Ochopee in the Glades between Ochopee and Paolita on Hwy. 41, Richard S. Mitchell 739 (FSU 64827). COLUMBIA

CO.— 4-30 1893, streams. Lake City, P. H. Rolfs (F 229018, MO 1983362).

1 May 1959, in ca. 1 ft. of water, swamp, 18 miles N of Lake City,

R. K. Godfrey 58518 (FSU 47947), 13 May 1961, marshes bordering swamp,

4 miles N of Lake City, separate pieces on extra sheets, this number

60832, each from a different stalk and gathered from widely spaced spots in "population", R. K. Godfrey and Richard D. Houk 60832 (FSU

64493-64499, 645-8-64515). 20 May 1964, marsh in clearing of cypress swamp, 6 miles N of Lake City, R. K. Godfrey 63755 (FSU 34701, 42020).

DADE CO.— +24 March 1899, Baldwin, H. B. Meredith (PH 624959). 1-9

Nov. 1901, the Everglades, west of Miami, J. K. Small and G. V. Nash

(NY). 31 Oct.-4 Nov. 1903, Nov. 1903, in everglades near the un­ finished railroad grade, between Cocoanut Grove and Cutler, J . K . Smal1 and J. J. Carter 666 (NY, PH 568265). 5 Nov. 1906, Everglades between

Miami and Snapper Creek, J. K. Small and J. J. Carter 2452 (NY). +Nov.

1906, Bet Miami & Suepper C?3, Small and Gatler C?] 114 (DUKE 25555). +18-26 Jan. 1909, Everglades, intersecting Long Key (Everglades),

J. K. Small and J. J. Carter 2941 (NY). 21-29 Sept. 1917, Wampee,

flowers bright blue, everglades, west of Park, W. E. Safford and C. A .

Mosler 115 (US 1035922) and 128 (US 1035935). I. 1923, Everglades,

Miami, D. C . Peattie (F 781047). +21 Jan.-[19]24, Long Key, J. K .

Small and J. J. Carter 302 (PH 570844). 28 Dec. 1927, in everglade

water, along Tamiami Trail, 25 miles west of Miami, flowers blue,

Harold N. Moldenke 3802 (NY). 1 April 1930, in water at edge of canal,

along Tamiami Trail, plants about 2 feet tall, flowers blue, Harold N .

Moldenke 844 (DUKE 09727, MO 1002674, NY). 18 Feb. 1933, very common

in streams, Miami, Delzie Demaree 10243 (DUKE 137365, MO 1047311).

DE SOTA CO.— =7 May 1917, shores of Lake Hicpochee, cruise of the

"Barbee1', Okeechobee Region, J. K. Small 8189 (NY). 14 March 1926,

bog, roadside, Arcadia, Mary H. Williams (PH 636872). 14 April 1961,

perianth white, this color form very local in extensive stand of blue-

flowered form, shallow water in hardwood swamp, e. of Fla. 72, w. edge

of Arcadia, S357, T37S, R24E, D. B. Ward B-22, with BTY 421 (FSU

94956, US 2604329). DUVAL CO.--May [no day or year], watery cypress

swamp, N. E. Florida, A. H. Curtis 2988 (BKL, F 132733, GH, MO 1983456,

NY-3, US 33956 and 937966). June, near Jacksonville, A. H. Curtiss

2988 (F 34840 and 132732). 19 April 1894, swampy places in pine barrens

near Jacksonville, A. H. Curtiss 4782 (F 108360, GH, MO 1983453, US

224476). 7 April 1897, So. Jacksonville, J. R. Churchill (GH). 16

April 1897, So. Jacksonville, Spring Glen, J. R. Churchill (MO 1053771).

28 April 1902, ditch, A. Fredholm 5126 (GH, US 717122). Spring 1932,

Jacksonville, M . E. Piemer (WIS). FRANKLIN CO.— +[no date], 121

Apalachicola, [Chapman ?] (MO 760555), 24 July 1957, in dense stand of small cypress, swamp along river, 11 miles south of Sumatra, plants

In shallow water, R. K. Godfrey 55715 (FSU 33678, GH). 5 Oct. 1963,

2 miles south of Crooked River bridge on SR 65, rooted plants In open roadside ditch. Plants 1-7, [8], 9-25, R. Christensen and Rex Kerstetter

13 (FSU 121616-121635). +[in partj 5 Oct. 1965, SR65 at Crooked River

Bridge, rooted plants near culvert, mostly shaded, Plants 1, 3-6, 10,

★Plants 2, 7-9, R. Christensen and R. Kerstetter 14 (FSU 121636-121638,

121640, 121642, 121645; *FSU 121639, 121641, 121643-121644). 5 June

1967, on Fla. 65, 4.6 miles n of U.S. 98 turnoff near Green Point, swampy, roadside ditch and field, Donald E. Stone 2255A long style form

(DUKE 183866) and 2255B short style form (DUKE 183866). GADSDEN CO.—

6 June 1956, along the banks and In the water of a roadside ditch, 1 mile west of the OckCh]lockonee River along U.S. 20, Paul L. Redfearn,

Jr. 2194 (FSU 29186, GH). GLADES CO.— ?+8 Nov. 1936, edge of pond near

Lakeport, Vida M. Lovett 186 (DUKE 38635). ?+10 June 1961, marshes,

3.5 miles N of La Belle, R. K. Godfrey and Grady W. Relnert 61013 (FSU

65221-65224). GULF CO*— 7+14 June 1960, swampy woodland. Cape San

Bias, each piece on separate sheets, RKG 59830, from a different stalk got wandering randomly across "population", R. K. Godfrey and J. N .

Triplett, Jr. 59830 (FSU 58080-58083, 58087-58091). HAMILTON CO.—

1 May 1959, in 6-12 Inches of water, swamp, 2 miles E of Jasper, R. E .

Godfrey 58508 (FSU 47952-47953). HIGHLANDS CO.— 10 March 1945, plentiful in grassy shallows of a quiet bay, flowers violet, Bear

Point, Lake Childs, 1.. J. Brass 14779 (GH). +20 March 1945, marsh In flatwoods 1 mile west of Archbold Biological Station, gregarious in 122

masses scattered In water and on wet margins of marsh, flowers blue,

conspicuous, L. J. Brass 14881 (US 2065223), 3 April 1960, flowers

bluish, common, shallow water off n. edge of Lake Placid (Lake Childs),

1 mi, s. of town of Lake Placid, S7, T37S, R30E, D, B. Ward 2-14 with

BTY 421 (DUKE 168873). HOLMES CO.— +20 July 1956, frequent on wet

sandy-peat of pondshore and in shallow standing water, 1 m. W of

Westville, R. Krai and P. L. Redfearn 2877 (DUKE 166056, FSU 29172,

GH). 1 May 1968, on U.S. 90, 0.2 mi. e. of Florida 81 in Ponce de

Leon, 3 style forms of Pontederia, D. E. Stone, short-style form 2589

(DUKE 199478, NY), long-style form 2590 (DUKE 199477, NY), and mid­

style form 2591 (DUKE 199476, NY). INDIAN RIVER CO.— +18 74, Indian

River [CO. 7], Edward Palmer 538 (GH, MO 126182, NY). 22 Feb. 1957,

headwaters marsh of St. Johns River, about 10 miles west of Vero

Beach, Robert J. Lemajre 149 (FSU 32157). 9 April 1962, marshy clearing

of swamp, ca. 4 miles S of Vero Beach, separate pieces, my number 61831,

from different stalks ca. 3 m. apart across "populationt’, R. K. Godfrey

and Grady W. Reinert 61831 (FSU 72302-72306). +[in p a r O 10 April

1962, marsh, 3 miles SE of Yehaw Junction, separate sheets of 61848,

each piece of plant from a separate Btalk widely scattered in "popu­

lation", R. K. Godfrey and Grady W. Reinert 61848 (FSU 72218-72225).

JACKSON CO.— ?+[in part, no date), leaf only, Herb. John A. Lowell

(GH). +10 April 1929, shallow pools, prairies, near Grand Ridge,

E. J. Palmer 35277 (GH, MO 983120). 17 March 1937, Sneads, Exploration

Party 1937 (DUKE 50646). +5 April 1957, in about 18 inches of water,

open cypress depression, Grand Ridge, R. K. Godfrey 55472 (FSU 33592,

GH, NY). +20 May 1960, very abundant, open gum pond, 2 miles E of Grand Ridge, this number 59539, each piece of plant taken from a dif­

ferent stalk about 4 paces apart zigzagging across pond, R. K. Godfrey

59539 (FSU 56338-56341, 56346-56348, 56360-56364). +[in part] 21

Sept* 1963, roadside ditch 5 miles west of Sneads on U.S. 90, rooted

plants in generally exposed habitat, Plants 1-2, 5-32, R. Christensen

and Rex Kerstetter 3 (FSU 121499-121518). [in part]+21 Sept. 1963,

roadside cypress swamp, 2.6 miles south of Cottondale on U.S. 231,

exposed with some shade, Plants 1-10, 12-26, R. Christensen and Rex

Kerstetter 4 (FSU 121519-121540). LAKE CO.— +16-30 April 1894, col­

lected in vicinity of Eustls, ditches, Geo. V. Nash 450 (F 37934,

GH, MICH, MO 1983455, US 227932 and 937965). +May 1894, collected in

vicinity of Eustls, swamp, flowers white, Geo. V. Nash 678 (NY).

+June-July 1894, vicinity of Eustls, A. S. Hitchcock (F 233269 and

233270, MO 1983457). +1925, Orlando, H. E. Wheeler (F 554777). +17

May 1960, Ocala Nat'l Forest along Rd. 72, dry longleaf pine-live

oak-turkey oak-palmetto sand ridge se. of Lake Dorr [Dorra], James D .

Ray, Jr. and George R. Cooley 9724 (DUKE 147989). 10 June 1961, marshy shores of lake, 13 miles S of Leesburg, this number RKG 61036,

separate sheets, each piece of plant from a separate stalk taken at widely spaced places in "population", R. K. Godfrey and Grady W .

Reinert 61036 (FSU 65262-65267, 65290-65292). LAKE OR ORANGE CO. 7 —

25 April 1930, marsh, Lake Apopka, collected by Hugh O 1Neill for F. S .

Blanton 6442 (GH). LEE CO.— 15 August 1963, low roadside, along Fla.

#80, about 3 miles east of Tice, Norlan C. Henderson 63-1590 (FSU

62344). LEON CO.— [in part] +18 Sept. 1963, west side of Lake Jackson on U.S. 27, rooted plants on exposed lake shore, young plants in deep 124 water (1% ft*) had very long petioles and very small lanceolate leaves,

Plants 1-15, [3 specimens without Plant numbers!, R. Christensen and

Rex Kerstetter 1 (FSU 121470-121482). [in part]+25 Sept. 1963, north­ west side of Lake Mlccosukee at Leon Co., Public Fishing Grounds, exposed lake shore, Plants 1-4, [4 sheets without Plant numbers labelled misc.], Christensen, Kerstetter. and McDaniels 8 (FSU 121564-

121568, *121563, 121562 [*in part]). 7+2 Oct. 1963, west side of Lake

Jackson on U.S. 27, rooted plants on exposed lake shore, Robert

Christensen and Sid. McDaniels 9 (FSU 48856). LEVY CO.--7+27 April

1959, section 7, T17S, R17E, in fresh water, flowers blue, Richard J .

Eaton and George R. Cooley 6590 (GH). LIBERTY CO.— [in part]+12 June

1960, very abundant in ca. 1 ft. of black water, swamp, Apalachicola

National Forest, New River, on separate sheets, my number 59735, each portion of plant is from a separate stem got more or less randomly across ’'population", R. K. Godfrey 59735 (FSU 57742-57743, 57761-57769).

MARTIN CO. [?]— 11-25 Nov. 1913, hammocks on sand-ridge, eastern shore of Lake Okeechobee, Pelican Lake to Cypress Creek, J. K. Small and

G. K. Small 4320 (NY). NASSAU CO.— +2 June 1958, In shallow water, peaty drainage ditch, Yulee, R. K. Godfrey and William Lindsey 56836

(DUKE 140566, FSU 40336). 23 May 1964, cypress swamp, 10 miles E of

Hilliard, R. K. Godfrey 64093 (FSU 87291). OKEECHOBEE CO. [?] —

11-25 1913, shores at mouth of Kissimmee River northern end of Lake

Okeechobee, J. K. Small and G. K. Small 4383 (NY). ORANGE CO.— +19

July 1894, Lake Brantley, Fred'k L. Lewton (NY). OSCEOLA CO. [7]—

+27 April 1901, Kissimmee River at Alligator Bluff, Edgar A. Mearns

(US 391087). PALM BEACH CO. [?]— +11-25 Nov. 1913, Fort Lauderdale to Lake Okeechobee, J. K. Small and G. K. Small 4132 (NY). +11-25 Nov.

1913, shores of Pelican Lake, Lake Okeechobee, J. K. Small and G. K .

Small 4140 (NY). 29-30 March 1952, Loxahatchee Wild Life Refuge, west of Delray Beach, Henry Field and Yusuf Lazar (F 1577359, US 2097121).

PASCO CO.— +13 April 1927, Fish Lake, shallow water, Rev. Hugh O'Neill

(MO 948636). POLK CO.— +12 May 1894, marsh, L. B. OhllnRer 520 (F

230520, MO 1983454). +March 1898 [?], J. M. Milligan (US 504163).

21 March 1952, Mountain Lake, Lake Wales, Henry Field and Yusuf Lazar

(US 2097119). 14 July 1956, abundant in standing water around the edge of phosphate pools, about 4 miles SW of Bartow, P. L. Redfearn,

Jr. 2696 (FSU 31806, GH). PUTNAM CO.— 18 April 1897 (specimen one on sheet), Aug. 1903 (specimen two on sheet), Palatka (specimen one),

Lake Ganoga (specimen two), C. S. Williamson (?). ST. JOHNS CO.— +22

May 1964, drainage canal bordering cypress swamp, 7 miles SE of Orange- dale, R. K. Godfrey 64002 (FSU 34381). SARASOTA CO. — tin part]+l

August 1950, Myakka River State Park 18 miles southeast of Sarasota,

1.5 m. high, perianth tube reddish at base and violet above like the lobes, yellowish green spot on the largest lobe, filaments and anthers violet, R. L. Wilbur and G. L. Webster 2505 (GH, NY, US 2067565)

SQ1IN0LE CO.--+4-1 1956, edge of Lake Adelaide, My Home, Herb, of Paul

O. Schallert 21916 (WIS). VOLUSIA CO.— +12 April 1911, Lake Helen, shallow water, Ex Herb. S. C. Hood (GH). WAKULLA CO. — ?+22 July 1957, shallow water of pool in cypress-gum clearing, 4.5 miles N of Craw- fordville, R. K. Godfrey 55654 (FSU 33754). Cfor the most part3+19

May 1960, in up to 1-1.5 ft. of water, openings in swamp, between

Medart and Sopchoppy, my number 59531, the separate sheets, each piece 126 from a separate stalk got randomly across "population", R. K. Godfrey

59531 (FSU 56342-56345, 56349-56357). [in part] 5 Oct. 1963, modified cypress swamp, 1 mile north of Crawfordville on SR 369, plants rooted and in partial shade, Plants 1-3, 5-8, 11-20, (7 misc. sheets), R.

Christensen and Rex Kerstetter 11 (FSU 121569-121588). [In part]

5 Oct. 1963, roadside ditch 4 miles south of Crawfordville on SR 369, plants fully exposed and had been mowed, Plants 1-4, 6-8, 10-27,

29-30, [and one sheet without a Plant no.],[*five sheets without

Plant nos.]. R. Christensen and Rex Kerstetter 12 (FSU 121589,

121591-121594, 121596-121605, 121607, 121610-121615; *FSU 121590,

121595, 121606, 121608-121609). FLORIDA CO. ?— +[no date], Southern

Flora, Herb. Chapman (NY, US 33958). +[no date], Chapman (NY).

+[no date], J. F. Powell (US 33957). +[no date], G. V. Nash (NY).

+[no date], Cape Florida (NY). + [no date], near Fort King, Lt. Alden

(NY). +[no date], label says Types of the Flora of the Southern States,

A, W. Chapman on same sheet as specimen from Georgia saying "Leconte"

(PH). +[no day or month], Rugel 196 (MO 1983458, US 512103). +March

1658, Ponds, Magnolia, Herb. Wm. M. Canby (NY). +June-July 1898,

Jaxa [?], A. S. Hitchcock (F 231899). +1900, North Florida, A. M.

HuRer 3 (MO 1983357). =5/29 1901, Danford [?], (MO 1983459). +28

Dec. 1910, Banks, J. W. Hershberger (NY). +13 March 1919,

Lardcan Camp, Big Cypress, Perley Poore Sheehan (NY). GEORGIA:

GRANTLEY CO.— 18 August 196 7, emergent in ditch through pine falt- voods along Ga. Rts. 15 & 121, 1 mile south of Hoboken, A. F. Clewell

2745 (FSU 108699). BROOKS CO.--[for the most part]+64 June 1959, in standing water of roadside ditch, ca. 4 mi. s of Quitman on U.S. 221, William P. Adams 176 (FSU 48844-48851). 6 June 1959, standing water of roadside ditch and adjacent damp soil, ca. 1 ml. nw of Morven,

William P. Adams 182 (FSU 48855). Tin part3+6 June 1959, in water along edge of small stream along roadside, ca. 6 ml. east of Pavo, colony appearing rather uniform, William F. Adams 184 (FSU 48757,

48759-48 760). CAMDEN CO.— +12 May 1930, in wet ditch, Colesburg, plants about 2 feet tall, flowers blue, Harold N. Moldenke 1182

(DUKE 09734, MO 1000848, NY). CARLTON CO. — *38 ? [?], near Camp

Cornelia in Okefenokee Swamp, Blrma Jarrard 2226 (DUKE 49558). Tin part3 2 May 1959, marsh, In ca. 1.5 ft. of water, Okefenokee Swamp,

W of Camp Cornelia, R. K. Godfrey 58551b (FSU 48215, 48226-48229).

9 May 1965, Okefenokee Swamp, roadside ditch ca. 1 mile from tourist area at end of park road, Jean Wooten 143 (FSU 96153). CARROL CO.—

+18 July 1959, along the shore but in the shallow water, small lake between Franklin and Carrolton, Sam Jones (FSU 51372). CHATHAM CO,—

[no date3 Savannah, (Pontederla *lanceolata Nutt.), (PH). CLINCH CO.—

4/10 1935, edge of pond, Dupont, Vida M. Louett [?] (DUKE 28613).

6/22 1936, moiBt open soil, 3 miles south of Homerville, D. S. Correll

5535 (DUKE 134924, FSU 21974). DeKALB CO.— ? 24 May 1897, Lakes,

H. Eggert (MO 1983407). DOOLY CO.— April 1845, In einem Sumpfe 1m

Dooly County unter Vienna, F. Rugel 24 (MICH). ECHOLS CO.— 17 August

196 7, ditch along Georgia Rt. 94, ca. 5 miles northwest of Statenville,

A. F. Clewell 2528 (FSU 108745). GLYNN CO.— 4/16 1961, 7 miles north­ west of Brunswich on hwy. 341, growing In 12" of water sandy bottom,

Steve E. Wright 41 (GH). LOWNDES CO.— 4/10 1938, Valdosta, W. B. Baker

2978 (DUKE 49557). 2 May 1959, in ca. 1 ft. of water, swamp, between Stockton and Valdosta, R. K. Godfrey 58570 (FSU 48224-48225, 48231-

48232). 5 May 1963, drainage canals through pine flatwoods, 5 miles

E of Valdosta, each piece of plant RFG 62760, from different stalk,

gathered one ca. every 3 paces along canal bank, R. K. Godfrey and

Richard D. Houk 62760 (FSU 78651-78662). McINTOSH CO.— +24 May 1909,

Darien, Altamaha Grit, Huron H. Smith 2110 (F 247759, WIS). RICHMOND

CO.— 13 June 1924, marsh south of Tubman Home, Augusta, S. F. Hilde­

brand (DUKE 09732). 19 August 1924, Tubman Home swamp, Augusta,

S. F. Hildebrand (PH 779671). WARE CO.— 2 August 1902, shallow pine-

barren ponds between Waycross and Ruskin, geological formation,

neocene, overlaid by Lafayette and Columbia, alt. 125 feet, Roland M .

Harper 1469 (F 141009, GH, MO 1983406, NY, US 431774). +12 May 1930,

very common, fl. deep blue or purplish, cypress pond, Suwannee Lake,

Jean Sherwood Harper 68 (DUKE 25553, PH 726 750). 25 April 1936, in

water, wet pine lands, ca. 10 m. s.w. Waycross, Arthur N. Leeds and

FranclB Harper 2793 (DUKE 38080, PH 740562). 17 August 1967, common

in 1-2 feet of water in a ditch along US Rt. 84 at Manor, A. F .

Clewell 2550 (FSU 108447). WAYNE CO.— 4 June 1893, Jesup, T. H .

Kearney, Jr. (OS). June 1893, margin of ponds, Jesup, A. Ruth (MO

773032). 5 June 1893, shallow ponds, Jesup, T. H. Kearney, Jr. (F

122674). GEORGIA: CO. ?— +tno date], MLeconteM, on same sheet as

specimens collected in Florida by A. W. Chapman, sheet with label

saying Types of the Flora of the Southern States (PH). + Cno date],

(herb Schw), Keb nom. [?], ”P. angustifolia Georgia Bald", (PH).

+Cno date], Georgia, C. W. Short, M.D. (MO 1983452). +[in part; no date], Milledgevllle, Georgia £?], Dr. Samuel Boykin (PH 781424). 129

+tno date!, (label perhaps Interchanged with Boykin from Georgia),

(PH 781420). + [no date], Georgia in fossis aquosis (HO 1983451).

INDIANA: LAKE CO.— 24 June 1896, sloughs, Miller, L. M. Urn bach (PH

669238). 18 July 1896, sloughs, Miller, L. M. Umbach (MICH). 7+7

July 1897, border of slough, Edgemoar Lake Co., Agnes Chase (? 703736).

7+8 July 1897, sloughs, Miller's, L. M. Umbach (F 434873, GH, MICH,

Mo 1983460 and 1983461, WIS). 9 July 1897, sloughs, Miller’s, L. M .

Umbach (F 85980, US 351282). MAINE: OXFORD CO.— 20 July 1934, emergent in Roxbury Pond, Byron, J. F. Reed 440 (DUKE 24665, PH

720794). CO. ?— +[in part] 22 August 1867, Spencer Pond, Herb, of

Charles E. Smith (PH). ?+[no day or month] 1891, "Cherno", Kate

Furbish (NEBC). +August 1901, Norcross [?] Pond, So. Ches. [S.

Chesterville], L. 0. Eaton (GH). MASSACHUSETTS: BARNSTABLE CO.—

7+12 Sept. 1897, Nine Mile Pond, Centreville, Emile F. Williams (GH).

7+27 July 1914, shallow water of pond-hole, Harwich, C. A. Weatherby

(NEBC). FRANKLIN CO.— 7+24 August 1925, Leverett Pond, Leverett,

Plants of Amherst & vicinity, J. R. Sanborn & F. C. Seymour 2529 (DUKE

66212). HAMPSHIRE CO.— tin part] +30 June [no year], Belchulown

CBelchertown] ponds, W. H. Blanchard (NEBC). 7+13 July 1931, Green­ wich Lake, Greenwich, A. S. Goodale, S. F. Potsubay and F. R. St. John

(NEBC). MIDDLESEX CO.— [no date; in part]. Fresh Pond, Cambridge,

Ball (MO 19883335). tno date; in part], Fresh pond, Cambridge, G. Ball

(F 814323). NANTUCKET CO. — [in part] +26 August 1962, Polpis, Nan­ tucket Island, slimy, dangerous quaking bog, Frank C. MacKeever N687

(NY-2). WORCHESTER CO.--28 July 1886, East Templeton, Miss Emna

Partridge (BKL). 7+31 August 1927, Brookfield, Quabog Lake, Alfred S . Goodale (DUKE 66220). 7 17 August 1930, S. Spectacle Pd., Athol,

Alfred S. Goodale, Walter C. Markert, and Randall H. Piper (DUKE 66223)

MICHIGAN: ALLECAN CO.— +7/23 1937, T2N R15W Sec. 26, Ely Lake, ata. no. 1, 2 ft. water, 42° 31' N. Lat., 86° 02' W. Long., D. C. Chandler

141 (MICH). ?+7/24 1937, T2N R15W Sec. 25, Little Tom Lake, sta. no.

1, 10" of water, 42° 31* N. Lat., 86° 01' W. Long., sand bottom,

E. L. Cheatum 158 (MICH).NEWAYGO CO.— 7+6-17 1937, T14N R13W Sec. 12-13

Diamond Lake, sta. no. 4, 2 ft. of water, 43° 35* N. Lat., 85° 49* W.

Long., pulpy peat bottom, J. Bailey 5 (MICH). 7+6/19 1937, Nichols

Lake, T15-16N R13W Sec. 31-32, sta. no. 2, 6" of water, 43° 43' N.

Lat., 85° 55' W. Long., sandy marl bottom, J. Bailey (MICH). OAKLAND

CO.— tin part] +23 June 1955, west side of Indian Lake, northeast of

Pontiac, H. E. & V. L. Bailey (WIS). SCHOOLCRAFT CO.— 6/25 1937,

X41-42N R16-17W, Indian Lake, sta. no. 4, 2 ft. (snails 32a) of water, about 46° N* Lat., 86° 20* W. Long., detritus bottom, W. C. Beckman 32

(MICH). NEW HAMPSHIRE: CARROLL CO.— 3 August 1947, Choeorus Lake, 2' of water, Tamworth, Frederic L. Steele (NEBC). NEW JERSEY: MORRIS CO.

1 August 1894, pond, Greed Pond, Wm. M. Van Sickle (BKL). 7 1 August

1894, Green Pond, frequent along shores, Herb, of Wm. M. Van Sickle

(GH, US 309049). NEW YORK: ERIE CO.— ?+[no date], Buffalo, G. W .

Clinton (NY). +3 August 1877, Morrorville [?] L.I., E. S. Miller

(F 134345). ? [In part] +5 July 1873, Saw Mill Brook, Agreebogun [?], flora of Long Island, Ex Herb. H. W. Young (F 557025). NORTH CAROLINA:

BRUNSWICK CO.— 7 [in part] +18 May 1957, coranon in ditch beside low moist sandy savanna on Route 904 about 1*5 miles east of Pireway, perianth deep violet, R. L. Wilbur 5864 (DUKE 141904). OHIO: STARK 131

CO.— ?+July 1836, Gralen und Lecufer, circa Canton, Riehl (NY).

SUMMIT CO.— Cin part} +3 July 1899, Cuyahoga Falls, W. A. Kellerman

(OS 9875 and 9891). PENNSYLVANIA: LUZERNE CO.— ?+July 1896, Lakelugh

[?], Herb, of Stewardson Brown (? 502114). RHODE ISLAND: PROVIDENCE

CO.— ?+7 July 1913, shallow water of muddy pond, Foster, G. S. Torrey

696 (NEBC). +5 Sept. 1920, shallow water of Round Fond, Burrillville,

C. A. Weatherby (NEBC). WASHINGTON CO.— 22 Sept. 1920, Exeter,

Ieppican Pond [?], C. B. Graves and R. W. Woodward (GH, NEBC). 21 July

1928, Worden Pond, South Kingstown, J. Franklin Collins (NEBC).

SOUTH CAROLINA: COLLECTON CO.--18 July 1927, ditch, in sand, 8 miles

southeast of Walterboro, K. M. Wlegand and W. E. Manning (GH). JASPER

CO.— 7+9 May 1956, roadside ditch, 4.7 miles northeast of Hardeeville on U.S. Rt. 17, C. R. Bell and Harry E. Ahles 12255 (GH). SUMTER CO.—

25_May 1914, vicinity of Sumter, Witroer Stone 438 (PH 554350). [in part] +25 May 1914, vicinity of Sumter, Witmer Stone 440 (PH 554352).

TENNESSEE: COCKE CO.--17 June 1939, margin of small pond on south

side of highway, Newport to Greenville, five miles east of Newport, elev. 1200', A. J. Sharp and H. M. Hennison 342 (BKL, MO 1258381).

COFFEE CO.— [in parti wl August 1938, shallow water of pond south of

Manchester, H. K. Svenson 8775 (BKL, GH, PH 796870). CO. 7 — [in part] +July 1898, shallow ponds, Wolf Creek, near Smoky Mts., E. Tenn.,

Albert Ruth 128 (US 345311), 135 (MO 1983382), and 1_36 (NY). TEXAS:

BRAZORIA CO.— ?+ [in part] 30 Sept. 1914, Columbia, margin of swampy lake, E. J. Palmer 6682 (MICH, MO 754346). CHAMBERS CO.— [in part]

5-16 1937, Anahuac, V. L. Cory 22410 (GH). HARRIS CO.— [in part]

+VIII 1904, swamps Houston, 0. Kye (NY). 9/10 1926, Cob Pond, Humble, B. C. Tharp 4314 (US 1285822). JACKSON CO.— 7+8/7 1920, Horseshoe

Lake, J. H. Drushel 4140 (MO 855426). CO. 7 — [in part] +Cno date],

Herbarium Berlandlerlanum Texano-Mexicanum 2516 (F 310232, GH, US

33954 and 937960). +[no date], 434 (GH). +July [no year], waters of

the San Bernard, Mr. Charles Wright (US 937961). +July 1842, banks

of brakish water, near Galveston bay, Buffalo bayou, F. Lindhelmer

(MO 1983423). +July 1842, Lynchburg, F. Lindhelmer (GH). + [no day

or month] 1843, Fasc. I., F. Lindhelmer 194 (GH, MO 1983430, PH 781425).

+ [no day or month] 1844, Dr. Eman. Meyer (NY). VERMONT: NORFOLK CO.—

7+20 August 1961, W. side of North Landing Creek, in pond-pine swamp w. of Creeds, R. Krai 13875B (FSU 68599). WISCONSIN: SHOWANO CO.—

7+26 June 1949, Loon Lake, Sec. 11 T27N R16E, D. L. Grether [7] 7459

(WIS). STATE 7 CNO DATE]: +(GH). +Pontederia lanceolata, "always in ponds, or stagnant water, £. cordata in living or moving water", Dr.

Boykin (NY). +Herb. of Texas Experimental Station, A. M. Hildebrand

(US 329984). WEST INDIES: CUBA: + [no date], Plantae Cubenses

Wrlghtianae, C. Wright 3260 (NY-2, US 33961). +1860-1864, fl. blue- purple, in the river Yaco Yaco introduced C7] fr. los Almacegos, Niel

Labajo C7], Sept. 17, Plantae Cubenses Wrlghtianae, C. Wright 3260

(GH). +1860-1864, Plantae Cubenses Wrlghtianae, C. Wright 3260 (MO

198346 7). 28 Feb., 2 March 1911, 1 m . , corolla 1 cm long, blue purple,

The New York Botanical Garden Exploration of Cuba, Province of Pinar

Del Rio, vicinity of Coloma, N. L. Britton and J. F. Cowell 9693 (GH,

NY, US 696059). 14 August 1920, in water, Cienaga de Zapata, al norte de la Bahia de Cochinos (Sta Clara), Bro. Lebn and A. Loustalot 9530

(NY). 1 May 1940, in a small pond, near La Coloma, Prov. Pinar Del Rio, Bros, Lcfen, M. VictorIn 6 Alain (GH). SOUTH AMERICA: ARGENTINA:

PROVINCIA BUENOS AIRES— Jan. 1852, Expedltio navis "Eugenie" 1851-53,

Buenos Aires, N. J. Anderason (US 1690300). 11/XII 1891, Buenos Aires,

Otto Kuntze (NY-2, US 701639). +1-1914, Delta, Rio Chana, A. C. Scala

67 (NY). 29 III 1931, Isla Santiago, cerca de la Plata, obs.

Heloflta junto a la rlbera, A. L. Cabrera 1705 (NY). 30 IV [19332,

Ribera del Rio de la Plata, Isla Santiago, A. L. Cabrera 2161 (F

670276, US 1574497). ?+12 XI 1939, Bosque de Punta Lara, Genevieve

Pawson 852 (GH, NY). +21 Dec. 1942, Delta marshland, Tigre, H. H .

Bartlett 19274 (MICH). +22 IV 1945, Punto Chico, F.C.C.A. [?], R. A .

Wosez [?] (GH). PROVINCIA CORRIENTES— +6 X 1949, Dept. Ituzaingo,

Salto Apipe, flor celeste, G. J. Schwarz 6130 (MO 1709444, US 2172780).

+27 10 1950, Estancia "Santa Teresa," Canada Portillo, Dept. Mburucuya,

Prov. Corrientes, Troels Myndel Pedersen 798 (US 2141472). [in part]

+25 III 1967, Dept. Concepcion, IV sec., Estancia "Palermo", planta acuatlca, nombre vulgar Camalote, Marla M. Arbo 44 (MO 1964951, WIS).

BRAZIL: PROV. MINAS GERAIS— 1868, Brasilia, prov. Minas Gerais, cldade de Caldas, G. E. Henschen (US 209442 and 209443). [no date], Brasilia, prov. Minas Gerais, A. F. Regnell (US 204681). PARANA--+29 11 1910,

Deslro Ribas, in paludosls, P. Dusen 10849 (GH, NY, US 1280857). +16

11 1960, procedencia Pinhaes-Parana, Banhodo IHN 6271, Eduardo A .

Moreira 40 (US 2440568). RIO DE JANEIRO— 17 April 1952, estado do Rio de Janeiro, municlpio de Cabo Frio, Cabo Frio, Praia do Pontal (22° 56'

48" S-42° I* 54" W. Grw.) ao nivel do mar, Lyman B. Smith 6595 (US

2197793). 30 May 1953, St. of Rio de Janeiro, Sao Joao da Barra Co., along the road, between Sao Joao da Barra-Atafona, brejo turfoso can 134

30 a 60 cm de agua, completamente coberto de gramineas, Restlnga-I,

F. Segadas-Vlanna, L. Dau, W. T. Ormond, G. C. Machllne and J. Loredo,

Jr. 416 (GH, US 2283941). RIO GRANDE DO SUL— +31 I 1948, Belem Novo,

Costa Rto Guahyba, M. A. Palacios and A. Cuezzo 416 (MO 1634336). 18

III 1948, Quarai, Palaclos-Cuezzo 1986 (MO 1634246). 2 12 1957, I.A.S.

(Institute Agronomico Do Sul) Pelotas, Canal I.A.S., Sao Goncalo,

Escassa, Jose da Costa Sacco 748 (F 1549223). BRAZIL— [no date],

Burchell, Catalogus Geographicus Piantarum Braslllae Troplcae No. A156

(GH). 6 Sept. 1936, Capao Bonito, Campo Grande, Mato Grosso, flores

azues, W. Archer and Aug. Gehrt 109 (NY 2699, US 1740798). 8 IX 1949,

Do Varjao, Rio Paranalba [Minas Gerais], flores azues, Loc. S.

Terezinha, mun. Huintalu [?], estado Minas, A. Macedo (MO 1600292, US

2196745). 7/10 1953, Mato Grosso, Mun. de Corumba, Fazenda Marilandia,

flores esbranquicadas, aquaticas, E. Pereira 225 (NY) and flor violeta-

claro 252 (NY). 2 Dec. 1961, State of Sao Paulo, Municipio de Sao

Carlos, porto Pulador on the Rio Moji-Guacu, 8.9-9.0 km. NNE of the

RR station at Santa Eudoxia, alt. 500 m. on narrow levee between the

river and a marsh, in open spot in mud of small brook bisecting the

levee, the brook flows through a forest, corolla tube light blue out­

side, inside on the upper part yellow, style light blue, George Eiten

and J. M. de Freitas Campos 3497 (US 2555590). COLOMBIA: COMMISSARIA

MAGDALENA— +2 Nov. 1924, swamp, Poponte, Magdalena Valley, height 3 ft.,

fl. white, Cyril Allen 786 (F 1391708, MO 1015052). CCMMISSARIA

PUTUMAYO— Dec. 1930, remedy for "caracha", perianth white, swamp,

Umbria, 0° 54* N, 76° 10' W, alt. 325 meters, forest, G. Klug 1877

(F 642745, GH, MICH, M0 1004G56, NY, US 1456627). PARAGUAY: 1901- 135

1902* iter ad Paraguarlam septentrlonalem, in regione cursus superioris

flumlnis Apa, Dr. E. Haasler 7849 (GH, MICH, MO 1573413, NY). URUGUAY:

DEPARTMENT 7— +XII 1943-1 1944, Nueva Palmina [7] (Dto. Colonia),

alrededores de la poblacion, n.v. aguaipe, Rosa Scolnik 32 (NY 3641).

lc. Pontederia cordata L. var. ovalis (Mart, in Roemer & Schultes) Solms In DC., Monog. Phan. 4:533. 1883.

Pontederia ovalis Mart, in Roemer & Schultes, Syst. Veg. 7:1140. 1830. Type locality cited below.

Pontederia lanceolata Nutt. var. vlchadenals Hermann, Caldasia 5:39. 1948. (Habitat, Colombia, "Vichada: mucky edge of stream in open forest on llanos bordering the Rio Vichada, alt. 100m., ca. 18 km. NE [E.N.E.3 of San Josh de Ocunh," 21 Jan. 1944, F. J. Hermann 11045, holotype US 2169640!)

Pontederia lanceolata Nutt, forma ovalis (Mart.) Castell., Rio de Janeiro Jardln Botanico 15:62. 1958.

Plant maximum height 9 dm. (Fig. 16); inflorescence (Fig. 14)

length 2.7-16.0 cm.; perlgone white or blue; peduncle length (in­

florescence base to floral shoot leaf base) 12.3-28.0 cm., usually densely pubescent next to inflorescence base (Fig. 13); leaves ovate

(Figs. 13, 14, 16), leaf width 2.2-21.0 cm.; spathe (terminal phyllode of floral shoot) length 2.2-7.4 cm.; floral shoot leaf petiole (includes

sheath base of petiole) length 5.3-31.0 cm.; petiole length of other

leaves 28.7-30.6 cm., maximum ligule length of petiole sheath 4.0 cm.;

sheath (axil of floral shoot) maximum length 22.3 cm.

Type locality: Habitat, Brazil, "Crescit in Brasiliae mediterraneae

stagnis," holotype M. Photograph of type seen US!, Figure 13. Habitat, prov. Goyazanae, 1836-1841,

gardner 3483, NYJ, cited with type, Figure 14.

Distribution: United States and South America (Figs. 6, 15). Fig* 13* Holotype of Pontederia ovalis Mart* in Roemer & Schultes, Brazil (M), photograph (US), showing ovate leaves.

137 ...+ >•

I u n ite d s t a t e s n a t io n a l m useum Fig. 14. Specimen cited as Pontederia ovalis Mart. InMartlus, Flora Brasiliensis, collected by George Gardner 3483 (NY).

139 U .v o tn Fig. 15. Geographical distribution of Pontederia cordata L. var. cordata (dots), var. lancifolia (Muhl.) Torr. ("triangles) and var. oval is (Mart.) Solms (stars) in South America (map used, National Geographic's, Atlas of the World, enlarged second edition, map of South America).

141 oar

S*lT Fig. 16. Holotype of Pontederia lanceolata Nutt. var. vlchadensis Hermann, 21 Jan. 1944, Colombia, Comisaria del Vichada, San Jose de Ocune, F. J. Hermann 11045 (US 2169640).

143 1U2

SPECIMENS EXAMINED:

UNITED STATES: FLORIDA: MARION CO.— 18 May 1949 Juniper Creek, Ocala

Natf 1 Forest, D. W. Mather M-277 (GH). NORTH CAROLINA: JACKSON CO.—

18 June 1929, Sapphire Lake, S. 0. Trentham (DUKE 09725). SOUTH

AMERICA: ARGENTINA: PROVINCIA OF BUENOS AIRES— +2 XII 1944, Part.

t?3 La Plata, entre Rio Santiago cf [?3 Balo Blanco, Rodriguez V. 571

(GH). PROVINCIA OF CORRIENTES--24 IV 1944, Dept. Concepcion, Con­ cepcion, planta acuatica, C. Ybarrola [?3 322 (GH-2). BOLIVIA:

DEPARTMENT OF BENI— 27 Oct. 1921, Reyes, alt. 1000 ft., H. H. Rusby

1389 (BKL, GH, MICH, NY, PH, US 1232357). BRAZIL: PROV. MINAS GERAIS

19-XI 1937, Dentro de lagoa, Dattas de Cima, Mun. Dlamantina, pouco, fl. olva, herbacea, Mello Barreto 9817 (F 1007286). 8/11 1952, entre

Tiradentes e Sao Joao Del Rey estado de Minas Gerais, planta de lagoa formadas pelo Rio Das Mortes, A. P. Duarte 4079 (NY-2). [in part}

+19 Sept. 1964, vicinity of Sobredinho, marshy ground by stream, aquatic, corolla white, uppermost lobe with yellow patch on inside,

G. T. Prance and N. T. Silva 59082 (MO 1811483, NY, US 2457328). +3

Oct. 1965, aquatic, rooted in mud, shallow water, flowers white, unlcate, gallery forest and adjacent meadow and cerrado, vicinity of

Planaltlna, D.F., elevation 950 m., H. S. Irwin, R. Souza and R. Reis dos Santos 8904 (NY). +14 IV 1936, Brasilia, civit. Parahyba, Sao

Goncala, Varzea, Ph. von Luetrelburg 28014 (F 912233, NY, US 1700345).

PARANA— +28 XI 1959, Est. Parana, Mun. Arapoti, Arapoti, das aguas de corrego, flor alva comacula amarela, G. Hatschbach 6541 (US 2324918).

RIO GRANDE DO SUL— +10/11 [19304, Neu. Wurhemburg, in 450 m, von

Alfred Bornmuller 356 (GH). +24 I 1949, Guaiba, In palude ad flumen, B. Rambo 40141 (GH, US 2046893), 11 XI 1949, Navegantes, prope Porto

Alegre, B. Rambo 44357 (US 2026916), 31 III 1950, pared, prope

Montenegro, B, Rambo 46524 (F 1494640). BRAZIL— +[no date], Burchell,

Catalogue Geographlcus Plantarum Braslllae Troplcae No. A157 (GH).

+(no date], Burchell, Catalogue Geographlcus Plantarum Braslllae

Troplcae No. A163 [in part] (GH). 28 August 1945, 12 k. south o£

Crlstaes CCrateus ?], Ceara, fls. white with deep yellow spot on Inside

of upper petal, anther light yellow with dark area at tip, in pond,

common in this one, Hugh Cutler 8398 (US 1989761). +10/11 1946, S.

Leopoldo [Rio Grande Do Sul], RGS, in palude, E. Henz SJ 35412 (MO

1576735). +29 Sept. 1963, plants of the Belem-Brasllia- Highways,

Estados Para, Goias, Mato Grosso, Terr. Pondonia 1 m. high, flowers

white, in bog in cerrado, 300 km. past Cuiaba en route to Goiania,

600-700 m. alt., Mato Grosso, Bassett Maguire, J. Murca Pires, Celia

K. Maguire, and Nilo T. Silva 56931 (US 2513755). COLOMBIA: CGM-

MISSARIA BOYACA— 29 March 1939, growing in mud along margin, Cano

Orocue, flowers white, one petal with yellow spot, near Orocue, alt.

140 m., los llanos, Oscar Haught 2716 (GH, US 1707263). CCMMISSARIA

VALLE [DEL CAUCA]— +28 Jan.-10 Feb. 1944, hoja verde glauca oscura,

flor azul, costa del Pacifico, rio Yurumangui, Veneral, bosques, 5-50 m. alt. J. Cuatrecasas 16153 (F 1354674 and 1354675). CCMMISSARIA

VAUPES--1-3 June 1953, in headwaters of Rio Paca, flowers white, in water, Rio Paca (tributary of Rio Papuri), alt. 650 feet, Lat. 0° 30'

N, Long. 70° 10' W, Richard Evans Schultes and Isldoro Cabrera 19526

(GH, NY, US 2172436 and 2172437). PARAGUAY: [date ?], in altoplanitie et decliviis, Sierra de Maracayu, Dr. E. Hassler 5112 (GH, NY). 147

URUGUAY: DEPARTMENT CERRO LARGO— +Dec. 1937, Rio Negro, Palleros, laderas pratenses, GallInal, Ara&one, Bergalll, Campal, and Rosengurtt

1239 (GH, US 1742783). DEPARTMENT CANELONES— 26 Nov. 1943, Arroyo del

Sauce, 1 km. south of km. 78 on road west of Solis, H. H. Bartlett

20865 (GH, US 1930147). DEPARTMENT TREXNTA Y TRES— Dec. m. 1932, Loc.

Diet. Vergara, W. G. Herter 1719 (GH).

2. Pontederla saglttata Presl, Reliquiae Haenkeanae 1:116. 1827.

Pontederla cordata L. forma sagittate (Presl) Solms in DC., Monog. Phan. 4:533. 1883.

Pontederla cordata L. var. sagittata (Presl) Schery, Ann. Missouri Bot. Card. 31:157. 1944.

Trlstylous (Table 1); plant maximum height 20 dm.; inflo­

rescence (Fig. 17) length 2.5-15.5 cm.; perlgone blue purple,

pale blue to blue or white blue; hardened perlgone bases smooth

ridged (Plate IV, a); anthers blue; peduncle length (Fig. 18,

Inflorescence base to floral shoot leaf base) 13.0-40.0 cm.;

leaves sagittate, shallow to deeply aurlcled (Fig. 19), leaf

width 4.0-31.5 cm.; spathe (Fig. 21, terminal phyllode of floral

shoot) length 4.0-11.5 cm.; floral shoot leaf petiole (includes

sheath base of petiole) length 6.5-21.5 cm.; petiole length of

other leaves 19.0-68.0 cm., maximum ligule length of petiole

sheath 10 cm.; sheath (axil of floral shoot) maximum length 35.0

cm.; fruit and seeds reniform.

Type locality: Mexico, "Hab. in Mexico," holotype M. Photograph

of type NY1

Distribution: Mexico, Guatemala, Honduras and Costa Rica (Fig. 20). 148

Fig. 17. Numerous-flowered Inflorescences of Pontederla saglttata Preal growing In association with Saglttarla, Honduras, Dept. Cortes, Puerto Cortes, 3 Ave.-5 Calle 0 (photographed, August 1970, R. M. f Lowden).

/

/

Fig. 18. Pontederla saglttata Presl with drooping fruit clusters and sagittate leaves in a woody habitat, Mexico, State of Vera Crui, Laguna Catemaco, Arroyo Agrlo, 6 kins, from Catemaco town between Coyame and San Andres Tuxtla (photographed, July 1970, R. M Lowden). *

Fig- 19. Pontederla saglttata Presl, sagittate flowering stalk leaves, Mexico, State of Vera Cruz, along the Trans Isthmian highway (route 185), 17 kilometers northeast of Minatitlan, R. M. King 933 (US 2397514),

149 150

« Fig, 20. Geographical distribution of Pontederla saglttata Presl in Mexico and Central America (map used, Countries of the , 1960, National Geographic Society, Dept, 87, Washington 6, D.C.).

151 t

MILES Fig. 21. Young plant specimen of Pontederla saglttata Presl, showing short petioles of flowering stalk leaf (arrow) and long petioles of leaves attached to rhizome, Mexico, State of Chiapas, fields along route 225, ca. 1 mile north of Suchiate, R. M. King 3430 (MICH).

153

155

SPECIMENS EXAMINED:

MEXICO: STATE OF CHIAPAS— 10-12 July 1939, Rvina, Palenque, E. Matuda

3688 (F 1027546, GH, MICH). 21 June 1947, Aguas Calientes, Eeculntla,

In swampy wet sunny field, E. Matuda 16623 (F 1283974). 22 Feb. 1948,

Macultian, Acapetahua CAcapetagual, v.n. Lilio de Laguna, in swamp,

E. Matuda 17469 (F 1273172), 19 March 1955, near junction of the Rio

Perlas and Rio Jatatfe at San Quintln and near Laguna Miramar, alt.

200 m., growing in water, flowers blue, Ernest R. Sohns 1658 (MICH,

NY, US 2188730). 20 July 1960, fields along route 225, ca. 1 mile north of Suchiate, abundant in shallow water, open sun, flowers pale blue with yellow centers, Robert Merrill King 3430 (MICH, NY, US

2363031). 4 Sept. 1970, El Ejido de Santa Ana, Distrito de Huixtla,

6 kms. de Huixtla, R. M. Lowden 22 (OS-13). STATE OF OAXACA— 8 Nov,

1917, Laguna de Tonameca, Oaxaca, B . P. Reko 3451 (US 842533). 10

Dec. 1921, Jamiltepec, 250 m. Distrito Jamiltepec, Oaxaca, C. Con­ ga tti 4434 (NY). July 1940-Feb. 1941, District of Tuxtepec, Chiltepec and vicinity, alt. about 20 m., in llanos, flowers blue-purple,

G. Martinez-Calderfen 76 (GH, US 1842007) and 494 (GH, US 1842096).

13 Feb. 1965, near San Gabriel Mlxtepec, Mpio. de Juquila, 2 km. north of San Gabriel, swift rocky stream in tropical subdeclduous forest, elevation 725 m., local, in muddy places, flowers violet, one petal with yellow spot, Rogers McYaugh 22416 (MICH). 25 July 1965, zone of tropical deciduous vegetation, foothills of Sierra Madre del Sur,

53 km. N of Puerto Escondido on road to Zimatlan at km. 221, alt. ca.

1000 m., 16° 10' N, 97° 05* W, growing in water under streamside shrubs, disturbed open roadside, moist grazed pasture, fls. lavender, Keith Roe, Eunice Roe and Scott Mori 559 (F 1646274, WIS). STATE OF

TABASCO— 6 Jan. 1889, In paludosls de Curahueso, Tabasco, fule-

incolarum, Herbarium J. N. Revlrosa 334 (NY-2, PH), 1-6 June 1939, la Palma, Balancan, fls. purplish, In lakeside, E. Matuda 3302 (GH,

MICH). August 1962, fresh-water aquatic, marsh, Popal Grande on road between Frontera and Villahermosa, flowers purple, F. D. Barlow (MICH,

WIS). 23 May 1963, fresh-water marsh, savanna, west-Bouthwest of

HulmangulUo, F. D. Barlow 30/52 (WIS). STATE OF VERA CRUZ— Feb. [no day or yearD, Vera Cruz, in paludlbus inter urbem et Santa Fe et prope

Iob Cocos, Schiede and Deppe 981 (MO 1983463, US 1205363). [no datel, provCe de Vera Cruz, H. Galeotti 5562 (F 195969, GH, NY, US 573154).

A° 1853, Vera Cruz, Fred. Muller 2145 (NY). 1858, Wartenberg, near

Tantoyuca, prov. Huasteca, L. C. Ervendberg 277 (GH, PH 781564).

8 Jan. 1895, Coatzacoalcos, isthmus of Tehuantepec, Charles L. Smith

1034 (F 1524914, GH, MICH, MO 1983464, NY-3, US 312654, WIS).

9-1-1903, Vera Cruz, Tuxpam [Tuxpan}, im Fluss, Rio de Vinasco [Rio

Vlnazco], in Wasser, Ex Museo botanico Berollnensl 3731 (GH, US

1206007). 23 Jan. 1906, Laguna near the city of Vera Cruz, J. M .

Greenman 19 (F 189484-2, GH). 7-III-1907, Vera Cruz, Distr. San Andres

Tuxtla, Laguna de Catemaco, in Wasser, Ex Museo botanico Berolinensi

5005 (GH). 16 Feb. 1910, Jalapa, Vera Cruz, C. R. Orcutt 2874 (F

279013, GH, MO 1983466, US 464897). March 1923, Barranca de Panoaya,

C. A. Purpus 9041 (F 553029, GH, M0 913403, NY, US 1206862). 12

August 1926, Vera Cruz, alt. 20 ft., Geo. L. Fisher 108 (US 1268703).

13 Feb. 1943, Mun. Puente Nacional, km. 394 on Jalapa-Vera Cruz high­ way, between Rinconada and Puente Nacional, C. L. Gllly, Sr., Robert F. Simpson and Donald Dodds 90 (MICH). 27 Feb. 1944, along the side of Rio de Cosolapa, Le Gartera, aquatic herb, fls. light purple, Ejldo de San Agustln, 5 km. northeast of Campo Experimental de Hule, El

Palmar, Zongollca, Josfe Vera Santos 2821 (MICH). 1 August 1958, along

the Trans-Isthmian highway, route 185, 17 km. northeast of Minatitlan, elevation 50 m. or less, common, growing in mud, petals lavender to pink, Robert Merrill King 933 (MICH, US 2397514). 8 Dec. 1958, Vera

Cruz, abundant in swamps in and around Vera Cruz, George Neville Jones

22660 (WIS). 26-111-1964, orillo de rio, Desembocadura del Rio

Mhquina, municipio de San Andrfes Tuxtla, L. Gonz&lez Quintero 699

(MICH). 29-VIII-1964, Rio Los Tuxtla, municipio de Santiago Tuxtla, orilla de rio, L. Gonz&lez Quintero 1409 (MICH). 5-VIII-1965,

Sontecomapan, municipio de Catemaco, orilla de arroyo, alt. 100 m., planta de 1 m. de alto, flores moradas, J. Rzedowshl 20368 (MICH).

25 July 1970, Vera Cruz, El Coyol, Prolongacion Carretera Aleman, long time ago people used the leaves to wash milk bottles and as decoration in the house, v.n. Planta de Tlapa, Flora de Pantanos,

Lirio de Agua, R. M. Lowden 6 (OS-30). 26 July 1970, Carretera

Nacional 180, El Puente de Teculapilla, 5 km. ante de llegar a nw.

Lerdo on route 180, R. M. Lowden 7 (OS-8). 27 July 1970, Laguna

Catemaco, south edge of Laguna off route 180, R. M. Lowden 8a-b

(OS-4); Laguna Catemaco, Arroyo Agrio, 6 kms. from Catemaco Town between Coyame and San Andres Tuxtla, R. M. Lowden 9 (0S-12).

STATE ?— 1841-43, ad ripas fluvii inter el Morro & Rancho Nuevo,

Plantae Mexlcanae Llebmann 1618 (US 1092128). Winter of 1877,

Thomas Hogg (NY). 27-11-1899, San Luis, PI. aquatique, Racine 158

tracante, fleurs mauves, tachfees de jaune, se fermant aprfis midi,

Marais avolelnant la mer, n.v. Platanillo, E. Langlassfe 928 (GH, US

386283). CENTRAL AMERICA: COSTA RICA: PROVINCE OF LIMON— IX-1896, marais de Matlna, os lea fosses le long de la vole ferree, fleuls litas,

aquatique, alt. 0-20 m., H. Ptttier 10301 (US 577317 and 938675).

GUATEMALA: DEPARTMENT OF ALTA VERAPAZ— M Dec. 1886, Coban, alt.

4800 pp., H. von Turckheim 547 (GH, NY-2, PH, US 33959 and 937968).

M. Apr. 1901, Cubilquitz [Gubilquitz], alt. 350 m., H. von Tuerckhelm

8025 (US 937953). DEPARTMENT OF IZABAL— M. Mart. 1889, Rio Dulce, alt.

0 pp., John Donnell Smith 1652 (GH, US 937967), 29 Dec. 1904, vicinity

of Puerto Barrios, near sea-level, William R. Maxon and Robert Hay 3061

(US 473079). 18 Jan. 1905, Livingston, W. A. Kellerman 5131 (OS, US

576281). 27 May 1905, Livingston, in swampy places along seashore,

H. Pittier 357 (NY, US 473022). 2 June 1919, Izabal, shallow water

in Rio Mosinga, very pale lavender, upper lip with yellow spot, anthers yellow, stem erect, S. F. Blake 7864 (GH, US 1168965). 25 Dec. 1920,

Jocolb, Izabal, Lago Izabal along shore growing in the water, Harry

Johnson 1053 (US 1792253). 15-31 May 1922, vicinity of Quiriguh, alt.

75-225 meters, herb 3-4 ft.* fie. pale purple, in marsh, Paul C .

Standley 24029 (GH, NY, US 1139695). 2-6 June 1922, vicinity of

Puerto Barrios, at sea level, in swamp, common, Paul C. Standley

25005 (US 1150606). 15 May 1937, Rio Dulce, shallow water, W. C .

Muenscher 12623 (F 904863). 25 Apr.-6 May 1939, near Puerto Barrios,

at sea level, in marsh, erect 2-3 ft., flowers pale purple, abundant,

Paul C. Standley 73170 (F 991651). 26, 27 Apr. 1939, near Quirigufc,

alt. 72-150 m., in marsh, common, flowers pale purple, Paul C . 159

Standley 72425 (F 991428). 28 March 1940, between Bananera and "La

Presa" In Montana del Mlco, alt. 40-300 m . , stems 3-5 ft. tall, stems

green to purple, leaves fleshy, membranaceous, rich grass green above,

paler beneath, bracts hairy, flowers lilac to lilac-lavender, swamp,

Julian A. Steyermark 38050 (F 1660124, US 1842208). 19 April 1940,

shores of Lago Izabal, opposite San Felipe, between San Felipe and mouth of Rio Juan Vicente, alt. 50 m . , common along shore of lake In

swampy places, petals pale lilac, leaves olive-green, Julian A .

Steyermark 39688 (F 1055462). 1 May 1966, vicinity Lago Izabal, Long.

89° O'-89° 25', Lat. 15° 15'-15° 35', alt. sea-level-600 m. , herb

1.4 m., rooted in 0.8 ra.of water in Rio Izabal ca. 50 m. from river mouth, village of Izabal, at sea level, flowers light purple with yellow markings, Gayle C. Jones and Lynden Facey 3213 (F 1665508 and

1665829, NY). 31 July 1970, Quiriguh, swampy area by railroad, in front of abandoned hospital, R. M. Lowden 10 (OS-9). 1 August 1970,

Puerto Barrios, in swampy areas all around Puerto Barrios, R. M .

Lowden 11 (0S-11). DEPARTMENT OF SAN MARCOS — 18 March 1940, Rio

Suchiate, just west of Ayutla, alt. 20 m., flowers lilac, wet swampy depressions, Julian A. Steyermark 38034 (F 1045744). DEPARTMENT OF

PETEN— 26 April 1932, El Faso, from San Pedro River, C. L. Lundell

1564 (MICH-2, NY). DEPARTMENT OF ZACAPA— 15 Jan. 1905, Gualan, elevation 420 feet, 2% feet, Charles C. Deam 208 (F 200633, GH, MICH,

MO 1983465, NY, US 576916). DEPARTMENT 7— 1885, eastern portions of

Vera Paz and Chiquimula, Sereno Watson 28, 33, and 386 (GH). Feb.

1912, N. P. Cockerell 10 (US 861329). HONDURAS: DEPARTMENT OF

ATLANTIDA— 18 Jan. 1903, shore of Tela River, (Tela) Puerto Sierra, Percy Wilson 70 (F 186033, NY-2, US 937954). 26 Sept. 1916, Ceiba,

In pond near beach, leaves 4-6 ft. high, F. J. Pyer A92 (US 866203).

1923, Tela, fls. violet, A. van Severfen 5 (US 1168279). 14 Dec.

1927-15 March 1928, vicinity of Tela, at sea level, in edge of river, rank erect herb 3-4 ft., abundant, fls. pale purple, Paul C. Standley

53607 (F 582340, US 1407830). 21 July 1934, in swamp near Micos

Lagoon, 12 km. west of Tela, flowers pale purple, T. G. Yuncker 4684

(F 749630, MICH). 6 July 1938, in boggy area, near the seashore, vicinity of Ceiba, T. G. Yuncker, J. M. Koepper and K. A. Wagner 8249

(F 962391, GH, MICH, MO 1194879, NY, US 1747694). 6 Nov. 1948, La

Ceiba, "Platanillo" edge of pond, very common, Antonio Molina R. and

E. C. Becker 1 (F 1373135). 10 August 1970, Tela, drainage ditch by railroad track where the main road passes over the track, MLa Curva",

R. M. Lowden 15 (OS-10); Santiago, 11 miles west of Tela, drainage ditch off main road, R. M. Lowden 16 (OS-12). DEPARTMENT OF CORTES—

M. Mart. 1889, San Pedro Sula, Depart. Santa Barbara, alt. 600 pp.,

Carl Thleme 5510 (GH, US 937955). 9 August 1970, Puerto Cortes,

3 Ave.-5 Calle 0, R. M. Lowden 13 (OS-5); drainage ditch at side of main road to Puerto Cortes, 13,5 miles inland from Puerto Cortes, 1 mile Inland from Bijao-Cemento Hondurenas Co., R. M. Lowden 14

(OS-14).

SPECIMENS FOR WHICH IDENTIFICATION WAS NOT POSITIVE:

GUATEMALA: Department of Alta Verapaz— 10 May 1942, corolla lilac with yellow spot, in water along margin of lake (La Laguna), along route no. 5, between Semococh and La Laguna on road to Chajmayic, alt. 500 m., Julian A. Steyermark 46362 (F 1127347). Department of 161

Peten--28 March 1932, Isabilito, Peten, water plant In stream, C, b .

Lundell 1445 (MICH, US 1494619), 26 April 1932, El Paso, Peten, from

San Pedro River, C. L, Lundell 1564 (US 1494659), 10 April 1933,

Santa Teresa, Subin River, C. L. Lundell 2709 (F 738066, MICH), 3 Jan,

1970, flores lilas, en la orilla de Laguna Peten Itza, frente Playa

Blanca, Santa Elena, Dept, Peten, Rolando Tun Ortiz 512 (F 1678935).

3, Pontederia parviflora Alex., N. Am, Flora 19:59. 1937.

Pontederia cordata L. var. parviflora (Alex.) Schery, Ann. Missouri Bot. Gard. 31:156. 1944.

Homostylous (Table 1), style length equivalent to the lengths

of the 3 longest stamens; plant maximum height 1 dm.; inflorescence

(Fig. 23) length 3.0-9.5 cm.; perigone white to white green;

hardened perigone bases toothed ridged (Plate IV, b); anthers

brown to black; peduncle length (inflorescence base to floral shoot

leaf base) 21.0-35.0 cm.; leaves subcordate (Fig. 22), ovate

lanceolate or lanceolate, leaf width 1.8-10.5 cm.; spathe (Fig. 24,

terminal phyllode of floral shoot) length 5.0-10.5 cm.; floral

shoot leaf petiole (includes sheath base of petiole) length 5.5-

19.0 cm.; petiole length of other leaves 19.0-45.0 cm.; sheath

(axil of floral shoot) maximum length 34 cm.; fruits (Fig. 26)

and seeds reniform (Plate I, d; Plate II, a).

Type locality: Habitat, Panama, Camino del Boticario, near Chepo,

province of Panama, altitude 30 to 50 meters, Oct.

1911, H. Pittier 4556, holotype NYi, isotype US

679717i Habitat of fruiting specimen cited with 162

holotype, Panama, Aguadulce, province of Coclfe,

in savannas, near sea level, 3-6 Dec. 1911,

H. Plttier 4915, GH! NY! US 715090-715091!

Distribution: Panama and Colombia (Fig. 25). Fig- 22- Numerous-white- * I t flowered inflorescences of I . & Pontederia parviflora Alex., shallow cordate based leaves* f t Panama, Province of Herrera, Los Llanos de Santa Maria (photo­ graphed, August 1970, R. M. Lowden). 9

Fig. 23. Pontederia parviflora Alex, in wet field habitat, Panama, Province of Herrera, Los Llanos de Santa Maria (photographed, August 1970, R. M. Lowden). Fig. 24. Flowering specimen of Pontederia parviflora Alex., showing clasping subtending spathe and flowering stalk leaf, Panama, Province of Herrera, Los Llanos de Santa Maria, Lowden 20 (OS).

164 TV|V *rt

TVfVlfir'* I”1!’ H-’rr-Tn; I - * I.lv t ^V* ■' '11 f i i Fig. 25. Geographical distribution of Pontederia parviflora Alex, in Panama and Colombia, stars represent sighted records (maps used. Highway Map of Panama, The Engineer Reproduction Division, Usarcarib, Road Classifi­ cation, revised Sept. 1962 and Countries of The Caribbean, 1960, National Geographic Society, Dept. 87, Washington 6, D.C.).

166

Fig. 26. Fruiting specimen of Pontederia parviflora Alex., Panama, Province of Herrera, Los Llanos de Santa Maria, Lowden 20 (OS).

168 169

Fqn»«J«rl> a L. v*r- ptnrtflcw (A l» .) Sc Im t v

rtJOPA CT P*K;*'

TT'>rrrtr n : I .< « L1*r.- * i, , * ■■' ^nt-' a 'h . v M A ■•■•'

. I 1 •■ f< *' I, SPECIMENS EXAMINED: 9 CENTRAL AMERICA: PANAMA: PROVINCE OF COCLE— 3-6 Dec. 1911, Aguadulce,

In savannas, near sea level, H. Plttler 4915 (GH, NY, US 715090 and

715091). 12 July 1938, pools and their margins In wet llanos between

Aguadulce and Ant&n, ca. 15-50 m., fls. greenish white, upper petal

yellow, R. E. Woodson, Jr., P. H. Allen, R. J. Seibert 1208 (GH, MICH,

MO 1171740, NY). PROVINCE OF HERRERA— 13 Sept. 1938, Santa Marla, ca.

20 m . , herb 1 m . , fls. white with yellow blotch, stamens 3, purple tip,

P. H. Allen 790 (MICH). 10 August 1962, bog near Dlvisa, 4 ft. high,

flowers white, John D. Dwyer 2468 (US 2413647). 24 August 1970, Los

Llanos de Santa Marla, 2 km. at Santa Marla and 3.5 km. at El Escota

from the International highway, R. M. Lowden 20 (OS-40). PROVINCE OF

PANAMA— Oct. 1911, Camino del Boticario, near Chepo, altitude 30 to 50

meters, H. Plttler 4556 (NY, US 679717). 11 Dec. 1923, near the big

swamp east of Rio Tecumen [Rio Tocumen], In swamp, erect, flowers

white, common, Paul C. Standley 26656 (US 1153615) and in mud of swamp,

erect, flowers white, abundant, Paul C. Standley 26495 (US 1153575).

29 Nov. 1934, along road between Panama and Chepo, ponds, flowers

white, C. W. Dodge, A. A. Hunter, J. A. Steyermark, and P. H. Allen

16702 (MICH, MO 1119882). 1 August 1938, boggy grasslands and marginal thicks, between Pacora and Chepo, ca. 25 m., fls. yellowish- white, small sluggish streams in savannas, R. E. Woodson, Jr., P. H.

Allen, R. J. Seibert 1661 (GH, MICH, NY). 26 August 1970, along inter­

national highway, between Pacora and Chepo, 7 miles from Chepo,

Tapagara, roadside, R. M. Lowden 21 (OS-38). SOUTH AMERICA: COLOMBIA:

DEPARIMENT OF MAGDALENA— 1845, Rio de Hacha [Colombia], near Mollno, 171

Sa. Martha, PurCchl], Hooker. 31 Sept. 1961, Costa Del Caribe, oeste de Los Venados (10° N., 73° 42’ W.), llanuras subxerofiticas de sabana ablerta con "matas de monte'1 esparcidas (grupos alslados de ferboles, arbustos y bejucos) 60 m. s. n. m., A. Dugand 5834 (US 2455739).

II. Pontederia subg. Re_p3s_la (Endl.) Lowden, comb. nov. ReusBla Endl. (nom. cons.), Gen. PI. 139. 1836. (Type of subg.: Pontederia trlflora (Endl.) Lowden, Reuseia triflora Endl. ex Seub., Gen. PI. 139. 1836.)

Hardened perigone bases spinulose ridged (Plate IV, c-d);

floral bearing shoots prostrate; main stems with long internodes;

underground rhizomes short (mature plants).

4. Pontederia rotundifolia L. f., Suppl. 192. 1781.

Pontederia cordlfolia Mart, in Roemer & Schultes, Syst. Veg. 7:1142. 1830. (Habitat, Brazil, "Crescit in Brasiliae mediterraneae stagnis," holotype M. Photographs of holotype NY! and US!)

Pontederia braslllensls Willd. Roemer & Schultes, Syst. Veg. 7:1145. 1830. (Habitat, Brazil, "Religu. Willd." "In Brasilia prope Para. Com. de Hoffmannsegg." Type not seen.)

Unlsema orbleulata Raf., Medical Flora 2:108. 1830. (Habitat, Raf. 1830, "... grows in South America . . ." »P. rotundifolia L. f. Type not seen.)

Pontederia erlantha Miquel, Linnaea 17:60. 1843. (Habitat, Surinam, "Crescit Surinam!, ad rlpas fluminls Commewyne (Focke)." Holotype GH-2!)

? Pontederia renntformis Larranaga, Pub. Inst. Hist. Geog. Uruguay 2:134. 1930. (Cited as "Foliis reniformibus, floribus spicatis.'1)

Reussia grazlelae Machado, Revista Brasil. Biol. 7:177. 1947. ("Habitat: Amazonas (?) Bahia (?), in acquis. Collegit: Ducke, ano 1937. Herb. J. Bot. R. Jan. 55.083 (Type)" Type not seen. Photograph of type, shown by Machado 1947, Fig. 3.) 172

Reussla rotundifolia (L. f.) Castell., Lllloa 25:593. 1951.

Tristylous (Table 1); plant maximum height 12 dm. (Fig. 28);

Inflorescence (Fig. 27) length 2.0-7.5 cm.; perigone pale blue to blue or white blue; hardened perigone bases spinulose ridged

(Plate IV, c); anthers blue; peduncle length (inflorescence base to floral shoot leaf base) 6.0-25.0 cm.; leaves reniform (Fig.

29), sagittate (Fig. 31, can be deeply auricled) or cordate, leaf width 3.2-22.0 cm.; spathe (terminal phyllode of floral shoot) length 2.5-4.5 cm.; floral shoot leaf petiole (includes sheath base of petiole) length 4.7-45.0 cm.; petiole length of other leaves 17.0-54.0 cm., maximum ligule length of petiole sheath 7.5 cm.; sheath (axil of floral shoot) maximum length

24.0 cm.; fruit and seeds ovoid (Plate I, c; Plate II, d).

Type locality: "Habitat in Surinamo. C. G. Dalberg." Type not

seen, the possibility of a type in the Linnaean

Herbarium is being investigated.

Distribution: Central and South America (Figs. 30, 33). 173

Fig. 27. Fewer-flowered Inflorescence of Pontederia rotundifolia L. f. showing two yellow-orange spots on the largest perianth lobe, El Salvador, Dept. La Libertad, Laguna de Zapotitan (photographed, March 1970, R. M. Lowden).

Fig. 28. Pontederia rotundifolia L. f. In a swampy habitat at Pito Solo, south end of Lago Yojoa, Dept. Comayagua, Honduras (photographed, August 1970, R. M. Lowden). Fig. 29. Reniform leaves and short inflo­ rescence of Pontederia rotundifolia L. f., Brazil, Brazilian Amazonia, Territory of Rondonia, Prance, Forero, Wrisley, Ramos & Farias 5914 (NY).

174 T • I) U \% I# 7 '* !* ' C t)t i l l r j 1 4 m | Fig. 30. Geographical distribution of Pontederia rotundifolia L. f. in Central America, arrow points to insert of canal zone (map used, Countries of The Caribbean, 1960, National Geographic Survey, Dept. 87, Washington 6, D.C.).

176 I

hJ -i -'j Fig. 31. Sagittate leaves of Pontederia rotundifolia L. f. attached to trailing prostrate shoot, Nicaragua, Dept. Granada, Lago Nicaragua, Lowden 18 (OS).

178 FTjOHA o f w i c m a t v i a -'n' r f t>v•f M -"xr*. rr**ri' i [a ^»rriiv»

», ” T. rf! •■

ro* -imlf L f. SPECIMENS EXAMINED:

CENTRAL AMERICA: BRITISH HONDURA: TOLEDO DISTRICT— 9 Oct. 1941,

Monkey River, In river, "water Illy", flowers mauve with yellow spots,

Percy H. Gentle 3700 (F 1504548, CH, MICH, MO 1213584, NY 2265, US

2087540). COSTA RICA: PROVINCE OF ALAJUELA— 1 August 1949, low tropical rainforest, with palms prominent, vicinity of Los Chiles,

Rio Frio, 11° 42' N., 84° 441 W . , 30-40 m., flowers violet-blue, edge of Lago near Los Chiles, Richard W. Holm and Hugh H. Iltis 823 (GH).

PROVINCE OF GUANACASTE— 5-V-1923, Rio Arenal, alt. 500 m. and 5-IX-

1923, Arenal, alt. 600 m., n.v. Lirio de Agua, Juvenal Valerio 4

(US 1206150) and 360 (US 1206175), respectively. 18 August 1970, in field, along main road to Arenal, 1 km. from Arenal, 2 km. from

Tronadora, R. M. Lowden 19 (OS-27). PROVINCE OF LIMON— 18-19 Feb.

1926, Finca Montecristo, on the Rio Reventazbn below Cairo, edge of stream, herb 2-3 ft., flowers white, alt. about 25 m., Paul C.

Standley and Juvenal Valerio 49024 (US 1305856). 8 March 1965, Barro

Colorado de Norte, R. S. Blalsdell 267 (FSU 96644). EL SALVADOR:

DEPAR1MENT OF LA LIBERTAD— 9 Nov. 1953, bed of nearly extinct Laguna de Zapotitan, flowers lilac, with 2 yellow eyes, N. C. Fassett 29320

(M0 1659606, WIS). 26 March 1970, extinct Laguna de Zapotitan, R. M .

Lowden 23 (OS-24). DEPARTMENT OF SAN SALVADOR— 1905, vicinity of San

Salvador, Carlos Renson 273 (NY, US 399566). 20 Dec. 1921-4 Jan. 1922, vicinity of San Salvador, alt. 650-850 m . , "Balsa", fl. lilac, in spring, Paul C. Standley 19143 (GH, NY, US 1135051). 30 March-24

April 1922, vicinity of San Salvador, alt. 650-850 m., fls. pale purple, edge of stream, common, Paul C. Standley 23284 (GH, US 1138974). 1922, San Salvador, Salvador Calderon 411 (NY, US 1151391). 30 Oct.

1950, In about 1 ft. water, shaded place on delta at Aslno, west end of Lago Ilopango, alt. 1500 ft., corolla light lilac, with bright yellow spot on upper lip, N. C. Fassett 28364 (F 1443311, MO 1678840,

US 2086460, WIS). 8 Jan. 1951, in shallow water on delta at Asino, west end of Lago Ilopango, alt. 1500 ft., N. C. Fassett 28582 (F

1443316, GH, US 2086469, WIS). 7-1-1951, Lago de Ilopango, W-Seite, bei Asino, alt, 450 m u/M, in der Trockenzeit sumpfiger Boden, in der

Regenzeit durch fliessendes wasser uberschwemmt, Otto Rohweder 566

(F 1512411). GUATEMALA: DEPARTMENT OF ALTA VERAPA2— 1 May 1906,

Panzos, F. L. Lewton 400 (US 2521442). 19 Nov. 1920, Panzos, Harry

Johnson 1028 (F 1072430, US 1792248). DEPARTMENT OF IZABAL— 1 June

1919, Izabal, bank of Rio Izabal, stems floating up to 6 m . long, very pale violet, upper lip with yellow spot, n.v. "lechua del agua",

S. F. Blake 7844 (GH, US 1165501 and 1165502). HONDURAS: DEPARTMENT

OF COMAYAGUA— 3 Sept. 1932, Lake Yojoa, Pito Solo, alt. 2000 feet, a floating plant, flowers lavender with yellow spot, J. B. Edwards AQl

(F 663806, GH). 7 July 1934, marsh near shore of Lake Yojoa, Cortez,

2100 ft. alt., T. G. Yuncker 4840 (F 749088, MICH, MO 1087014). 18

April 1945, Pitosolo Lago Yojoa, alt. 500 m., Juvenal Valerio Rod- rlRuez 2917 (F 1164523). 8 August 1970, south end of Lake Yojoa,

Pito Solo, R. M. Lowden 12 (OS-18). DEPARTMENT OF CORTES— 16 August

1951, Agua Azul tract No. "B", north shore of Lake Yojoa near the canyon, stem sprawling, mostly submerged, perianth sky-blue with 2 yellow spots, inflorescence and leaves emerged 1-3 dm., open water at edge of floating sod, or with Eichhornia crasslpes, P. Kamb 2094 (GH). NICARAGUA: DEPARTMENT OF GRANADA— 11 Feb. 1903, Granada,

extensive colonies from erect tangled stalks, about borders of pools,

C. F. Baker (GH, MO, NY). 8 March 1922, near Granada, along the shore

of Lake Nicaragua, fls. lavender with yellow spot, J. M. Greenman and

M. T. Greenman 5746 (MO 907819 and 1881510). 14, 16 August 1970, Lago

Nicaragua, frente a la Terraza, near Granada, R. M. Lowden 17 (OS-24)

and 18 (OS-23). DEPARTMENT 7-1853-56, Herbarium of the U.S. North

Pacific Exploring Expedition under Commanders Ringgold and Rodgers,

C. Wright (GH, US 33960). PANAMA: CANAL ZONE— 28 Oct. 1859, Gatun

Station, Panama Railroad, in water, Sutton Hayes 89, 93 (GH) and 99

(NY). 9 March 1905, Ahorca Lagarto to Culebra, J. F. Cowell 379 (NY).

26 Jan. 1911, black swamp between Gatun and Lion Hill, alt. 10-20 m.,

H.Plttler 2572 (GH, NY, US 677228). 17 Feb. 1911, banks of the

Chagres River, below Gatun, near sea-level, William R. Maxon 4883 (US

6 75255). 24 July 1927, Barro Colorado Island, L. A. Kenoyer 602 (US

1319216). 28 Feb. 1931, Barro Colorado Island, aquatic plant In

shallow water near end of Chapman Trail, C. L. Wilson 74 (F 709295).

July 1931, Barro Colorado Island, in water of lake, flowers lavender,

D. E. Starry 120 (F 703208). 15 Nov. 1931, Barro Colorado Island,

Barbour Trail, Otis Shattuck 408 (F 703117). 11 Feb. 1932, Barro

Colorado Island, in water, flowers light blue, cove w. of end of T.

Barbour Trail, R. H. Woodworth and P. A. Vestal 479 (F 707952, GH).

31 July 1934, Barro Colorado Island, Pena Blanca Bay, Otis Shattuck

1099 (F 737754, MO 1777676). 27 Feb. 1935, Miraflores Lake, vicinity of former town Empire, Culebra Cut and vicinity, 80 m., A. A. Hunter and P. H. Allen 776 (MO 1119866). Jan. 1939, Barro Colorado Island, Majorle Brown 30 (F 1005021). 25 August 1939, Rio Chagres, near

Gamboa, alt. 30 m . , flowers pale lavender, 2 orange spots, P. H .

Allen 1963 (GH, MICH, MO 1210417). 3-20 August 1940, Madden Lake,

flowers lavender, upper petal with 2 yellow spots, R. E. Woodson, J r .

and R. W. Schery 949 (MO 1204958, NY, US 1822615). 17 June 1960,

Frijoles, open sunny area near boat dock, growing in water, light blue

flowers, upper lips yellow, John E. Eblnger 69 (MO 1819208). 28 June

1960, Barro Colorado Island, near dock, flowers light purple with

yellow on upper petal, John K. Ebinger 209 (MO 1819206, US 2561671).

23 Dec. 1963, Barro Colorado Island, frequent along shoreline beside

the main boat dock, flowers light purple, Shirley A. Graham 212 (GH,

MICH). July 1965, along shore of Gatun Lake, frequent, flowers light

blue with yellow on largest petal, M. Victoria Hayden 123 (MO 1827198).

25 June 1966, Barro Colorado Island, in wet area of lake edge near

boathouse, fl. lavender, Edwin L. Tyson 4198 (FSU 104059, MO 1832417).

14 July 1966, Gamboa, aquatic, flowers mauve, Edwin L. Tyson, J. Dwyer,

and K. Blum 4604 (FSU 105857, NY, US 2572190). 2 June 1969, in

Chagres River-N of Gamboa, all along river bank, all material seen

sterile, R. Lazor, M. Correa, and M. Boreham 3739 (FSU 120428).

PROVINCE OF DARIEN— April 1966, Rio Pirre, floating aquatic, flowers

blue, J. A. Duke with Narclso Brlstan 8307 (MO 1834688). SOUTH

AMERICA: ARGENTINA: +May 1851, prope Barra, Prov. Rio Negro, R.

Spruce 1486 (NY). 2 IV 1931, Provincia de Buenos Aires, Flora Del

Portido de La Plata, loc. entre Ensenada 6 Punta Lara, flotante en las

charcas riberenas, A. L. Cabrera 1708 (NY). I II 1936, Loc. Camino

de Santa Fe a Laguna Stubal's Guadalupe, Prov. de Santa Fe, M. M. Job 184

679 (NY). PROVINCE OF CORRIENTES— 2/4 1954, Estancia "Santa Teresa",

Dep. Mburucuyb, Troels Myndel Petersen 2657 (MO 1731124, US 2169238).

+18 IV 1964, Buenos Aires, Rio Santiago, flotante, flor azul, A. L .

Cabrera 15915 (NY). BOLIVIA: DEPARTMENT OF BENI— 25 Oct. 1921, Reyes,

alt. 1000 ft., O. E. White 1513 (BKL, NY). 3 Nov. 1921, Pampas near

Lake Rogagua, alt. 1000 ft., M. C&rdenas 1417 (BKL, MICH, NY, US

1232362). VIII 1926, Rio Chaparfe-Mamorb, ca. 250 m. ii. M., Reise durch Ost-Bolivia, E. Werdermann 2198 (MO 999438). DEPARTMENT OF

LA PAZ— 21 Dec. 1921, Ixiamas, alt, 800 ft., M. Cbrdenas 2031 (BKL,

MICH, NY). DEPARTMENT OF SANTA CRUZ— VI 1892, Yapacani 400 m.,

[Dept, of Santa CruzD, Otto Kuntze (NY-2). 6 Feb. 1925, Prov. Para,

450 metros, rastrera, cleve .10 b-.20 metros, color del tallo alter- nativamente vislaceo y verde-clara, flor lila, Jose Stelnbach 6977 (GH).

BRAZIL: STATE OF AMAZONAS— 15-17 Nov. 1934, Basin of Rio Madeira, municipality Humayta, between Monte Christo and Santa Victoria on Rio

Iplxuna, margin of river, aquatic herb, in river, B. A. Krukoff's 5th

Expedition to Brazilian Amazon!a 7129 (F 874379, GH, MICH, MO 1280512,

* NY, US 1661059). STATE OF PARA— II-VI-1927, Amazonas-Gebiet,

Taperlnha bel Santarem, Waeserpflans enformation im, uberschwanmungs gegiet des Amazonas armes Ayaya, A. Ginzberger and H. Zerny 757 (F

934772). March-May 1929, Belem, B. E. Dahlgren and E. Sella 525 (F

60254 7, US 1471340) and 557 (F 602197, GH). 28-29 August 1934, Region of the Lower Amazon, Bocca do Paru, on varzea land, aquatic herb,

B. A. Krukoff's 5th Expedition to Brazilian Amazonia 5935 (GH , NY).

4 July 1935, in shallow water. Villa Carmo, municiplo de Cametb,

Francis Drouet 1999 (GH, MICH, US 1594688). 28 Sept. 1945, Plantas da Amazonia, flores roxas, J. Murca Plres and G. A. Black 770 (GH).

9 IV 1961, State of Parana, Hun. Gualra, Rio Parana, Barra do Rio

Piquiry, aquatica, flor lilaz com macula amarela, G. Hatschbach 8078

(US 2449448). 12 July 1968, Brazilian Amazonia, Territory of Rondonla,

Basin of Rio Madeira, east bank of Rio Madeira at Abuna, muddy river

bank, in small stream, corolla white, upper lobe with yellow patch,

G. T. Prance, E. Forero, B. L. Wrigley, J. F. Ramos & L. G. Farias 5914

(NY, US 2573101). 3 August 1968, Brazilian Amazonia, Territory of

Rondonia, Basin of Rio Madeira, Rio Pacaas Novos, 4 km. above mouth,

river bank, floating aquatic, corolla white, upper lobe with yellow

patch, G. T. Prance, E. Forero, B. L. Wrigley, J. F. Ramos & L. B .

Farais 6 764 (NY, US 2573100). COLOMBIA: CCMMISSARIA PUTUMAYO— 13

Dec. 1940, Frontera Colombo-Ecuatoriana, selva higroflla del rio San

Miguel, margenes del rio entre los afluentes Bermeja y Conejo, 360-

300 m. alt., hierba rastrera, flores blancas, J. Cuatracasas 11054

(F 1241295, US 1798471). 10 Oct. 1965, Mocoa, camino de herradura

antlguo y rio Rumiyaco, alt. 700-850 m. , n.v. "Amarfen barrachero” ,

planta acuatica, flores blancas, se usa para qultar parallels fasciales,

H. Garcia-Barrlga, Yohel Hashimoto, and Motosuke Ishikawa 18685 (NY).

Nov. 1945, Amazonas, Trapecio Amazonico, Loretoyacu River, alt. about

100 m., fl. white, Richard EvanB Schultes 6935 (US 1952881). Sept.

1946, Amazonas, Trapecio amazonico, , Leticia, alt. about

100 m . , Richard Evans Schultes 8188 (US 1996266). GUIANA: Martin,

Ex Herbario Musel Brltannicl (F 639198). PARAGUAY: April-May 1845,

M. A. Weddell 3156 (NY), [no day or month] 1885-1895, Plantae

Paraguarlenses, Lag. Ypacaray, E. Hassler 1196 (NY). Dec. 1900, 186

Plantae Paraguariensis, Lacus Ypacarai, E. Hassler 3693 (NY). 1913,

Paraguarla Centralis, In regione lacus Ypacaray, E. Hassler 11861 (F

690764, GH, HO 847474, NY, US 1057094). 4 II 1951, Dept. Cordillera,

San Bernardino, laguna, flor rosada con "ojoM naranja, Sparro and

Vervoorst 2316 (GH). : DEPARTMENT OF LORETO— 29 July 1929, Pebas

on the Amazon River, aquatic plant, Llewelyn Williams 1854 (F 615101).

5 Sept, 1929, La Victoria on the Amazon River, La Victoria-Peru-Brazil,

Llewelyn Williams 3097 (F 617075, US 1515826). 22 August 1965,

Requena, Prov. Requena, Dept. Loreto, margen derecha del Ucayallo

Fangoso, flores blancas, macula amarilla, alt. 150 m.s.m., A. Sa-

gastequl and A. Aldave 5757 (US 2483257). SURINAM: (Focke) (GH-2).

+23 XI 1948, herb, half creeping in mud, flowers very pale blue with

yellow mark on upper lip, via secta ab Wla-bank ad Grote Zwiebelzwamp,

in swamp near km. 7, 9, J. Lanjouw 6 J. C. Llndeman 1236 (NY).

Venezuela: Dec, 1952, acuatico, alt. 0, State of Delta Amacuro, Loc.

C. Araguo CBoca Araguaoj, Hermano Gines 4881 (US 2222898). Jan. 1961, planta acuatica, flores moradas, Pto. de La Ceiba, sur lago de Maracaibo,

Edo. Trujillo, L. Arlateguiseta 4443 (US 2343685). +27-30 August 1966,

Estado Tachira, swamps, 12 km, southwest of Punta de Piedra, towards

Sacramento, alt. 175 metros, Julian A. Steyermark & Marvin Rabe 96610

(US 2534925).

5. Pontederia suhovata (Seub. in Mart.) Lowden, comb. nov.

Eichhornia subovata Seub. in Mart. FI. Bras. 3 (l):9l. 1847. Type locality cited below.

Reussia triflora Endl. ex Seubert in Martius, Fl. Bras. 3 (1^:96. 1847. (’'Habitat in Brasilia: Pohl, Sellow". Type not seen.) 187

Pontederla schomburgklana Klotzsch In Schomburgk., Vers. Faun. & FI. v. Brit. Guiana 1118. 1848. (Habitat, British Guiana, "Im See Venturi und auf strom- losen Stellen des Flrara in der Nahe seiner Quelle.*' Type not seen.)

Pontederla lagoensls Wanning in Vldenskab. Meddel., p. 323, T. VI. 1871. (Habitat, Brazil, "Tab. VI, fig. 1-8. In lacu Lagoa Santa (Min. Geraes, Braslllae) . . •" Type not seen.)

Reusela subovata (Seub. In Mart.) Solms in DC., Monog. Phan. 4:534. 1883.

Reusela lagoensls (Warm.) Castell. non Solms, Rio de Janeiro Jardin Botanlco 16:209. 1958.

Trlstylous (Table 1); plant maximum height 30 cm.; inflo­

rescence (Fig, 32) length 3.0-8.0 cm.; perigone blue purple, blue green or white; hardened perigone bases spinulose ridged (Plate

IV, d); anthers blue; peduncle length (inflorescence base to floral shoot leaf base) 5.0-14.0 cm.; leaves subovate (Fig. 32) or ovate lanceolate (Fig. 34), leaf width 0.2-4.4 cm.; spathe

(Fig. 32, terminal phyllode of the floral shoot) length 1.9-5.0 cm.; floral shoot leaf petiole (includes sheath base of petiole) length 4.0-10.0 cm.; petiole length of other leaves 6.5-25.0 cm., maximum llgule (Fig. 32) length of petiole sheath 3.5 cm.; sheath

(axil of floral shoot) maximum length 6.5 cm.

Type locality: "Habitat in Braslllae prov. Goyazana: Gardner

(n. 4022); Bahiensi: Blanchet (n. 2720)."

Lectotype, 1836-1841, George Gardner 4022, NYI

Distribution: South America (Fig. 33). Fig. 32. Fewer-flowered inflorescence and sub- ovate leaves of Pontederla subovata (Seub. in Mart.) Lowden, arrow points to a ligule of a petiole sheath, Paraguay, in regione lacus Ypacaray, E* Hassler 12503 (MO 847628).

188

Fig. 33. Geographical distribution of Pontederla rotund1folia L. f. (dots) and Pontederla subovata (Seub. in Mart.) Lowden (triangles) in South America (map used, National Geographic's, Atlas of the World, enlarged second edition, map of South America).

190 191 Fig. 34. Narrow subovate leaf variant of Pontederla subovata (Seub. in Mart.) Lowden, Brazil, Minas Gerais, Munlciplo Lagoa Santa, Lagoa Santa, L. B. Smith 6702 (US 2173371).

192 Srttil. tfiitftB G«r«i». Iftintclpig ,s',irititr L a ( o a .Miir*, lift 4 0 ' S-43 J V W' ^r« l rty.r a|titud<*. 3d*»*i>' 1^' Lr*an », S»tth_»lL* w* * I*!*!- L 1"11 t SPECIMENS EXAMINED:

SOUTH AMERICA: ARGENTINA: +? 4 1919, Flora del Terrltorio de Formosa

. . . , P. Jorgensen 3347 (GH, US 1065711). 10 II 1941, Prov. Chaco,

Loc. Colonla Benitez, alt. 55m., acuatica, flor azul, C. Meyer 3864

(F 1519935, GH, NY). 4 1 1952, Estancla, "Santa Teresa," Dep.

Mburucuya, Prov. Corrientes, Troels Myndel Petersen 1427 (NY, US

2170070). BOLIVIA: +? June 1886, Reis, 1500 ft., H. H. Rusby 553

(GH, NY-2). 7 March 1922, Trinidad [Dept, of Beni], alt. 700 ft.,

M. Cardenas 27 special (GH, NY). +? II 1950, Santa Cruz, Chiquitos, between El Carmen and Palmito-Chiquitos, height 20-40 cm*, flowers light blue, M. Cardenas 4488 (US 1989924). BRAZIL: +1836-1841,

George Gardner 4022 (NY). 1 2 1949, Pantanos de Canoas, Instituto

Geoblologlco, La Salle, Canoas-Estado do Rio Grande do Sul, Irmao

Gllberto Miguel 23 (F 1337493, GH, US 1953760). 23 IV 1950, aquatica, azues, Loc. S. Tereziuha, Mun. Ituiutaba, Estado Mg [Minas Gerais],

A. Macedo 2326 (NY). 18 II 1951, aquatica, flores azues, Loc. S.

Terezuiha, Estado Minas, Mun. Ituiutaba, A. Marcedo 3168 (US 2026533),

3 May 1952, Minas Gerais, Municipio Lagoa Santa, Lagoa Santa (19° 40'

S-43° 55' W. Grw.), 835 m. alt., Lyman B. Smith 6702 (F 1489846, GH,

NY, US 2173371). BRITISH GUIANA: 7 22 XI 1957, Rupununi, 20 mis. S. of Lethem on road to Wichabi, 18 inch deep pool on sand, alt. - 300 ft herb with floating leaves, inflor. erect, petals blue, yellow at base,

C. D. K. Cooke 197 (NY). + 24 XI 1957, Twinpools South Sand Creek crossing shallow water, blue flowered aquatic, C . D . K. Cooke 205 (NY)

30 Sept. 1963, Rupununi Northern Savanna, aquatic, Mauritia swamp, k m. N. Waruma, grassland with scattered trees, alt. ca. 350 ft., 195 dominants, Curatella, Byrsonlma, Trachypogon and Flmbrletylls, R. Good- land 897 (US 2548129). PARAGUAY: +? 1888-1890, Pilcomayo River,

Thos. Morong 859 (NY). Sept. 1901/2,Iter ad Paraguariam Septentri- onalem, prope Concepcion, E. Hassler 7352 (GH, NY). Feb. 1913,

Paraguaria Centralis, in regione lacus Ypacaray, E. Hassler 12503

(GH, MO 847628, NY, US 1057399). VENEZUELA: 15 1 1939, State of

Cojedes, near San Carlos, margin of pond on llanos, bracts pale blue, flowers yellow, A. H. G. Alston 6311 (WIS). 13 II 1941, "Bora,"

Apure, near Cunaviche, lake, C. E. Cherdorn [7] 254 (US 1801387).

TABLE 1 REPRESENTATIVE LENGTH MEASUREMENTS (MILLIMETERS) OF PERIGONES, STAMENS AND STYLES IN PONTEDERIA L.*

Ovulary Anther

Perigone Style Stamen Taxon Perigone Tube Short Medium Long Short Medium Long

P. cordata var. cordata 14.0 6.5 2.0 0.8 0.8 1.0 10.0 15.0 15.0 7.0 2.0 0.9 1.0 6.0 5.0 15.0 15.0 7.5 2.0 0.8 0.8 13.0 5.0 10.0

var. lancj- 13.0 7.0 1.5 0.6 0.8 folia 1.0 8.5 13.0 13.0 6.5 2.0 0.9 1.0 5.0 5.0 12.0 14.0 6.5 1.5 0.4 0.4 12.5 3.0 9.2

var. ovalis 12.0 6.0 2.0 0.8 0.9 0.8 7.5 12.0 12.0 6.0 2.5 0.8 1.0 5.0 3.5 11.0 11.5 6.0 2.0 0.8 0.9 10.0 4.5 8.0 196

TABLE 1 (continued)

Ovulary Anther

Perigone Style Stamen Taxon Perigone Tube Short Medium Long Short Medium Long

P. eagittata 12.0 5.0 1.5 - - 0.9 0.7 0.9 8.0 11.0 11.0 5.5 - 2^0 - 0.8 - 1.0 A.O 375 10.0 13.0 7.0 - - 2.0 (V7 0^9 11.0 5.0 8.0

P. parvlflora 13.0 7.0 - - 2.0 0.5 - 0.8 5.0 9.9

_P. rotundifolla 20.0 12.0 1.5 - - 1.0 1.2 2.0 11.0 16.0 11.0 6.5 - jM) - h O - L 4 5.2 A.5 9.0 10.0 5.0 - - JM) L 0 U 0 6.2 3.5 6.8

P. B u b o v a t a 19.0 10.0 2.2 - - 1.0 1.0 2.1 10.0 17.0 21.0 12.0 - _U9 - - 1.0 4.8 5.0 11.0 20.0 11.0 - - 2.6 0.9 1.0 1A.0 5.0 11.5

★ Pontederia species are tristylous except Pontederla parvi- flora which is homostylous (see section. "A Tristylous Genus With An Isolated Floral Form: Population Reproductive Biology"). Perigone tube and stamen lengths are estimated; stamen length is from the base of the perigone to the tip of the anther, instead of the place of stamen adnation on the perigone to the anther tip. Sample size, at most, three populations or three herbarium specimens were examined for a taxon. Disposition of Excluded Names

Marantaceae

Pontederla ovata L., Sp. PI* 288. 1753.

Naruklla ovata (L.) Farwell, Pap. Michigan Acad. Sci. 3:91. 1924.

Pontederiaceae

Elchhornia Kunth (nom. cons.)* Enum. PI. 4:129. 1843.

Pontederla aquatica Veil.* FI. Flum. 3, tab. 164, 1827; 144, 1825.

Pontederla azurea Hook, non Sw., Bot. Mag. pi. 2932, excl. syn. 1829.

Pontederla azurea Sw., Fl. Ind. Occ., I, p. 609. 1797.

Pontederla crasslcaulls Schlecht., Abh. Naturf. Ges. Halle 6: 177. 1862.

Pontederla crasslpea Mart. Nov. Gen. & Spec. I, 9, t. 4. CRoemer & Schultes, Syst. Veg. 7:1137. 1830.]

Pontederla elongata Balf., Proc. Bot. Soc. Edinburgh. 50, 1855.

Pontederla martluslana Schult. fil. in Roemer & Schultes, Syst. Veg. 7:1143. 1830. [Pontederla Martlana Schult. fil., Martius Fl. Bras. 3 (1):92. 1947*7]

Pontederla natans Pal de Beauv., Fl. d'Oware, II, p. 18, tb. 68, fig. 2.

Pontederla paniculate Spreng., Neue Entdeck. Pflanzenk. 3:18. 1822.

Pontederla paradoxa Mart, in Roemer & Schultes, Syst. Veg. 7: 1144. 1830.

Pontederla tricolor Mart, ex Seubert, Fl. Bras. 3 (l);92. 1847.

Heteranthera Ruiz & Pavon (nom. cons.), Prodr. 9. 1794.

Pontederla llmosa Sw., Prodr. 57. 1788.

Monochoria Presl, Rel. Haenk. 1:127. 1827.

Pontederla dilatata Buchan, in Roemer & Schultes, Syst. Veg. 7: 1147. 1830. 198

Monochorla (continued)

Pontederla dllatata Andrews Reposit, VII, tb. 490* 1807. [Solms in DC. Monog. Phan. 4:524. 1883.3

Pontederla dubia Blume, Enum., I. p. 33. [Roemer & Schultes, Syst. Veg, 1148. 1830.3

Pontederla hastata L., Sp. Pi. 288. 1753. [Roemer & Schultes, Syst. Veg. 7:1147. 1830.3

Pontederla linearis Hassk., Cat. Bog. 28. 1844.

Pontederla loureirlana Roemer & Schultes, Syst. Veg. 7:1145. 1830.

Pontederla pauclflora Blume, Enum., I, p. 32. [Roemer & Schultes, Syst. Veg. 7:1147. 1830.3

Pontederla plantaginea Roxb., Fl. Ind., II, p. 123.

Pontederla saglttata Roxb., Fl. Ind. II, p. 123,

Pontederla vaginalis L. Mant. [Roemer & Schultes, Syst. Veg. 7: 1145. 183071 A GENERIC BASIS: CYTOLOGY

Methods

Young Pontederla Inflorescences still enclosed by the spathe and the petiole base of the flower stalk leaf (Plate I, a) were col­ lected for cytological investigations of microsporocytes undergoing meiosis. These inflorescences were placed in 1007. ethanol and glacial acetic acid in the proportions 3:1 respectively. After 24 hours, they were removed and placed in 707. ethanol for storage at room temperature.

Anthers were removed and stained with iron aceto-carmine. After squashing, slides were made permanent by freezing them on dry ice.

The cover slip was removed (using a razor blade) and the slides were rapidly dipped first in 95% and then in 1007. ethanol. A drop of euparal was placed over the specimen and a new cover slip added.

Slides were allowed to dry on a warming table (38°C) until the euparal hardened.

Discussion

The earliest chromosome counts in the Pontederiaceae of the

Americas were recorded by R. W. Smith in 1898. These chromosome counts were reported as: n=16, 2n="not far from thirty" in Eichhornia crassipes and n=8, 2n=16 in Pontederla cordata. Taylor (1925) reported

Eichhornia crassipes with a somatic chromosome count of 2n=32. In 1945

Bowden’s cytological investigations of the Pontederiaceae showed

Eichhornia martlana Seub. with n=8 and E. azurea Kunth with 2n=32; 199 200

Fontederla cordata with n**8, P. cordata L. var. angustlfolia Torr.

(■ typical P. cordata) with n»8 and 2n=16; and Heteranthera dubia

(Jacq.) MacM. with 2n*>30.

These earlier haploid reports for Pontederla cordata were re­

confirmed in the present study. Haploid chromosome counts and col­

lection sites of cytological material investigated are recorded in the

following list:

Haploid Taxon Counts Collection Sites and Vouchers

subg. Pontederla typical P. cordata 8 U.S., Ohio, Ottawa Co., Vinous Point, 3^ miles sw. of Port Clinton off State Route 53, Lowden 36 & 37 (OS); New Hampshire, Strafford Co., Bellamy River, Lowden 2 (OS). British Honduras, Belize District, 9 miles north of Belize City, along the Northern Highway, Lowden 24 (OS).

Z* sagittate 8 Mexico, State of Vera Cruz, Vera Cruz, Lowden 6 (OS), Lerdo, Lowden 7 (OS) and Laguna Catemaco, Lowden 9 (OS); Guatemala, Dept. Izabal, Quirigua, Lowden 10 (OS) and Puerto Barrios, Lowden 11 (OS); Hon­ duras, Dept. Cortes, Puerto Cortes, Lowden 13 (OS) and 13.5 miles inland from Puerto Cortes, Lowden 14 (OS), Dept. Atlantida, Tela, Lowden 15 (OS) and Santiago, 11 miles west of Tela, Lowden 16 (OS).

P. parviflora 8 Panama, Province Herrera, Los Llanos de Santa Maria, Lowden.20 (OS) and Province Panama, between Chepo and Pacora, 7 miles from Chepo, Lowden 21 (OS). subg. Reussla P. rotundifolia 16 Honduras, Dept. Comayagua, Lago Yojoa, Pito Solo, Lowden 12 (OS); Nicaragua, Dept. Granada, near Granada, Lowden 17 & 18 (OS); Costa Rica, Province Guanacaste, 1 km. from Arenal, 2 kms. from Trenadora, Lowden 19 (OS); El Salvador, Dept. La 201

Liberated, Laguna Zapotitan, Lowden 23 (os).

Material examined of subgenus Pontederla and subgenus Reusela

Indicates that melosls is normal except for the occasional occurrence of secondary associations of either U or 6 chromosomes (Plate V, b).

Haploid chromosome counts were made of microsporocytes in diaklnesis

(Plate V, a-d). Separation of microspore tetrads occurs after melosls

II. Pollen grains are reniform and smooth walled.

Cytological Investigations indicate a base number of x=8 in

Pontederla. Subgenera of Pontederla may be distinguished cytologically by base numbers, x=8 in subg. Pontederla and 2x=l6 In subg. Reussla.

The polyploid _P. rotund!folia and possibly P. subovata (no cytological material was examined) most likely evolved from an x=8 ancestor.

Pontederla (x=8) has a closer affinity to Eichhornia (x=8) than to

Heteranthera (?n=15, 2n=30 in Heteranthera dubia). Perhaps

Pontederla, Eichhornia and Heteranthera evolved from an ancestral stock of x=8. PLATE V

Chromosome Configurations in Mlcrosporocytes

A. Chromosome configurations (diakinesis) of eight bivalents in Pontederla cordata L. var. cordata from U.S., Ohio, Ottawa Co., Wlnous Point (Lowden 36 & 37).

B. Secondary association of six chromosomes (arrow, three bivalents) in Pontederla cordata L. var. cordata from U.S., Ohio, Ottawa Co., Wlnous Point (Lowden 36 & 37).

C. Chromosome configurations (diakinesis) of sixteen bivalents in Pontederla rotundlfolia L. f. from Nicaragua, Dept. Granada, near Granada (Lowden 17 & 18).

D. Diagramatlc representation of C above, showing sixteen bivalents in Pontederla rotundifolla L. f. from Nicaragua, Dept. Granada, near Granada (Lowden 17 & 18).

202 203

C GEOGRAPHICAL DISTRIBUTION OF FONTEDERIA L. THE AMERICAS

The greater geographical range is occupied by subgenus

Pontederla which Is found 50° North to 35° South of the equator In the

Americas. In North America typical Pontederla cordata (= variety cordata) Is restricted to the eastern provinces of Canada— Ontario,

Quebec, New Brunswick, Prince Edward Island, and Nova Scotia (Fig. 5), whereas In the United States typical P. cordata has an eastern to mid-western distribution (Fig. 6) with greatest concentration along the Atlantic Coastal Plain and the Great Lakes Region as compared with the Mississippi embayment. Pontederla cordata var. lanclfolla is found primarily in the southeastern states (Fig. 6) with extensions into the northern Atlantic seaboard states. In Central America typical

P_. cordata is known only from British Honduras (Figs. 7, 8) with the narrow leaf var. lanclfolla (Fig. 12) found in Cuba, West Indies (Fig.

1 1 ) . In South America, P. cordata and its varieties (Fig. 15) are concentrated along the coasts of southeastern Brazil, southern Uruguay and east central Argentina. In Mexico P. saglttata (Figs. 18, 19, 21) is somewhat restricted to the Isthmus of Tehuantepec between the Bahia de Campeche and Golfo de Tehuantepec (Fig. 20) having an eastern range along the Caribbean coastal plain as far south as Costa Rica in Central

America. Pontederla parvlflora (Figs. 22, 23) is known only from the

Pacific coast of Panama, Golfo de Panama and the Caribbean coast of

Colombia (Fig. 25). 204 205

Of limited geographical range Is subgenus Reussla (Figs. 30,

33) which reaches Its northern limits in British Honduras and extends

Into South America as far south as east central Argentina. Pontederla rotundlfolla (Figs* 27, 31) is found more frequently in lagoon and lake habitats of Central America (e.g., Lago Izabal, Guatemala; Lago

Yojoa, Honduras; Laguna Zapotitan, El Salvador; Lago Nicaragua,

Nicaragua; and Lago Arenal, Costa Rica) than,the coastal plain taxa of

Pontederla sagittata and parviflora of subgenus Pontederla. In South

America P_. rotundlfolla (Fig. 29) is primarily a floristic component of the tributaries of the (Fig. 33). Pontederla subovata is known only from South America (Fig. 33).

Schwartz (1928) made the earliest cartographic attempt to show the distribution of Pontederla in the Americas. Muenscher (1944) mapped the distribution of typical Pontederla cordata and the variety lanclfolla in the United States. In both cases, these maps represent circumscriptions of distributions. Castellanos (1958) mapped the distributions of some Pontederla taxa (typical P. cordata and its var. ovalis, P. subovata) in South America. Geographical distributions

Illustrated in Figures 5, 6, 11, 15, 20, 25, 30, and 33 represent known distributions of Pontederla taxa based on specimens examined from the herbaria cited in the acknowledgments. These distributions are not inclusive and may be supplemented by further herbarium studies with reference to available written floras (e.g., in the United States,

C. C. Dears's Flora of Indiana, A. E. Radford's et al Manual of the

Vascular Flora of the Carollnas, E. L. Braun's The Monocotyledonae

Cof Ohio], J. A. Steyermark's Flora of Missouri and G. N. Jones & G. D. 206

Fuller's Vascular Plants of Illinois), for more complete accounts of distributions within the areas already circumscribed by the study of specimens in these herbaria.

Primarily a genus of the tropical and subtropical Americas.

Pontederla inhabits a wide range of aquatic habitats probably held in check by latitude (temperature) and altitude (elevation). Mountaip ranges as exemplified by the Appalachian (Core, 1966) and Rocky Moun­ tains In North America, Sierras Madre Oriental and Occidental in

Mexico, Sierras del Mico and Minas in Guatemala, Mayan Mountains in

Guatemala and British Honduras, Cordillera Isabella in Honduras,

Cordillera de Talamanca in Costa Rica and the in South America are and have been natural physiographic barriers to the spread of

Pontederla in the Americas. Geographical isolation of taxa is greatest in subg. Pontederla especially of taxa with a Central American range, typical P. cordata (Fig. 11), P_. cordata var. lanclfolla (Fig. 11),

P. saglttata (Fig. 20) and £. parviflora (Fig. 25).

The genetic amplitude of Pontederla is probably such that it

Inhabits the inland freshwaters and marshes transitional to salt waters along the coasts (Chapman, 1960). Field observations support the tolerance of Pontederla to brackish water along the North and Central

American coasts. No populations were observed to grow in salt water of ocean Inlets.

As a member of a family of transhemispheric movements

(Dansereau, 1957), the wide distribution of Pontederla in the Americas, considering the above limitations (latitudp, altitude and aquatic habit), is in part attributed to its mode of seed dispersal by water. 207

The utricle of Pontederla (Plate I, c 6* d) owes its buoyancy and consequently its wide distribution to the light aeriferous tissue of the perigone base (Plates III & IV) surrounding it. Once the cluster of fruits from a single floral stalk ripens, either the peduncle re­ flexes from the weight (Fig. 18; Muenscher, 1944, reports 20,000 seeds per pound), lowering the fruits (Arber, 1920) to the water surface, or the mature fruits abscize and fall into the water. They float for about a week or more (Ridley, 1930) until the light fleshy perigone bases become water logged and sink to the mud below to decay.

Schulz (1942) in "Las Pontederiaceas de Argentina" reports a flotation period longer than fifteen days for seeds enclosed in perigone bases of Pontederla cordata:

En esta especie, la base acrescente del perlgonlo sirve de flotador al fruto, que en esta forma puede ser llevado largo tiempo por las corrientes fluviales, segun demostro A. Burkart (Observac* dlsem. hldrof. de Mimosa, en Notas prellmln. Mus. de La Plata, t. II, pag. 161-162, en Nota; 1934). Este autor obsero que ^tfrutos^ normales flotan mas de 15 dias, mientras los que habian sido

As the seed (Plate II, a & d) germinates, the embryo (Plate II) is nurtured by copious endosperm (Plate II, b). The floating period

(Ridley, 1930) is long enough for the fruit to travel a considerable distance. The presence of fleshy perigone bases surrounding the utricles (Table 2) and the copious endosperm contained Inside greatly

Increase survival expectancy over long distances traveled.

The process of plant establishment by floating seeds (Ridley,

1930):

It is due to the fall of the water that the seeds of plants dispersed by floating become stranded in spots favourable for growth. The rise in the rains, or flood-time, conveys them 208

to the banks and river edge, the fall of the water leaving then to grow. When once rooted, the little plants are Clittle affected] by any rise of which may temporarily cover them.

The distribution of Pontederla rotundlfolla in South America (Fig. 33) reflects and Is representative of this kind of plant establishment process in the genus. When the tributaries of the Amazon Basin flood

the utricles float great distances before the water drops and the seedlings root. The ridged perigone bases, characteristic of

Pontederla taxa (Plates III & IV), "may serve to anchor" utricles

(Ridley, 1930) in the mud.

Seed dispersal by avian and terrestrial animals is thought to be of less frequent occurrence. Seeds of typical Pontederla cordata have been found in the stomachs of Wood Ducks (Aix sponsa) occurring

in North America, Cuba and Mexico, but not recorded as food for other ducks (Ridley, 1930). Carlson and Moyle (1968) report Pontederla cordata as having slight to fair food value for waterfowl. In con­ sidering the biotic importance of hydrophytes, Sculthorpe (196 7) indicates that fruits and seeds of Pontederla have moderate importance as food for ducks; vegetative parts are of moderate Importance to fish as food producers, shade and shelter, of moderate importance as food for cattle, goats and sheep, being of great importance as food to pigs and muskrats. Field observations of Pontederla rotundlfolla at the extinct Laguna Zapotitan, Dept. La Liberated, El Salvador and in the fields of Lago Arenal, Province Guanacaste, Costa Rica suggest ter­ restrial dispersal by animals. Both areas were Inhabited by cattle on which the spinulose perigone bases could easily become attached. 209

Vegetative reproduction by trailing prostrate branched stems

(in subg. Reussia) and by rhizomes (in subg. Reussla and subg. Pontederia)

Is of frequent occurrence throughout populations. Clones (in subgenus

Pontederia) are recognized by clumping of plants within populations

(Fig. 2). Rhizome fragmentation accounts for the establishment of new clones within close proximity. Field observations indicate that the prostrate branched stems of P_. rotundifolta with adventitious roots at each node may easily become severed and rooted. Trailing stems and rhizomes are able to survive harsh environmental pressures as organs of food accumulation during the cold periods of temperate regions and drought periods of the tropics.

Dispersal by water of Pontederia utricles is thought to be the main mode of long range dispersal with vegetative reproduction a j prominent factor in population establishment. In the closely related genus Elchhornla, the air inflated petiole bladders of Eichhornla crasslpes have (Ridley, 1930) been of greater importance in^ong range dispersal of whole plants by water and man. The reproductive biology of Pontederia populations will now be considered in light of a highly adaptable and flexible breeding system, heterostyly. 210

TABLE 2 SEED GERMINATIONS IN GROWTH CHAMBER CONDITIONS (PHOTOPERIOD, 12 HOURS; TQ1PERATURE, 78°F) WITH AND WITHOUT HARDENED PERIGONE BASES

Germinations Without Hardened Perigone Bases

Taxon & Location of # of Days Before Germination Germinations Per Seed Collections A 5 6 7 8 9 13 1A 33 Total Seeds Used

P. cordata var. cordata U.S., Ohio, Ottawa ----11 6/36 Co., Wlnous Point, 3*f miles sw. of Port Clinton off Route 33, Lowden 36 & 37 (OS). Ohio, Licking Co., 0/15 Cranberry Bog, Buckeye Lake, Lowden 33 (OS). ?. sagittate Mexico, State of 1 A 9 3 1 20/27 Vera Cruz, Vera Cruz, Lowden 6 (OS). Honduras, Dept. ---121 11/2A Atlantida, Tela, Lowden 15 (OS). P. parviflora Panama, Province 0/27 Panama, between Chepo and Pacora, 7 miles from Chepo, Lowden 21 (OS). P. rotundifolla Nicaragua, Dept. 0/3 Granada, Lago Nicaragua, near Granada, Lowden 17 & 18 (osH Germinations With Hardened Perigone Bases P. cordata var. cordata (33 days without any germinations) U.S., Ohio, Ottawa 0/50 Co., Winous Point, 3% miles sw. of Port Clinton off Route 53, Lowden 36 & 37 (OS). A TRISTYLOUS GENUS WITH AN ISOLATED FLORAL FORM:

POPULATION REPRODUCTIVE BIOLOGY

The flower Is both adapted to Its present method of functioning and may bear structural evidence of its past. There is far more than the immediate past to be learned from it . • . Francis W. Pennell (1948)

Ornduff (1969) suggested "taxonomists should make an attempt to understand the reproductive methods of the plants with which they work, since such a understanding will strengthen the foundation upon which taxonomic judgments are made." With this in mind, population studies in the United States and Central America were conducted in order to see if the heterostylous condition (tristyly) unique in the

Pontederiaceae might implement strength to the taxonomy of Pontederia.

Vuilleumier (1967) points out that Hildebrand (1867) first used the term heterostyly to describe the breeding mechanism of plants of the same species which have two (distyly) or three (tristyly) different ratios of style length to stamen length present in the flowers. Hildebrand indicated that such plants were self-Incompatible and the difference In the length of floral parts Is a mechanism of insect cross-pollination. Darwin (1877) adopted Hildebrand's termi­ nology and distinguished between legitimate (pollen transfer from a stamen of one floral form to the stigmatic surface of a pistil of equivalent length of another floral form) and illegitimate (pollen transfer from a stamen to the stigmatic surface of a pistil not the 211 212

equivalent length, within or between floral forms) pollinations (Fig.

35). Crosses made by Darwin indicated that legitimate pollinations

produce more viable seeds than illegitimate pollinations.

Accompanying this difference in stamen and style length is a

difference in pollen sire reported in Pontederia by Muller (1871),

Leggett (1875a, b), Darwin (1877), Halsted (1889), Hazen (1918), and

Ornduff (1966). The present study confirms these reports, the largest

pollen being produced by the longest set of stamens. Intermediate

pollen being produced by the medium set of stamens and the smallest

pollen being produced by the shortest set of stamens. A physiological

incompatibility system (Ornduff, 1966; Vuilleumier, 1967) is associ­

ated with tristyly, accounting for legitimate pollinations having greater seed productivity over illegitimate pollinations,

Pontederia is entomophilous; this is characteristic of the

Pontederiaceae (Sculthorpe, 1967). Lovell (1898), Bembower (1911) and Hazen (1918) report four major groups of insects visiting typical

Pontederia cordata populations; Lepidoptera, Hymenoptera, Diptera and Coleoptera, The success of illegitimate and legitimate pollinations

is certainly Influenced by the numerous and wide variety of insect pollinators visiting floral forms,

Vuilleumier*s (1967) survey of heterostyly in the Angiosperms

indicates "that it must have arisen Independently in many different groups. Yet despite Its polyphyletic origin, heterostyly is similar enough in different taxa to strikingly illustrate parallel evolution under (presumably) comparable selection pressures," This is even more

striking since the occurrence of tristyly in the Angiosperms is known (Ornduff* 1969) only in Pontederiaceae (the only Monocotyledonous family)* Oxalidaceae and Lythraceae (Vuilleumier* 1967* reports possibly a fourth occurrence of tristyly in the Rubiaceae). The genetics of distyly is discussed by Vuilleumier (1967); the genetics of tristyly awaits further investigation.

Tristyly, as the term relates to Pontederia, is the type of heterostyly in which three kinds of plants occur in a species: (1) plants with flowers having a SHORT STYLE, 3 medium stamens and 3 long stamens (Sml); (2) plants with flowers having 3 short stamens, a

MEDIUM STYLE and 3 long stamens (sMl); and (3) plants with flowers having 3 short stamens, 3 medium stamens and a LONG STYLE (smL).

These three floral forms are photographed in Plate VI, including a fourth floral form known only, exclusive of the other floral forms, in Pontederia parvlflora of Panama and Colombia (Figs. 23, 25). This fourth floral form (Plate VI, d) has 3 short stamens, 3 long stamens and a LONG STYLE (slL).

Ornduff (1969) states that "in almost every distylous or tri- stylous genus there are plants which have anthers and stigmas at the same level and are self-compatible if not actually self-pollinating."

Even though this fourth floral form (slL) has only three stamens the same height as the LONG STYLE, it is possible that it has been derived from the smL floral form of a heterostylous ancestor. Vuilleumier

(1967) states that "heterostyly has, and is, evolving in two directions one toward dloecy and subdioecy, and the other to homostyly and self­ compatibility." The slL floral form of Pontederia parvlflora might represent a step toward homostyly and increased self-compatibility in 21 4

the genus. Field observations (In Panama) of P. parvlflora (slL) show

that a compatibility system within a single floral form la operating

due to the high seed set of infructescences, however, seed germi­

nations under growth chamber conditions (Table 2) do not reflect the

expected success of such a compatibility system in nature.

Tristyly (Ornduff, 1969) could serve as "a source of taxonomic

confusion because of the striking and discrete nature of the morpho­

logical differences between floral forms of the same species." It

can even lead to descriptions of separate species described from the

three floral forms of the same species (Ornduff, 1964). Tristyly In

the nomenclature! sense has not been a source of confusion in the

taxonomy of Pontederia and this study Indicates that the slL floral

form in Pontederia parvlflora (homostylous member of the genus) is an

excellent key character (since other morphological and geographical distinctions are of little aid in identifying dried specimens) to distinguish this taxon from its closely related heterostylous counter­

part, typical Pontederia cordata.

Ornduff (1966) confirmed Darwin's findings on heterostyly

that legitimate pollinations are more productive of seeds than il­

legitimate pollinations. The results of Ornduff's breeding program using typical Pontederia cordata indicate that self-incompatibility

(self and own-form pollinations ore illegitimate. Fig. 35) is "con­

siderably stronger" in smL (maximum of 18.7% seed production) and

Sml (maximum of 12.7% seed production), than in the sMl (maximum of

53.9% seed production) floral form (since there Is one seed per utricle,

"fruit production" used by Ornduff is interpreted as seed production). 215

Regarding legitimate pollinations (Fig. 35), Ornduff found that the sMl floral form had the greatest percentage seed production (82.7%), followed by smL (71.17.) and Sml (61.3%). Because there was hardly any difference (Ornduff, 1966) In the percent seed production from each of the two possible legitimate pollinations of each floral form

(Fig. 35), the results indicated that pollen from one anther set was physiologically equivalent to pollen from the same-level anther set of another floral form (e.g., pollen from 3 long stamens of Sml equals that of the 3 long stamens of sMl).

Through the study of the frequency of floral forms within populations of different Pontederia taxa (Table 3), an indication of the breeding system of individual populations may be extrapolated from the results of Ornduff's research with typical Pontederia cordata in the United States. Field and herbarium studies in the United States and Central America (Table 3) indicate the presence of three floral forms (Sml. sMl and smL) in typical Pontederia cordata and its varieties, P. sagittata, P. rotundifolia and P. subovata, exclusive of P. parvlflora which has only the slL floral form. Pontederia taxa are tristylous except P_. parvlflora which is homostylous.

In populations where all three floral forms are present (Table

3), it is difficult to say if any one floral form has greater breeding value compared with another floral form in the same population, since the samples were quite small. However, In Pontederia rotundifolia, three out of four populations (Table 3) had the frequent occurrence of one floral form in the absence of the other two. In each case, it was not the same floral form that was of frequent occurrence (e.g., smL 216

floral form Is found at Lago Nicaragua, Granada, Nicaragua and is

absent at Lago Arenal, Costa Rica where Sml floral form was found).

Barring the possibility that the samples are Inadequate, perhaps these

populations represent clonal establishment of a single floral form.

Vuilleumier (196 7) reports that "separate populations of one

species and individuals within one population which are either homo-

or heterostylous, indicate that heterostylous species can maintain a

balance between inbreeding and outbreeding." In Pontederia rotundi­

folia populations where one floral form was found, a certain amount

of inbreeding must be occurring through Illegitimate pollinations

(selfing and own-form pollinations). Even though an incompatibility

system Is operative there is not total incompatibility in a population

containing a single floral form. Ornduff's results (1966) indicated

for typical Pontederia cordata a maximum of 53.9% seed production for

an average of self and own-form cross pollinations of the siMl floral

form. This flexibility in P. rotundifolia and P. parvlflora is

indicative of heterostyly. a highly adaptable reproductive system

(Vuilleumier, 1967).

In Pontederia heterostyly functions in juxtaposition with an

effective mode of vegetative reproduction, stem fragmentation; a

reproductive system has evolved which is highly tolerant to different

pressures of the environment. Subgenua Reussla is characterized by having fewer flowered inflorescences (implying less seed production)

and long trailing stems (Figs. 27, 32). Vegetative reproduction

especially in populations having one floral form (Pontederia rotundi­

folia) has greater Immediate survival value than reproduction through 217

a heterostylous breeding system# In Bubgenus Pontederia just the reverse has occurred. Inflorescences are many flowered (implying greater seed production) and the erect above ground stems are much

shorter. In this subgenus tristyly and possibly homostyly (P_. parvl­

flora) have greater long range survival value.

In the foregoing discussion, tristyly was reported in all

Pontederia taxa except the homostylous P_. parvlflora. The smL floral

form of a heterostylous ancestor is suggested as the origin of the slL

floral form restricted to P. parvlflora. The discovery of the slL

floral form has aided in the distinction of P_, parvlflora from its heterostylous counterpart, typical Pontederia cordata. Heterostyly as a highly flexible breeding system and vegetative reproduction as

an effective agent of population establishment are considered eminent

factors in the survival of this widely dispersed aquatic plant genus. I

Fig. 35. Legitimate and illegitimate pollinations in Pontederia L. floral forms, arrows represent the transfer of pollen from anthers to stigmas, code to floral forms (Sml) SHORT STYLE, 3 medium stamens, 3 long stamens; (sMl) 3 short stamens, MEDIUM STYLE, 3 long stamens; (smL) 3 short stamens, 3 medium stamens, LONG STYLE; (slL) 3 short stamens, 3 long stamens, LONG STYLE.

218 LEGITIMATE

ILLEGITIMATE PLATE VI

Floral Forms

A. SHORT STYLE, 3 medium stamens, 3 long stamens (Sml): flower of Pontederia rotundifolia L. f. from Lago Arenal, Costa Rica, (Lowden 19), length 10 mm.

B. 3 short stamens, MEDIUM STYLE, 3 long stamens (sMl): flower of Pontederia rotundifolia L. f. from Pito Solo, Lago Yojoa, Honduras (Lowden 12) , length 13 mm.

C. 3 short stamens, 3 medium stamens, LONG STYLE (smL): flower of Pontederia rotundifolia L. f. from Granada, Lago Nicaragua, Nicaragua (Lowden 17 & 18), length 11 mm.

D. 3 short stamens, 3 long stamens, LONG STYLE (slL): flower of Pontederia parvlflora Alex, from between Pacora and Chepo, Panama (Lowden 21), length 7 mm.

220 2 2 1 TABLE 3 INDICATION OF FLORAL FORM FREQUENCY IN PONTEDERIA L. BASED ON POPULATION AND HERBARIUM STUDIES

Floral Form & # (%) Inflorescences Population Study Taxon Site (vouchers) Sml sMl smL slL Totals

P. cordata U.S., Ohio, Ottawa Co., 8(33.3) 5(20.8) 11(45.8) 0(0.0) 24(99.9) var. Winous Point, 3^ miles cordata SW of Port Clinton, western end of Sandusky Bay, Bay Twp. Cent. SE k Sec. 22 (Lowden 36 A 37; 27 June, 10 August 1969). British Honduras, C.A.: 5(62.5) 2(25.0) 1(12.5) 0(0.0) 8(100.0) 9 miles N of Belize City along Northern Hwy. (Lowden 24, 24 Dec. 1969); 29 miles N of Belize City 0(0.0) 0(0.0) 3(100.0) 0(0.0) 3(100.0) along Northern Hwy., Cow- head Creek (Lowden 26; 30 Dec. 1969); 32 miles N of Belize City 2(66.7) 1(33.3) 0(0.0) 0(0.0) 3(100.0) along Northern Hwy. (Lowden 27; 30 Dec. 1969); Hattieville-Burrell Boom 0(0.0) 2(40.0) 3(60.0) 0(0.0) 5(100.0) Road, 5h miles from Burrell Boom (Lowden 28, 30 Dec. 1969). 222 TABLE 3 (continued)

Floral Form & # (7.) Inflorescences Population Study Taxon Site (vouchers) Sml sMl smL slL Totals

var. herbarium specimens 4 land* U.S., Florida, Holmes folia Co., on U.S. 90, 0.2 ml. E of Florida 81 In Ponce de Leon, D. E. Stone 2589 Sml (DUKE 199478, NY), 2591 sMl (DUKE 199476, NY), and 2590 smL (DUKE 199477, NY), 1 May 1968. var. herbarium specimens 4 ovalis Brazil, Mato Grosso, 300 km. past Cuiaba in route to Goiania, 600-700 m. alt., B. Maguire, Plres, C. K. Maguire and Silva 56931 Sml (US 2513755), 29 Sept. 1963; Colombia, Dept. Boyaca (eastern), near Orocue, alt. 140 m., Oscar Haught 2716 sMl (US 1707263), 29 March 1939; Brasil, exploration of New Brazilian Highways, Brasilia, Federal District, vicinity of Sobredlnho, G. Prance and N. Silva 59082 smL (US 2457328). 19 Sept. 1964. 223 TABLE 3 (continued)

Floral Form & # (7*) Inflorescences Population Study Taxon Site (vouchers) Sml sMl smL slL Totals

£. Mexico, State of Vera 46(33.8) 52(38.2) 38(27.9) 0(0.0) 136(99.9) sagittate Cruz: Vera Cru2 , El Coyol, Prolongacion Carretera Aleman (Lowden 6 ; 25 July 1970); Carretera Nacional 180, 5(55.6) 1(11.1) 3(33.3) 0(0.0) 9(100.0) El Puente de Teculapilla, 5 kms. ante de llegar a nw. Lerdo (Lowden 7; 26 July 1970); Laguna Catemaco, Arroyo 10(76,9) 3(23.0) 0(0.0) 0(0.0) 13(99.9) Agrio, 6 kms. from Catemaco Town between Coyame and San Andres Tuxtla (Lowden 9; 27 July 1970). Guatemala, Dept, Izabal: 2(9.1) 12(54.5) 8(36.4) 0(0 .0) 22(100.0) Quirigua, swampy area by railroad, in front of abandoned hospital (Lowden 10; 31 July 1970); Puerto Barrios, in swampy 10(32.3) 14(45.2) 7(22.2) 0(0.0) 31(99.7) area (Lowden 11; 1 August 1970). TABLE 3 (continued)

Floral Form & # (7.) Inflorescences Population Study Tax on Site (vouchers) Sml sMl BmL slL Totals

Honduras: Dept, of 0(0.0) 6(37.5) 10(62.5) 0(0,0) 16(100.0) Cortes, Puerto Cortes, 3 Ave*-5 Calle 0 (Lowden 13; 9 August 1970); 13.5 miles inland from 16(47.1) 7(20.6) 11(32.3) 0(0.0) 34(100.0) Puerto Cortes, 1 mile inland from Bijao- Cemento Hondurenas Co., along main road (Lowden 14; 9 August 1970). Dept. Atlantlda, Tela, 15(34.1) 16(36.4) 13(29.5) 0(0.0) 44(100.0) drainage ditch by rail­ road, where main road passes over the track, "La Curva" (Lowden 15; 10 August 1970); Santiago, 11 miles W 21(55.3) 12(31.6) 5(13.2) 0(0.0) 38(100.1) of Tela, drainage ditch along main road (Lowden 16; 10 August 1970T. P. Panama: Province of 0(0.0) 0(0.0) 0(0.0) 89(100.0) 89(100.0) parvlflora Herrera, Los Llanos de Santa Marla, 2 kms. at Santa Marla and 3.5 kms. at El Escota from the International Highway 225 (Lowden 20; 24 August 19707: TABLE 3 (continued)

Floral Form & # (%) Inflorescences Population Study Taxon Site (vouchers) Sml sMl smL slL Totals

Province of Panama, 0(0.0) 0(0.0) 0(0.0) 83(100.0) 83(100.0) along International Hwy., between Pacora and Chepo, 7 miles from Chepo, Tapagara (Lowden 21; 26 August 1970). 17 flowering herbarium specimens cited under specimens examined. P_. rotun- El Salvador, Dept, of La 0(0.0) 0(0.0) 29(100.0) 0(0.0) 29(100.0) dlfolla Libertad, extinct Laguna Zapotitan (Lowden 23; 26 March 1970). Honduras, Dept, of 0(0.0) 30(81.1) 7(18.9) 0(0.0) 37(100.0) Comayagua, S end of Lago Yojoa, Pito Solo (Lowden 12; 8 August 1970). Nicaragua, Dept, of 0(0.0) 1?(0.7?) 145(99.3) 0(0.0) 146(100.0) Granada, frente a la Terraza, near Granada (Lowden 17 & 18; 14 & 16 August 1970). Costa Rica, Province of 184(100.0) 0(0.0) 0(0.0) 0(0.0) 184(100.0) Guanacaste, along main road to Arenal, 1 km. from Arenal, 2 kms. from Tronadora (Lowden 19; 18 August 1970). TABLE 3 (continued)

Floral Form & # (7.) Inflorescences

Taxon Site (vouchers) Sml sMl smL slL Totals

P. herbarium specimens + + + subovata Paraguay, prope Con­ cepcion, E. Hassler 7352 Sml (NY), Sept. 1901/1902; Brazil, Estado Minas, Mun. Ituiutaba, Loc. S. Tere- zulha, A. Marcedo 3168 &11 (US 2026533), 18 Feb. 1951; Argentina, Proy Chaco, Loc. Colonia Benitez, alt. 55 m., C. Meyer 3864 srnL (F 1519935), 10 Feb. 1941.

Totals 324(30.3) 164(25.0) 294(34.7) 172(10.0) 954(100,0) Sub Total 324(33.7) 164(27.7) 294(38.5) 782(100.0)

Code to floral forms: (Sml) SHORT STYLE, 3 medium stamens, 3 long stamens; (sMl) 3 short stamens, MEDIUM STYLE, 3 long stamens; (smL) 3 short stamens, 3 medium stamens, LONG STYLE; (slL) 3 short stamens, 3 long stamens, LONG STYLE. A plus sign (+) indicates the occurrence of a floral form, whereas a hyphen (-) indicates a floral form unknown in a taxon. 227 STATISTICAL ANALYSIS OF PHENOLIC QUALITATIVE

FEATURES: .CHROMATOGRAPHIC DATA

In modern systematica, variation la often a valuable asset . . . Ralph E. Alston (1967)

In recent years considerable attention has been given to phenolic compounds, their occurrence, biochemistry and biogenetics.

Phenolic compounds compose a large group of substances possessing an aromatic ring bearing a hydroxyl substituent (Thomson, 1964), The combination of simple phenolic units are referred to as polyphenolics which are collectively known as flavonolds. Phenolic compounds have been the subject of a large number of biochemical systematic studies

(Alston, 1967), owing to their abundance, chemical diversity, metabolic stability, biogenetlcal relatedness and wide distribution in vascular plants. McClure's (1970) synopsis of "Secondary Constituents of

Aquatic Angiosperms" includes the following compounds found in the

Pontederiaceae; Pontederia cordata, alkalolds-none, terpenoids-no saponins (Hegnauer, 1963); P. lanceolata,simple phenolics-caffeic acid, para-coumaric acid, slnaplc acid, ferulic acid, flavonoidB- cyanidin, delphinidin (Bate-Smith, 1968); Eichhornia speclosa, alkaloids-conflictlng reports, terpenoids-triterpenes (a conflicting report), simple phenolics-caffeic acid, ferulic acid, flavonoids- dephinidin 3-diglucoside, cyanidin, leucoanthocyanin ? (Hegnauer,

1963; Willaman & Schubert, 1961; Bate-Smith, 1968; Shlbata, 1965;

228 229

Bate-Smith & Swain, 1965).

The environment of aquatic plants plays an important role in the metabolic changes influencing qualitative and quantitative pro­ duction of secondary constituents. The capacity of an organism to withstand anaerobic conditions (Sculthorpe, 196 7) would certainly have an effect on the production of secondary constituents (McClure,

1970). Laing (1940) reports remarkably prolonged endurance of

Pontederia cordata rhizomes to anaerobic conditions, in which even new shoots developed in pure nitrogen.

Aquatic plants lend well to phytochemical systematic studies

(McClure, 1970), since they "do not appear to produce the bewildering array of secondary constituents found in terrestrial plants." In the

Lemnaceae phytochemical studies have contributed much to the bio­ synthesis and biochemical systematica of secondary metabolites in a group of aquatic plants (McClure, 1970).

Much of the earlier work (Alston, 1967) using chromatographic data of phenolic compounds does not include chemical characterization.

The merit and validity of empirical chromatographic data is that color and position of phenolic compounds, evidenced as spots on chromato­ grams, can show systematic relationships without necessitating chemical characterization. Objectives of this applied empirical approach in the following chromatographic study of phenolic compounds in Pontederia were (1) to learn the degree of variation in phenolic patterns in indi­ vidual plants within and between populations of Pontederia taxa, (2) to discover if there is significant identity between compared phenolic patterns in Pontederia taxa, (3) to see if there is any similarity of 230

phenolic patterns found In Pontederia compared with those found In

Elchhornla and Heteranthera (genera of the same family) and (4) to note the significance of phenolic patterns as they relate to pre­

conceived taxonomy suggestive of morphology, reproductive biology,

geographical distribution and cytology.

Methods

The descending two-dimensional chromatographic procedure

followed in the extraction of phenolic compounds is essentially that

outlined by Mabry, Markham and Thomas (1970). Phenolic compounds were

extracted from dried leaf material (in practically all cases it was

the flower stalk leaf, Fig. I) in 50% aqueous-methanol. Vials of

extracts were placed on a shaker for at least 24 hours before 4 ml of

individual leaf extracts were spotted, in the conventional manner, on

Whatman 3MM chromatographic paper (46 x 57 cm). Jet streams of warm

air from hair dryers aided in the drying of spots during the spotting

process. The two-dimensional solvent systems employed: (1) TBA 3 parts reagent-grade tertiary butanol:1 part reagent-grade glacial

acetic acld:l part distilled water and (2) HOAc 15 ml of reagent-grade

glacial acetic acid mixed with 85 ml of distilled water. The solvent

systems were placed in trays of TBA and HOAc designated chromatocabs

in which a maximum of eight chromatograms were run. It took approxi­ mately 25 hours for the TBA solvent system to run the first dimension.

Chromatograms were removed and dried before the second dimension run

in HOAc. This second run took approximately 5 hours. Chromatographic

patterns were first read using no reagent in visible and ultraviolet 231 light, after which they were read in the presence of NH40H vapor in ultraviolet light*

Procedure and Results

Twenty-nine spots (Fig. 36) characterized by their position and color (Table 4) were discernible after each chromatogram had been read using no reagent in visible and UV light, and with NH40H vapor in UV light. Color characterizations (Table 4) were uniform throughout chromatograms, however for spots with more than one color indicated, it is possible that more than one phenolic compound is present having approximately the same value

distance between origin and the center of concentration of the (R^. » ______phenolic spot ) . distance between origin and solvent front

Rj values and rechromatography of eluates for composite color spots were not attempted. Spot occurrence of phenolic compounds for indi­ vidual leaves and collection sites for taxa are recorded in Appendix

I (Table 7). It was necessary to subdivide numbers 11 and 15 (Fig. 36) because color and close proximity of spots prevented contrasting delineations.

Summary phenolic patterns for each taxon were compiled using the total number of spots found for each taxon (Table 5)• Nine spots were common to Pontederia taxa examined and seven out of these nine spots were shared equally with Heteranthera and Eichhornia taxa.

Spots 15b and 22 (Table 5) were found to occur in single taxa. No spots were added to the survey by the addition of Eichhornia and

Heteranthera taxa. The average number of spots per chromatogram 232

(Table 5) was considerably less than the total number of spots representative of summary phenolic patterns for each taxon.

Infra- (Pontederia only) and inter- generic comparisons

(Pontederia with Eichhornia and Heteranthera, Table 6) of summary phenolic patterns (Table 5) were based on (1) positive matches (a spot occurring in two taxa compared; a spot restricted to two taxa only is a qualified positive match), (2) negative matches (the absence of a spot in two taxa compared) and (3) differences (a spot occurring in one of the taxa compared but not in the other). The degree of re­ lationship between pairs of taxa was calculated in two ways, (l) the coefficient of similarity ^ P ^ (or paired affinity index) and (2) the matching coefficient p ^ n"+"d * eac^ coefficient, p represents the positive matches, n the negative matches and d the differences.

Each coefficient may vary from 0 (no resemblance) to 1 (identity).

Despite the consistent higher matching coefficients, the coefficient of similarity, in which positive matches have more weight over negative matches, is considered to have precedence from a biological point of view. A negative match is a weaker indication of chemical identity

(Runemark, 1968) due to different inherent features unpredictable both

In the plants and the system of analysis.

Discussion

Chromatographic patterns were responsive to analytical inter­ pretation, however limitations of Infra- and inter- specific variation in Pontederia were not possible. Perhaps variation in individual phenolic patterns (Appendix I, Table 7) reflect leaf polymorphism, since similar leaf shapes were found in each of the major taxa (e.g., 233

compare the sagittate leaves of Pontederia sagittata, Fig. 21 and

Pontederia rotundifolia. Fig. 31). This might account for a qualified

positive match (spot 26) between these two taxa (Tables 5-6). This

relationship was not investigated.

The close morphological resemblance between Pontederia taxa

was reinforced by the high matching and similarity coefficients

(Table 6; Fig. 37). Infrageneric (Pontederia) comparisons (Table 6;

Fig. 37) indicate higher resemblances between major taxa (typical P.

cordata, £. sagittate, P. parvlflora and P. rotundifolia) than minor

taxa (P. cordata varieties lanclfolla and oval is). Intergeneric

comparisons (Table 6) indicate a higher identity between Pontederia

and Eichhornia, than Pontederia and Heteranthera. This relationship

is further substantiated by a qualified positive match (Table 5, spot

21) between typical Pontederia cordata and Eichhornia (E. azurea and

E. craasipes).

The greatest variation in phenolic patterns is found in typical

Pontederia cordata and £. sagittata, each containing an isolated spot

(Table 5). spot 15b and 22 respectively (Fig. 36). These spots are

not thought to represent Isolated metabolic systems since in other

taxa it is very possible they might be present in smaller amounts, masked by overlapping adjacent spots (Fig. 36).

Seven spots were common to all taxa (Table 5) and no spots were added to the survey by members of Eichhornia and Heteranthera.

Undoubtedly similar metabolic systems have evolved in the Pontederi-

aceae, perhaps divergently from an ancestral phenolic system comparable

to that found in Pontederia cordata and P. sagittate. Correlation of phenolic patterns with partial geographical isolation of taxa (Figs.

5, 6, 11, 15, 20, 25, 30, 33) indicates a possible loss of metabolic systems, indicative of fewer numbers of common spots in Pontederla taxa as compared with typical P_. cordata and saglttata. A thorough survey of phenolic systems in familial taxa, beyond the empirical approach is necessary to the proof of this evolutionary affinity. f

Fig. 36. Summary two-dimensional paper chromatographic profile of phenolic compounds found in Fontederla L. and related genera.

235 1

11a ✓

11b 237

TABLE 4 SPOT COLORS OF PHENOLIC COMPOUNDS IN PONTEDERIA L. AND RELATED GENERA

No Reagent Reagent NH,OH Spot No. Visible Light UV Light UV Light

I aqua green (in part) 2 - aqua green (in part) 3 — dark purple yellow-green-brown (in part) 4 - cream-yellow - 5 - blue purple - 6 - green blue-cream purple - 7 - white blue - 8 - aqua - 9 - dark blue (royal blue) - 10 - aqua green (sometimes) 11a - a) rose cream - lib - b) blue green-pale blue - 11c - c) blue purple - 12 green brown green yellow* red brown- pink purple 13 green brown pink purple-bright pink - 14 blue purple* blue-white blue - 15a - a) yellow green brown - 15b - b) yellow - 16 pink (some­ bright pink-dull pink — times) 17 - dark purple - 18 - green blue cream-cream purple - 19 - dull pink - 20 — cream yellow green brown — green blue 21 - salmon - 22 - yellow green brown - 23 - yellow-gray cream - 24 - pale blue - 25 - dark yellow green brown Urn 26 - dull pink purple - TABLE 5 SUMMARY OF SPOT OCCURRENCE OF PHENOLIC COMPOUNDS IN PONTEDERIA L. AND RELATED GENERA*

Spot Number Taxon 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b 16 17 18 19

Pontederla cordata + + + + + + + + + + + + + + + + + + + + + + jvar. cordata Pontederla cordata + + + + + + + + + + + - - + + - + - + - - - var. lancifolla Pontederla cordata + + + + + - + + + - - + + + + - - - + var. ovalIs Pontederla saglttata + + + + - + + + + + + + + + + + + - + - + + Pontederla parvlflora + + + + + + + + + + + - + + + + - - + - - - Pontederla rotund If ol la + + + + + + + + + + + - - + + + - - + + - + Heteranthera ltmosa + - - + - + + + + + + + - - - + - + - Elchhornla azurea & + + + + - - + + + + + + + - - - + - - + ]S. crasslpes spots shared by + + + + - - + + + + + ------Pontederla taxa spots shared by + - - + - - + + + + + ------Pontederla taxa, Heteranthera llmosa and Elchhornla azurea & E, crasslpes spots occurring in a ------+ .... single taxon 238 TABLE 5 (continued)

Spot Number Number of Average Number of Spots Taxon 20 21 22 23 24 25 26 Spots Per Chromatogram

Pontederla cordata + + - + + + - 27 15.90 var. cordata Pontederla cordata - - - + - + - 17 12.75 var. lancifolia Pontederla cordata + - - - - - 14 14.0 var. ovalis Pontederla sagittate + - + + + + + 25 10.41 Pontederla parviflora + - - + + - 19 7.62 Pontederla rotundifolia + - - - + + + 21 13.52 Heteranthera limosa - - - - + - 12 11.0 Elchhornla azurea & + + -- - - 15 11.75 E. crasslpes spots shared by * _ ■ • 9 Pontederla taxa spots shared by 7 Pontederla taxa, Heteranthera limosa and Elchhornla azurea & E. crasslpes spots occurring in a • + m • • 2 single taxon 239 *+ sign represents the presence of phenolic compounds Based on position and color of spot on chromatogram. TABLE 6 f A COMPARISON OF TWO DIMENSIONAL PAPER CHROMATOGRAPHIC PROFILES OF PONTEDERLA L. AND RELATED GENERA BASED ON SUMMARY OF PHENOLIC SPOT OCCURRENCE (TABLE 7)

Positive Matches Negative Coefficient of Matching Taxa Compared Normal Qualified Matches Differences Similarity Coefficient

Pontederia cordata var. cordata-var. lancifolia 17-2 10 0.63 0.66 var. cordata-var. ovalis 14-2 13 0.52 0.55 var. cordata-sp. sagittata 23 6 0.79 0.79 var. cordata-sp. parvlflora 19-2 8 0.70 0.72 var. cordata-sp. rotundlfolia 20-1 8 0.71 0.72 var. cordata-Heteranthera limosa 12-2 15 0.43 0.48 var. cordata-Elchhornia azurea 15 1 2 12 0.56 0.59 & E. crasslpes var. lancifolia-var. ovalis 10 - 8 11 0.48 0.62 var. lancifolla-sp. sagittata 16 - 3 10 0.62 - 0.66 var. lancifolia-sp. parviflora 15 - 8 6 0.71 0.79 var. lancifolia-sp. rotundifolia 15 - 6 8 0.65 0.72

var. lancifolia-Heteranthera 11 - 11 7 0.61 0.76 limosa

var. lancifolia-Eichhornia 12 - 9 8 0.60 0.72 azurea & E. crasslpes

to O TABLE 6 (continued)

Positive Matches Negative Coefficient of Matching Taxa Compared Normal Qualified Matches Differences Similarity Coefficient

var. ovalis-sp. sagittata 12 - 2 15 0.44 0.48 var. ovalis-sp. parvlflora 12 - 8 9 0.57 0.69

var. ovalis-sp. rotundifolia 13 - 7 9 0.59 0.69

var. ovalis-Heteranthera limosa 7 - 10 12 0.37 0.59 var. ovalie-Eichhornia azurea 10 10 9 0.53 0.69 & E. crasslpes

Pontederla sagittata sp. saglttata-sp. parvlflora 18 3 8 0.69 0.72 sp. saglttata-sp. rotundifolia 19 1 2 8 0.70 0.72 sp. saglttata-Heteranthera limosa 12 - 4 13 0.48 0.55 ap. sagittata-Eichhornla azurea 14 3 12 0.54 0.59 6 E. crasslpes

Pontederla parvlflora sp. parviflora-sp. rotundifolia 18 * 7 4 0.82 0.86 sp* parviflora-Heteranthera 11 - 9 9 0.55 0.69 limosa sp. parviflora-Elchhornla azurea 13 - 8 8 0.62 0.72 & E. crassy>es TABLE 6 (continued)

Positive Matches Negative Coefficient of Matching Taxa Compared Normal Qualified Matches Differences Similarity Coefficient

Pontederla rotundifolia sp. rotundifolia-Heteranthera 11 7 11 0.50 0.62 limosa sp. rotund!folia-Eichhornla 14 7 8 0.64 0.72 azurea & E. crasslpes

Heteranthera limosa-Eichhornia 9 11 9 0.50 0.69 azurea & E. crasslpes Z*?Z Fig. 37. Graphic representation comparing two dimensional paper chromatographic profiles of Pontederla L. taxa based on coefficients of similarity (Table 6).

243 CORDATA

.79

ROTUNDIFOLIA

.52

.63 ovalis

SAG ITTATA lancifolia .82

.69

p a r v if l o r a BIOSYSTEMATIC ASPECTS: EVOLUTIONARY IMPLICATIONS

A mere speculative hypothetical positioning of Pontederla,

Elchhornla and Heteranthera in relation to a common line of origin is attempted based on the present investigations. The immediate ancestral stock is hypothesized as aquatic, tropical in origin, in­ florescence a many-flowered raceme, fruit a many-seeded capsule, axile placentation, heterostylous, 6 stamens, flowers actlnomorphlc and perianth parts mostly equal and separate. Cytological evidence indi­ cates a haploid base chromosome number of Xs8. Chromatographic pat­ terns of phenolic compounds suggest that similar metabolic systems have evolved within these genera of the Pontederiaceae. These same studies indicated a closer relationship between Pontederla and

Elchhornla. A conceivable line of evolution (Fig. 38) follows, for which the arrow indicates a progression from a primitive to a derived condition. Characters in parentheses represent divergent evolution

(Fig. 38) from the above hypothetical ancestor:

Primitive Derived______m

Heteranthera Elchhornla Pontederla (mostly equal perianth (unequal perianth parts (unequal perianth parts basally connate) basally connate) parts basally connate) (3 stamens) 6 stamens 6 stamens many seeded capsule many seeded capsule (1 seeded utricle) (parietal placentation) axile placentation (parietal placentation) actlnomorphlc flowers (zygomorphic flowers) (zygomorphic flowers) l^many flowered inflo- many flowered inflo- numerous flowered in- rescence rescence florescence

245 246

Paleolimnological studies have traced Pontederla to the Eocene.

Knowlton (1922, Pontederltes Knowlton, Plate XXXVI, fig. 6, based on a

’’well-preserved upper part of a leaf") and Bradley (1963) found

Pontederla part of the lake margin fossil flora of the Green River

Formation (Kulp, 1961, of early and middle Eocene age, about 50 million years ago) in northwestern Colorado and the Gosiute Lake in south­ western Wyoming. Bradley (1929, 1963) had deduced an average annual temperature of 66.5°F. "This was based on an elaborate statistical

Btudy by Brooks (1926, . . . ) of the effects upon climate produced by the relative distribution of land and sea, the altitude of the land, the direction and strength of ocean currents, and the relative amount of volcanic activity. The large and varied flora of the Green River

Formation, according to paleobotanists, indicates a climate comparable > to that now found in the Gulf Coast States. It is interesting, there­ fore, to note that the present mean annual temperature along the Gulf

Coast is about 70°F."

By the Lower Eocene, Pontederla had spread forming a continuous range from South America into North America via the Middle American landbridge. On several occasions (Schuchert, 1935, from the Upper

Eocene to the Lower Pliocene) portions of Middle America were under water. Pontederia probably evolved in isolated areas as this isthmian landbridge rose and fell. The environment acted as the promoter of speciation. This Is especially the case in Middle America where speciation took place in areas of tension (Baker, 1970, from which the following statements are paraphrased) as a result of climatic instability, mountain-building, and volcanic activity. Local 247 conditions changed rapidly so that geographical distributions of

Pontederla also changed rapidly, Taxa were alternately isolated and brought into contact, giving opportunities for hybridization under strong selective stresses. Under these circumstances, speciation could have taken place at a high-rate although extinction could have occurred frequently. Some of the products of this speciation in the tension areas were suitable for life in more stable areas nearby into which they migrated and persisted until today.

In North America Pontederla had a more western distribution as indicated by the Green River Formation of Eocene Age. According to Knowlton (1922) the Green River Formation was '’bordered on the north and west by what are now lofty mountain ranges CRocky Mountains], though they were then of far less elevation. Its containing barriers on the east and south are less recognizable . . •" During the Eocene the Gulf Coastal Plain was under water (Schuchert, 1935). As the waters receded Pontederla was redistributed to the warmer subtropical climate of the Gulf Coastal Plain where it spread throughout the

Atlantic Coastal Plain of North America. The adjacent Appalachian

Mountains were and still are the main obstacle to its spread into the interior of eastern United States (Core, 1966).

The advent of the Pleistocene brought deleterious climatic changes resulting in a restricted southerly distribution of Pontederla toward the warmer climate of the Gulf Coast. As the climate fluctuated during glacial and interglacial periods, fluctuations also occurred in the distribution of Pontederla along the Atlantic Coast. Glacial withdrawal (Sculthorpe, 196 7) brought an Increasing warmth accompanied 248 with an optimum post glacial maximum warmth, culminating In today's climatic conditions. The retreat of the last lce-sheets left behind numerous lakes, bogs and ponds. Since the Wisconsin glacier,

Pontederla has spread and thrived throughout these aquatic glacial remanents of eastern North America reaching Its present northern and northwestern limits within relatively recent times.

4 Fig. 38, Speculative scheme of evolution. Arrows represent divergent evolution from a hypothetical ancestral stock.

249 250

?

\

Heteranthera

Eichhornia

Pontederia

P. cordata R subovata var. laneifolia P. sagittata Rparviflora fvar. oval is R rotundifolia var. cordata APPENDIX I

TABLE 7 SPOT OCCURRENCE OF PHENOLIC COMPOUNDS IN PONTEDERIA AND RELATED GENERA AS EXEMPLIFIED BY POPULATION STUDY AND HERBARIUM SPECIMENS EXAMINED

251 TABLE 7 SPOT OCCURRENCE OF PHENOLIC COMPOUNDS IN PONTEDERIA AND RELATED GENERA AS EXEMPLIFIED BY POPULATION STUDY AND HERBARIUM SPECIMENS EXAMINED*

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

Pontederla cordata u + 4 4 4 4 4 4 4 4 4 4 -- 4 4 4 4 - var. cordata IB + 4 4 4 4 4 4 4 4 4 4 - - 4 4 - 4 - U.S., Maine, York Co., 2A 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 - North Berwick, Lowden 1 (OS), New Hampshire, 2B + 4 4 4 4 4 4 4 4 4 4 4 - 4 4 - 4 - Strafford Co., Bellamy 2D + 4 4 4 4 - 4 4 4 - 4 4 4 4 4 4 4 - River, Lowden 2 (OS), Maine, York Co., Kenne- 2E + 4 4 4 4 4 4 4 4 4 4 4 - 4 4 4 4 -

’4A + 4 4 4 4 4 4 4 4 4 4 - - 4 4 - 4 - Alabama, Baldwin Co., 4 miles E of Mobile, 4B + 4 4 4 4 4 4 4 4 4 4 - - 4 4 - 4 - Lowden 5 (OS); British 5A 4 4 4 4 - - 4 4 4 4 4 -- 4 4 - - - Honduras,Belize Dist., 9 miles N of Belize 5D + 4 4 4 -- 4 4 4 - - - - 4 4 4 - - City, Lowden 24 (OS), 24F + 4 4 4 - - 4 4 4 4 4 4 - 4 4 7 - 4 29 miles N of Belize 24H + 4 4 4 4 - 4 4 4 - 4 4 - 4 4 4 - 4

241 4 -- 4 - 4 - - 4 - 4 - - 4 4 - - - Lowden 27 (OS), Hattie- ville-Burrell Boom Rd., 24J + -- 4 - 4 - - ? - 4 - - 4 4 - - - Lowden 28 (OS); U.S., 24K + __ 4 4 - ? — 4 - 4 4 4 - Maine, Sagadahoc Co., 1 mile from Topsham, Lowden 29 (OS), Foreside road near Topsham, Lowden 30 (OS), Lincoln Co., 3 miles S of New­ 252 castle, TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

1A + 16 IB + ------+ - - - 16 2A + - 7 + + -- - - - m 20 2B + - - - •* -- 7 -- - 16 2D + - -- ? -- + - -- 17

2E + - - - + ------18 4A + --- - + --- - - 16 4B + - - - + - -- - - 16 5A + ------+ - - 13 5D + 11 24F + - - + + ------16 24H + -- + ------16 241 + 8 24J 6 24K + * + + . _ • ___ 10 253 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

Lowden 31 (OS), New 24l + + + + _ + + _ + + _ + + _ • a. Hampshire, Rockingham 26A + + + + + + + + + + + + Co., Exeter River, Phillips Exeter Acad., 26b + + + + + - + + + + - m - + + + + + Exeter, Lowden 32 (OS), X xT x Tx x x x x xTT x TX x x Ohio, Licking Co., Cranberry Bog, Buckeye 26E + + + - + - + + + + - -- + + + + +

Lake, Lowden 33 (OS), o * j i A 9 i. ID + + + T + I Tx xT + + + Defiance Co., Little T Pond off route 49, 27C + + + - + - + + + • - -- + + + + + Lowden 34 (OS) and Big 27D + + + + + * + + + * + + + + + Pond, Lowden 35 (OS), Ottawa Co., Wlnous Pt., 28A + + + + + - + + + + + + - + + + + + 3*i miles sw of Port £OD + +i T i Tx TTTx X _L TxT x x x Vx x Clinton, Lowden 36 & 37 (OS). 28C + + + + + - + + + + + -- + + + + +

28F + + + + - - + + + + - + - + + + + +

29B + + + - - + + + + - + - - + + - + - 29C + + + + + + + + + - + -- + + - + - 29D - + + + + + + + + + + + - + + - - 3 OB - + + + + + + + + - + - - + + - + -

30C _ + + + + + + _ + • _ + + __ 254 TABLE 7 (continued)

Taxon* Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

24L + - + + ------12 26A ? - - - + ------13 26B + - - - + - - - + - - 17 26C - - - - + - - - + -- 15

26E - - - - + - - - + . -- 15

27b + - - - + - - - + - - 14 27C + - - - + ------14 27D + + - - + - - - + - - 17 28A + - - - + - - - + - - 19 28B + -- - + - - - + - - 17 28C + -- - + - - - + - - 18 28F + - -- + -- - + - 17 29B + - + -- -- + - - - 14 29C + ------+ - - - 15 29D + - + 15 30B + - + - + ------15 30C + 10 255 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

30D + + + + - + - + - + + + + + p + - 3 OF + + + + + + + + + - + + + + + + - 30G + + - + + + + - + - + + + - 30H + + + - + + + - + + + + + + 31A + + + + + + + + - + + + + + + - 3 IE + + + + + + + - - - - + + - 32A + + + + + - + + + - + - + + + - 32C + + + + - - + + + - + - + + + - 32D + + + + + + + + + - + - + + + + - 32F + + + + + + + + + + + - + + - + - 33A + + + + + + + + + + - + + + + -

33b + + + + + + + + + + + - + + + + -

33c + + + + + + + + + + + - + + + + -

33D + + + + + + + + + + + - + + + + - 34B + + + + + + + + + + + + + + + + + - 34C + + + + + + + + + + + + + + + + + - 34D + + + + - ? + + + + + + + + + - 35A + + + + + + + + + - + - + + + + - 35B + + + + - - + + + - + + + + - -

_ 35C + + •• + + + + W + + + 256 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

30D + - + ------14 30F + - + 17 30G + 11 30H + - + 14 31A + - + - - - - ? --- 16 31E + - -- + - + --- 13 32A + 13 32C + - + 13 32D + - + -- -- ? -- 16 32F + m + - - - + - - 17 33A ' + m - + --- - - 17 33B + m - + ----- 17 33C + - - - + ---- - 17 33D + - - + - - -- 17 34B + - - + -- - -- 19 34C + - - + - + - - - 20 34D -- -- + - ---- 14 35A + -- - + - --- - 16 35B + -- m + - + --- 14

_ m wm _ -- • 35C + + 13 257 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9- 10 11a lib 11c 12 13 14 15a 15b

35D + + + + - - + + + - - - - + + - + - 36 B + + + + + + + + + + - - + + - + - 36C + + + + + + + + + + + - - + + + + - 36F - + + - + + + + + + + - - + + + + - 36H - + + + + + + + + + - - - + + - + - 361 + + + + + + + + + + + - - + + + + - 36M + + + + - + + + + + + - - + + - + - 360 + + + + + + + + + + - -- + + + + - 36P + + + + + + + + + + + - - + + + + - 37A + + + + + + + + + + + - - + + + + - 37B + + + + + + + + + + - - + + + + + - 37C + + + + - + + + + + + - ** + + + + - 37D + + + + + + + + + + + - - + + + - 37E + + + + + + + + + + + + - + + + + - 37F + + + + + + + + + + + - - + + + + - 37G + + + + + + + + + + + - - + + + + - 37H + + + + + + + + + + + - - + + - + - 371 + + + + + + + + + + - + - + ♦ + + - 37J + + + + + + + + + + - - + + + + -

37K + + + + + + + + + + + + _ + + + + _ 258 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

35D ---- + - 4 - - - 12

36B 4 - - - + - - - - - 16

36C + 4 -- + - 4 - - - 19

36F + - - - + - 4 - - - 16

36H + + - + + - 4 - - - 17

361 + - - - + - 4 -- - 18

36M + + - - 4 - 4 - - - 17

360 + -- - 4 - 4 - - - 17

36P - - - - 4 - 4 -- - 17

37a + + - - 4 - 4 - -- 19

37b + -- - 4 - 4 - - 18

37C + + - - 4 - 4 - - - 18

37D + - ? - 4 - 4 - - - 18

37E + - - - 4 - 4 - m - 19

37F + - ? - 4 - 4 - 4 - 19

37G + - ? - 4 - 4 - 4 - 19

37H + --- 4 - 4 - - - 17

371 + - - - 4 - - -- 17

37J + -- - 4 - 4 - - - 18 _ _ 37K + 4 4 19 259 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

37L + + + + + + + + + + + + + + + + 37M + + + + + + + + + + + - - + + + + - 37N + + + + + + + + - - + + + + - 370 + + + + + + + + + + + “ - + + + + -

37P + + + + + + + + + + + - - + + + + -

Pontederia cordata var. A + + + + + + + + + - - + + + - lanclfolla B + + + + + - + + + + - - + - + U.S., (A) Florida, Columbia Co., Godfrey C + + + + + + + + + - - + “ -- _ \ 63755 (FSU 34701), (B & C) + + + - + . + + + + -- + + - - - Jackson Co., Godfrey 591^.,1 D (FSU 56361, 56362); (d ) Georgia, Lowndes Co., Godfrey & Houk 62760 (FSU 78653). Pontederia cordata var. ovalls Brazil, plants of the Planaito do Brazil, Serra dos Pirineus, Golas, in Valley of Rio Corumba, Irwin, Maxwell & Waas- hausen 18540 (OS). TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

37L + - - - + - - + - - - 19 37M + + - - + - - + - + - 20

37N + - ? - + -- + - + - 18 370 + + -- + -- + -- - 19 37P +- - - + -- + -- - 18 A+------+ - 14 B+------+ - 13 C + ------11 D + ------+ - + - 13

14 261 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 8 9 10 11a lib 11c 12 13 14 15a 15b

Pontederia saglttata 6 + + + + + + + Mexico, State of Vera Cruz *6 + + + + + + + Vera Cruz, Lowden 6 (OS), Lerdo, Lowden 7 (OS), 6 + + + + + + Laguna Catemaco, Lowden 9 6 + + + + (OS); Guatemala, Dept. Izabal, Quirigua, Lowden 6 + + + + 10 (OS) and Puerto Barrios ’6 + + + + + + Lowden 11 (OS); Honduras, Dept. Cortes, Puerto 6 + + + + + + Cortes, Lowden 13 (OS), 6 + + + + + + + 13.5 miles inland from Puerto Cortes, Lowden 14 6 + + + + + + + (OS), Dept. Atlantlda, 6 + + + + + + Tela, Lowden 15 (OS), Santiago, 11 miles W of 6 + + + + + Tela, Lowden 16 (OS); 6 + + + + + Mexico, 6 kms. de Hulxtla, Lowden 22 (OS); (T & V) 6 + + + + + + seedlings grown from seed 6 + + + + + + ? under culture chamber conditions, from Tela, 6 + + + + + + + Honduras and Vera Cruz, 6 + + + + + + + 7 Mexico, respectively (Lowden 6 & 15). 6 + + + + + + + 6 + + + + + + + 6 + + + + + + +

6A + + + + + + 262 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

6 + - - - + ------13

6 + - - - + 4* - - - - - 13 6 + - - - + ------12

6 + - - - + ------11 6 + m -- + ------11 uft T inXW 6 + --- + -- • --- 12

6 + - - - + - - - - - 12

6 + - - - + ------12

t D Tx 10X v/

6 -- - + + ------11

6 + 11 6 + • • * 11 6 + m + 13

6 + - -- + - - - - - 12 6 + -- - + ------14

6 + --- + ------12

6 + - - - + ------15 6 + - - - + ------13

_ _- . - m 6A + + + 14 263 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

6B + + - + - + + + + + + - - + + - - - 6C + + - - - + + + + - + + - + + - - - 61 + - - + - + + - - - + - - + + - - - 6J + - - - + - - - - + - - + - - - - 6K + + - + - + + + + + + + - + + - - - 6R + + + - - + + + + + + - - + + - - - 6U + + - + - - + + + - + - - + + - - - 6W + - - + - + ------+ + - - - 6x + -- - - + + + - - + + - + + - - - 7 + + " + + + *• “ + + “ 7 + 7 + + + - + + - - - 7 + ------+ - + - - - - 7B + + - + - + + + + + + + - + + - - - 9 + - - - + ------+ - - - - 9 + • + “ • + "■ “ * 9 + 9B + + + + - + + - + + - - - 9E + - - - - + + + - - + - - + + - - -

9F + + + + _ + ___ 264 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

6B + - - + ------13 6C + - - + + - - - - + 14 61 - - - + ------8 6J - - - + ------5 6K + - - + ------14 6R + -- - + ------13 6U - - - + + ------11 6W + - - + ------7

6X + - - + ------10 7 - - - + + ------9 7/ + 7 + 6 7/ + / 7B + _ _ + + _____ 15 9 + 4 9 3 9 2 9B + - ? + ------10 9E + 8

9F 5 265 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a 11b 11c 12 13 14 15a 15b

9K + -- + - + - - + + + - + + - - - 9L + + + - - -

10 + + - - - - + + + - - - + + - - -

10 + + + - - -

inL U +T T

10A + + - + • - + + - + + + + + - -- 10E + + - + - - + + - + - - + + - 7 -

10F + - - - - + + + + + - - + + - - -

10G+ + + - + - - + + + + + - + + -- -

10HLun t+

11B + -- + -- + + + + + + - + + + --

lie + + - - - - + + + + + - - + + - - -

H E + - - + - - - - - + + + + + + - 11F + + - + - - + + - + - - + + -- - 11G + + + + - - + + + + + - - + + -- - 11H + + - + -- + + + + - - + + - - - 13A + + - - - - + + + + + - - + + - - -

13B + + - + - - + + + - - - + + - - - 13C + + - + - - + + - + + - - + + - - -

13D + + + ___ + + + + + _ • + + • __ 266 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

9K + 9 9L + - - + ------6 10 - - - - + ------8 10 + - - - + - - - - - 5 10 3 10A + + 12

10E + - ? - - - + - -- + 11 10F + 9 10G + 11 10H + 4 11B + - - + -- + - - - + 15

11C - - - - + ------10 HE + - ? + - - + - - - + 12 11F + 9 11G + - 12 11H + - 10 13A + - -- + - - - ? - - 11 13B + -- - + - - - ? - - 10 13C + - - - + - - + + -- 13 13D + _ ? __ • 11 o\ > 4 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

13E + +■ + 4- - - + + - + + - - + + - - -

14 + + + - - + + + - - - m + + - - -

14 + + + - - - + + + + - - - + + - - -

14 + + + wm - - + + + + + - - + + -- -

14A + + - - - + + + + + + * + + - - - 14B + + + - 7 - + + - + + + - + + - - - 14C + + + - • + + + + + - - + + - - - 14D + + + - - - + + - + + - - + + - - 14E + + - - - + + + - + - - + + - -

14F + + + -- - + + + + + - - + + -- - 14G + + + - - - + + + - + - - + + -- - 14H + + + + - - + + + + + - - + + -- -

141 + + + - + + -- -

14J + + + + ? - + + + + - - - + + -- - 14K + + + -- - + + - + + - - + + -- -

15 + + -- - + + + + + - - + + - - -

15G + + - - - + + + - + - - + + ---

15H + + + -- - + + + - - - - + + - - - 16A + + + - - - + + + + + -- + + - - -

16B + . + + - - - + + - + + - - + + - + - 268

16C + + + • _ + + + + __ + + ? + — TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

13E + -- - + • ? - - 4 13

14 + - -- 4 - - - - 10

14 - - - - + - ---- 10

14 4 - - - 4 - ? 7 - - 12

14A + - - - 4 -- -- - 12

14B + --- 4 - ? 7 - 4 13

14C + - -- 4 - 7 7 -- 12

14D + - - - 4 - 4 - -- 12

14E + - -- 4 - 7 - - - 10

14F + -- - 4 - 7 - -- 12

14G - - -- 4 - 7 - - - 10

14H + -- - 4 - ?- - - 13 141 + 6

14J + - -- 4 - 7 --- 12

14K + - - - 4 - ? 7 -- 11

15 ---- 4 - -- m - 10 15G + -- -- - 7 - - - 9

15H + - - - 4 - 7 7 - - 10

16A ---- 4 - ? m - - 11

16B + - - m 4 - - - - 4 13 269

16C + • + mm _ 4 as 4 4 15 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

16D + + + + - - + + + + + - - + + - - - 16E + + + - - + + - - + - - + + - + - 16G + + - - - - + + - + + - + + - - - 16H + + - - - + + + + + - - + + ? - - 16K + + - + -- + + + - + - - + + - + - 16L + + - - -- + + -- + + - + + - + - 22A + - - + ------+ + - + + - - - 22B + + - + - - + + + + + + - + + ?- - 22C + - - + - + - - - - + + - + + - •• - 2 2D + - -• - - + ---- + + - + + -- - 22F + + - + - + + + + + + - - + + - -- 22G + + - + -- + + + + + + - + + -- - 22 J + + - + - - + + + + + + - + + 7 - - 22K + + - + -- + + - + + + - + + - - - 22L + + - + -- + + + + + + - + + ? - - 22M + + - + - - + + - ? + + - + + -- -

T + + + - - - + + + + - - - + + ? - - T + + --- - + + - + - - - + + ? - -

V + + + __ _ + + + + • __ + 7 ? 270 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

16D 13 16E + - + + - - + - - + + 15 16G + — — — + — “ — “ — + 12 16H + 12 16K 14 16L 13 2 2 A + ...... 7 22B + 12 2 2 C ...... 7 2 2D ...... 7 2 2 F ...... 11 22G+------12 22J 12 2 2 K + - 12 22L 13 22M + - - - + - ...... 11 T? - 10 T7 ------7 V?------8 271 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a I5t

Pontederia parvlflora 20a + -- - + ---- + - + + + + -- Panama, Province Her­ 20b rera, Los Llanos de Santa Maria, Lowden 20 20C - - - + ------+ - + + + + - * (OS) and Province 20E + - - + - + - - - - + - - + + + - - Panama, between Chepo and Pacora, 7 miles 20F + m - + - + - -- - + - + + + + -- from Chepo, Lowden 21 20G + + + + + + - - (OS). - 2 OH + + - + + + + - - 201 + - - + ------+ - - + + + - 20J + + - + - + - - - - + - + + + + - m 20K + - - + ------+ - + + + + - - 20L + - - - - + - - - - + - + + + + - - 2GM + + - + - + - - - - + - + + + + - - 2 ON + - - + - - - - - + - - + + + - - 200 + - - + - + - - - + - + + + + - - 2 OP+ - -- - + - - - - + - + + + + - 20Q + - - - - + - - - - + - + + + + - - 20R + - - + - + - - - - + - + + + + - - 20S + - + - + - - - - + - - + + + - - 20T + - - + - + - - - - + - + + + + - - 21A+ + 7 + _ + - + -_ + + + -- TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

2 0 A ...... - 7 2 0 B ...... 1 20C ------6 20E ------7 2 0 F ...... 8 2 0 G ...... 6 20H ------6 2 0 1 6 2 0 J ...... 9 20K ------7 2 0 L ...... 7 2 C M ...... 9 20N ------6 200 ------8 20P ------7 20Q...... 7 2 OR...... 8 20S...... 7 20T ------8

21A...... 10 273 TABLE 7 (continued)

Taxon* Location and Spot Number Leaves 1 Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

21B + + - + 7 + + + + + + - - + + + - - 21C + - - + + - - + + - - + + + - - 21E + - - - 7 - - - + + - - + + - - - 21F + + - + - + - - - + - - + + + - -

21G + -- + - - - - - + - + + + + - -

21H + - - a* - - - - + + - - + + + - - 211 + + + - -

21J + + - + - + + - + + - - + + + - -

21L + + + + - + + - + - - + + + - - 21M + - - + - + + - + + - - + - + - -

21N + - + - - + -- + - - + + + -

210 + + + + aw + + - + + - - + + + - -

21P + + + + - + + + + - - + + + - - 21Q + - + + - + + - - + - - + + + - - 21R + ------+ + - - + + + - - 21S + - - + - - - - - + - + + + + - - 21T + - - - + - - -- + - + + + + - - Pontederia rotundifolla 12A + + + + + + + + + + - - + + - - - Honduras* Dept. 1 9ft X*XX -i X TXXXXj _ ± . A x XTXX Comayagua, Lago Yojoa,

Pito Solo* Lowden 12 12C + + - + + + + + + + - - + + - - - 274 (OS); Nicaragua, Dept. 12D + + _ + + + + + + + _ _ + + _ _ - TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 '21 22 23 24 25 26 Spots

21B + - - - + ------14 21C 8 21E 5 21F 8 21G 7 21H 6 211 3 21J 10 21L - - - - - m - - + - - 11 21M 8 21N 7 210 11 21P - - - - + ------12 21Q + - - - + - - - - + - 12 21R 6 21S 7 21T 7 12A + -- - + - - - + ? - 15 12B + - - - + ------14

12C + - - - + ------13 275

12D + __ + __ _ __ 13 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 A 5 6 7 8 9 10 11a 11b lie 12 13 14 15a 15b

_ Granada, near Granada, 12E + + + + + + + “ ■*

Lowden 17 & 18 (OS); t x x itr T T T TiTTTT i _i_ i _i_ x.TTx T ' Costa Rica, Province Guanacaste, 1 km. from 12G + + + + + + + + + + - - m + + --

Arenal, 2 kms. from 1 * * i A 1 1Lcl 9 T + + + + + + T T- + + T Tronadora, Lowden 19 (OS); El Salvador, Dept. 12 J + + + + + + + + + + - - - + + - - La Llbertad, Laguna 12K + + + + + + + + + + + + ' Zapotltan, Lowden 23 (OS). 17A + + + - - - + + + + ? - M + + - -

17B + + + m - - + + + + + - - + + - -

17C + + + ? ? ? + + + + + - W + + - -

17D + + + - 7 - + + + + - a* - + + - - 17E + + + - ? - + + + + ?- - + + - - 17F + + + + + - + + + + + - - + + - - 17G + + + + + - + + + + - - + + - -

17H + + + - - + + + + + - - + + - 171 + + + - + - + + + + - - + + -- 17J + + + ? + - + + + + + - - + + - -

17K + + + + 7 - + + + + + - m + + - - 17L + + + + + - + + + + + - - + + - - 17M + + + + + + + + + + + - - + + - - + + + - + + - - 17N + + + + + - + + - 276 170 + + + + _ + + + + + _ + + _ TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

12E + - - - + - - - + - - 10

12F + - - - + - “ - ? + - 14

12G + - -- + -- -?- - 14 121 + - - - + - - - - + - 15

12 J + - -- + - - - - + - 15

12K + - - - + - - - - + - 15

17a + - - - + - - - - + + 14

17B + - -- + - - - + 7 + 15

17C + - - - + - - - + - - 14

17D ? - - - + - -- + - - 12

17E + - - - + - - - - 7 7 12

17F + - - - + - - - “ - ? 15 17G + - - - + - - - ? - - 15 17H + - - - + - - - - - + 15

171 + -- - + - - - ? - - 13 17J + - - - + - - - ? - - 14 17K + - - - + - - - 7 - - 14

17L + -- - + -- - 7 - - 15 17M + - - - + - - - ? 7 - 16

17N + - - - + - - - - - 7 15 277

170 + ■ + _ —__ 14 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

17P + + + 4 4 - 4 4 4 4 4 - - 4 4 ? --

17Q + + + ? -- 4 4 4 4 ? - - 4 4 4 - -

17R + + + 4 - - 4 4 4 4 4 - - 4 4 4 - -

17S + + + - 4 m 4 4 4 4 4 -- 4 4 4 --

17T + + + 4 4 - 4 4 4 4 4 mt - 4 4 ?--

ISA + + + 4 4 4 4 4 4 4 4 - - 4 4 4 - -

18B + + + 4 - - 4 4 4 4 4 - - 4 4 4 - -

18C + + + 4 4 - 4 4 4 4 4 - - 4 4 4 -

18D + + + 4 4 4 4 4 4 4 - - 4 4 4 --

18E + + 4 4 4 - 4 4 4 4 4 - - 4 4 4 - -

18F + + 4 4 4 - 4 4 4 4 4 - - 4 4 4 --

18G + + 4 4 -- 4 4 4 4 4 - - 4 4 4 - -

18H + + 4 4 4 - 4 4 4 4 4 -- 4 4 4 - -

181 + + 4 4 - - 4 4 4 4 4 -- 4 4 4 - -

18J + + 4 4 4 4 4 4 4 4 4 - - 4 4 4 - -

18K 4 + 4 4 - 4 4 4 4 4 4 - - 4 4 4 - - 00 r~i + + 4 4 4 - 4 4 4 4 4 - - 4 4 4 --

18M + 4 4 4 4 4 4 4 4 4 - - - 4 4 4 - -

18N + 4 4 4 ? 4 4 4 4 4 4 - - 4 4 4 - -

18P + + 4 4 -- 4 4 4 4 4 - - 4 4 4 - - 278

18Q + + 4 4 4 4 4 4 4 4 4 - - 4 4 4 -- TABLE 7 (continued)

Taxon* Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spot 8

17P + -- - + * -- + -- 15

17Q + - - - + - -- + - - 13

17R + - - - + ------14

17S + -- - + - - - + - - 15

17T + - - - + --- - - + 15

18A + + -- + -- - ? + + 19

18B + -- - + -- - -- 7 14

18C + -- - + -- - ?- 15

18D + -- - + - -- ? ? 7 15

18E + - - - + - - - 7 - ? 15

18F + - - - + - - - 7 ? 7 15

18G + -- - + -- - ? - 14

18H + - - - + - - - - - ? 15

181 + - -- + - -- + + 16

18 J + - - - + - - - - + 7 17

18K + -- - + - - -- + 7 16

18L + - -- + m -- - - 7 15 18M + - - - + - - - 7 - - 15 18N + - -- + ------15 18P - - _ + --- 14 + - -- 279

18Q + --- + --- ? - - 16 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

18R + 4 4 4 - - 4 4 4 4 4 - - 4 4 4 - -

18S + 4 - 4 - - 4 4 4 4 4 -- 4 4 4 - -

19A + 4 -- - m 4 4 4 - 4 a* - 4 4 - - -

19B + 4 4 4 4 - 4 4 4 4 4 -- 4 4 - - m

19C + 4 4 4 4 4 4 4 4 4 - - 4 4 - - -

19D + - - 4 ------4 - - 4 4 - - -

19E + 4 - 4 4 - 4 4 4 4 4 - - 4 4 4 m - 19F + ------4 - - 4 4 - - -

19G + 4 - - 4 - 4 4 4 4 4 - - 4 4 - --

19H 4 4 - 4 4 - 4 4 4 - 4 - - 4 4 - m -

191 4 4 - 4 4 - 4 4 4 4 4 - - 4 4 - m -

19 J 4 4 - - 4 - 4 4 4 4 4 - - 4 4 - - -

19K 4 4 - 4 4 - 4 4 4 4 4 -- 4 4 - - -

19L 4 4 4 4 4 - 4 4 4 4 4- - - 4 4 - - -

19M 4 4 4 4 4 4 4 4 4 4 -- - 4 4 - - -

23A 4 4 - -- - 4 4 4 -- -- 4 4 - - -

23B 4 4 m - - - 4 4 4 4 --- 4 4 - - -

2 3D 4 4 4 4 4 4 4 4 4 4 4 -- 4 4 ---

23E 4 4 4 - 4 - 4 4 4 4 - - - 4 4 - --

23F 4 4 4 4 4 - 4 4 4 4 - -- 4 4 - - - 280

23G 4 4 __- 4 4 4 - 4 _— 4 4 _ - - TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

18R + --- + ------14

18S + -- - + -- - ? - - 13

19A + - - - + - - - - + ? 11

19B + - + + - - - - - + 16

19C + - - + + ------15

19D - - - + ------6

19E - • - + + ------14

19F + + - - + ------7

19G + - - + ------12

19H + - + ------12

191 - - + + ------13

19J ?- - - + ------11

19K + •• - - + ------13

19L + - - - + ------14

19M + - - - + - - - + - - 15

23A + -- - + ------9

23B + m -- + ------10

23d + - - - + ------15

23E + m - - + ------12

23F + - - - + ------13 261 23G + _ + _ _ — _ 10 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined 1 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

23H + + - + - - + + + + + -- + + - --

231 + - + -- + + + + + + + - --

23J + + - + ? ? + + + + + m m + + - --

23K + + + + - + + + + + -- + + - --

23L + + + + - - + + + + m - + + - - -

mm 23M + + - + - - + + + + - - + + - - -

23N + + - + - - + + + + + m - + + - - -

230 + + + - -- + + + + + -- + + - -- 23P + + + - - - + + + - - - - + + - - - 23Q + + + + + - + + + + - - - + + - - - 23R + + + - - - + + + + - - + + - --

23S + + + + - - + + + - m m + + - --

23T + + + + - - + + + + + -- + + - -- 23U + + - + - - + + + + + -- + + - --

23V + + + + - + + + + - - - + + - -- 23W + + + + + - + + + + + -- + + - --

23X + + + + - + + + + + + - - + + - - -

23Y + + + - -- + + + + -- - + + - - -

23Z + + + ___ + + + + + _ + + _ _— 282 TABLE 7 (continued)

Taxon* Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

23H + - - - + ------12

231 + - - - + ------12 23J - - - + ------12

23K + - - - + ------13

23L + - - - + ------12

23M + - - - + ------11

23N + - - - + - - - - - ? 12

230 + - - - + ------12

23P + - - - + - - - + - - 11

23Q + - - - + - - - + - - 14

23R + - - - + - - - + - - 13

23S + - - - + ------12

23T - - - + ------13

23U + - - - + - - - + - 13

23V + - - - + - - - + - - 14

23W + --- + ------14

23X + - - - + ------14

23Y + - - - + - - - + - - 12

23Z + ___ + ___ + _- 13 283 TABLE 7 (continued)

Taxon, Location and Spot Number Leaves Examined l 2 3 4 5 6 7 8 9 10 11a lib 11c 12 13 14 15a 15b

_ Heteranthera limosa A + + + + + + + + +

(A & B) Panama, * i ■>_ i D + + T + + + + * + + Province Herrera, Los Llanos de Santa Maria, + + - + -» - + * + + -- + + -- Lowden 38 (OS). Eichhornia azurea Mexico, State of Vera Cruz, Carretera Naclonal 180, El Puente de Teculapilla, 5 kms. ante a llegar a nw. Lerdo, Lowden 41 (OS).

Eichhornia crasslpes A + + + + — + + + + + _- + + --_ & (A c) El Salvador, B x * > _ X i o T T + ■ T T T Lago Illopango, oeste lado de Apulo, Lowden c + + + + - - ? + + + - - + + -- 39(0S); (B) Honduras, Lago Yojoa, Pito Solo, Lowden 40 (OS).

+ sign represents the presence* ? the doubtful presence, and - sign the undetected presence of phenolic compounds based on position and color of spots on chromatograms. 284 TABLE 7 (continued)

Taxon, Location and Spot Number Number of Leaves Examined 16 17 18 19 20 21 22 23 24 25 26 Spots

A+- + ------+ - 12

- 10

+ - 11

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