Hodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON RADCLIFFE BARNES PIKE ^ Associate Editors VOLUME 67 1965 Corda Enne c Lad. m Arr P Lie TL vel . GB. The New Lugland Botanical Club, Jue. Botanical Museum, Oxford St., Cambridge 38, Mass. FARLOW REFERENCE LIBRARY 1965 Hovdova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL 3 STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER Associate Editors ROLLA MILTON TRYON f RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. J Vol. 67 January-March, 1965 No. 769 CONTENTS: Monograph of the Genus Elodea: Part 4 and Summary IEE OE SE OB EOE PRIE e RE, | A New Species of Lagenophora (Compositae) from Guatemala John H. Beaman and D. C. D. DeJong EN 86 Genotypic Variation in the Phacelia hirsuta Complex Georgo W: Gilet Ee ee . 42 Eulophia ecristata and Epidendrum conopseum in Mississippi BEN NOI iion orc A 48 Determination of the Developmental Pattern of Angiospermous Leaves Edward L. Davis nesses 49 Errata For Rhodora Volume 66 ......... eee metes e 644 (Continued on Inside Cover) The New Zoning Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge 38, Mass. CONTENTS: — continued Natural and Artificial Hybrids of Cacalia atriplicifolia and C. muhlenbergii James R. Coleman ............ emm 55 Notes on Varieties of Bidens connata and a Hybrid with B. cernua Earl Edward Sherf e 59 A Revision of the North American Species of Helianthemum (Cistaceae) H. S. Daoud and Robert L. Wilbur ............—.— 63 An Earlier Name for Ludwigia natans (Onagraceae) Peter H. Raven eee nnne nene nente nnne ennt nennen nennen 83 Mountain Flowers of New England (Review) A. R. Hodgdom ........ eene nennen entente nnns 84 A Second Character Distinguishing Heterotheca s. str. from Chrysopsis (Compositae: Astereae) Vernon L. Harms ........ 86 Tulostoma brumale Pers. A New Record for New England Wesley N. Tiffney and Wesley N. Tiffney, Jr. ees 89 New Records of Arctic Species in Southeastern New Brunswick Patricia R. Roberts .........., 25e 92 Common Trees of Puerto Rico and the Virgin Islands (Review) R. B. Pike .ssssssssesrsrrereseerererersssersessererreenereeeeeeerereeenee 93 Dissecting Equipment and Materials for the Study of Minute Plant Structures Richard W. Pohl ...............4.... 95 RHODORA.—A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price, $6.00 per year, net, postpaid, in funds payable at par in United States currency in Boston; single copies (if available) $1.80. Back vol. umes 1-58, with a few incomplete, can be supplied at $5.00 per volume, Volume 59— available at $6.00. Somewhat reduced rates for complete sets can be obtained upon application. Scientific papers and notes, relating directly or indirectly to the plants of North America, will be considered by the editorial com- mittee for publication. Articles concerned with systematic botany and cytotaxonomy in their broader implications are equally accept- able. All manuscripts should be double-spaced throughout, Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Rbodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 67 January-March, 1965 No. 769 MONOGRAPH OF THE GENUS ELODEA: PART 4 AND SUMMARY. HAROLD ST. JOHN! I. MONOGRAPH OF THE GENUS ELODEA, PART 4: THE SPECIES OF EASTERN AND CENTRAL NORTH AMERICA. INTRODUCTION A preliminary discussion was given in part one of this monograph and it does not need repetition here. Each of the four parts of this study covers the species of a large natural area. Since the species flower but briefly, and have deli- cate, largely evanescent flowers, they are poorly represented as dried herbarium specimens. Hence, for practical use, an artificial key, based mostly on foliage characters, is here included. With it, plants having well developed median and upper leaves can be identified with reasonable surety. Il- lustrations of typical leaves are here provided. The other parts of this monograph have been printed elsewhere, so written that they can be assembled into a single whole. Part 1, The Species Found in the Great Plains, the Rocky Mountains, and the Pacific States and Provinces of North America, Res. Stud., Washington State Univ. 30 (1962) 19-44, fig. 1-5; Part 2, The species found in the Andes and western South America, Caldasia 9 (1964) 95-118, figs. 1-8, is in press in Mutisia; Part 3, The species found in northern and eastern South America, Darwiniana 12 (1963) 639-652, fig. 1-3, tab. 1. Also two species formerly 'Botanist, B. P. Bishop Museum, Honolulu, Hawaii, 96819, U.S.A. 1 2 Rhodora [Vol. 67 placed in Elodea were treated in the author's monograph of the genus Egeria Planchon, Darwiniana 12 (1961) 293-307, figs. 1-2; 12 (1962) 525. ARTIFICIAL KEY TO SPECIES OF NORTH AMERICA, FROM THE ATLANTIC SLOPE TO THE GREAT PLAINS. 1. Middle and upper leaves in whorls of 4-b; staminate spathe 2-4- flowered; filaments glandular above, at least 3 times as long as the anthers; pistillate petals 8 mm. wide. This species of Egeria is here included because of its superficial similarity to Elodea, and because it was long considered a member of that genus. ........ Egeria densa. NEELAM late petals 0.5-1.5 mm. wide. seen Genus Elodea. 2. Middle and upper leaves opposite; pistillate spathe 3-7 cm. long. vececcccoscoccecanevescsonecsssusssssssssassusoosscssscessesenes 21. E. longivaginata, p. 34. 2. Middle and upper leaves in whorls of 3 (-4); pistillate spathes shorter. 3. Middle and upper leaves less than 1 mm. wide; stamens equal and equally attached on the hypanthium. e TTT 20. E. linearis, p. 29. 3. Middle and upper leaves all, or some of them, more than 1 mm. wide. 4. Middle and upper leaves 0.3-1.5 mm. wide, flaccid, diver- gent, not imbricate at tip of stem; pistillate sepals 1.1 mm. long; staminate flowers sessile, at anthesis detached and floating. ENEE 18. E. Nuttallii, p. 6. 4. Middle and upper leaves 1-5 mm. wide, firm or subflaccid ; all flowers long-stalked by the hypanthium base, not liberated. 5. Upper leaves usually 2-5 mm. broad, firm, dark green, closely imbricate; flowers dioecious; pistillate sepals 2-2.2 mm. long. .......2. errem 19. E. canadensis, p. 12. Upper leaves 1-2 mm. broad, subflaccid, pale green, loose- ly imbricate; flowers perfect; sepals 2.7-3 mm. long. AN 17. E. Sehweinitziüi, p. 2. c DESCRIPTION OF SPECIES 17. Elodea Schweinitzii (Planch.) Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857 (1857) 46; also in Pringsheim's Jahrb. Wis- sensch. Bot. 1 (1858) 468; Apalanthe Schweinitzii Planch., Ann. Mag. Nat. Hist. II, 1 (1848) 87; Ann. Sci, Nat. Bot. III, 11 (1849) 76. Fig. 3 a-c, 4. 1965] Elodea — St. John 3 Description of all specimens examined: Submerged aquatic; stems slender, dichotomously branched; lower leaves opposite, ovate- lanceolate to lanceolate, smaller than the upper; medium and upper leaves 8-14 mm. long, 1-2 mm. broad, in whorls of 3, linear to lance- linear, finely serrulate, subflaccid and pale green; spathe 10-16 mm. long, narrowly urceolate, sharply bidentate at apex; flower perfect, exserted from the spathe by the thread-like elongating hypanthium 2-6 cm. long; sepals 2.7-3 mm. long, 1-1.2 mm. broad, elliptie, dark striate; petals 2.7 mm. long, 0.9 mm. broad, white, delicate, spatu- late; stamens three, filaments 0.3 mm. long, stout; anthers 0.8-1.8 mm. long, elliptic; ovary narrowly cylindrie; style slender, equaling the hypanthium; stigmas 3-4, linear-oblanceolate, entire (or bifid), usually one of them shorter than the others which are twice the length of the sepals; capsule and seeds unknown. Holotype: “in Americae septentrionalis provinciis confederatis [United States], loco proprio non indieato, Schweinitz in Herb. Hook." The holotypic specimen bears only the following data: North America, Schwein. (K)! Type examined! An apparent iso- type lacks the locality and bears only the abbreviation Schwein. of the collector's name (K)! UNITED STATES. NEw JERSEY: N. Caesar. 1834, Torrey (P); without locality, Gray (M); without locality or collector (K). PENN- SYLVANIA: Bet. [hlehem], 1832, Schweinitz (K), sterile, but likely an isotype; Nazareth, [Schweinitz], (PH), from Schweinitz’s own herbarium, doubtless an isotype; ubi, 1829, Schweinitz (BR), likely an isotype; Bethlehem, unio itiner, 1832, Moser (B, CAS, G, NY, P); in der Lecher bei Bethlehem, Juli 1832, Prine Wiedensis (BR); Beth- lehem, Wolle (Mo, PH). American Borealis: without data, (PH). Elodea Schweinitzii is a member of the subgenus Apa- lanthe, a small group containing only two other species: E. Brandegeae St. John of California, and E. granatensis Humb. & Bonpl. of South America. Their distinctive fea- ture is the possession of perfect flowers. Elodea Schweinitzii is of very local occurrence. The holo- type was collected by L. B. Schweinitz, but its label gave no locality data. There are, however, several other speci- mens collected by him, and doubtless some or all of them are isotypes. The one retained in his own herbarium was labeled Nazareth, while others are marked Bethlehem. Bethlehem, Pennsylvania was a small city and Nazareth an adjacent village. Schweinitz spent his life there, and made most of his collections in that immediate vicinity. It seems 4 Rhodora [Voi. 67 that this Elodea with perfect flowers was then locally com- mon there. It will be noted that the last collection of it was made in 1832. In 1940 the writer drove to eastern Penn- sylvania and explored the region of Bethlehem and Naza- reth. Unfortunately by this time Bethlehem had long been a center of steel mills and the streams of the vicinity were opaque and foul from the waste discharged by the mills. No aquatic flowering plants of any kind were to be seen in the despoiled streams. The three collections recorded from New Jersey give no more precise locality. John Torrey certainly collected in New Jersey, but Asa Gray did so little if at all. No sub- sequent collections are known to confirm the New Jersey locality. The writer doubts it and thinks that the specimens probably came from Bethlehem, Pennsylvania and were labeled in the casual, inexact manner so often characteris- tic of botanists of that century. The floras of eastern Pennsylvania and of New Jersey are very well known. For some 150 years these regions have been intensively botanized by numerous local, resident botanists. No one of them in the last 130 years has re- discovered Elodea Schweinitzii. It seems that this species is extinct in Pennsylvania, and that its report from New Jersey was erroneous. In the other species of Elodea the morphology of the flowers is invariable. Not so in E. Schweinitzii, as the flowers show considerable variability. Planchon described in his Latin diagnosis that the flowers had: “three stamens ; filaments slender, longer than the anthers; anthers obovate, compressed, . . . stigmas three, bipartite (?), the lobes re- curved twice longer than the outer parianth lobes." The usual flowers have three stamens, three staminodia, and three or four undivided stigmas. Caspary described one specimen of Moser's 1832 collection as having 7 stamens and no stigmas. The writer has not found this specimen and cannot confirm the statement. Although Caspary main- tained Elodea Schweinitzii (Planch.) Casp. he did so largely in confidence on the work of Planchon. Caspary studied the type specimen in Herb. Hooker, but found the only two 1965] Elodea — St. John 5 flowers partly destroyed. In one of these he could observe 2 petals and 3 stigmatic lobes, but he could not confirm the flowers as bisexual. He did see elsewhere several specimens of the bisexual plants from Bethlehem, collected by Schweinitz and by Moser, but he classified them as E. cana- densis, a species as he interpreted it having either herma- phroditic or dioecious flowers. It is obvious now that these specimens with hermaphroditic flowers collected at Beth- lehem were the genuine E. Schweinitzii. One of the flowering specimens collected by Wolle (MO) has a loose staminate flower tangled in the leaves. On dissection this proved to have 9 stamens and to be a stami- nate flower of Elodea canadensis. The other specimen on this Wolle collection has an abnormal flower with a large, coriaceous, swollen hypanthium, probably due to insect in- jury. The "New Jersey" specimen, Gray (M) has 3 good flowers, one of which has one stigma deeply split, apparent- ly accidentally so, down the median submembranous tissue. In the Delessert Herbarium (G) there are two sheets of the 1832 Moser collection. One has a flower with 3 good sta- mens, but the 3 stigmas are bifid 3/4 way down. Their lobes are narrow, but flat, and more delicate in texture than those of E. canadensis. The duplicate of this (P) bears branches with one old and two good flowers. Of these one has 4 entire stigmas, while the other has 4 stigmas of which only one is bifid for 0.7 mm. On the Torrey speci- men “N. Caesar." (P), of the three flowers one has 4 entire stigmas, and the other good flower has 4 stigmas, three entire and one shortly bifid. The 1829 Schweinitz (BR) specimen has the stigmas bifid 3/4 way. In sum it can be seen that this species is unusual in its variability in floral morphology. A possible explanation would be that the species was of recent origin by hybridiza- tion and that it was not fully homozygous. Opposed to that hypothesis is the fact that in the general area there are only two other species, E. canadensis, and E. Nuttallii. Both of these species have dioecious flowers and in their pistillate flowers have the male element represented by sterile stami- nodia. Obviously they were derived from ancestors with 6 Rhodora [Vol. 67 perfect flowers. The species with perfect flowers represent the primitive state in the genus. Consequently it is not reasonable to look upon E. Schweinitzii, a species with per- fect flowers, as a putative hybrid of two species with dioe- cious flowers. 18. (7. of part 1.) Elodea Nuttallii (Planch.) St. John, Rhodora 22 (1920) 27-28, as to basionym, non sensu St. John (1920); E. oc- cidentalis (Pursh) St. John, Rhodora 22 (1920) 27-29, sensu St. John (1920), not as to basionym; E. minor Farw., Rept. Mich. Acad. Sci. 17 (1916) 181; E. canadensis Rich. in Michx. var. angustifolia (Britton ex Rybd.) Farw., Am. Midl Nat. 10 (1927) 203; Ana- charis Nuttallii Planch., Ann. & Mag. Nat. Hist. II, 1 (1848) 86, and Ann. Sci. Nat. Bot. III, 11 (1849) 74, A. occidentalis (Pursh) Vic- torin, Contrib. Lab. Bot., Univ. Montréal 18 (1931) 40; Serpicula occidentalis Pursh, Fl. Am. Sept. 1 (1814) 33, a superfluous epithet since Pursh should have adopted the epithet canadensis; S. verticil- lata L. f. B var. angustifolia Muhl., Cat. Pl. Amer. Sept. (1813) 84, nomen nudum; Philotria minor Small, Fl. S. E. U. S. (1903) 47; P. Nuttallii (Planch.) Rybd., Bull. Torrey Bot. Club 35 (1908) 461- 462, 465, a provisional name; P. Nuttallii (Planch.) Rybd. ex Brit- ton & Brown, Ill. Fl. N. E. U. S. ed. 2, 1 (1913) 105; P. occidentalis House, Bull. N. Y. State Mus. 243-944 (1923) 55. Fig. 2 e-i, 4. Description of all specimens examined: Submerged aquatic; stems slender, dichotomously branched, often freely so; lower leaves op- posite, lance-ovate, smaller than those above; median and upper leaves 6-13 mm. long, 0.3-1.5 mm. wide, whorled in threes or occasionally in fours, linear or narrowly lance-linear, pale green and flaccid, finely serrulate; staminate spathes borne at the median axils, sessile, ovoid, apiculate, 2-parted to well below the middle, the two acuminate teeth often twisted, forming the apiculate tip, the body 2 mm. long; staminate flower single in each spathe, sessile, at maturity breaking loose, floating to the surface and there opening; sepals 1.9-2.1 mm. long, 1.5-1.7 mm. wide, ovate, sometimes reddish-tinged; petals usually wanting, or when occasionally present 0.5 mm. long, ovate- lanceolate; stamens 9, and 1.2 mm. long, the 3 central ones slightly elevated on a common stalk, the 6 outer ones at a lower level and with separate filaments, the anthers 1 mm. long, 0.6 mm. wide, broad- ly ellipsoid; pistillate plants bearing in the upper axils spathes 9-25, but usually 10-15 mm. long, narrowly cylindric, somewhat ovoid at base, slightly enlarged at the bifid tip, the two acuminate teeth slightly spreading; pistillate flower stalked by a slender, thread- like, elongating hypanthium as much as 9 cm. in length; sepals 1.1 mm. long, 0.5 mm. wide, obovate, dark-striate; petals 1.8 mm. long, 1 mm. broad, white, delicate, broadly obovate; staminodia three, 0.5 mm. long, acicular; capsule 5-7 mm. long, 1.5-2 mm. in diameter, ses- 1965] Elodea — St. John fi sile, narrowly ovoid to fusiform; seeds 3.5-4.5 mm. long, 0.5-0.8 mm. in diameter, cylindrie, short beaked, pilose; style slender, equaling the hypanthium; stigmas 3, slender, bifid, somewhat exceeding the sepals. Holotype: “in America septentrionali." The type specimen is Herb. Thomas Nuttall, Udora canadensis, Phil.[adelphia], (BM). Specimen examined! It has the narrow leaves and the small pistil- late flower of the species formerly called E. occidentalis (Pursh) St. John. Range: In fresh (or rarely brackish) waters, Quebec to North Carolina, westward to Minnesota, Kansas, and Colorado, then in northern Idaho. NORTH AMERICA: 1825, Barrett (w); Brendel (B); en 1868, Durand (P); Engelmann (G, K, P, W), with staminate flowers, la- belled Udora verticillata minor, and probably isotypes of it; ditto, but with pistillate flowers, Engelmann (BR, K, S) ; Leconte (P) ; Wolz (zT). CANADA. QUEBEC: cult, Jard. Bot. Montréal, ex l'ile Ste.-Thér- ése, Richelieu, Rolland-Germain 7498 (FSU, S); Saint-Jerome, Lau- rentides, Victorin 20322 (GH). UNITED STATES. J. Blake (BM). MAINE: Cobossee Contee L., Battey (NEBC); Unity Pond, Burnham, Bean (NEBC); Messalonskee R. Waterville, Chamberlain 774, Fernald 2750 (BRU, GH, NEBC); Cathance R., Bowdoinham, Fassett 14 (F), 94, (NY), Fernald & Long 12748 (NEBC, PH); S. Poland, Furbish (NEBC); E. Livermore, Fur- bish (NEBC); Androscoggin L., N. Leeds, Furbish (NEBC); Haley Pond, Rangeley, Furbish (NEBC); L. Auburn, Merrill 508 (NEBC), 963 (US), s.n. (NY). NEW HAMPSHIRE: Connecticut R., West More- land, Cheshire Co., Krochmal 1227 (NHA); Beard's Brook, Durham, Strafford Co., Hodgdon 5889 (NEBC, NHA); tidewater, Durham, Hodg- don 5893 (NHA); Beard's Brook, Durham, Hodgdon & Hooghkirk 5890 (NHA); Ashuelot R., Hinsdale, Cheshire Co., Hodgdon & F. Steele 10048 (NHA); Province L., Wakefield, Carroll Co., Pease 29912 (NEBC); Pontook Res., Dummer, Pease 36781 (NEBC, NHA); Cres- cent L., Wolfboro, Carroll Co., Seymour 4824 (NEBC); S. Charles- town, Sullivan Co., Seymour 21058 (NEBC); Conway L., Conway, Carroll Co., F. L. Steele 3374 (NEBC). VERMONT: without locality, Chapman (UC); Windsor, Eggleston 2085 (GH). MASSACHUSETTS: Somerville, Bailey (NEBC) ; Congamond L., Congamond, Blewett & Harger 6631 (PH); Alewife Brook, Medford, Boott (GH); Harwich, F. S. Collins 2445 (NEBC); Back Bay Fens, Boston, Collins 3639 (NEBC), s.n. (NEBC); Fresh Pond, Cambridge, Deane (NEBC), Deane and E. & C. E. Faxon (GH, NEBC); Great S. Pond, Plymouth, Fernald, Hunnewell & Long (NY); Agawam R., Wareham, Fernald & Svenson 758 (NY); Muddy R., Brookline, Forbes (NEBC) ; Great S. Pond, Plymouth, Gliick (NEBC); Heard’s Pond, Wayland, Kennedy (GH); Beaver Brook, Middlesex Co., Kidder (Ny); Spot 8 Rhodora [Vol. 67 Pond, Stoneham, Kingman (GH); Fresh Pond, Cambridge, Morong (NY), Pease 2063 (NEBC) ; L. Cochichewick, N. Andover, Pease 2638 (NEBC); W. Cambridge, St. John 679 (NEBC, PH); Beaver Brook, Waverley, Belmont, Seymour 3620 (s); Agawam R., Wareham, L. B. Smith & H. K. Svenson 825 (NY); Fresh Pond, Cambridge, Young (NEBC). RHODE ISLAND: Olney Pond, Lincoln, J. F. Collins (NY); Mill Pond, Lonsdale, Greene (BRU). CONNECTICUT: Waterbury, Blewitt 573 (NEBC); Farmington R., New Hartford, Blewitt 586 (NEBC); Connecticut R., Hartford, Blewitt 609 (NEBC); Pistapaug Pond, Durham, Blewitt 1680 (NEBC) ; Stony Brook, E. Haven, Blewitt 1980 (NEBC); Congamond L., Suffield, Castle & Dann 114 (CU); Kensington, Cowles (GH): Beaver Cr., Milford, E. H. Eames (NEBC) ; Bantam L., Litchfield, Eaton (GH); without locality, Gray 4606 (BM), s.m. (CP, 0); Housatonic R., Orford, Harger (Uc) ; Simsbury, Holcomb (GH); without locality, M. Mitchell (G); Norwich, Setchell (UC); Middletown, Ware 3472 (NEBC); Boardman’s Pond, E. Hartford, Weatherby 367 (NCSC); Pistapaug Pond, Wallingford, Weatherby 3373 (NEBC); E. Haven, Weatherby 3591 (NCSC); Putnam, Weatherby 3777 (NEBC); Old Lyme, Woodward (GH); Hartford, Wright (oni: Wethersfield, Wright (GH); without locality, Wright, in part (MO); without locality, Herb. D. Don, Herb. Martii (BR). NEW YORK: Jamaica Reservoir, Bicknell 176 (NY) ; Hudson R., Hastings, Bicknell (NY); S. Pond, Bedford, Cushman & Wood (NY); Fall Cr., Ithaca, A. J. Eames & K. M. Wiegand 13351 (CU); Woodside, Ferguson 4147 (NY); Cold Spring Hbr., Ferguson 4414 (NY); Oyster Bay, Ferguson 4832 (NY); Millneck, Ferguson 7843 (NY); Great S. Pond, Babylon, Ferguson 8027 (NY); Baldwin, Ferguson 8154 (NY); Mill- neck, Ferguson (NY); Salmon R., Selkirk, Fernald, Wiegand & Eames 14123, 14124 (CU, GH); Wading R., Glück (Glück Herb.) ; Watertown, Gray, in part (NY); Troy, J. Hall (F); Clove L., Staten L, Hollick (NY), Heuser (B); Long L, Hennecart (P); Black R. Bay, Keyes (uc); Quogue, Knight (NY); Mohegan L., Leggett (F); Peek- skill, Leggett (NY); Ithaca, Muenscher 13352 (cv); Hudson R., Catskill, Muenscher & Curtis 5570 (cu); Great Pond, Riverhead, St. John 2548a (CU, GH); Pocantico Hills, W. Chester Co., N. Taylor 847 (NY); Hotaling I., Hudson R., New Baltimore, Taylor 1372 (NY); 3rd. Ave, (New York), Torrey & Gilman (NY); Ontario See, Charlotte, Weinland (B); N. Fairhaven, Wiegand 13358, in part (cU). New JERSEY: Oldmans Cr. Woodstown, Adams 263, 264 (GH, PH); Newton Cr., Collingswood, Adams 294 (PH); Camden, Boice (PH); Bayhead, Cannon (CAS); without locality, Enneandr (M0); Fishhouse Sta., (Delaware R.), Glück (Glück Herb.) ; Brindle- town, Grove 490 (PH); Silver L., Belmar, Harshberger (PH); Rah- way, Heuser (B, M); Paulsboro, Jahn 167 (PH); Swedesboro, Lippincott (PH); Crosswicks Cr., B. Long 6036 (PH); Delaware R., Delair, Long 6293 (PH); Budd's L. Morris Co., Mackenzie 1505 1965] Elodea — St. John 9 (NY); Barnegat Bay, Bay Head, Mackenzie 4731, 5095 (NY); Dela- ware R., Bordentown, Mackenzie 6833 (NY); Delaware R., Delair, Mackenzie 7352 (NY); Hibernia, Mackenzie (NY); Swartswood L., Sussex Co. Mackenzie (NY); Camden, Martindale (L, PH); without locality, Morong, in part (MO); Delaware R., Delair, Pennell 6498 (NY); without locality, Pursh, in part (K); Sussex Co., Rusby (BRU); Princeton, Schott (F, US); without locality, Torrey (K); Budd’s L., Vail (NY). PENNSYLVANIA: Lagrange, Adams 76 (PH); Neshaminy Falls, Adams 99 (PH); Darby Cr., Delaware Co., Barker (PH); Stroudsburgh, Bicknell (NY); York Furnace, Brown (PH); Falls of Schuylkill, Carson (PH); Fulton Twp., Carter (PH); York Furnace, Crawford (PH); Perkiomen Cr., Schwenkville, Driesbach 1598 (PH); Philadelphia, Engelmann (B); Darby Cr., Adele, Fogg (PH); Pt. Pleasant, Fretz (PH); Cobbs Cr., Githens (PH); Phila- delphia, Griffith 260 (PH), s.m. (P), James (GH); York Furnace, Keller (PH); Neshaminy Cr., Long 4595 (PH); Wissahickon Cr., Penllyn, Long (PH); Cobbs Cr., MacElwee (PH); Fishing Cr., Colum- bia Co., Meredith (PH); Greenlane, Mumbauer 313 (PH); Philadel- phia, Nutt(all), perhaps an isotype of E. Nuttallii (PH); Brinton's Bridge, Brandywine Cr., Chester Co., Pennell 2280 (NY), 5110 (PH); White Clay Cr., Avondale, Pennell 5014 (PH); Chester Cr., Darling, Middletown Twp., Pennell 5058 (PH); Unionville, Porter (PH); Allentown, Pretz 5175, 6227 (PH); Saucon Cr. Center Valley P. O., Pretz 7279 (PH); McCalls Ferry, Small (NY); river, Harrisburg, Small (NY); Collen Brook, B. H. Smith (PH); canal, Belmont, C. E. Smith (PH); Philadelphia, Watson (L); Dunning Cr., Cessna, W. F. Westerfeld 6357 (FSU); Steelton, C. S. Williamson (PH); Martins Cr., Williamson (PH); Del(aware) R., above Richmond, Wister (PH); Delaware R., Philadephia, Zantzinger (MO). DELAWARE: Wilmington, Herb. Canby (GH); New Castle City, Commons (NY); Brandywine Mill Race, Wilmington, 1874, Commons (MO, NY), and in 1896 (PH); Red Clay Cr., Greenbank, Commons (PH); Wilmington, Rhoads 831 (PH); Middleton, Van Pelt & Long (PH). MARYLAND: Miles R., Easton, Talbot Co., Earle 4281 (DUKE); Plummers L, Killip 32656 (US); by Canal, Maxon 6329 (US); Annapolis, Rothrock (F); Havre de Grace Light, Shull 146 (GH, MO, US); Baltimore Co., K. A. Taylor (NY); Great Falls, Ward 126 (US), without collector (US). DISTRICT OF COLUMBIA: Eastern Branch, Potomac, Boettcher 2543 (CAS); Potomac pools, Morris (F); Georgetown, Van Eseltine & Moseley 203 (US); Potomac, Washington, E. S. Steele (Us); Washington and vicinity, Steele (C, CP, DUKE, G); Great Falls, Ward (vs); Washington, Ward (US). WEST VIRGINIA: Cacapon R., Hardy Co., Moldenke 6765 (NY). VIRGINIA: Appomatox R., Hopewell, Fernald, Long & Smart 5592 (GH); Suffolk, Kearney 1697 (US); Great Falls, McAtee 2311 (us); Hunting Cr., McAtee 2339 (Us); Black Pond, Fairfax Co., Muenscher 3614 (cU); Mountain L., Salt Pond Mrt. 10 Rhodora [Vol. 67 Giles Co., Thorne 11395 (FSU); Four Mile Run, Alexandria, Tide- strom 94 (S). NoRTH CAROLINA: Jordans Mill Pond, Seaboard, Beal 3737 (NCSC); Garysburg, Beal 3816 (NCSC); Knotts I. Currituck Co., Beal 3956 (NCSC); Wiggins Rd., Wilson Co., Beal 4114 (NCSC); Moore’s Pond, Youngsville, Beal 5690 (NCSC); L. Logan, Sunburst, Pisgah Natl. For., Beal 5835 (ncsc); Linville R., Pineola, Beal 6006 (Ncsc); Alligator R., Fort Landing, Tyrrell Co., Radford 4616, 5409 (NCU); Perquimans R., Hertford, Radford 4686 (NCU); State Fish Hatchery, Marion, McDowell Co., Radford 4886 (NCU); Pensa- cola, Yancey Co., Radford 4956 (cul: Currituck Sound, Duck, Dare Co., Radford 5436 (NCU); Boone’s Mill Pond, Jackson, Radford 5739 (ncu); Island Cr., Trent R., Jones Co., Radford, Haesloop & Miller 7683 (NCU); Jack Smith Cr., New Bern, Craven Co., Whitford 231 (ncsc); Avery Co., Whitford (ncsc); L. Myra, Wake Co., Whitford (Ncsc). ALABAMA: Little Bay Batte, Lower Mobile Bay Delta, Lueth 53 (DUKE). MICHIGAN: Jacobsville, Herman 7642 (K). OHIO: without locality, Læ (= Lesquereux), (BERN, G); Loraine, Herb. Oberlin College (Us); Fox L., Wayne Co., Selby & Duvel 1321 (NY); Columbus, W. T. S(ullivant), (G, M); Geauga L., Portage Co., Webb (Gu), INDIANA: Blue R. Lake, Whitley Co., Deam 14458 (Deam Herb., Uc); Wolf L., Noble Co., Deam 20721 (Deam Herb.); Wabash R., Fountain Co., Deam 22982 (Deam Herb.). KENTUCKY: Kentucky R., Peter (K, L, NY); Lexington, Short (K, P). MississiPPI: Horse- shoe L., Holmes Co., F. A. Cook (Us). WISCONSIN: Wisconsin R., Merrill, Fassett 7376 (WIS); Mineral L., Mellen, Fassett 7377 (WIS) ; L. Superior, Oronto, Gillman (GH); Madison, Hale (F); Milwaukee, Hase (NY); Whitewater, Kleeberger (cas); Milwaukee, Lapham (G); L. Mendota, Madison, Mahoney (WIS); Sullivan, Shallert (ZT); Whitney’s Slough, Green Bay, Shuette (F); Point Sable, Green Bay, Shuette (F, K); St. Francis R., s. Wisc., Widman (MO). ILLINOIS: Cook Co., Babcock (mo); Indian L., Eggert (CAS, F, MO, UC, US); Keokuk Reservoir, Niota, Eyles 318 (NCSC); Chicago, Gates 632 (F); Chicago R., Bowmanvil, Gates 913a (F); Athens, E. Hall (F, G, P), 1079 (GH) ; without locality, Hall (Gu) ; Oquawka, Patterson (F, NY); Murphys (boro), Shuette (F, UC); Pecatonica, Swezey (CAS). MINNESOTA: Winona, Holzinger (us); L. Winona, Holzinger (NY); without locality, Holzinger (cs); Swan L., Nicollet Co., Metcalf 6 (cu, Us); L. Chisago, Metcalf 1232 (GH); Crystal L., Blue Earth Co., Moore 23682 (S); Center City, B. C. Taylor (B, BM, G, NY, RM, s, UC, US, ws). IOWA: Pike Cr., Muscatine Co., Davidson 4409 (Ncsc); Harper's Ferry, D. E. & M. S. Eyles 338 (NCSC); Fayette, Fink (GH); Johnson Co., Fitzpatrick (MO) ; without locality, Harvey (GH); Swan L., Madison, Thorne 10472 (NCSC). Missourr: Kennett, Bush (Mo); Carondelet, Dewart (MO), without collector (WU); Poplar Bluff, Dewart, in part (MO); Gascondy, Emig 218 (MO, US) ; American bottom, opposite St. Louis, Engelmann (MO); St. Louis, 1965] Elodea — St. John 11 Engelmann, holotype and isotypes of Udora verticillata Spreng. var. minor Engelm. (B, NY); St. Louis, Gurney (MO); Pink’s slough, Allentown, Kellogg (Mo); Jefferson City, Krause (M0) ; Ha-ha-tanka, Camden Co., Metcalf 924 (Us); Rich Hill, Bates Co., Metcalf 992 (GH); Wayland, Metcalf 1089 (Ny, US); Forest Mill, E. J. Palmer 3765 (MO, NY), 3766 (GH, US); Flat Cr., Barry Co., Redfearn d T. Stombaugh 4210 (FsU). NEBRASKA: Carter L., Omaha, Kiener 17601 (G); Enders L., Thomson 9 (US); Sweetwater Lakes, Thomson 177 (GH, NY, US). KANSAS: Linn Co., Clothier 1052 (GH, MO, NY, P, US, wu); Leavenworth, Fendler (GH). COLORADO: Larimer Co., Cowen (RM). IpAHO: L. Coeur d’Alene, Epling & Houck 10053 (LA); Fernan L., Coeur d'Alene, Rust 373 (Us). WITHOUT LOCALITY: Goldie (KX); Torrey (K). The sterile specimens formerly listed for Oregon (Rho- dora 22: 29, 1920) are now known to be Elodea columbiana St. John. The record for Alabama, Mobile Bay Delta, Lueth 53, seems far out of range, and its occurrence there seems best explained as an escape from aquarium culture. Prof. J. Ewan agrees with this interpretation. Again, the occurrence in northern Idaho, in and near Lake Coeur d'Alene, is remote from the rest of the range. The shores of the lake are a favorite hunting ground, and local sportsmen have imported and established duck food plants to attract the birds. They have successfully estab- lished Zizania interior, and it seems probable that Elodea Nuttallii was introduced in the same manner. Cultivated specimens of the species are known from Germany, and the species is established in England, Hol- land, and probably in some other European countries. Elodea Nuttallii is the type and only member of the sec- tion Natator. Its sessile staminate buds which are liberated to expand and float on the surface of the water are unique. They usually lack petals but have 9 stamens in two series, 6 outer, and 3 central. Planchon's species was tranferred and adopted by Ryd- berg as Philotria Nuttallii (Planch.) Rydb., but only as a provisional name. Later, in 1913, it was validly published, as P. Nuttallii (Planch.) Rydb. ex Britton & Brown. It is evident that Rydberg did not examine Planchon's holotype. 12 Rhodora [Vol. 67 Rydberg cited six collections as this species, from New York, New Jersey, and Virginia. These specimens, in the New York Botanical Garden, have been reexamined. They differ in foliage and in flowers from the true E. Nuttallit. Instead, there is no basis of separation in the acuteness of their leaves or in the structure of the staminate flowers from the older E. canadensis. The writer positively refers these specimens, which represent the sense of Philotria Nuttallii Rydb. (but not of Planch.), to the synonymy of E. canadensis. 19. (3. of part 1.) Elodea canadensis Rich. in Michx., Fl. Bor.-Am. 1 (1803) 20; Richard, L. C., Mém. Inst. de France 12 (2), (1811-1812 — 1814) 3-4; E. canadensis var. latifolia (Casp.) Aschers. & Graebn., Syn. Mitteleur. Fl. 1 (1897) 403; E. canadensis var. latifolia var. repens Sanio, Verh. Bot. Verein Prof. Brandenburg 32 (1891) 121, an illegitimate epithet; E. canadensis var. latifolia subvar. repens Aschers. & Graebn., Syn. Mitteleur. Fl. 1 (1897) 404; E. canadensis var. Planchonii (Casp.) Farw., Am. Midl. Nat. 10 (1927) 203; E. Planchonii Casp., Monatsber. Kgl. Preuss, Akad. Wissensch, 1857 (1857) 47; also in Prinsheim's Jahrb. Wissensch. Bot. 1 (1858) 468-469, 500-501; E. latifolia Casp., Monatsber. Kg. Preuss Akad. Wissensch. 1857 (1857) 46, (and reprint p. 11, 1857) ; also in Pring- sheim's Jahrb. Wissensch. Bot. 1 (1858) 467-468; E. oblongifolia Michx. ex Casp., Pringsheim’s Jahrb. Wissensch. Bot. 1 (1858) 462; E. iowensis (as Iowensis) Wylie, Proc. Iowa Acad. Sci. 17 (1910) 82, a provisional name; and in Science n.s. 33 (1911) 263; E. ioensis Wylie, State Univ. Iowa, Nat. Hist. Bul. 6(4) (1913) 48, pl. 1-2; Anacharis Alsinastrum Babington, Ann. & Mag. Nat. Hist. II, 1 (1848) 83-84, 86; also in Ann. Sci. Nat. Bot. III, 11 (1849) 74; A. canadensis Planch., Ann. & Mag. Nat. Hist. IT, 1 (1848) 86, and in Ann. Sci. Nat. Bot. IIT, 11 (1849) 75; A. canadensis Planch. var. Planchonii (Casp.) Victorin, Univ. Montréal, Contrib. Lab. Bot. 18 (1931) 40; A. Planchonii (Casp.) Rydb., Fl. Prairies & Plains Cent. N. Am. (1932) 57; A. Planchonii (Casp.) Peck, Man. Higher PI. Ore. (1941) 76-77; Philotria canadensis (Rich. in Michx.) Britton, Science II, 2 (1895) 5; P. Planchonti (Casp.) Rydb., Bull. Torrey Bot. Club 35 (1908) 462-463; P. iowensis (as Iowensis) Wylie, Proc. Iowa Acad. Sci. 17 (1910) 82; and Science n.s. 33 (1911) 263; Udora canadensis (Rich. in Michx.) Nutt., Gen. N. Am. Pl. 2 (1818) 242, as to basionym, but excluding the linear-leaved Philadelphia speci- men which is the holotype of E. Nuttallii; Serpicula canadensis (Rich. in Michx.) Eaton, Man. Bot. ed. 5, (1829) 5390. Fig. 1, 2 a-d, 5. Description of all specimens examined: Submerged aquatic, root- 1965] Elodea — St. John 13 PLATE 1308 Fig. 1. Elodea canadensis Rich. in Michx. Holotypic sheet in the, Museum National d'Histoire Naturelle, Paris. T T Seen) b H 10 15 mm. mm. 01mm o LL Fig. 2. Elodea canadensis Rich. in Michx., staminate flower, from L. N. Gooding 597, X 5; b, pistillate flower, from Cayuga Lake, W. R. Dudley, X 5; c, leaf from holotype, X 5; d, leaf apex of same, X 50; e, Elodea Nuttallii (Planch.) St. John, staminate spathe and bud, from Okuawka, Ill, H. N. Patterson, X 5; f, expanded staminate flower, from Old Lyme, Conn. R. W. Woodward, X 5; g, pistillate flower, from Center City, Minn, B. C. Taylor, X 5; h, leaf, from H. St. John 1,827, X 5: i, leaf apex of same, X 50. 1965] Elodea — St. John 15 ing in the mud; plants dioecious; staminate plants rarely observed, their stems 2-10 dm. long; lower leaves opposite, small, lance-ovate; median and upper leaves in whorls of 3 and 7-17 mm. long, 1-4 mm. broad, linear, lance-linear, or lance-oblong, bright green, thin, firm or subflaccid, finely serrulate; staminate spathes borne in the upper axils, 15 mm. long, the lower half narrowed into a peduncle-like base, the upper half 6-8 mm. long, 4 mm. wide, inflated, ellipsoid or ovoid, the apex resembling a gaping mouth, due to the two acute, salient lobes; staminate flowers peduncled by the slender, thread-like, elongating base of the hypanthium which is 3-20 em. long; sepals 3.5-5 mm. long, 2-2.5 mm. wide, elliptic, dark striate; petals 5 mm. long, 0.3-0.7 mm. wide, white, very delicate, the basal half a linear claw, the apical half linear lanceolate; stamens 9, the 3 inner raised on a common, fused stalk; anthers 2-3.5 mm. long, 1 mm. wide, oblong-ellipsoid; pistillate plants with stems slender, dichotomously branched; lower leaves opposite, ovate, much smaller than the upper ones; median and upper leaves in whorls of 3, and 6-13 mm. long, 1-5 mm. broad, oblong, ovate-oblong or lance-ovate, firm, dark green, serrulate, crowded and strongly imbricate at tip of the stem; pis- tillate spathe 10-20 mm, long, borne in upper axils, cylindric, the apex bidentate and with two broad teeth; pistillate flower exserted from the spathe by the thread-like, elongating base of the hypan- thium which is 2-28 cm. long; sepals 2-2.2 mm. long, 1.1 mm. broad, oblong elliptic, dark striate; petals 2.6 mm. long, 1.3 mm. wide, broadly elliptic-spatulate, white, delicate; staminodia three, 0.7 mm. long, acicular; style equaling the hypanthium; stigmas three, 4 mm. long, broad, 2-cleft at apex; ovary 3 mm. long, lance-ovoid; ovules 3-4, erect; capsules 6 mm, long, 3.2 mm. in diameter, ovoid; seeds 4.5 mm. long, narrowly cylindric, glabrous. Holotypus: “in rivulis Canadae.” Environs de Montreal, Elodea oblongifolia, A. Michaux (P)! Type examined! Though the speci- mens bear the name E. oblongifolia, it was published as E. canadensis Range: In fresh ponds and streams, especially in calcareous areas, from Quebec, abundant southward to North Carolina, Alabama, and Arkansas, westward to Manitoba, British Columbia, Colorado, and central California, but absent in the higher Rocky Mountain region. NORTH AMERICA, Herb. Durand (P); in 1823, Greville (K); Michaux (P); in 1827, Schweinitz (B); donné par W. Hooker (P). CANADA: Cleghorn (K); Ny. He. [Norway House] to Canada, R. D. (K); Herb. Sprengel (B). QUEBEC: Corris Sta., Cleveland, Chamberlain & Knowlton (GH); L. Memphremagog, Churchill (GH, K), Aug. 18, 1906, Churchill (GH) ; L. Bernard, Farrellton, Hull Co., Rolland-Germain 6472 (S); flum. St. Laurent, Mont Real, Hennecart (P); Magog L., Orford, Knowlton (GH); R.-aux-Serpents, Deux- Montagnes Co., Pere Louis-Marie (s); Hull, Macoun 27001 '(CAN) ; 16 Rhodora [Vol. 67 Farm Pt. Gatineau R., Macoun 61033 (CAN); Ste. Anne de Beau- pré, Macoun 68806 (CAN, GH); Hull, Macoun 85553 (CAN); Pickanock R., Macoun (GH); Ottawa R., Hull, Macoun (F); St. Lawrence R., Caughnawaga, Victorin 8171 (BM, DEAM HB.); R. St.-Laurent, Mon- tréal, Victorin 9910 (P); Longueuil, Victorin 11256 (DEAM HB., P, US); Montréal, Victorin 20318 (GH, P, S); Longueuil, Victorin 24725 (s); Montréal, Victorin 28288 (K, s); Pt. à Bison, Co. de Beauhar- nois, Victorin 28393 (K, S); rapides Richelieu, Chambly, Victorin & Germain 33905 (s); R. Richelieu, St.-Mathias, Chambly Co., Victorin & Germain 7310 (K, LD); R. Richelieu, St.-Mathias, Chambly Co., Victorin & Germain 7410 (FSU, S) ; R.-des-Prairies, Hochelaga Co., Victorin & Germain 33996 (s); Ile Ste.-Therese, St.-Jean Co., Victorin & Germain 45631 (C), 49140 (G), 49147 (W); Ste.-Rose, Laval Co., Victorin & Germain 49306 (c, w); Ile Ste.-Therese, Sainte-Jean Co., Victorin & Germain 49361 (G, M, W); Melocheville, Saint-Laurent R., Victorin, Germain & Boivin 4302 (S), Lac Olive, Reg, de l'Abitibi, Victorin, Germain & Meilleur 44974 (c); Ile Calu- met, Ottawa R., Victorin, Germain & Rouleau 2017 (W); Ste.-Anne de Sorel, Richelieu Co., Victorin, Germain & Rouleau 6534 (C, S). ONTARIO: Kapuskasing R., Cochrane Dist., Baldwin & Breitung 3515 (K); Cochrane, Lillabele L., Baldwin & Breitung 3898 (K); Ger- man Twp., Timmins, Cochrane Co., Baldwin & Breitung 4062 (s): Detroit R., Herb. Boott (GH); London, Burgess 2208 (BM); Frog L., Hardwick, Garton 1676 (G, w); Chelsea, Harrington (CAN); St.- Thcmas, Elgin Co., L. E. James 1711 (C, G, L, W); Sparrow L., Kirk 679 (L); Stokes Bay, Bruce Penin., Krotkov 8649 (NY); St. Clair Flats, McAtee 3424 (Us); Kaministiqua R., Neeling, Thunder B. Dist., Mayall & Cormack 24 (w); Lake Region, Macoun (K); Bay of Quinte, Macoun (CAN); Niagara Falls, Macoun 82331 (CAN); Kenora, Lake of the Woods, Ostenfeld 552 (c, cP); Ccldwell, Osten- feld 584a, 584c (C, CP) ; L. Superior, Coldwell, Rendle & Good (BM); Toronto, Scott (CAN); Jock R., Beckwith, Lanark Co., Senn 552 (S); Moose Factory, Spreadborough 62633 (CAN); Toronto, Sturtevant (Mo); Pancake R., Algoma Dist., Taylor, Hosie et al. (s); L On- tario, Belleville, Hastings Co., Victorin, Germain & Jacques 45855 (GB); Niagara, C. S. Williamson (PH). MANITOBA: R. des Marais, Otterburne, Bernard 58284 (S): Brandon, Macoun 13724 (B, CAN, F); Souris R, N. W. T., Macoun 27002 (CAN, GH); Wekuske L, Scoggan 6680 (C, K, S); Grassy R., Tramping L., Scoggan 6900 (C, K); Whiteshell For. Res., Scoggan 8723 (K); Minnedosa R., El- phinstone, Scoggan 10268 (C, S). SASKATCHEWAN: Carleton House without collector, (GH): without lccality, Drummond, type of E. Planchonii Casp. (GH, K). BRITISH COLUMBIA: Mara L., Enderby to Sicamous, Calder & Savile 11838 (w). UNITED STATES. Marne: L. Auburn, Auburn, 1953, Bean (NEBC); Benton Falls, Sebasticook R., 1959, Bean (NEBC); Lily Pond, Rockport, C. A. E. Long 906 (NEBC, NY); Rockport, Long 1965] Elodea — St. John 17 (NEBC, NY); calcareous water, Lily Pond, Rockport, Rossbach 96 (NEBC) ; Unity Pond, Unity, Rossbach 97 (NEBC). NEW HAMPSHIRE: Harris Pond, Pelham, Hillsborough Co. Dupee (NHA); Partridge Pond, Littleton, Grafton Co., Sheehan & Krochmal 53 (NHA). VER- MONT: Winooski R., Essex Jct., S. F. Blake 2210 (NEBC, PH); N. Hero, Brainerd (P); Herb. Chapman (uc); Fern L., Leicester, Dut- ton (CU, MO); Catharine Wells, Dutton & C. W. Dodge (NY); L: Bomoseen, W. Ferrisburg, Eggleston (GH); Little Otter Cr., Fer- risburg, Eggleston & Grout (Gu); Ferrisburg, Eggleston (F). MASSA- CHUSETTS: Andover, J. Blake 2210 (NEBC); Pontoosuc L., Lanesboro, Churchill (M0) ; Tyringham, H. T. Clark (BR); L. Garfield, Monterey, Hoffman (NEBC) ; Sheffield, Hoffman (NEBC) ; Fresh Pond, Cambridge, Morong (MO); Pontoosuc L., Pittsfield, Kennedy (GH); L. Quannapo- witt, Wakefield, Ripley 16138 (GH), 17494 (GH, NEBC) ; Great Brook, Southwick, Seymour 257 (MO, NY). RHODE ISLAND: N. Providence, Olney (BRU, S), Curtiss (G), Hope (NY), Olney (BRU), Olney (S). CONNECTICUT: L. Saltonstall, Blewitt 1981 (NEBC); Norwalk, E. H. Eames 5265 (NY); L. Congamond, Eames 8457 (GH); Boardman’s Pond, E. Hartford, Weatherby 367 (NCSC) ; Housatonic R., N. Canaan, Weatherby 2700 (NEBC); E. Haven, Weatherby 3591 (NCSC) ; Housa- tonic R., Huntington, Harger & Weatherby 3595 (NCSC) ; Huntington, Weatherby 3604 (NEBC), 3595 (WEATHERBY HERB.); Branford, Setchell (uc); E. Haven Pond, Herb. Torrey (NY); Wright in part (MO). NEW YorK: Fall Cr., Ithaca, Bechtel 13364 (CU) ; New York, Bicknell 174 (NY); Thousand Is., Bicknell 177 (NY); Chilton L., Brit- ton (NY); New York, Britton (G); Kendall, Burnett (NY); Harris Bay, L. George, Burnham 53 (us); Fall Cr., Ithaca, Clausen 2386 (G); Buffalo, Clinton (F, MO, NY); Grand I., Niagara R., Coville (us) : Cayuga L., Dudley (DS); Springport, A. J. Eames 13362 (cu); Ren- wick, Eames & Wiegand 13353, 13353a (cU); Cayuga L., Eames & Wiegand 13356 (CU); Mattituck, Ferguson 2405 (NY): Oakdale, Fer- guson 7879 (NY); Cayuga L., Fernald & Wiegand 14548 (CU, GH); Watertewn, Fernald, Wiegand & Eames 14125 (cU, GH); Ossawegat- chee, Fernald, Wiegand & Eames 14126 (cv, GH); Clayton, Fernald, Wiegand & Eames 14127, in part (CU, GH); Selkirk, Fernald, Wiegand & Hames 14128 (CU, GH); Watertown, Gray (NY); w. N. Y., Gray (GH); Gray (K, M); Renwick, R. Hitchcock & A. R. Bechtel 11218 (cU); Pecksport, House 6561 in part (GH); Pierrepont Pond, Wood- ville, House 8223 (NY); Irondequoit Bay, Killip 1063 (M0) ; Oswego R., Oswego, Killip 12533 (us); Jennings Pond, Danby, Lawrence 831 (S); without locality, 1821, Mitchell (G); L. Ontario, Morong (Mo, NY); Cayuga L., Ithaca, Morong (BM, NY); Danby Pond, Danby, Muenscher 19584 (S); Fall Cr, Ithaca, Muenscher 13354, 13363 (CU, GH) ; Oneida L., Muenscher 14547 (CU); Fall Cr., Ithaca, Muenscher & Bechtel 50 (oi: Cayuga L., Ithaca, Muenscher & Bechtel 51 (0); Geneva, Muenscher & Burkholder 16366 (ws); Conesus L., Muenscher & Burkholder 16367, 16370 (ws); Silver L., 18 Rhodora [Vol. 67 Wyoming Co., Muenscher & Burkholder 16368 (WS), 16369 (GH, WS) ; Irondequoit Bay, Muenscher, Wiegand & Wright 15197 (CU); Sodus Center, Wiegand & Wright 15198 (cU); Thousand Is., J. I. & G. R. Northrup (NY); Junius marlbog, Seneca Co., Ostenfeld (c); Tiough- nioga R., Cortland, E. L. Palmer 73 (cu); Seneca Falls, Cayuga L., Palmer & Eames 72 (cU); Grass R., Canton, Phelps 279 (CU, GH); P Y = (Penn Yan), Sartwell (F); Cayuga L., H. von Schrenk (MO); Fall Cr. Ithaca, von Schrenk (M0); Albany, Herb. Schinz (Z, ZT); Bainbridge, Topping (US); Oneide by New York, Torrey (P); Ithaca, Trelease (MO); Fish Cr., Oneida L., Underwood 3211 (NY); Lake View, L. Ontario, Jefferson Co., Underwood (NY); Charlotte am Ontario See, Weinland 49 (B); North Pond, Sandy Cr., Oswego Co., Wiegand 13355 (CU); N. Fairhaven, Wiegand 13358, in part (CU) ; McMullen Pond, Wiegand 13359 (cu); Renwick, Wiegand 13360 (cu); Fall Cr., Ithaca, Wiegand 13361, in part (CU); Spencer, Wiegand & Fernald 14546 (CU, GH); Cayuga L., Ithaca, Wiegand & C. C. Thomas 1533 (cv); Wellesley I., St. Lawrence R., Ward (us); L. Katrine, Ulster Co. Wilson (NY). NEW JERSEY: Foul Rift, Warren Co., Bartram (PH); Muscanetcong R., Asbury, Warren Co., Godfrey 62102 (FSU); Delaware R., Delaware Co. Mackenzie 7233 (NY); New Brunswick, Mackenzie 7258 (NY); Green Pond, Warren Co, Mackenzie 7732 (NY); Raritan R., New Brunswick, Mackenzie (NY); Rockaway, Mackenzie (NY, S); Stillwater, Mac- kenzie (NY); Clifton, Nash 1 (NY); Morris Co., Rusby (MO); Sussex Co. Rusby (BRU); Budd's L., Sussex Co., Small (C, F); Pensauken Cr., Trudell (NY); Foul Rift, Van Pelt (PH); Cooper's Ferry, opp. Kensington, without collector (PH). PENNSYLVANIA: Ingham Sprs., Bucks Co., Bassett (s); W. Chester, W. D[arlington] (K, LD); Peach Bottom, Eby (Mo); Sellersville, Fretz (UC); without locality, Herb. Guthnik (BERN); Flinton, Jennings (CU, MO); Churchtown, Long (PH); Bloomsburg, Meredith (PH); Bethlehem, Moser (L, NY); Lancaster, Mühlenberg (M); without locality, Mühlenberg (S); Philadelphia, Pickering (PH); Erie, Porter 91 (PH); Pot Rock, Easton, Porter (PH); Saucon Cr., Friedensville, Pretz 4183 (PH); Allentown, Pretz 4784 (PH); Cedar Cr., Griesemersville, Pretz 6962 (PH); Centre Valley Sta., Pretz 11212 (PH, ws); Juniata R., Roth- rock (F); without locality, L. B. de Schweinitz, in part (BR); Bethle- hem, Schweinitz, in part (PH); Harrisburg, Small (F); Telford, Strohm (PH); Bethlehem, without collector, in part (BR). DELAWARE: Delaware City, Commons (GH, PH); Wilmington, Tatnall (MO). MARYLAND: Susquehannah, Herb. Buckley (M0); C. & P. Canal, near Lock 12, Killip 32129 (P); Ipesutic L, J. D. Smith (us). DISTRICT OF COLUMBIA: Rob’s Pt, Potomac R., Coville 120 (US) ; Washington, Vasey (US), Ward (US). WEST VIRGINIA: Booth Cr., Uffington, Monongalia Co., Davis 205 (ncu); Kanawha Co., Holton (F); Wallace Switch, Small (Ny). VIRGINIA: without locality, Bey- rich (B); Big Walker Cr. Poplar Hill, Giles Co., Kral 13971 1965] Elodea — St. John 19 (rsv); V. P. I. grounds, Montgomery Co., Massey (NCU); Hunting Cr., McAtee 2281 (US); Waynesborough, Murrill (NY); Four-Mile Run, Shull 467 (us); New R., Goodwin Ferry, Thorne 17862 (FSU); Four Mile Run, near Alexandria, T'idestrom 94 (P); The Dyke, Potomac R., Tidestrom 7185 (US). NORTH CAROLINA: shallow pond, Perquimans Co., 1932, Glasson (DUKE); University L., Chapel Hill, Hueske (NCU); Snyder’s L., Blowing Rock, Watauga Co., Radford 4950 (NCU); pools, State Fish Hatchery, Marion, McDowell Co., Radford 5279 (NCU). MICHIGAN: Detroit, Bigelow (M0); R. Rosin, Monroe, Chandler (US); Wolf L., Chase 1711 (US); Arenac Co., Dodge (US); Black R. near Black L., Cheboygan Co. Ehlers 5902 (S); Ocquecoc L., Presque Isle Co., Ehlers 6256 (S); Douglas L., Cheboygan Co., Ehlers 6280 (S); Hook Pt., Mich. Biol. Sta., Cheboy- gan Co., Gates 14143 (S); Douglas L., F. C. & M. T. Gates 10592 (F, MO); Sault St. Marie, Gillman (GH); Fort Gratiot, Gillman (GH); Sailor’s Encampment, E. T. & S. A. Harper (B); Liver-Light Lakes, Metcalf 2222 (c, cP); Manistee, Morong (GH, NY); Kewenah BE L. Superior, Robbins; Whitmore L., Stearns 305 (C, CP, G, GH, K, LD, NCSC, NY, 0, ZT); Reed's L., Grand Rapids, C. F. Wheeler 7663 (cs); Swan Cr., Old Fort, Wight 41 (us); Lower Scott L., Lee, Wight 73 (us); Ann Arbor, Winchell 7760 (M0); Sault Ste. Marie, no collector (PH). On: Cincinnati, J. Clark (BM); unio itiner., Frank (G, K, P); Clear Cr., Hamilton Co., J. F. James (Ds, US); Sandusky, Moseley (F), R. C. O. & E. B. W. (Deam Herb.) ; Put-in- Bay, Pieters (Us); East Harbor, Pieters (us); Braceville, A. N. Reed & Webb 929 (GH); without locality, J. L. R. (P); Newark, E. B. Williamson (Deam Herb.). INDIANA: Wolf L., Chase 1711 (c, F, Mo, 0); L. Maxinkuckee, H. W. Clark 1 (Us), 1593 (US); Rochester, Deam 11765 (Deam Herb.); L. James, Steuben Co., Deam 20245 (Deam Herb.) ; Princeton, Deam 25520 (Deam Herb.); Pendle- ton, Deam. 25599 (Deam Herb.); Deshee R., Decker, Deam 2922, (Deam Herb.); Waldron, Deam 30087 (Deam Herb.) ; Knox, Deam 42177 (Deam Herb.); Harrison, Deam (Deam Herb.) ; L. Maxin- kuckee, Evermann 1070 (US); 5-Mile Pond, Terre Haute, Evermann (F, US): Wolf L., Greenman (MO); Tippecanoe R., Talma, Fulton Co., N. C. Henderson 61-471 (FSU); Bedford, Kriebel 3479 (GH); E. Chicago, Lansing 1767 (F); L. Maxinkuckee, Scovell & H. W. Clark 1070 (ps, F); Mishawaka, E. B. U. (F); Whitings, no collector (PH). KENTUCKY: Reelfoot L., Alexander 323 (us); Kentucky R., Peter (PH); without locality, Herb. C. W. Short. ALABAMA: Lightsey's Pond, Centerville, R. M. Harper 3287 (NY, PH, US). WISCONSIN: Lauderdale, R. Bebb 1531 (F); Prairie du Chien, Fassett 5049 (Glück Herb., Heidelberg); Kawaquesagon, Minocqua, Fassett 5328 (Glück. Herb.) ; Beloit, Fassett 5691 (Glück Herb.) ; St. Croix Falls, Fassett 7378, 7379 (wis); Nemakagon R., Cable, Fassett 7380 (wis); L. Wiehe, Eliot, Fassett 7381 (WIS); Tomahawk R., Fassett (Gliick Herb.); Trempealeau, Fassett & Wilson 5050, 5051 20 Rhodora [Vol. 67 (Glück Herb.); Bear L., Barron Co., Goessl 6907 (B); Nobleton, Washburn Co., Goessl 7471 (B); Cameron, Barron Co., Goessl 8785 (B); Milwaukee, Hasse 2531 (PH); Whitewater, Kleeberger (CAS); without locality, Lapham (M0); Prairie du Chien, H. H. Smith 7422 (B). ILLINOIS: Mound Cr., Ogle Co., M. S. Bebb (PH); Ogle Co., Bebb (F); Peoria, Brendel (P); “Canes,” De Selm 717 (F); Grass L., Lake Co., Gates 1758.4 (Deam Herb.); Oregon, Waite (US). MINNESOTA: Prior’s L., Scott Co., Ballard (NY); Clearwater Co., Buell 1650 (NCSC); E. Twin Pond, Itasca Park, Clearwater Co., Buell 1651 (NCSC); Ft. Snelling, Campbell (F, MO); L. Winona, Holzinger (NY, us); L. of the Woods, MacMillan & Sheldon 572 (CU, GH), 3334 (NY); Ft. Snelling, Mearns 804 (us); Squaw L., Cass Co. Metcalf 2364 (GH); N. W. Territory, Nicollet 415 (PH); Thompson, Sandberg 563 (ws); Hennepin Co., Sandberg (F, UC, US) ; Center City, B. C. Taylor (0). Iowa: Fayette Co., Fink 552 (US); Spirit L., Dickinson Co., Anderson (C, G, GH, L, LD, NCSC, S, W); Clinton Co., Butler 27 (MO, P); Des Moines R., Emmet Co., Cratty (NY); L. Okoboji, Wolden (GB); Pilot Knob State Park, Hancock Co., Thorne 14595 (NCSC); Cheever L., Estherville, Emmet Co., Thorne (NCSC); E. Okoboji L., Dickinson Co., Wylie, type of E. iowensis Wylie (GH, NY). MISSOURI: Neosho, Bush (MO); St. Louis, Engelmann (NY); Phillips Spr., Van Buren Cc., Redfearn 5742 (FSU) ; Mill Spr., Wayne Co., Russell: Trc- lease (MO). ARKANSAS: Mammoth Spr. Trelease (MO). NORTH Dakota: L. Metigoshe, (C. E.) Lee (RM); Minot, Lunell (Deam Herb., DS, NY, RM, US); Jim L., Pingree, Mabbott 321 (us); Fargo, Waldron (MO). NEBRASKA: Loup Fork, Hayden 1553, 1554 (MO); without locality, Hayden (NY); Jackson L., N. Platte, Kiener 17514 (G); Kingsley Dam, Keith Co., Kiener 15537 (G): Fremont, Williams 373 (US). MONTANA: Lindberg L., Swan R., Missoula Co., C. L. Hiteh- cock & Muhlick 13712 (S); S. Pablo Reservoir, Lake Co., Hotchkiss 6349 (US). IDAHO: L. Pend d'Oreille, Leiberg 9 (C). WYOMING: Fish Hatchery, Nelson 1429 (RM), 5374 (BM, GH, MO, NY, RM, US). COLORADO: Alamosa, Biltmore Herb. 105a, Colo. Exped. 1-897 (US); Rio Grande, Alamesa, Shear 8746 (NY). UTAH: Panguitch L., Jones 6018 (US); Fish L., Locke 4 (US). NEVADA: (formerly in Utah), Carson R., Ragtown, Remy (B, P). WASHINGTON: Seattle, Bardell (M0) ; Green L., Seattle, Congdon (MO); Seattle, Eyerdam (G); Clover Cr., Pierce Co., Eyerdam 1221 (G, s); 98th st., (Seattle), Eyerdam 6627 (G); Lake Chelan, Gorman (Us, Ws); Green L, Seattle, Piper (ws); Fish L., Spokane Co., Preston 860 (WS); Little Spokane R., Dartfcrd, St. John 3438 (Ds, K, WS), St. John & Warren 2423 (WS); Faleon Valley, Suksdorf 87 (0); Calispell L., Pend Oreille Co., Yocom (W); Seattle, Zeller (MO). OREGON: Harriman Lodge Sta, Abrams 9719 (Ds, MO); Crater L., Coombs (CAs); Sprague R., Bly, Coville & Leiberg 318 (US): DesChutes R.. Bend, Peck 4250 in part (W, WS). CALIFORNIA: Big Meadows, Austin 573 (MO, US); Feather R., Austin 1179 (US); Plumas Co., Austin (NY); 1965] Elodea — St. John 21 Egg L., Baker (UC); Sisson, Eastwood 1268 (CAS); Folsum, Sacra- mento Co., Johannsen 652 (FSU); Mt. Eady, Krautter (BM); Sisson, Krautter (K); Walker Cr. Klamath R., Siskiyou Co., Lee 1239 (G, K); San Dimas Canyon, San Gabriel Mts., “of very doubtful indigeniety," L. Wheeler 2325 (DS, CAS, LD); Mendocino Co., Vasey (US). Discussion: Caspary in his treatment of the Hydrilleen (1858) included a detailed monograph of Elodea, in which he aecepted 10 species. He treated E. canadensis Rich. in Michx. in great detail. It is noted that he describes plants with a floral morphology quite different from those of the present concept. In E. canadensis he included plants that were hermaphroditie, others dioecious, others polygamous (fide Torrey, and fide Gray). His “hermaphroditic” plants were Michaux's type specimens of E. canadensis (hermaph- roditic fide Rich. in Michx.); and Moser's and Schwein- itz's collections from Bethlehem, Pa. which are the true E. Schweinitzii and are genuinely hermaphroditic. His dioe- cious plants were E. Nuttallii, and Anacharis Alsinastrum which was based on pistillate plants of E. canadensis when established in England, and Udora occidentalis (Pursh) Koch, in part, which is a synonym of E. canadensis. In ad- dition Caspary accepted as a species E. Planchonii Casp. which is merely the staminate plant of E. canadensis. He also made and accepted E. latifolia, based on broad leaved, sterile material of E. canadensis. He accepted E. Schwein- itzii (Planch.) Casp., though he had some doubt about it. He had seen the holotype but its flowers were mutilated and he could not confirm its characters, so he accepted it, with confidence in the description by Planchon. However, he cited four good specimens of this species as hermaphroditic specimens of E. canadensis. In sum, Caspary did not have a good understanding of E. canadensis, as he made three species out of it, while reducing to the synonymy of E. canadensis itself two other excellent species. E. guyan- nensis Rich. was maintained by Caspary, though there are no actual differences to separate it from the earlier E. granatensis Humb. & Bonpl. The floral morphology of Elodea canadensis is in need of 22 Rhodora [Vol. 67 discussion, as it has often been misinterpreted. In the original diagnosis of the monotypic genus Richard in Mi- chaux included among other details, the following: that the three stamens, with thick filaments, bore terminal, cordate anthers. In part 1 of this monograph (1962: 34-36) it is shown that of the existing 8 good, open flowers on the holotypic sheets in Paris and the isotypic sheet in Geneva, all are pistillate, with 3 acicular staminodia, all with- out anthers. The type locality was “in rivulis Canadae,” and the label adds, “environs de Montreal.” The species still grows commonly there, and the pistillate plants from the type locality have subsequently been described in detail by Victorin (1931: 15-19). Their pistillate flowers are identical in structure with those of the holotype collection, and with all other normal flowers of all other collections of wild, indigenous specimens from Canada and the United States. Their flowers are dioecious, as the functionally pis- tillate ones have 3 bifid stigmas and 3 acicular staminodia, but no anthers. These pistillate plants were the kind that in 1836 escaped from cultivation in Ireland, and with re- markable rapidity spread to aquatic habitats of most of Europe. Rydberg (1908: 458-459) stated that ‘““Caspary, who saw the original Michauxian material at Paris, stated that the two flowers found there were hermaphrodite as described.” The present writer in 1935 found Michaux’s holotype in Paris to consist of five sheets, and in Geneva one isotypic sheet, the holotype with a total of eight good flowers, but none of them with anthers. Caspary (1858: 462) stated that he examined two of Michaux’s specimens. For one he quoted from the label, “Triandr.;” for the other he said, “mit schlechter bliithe.” So, Rydberg notwithstanding, Cas- pary did not say that he saw two flowers of the type ma- terial and that they were hermaphrodite. It is clear that the existing abundant holotypic material has only pistillate flowers with 3 staminodia. The staminate plants and flowers of this species were not at first recognized as of the species Elodea canadensis. They were described as E. Planchonii Casp. (1857: 47), 1965] Elodea — St. John 23 based upon “Suskatschawan, Drummondio; Canada, Cleg- hornio.” The former specimen, Saskatchewan, Drummond (K), which has good staminate flowers, is here chosen as lectotype. These staminate flowers are strikingly different from the pistillate flowers, but are produced from the same sort of spathe which has a peduncle-like base and a biden- tate ellipsoid apex. The sepals are larger, 3.5-5 mm. long; so also are the petals, being 5 mm. long. The 9 stamens consist of 6 lower outer ones and 3 inner ones raised on a common stalk. On the holotype and on most of the stami- nate plants the leaves of successive nodes are more widely spaced, narrower, and thinner, than on the pistillate plants of E. canadensis. Since their description as E. Planchomii in 1857, these staminate plants have by most botanists been accepted as a distinct species, but by a few as a variety, Anacharis canadensis var. Planchonii (Casp.) Victorin. The first good evidence that Elodea Planchonii really represented only the staminate plants of E. canadensis came from the studies of Wiegand and Eames (1926: 55). They reported that these staminate plants grew together with the pistillate E. canadensis and they classed them as the two sexes of a single species. Several of their collections of flowering staminate plants had broad, firm, imbricate leaves, indistinguishable from those of genuine, pistillate E. canadensis. The collection from New Jersey, Morris Co., July 1879, H. H. Rusby, has the leaves as narrow as in Elodea linearis, but the staminate flowers as in E. Planchonii. The speci- men from Quebec, Longueuil, Aout 1920, Marie-Victorin 11256 (US); and that from North Dakota, Jim Lake, Pin- gree, Aug. 14, 1917, D. C. Mabbott 321 (US) have the typical staminate flowers but the leaves as short and broad and imbricate as in the pistillate E. canadensis. In 1926 to confirm the observations of Wiegand and Eames, the writer followed Dr. W. C. Muenscher, and at Ithaca, New York, waded in the estuary of a stream flow- ing into the south end of Cayuga Lake. There, flowering pistillate E. canadensis and flowering staminate E. Plan- chonii were growing in 3-4 feet of water in both adjacent 24 Rhodora [Vol. 67 and intermingling luxuriant colonies. At first glance they seemed different in herbage. That is, of course, to be ex- pected between plants of different species, but it is also possible between the two sexes of a dioecious species. An extended survey in this large colony showed that besides the different looking plants, there were also plants with in- termediate and ones with identical foliage characters. We agreed that there were no constant differences or usable distinctions in the foliage of the male and the female plants. Though the male flowers are much the more conspicuous, they have seldom been collected. However, their total range in North America falls within that of Elodea canadensis, and at almost every locality for the staminate, the pistillate is also known to occur. From these considerations it seems clear that the staminate E. Planchonii is not a distinct taxon, but is merely the staminate sex of the wide ranging E. canadensis. Staminate plants of Elodea canadensis have rarely been found in Europe. A second botanical collector with the famous name D. Douglas found and published (1880: 227- 229) on the occurrence of these plants at the Braid Hills, Edinburgh, Scotland. In the Kew herbarium there are three of his sheets from the above locality, collected in Aug. 1881. They are vigorous shoots with oblong leaves 4-5 mm. wide, and with normal, 9-anthered staminate flowers. No later collections of this plant have been seen or reported. It did not rapidly spread over Europe, instead it apparently died out. There seems to be no documentary evidence of its advent in Seotland. It may have been privately imported from North America by some individual for aquarium culture. Then it may have been wholly or in part discarded by a stream or pond, where it then became established. The other possible origin of these staminate plants in Scotland is that a flowering pistillate plant may have set some parthenogenetic seed. Some of these seeds could have grown into staminate plants. If this happened, it has been a rare occurrence. The other possible method of advent, the direct importation of staminate plants, seems the more probable explanation of its occurrence in Scotland, 1965] Elodea — St. John 25 The description of the fruit given by Victorin (1931: 19) is rather different from the writer’s. His description when translated is, “capsules 10-15 mm. long, ovoid, acuminate and with a persistent style; seeds 1-2, rarely more, lance- ovoid.” It would seem that he included the style when measuring the capsule. The details and measurements here presented by the writer came from study of a specimen from North Dakota, Lake Metigoshe, July 9, 1891, Lee (RM). In the southeastern states of the U. S. A., Elodea cana- densis has been considered absent. Here is accepted as that species a sterile collection from Centerville, Alabama, Har- per 3287. The leaves are broad and imbricate, and so typi- cal that no hesitation is felt in accepting this record. In the Kew herbarium there is another specimen labeled Ala- bama, Schweinitz. He was a Pennsylvanian botanist who collected intensively around Bethlehem, Pa., but is not known to have collected in Alabama. The flowers are small and resemble those of E. canadensis, but the herbage looks like that of E. Schweinitzii, a species found only near Bethlehem, Pa. It seems best to leave this specimen unde- termined and to question its locality. Elodea canadensis has been recorded as occurring in Texas by Coulter (1894: 421); and as Amacharis canaden- sis by Cory and Parks (1937: 14). Having seen no speci- mens of this or other indigenous species from that state, inquiries were made. V. L. Cory replied on Dec. 16, 1940, "[ regret to inform you that the inclusion of Anacharis canadensis in our Catalogue was apparently based on the report given in Coulter's Botany of Western Texas. As for myself, I have not seen this plant in Texas, nor have I Seen any material of it collected by anyone else in this State." Prof. B. C. Tharp, curator of the herbarium of the University of Texas, replied on Jan. 14, 1941, that concern- ing, "specimens of Elodea, I regret to say that I do not find any specimens in the herbarium.” Recent collections in the herbarium of the Texas Agri- cultural and Mechanical College, from Brazos, Robertson, Hays, and Leon Counties prove to be not Elodea, but the 26 Rhodora [Vol. 67 introduced Egeria densa. Thus, it appears that no indigen- ous species of Elodea occur in Texas, or in South Carolina, Georgia, Florida, or Louisiana. There is a collection of Elodea canadensis from Mexico. Its data is Morelia, Laguno, in aqua, 2,000 m. alt., 1.9.1911, Arséne (B). This locality is 130 miles slightly north of west of Mexico City, and is remote from the known range of the species. To an inquiry, Dr. F. Miranda of the Uni- versidad Nacional Autónoma de México replied on 23 Nov. 1962, *No conazio ejemplares verdaderamente indígenas de ninguna especie de Elodea (o Egeria) en México. Elodea densa, es frecuente en estanques de jardines püblicos o culti- vada en acuarios de esta ciudad." It can be safely con- cluded that the collection of E. canadensis from Morelia represents a plant cultivated or adventive in that area. One other locality record needs discussion. In the Berlin herbarium there is a specimen of E. canadensis labeled as from Labrador. It was determined as Anacharis Alsinas- trum which is a synonym of E. canadensis. It is marked "ex Mus. Parisiensis," but the bottom of the label is cut away so that the collector's name is shown only by the remaining tops of a few taller letters. Later the originals were found in the museum in Paris, three similar sheets of E. canadensis, and one of E. Nuttallii. They all have the printed label form: Amerique du Nord, M. Lamarre- Picquot, and on it in handwriting is the locality Labrador. He was sent in 1848 to North America to collect economic plants for the Ministére de l'Agriculture. From his report (1849) we learn his itinerary. He landed on the 24th of June 1848 at New York. He journeyed up the Hudson River to Lake Erie, Detroit, across Michigan, and Indiana to Chicago, Galena, Saint-Paul, Mendota at the junction of the R. Saint-Pierre with the R. Mississippi, to Lac-qui- Parle, Mendota, Saint-Paul, Buffalo, Erie Canal, Albany, and he sailed from New York on Oct. 3, 1848 to Europe. As edible plants he brought back to France Psoralea escu- lenta, Pursh and Apios americana, Medic. (A. tuberosa). His specimens of Elodea are genuine, but his locality Labrador is incredible. On his voyage he nowhere ap- 1965] Elodea — St. John 27 proached Labrador. No species of Elodea is known within many hundreds of miles of Labrador. The present writer explored the full length of the southern coast of the Labra- dor Peninsula, and wrote his doctor’s thesis on its flora. Others have explored the eastern and other sections of Labrador. This record of Elodea from that area is rejected as an undoubted error. There is a specimen of Elodea from Nicolett’s North- Western Expedition, ponds up Riviére aux Serpent near Oanus River, July 10, 1883, Charles A. Geyer (US), that was omitted in the enumeration of specimens. The plant is E. canadensis, but the locality is obscure to the writer and to the present staff of the Missouri Botanical Garden. How- ever, Dr. J. M. Greenman noted on the sheet that it was “probably from Nebraska." In the British Museum of Natural History is a specimen of Elodea canadensis, with the data: Hawaiian Ids., Udora*, the asterisk indicating a new species, then follows a specific epithet meaning broad-leaved. The epithet is not quoted here, as this would constitute the publishing of a nomen nudum. The locality Wahoo is in T. Nuttall’s writ- ing, then in another hand is added, Herb. Nuttall. The label is like Nuttall's other labels for plants that he collected on Wahoo (Oahu L, Hawaiian Is.). A confusion is suspected here, since Elodea is not indigenous to the Hawaiian Is- lands, is not established there now, nor does it ever seem to have been. The sheet bears three small, but genuine, branches of E. canadensis, but the locality stated seems to be in error. In 1836, Elodea canadensis was found introduced in Ire- land, and in 1842 in Scotland, and in 1847 in England. It spread to continental Europe and soon became abundant and a pest in the watercourses of much of the continent. As the plants were pistilate only, its spread was due to vegetative fragmentation. The species spreads rapidly by its own means, and man has doubtless been responsible for its arrival on distant continents or regions. It is now established in Europe, Asia, Africa, Australia, and New Zealand. From these 28 Rhodora [Vol. 67 areas it is abundantly represented in herbaria, but there seems no good reason for listing these records in this monograph. There is also a very extensive literature re- porting its occurrence as an exotic in the above named areas. Also, there are many articles on its anatomy, cy- tology, geneties, and physiology, but they are not pertinent here. J. K. Santos (1923, and 1924) published two studies upon the genus Elodea, issued as Contributions 302 and 317 of the Hull Botanical Laboratory. They are both largely cy- tological studies on the cell division in the formation of the pollen grains. However, being studies of the sexuality of Elodea plants, they touch in several places on the charac- ters used by taxonomists for the differentiation of the spe- cies of this genus. In the first article Santos deals particularly with E. gigantea which he inadvertently pub- lished as a new species, Elodea gigantea Santos. Evidently this is a synonym of the cultivated Egeria densa Planch. He states (1923: 44) that E. canadensis “is strictly dioe- cious.” His second paper deals principally with what he calls E. canadensis. His material came from Wolf Lake, Indiana. He again describes it as dioecious, saying (1924: 353), "both male and female plants are abundant." His papers do not record his means of determination of the species. By correspondence it is clear that Gray's Manual was the only source consulted. On page 85 of the 7th edition, this gives Elodea canadensis as the only species in northeastern North America. As synonymy it gives Anacharis Planch., and Philotria Britton. Britton & Brown's Illustrated Flora (ed. 1913) gives four species for the same area, and in the key leading to Philotria (Elodea) canadensis says ‘“‘staminate flowers unknown.” Also there are given references to the monographs by Planchon, Caspary, and Rydberg. Santos gives a bibliography of 36 titles in his first paper, and one of 53 in his second. In neither does he include any of these four monographs dealing with the classification of Elodea, or give any indication that he had consulted them. If he had, he could not have determined his plants all as E. canadensis. 1965] Elodea — St. John 29 No voucher specimens of the Elodea studied by Santos were preserved, which makes it more difficult to verify his determinations, but both E. canadensis and E. Nuttallii are known to occur in Wolf Lake, Indiana, the source of the specimens investigated by Santos. In his final discussion of the determination of sex, Santos says (1924: 368) any experimental studies should be accompanied with a critical cytological investigation. The effect of the environmental condition should not be looked upon from the external character of the individual only, but also in its effect upon the structure of the cell. One ap- parently may get some changes in form, but in reality the changes may be produced by changes in the chromatin." On this the writer would comment, any such experimental study should be preceded by a critical and accurate identifi- cation of the species studied. 20. Elodea linearis (Rydb.) comb. nov. Philotria linearis Rydb., Bull Torrey Bot. Club 35 (1908) 464; Anacharis linearis (Rydb.) Vietorin, Contrib, Lab. Bot., Univ. Montréal 18 (1931) 41. Fig. 3 d-f, 5. Diagnosis of Holotype: Submerged aquatic; stems slender, dicho- than 1 mm. wide, in whorls of 3, linear, acute, finely serrulate; tomously branched; middle and upper leaves 5-12 mm. long, less staminate spathes borne in the upper axils, about 1 cm. long, the lower half 3-5 mm. long, narrowed and peduncle-like, the upper inflated ovate or obovate, the apex like a gaping mouth, with 2 salient teeth 3-4 mm. long; staminate flowers peduncled by the slender, thread-like, elongating hypanthium; sepals 2.5-4 mm. long, 1.5-2 mm. broad, obovate, or oval, dark striate; petals 4.5 mm. long, 0.5 mm. broad, linear lanceolate, white and filmy; stamens 9 and equal, 2 mm. long, all attached at the summit of the hypanthium: anthers oblong; pistillate plants and flowers unknown. Holotype: Tennessee, swamps bordering on Cumberland River, vicinity of Nashville, A. Gattinger (NY). Type examined! Isotypes (DS, MO, NY, US). Discussion: Several of the characters mentioned by Dr. Rydberg in the original description do not hold after exami- nation of the holotype and the isotypes. Instead of being entire, the leaves are finely serrulate as in the other species of this genus. The peduncle-like base of the spathe reaches 5 mm. in length. The inflated apex of the spathe is ovate Fig. 3. a, Elodea Schweinitzii (Planch.) Casp., perfect flower, from holotype, X 5; b, leaf, X 5; c, leaf apex of same, X 50; d, Elodea linearis (Rydb.) St. John, staminate flower, from holotype X 5; e, leaf, X 5; f, leaf apex of same, X 50; g, Elodea longivaginata St. John, staminate flower, from C. S. Crandall 2,528, X 5; h, staminate spathe of same, X 5; i, pistillate flower of holotype, X 5; j, staminate bud, from Crandall 2,528; k, fruit, from Crandall & Cowen 6,720, X 1; l opened capsule and seeds of same, X 2; m, leaf, from C. S. Crandall, 2,423, X 5; m, leaf apex of same, X 50. 1965] Elodea — St. John 91 when young, but appears obovate when mature and opened. The sepals reach 3 or 4 mm. in length. The petals instead of being oblong, are clawed and narrowly linear lanceolate. The specimen from Center City, Minnesota, 1892, B. C. Taylor, the writer considers to be E. Nuttallii, and not the pistillate plant of E. linearis. Elodea linearis is a member of the section Elodea. In many ways it closely resembles staminate E. canadensis, but it differs in having the leaves less than 1 mm. wide, and in having the stamens equal in length and equally attached on the hypanthium; while the latter species has the leaves 1-4 mm. broad, and the 3 inner stamens raised on a joint stalk. Here is presented a copy of a note on the specimen, Tennessee, Nashville, Sept. 1875, A Gattinger (NY). This is apparently the same collection as that distributed with a printed label and made the type of P. linearis Rydb. It has several staminate flowers still in the spathes, but more mature than those described by Rydberg. The 3 central anthers are slightly the longer, but attached as shown in our illustration. The flower examined had only one petal, it being short and bearing a single anther sac containing pollen, attached along one edge of the petal. The following note is in a pocket on the sheet. It is in T. Morong's handwriting, but is evidently a copy of field observations, probably Gattinger's. “Anacharis Canadensis Planch. Male flowers apt. in the spathe, undeveloped. There seem to be about 9 stamens (as Wood says) which are contained in a hyaline sack, and this in a spathe; the spathe on a pedicel !4^ or 14’ long, opening in a bifid mouth, and suffering the stamens with their sack to escape, when mature. These stamens rise to the surface, on which they float, and there fertilize the pistillate flowers, which rise to the surface on a long and very slender tube. The pistillate flowers are also furnished with stamens, and these in the time of maturity seem to have an abundance of pollen, as I have often seen when collecting the plant in August." “Why Wood and Gray should both call these staminate flowers *minute', I do not understand, as the sack is at least [Vol. 67 Rhodora 32 "Huot “FS Cuoue[q) VMN “A ‘UYOF “Ig Wmbvarbu0] DIPO jo uonnqruüsip ƏY} Jo dew ^p 214 "ASEO (wouv[q) uzjmanyog * pue E EECHER ER TMIY32TTN FO ; gvpuwbembuo Fw / Y300137-. 33 John Elodea — St. 1965] Snow] AN pu "Huot "s (Copa) H "Su Ul YNY sisuoppuno Dopo]4g jo uonnqrujstip əy} jo dep ce “By F3 E] mur E? EE cuau Jo ; "P-ssuap.u?»'3e | Y3001377- 34 Rhodora [Vol. 67 1” long & 1” wide, & the spathe on a pedicel 3” or 4” lines long, fully as large as the fertile flowers.” “The staminate flowers seem to be without corolla, so far as I can make out. This does not appear to be the theory of the books, but from this specimen I judge it to be so.” These notes seem to contain partly material derived from the books by Wood, and by Gray, and partly actual field observations by Gattinger. Characteristics of more than one species are included. The sessile staminate flowers which are liberated and float to the surface are known only in E. Nuttallii. He recorded seeing perfect flowers, but none of his numerous specimens have any such, and no other species with bisexual flowers is known in the area. 21. (5 of part 1.) Elodea longivaginata St. John, Research Studies, Washington State Univ. 30 (1962) 38-40, fig. 2, e-j, 4, c-d, 5. Fig. 3 g-n, 4. Description of all specimens examined: Submerged aquatic, root- ing in the mud; stems elongate, slender, sparsely dichotomously branched; leaves opposite, linear, obtuse or acute, finely serrate near the tip, the middle and upper ones 5-24 mm. long, 0.5-2 mm. wide, bright green and flaccid, scarcely imbricate; apparently the lower leaves do not differ from the middle and upper ones, at least this is so down to the 21st. leafy node; staminate spathe 20-150 mm. long, ovoid, inflated near the tip, before opening with two small apical teeth, the inflated portion 8-10 mm. long, 2.5-4 mm. broad, finally with two large salient teeth at the open mouth-like apex; staminate flowers peduneled on the elongating, thread-like hypanthium which is up to 30 em. in length; sepals 3.5-5 mm. long, 1.9-2.5 mm. wide, elliptie, dark-striate especially near the tip; petals 5 mm. long, 0.6 mm. wide, white, delicate, linear; stamens 9, all attached at the summit of the hypanthium, 3.1 mm. long, linear-eylindric; pistillate spathe 30-70 mm. long, linear-cylindric, slightly larger near the tip, the apex with 2 salient teeth, the base split by the ripening capsule; pistillate flowers stalked by the thread-like base of the hypanthium which elongates even to several decimeters, bearing the flower to the surface of the water; sepals 2.8 mm. long, 1.8 mm. wide, elliptic, dark-striate especially near the tip; petals 4 mm. long, 1.8 mm. wide, white, delicate, spatulate; staminodia 3, strap-shaped; style slender equaling the hypanthium; stigmas 3, oblong, undivided, only half the length of the sepals; ovary commonly setting fruit and becoming large and conspicuous, ovoid; capsule 10 mm. long, 3 mm. in diameter; seeds 6 mm. long, cylindrie, surrounded by jelly, one attached on each parietal placenta, 6 attached basally. 1965] Elodea — St. John 35 CANADA. ALBERTA: in a lake at the northern edge of the Milk River Ridge, where the Lethbridge Trail leaves the summit, [holo- type], July 20, 1895, J. Macoun 13725 (CAN)! UNITED STATES. NoRTH DAKOTA: slough 7 miles e. of Bismarck, Metcalf 358 (US); Ward L., Kenmare, Burke Co., Metcalf 569 (cv). MONTANA: n. e. of Loring, Reservoir 158, Phillips Co., Hotehkiss 6348 (US). WYOMING: Seven Mile L., Albany Co., Gooding 597 (G, P); Teapot Dome, Reservoir, Casper, Porter 6259 (UPS); L. De Smet, Johnson Co., Porter 6285 (UPS); Two-Ocean L., Teton Co., Porter 6365 (UPS). COLORADO: Lee’s L., Fort Collins, 5,000 ft. alte Crandall & Cowen 6720 or Crandall 6720 (cs); Lee’s L., Crandall 2528 in part (WS), Crandall 2423 (B, CS, F, GH, LD, NY, P, PH, RM, US, WS); Fort Collins, Cowen (Mo, UC); Lee's L., ex Herb. State Agric. Col. 2421 (NY, PH, US, WS). UTAH: drift, s. end, Strawberry Reservoir, Wasatch Co., Hotchkiss 4933 (us). NEW Mexico: L. La Jara, Jicarilla, Apache Reservation near Dulce, Standley 8274 (us). Rocky Mts., 1868, G. Vasey (us). (To BE CONTINUED) CORRECTION In volume 65, p. 241 (1963) of Rhodora I reported Habenaria conopsea (L.) Benth. as having been collected by the late W. C. Coker in *Abisco," Labrador. Dr. A. E. Porsild has kindly called my attention to the geographical error of placing Abisko, Lapland in Labrador, and I have since learned from Dr. H. R. Totten that, in 1921, Dr. Coker was truly in Lapland. I must apologize for the bad labeling and my poor geography that resulted in this erroneous report. This species is now dubiously known from North America only in Connecticut. D. S. CORRELL, TEXAS RESEARCH FOUNDATION, RENNER, TEXAS A NEW SPECIES OF LAGENOPHORA (COMPOSITAE) FROM GUATEMALA' JoHN H. BEAMAN AND D. C. D. DE JONG Lagenophora cuchumatanica Beaman & De Jong, sp. nov. Herba perennis humilis cum 1-4 caulibus adscendentibus vel decum- bentibus usque ad 20 em longis, usque ad capitula foliosis vel cum pedunculis usque ad 6.5 em longis; folia basalia cum petiolis 0.7-3.3 em longis, lamina ovata-elliptica ad orbiculata, margine crenato- dentata, ciliata; folia caulina media oblanceolata vel spatulata, pauci- serrata; folia caulina superiora oblanceolata, sessilia, margine 1-4-serrata, folia suprema subulata, integra; capitula parva, termi- nalia, solitaria; involucra campanulata, 4.0-5.0 mm alta, 6.0-7.0 mm lata; phyllaria 3-seriata, herbacea, oblongo-lanceolata ad anguste- lanceolata, apice ciliato-fimbriata; flores radii 10-20, 1-seriata, ligula reflexa, viridi-alba, 2.0-2.2 mm longis, apice 2-dentata vel 3-denticulata; flores disci 8-14, corollae viridi-flavae cum lobis paten- tibus circa 0.7 mm longis; achaenia radii et disci obovata, 3.2-4.0 mm longa, 1.0-1.5 mm lata, glabra, brunnea, margine crassi-nervata, apice annulari glanduloso; pappus nullus. Low perennial herb from a short, erect or oblique rhizome with fibrous lateral roots; stems 1-4, simple, to 20 em long, rarely longer, ascending but more commonly decumbent, green to reddish, grooved, flattened or subterete, pilose with short to long, spreading to ap- pressed often purple-based multicellular hairs, leafy throughout or with peduncles gradually elongating in age to 6.5 em, rarely longer, the summits of the stems somewhat enlarged and densely pubescent below the heads; basal and cauline leaves pubescent with multicellular hairs, the margins ciliate and faintly revolute; basal leaves few (less than 10), the petioles 0.7-3.3 em long, densely pilose with spreading hairs, less so toward the sheathing base, the lamina ovate-elliptic to orbicular, 0.9-1.8 em long, 0.7-1.4 em wide, the margins crenate- dentate with 3-5 pairs of teeth with callous-tipped apices, pubescent below with appressed to spreading hairs, minutely white-dotted and glabrate to appressed-pubescent above with scattered hairs often in rows near and paralleling the margins; lower cauline leaves similar, smaller, soon shriveling; middle and upper cauline leaves sparsely appressed-pubescent below, glabrate above, minutely white-dotted only toward the apex; middle cauline leaves oblanceolate to spatulate, ‘Supported by grants G-9045 and G-23187 from the National Science Foundation. We are indebted to Dr, Mladen Kabalin, Michigan State University Science Librarian, for editing the Latin diagnosis. The U. S. National Herbarium loaned an isotype of Lagenophora andina Badillo for comparison with the new species. 36 1965] Lagenophora — Beaman and De Jong 37 serrate with a few callous-tipped teeth; upper cauline leaves oblan- ceolate with broad, sessile bases and margins with 1-4 small teeth, the uppermost subulate, entire; heads terminal, solitary, rarely more than 1 per stem, often subtended by a single phyllary-like bract; involucre campanulate, 4.0-5.0 mm high, 6.0-7.0 mm wide; phyllaries imbricated in about 3 series, distinct to the base, essentially herbaceous but with very narrow scarious margins and prominent midribs, often carinate near the base, shiny, glabrate to sparsely pilose on the back, the margins sparsely long-ciliate toward the base, the apices ciliate- fringed with often purple, sometimes sub-glandular hairs; outer phyllaries lance-oblong, 3.0-3.8 mm long, about 0.6-0.7 mm wide, minutely white-dotted near the obtuse apex, the middle phyllaries similar in outline and texture, 3.5-4.2 mm long, about 0.9 mm wide, the inner phyllaries thin, narrowly lanceolate, acute, about as long as the outer; receptacle 1.5-2.0 mm across, somewhat concave, naked, with whitish achenial attachment points; ray florets 10-20, pistillate, in one series, the tube ca. 0.1 mm long, its abaxial surface with short, thick glands, the ligule reflexed, 2.0-2.2 mm long, with 2-dentate or 3-denticulate apex, greenish-white, turning purple in age; disk florets 8-14, perfect, the limb greenish-yellow, campanulate, 1.9-2.4 mm long, 5-lobed with spreading, apically thickened lobes ca. 0.7 mm long, turning purple in age; style branches of the disk about 0.6 mm long, pubescent on the outside with short, blunt collecting hairs, glabrous within, the ovate stylar appendages about as long as the stigmatic lines; stamens with minute, blunt apical appendages and rounded anther bases; achenes of disk and ray similar, obovate, those toward the periphery increasingly oblique, compressed, with thick-nerved margins, the faces with minute ridges near the base, shiny, glabrous. brown (greenish-purple when immature), 3.2-4.0 mm long, 1.0-1.5 mm wide, the apices produced into a sticky glandular ring, this 0.4 mm high in the ray, 0.2 mm high in the disk; pappus absent. Plate 1309. GUATEMALA. HUEHUETENANGO: Sierra de los Cuchumatanes, between Chemal and Tojiah at Km 319.5 on Ruta Nacional 9 N, ca. 3,865 m alt, among low forbs in Pinus rudis forest, frequent in a local area, 29 July 1960, Beaman 3756 (Msc 172203 holotype; F, GH, K, TEX, UC, US, isotypes) ; same locality as type, 26 Aug 1961, De Jong 1145 (MSC); Sierra de los Cuchumatanes, between Km 322 and 323, Ruta Nacional 9 N, ca. 3,200 m alt, 27 Aug 1961, De Jong 1147 (MSC). TOTONICAPAN: on the Tecum Uman Ridge at Km 154 on Ruta Nacional No. 1, ca. 20 km east of Totonicapan, ca. 3,340 m alt, in pine forest, 14 Aug 1960, Beaman 4170 (MSC). The genus Lagenophora includes upwards of 20 species distributed from Malaysia and the south Asian coast south- ward around the Pacific rim through Australia, Tasmania, 38 Rhodora [Vol. 67 and New Zealand into southern South America, the Ant- arctic Islands, and north into the high mountains of Vene- zuela and Central America. Three species also have been described from Hawaii. Discovery of a species in Guatemala extends the Central American range of the genus northward from Panama. South Pacific distribution patterns of this type have been recognized since Hooker's (1853) classical account of the relationships of the flora of New Zealand. Several of the genera noted by Hooker (eg. Drimys, Colo- banthus, Acaena, and Oreomyrrhis) have ranges which closely parallel that of Lagenophora. Members of the South Pacific element such as Lageno- phora which range into tropical latitudes in the western hemisphere occur as isolated populations at high altitudes. In Central America their distribution always must have been discontinuous because the area has not been traversed by a high-mountain system in Cenozoic time (Schuchert, 1935). Long-distance dispersal seems to provide the most plausible explanation for the presence of Lagenophora in Guatemala (and probably also in Panama, Venezuela, and Hawaii). The sticky glandular achenial apices may have facilitated its dispersal. Although no data on the breeding system of Lagenophora are available, one might suspect self-compatibility on the basis of the inconspicuous heads and ray corollas (Plate 1309). If it is self-compatible, this could also be a key factor in its distribution. As Baker (1955) has noted, autogamy makes possible the establish- ment of a new population from a single propagule. A strong correlation between wide disjunction and self-compatibility has been demonstrated by Raven (1963) for amphitropical species. The distribution of other groups like Lagenophora which cross the tropies on high mountains similarly may be related to autogamy and long-distance dispersal. The two mountains upon which Lagenophora cuchuma- tanica has been found have different geological histories. The Sierra de los Cuchumatanes is made up chiefly of lime- stones which Schuchert (1935) suggests are of Cretaceous age. The Tecum Uman Ridge is composed of Tertiary rhyolitie lava domes (Williams, 1960). Both areas are of 1965] Lagenophora — Beaman and De Jong 39 P j Y t Plate 1309. Portion of the holotype of Lagenophora cuchumatanica. Insert, from left to right, shows outer phyllary, ray floret with reflexed ligule, and disk floret (X 8.3). 40 Rhodora [Vol. 67 considerably greater geological age and have much richer high-altitude floras than the neighboring Quaternary vol- canic cones. Although the geology differs, the habitats of the two populations are similar. Both localities are on nearly level terrain, in pine forests which are interspersed with large meadows (llanos), and have many other species in common. Slight morphological differences are evident between specimens from the Sierra de los Cuchumatanes and the single plant (Beaman 4170) from the Tecum Uman Ridge. The middle cauline leaves of the latter are conspicuously spatulate with 4-5 marginal teeth whereas the Cuchuma- tanes specimens have oblanceolate middle cauline leaves with 3-4 teeth. This species, Lagenophora cuchumatanica, is closely related to the Venezuelan L. andina Badillo. From Badillo's (1947) description and illustrations we thought that the two taxa might be conspecific, but an examination of isotype material of that species revealed several important differ- ences which are summarized in Table I. These two species also are related to L. panamensis Blake, but the latter has a heavier caudex, more elongate basal and cauline leaves, larger heads, and more numerous rays. As noted by Blake (1939) L. panamensis seems to be closer to the Hawaiian L. mauiensis than to the southern South American species which are scapose or nearly so. The Central and northern South American species seem to have more characters in common among themselves than with other members of the genus. The genus Lagenophora was included in the subtribe Bellidinae of the Astereae by Hoffmann (1894). In a re- evaluation of this group, De Jong (1964) concludes that it is artificial and suggests that its members be assigned to other subtribes of the Astereae. He considers Lagenophora, along with the related Myriactis, Rhynchospermum, and Solenogyne, to belong to the Grangeinae. DEPARTMENT OF BOTANY AND PLANT PATHOLOGY, MICHIGAN STATE UNIVERSITY, EAST LANSING DEPARTMENT OF BOTANY, MIAMI UNIVERSITY, OXFORD, OHIO. 1965] Lagenophora — Beaman and De Jong 41 LITERATURE CITED BADILLO, V. M. 1947. Una especie nueva del género Lagenophora en los Andes de Venezuela. Darwiniana 7: 331-332. BAKER, H. G. 1955. Self-compatibility and establishment after “long-distance” dispersal. Evolution 9: 347-348. BLAKE, S. F. 1939. Compositae in Contributions toward a flora of Panama, III, by R. E. Woopson, JR. and R. J. SEIBERT. Ann. Mo. Bot. Gard. 26: 265-324. DE Jone, D. C. D. 1964. The taxonomy of the genus Astranthium (Compositae-Astereae). Ph. D. thesis, Michigan State University Library. HOFFMANN, O. 1894. Compositae in Engler & Prantl, Natürl. Pfanzenfam. 45: 87-391. Hooker, J. D. 1853 (1852-1855). Flora novae-zelandiae. 2 vols. The botany of the Antarctic voyage. II. RAVEN, P. H. 1963. Amphitropical relationships in the floras of North and South America. Quart. Rev. Biol. 38: 151-177. SCHUCHERT, C. 1935. Historical geology of the Antillean-Caribbean region. John Wiley & Sons, Inc, New York. 811 p. WirLLiAMS, H. 1960. Volcanic history of the Guatemalan highlands. Univ. Calif. Publ. Geol. Sci. 38: 1-87. Table I. Characters which distinguish Lagenophora cuchumatanica from L. andina. L. cuchumatanica L. andina (Steyermark 57501, US) 1. Basal leaves pilose below, gla- 1. Basal leaves hirsute above and brous above except for pilose below. rows near the margins. 2. Basal leaves broadly ovate to 2. Basal leaves obovate, blades orbicular, blades 1.2-1.5 em 2.0-3.0 em long. long. 3. Cauline leaves, at least the 3. Cauline leaves oblanceolate to lower, spatulate. subspatulate. 4. Stems unbranched. 4. Stems with 1-2 lateral branch- es above. 5. Ray florets in 1 series, ca. 10- 5. Ray florets in 2 series, ca, 35- 20. 45. 6. Tube of the ray ca. 0.1 mm 6. Tube of the ray ca. 0.5 mm long. long. 7. Immature achenes (for com- 7. Immature achenes ca. 2 mm parison with L. andina) ca. 3 long. mm long. GENOTYPIC VARIATION IN THE PHACELIA HIRSUTA COMPLEX! GEORGE W. GILLETT The Phacelia hirsuta complex is here defined to include P. gilioides Brand as well as P. hirsuta Nutt., and inter- mediate populations. Field and laboratory studies of these were carried out in conjunction with a biosystematic in- vestigation of 10 species in the Cosmanthus group. The initial paper of this study (Gillett, in press) includes intro- ductory comments and basic literature citations. Both Phacelia hirsuta and P. gilioides are outcrossers and have 9 pairs of chromosomes. Experimental F, and F, hybrids have been produced. Each of these species is inter- sterile with the other 8 species of the Cosmanthus group. The P. hirsuta complex is separated from the other Cosman- thus phacelias by differences either in seed structure or in chromosome number (Gillett, op. cit.). The P. hirsuta complex has a distinct geographical status so that the combined genetical, cytological, morphological, and geo- graphical information portrays this complex as a distinct evolutionary line. The purpose of this paper is to show the variability of certain genotypic character differences between Phacelia hirsuta, and P. gilioides and through these more clearly to portray their relationship. These two species and their intergrades occur in an area extending nearly 700 miles north and south in the region between the eastern margin of the Great Plains and the Mississippi River. The more northern populations of central Missouri and southeastern Kansas are usually closer to the description of P. gilioides, while the southern populations usually fall within the circumscription of P. hirsuta. How- ever, many populations in Arkansas, eastern Oklahoma, and southern Missouri cannot readily be assigned to either species. 'Aided by a grant from the National Science Foundation. 42 1965] Phacelia hirsuta — Gillett 43 Phacelia gilioides is usually found on sparsely vegetated habitats in clearings of deciduous forests and frequently occurs on rocky barrens or limestone outcrops. On the other hand, P. hirsuta is usually found in more densely vegetated habitats in close association with grasses and weeds on stony to fine-textured soils of woodland clearings and roadsides. However, populations of the complex occur on à broad spectrum of habitats between these weakly- defined extremes. The two species differ by several flower characters in- cluding corolla shape and color. However, because of the desirability of using herbarium specimens for the study of variability, only those characters readily apparent in dried material were studied. Those selected were: 1) the margin of the corolla lobes; 2) the width of the sepals; and, 3) the orientation of the cauline hairs (spreading to appressed). These are illustrated in Fig. 1. A useful summary comparison of Phacelia gilioides and P. hirsuta is given in the following table: Phacelia gilioides Phacelia hirsuta Corolla tube rotate at anthesis. Corolla tube open-campanulate at Tube with distinct to faint purple anthesis. mark on each lateral vein, or Tube with distinct purple mark marks lacking. on each lateral vein, or these Petal margins fimbriate. merging across the mid-vein. Sepals from % to % mm broad. Petal margins entire. Appressed cauline hairs. Sepals from 34 to 2% mm broad. Spreading cauline hairs. The variability in each of the last three characters of the above table was determined either by direct measurement or by comparison. Voucher specimens were prepared from the parents employed in erosses and from one of the experi- mental F, hybrids and were used for scoring herbarium specimens. 'lThree classes of expression were recorded for each character and these were plotted on the distribution map of Fig. 2. The expressions classified for each character included the two extremes, each with a representation of near-extremes, and an intermediate generally comparable to the expression in F, hybrids, but including some near- 44 Rhodora [Vol. 67 Fig. 1. Corolla margins (a), sepals (b), and cauline pubescence (c), expressed in P. gilioides (Pı), Gillett 1260; P. hirsuta (P2), Gillett 1215; and experimental F; hybrid (center). Map symbols are below the corollas, to the right of the sepals, and above the stems. 1965] Phacelia hirsuta — Gillett 45 intermediates on either side of this central expression. In other words, all three characters are metrical with several degrees of expression for each. These three categories are represented in map (Fig. 2) symbols as follows: the Phacelia gilioides extreme by the open circle; the inter- mediate category, similar to the expression in F, hybrids, by a line 1 mm long; and the P. hirsuta extreme by a line 2 mm long. The left-hand ray portrays cauline pubescence, the vertical ray corolla margin, and the right-hand ray sepal width. The three recognized classes of expression permitted a crude graphical portrayal of the variability for a particu- lar character over the entire range of the complex. The scoring was accomplished by the procedures enumerated below. COROLLA LOBES: This character was scored by direct comparison. The fimbriate margins with well-defined teeth characterize one extreme. This expression occurs in central and southeastern Missouri. The intermediate expression is found from the northern limit of the complex to north- eastern Oklahoma and central Arkansas. The other extreme, the entire corolla margin, occurs in populations from south- western Missouri to Texas and Louisiana. SEPAL WIDTH: This character was measured by the ocular micrometer of a stereoscopic microscope. Normally, 11 sepals were measured on a given plant, these representing 5 mature flowers. The mean value was taken as the ex- pression of central tendency. The narrow extreme (sepals .5 -.75 mm broad) is closely restricted to central and south- eastern Missouri. However, the intermediate expression (sepals .75 - 1.00 mm broad) and the other extreme (sepals 1.0 - 2.5 mm broad) have remarkably wide distributions, each extending over almost the entire range of the complex. CAULINE PUBESCENCE: This character was scored by direct comparison. The pubescence was recorded from the central axis of the plant, just below the mature inflorescence. One extreme is characterized by relatively short hairs that are abruptly bent at the base and have an appressed-as- cending orientation with the stem. This expression is found in populations of central and southeastern Missouri, and in 46 Rhodora [Vol. 67 0 25 50 100 50km 0 25 ap 100 150 miles E— 32 & EXTREME FORM OF PHACELIA HIRSUTA A 30° EXTREME FORM OF PHACELIA GILIOIDES O VARIATION IN THE PHACELIA HIRSUTA COMPLEX SCORING SYMBOLS CAULINE PUBESCENCE COROLLA MARGIN SEPAL WIDTH (MM) APPRESSED-ASCENOING— O FIMBRIATE - O 5-75 o INTERMEDIATE —— — — Ò EROSE — Ò 75-100 Ó D ENTIRE —— b 100-25 o SPREADING Fig. 2. Map showing distribution of variation in the Phacelia hirsuta complex. 1965] Phacelia hirsuta — Gillett 4T isolated populations of eastern Oklahoma. The other ex- treme is typified by a pubescence of relatively long, spread- ing hairs and occurs in populations of south-central Missouri to Texas and Louisiana. However, the intermediate ex- pression of cauline pubescence occurs in populations of central and southwestern Missouri, over a narrower range than either of the extremes. The variation portrayed by this complex is suggestive of introgressive hybridization (Anderson, 1953; Stebbins, 1950), with the more northern P. gilioides and the southern P. hirsuta converging in southwestern Missouri and north- ern Arkansas, and subsequent bidirectional gene flow into both of the outbreeding parental stocks. The variation outlined above is suggestive of that found in the Phacelia sericea complex (Gillett, 1961). In both cases it is impossible to make clear-cut taxonomic dis- tinctions because of hybridization and the independent segregation of the genes that regulate character differences. The geographical variation in the P. hirsuta complex is so extensive that one feels obliged to recognize that the majori- ty of populations of either species carry some genes of the other. It seems necessary, therefore, to distinguish these species in terms of relative differences imposed by relative gene frequencies rather than to take refuge behind the time-worn cliché “separated only with great diffculty” which implies that the taxa involved are indeed quite sepa- rable, but only with remarkable intuition. In this material it seems preferable to follow the suggestion of Dobzhansky (1951) and emphasize the geography of the genes rather than to impose a taxonomic distinction based on arbitrary judgment or hypothetical statistical differences, both of which tend to conceal the problem rather than to elucidate it. DEPARTMENT OF BOTANY, UNIVERSITY OF HAWAII LITERATURE CITED ANDERSON, E. 1953. Introgressive hybridization. Biol. Rev. 28:280- 807. DoBzHANSKY, T. 1951. Genetics and the origin of species. New York. Columbia University Press. 48 Rhodora [Vol. 67 GILLETT, G. W. 1961. An experimental study of variation in the Phacelia sericea complex. Am. Jour. Bot. 48:1-7. In Press. Genetic barriers in the Cosmanthus phacelias (Hydrophyllaceae). Rhodora. STEBBINS, G. L. 1950. Variation and evolution in plants. New York. Columbia University Press. EULOPHIA ECRISTATA AND EPIDENDRUM CONOPSEUM IN MISSISSIPPI According to the latest authority available to me (Native Orchids of North America North of Mexico, Donovan Stewart Correll, 1950) the present known range of Eulophia ecristata (Fernald) Ames is extremely disjunct with sta- tions in North Carolina, Florida and Louisiana. I recall collecting it some fifteen years ago in Georgia. It was almost a certainty that sooner or later it would turn up in the intervening territory. This past summer while collect- ing in a low lying, grassy savanna within the western limits of the town of Picayune I found a small colony of three plants. Close associates were various species of Panicum and Paspalum while nearby were such genera as Stokesia, Eupatorium, Gerardia, Chondrophora, Solidago, and Helian- thus. The Eulophia was in bloom on Aug. 21st. A few days later I found a single specimen about eight miles to the northwest in open, grassy pineland. A specimen has been deposited at the University of Mississippi. Curiously enough almost the same pattern applies to the range of Epidendrum conopseum R. Br. This past winter I noticed its leaves about ten feet up on the trunk of Nyssa in a wooded overflow swamp two miles west of Picayune. About the middle of July, in company with Dr. Thomas M. Pullen, it was collected in flower. F. H. SARGENT PICAYUNE, MISSISSIPPI DETERMINATION OF THE DEVELOPMENTAL PATTERN OF ANGIOSPERMOUS LEAVES: EDWARD L. DAVIS In studies on the development of fruits in curcubits, Sin- nott (1936) has shown the consequence of differential growth rates in two dimensions on the ultimate fruit form. Where the relative rate of growth of width and length remains constant throughout maturation, but width is in- creasing more, or less, rapidly than length, there will be major differences between the form of small and large fruits. The resulting difference in ultimate form may serve to mask what is a high degree of genetic uniformity. Under these circumstances, a single variable, the duration of growth, can produce marked differences in appearance. Comparable conditions certainly operate in respect to leaf development (Hammond 1941, Stephens 1944, Whaley and Whaley 1942). In the taxonomic analysis of species popu- lations of angiospermous plants collected over a wide geo- graphical range, and under different conditions of photoperiod and growing season, there is a wealth of potential information to be obtained from the developmental pattern of leaves. Unfortunately, most methods of studying leaf development require extensive periodic measurements, or anatomical sections. In order to reduce the time required for accumulation of such growth information, we have been experimenting in our laboratory with 3-amino triazole as a rapid method of showing the developmental pattern of leaves. Three-amino triazole (3-AT) interferes with the synthesis of chlorophyll in leaves (Pyfrom et al. 1957, Jukes 1963) but not with pre-existing chlorophyll molecules, and as a consequence, marks newly developed parts of the leaf through the absence of chlorophyll. From the interpreta- tion of leaves caught at various stages of development at the time of application of 3-AT, the pattern of development — basipetal, acropetal or diffuse, — can readily be deter- "This research was supported, in part, by a grant from the National Science Foundation, NSF-GB1270. 49 50 Rhodora [Vol. 67 mined. Moreover, by treating some leaves early in develop- ment, and others at later stages, it is possible to see the way in which leaf shape is changing, much more readily than through a comparison of unmarked leaves. For many plants so far examined, an aqueous solution of 3-AT at concentrations of 0.13 grams per liter can be ap- plied by spray or dropper directly to the apex of the plant without significantly altering the leaf size or form. The developmental pattern for a leaf of tomato, Lycopersicon esculentum Mill. variety Marglobe is shown in Fig. la. The pattern of development, which is basipetal both for the entire compound leaf, and for each lobe, is shown by the development of chlorophyll-free areas within the leaf. In a more detailed treatment of tomato, Breil (1963), Briel and Davis (1964) has shown that the pattern illustrated by 3-AT fits the developmental pattern as determined from anatomical studies and mitotic counts. Daily measurement of growing leaves of plants of Tia- rella cordifolia L., from New England and North Carolina, show the maximum length to maximum width ratio to re- main almost constant at a value of 1 during the eight day period of principal leaf growth. During that period of growth, differences in leaf form, although minor, become accentuated. Tiarella plants from North Carolina show in- creased crenations on the leaf margin. Figure 1b shows a leaf in which 3-AT was applied early in development, and the effect on chlorophyll synthesis was wearing off in the final stages of leaf expansion. Consequently, chlorophyll appears only in the margin, which was the last part of the leaf to develop. Fig. 1c represents a drawing of a similar leaf, with the chlorophyll containing parts of the leaf in black. Tiarella plants from New England tend to stop expansion prior to this last phase of leaf growth, which occurs only in pockets at the margin of the leaf, and conse- quently show less pronounced crenations. Equally as important as determining where leaf form does change during enlargement is establishing leaf charac- teristics which are constant throughout development. Figure 1d shows a photograph of a much dissected leaf of Humulus PLATE 1310 Fig. 1. (a) Leaf of tomato treated with 3-AT; chlorophyll deficient areas at the base of lower lobes. (b) Treated leaf of Tiarella, chloro- phyll containing areas at leaf margin. (c) Diagram of treated leaf of Tiarella, black areas corresponding to chlorophyll containing zones. (d) Treated leaf of Humulus lupulus from Rocky Mountains; central and right lobes showing normal growth; lateral lobes on left developed assymetrically and later than usual as indicated by absence of chloro- phyll. 52 Rhodora [Vol. 67 lupulus L. from the Rocky Mountains, treated with 3-AT at the mid-point in its development. The developmental sequence of the hop leaf is shown to be basipetal with the lobes maturing first, followed by the “palm” of the leaf. The form of the leaf has taxonomic significance for several hop varieties have been distinguished by the relative size of the lobes of the leaf. Daily measurements of the length of the lobes of the expanding leaf during a thirteen day period, from the time the leaves are first large enough to measure to completed expansion, show the ratio of the central lobe to the adjacent lateral lobe to be nearly constant within the range of 1.1 to 1.2 (Fig. 2), and its taxonomic use is inde- pendent of leaf age. This same information can be obtained more readily by measuring treated leaves at different stages L6] 151 144 o EEN iol 91 B+ *——58——34——$——8———1-—— 1 DAYS Fig. 2. Ratios of the length of the central lobe to the length of the adjacent lateral lobe plotted on the graph are for a single hop leaf, although ten leaves were measured in the experiment with comparable results, since the ratios are compared to a paired leaf treated with 3-AT. (1) ratio of green tissue at the time of treatment (prior to 3-AT effect) and (2) ratio of chlorophyll deficient tissue at time of leaf collection. 1965] Angiospermous Leaves — Davis 58 of expansion and determining on each leaf the ratio of the chlorophyll containing zones of the central and adjacent lobes, and making the same determination for the chloro- phyll-deficient zones on these lobes. Where the expansion rate (as in hops) is uniform, calculations of the ratio of the tissues in each lobe formed before, or after, treatment for leaves at different stages of development give the same value. Fig. 2 also shows the ratios, determined by measuring green and colorless zones on a paired treated leaf at the time of treatment and at the time the leaf was collected 3 days later. A comparison of the ratio during expansion of the central lobe to the second lateral lobe shows à marked decline as the leaf enlarges, as would be predicted from the treated leaves (cf. Fig. 1d) which show the basal part of the leaf to develop much later than the apical central portion. Thus with 3-AT it is possible to determine very rapidly those ratios for the leaf which may be independent of the duration or extent of leaf expansion. It is not yet clear at what cellular stage 3-AT affects chlorophyll synthesis in developing leaves, Experiments are presently underway using tritiated thymidine to determine if only dividing cells, or both expanding and dividing cells are affected. Whatever the case, the use of an easily applied marker in leaf development, which permits a rapid recog- nition of newly expanding parts of the leaf blade, should make it possible to include more developmental information in taxonomic studies of plant populations. DEPARTMENT OF BOTANY, UNIVERSITY OF MASSACHUSETTS, AMHERST LITERATURE CITED BREIL, D. 1963. Pattern of leaf development in Tomato using 3- amino-1,2,4 triazole. M.A. Thesis Univ. of Mass. BREIL, D. A. and E. L. Davis, 1964. Morphogenetic effect of amino triazole on the developing leaves of Lycopersicum esculentum Mill., var. Bonny Best. HAMMOND, D. 1941. The expression of genes for leaf shape in Gossypium hirsutum L. and Gossypium arboreum L. Am. Jour. Bot. 28:124-150. 54 Rhodora [Vol. 67 JUKES, T. 1963. Some biological effects of Amino triazole. Ec. Bot. 17:238-240. PvrROM, H. T. APPLEMAN, and W. G. HkrM. 1957. Catalase and chlorophyll depression by 3-amino-1,2,4 triazole. Plant Physiol. 32:674-6'16. SiNNoTT, E. W. 1936. A developmental analysis of inherited shape differences in cucurbit fruits. Amer. Nat, 70:245-254. STEPHENS, S. G. 1944. The genetic organization of leaf shape de- velopment in the genus Gossypium. Jour. Genet. 46:28-51. WHALEY, W. G. and C. Y. WHALEY, 1942. A developmental analy- sis of inherited leaf patterns in Tropaeolum. Amer. Jour. Bot. 29:195-200. Errata for Rhodora Volume 66 . 218 line 33 for Fucus vesiculous read Fucus vesiculosus 223 in note at bottom, for Pantoneura barii read Pantoneura bearii 234 line 20 for P. plocamiodes read P. plocamioides 309 line 37 for Basia read Bahia 312 line 9 for Diosopyros read Diospyros . 358 Fig. 7 is Plate 1506 405 Fig. 1 is Plate 1507 PTTTUUTUT NATURAL AND ARTIFICIAL HYBRIDS OF CACALIA ATRIPLICIFOLIA AND C. MUHLENBERGII JAMES R. COLEMAN In the summer of 1961, the author undertook, at the suggestion of Dr. Charles B. Heiser, Jr., an investigation of those species of Cacalia found in Indiana. This involves four species: C. atriplicifolia L., C. muhlenbergii (Sch. Bip.) Fern., C. suaveolens L. and C. tuberosa Nutt. The two species Cacalia atriplicifolia and C. muhlenbergu are morphologically rather similar, differing primarily in the following characters: C. atriplicif olia C. muhlenbergit . Stems glaucous 1. Stems not glaucous . Stems terete 2. Stems grooved and angular . Leaves white beneath 3. Leaves green beneath 4 5 A wD ka . Basal leaves triangular-ovate . Basal leaves reniform 5. Leaves essentially glabrous . Leaves pubescent Whereas Cacalia muhlenbergii occurs primarily in rich, shaded woods C. atriplicifolia occurs primarily in dry, open woods. The author found one locality (Cedar Bluff, Monroe Co., Ind.) at which populations of the two species were growing within 200 feet of one another. The area is a rich woods penetrated by an abandoned road. Cacalia muhlenbergii was growing primarily in the shaded areas to the sides of the road, and C. atriplicifolia was growing in the clearing of the road. One plant, occurring at the edge of the C. atriplici- folia population toward the C. muhlenbergii population, showed intermediacy for the characters separating the species. However, the stalk was broken off before sexual maturity was reached thereby preventing an examination of meiosis and pollen stainability. The rootstock of the putative hybrid was transferred into the greenhouse at Indiana University as were mature plants of the parental species. Crosses were made between Cacalia atriplicifolia and C. muhlenbergii with the former as the female parent. Of the 165 florets crossed, 7.5% developed seeds. 55 56 Rhodora [Vol. 67 The putative hybrid seeds were planted in the spring of 1962 and placed out-of-doors for a two week cold treatment. They were then transferred to the greenhouse where they germinated and appeared healthy but did not produce flowering stalks. 'The putative natural hybrid behaved similarly. The plants remained in the greenhouse until the fall of 1963 without flowering. In the fall of 1963 the putative natural hybrid and two putative artificial hybrids were transferred to the experi- mental garden at Indiana University. All three flowered in the summer of 1964. Pollen stainability with cotton blue (based on 200 grains) was 2.0% for the natural hybrid and 3.0% and 9.5% for the artificial hybrids. The lack of seed- set observed in these hybrids accounts for the failure to detect hybridization in the Cedar Bluff populations. An examination of meiosis, using acetocarmine squash methods, revealed both bivalents and univalents (Fig. 1). However, since the bivalents have a tendency to dissociate even in the parental species, the exact number of true uni- valents could not be accurately determined. Laggards were observed at anaphase (Fig. 2) as were “tetrads’ with micronuclei. The chromosome count of Cacalia muhlenbergii has been reported as 2n—50 (Afzelius in Darlington and Wylie, 1955), of C. atriplicifolia as n—25 (Jackson, 1962) and C. suaveolens as 2n—40 (Afzelius in Darlington and Wylie, 1955). Counts by the author confirm these reports. An examination of C. tuberosa revealed n=27 (Fig. 3) (voucher: Cabin Creek Bog, ca. 6 mi. n. of Modoc, Ran- dolph Co., Ind. Coleman 101 IND). Other numbers reported for the genus are 2n—52 (2 species) and 2n—60 (4 species) (Darlington and Wylie, 1955 and Takeshita, 1961). DEPARTMENT OF BOTANY, INDIANA UNIVERSITY, BLOOMINGTON Present address, INSTITUTO DE BOTANICA, SAO PAULO, BRAZIL Fig. 1. Diakinesis of natural hybrid showing bivalents and uni- valents. Fig. 2. Anaphase 1 of artificial hybrid showing laggards. Fig. 3. Diakinesis of Cacalia tuberosa, n=27. 1965] C mn m D LIU 1 acalia Hybrids — Col eman 57 e eo et iS = Oe e~" e x É m É- Se e d S 58 Rhodora [Vol. 67 LITERATURE CITED DARLINGTON, C. D. and A. P. WYLIE. 1955. Chromosome atlas of flowering plants. George Allen and Unwin Ltd. London. Jackson, R. C. 1962. In. Documented chromosome numbers of plants. Madrofio 16:268. TAKEsHITA, M. 1961. Cytological studies on Cacalia and its related genera I. The chromosome number of 3 species and one variety of Cacalia and one species of Miricalia. Jap. Journ. Genet. 36:(516) :217-220. NOTES ON VARIETIES OF BIDENS CONNATA AND A HYBRID WITH B. CERNUA EARL EDWARD SHERFF Some years ago I cited (Brittonia 11:190. 1959) Bidens connata, var. gracilipes Fern. as growing on the east shore of Eagle Lake, Kalamazoo County, Michigan. Later, I listed various additional localities for the same variety (Bidens connata var. gracilipes Fern. in Western Michigan, Rhodora 63:145. 1961). Still later (ibid. 64:23-28. 1962), under the caption, *Further notes on the distribution of Bidens connata vars. pinnata and gracilipes," I discussed the two varieties, giving an extended list of exsiccatae of var. pinnata for Wisconsin (pp. 24 & 25), also a list of ad- ditional ones of var. gracilipes for Western Michigan (pp. 27 & 28) and Wisconsin (p. 26). Recently, through the kindness of Dr. Gerald B. Ownbey, Professor of Botany and Curator of the Herbarium at the University of Minnesota, I was permitted to examine the entire collection of Minnesota specimens of Bidens in that herbarium. A careful search was made for the possible occurrence of var. gracilipes in Minnesota. Only five col- lections of the variety were found, all from the north or eastern part of the state: Olga Lakela 1860, (“leaves on mainstem divided" fide Lakela), Oatka Beach Addition, Bay Front, Duluth, Minnesota, Sept. 15, 1936 (MIN) ; Lakela 6202, St. Louis River estuary at New Duluth, Minn., Sept. 9, 1945 (MIN) ; Lakela 19270, shore swamp of Black Bay of Rainy Lake, Koochiching County, Minn., Aug. 16, 1955 (MIN); Carl O. Rosendahl 6503, shores of Deming Lake, Itasca Park, Minn., Sept. 8, 1933 (MIN); J. H. Sandberg, wet places, Hennepin County, Minn., August, 1889 (MIN). So far as known at present, Bidens connata var. pinnata is found growing only in Minnesota and Wisconsin, as pre- viously stated by me (N. Amer. Fl. ser. II: pt. 2: 91. 1955). To the list of exsiccatae of Bidens connata var. pinnata cited for Wisconsin, may be added the following found in the Herbarium of the University of Minnesota: all from localities in Minnesota: 59 60 Rhodora [Vol. 67 F. K. Butters & C. O. Rosendahl 6555, annual, shores of island, Form Island Lake, Aitkin County, Minn., Sept. 6, 1934; C. L. Herrick, Minneapolis, Minn., July, 1878; Olga Lakela 1858, in shallow water on bog shore, Oatka Beach Addition, Duluth, Minn., Sept. 7, 1936 ("leaves pinnate on the main stem"); Lakela 4869, roadside swamp south of Cloquet, Carlton County, Minn., Sept. 5, 1941; Lakela 5905, in shallow pond on Minnesota Point, Duluth, Minn., Sept. 10, 1944; Lakela 6231, swamp along St. Louis River at New Duluth, Minn., Sept. 15, 1945 (2 sheets examined); Bruce R. Ledin, edge of Hill Point Bog across from Schoolcraft Island, Itasca Park, Minn., Aug. 25, 1938; John W. Moore 24920, growing along the north shore of Loon Lake, Blue Earth County, Minn., Sept. 30, 1959; C. O. Rosen- dahl 6134, in dried-up pond near Pine Tree Lake, Washington County, Minn., Sept. 17, 1929. During the examination of Minnesota specimens of Bidens connata and its known varieties, however, two ad- ditional sheets of material were discovered, one of them bearing an obviously new variety of B. connata and the other bearing what I construed to be a hitherto undescribed hybrid. The new variety is set forth here: Bidens connata var. saint-vincentii var. nov. — Herba annua +4 dm. alta, glabra, pallida, suberecte ramosa, ramis tenuibus angulatis striatisque tantum circ. 1 mm. crassis. Folia principalia (pro unica planta visa) tripartita, petiolata petiolo tenui 2.5-3 em. longo, foliolis membranaceis ovatis ovato-lanceolatisve apice acutis basi plus minusve petiolulatis marginibus subintegris saepius acriter paucidentatis lateralibus 1-2.5 cm. longis terminali (petiolulo excluso) 1.5-4.5 cm. longo; alia folia saepius simplicia, minora, interdum integra. Involucri bracteae exteriores 4-8, lineari-spathulatae, adscendentes, usque ad circ. 3.5 em. longae et 5 mm. latae, apice acutae, marginibus pauciter antrorso-spinulosae alibi glaberrimae. Achaenia biaristata, corpore atra vel subatra, 7-11.5 mm. longa et 1.5-3.5 mm. lata, plana vel valde obcompressa, anguste lineari-cuneata, faciebus striata et remote subremoteve erecte adpresseque setulosa, marginibus de apice usque ad basim erecte setosa; aristis circ. 2.5-4 mm. longis suberectis, acriter retrorso-hamosis. Annual herb +4 dm. tall, glabrous, pale-green, suberectly branched, the branches delicate, angulate and striate, only about 1 mm. thick. Principal leaves (seen only for a single plant) tripartite, petioled with a slender petiole 2.5-3 cm. long; the leaflets membranaceous, ovate or ovate-lanceolate, apically acute, at base more or less petiolu- late, the margins subentire more often sharply few-dentate, lateral ones 1-2.5 cm. long, terminal one (petiolule excluded) 1.5-4.5 cm. long; secondary leaves more often simple, smaller, at times entire. Outer involucral bracts (phyllaries) 4-8, linear-spatulate, ascending, 1965] Bidens — Sherff 61 up to about 3.5 cm. long and 5 mm. wide, apically acute, at margins scantily antrorse-spinulose elsewhere very glabrous. Achenes biari- state, their body black or blackish, 7-11.5 mm. long and 1.5-3.5 mm. wide, flat or strongly obcompressed, narrowly linear-cuneate, on the faces striate and remotely or subremotely erectly and appressedly setulose, edges erectly setose all the way from apex to base; aristae about 2.5-4 mm. long, suberect, sharply and retrorsely barbed. Holotype: John W. Moore & John H. McAndrews 24810, growing on the river bank at St. Vincent, Kittson County, northwesternmost Minnesota, Sept. 17, 1959 (MIN., herb. no. 574691). The general habit of the foliage and fruiting heads, also the striate achenial surfaces, place this plant with the Bidens connata Muhl. complex. Indeed, the collectors had determined the type as “Bidens connata Muhl.” In my previously published key to the varieties of Bidens connata (North Amer. Flora ser. II: pt 2: 89. 1955), var. saint-vincentii will deserve a special line as a fourth line of the key, to read: Achenial aristae normally 2 var. saint-vincentit. The type, a moderately small specimen, doubtless selected in the field for ease of preserving, probably is somewhat atypic as to foliage characters of large, well developed plants. More specimens should be studied in the type locality. In the entire species, Bidens connata, consisting as it does of eight hitherto-known varieties (including the var. con- nata), no achenes have been known to be as here, constantly biaristate. The usual number of aristae is four, though in var. submutica Fassett they may be “none or rarely one or two." The above-mentioned undescribed hybrid may be set forth as: A putative hybrid of Bidens cernua L. and B. connata Muhl. Under her no. 2356, Miss Olga Lakela distributed as Bidens cernua L. a remarkably slender-leaved annual herb which she had collected at Duluth, Minnesota: Lakela 2356, in shallow water on Oatka Beach Addition, Duluth, Minne- sota, “leaves very narrow, bracts long," Sept. 6, 1937 (MIN). The plant examined by me is on herbarium sheet no. 338394. It is of slender, erect habit with slender sub- 62 Rhodora [Vol. 67 erect branches; the leaves are all simple, exceedingly nar- row (even graminiform) and 4-12 em. long by 1.5-7 mm. wide, very long-attenuate at both ends, the tip very acute, surfaces glabrate, margins entire or each --2-5-dentulate with slender, small and at times inflexed teeth, otherwise glabrate except for almost microscopic spinulae. The heads are radiate, about 2.5-3 cm. wide when expanded at anthesis, the disk (excluding the outer phyllaries) about 1.5 cm. wide; ray florets about 1.2-1.4 cm. long, ray yellow, linear- oblong and about 3-4 mm. wide. Outer phyllaries about 8, linear or linear-spatulate, +1.5 cm. long, about 3- or 5- striate, glabrous except for minutely spinulose edges. The achenes are of the type found on Bidens cernua but are grossly undeveloped, almost abortive, seemingly sug- gestive of hybridity. As the general aspect of the plant is unlike that of any Bidens cernua material ever heretofore seen by me, and as specimens of B. connata var. pinnata Wats. and B. connata var. gracilipes Fern. were collected by the same collector in the same small locality just a year earlier (vide Lakela 1858 and 1860, supra), I have con- cluded Lakela 2356 to be designated best as a putative hybrid between B. cernua L. and some variety, perhaps var. gracilipes Fern. or var. pinnata Wats., of B, connata Muhl. Years ago I wrote (The Genus Bidens. Bot. Ser. Field Mus. Nat. Hist. 16: 262, footnote 1. 1937): Bidens connata ap- pears to hybridize very rarely in nature. I have seen two sheets of material (Herb. W. C. Ferguson) collected by W. C. Ferguson, Plattsdale, Long Island, New York, Sept. 20, 1919, which appeared clearly to be of hybrid origin — B. connata X B. cernua. The leaves were tripartite as in typical B. connata; the heads were radiate and much like those of B. cernua. CHICAGO NATURAL HISTORY MUSEUM, CHICAGO 5, ILLINOIS A REVISION OF THE NORTH AMERICAN SPECIES OF HELIANTHEMUM (CISTACEAE) H S. DAOUD AND ROBERT L. WILBUR* The genus Helianthemum s.l. is composed of approxi- mately one hundred species found in both the New and the Old World. The species are particularly abundant about the Mediterranean Sea and in North America. The American species, as well as those of the rest of the world, were most recently revised in their entirety by Grosser (Engler's Pflanzenreich 14. (IV. 193.) 1903). Several regional treat- ments for the North American species have been since published but no comprehensive account of the genus in North America has appeared, Since Grosser's monograph, several additional American species have been proposed. The generic dismemberment of Helianthemum has con- tinued to be a source of considerable disagreement. This study, however, was not intended to evaluate the generic relationship of the American species with those of the Old World but to review the North American representatives. The troublesome question of generic limits deserves careful reappraisal but ought to be undertaken where collections of the entire generic complex are readily available. Twenty North American species of Helianthemum are recognized in this account. Our geographical area ‘North America’ includes the West Indies and Central America. This treatment therefore includes all the species in the New World except the three known from South America. These were excluded primarily because too little material was readily available to make their study feasible. One species, H. brasiliense (Lam.) Pers., is known from the eastern side of that continent, *The present study is based largely on the dissertation which the senior author presented to the Graduate School of Duke University, in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Duke University. The junior author’s role in this investigation was made possible by NSF Grant 18799 for which grateful acknowledgment is made. 63 64 Rhodora [Vol. 67 occurring in Brazil, Uruguay and Argentina, while two others, H. hirsutissimum Presl and H. spartioides Presl, are found in Chile. The last mentioned species is of particu- lar interest since its relationship is reported to be with the Californian H. scoparium Nutt. HISTORY OF THE GENUS IN NORTH AMERICA Although the names Helianthemum and Cistus were in use earlier, Tournefort (Inst. Rei Herb. 1700.) recognized two genera on the basis of their capsule differences: Helian- themum having but one locule and Cistus with 5(6-10) locules. Linnaeus (Sp. Pl. 523-529. 1753 and Gen. PI. 234. 1754.) did not accept Tournefort's Helianthemum. Instead he combined it with Cistus, which he placed in the first order (Monogynia) of his thirteenth class (Polyandria). He recognized 28 species of Cistus including one New World species, Cistus canadensis, Two years earlier, Linnaeus described Lechea major which he later included in Species Plantarum in the class and order Triandria Monogynia. Blake (Rhodora 20: 49-50. 1918), found the type of Lechea major to be conspecific with Cistus canadensis. Nomenclaturally, the genus is attributed to Miller (Gard. Dict. Arb. 1754.), whose Fourth Edition, although pre- Linnean in character, is still an acceptable source of generic names. Rafinesque (Car. Nuov. Gen. Sp. Sicil. 73. 1810.) pro- posed the generic name Xolantha to replace Helianthemum since the latter could be confused with Helianthus (Compo- sitae). Later Rafinesque (Chlor. Aetn. 9. 1815.) proposed the generic name Anthelis as another substitute for Helian- themum. Dunal (DC. Prodr. 1: 266-284, 1824.) recognized 112 species of Helianthemum and divided the genus into three "series" and nine sections primarily on the bases of differ- ences of stigma, style and stamens. Dunal's section Lecheoides contained only the seven species of Helianthemum adequately known to him from the New World (H. corymbosum, H. rosmarinifolium, H. 1965] Helianthemum — Daoud & Wilbur 65 glomeratum, H. ramuliflorum, H. canadense, H. brasiliense, and H. carolinianum). It would seem that the American occurrence of these species and their alternate cauline leaves were their principal claim to separate sectional ranking. Sweet (Cistineae. 1825-30.) adopted Dunal's classification and listed four more species with the seven American species which Dunal had assigned to his section Lecheoides. Three of these (H. obcordatum, H. tripetalum and H. asty- lum) Dunal previously had listed as unsatisfactorily known to him and they were known no better by Sweet. The fourth species, H. polygalaefolium Sweet, is generally considered a synonym of H. brasiliense. Sweet indicated that several New World species produced petaliferous flowers in the summer and apetalous flowers in the autumn, a feature he expected was characteristic of all American species. Spach (Ann. Sci. Nat. 2nd ser. 6: 357-375. 1836.) recog- nized sixteen genera in the Cistaceae. The known species of Helianthemum s.l. were placed in eight of these genera. Five of these eight genera were restricted to the Old World. The other three, Crocanthemum, Heteromeris and Taenios- tema, all newly described, occurred only in the New World. The primary points in Spach's account of the New World genera are as follows: Crocanthemum: flowers all petaliferous. (C. carolinianum (Walt.) Spach and C. brasiliense (Lam.) Spach were both definitely included and Helianthemum polygalae- folium Sweet was indicated as a possible addition.) Heteromeris: both petaliferous and apetalous flowers present. (Included in this genus were taxa which are today known as Helianthemum glomeratum, H. cana- dense, H. corymbosum and H. rosmarinifolium but which were in several cases provided by Spach with new epithets under Heteromeris.) Taeniostema: flowers all apetalous. (The only species included was T. micranthum to which he appended Helianthemum glomeratum as a questionable syno- nym.) A fundamental weakness of his last two genera was the 66 Rhodora [Vol. 67 inclusion of different floral states of H. glomeratwm within both taxa. Most American authors who accept generic status for the American species have taken up the generic name Crocanthemum. Although this genus was originally characterized by the constant petaliferous condition of its flowers, a feature not strictly true of either of the species originally included and certainly correct for only a minority of the species included by those who have accepted the genus, this in itself would not bar the employment of the name for the American segregate. The subgenus Horanthes was proposed by Rafinesque (New Fl. N. Am. 3: 30 + 31. 1838.) as a substitute for Dunal’s section Lecheoides since, in his opinion, the latter too closely resembled the generic name Lechea. Later in the same year he elevated (Sylva Tell. 132 + 133. 1838.) Horanthes to generic status along with four other segre- gates from the Old World: Stegitris, Fumana, Xolanthes (=Xolantha), and Psistina. Willkomm (Icon. et Descr. Pl. 2: 8-12. 1856.) proposed a new classification for Cistaceae dividing it into eleven genera. He accepted Spach’s genera, Crocanthemum, Hete- romeris and Taeniostema, to which both workers had as- signed all the known New World species. Crocanthemum and five Old World genera comprised his Tribe Normales possessing only chasmogamous flowers while Heteromeris and Taeniostemma (—Taeniostema) were placed in his Tribe Abnormes since they possessed cleistogamous flowers. Bentham (Gen. Pl. 1: 113 + 114. 1862.) maintained the genus in the broader sense, indicating that the characters used in separating the species of Helianthemum into sup- posed genera by Spach and others were minor and in- constant. The Synoptical Flora (Gray’s Syn. Fl. 1(1): 189-191. 1895.) accepted nine species of Helianthemum in the United States which were arranged in three groups. The first group contained the “Atlantic species, with dimorphous flowers” (H. canadense, H. majus (=H. Bicknellii), H. capitatum (=H. rosmarinifolium), and H. corymbosum). The second included the “S. Atlantic species with homomor- 1965] Helianthemum — Daoud & Wilbur 67 phous flowers" (H. arenicola, H. Nashii, and H. carolinia- num). Barnhart (Bull. Torrey Club 27: 589-592. 1900.) pointed out that H. arenicola and H. Nashii were apparently placed among the species with homomorphous flowers on the basis of their original description since both possess cleistogamous as well as petaliferous flowers. Gray's third group consisted of the “Pacific species: flowers homomor- phous" (H. scoparium and H. Greenei). Reiche (Nat. Pflanzenfam. III. 6: 304-306. 1895.), em- ploying Helianthemum in the broad sense, treated all of Dunal's sections as subgenera except Macularia which was omitted. All the New World species were placed in the subgenus Lecheoides (Dunal) Reiche. Grosser (Pflanzenreich 14 (IV. 193.) 1-161. 1903.) assigned all known species of Helianthemum s.l. to three genera: Halimium, Tuberaria, and Helianthemum. Tuber- aria and Helianthemum, both confined to the Old World, supposedly resemble one another in possessing either no style or one that is bent or curved and often elongate as well, somewhat swollen funiculi and only petaliferous (chasmogamous) flowers. Halimium (Dun.) Wilk., found according to Grosser’s concept both in the Old and New World, was reported to differ from them in its short, straight style, filiform funiculi and often apetalous (cleis- togamous) flowers in addition to the petaliferous. There were also differences reported for the embryos of the three genera. Grosser recognized three sections in Halimium. Section Spartioides, containing the Californian endemics, H. scopa- rium and H. occidentale (=Helianthemum Greenei), and the Chilean H. spartioides, was characterized primarily by its supposed broom-like habit. The Old World section Euhalimium was distinguished principally by possessing only chasmogamous flowers while the American section Lecheoides often has cleistogamous as well as chasmoga- mous flowers. Britton (Ill. Fl. N. U.S. 2: 539-541. 1913.) transferred the three species of Helianthemum then recognized within the range of the Illustrated Flora to Spach's Crocanthemum. In the same year Bicknell (Bull. Torrey Club 40: 613-616. 68 Rhodora [Vol. 67 1913.) accepted this generic segregate. However, Fernald (Rhodora 19: 58-60. 1917.) rejected the idea of separating the New World species into the endemic, segregate genus Crocanthemum. Ponzo (Nuovo Gior. Bot. Ital. 28: 157-173. 1921.) basing his classification on anatomical and morphological features, included the seven New World species known to him in Spach’s genus Heteromeris. Standley (Contr. U.S. Nat. Herb. 23: 832-834. 1923.), following Grosser, treated all nine Mexican species of Helianthemum known to him in the genus Halimium. Janchen (Österr. Bot. Zeitschr. 71: 266-270. 1922.) followed Britton and Bicknell in accepting the segregation of the New World species in Spach's genus Crocanthemum. He indieated that these species have alternate, pinnately nerved leaves while the Old World species of Halimium have opposite, three nerved leaves. Janchen (Engler & Prantl’s Nat. Pflanzenfam. 2 Aufl. 21: 289-313. 1925.) included all the species of Helianthemum (sensu lat.) in four genera: the New World Crocanthemum and three genera from the Old World. The species of Crocanthemum were aligned in two sections, Spartioides and Lecheoides, which were pre- viously recognized by Grosser (Engler & Prantl’s Nat. Pflanzenfam. 14. IV. 193. 33. 1903.) as sections of Hali- mium. Barnhart (in Smal's Man. Se. Fl. 878-880. 1933.) followed Britton, Bicknell, and Janchen in accepting Crocanthemum. He also arranged the nine species genus into three informally designated groups. Schreiber (Madrono 5: 81-85. 1939.) studied the genus Helianthemum in California, accepting the two previously recognized species and three varieties, and proposed a new species, H. suffrutescens. Fernald (Rhodora 43: 609-616. 1941.) for the second time questioned the status of the segregate genus Crocan- themum and the way it was applied to the species of the New World. He indicated that the characteristics proposed by various authors as a basis for the segregation of the New World species into a single genus were inconstant. 1965] Helianthemum — Daoud & Wilbur 69 MORPHOLOGY The HABIT of all American species is perennial. The genus contains herbaceous, suffruticose and _ fruticose growth forms. The ten species found in the eastern half of the United States, Canada, and the West Indies are typically herbaceous while the two Californian species are suffruti- cose. The Baja Californian species, H. nutans, is definitely shrubby although low and divergent. Three Mexican and Central American species (H. Pringlei, H. Coulteri and H. patens) may be either herbaceous or suffruticose, while the remaining Mexican and Central American species are all suffruticose. The habit of the species is of minimal value in identification. PUBESCENCE varies greatly in both quality and quantity on different parts of the plant. All the American species are typically pubescent although occasional specimens of H. patens are completely glabrous. The basic types of pubescence found among the North American species are (1) stellate hairs, the predominant type varying greatly in both length and density; (2) simple hairs, longer than the stellate hairs but less abundant and usually intermixed with them; (3) glandular hairs, very rare but typically found on the calyces of H. dumosum and always on the inflorescence as well as the upper part of the stem in H. Greenei and occasionally on the calyces of H. carolinianum, H. Coulteri and H. canadense. They are conspicuous only in H. Greene. Aside from habit and pubescence, few features of the STEM are taxonomically useful. Usually the stem is pubes- cent although in some species it is glabrate, or in a few specimens of H. patens completely glabrous. Among the species of the southeastern United States, H. carolinianum alone has stems sparsely covered with elongate, hyaline stellate pubescence. Among the Mexican species, the stems of H. chihuahuense and sometimes those of H. Pringlei are villous. Helianthemum propinquum characteristically arises from a creeping subterranean rootstock, and this dis- tinguishes it from the closely related H. Bicknellii which like most American species possesses a woody caudex. The LEAVES are simple, pinnately but often obscurely 70 Rhodora [Vol. 67 veined, estipulate, and alternate. STIPULES have been reported in certain species (e.g. Crocanthemum stipulatum Janch.) but these are the result of a mistaken interpretation of fasciculate axillary leaves. A few species possess basal leaves in rosettes or crowded at the base of the plant but these differ in no essential way from the lower cauline leaves. These have been reported as opposite but actually are alternate and separated by very short internodes. The presence of basal leaves is often characteristic of H. caroli- nianum and H. georgianum. Occasionally, however, H. rosmarinifolium or a few other species also have basal leaves. PETIOLES are present in a majority of the species. A few species may be either epetiolate or with a petiole up to 3 mm. long. Sometimes no definite distinction can be made between the blade and petiole. This renders the presence or absence of the petiole of negligible taxonomic value. The shape of the BLADES varies from linear to broadly obovate. Although the shape is characteristic for some species, it varies greatly in others. Also there is the ex- pected gradual decrease in size and change in shape of the lamina from the base of the plant upward. The shape of the leaf-blade, therefore, is of limited aid in distinguishing the species and has been employed only as a supplementary character. The PUBESCENCE of the leaves is essentially of two types: (1) the predominant stellate hairs which vary greatly in both length and quantity, and (2) the relatively uncommon simple hairs which are longer, less abundant and usually intermixed with the stellate type. (Simple foliar trichomes occur on the upper leaf surface of H. canadense, H. dumo- sum, and sometimes H. Coulteri, both surfaces of H. chihua- huense and sometimes also on H. Pringlei, and only on the leaf margin of H. nutans). The leaves of all species ex- amined are typically pubescent, but those of H. scoparium and H. Greenei sometimes are glabrate and occasionally those of H. patens are completely glabrous. The pubescence on both surfaces of the leaves is similar in quantity and quality in some species (H. Greenei, H. scoparium, H. Pringlei, H. chihuahuense, H. patens, H. 1965] Helianthemum — Daoud & Wilbur 71 argenteum, and H. nutans) and in these species the second- ary veins are not elevated below. In other species (H. canadense, H. dumosum, H. Bicknellii, H. propinquum, H. carolinianum, H. rosmarinifolium, H. georgianum, H. co- rymbosum, H. concolor, and usually H. Coulteri) the quality and the quantity of the pubescence are remarkably dis- similar on both surfaces of the leaves, and in these the secondary veins are prominently elevated beneath. Both leaf surfaces of H. glomeratum possess similar pubescence but its venation varies from obscure to prominent. Helian- themum arenicola has obscure secondary veins on the lower surface of the leaf but the pubescence differs somewhat (largely in color) on the two sides. The secondary veins of H. Nashii vary from obscure to prominent, and the pubes- cence differs on the two surfaces. Among the species with prominent secondary venation beneath only H. carolinianum and H. concolor have leaves sparsely stellate-pubescent while the remaining prominently veined species are so densely stellate-tomentose that the lower surfaces of their leaves are not visible. In spite of the indicated variability, the pubescence and the secondary veins are taxonomically helpful in dis- tinguishing the species. The MARGINS of the leaves of H. carolinianum are sub- denticulate while those of the remainder of the species are entire. The margins of some species are characteristically revolute. The INFLORESCENCES of the American species are re- markably diverse as to type, position, and structure. It may consist of: (1) all chasmogamous flowers varying from solitary (as in H. canadense) to numerous (as in H. Greenei); (2) many chasmogamous flowers and few cleis- togamous ones as in H. Coulteri; (3) few chasmogamous flowers and many cleistogamous ones as in H. corymbosum; and (4) all cleistogamous flowers, varying from a solitary flower as in H. argenteum to numerous as in H. glomeratum. A few species (H. scoparium, H. Pringlei, H. Coulteri) have unusually variable inflorescences. Other species (H. canadense, H. dumosum, H. Bicknellii, H. propinquum, and to a much lesser extent H. chihuahuense) have two types of 72 Rhodora [Vol. 67 inflorescence which mature at different times in the growing season and in different positions on the plant, resulting in great variability in aspect. However, the inflorescence is characteristic for most species and hence has proved to be a useful diagnostic feature. The BRACTS of H. corymbosum are obtusely spatulate- linear while those of the remaining species are acutely linear or lanceolate. The PEDICELS of the chasmogamous flowers range from shorter than to about five times longer than the calyx. As a taxonomic character, the pedicel has been greatly over- emphasized and its variability minimized. The cleistoga- mous flowers may be sessile or pedicellate. The FLOWERS are either chasmogamous (pollination apparently effected after expansion of the floral envelope) or cleistogamous (with pollination occurring within the closed perianth). When these two kinds of flowers are borne on the same plant, the CHASMOGAMOUS flowers are earlier and usually longer pedicellate. They are character- ized also by a larger calyx, five fugacious petals, more numerous stamens (10 or more), and by larger fruits which usually contain more numerous seeds. Their anthers are usually longer than wide and dehisce along their entire length. The CLEISTOGAMOUS flowers lack petals and are borne later (but may fruit earlier) and possess shorter pedicels, a smaller calyx, and fewer stamens (3-8). The anthers of the cleistogamous flowers are as long as wide or shorter and dehisce only by a break in the wall adjacent to the stigma to which they adhere. Four species (H. Greenei, H. scoparium, H. nutans, and H. patens) possess only petaliferous flowers. Cleistogamous flowers are very rare in H. carolinianum, while all of the remaining species are dimorphic, possessing both chasmoga- mous and cleistogamous flowers. The Mexican H. glom- eratum is the only species in which some of its members apparently produce only cleistogamous flowers. The CALYX is composed of five sepals in two dissimilar series: two narrower and usually shorter outer sepals and three inner sepals. The inner three constitute the prominent calyx. The outer and the inner sepals are united near the 1965] Helianthemum — Daoud & Wilbur 73 base in some species (e.g. H. Bicknellii, H. scoparium), while in others the fusion extends half or two thirds of their length (e.g. H. propinquum and H. Coulteri). The calyces are characteristically pubescent in all species, although they are sometimes glabrate in H. scoparium, or occasionally even completely glabrous in H. patens. The calyces of some species are covered with stellate pubescence of varying quality and density, while the calycine stellate pubescence of other species is intermixed with simple hairs and/or glandular hairs. The CHASMOGAMOUS OUTER SEPALS are variable in shape, length, and degree of fusion with the inner ones. They are useful in distinguishing species especially when combined with other characteristics. The southeastern H. corymbosum alone has spatulate outer sepals with the apices obtuse or nearly so, while the other species have linear or lanceolate sepals with acute apices. The amount of the fusion between the inner and the outer sepals and the length of the free portion of the outer ones are of significance in distinguish- ing the closely related H. Bicknellii and H. propinquum. The CHASMOGAMOUS INNER SEPALS are typically ovate or ovate-lanceolate with acute to acuminate apices except in H. Nashii whose inner sepals are asymmetrically oblique. The length of the inner sepals is exceedingly variable even within the same species and hence offers no serviceable diagnostic character. The CLEISTOGAMOUS OUTER SEPALS are mostly linear and attached near the base of the inner ones. In contrast the free portion of the outer sepals of H. canadense and H. propinquum is a rudimentary knob about twice as long as wide and fused to the middle edge of the inner sepals. The CLEISTOGAMOUS INNER SEPALS vary but little, being either ovate or ovate-lanceolate, with acute to acuminate apices. Two unrelated species, H. Nashii and H. canadense, are unusual in possessing sepals asymmetrically oblique at their apices. In most of the species the cleistogamous flowers are so small that most measurements were made at the fruiting stage. The COROLLA is present in the chasmogamous flowers and is fugacious. Cleistogamous flowers lack petals. The petals 74 Rhodora [Vol. 67 are essentially yellow but those of H. Pringlei are sometimes tipped or margined with purple. The shape of the petals is more or less uniform among the species and usually longer than the calyx at anthesis. The variation in petal size within a species is great. For example, the range is between 2.4-5.0 mm. long in H. patens, and 8-16 mm. long in H. carolinianum. ANTHER-DEHISCENCE is one of the best ways to distinguish chasmogamous from cleistogamous flowers. This is especial- ly true in fruiting specimens when the petals and some of the stamens have fallen which alone in the past have been relied upon to distinguish the two types of flowers. In the chasmogamous flowers stamen-number varies between 10- 50. The small-flowered species, such as H. argenteum, have 10-15 stamens, while the larger-flowered species, like H. carolinianum, have 20-50. The length of the filament as well as that of the anthers is quite variable. The anthers of the chasmogamous stamens, which neither cohere to each other nor adhere to the stigma at anthesis, are oblong to linear, (1-) 1.5-2 times longer than wide and dehisce throughout their entire length. In the cleistogamous flowers, the number of the stamens is also variable (3-8). Their anthers are coherent to one another and adherent to the stigma at anthesis and even in fruit for most species. These anthers dehisce by the rupture of their walls adherent to the stigma. There is such great variation in the size of the PISTIL and its parts that their size is of no taxonomic use. However the stellate-pubescent ovaries of H. N ashii and H. arenicola are most helpful in distinguishing them from all other species. The chasmogamous pistils are larger than the cleistoga- mous ones in the same species. The ovaries of the chasmoga- mous flowers are ovoid or nearly so and unilocular with few to many ovules. The styles of these flowers vary in length from wanting or nearly so in H. Coulteri to nearly as long as the ovary in H. glomeratum but never exceeding 1 mm. in length. The styles of the chasmogamous flowers, when present, are always erect and straight in American species and their stigmas are capitate and vary greatly in diameter. 1965] Helianthemum — Daoud & Wilbur 75 The cleistogamous pistils are so small that they usually have not been measured. However, their ovaries are ovoid and similar to the chasmogamous pistils of the same species in carpel-number and pubescence. Their styles are very short and are much exceeded in height by the ovaries of the same flower. The CAPSULES of both chasmogamous and cleistogamous flowers are loculicidally dehiscent. Previously the capsules of all species of Helianthemum were thought to be 3-valved. However H. Nashii characteristically is 2-valved. Only two species, H. Nashii and H. arenicola, possess stellate-pubes- cent capsules while those of the other American species are glabrous. The variation in size and shape of the chasmogamous capsules is so great as to be taxonomically useless. The capsules are shorter than the calyx (or rarely slightly longer in H. Pringlei). The chasmogamous capsules typical- ly persist throughout the growing season, but in both H. Bicknellii and H. propinquum they usually fall immediately after maturation. The largest capsules, often containing more than one hundred seeds, are found in H. caroliniamum and the smallest capsules, with only 1-3 (-6) seeds, occur in H. rosmarinifolium. Chasmogamous capsules are apparently produced by all species, although many specimens bear only cleistogamous capsules. The cleistogamous capsules vary in shape and size but are always shorter than the calyx. The largest cleistogamous capsules, containing up to 22 seeds are borne by H. Coulteri and the smallest capsules, with only one or rarely two seeds, are found in H. rosmarinifolium. Four species of course lack cleistogamous capsules. No cleistogamous capsules were seen in H. carolinianum, which is not surprising since less than one per cent of the specimens studied had cleis- togamous flowers. The number of the cleistogamous seeds per capsule is reasonably constant and hence helpful in identification. Except for the number and size, the SEEDS produced by both chasmogamous and cleistogamous fruits in the same species are similar. 76 Rhodora [Vol. 67 The seeds of twelve of the species studied are covered with a thin membrane, which is separable after the seeds are moistened with water for a few seconds. Such a membrane is usually yellowish or sometimes spotted with brown, but in H. nutans and H. argenteum white papillae occur. The seeds of the remaining species (H. canadense, H. dumosum, H. carolinianum, H. Coulteri, H. Bicknellii, H. propinquum, H. Nashii and H. arenicola) do not have a readily separable membrane. The seeds of H. canadense, H. dumosum, and H. carolinianum are papillate. The seeds of H. Coulteri are mostly papillate but in plants from Central America are often reticulate. Those of H. Bicknellii and H. propinquum are usually reticulate, while the seeds of H. Nashii and H. arenicola are pebbled to papillate. The sculpturing of the seed coats often provides useful diagnostic features. The seeds of most of the species have a somewhat trans- parent endosperm especially if examined when moist and after removal of the membrane. The embryos are curved, possess linear cotyledons, and are enveloped by the endo- sperm. They can easily be excised by rupturing soaked seeds. EVOLUTIONARY RELATIONSHIPS The American species of Helianthemum show numerous morphological trends which probably are best interpreted as evolutionary tendencies. Speculation as to evolution is of necessity highly subjective when facts are few but still it would seem desirable to record our feelings as to relation- ships. We are aware of the shortcomings of such a proposed *phylogenetic tree" as is presented in figure 1, but offer it in hopes that it will stimulate other investigations in this genus and thereby supplement the all too scanty morpho- logical data. The more obvious evolutionary tendencies are thought to be the development of cleistogamous flowers, the appearance of a hygroscopic testa and changes in inflo- rescence-type. There also seem to be established trends for decrease both in the size of floral parts and fruit accompa- nied by reduction in the number of seeds per capsule. In one species the carpel number has been reduced to two. Helianthemum — Daoud & Wilbur T1 1965] "wnwow(jupnioH jo ser eds ueotioury YON ey} Jo driugsuone[eri a[qissog "TI SA WnSO8WAHOO HO 102NO2 WNNYI9Y039 1319NIJd Wn1vN3Wo19 3SN30nHVhHIHO WniTO3INIHVWSOH WO31N394V 13N33489 wniuvdoos — N SNVLNN SN3LVd WONVINITONVS WNSOWNG 3SN3QOVNVO VIOSINAYV IIHSVN 193311002 1113853208 WNNONIdOd 78 Rhodora [Vol. 67 The supposedly more primitive species are presumed to possess only petaliferous, chasmogamous flowers. The presence of cleistogamous flowers is therefore considered an evolutionary advance. The four species lacking cleistoga- mous flowers are quite diverse and perhaps not closely related. There is no evidence that they are derived from taxa that once had cleistogamous flowers. Although possess- ing only chasmogamous flowers, it is felt that they still are evolutionarily more advanced than members (e.g. H. cana- dense and H. dumosum) that have cleistogamous flowers. The presence of a hygroscopic testa in these species would suggest a possible relationship with the supposed more advanced alliance. It is of course generally accepted that evolutionary progression does not proceed uniformly in all structures of any given taxon. Three principal evolutionary lines perhaps exist within the North American species. The four species which lack cleistogamous flowers possibly represent one line but their great diversity might be an indication that they are an unnatural assemblage of primitive species. Among the species which develop cleistogamous flowers two lines are apparent. The supposedly most advanced of the three lines consists of those species with a hygroscopic seed coat. The members of the presumed most primitive alliance have seeds lacking this hygroscopic testa. It has not appeared wise to recognize formally the three suggested evolutionary lines as taxonomic series or sections since each is character- ized by only one feature. It is felt that formally designated taxa ought to have more than one feature indicating their common origin. Since the arrangement presented here is admittedly tentative, there seems to be no urgency in formal designations. Eight species were recognized in the more primitive line characterized by papillate, pebbled or reticulate seeds lack- ing the hygroscopic seed coat. Helianthemum carolinianum is considered the most primitive American species for the following reasons: (1) the rarity of cleistogamous flowers; (2) the most primitive type of inflorescence (a scorpioid cyme) in this line; (3) the largest capsule which produces the greatest number of seeds; (4) the most numerous 1965] Helianthemum — Daoud & Wilbur 79 stamens. Barnhart (Bull. Torrey Club 27: 592. 1900.) has, however, indicated that this taxon seems to him “like a heteromorphous species which has lost its apetalous form of flowers" and he suspected that “if its origins could be traced, to find it had heteromorphous ancestors." Both H. dumosum and H. canadense are apparently very closely related and have been even treated as one species until this century. The chasmogamous calyces of these two species approach those of H. carolinianum in size. Helian- themum dumosum is here considered more primitive than H. canadense since it possesses both fewer and larger cleistogamous capsules containing more numerous seeds. Its chasmogamous flowers are usually larger than those of H. canadense. In addition the cleistogamous outer sepals of H. canadense are more reduced, being almost rudimen- tary. Also there is no sharp differentiation between the late chasmogamous flowers and the cleistogamous ones in H. dumosum. The fact that the range of H. dumosum is very restricted and confined entirely to a glaciated region in contrast to the widespread H. canadense, however, might argue for another interpretation. The unusually variable H. Coulteri often possesses a chasmogamous calyx nearly as large as those of H. cana- dense and H. dumosum. Its relationship to those two species is perhaps indicated by the frequent intermixture of stellate pubescence and pilose hairs on the chasmogamous calyx. Also the leaves sometimes resemble those two north- ern species in having the upper surface both stellate- pube- scent and pilose. Helianthemum Coulteri is judged as more advanced than either H. canadense or H. dumosum, in spite of its fewer but larger cleistogamous flowers. The southeastern H. arenicola and H. Nashii are unique among the American species in possessing stellate-pubescent ovaries and capsules. Of the two, H. arenicola is considered the more primitive since it has larger chasmogamous flowers, more seeds in each cleistogamous capsule and the characteristic 3-valved capsule rather than two which H. Nashii alone possesses in the genus. It is suggested that these two species are more advanced than the preceding members of this line because they have smaller chasmoga- 80 Rhodora [Vol. 67 mous flowers and fruit and fewer seeds/chasmogamous capsule. Also their outer sepals are narrower and shorter than the earlier-mentioned and supposedly more primitive species. The two closely related species, H. Bicknellii and H. propinquum, are perhaps more advanced than the other rough-seeded species since their numerous cleistogamous capsules are smaller and contain fewer seeds. Helianthemum propinquum is possibly more advanced than H. Bicknellii since it has fewer chasmogamous flowers and a much smaller free portion of the outer sepals. Helianthemum propinquum. has smaller cleistogamous outer sepals and generally smaller cleistogamous capsules. Although H. propinquum is ranked as the most advanced species in this evolutionary line, it is thought to be more primitive than some members of the other supposed phylads. The four Mexican and Californian species lacking cleis- togamous flowers (H. Greenei, H. scoparium, H. nutans and H. patens) perhaps represent an independent line but are so dissimilar that placing them together may be unwarranted. Among them H. Greenei, a narrow endemic to the Channel Islands of California, is treated as the most primitive be- cause of its broader, lanceolate outer sepals. The more widespread and variable H. scoparium is related to H. Greenei but has narrow, linear, outer sepals. The remain- ing strictly chasmogamous species, H. nutans and H. patens, possess a different habit and are believed to be more ad- vanced, as indicated by reduction in number of flowers and size of leaf. By similar reasoning H. patens would be con- sidered more advanced than H. nutans since its flowers are smaller with fewer stamens and fewer seeds/capsule. In addition the herbaceous habit of H. patens contrasted with the fruticose habit of H. nutans might be considered as an evolutionary advance. These four species possess a hygro- scopic testa which also characterizes the supposedly more advanced evolutionary line with cleistogamous flowers. The most primitive member of the line with a hygroscopic seed-coat is possibly H. corymbosum. Its more foliaceous outer sepals and bracts may indicate its relative primitive- ness. It also is the only species in this line possessing the 1965] Helianthemum — Daoud & Wilbur 81 prominent and persistent funiculi and placentae which are found also in the three supposedly more primitive species of the non-hygroscopic seeded alliance. It is suggested that H. concolor is another primitive species. It possesses the largest leaves and the longest outer sepals of any member of the hygroscopic-seeded alliance. It is treated here as more advanced than H. corymbosum because of its much narrower outer sepals and bracts and fewer seeds in each of the cleistogamous capsules. The next more advanced species within this line are perhaps H. georgianum and H. Pringlei. 'These two species have a supposedly reduced and hence more advanced in- florescence-type (a racemose cyme), and their outer sepals are reduced in size. Helianthemum georgianum may be more primitive than H. Pringlei as it has more numerous seeds in its cleistogamous capsule. It is felt that H. chihuahuense is more advanced than the two preceding species since its chasmogamous and cleis- togamous flowers are borne separately and in different positions on the plant. In addition the number of the cleistogamous flowers has been often reduced to a single flower in the leaf axil which seems to be clearly derived from the many-flowered clusters borne terminally on short axillary branches. Helianthemum glomeratum is thought to be more advanced than H. chihuahuense since the number of chasmogamous flowers is reduced to a usually solitary flower borne at the end of the branches and branchlets. Also chasmogamous flowers are occasionally lacking and the size of the cleistogamous capsule is reduced with only 1-3 seeds in each capsule. When chasmogamous flowers are present in H. glomeratum, they are to be found terminating both branches and branchlets while in both H. rosmarini- folium and H. argenteum the solitary chasmogamous flow- ers are to be found only at the tips of the major branches. Also in these two species the number of seeds/cleistogamous capsule is never more than one or two. In addition the cleistogamous capsules of both species are more reduced than in any other species. However, H. argenteum is con- sidered even more advanced than H. rosmarinifolium and also perhaps the most advanced species of the North 82 Rhodora [Vol. 67 American species of Helianthemum since (1) the chasmoga- mous flowers have been reduced to one at the tips of the major branches; (2) the more primitive condition of clus- tered cleistogamous flowers has been reduced to but one (or rarely two together) in the leaf axils; (3) the cleistoga- mous capsules are the smallest of all the species studied ; and (4) the leaf-size is greatly reduced. The primitive and advanced features are summarized below : More than 7000 herbarium specimens were examined of the Helianthemum collections of 53 herbaria to whose curators we are most indebted. These herbaria are indi- cated by their abbreviations following Lanjouw & Stafleu (Regnum Vegetabile, ed. 4. 15: 1959.) : A, ARIZ, BUS, CHRB, COLO, CU, DS, DUKE, F, FLAS, FSU, GA, GH, IA, ILL, IND, ISC, JEPS, KANU, KY, MICH, MIN, MISSA, MO, MSC, NCSC, NCU, ND, NO, NY, OKL, OKLA, OS, PENN, PH, POM, PUR, RSA, SBBG, SBM, SD, SMU, TENN, TEX, UARK, UC, UMO, US, SCAR (—Univer- sity of South Carolina), USF, VDB, WIS, WVA. PRIMITIVE ADVANCED 1. Possession of only chasmoga- 1. Possession of both chasmoga- mous flowers. mous and cleistogamous flowers or of cleistogamous flowers only. 2. Few but large cleistogamous 2. More numerous but smaller flowers. cleistogamous flowers. 3. Floral parts large. 3. Floral parts relatively small. a. Outer sepals of chasmoga- a. Outer sepals of chasmoga- mous flowers spatulate (and mous flowers linear (and relatively large). small). b. Outer sepals of cleistoga- b. Outer sepals of cleistoga- mous flowers large. mous flowers much smaller, often reduced to knob-like rudiments. c. Stigma large. c. Stigma small. 4. Fruits large and many-seeded. 4. Fruits small and few-seeded. 5. Numerous stamens in the . Fewer stamens in the chasmo- c chasmogamous flowers. gamous flowers. 6. Carpels 3. 6. Carpels 2. 7. Non-hygroscopic testa, 7. Hygroscopic testa. to be continued AN EARLIER NAME FOR LUDWIGIA NATANS (ONAGRACEAE) PETER H. RAVEN John Reinhold Forster's “Flora Americae Septentrion- alis, or a Catalogue of the Plants of North America .... ,” published in London in 1771, has in general been neglected. It is an enumeration in the form of a list with English names, localities, references, and occasionally economic notes on the plants known from North America up to that time, arranged according to the Linnaean system. It is not an original work but rather a compilation based on the published works of other students. Thus, Forster himself evidently considered its greatest utility to serve as a kind of index to the important early works of Kalm, Gronovius, Sloane, and Catesby. Forster did, however, propose new binomials for a few species described in the second edition of Gronovius' Flora Virginica (1762), a work which did not adopt Linnaeus’ binomial system of nomenclature. One such is Gronovius’ “Ludwigia caule repente, foliis obverse ovatis petiolatis" (p. 20), for which Forster (1771, p. 6) proposed “Ludwigia repens? N. S.,” quoting Gronovius’ description in full. It is often possible to typify Gronovius' names with collections made by Clayton. Clayton's specimen of this particular species, in the herbarium of the British Museum (Natural History), is Ludwigia palustris (L.) Ell. It cannot be taken as a type, however, as the description ("floribus . . . dilute luteis tetrapetalis fugacissimis . . .") can refer only to the species currently known as Ludwigia natans Ell. (1821). Clayton's mention of petals probably accounts for Linnaeus' not identifying this species with his Jsnardia palustris [= Ludwigia palustris (L.) Ell.], as subsequent authors have incorrectly done (cf. Fernald, Rhodora 37: 175-177. 1935). Thus, the partial synonymy for this species is as follows: Ludwigia repens Forst., Fl. Amer. Sept. 6, 1771. Ludwigia natans Ell, Sketch Bot. S. C. & Ga. 1: 581. 1821. 83 84 Rhodora [Vol. 67 Ludwigia repens Forst. is listed in the Index Kewensis (2: 123. 1895), but the page reference is given incorrectly as “22.” Five other new species are proposed by Forster in his work. Two on p. 7 are published under the genus “Menandra”’ and are therefore invalid, *Menandra" itself never having been published. A third proposed species, “Rhamnus volubilis," p. 11, is a nomen nudum. The remain- ing two, Potamogeton rotundifolium and P. oblongifolium, p. 7, are validly published and should, therefore, be taken up by the Index Kewensis and similar works. Gronovius' brief descriptions (Fl. Virg., ed. 2, 23. 1771) might make it possible for them to be identified. Unfortunately, there is apparently no existing herbarium material that can be identified with these names. This work has been supported by National Science Foun- dation Grant GB-141. DIVISION OF SYSTEMATIC BIOLOGY, STANFORD UNIVERSITY, STANFORD, CALIFORNIA MOUNTAIN FLOWERS OF NEW ENGLAND" This is an important illustrated and convenient guide to the alpine plants of New England and New York. The excellent illustrated series of articles by Stuart K. Harris, Plants of the Presidential Range, that appeared in Appalachia at intervals during the years 1940-1949 provided the principal basis for the present book. The area covered being somewhat larger in the present work, it has been necessary to define the boundaries of the alpine area very carefully; also, it was necessary to provide a complete and accurately selected list of species and to be as precise as possible about habitats. In these matters, the help of Frederic Steele has been particularly valuable. "Mountain Flowers of New England, by Stuart K. Harris, Jean Langenheim, Frederic L. Steele, and Miriam Underhill. Appalachian Mountain Club, Boston, Mass. 150 pp. April 1964. $4.50. 1965] Book Review — Hodgdon 85 The thirty-two plates in color at the end of the book are unusually well done and represent an admirable selection of the flora. Of these, twenty-eight are photographs of vascular plants illustrating some 129 species. Selected examples of mosses and lichens are depicted on three plates, while the final page shows six scenic habitats in the mountains of Maine and New Hampshire. This important photographic contribution was made by Miriam Underhill. The text on lichens and mosses was contributed by Jean Langenheim. With these groups there was no attempt made to be exhaustive, but a good beginning can be made by the amateur using the book and referring to the illustrations. The book is compact (4.5 X 7.5 in.), and with its profuse drawings, color photographs, readable descriptions, keys and glossary, will help materially to educate the mountain climber. A few suggestions for improvements in future printings or editions are perhaps in order. A comment from one of the authors refers to the desirability of providing infor- mation about dates of flowering. For example, certain species do not flower as late as July in most years, while others do not commence flowering until July. This infor- mation could be given in a special table or added to the discussion for any particular species. It seems to this reviewer that the four authors should be given more promi- nence by placing their names on the title page. The intro- duction seems to be an unusual place to have to look to locate the authors’ names. A. R. HODGDON, UNIVERSITY OF NEW HAMPSHIRE A SECOND CHARACTER DISTINGUISHING HETEROTHECA S. STR. FROM CHRYSOPSIS (COMPOSITAE: ASTEREAE) VERNON L. HARMS The sole criterion of the absence of pappus bristles in the ray florets of Heterotheca Cass. and their presence in Chry- sopsis has been used traditionally to separate the respective genera. An examination of numerous specimens of all included entities has revealed another character which also seems consistently to separate the two groups if the earlier cauline leaves have been retained on the specimens. The lowermost leaves of most Chrysopsis and of all Heterotheca s. str. species are distinctly petiolate and the upper ones sessile. In Chrysopsis there is simply a progressive re- duction of the petiole length with no enlargement of petiole bases from the lower to upper leaves (Fig. 4-8). But in Heterotheca s. str., the sequence from the lower petiolate to the upper sessile leaves is characterized by progressively more expanded petiole bases in successive leaves, resulting in a gradual proximal to distal lamination of the petiole (Fig. 1-3). The radical and very lowermost cauline leaves have long petioles without expanded bases. The leaves just above these begin to reveal small laminar petiole bases. These petiole bases or auricles become increasingly more expanded and conspicuous in successive leaves to merge eventually with the leaf blade proper obscuring all traces of the petiole. The peculiar lyrate to panduriform shapes of the middle leaves of this petiole lamination series in Heterotheca s. str. (Fig. 1-3) are quite distinctive of the group, never being present in any Chrysopsis species. This characteristic leaf sequence is apparent in all Heterotheca entities, but of course is not distinguishable on herbarium specimens if none of the lower petiolate leaves have been retained. In large, well-developed Heterotheca plants, especially in dense stands, most of the lower leaves may be deciduous and the remaining leaves predominantly of the upper sessile type. 86 1965] Heterotheca — Harms 87 009909000 : (0080006 E Omg Qu pum T ^ ^E de etd npn E. 10 cm. e Fig. 1-8. Comparative series of basal to upper leaves in various Heterotheca and Chrysopsis species. Fig. 1. Heterotheca latifolia, Clark County, Ga, G. L. Plummer. Fig. 2. H. subaxillaris Tift County, Ga, G. L. Plummer. Fig. 3. H. psammophila, Cochise County, Ariz., V. L. Harms 1849. Fig. 4. Chrysopsis camporum, St. Louis County, Mo., R. L. McGregor. Fig. 5. C. berlandieri, Harvey County, Kans, V. L. Harms 1324. Fig. 6. C. stenophylla, Woods County, Okla., V. L. Harms 1995. Fig. 7. C. fulcrata, Dona Ana County, N. Mex., V. L. Harms 1839. Fig. 8. C. villosa, Jackson County, S. Dak., V. L. Harms 2149. 88 Rhodora [Vol. 67 Yet in the field it is usually possible to find at least some shoots of almost every plant which reveal traces of this very characteristic petiole sequence. Various field botanists have previously noted and used the peculiar basal lobing of the petioles as a field character to help distinguish local Hetero- theca species. But this attribute has apparently never been recognized as a consistent diagnostic feature characteristic of all Heterotheca s. str. species in contrast to those of Chrysopsis. While this second diagnostic character improves the status of Heterotheca s. str. as a natural group and might possibly be construed as enhancing the continued generic separation of the two groups, such a conclusion is hardly warranted. Foliage characters just as diverse, if not more so, separate the section Pityopsis (C. graminifolia and its allies) from the other sections of Chrysopsis. Both morpho- logical data (Shinners, 1951, and Wagenknecht, 1960) and cytogenetic evidence (Harms, in press) seem to point to the congeneric status of Heterotheca and Chrysopsis, in which case the peculiar petiole lamination sequence reported here would constitute another sectional character, along with epappose ray florets, serving to distinguish Heterotheca sect. Heterotheca from the other sections of an enlarged genus. DEPARTMENT OF BIOLOGICAL SCIENCES, UNIVERSITY OF ALASKA, COLLEGE. LITERATURE CITED HARMS, VERNON L. In press. Cytogenetie evidence supporting the merger of Heterotheca and Chrysopsis (Compositae). Brittonia. SHINNERS, LLOYD H. 1951. The North Texas species of Heterotheca, including Chrysopsis (Compositae). Field and Lab. 19:66-71. WAGENKNECHT, BURDETTE L. 1960. Revision of Heterotheca section Heterotheca (Compositae). Rhodora 62:61-76, 97-107. TULOSTOMA BRUMALE PERS. A NEW RECORD FOR NEW ENGLAND WESLEY N. TIFFNEY AND WESLEY N. TIFFNEY, JR. In April 1963, we were collecting on Good Harbor Beach, Gloucester, Essex County, Massachusetts. Mr. Arnold Kenniston of Norton drew our attention to an unusual Basidiomycete. Later, we tentatively identified the plant as Tulostoma brumale Persoon of the Series Gasteromycetes in the order Sclerodermatales. This is a stalked puffball which produces its fruiting body in the fall as a small sclerotial swelling some two to three centimeters under the surface of the ground (Gaumann and Dodge, 1928). On maturation, it emerges on a slender stipe and sheds its exoperidium, exposing an endoperidium which opens by an apical mouth. Good Harbor Beach is a publie bathing area. Back of the broad, sandy storm beach is a series of low dunes which extend for about 350 yards to a public highway. This broad area of fine sand is a rarity on rocky Cape Ann. In 1963, thirty-five specimens of Tulostoma were collected from the north end of the beach ninety to one-hundred feet inland from the fore-dune. In 1964, thirty-three additional individuals were taken, nine from the previous station and twenty-four from a location sixty to seventy-five feet south of the 1963 site and about seventy-five feet from the fore- dune. The vascular plants associated with T'ulostoma were as follows: Agropyron repens (L.) Beauv. (common), Ammophila breviligulata Fernald (fairly common), Lathy- rus japonicus Willd. var. glaber (Sev.) Fernald (fairly common), Limonium Nashii Small (rather rare), Solidago sempervirens L. (rather rare), Sedum sp. (only a few plants), and Taraxacum officinale Weber (only a few). Specimens are on deposit at the Herbarium of the Universi- ty of New Hampshire. Our collection in Gloucester constitutes a new record for both the genus Tulostoma and the family Tulostomataceae in New England. The nearest station to our material for 89 90 Rhodora [Vol. 67 the genus is one reported by White (1901) in Ithaca, New York. The identification of this organism as Tulostoma brumale Persoon 1801 posed a considerable problem. We visited the Farlow Herbarium and the Herbarium of the New York Botanical Garden and examined their collections of Tulos- toma." We feel from our investigation that many uncertainties exist in the taxonomy of the genus. At present, no studies are available showing possible variation within a single large population. If such studies were systematically made for the genus, it might be possible to reduce to synonomy many existing species. Cunningham (1942) supports this conclusion when he states that of the eighty-five described species of Tulostoma, “not more than about thirty are valid." Our present collection is of a size to warrant making a statistical examination of Tulostoma brumale. Coker and Couch (1928) noted that Tulostoma simulans Lloyd was found at the same station for nineteen years. This indicates a perennial tendency in at least some species of the genus. Our material from Cape Ann appears to be perennial since it has persisted in the same locality for two years. If our organism can be collected annually, it is our hope to find eventually young specimens in the hypogenous stage and perhaps to verify Schroeter's figures of the basi- dia and the method of spore production. There is confusion in the taxonomic history of the genus Tulostoma and of the species T. brumale. Persoon es- tablished T. brumale in 1801 (Synopsis methodica fun- gorum, p. 139). Synonyms for Persoon's species are. T. imbricatum Pers. 1794 (N. Mag. Bot. (Rom.)1:86); T. squamosum Pers. 1801 (Syn. meth. fung., p. 139); T. mammosum Fr. 1829 (Syst. 3:42); and T. pedunculatum (L.) Schroet. 1876 (Beitr. Biol. Pflanz. (Cohn) 2:65). There has also been confusion concerning the spelling of "The authors are very grateful to Dr. Clark T. Rogerson, Curator of Cryptogamie Botany of the New York Botanical Garden, for the time and attention given to us during the two days spent at the herbarium. 1965] Tulostoma — Tiffney & Tiffney, Jr. 91 the generic name. Persoon established the genus in 1801 and spelled it Tulostoma. Sprengel in 1827 (Systema Vegetabilium 4(1) : 524) followed by Fries in 1849 (Sum- ma Vegetabilium Scandivaviae 440), Saccardo in 1888 (Syl. fung. vol. 7) and Long in 1946 (Mycologia 38:77), among others, questioned the etymology of the word and pointed out that the Greek upsilon should be transcribed by the Latin “y”, thus spelling the genus Tylostoma. We consulted Dr. Donald P. Rogers? of the University of Illinois on this matter, and he pointed out that there was “no law on the point and the practice was not invariable.” He stated that such names as “Tulocarpus, Tulodiscus and Buxus or Mucor” were equally vulnerable to the same type of logic. Thus, no “inadvertance or error on the part of the author" can be claimed. With this support, we advise the retention of Persoon’s spelling of Tulostoma. BIOLOGY DEPT., BOSTON UNIVERSITY, BOSTON, MASSACHUSETTS BOTANY DEPT., UNIVERSITY OF NEW HAMPSHIRE (Graduate School), DURHAM LITERATURE CITED CokKER, W. C., and J. N. CovcH. The Gasteromycetes of Eastern United States and Canada, University of North Carolina Press, Chapel Hill, N. C., 1928. CUNNINGHAM, G. H. The Gasteromycetes of Australia and New Zealand, John McIndoe, Dunedin, New Zealand, 1942. GAUMANN, E. A., and C. W. DopGE. Comparative Morphology of the Fungi, McGraw-Hill Co., New York, 1928. WHITE, V. S. The Tylostomaceae of North America, Bull. Torr. Bot. Club 28: 421, pls. 31-40, 1901. "We wish to thank Dr. Rogers for his aid with synonomy of the species and spelling of the generic name. NEW RECORDS OF ARCTIC SPECIES IN SOUTHEASTERN NEW BRUNSWICK Three arctic species with southern mainland limits in the mountains of the Gaspe Peninsula, according to Fernald (1950), are Dryas integrifolia Vahl, Salix myrtillifolia Anderss. and Solidago multiradiata Ait. The last named has been reported from St. Paul Island, Nova Scotia, by Perry in 1931. These three species were found in the vicinity of Hills- borough, Albert County, in south-eastern New Brunswick during 1964 associated with Anemone parviflora Michx., Shepherdia canadensis (L.) Nutt., and Potentilla fruticosa L., new records for this part of the province, and with Erigeron hyssopifolius Michx., Galium triflorum Michx., Mitella nuda L., Campanula rotundifolia L., and Cystopteris bulbifera (L.) Bernh., all common elsewhere in Albert County. The habitat is a crumbling cliff of white gypsum, less than half a mile long, 70-80 ft. high, with a north-facing slope of 70-159. The plateau at the top of the cliff is 300 ft. above sea level and bounded on at least three sides by the steep gypsum cliffs. It is dissected slightly by streams and honey- combed with funnel-shaped sinkholes ranging from 10 to 60 feet in depth and in orifice diameter. The plateau, which has been partially cut over, supports a somewhat stunted forest of Picea rubens Sarg., P. mariana (Mill.) BSP., Abies balsamea (L.) Mill., Betula lutea, Michx.f. and Tsuga cana- densis, (L.) Carr.; there is generally a heavy carpet of herbs, mosses and lichens beneath. Along the streams, for the most part, the same species are present but the forest growth is richer, particularly at the base of the cliffs. None of the newly recorded species are present in the forest al- though Salix myrtillifolia is abundant near the base of the north-facing cliff at the upper edge of the forest. Some of the other species do appear wherever gypsum occurs at the surface and the canopy is open. Dryas integrifolia, Solidago multiradiata and Anemone parviflora appear to be confined to the exposed north-facing cliff, forming, with the other 92 1965] New Records — Roberts 93 species, mats of up to two square meters in area on the otherwise loose talus. Other gypsum cliffs in Albert County investigated thus far do not support Dryas integrifolia, Solidago multiradi- ata, Anemone parviflora or Salix myrtillifolia although all of the other species are usually present. The nomenclature employed here follows Fernald (1950). Specimens have been deposited in the Herbarium of the University of New Brunswick, Fredericton, New Bruns- wick and in the Phanerogamic Herbarium of the Canada Department of Agriculture, Ottawa. This work is part of project G15, “The distribution of the flora of New Brunswick,” supported by the New Bruns- wick Research and Productivity Council. The help of Dr. Bernard Boivin of the Canada Department of Agriculture, Ottawa, in checking identifications is gratefully acknow- ledged. PATRICIA R. ROBERTS, UNIVERSITY OF NEW BRUNSWICK, FREDERICTON, NEW BRUNSWICK. LITERATURE CITED FERNALD, M. L. 1950. Gray's Manual of Botany. 8th ed., American Book Co., N. Y. Perry, L. M. 1931. Vascular Flora at St. Paul Island, Nova Scotia. Rhodora 33: 105-126. COMMON TREES OF PUERTO RICO AND THE VIRGIN ISLANDS Visitors to the West Indies interested in trees usually come away with much misinformation acquired from chauf- feurs and local guides. Even the best “official guides" in the "Common Trees of Puerto Rico and the Virgin Islands. By Elbert L. Little, Jr., and Frank H. Wadsworth. U. S. Department of Agricul- ture, Agriculture Handbook No. 249, 548 pages, illus. 1964. For sale by the Superintendent of Documents, U. S. Government Printing Office, Washington, D. C. 20402. Price $4.25 (cloth). 94 Rhodora [Vol. 67 several botanie gardens often confuse rather than inform the interested tourist. This is particularly true if several islands are visited where different languages and cultural backgrounds have served to give diverse and often over- lapping common names to the native and introduced trees. Even a visitor with considerable botanical background finds himself fairly helpless when confronted for the first time with numbers of unfamiliar tropical and subtropical trees and is usually frustrated in identifying any except a handful of well known ornamental and economic species. The preparation of this guide by the Forest Service and USDA offers at least partial relief to this situation by describing in nontechnical language 250 species of the com- moner native and introduced trees of Puerto Rico and the Virgin Islands. Full page line drawings, as near natural size as space permits, of foliage, flowers and fruit face each description and permit identification by this means alone. In addition, several categories and nontechnical listings as well as keys are included to enable even the novice to identify the described trees. A valuable part of each treatment is the listing of common names used in various Carribbean islands and countries, With both English and Spanish editions available this volume will be helpful in the whole Carribbean and Central American area as a great many of the species described occur throughout. Information is included indicating economic value and ornamental use of the trees described which should make this book valuable to foresters, horticulturists and other residents of the area. Biology teachers from grade school to college level will find this an almost necessary reference book. Both scientific and common names are included in one complete index. A second volume is planned to include several hundred less common and rare tree species (27 feet or over) which will presumably complete the treatment for these islands. R. B. PIKE, UNIVERSITY OF NEW HAMPSHIRE DISSECTING EQUIPMENT AND MATERIALS FOR THE STUDY OF MINUTE PLANT STRUCTURES' At one time the agrostologist's favorite bit of dissecting equipment was a carefully cultured fingernail. However handy this might be, it lacked the precision that artificial equipment may possess. Unfortunately, the average bot- anist's collection of dissecting equipment is ill-suited to the tasks of dissecting small plant structures and rarely is kept in proper condition. Hooked-tip dissecting needles or blunt prods make the ticklish job of manipulating spikelet parts or similar small structures almost impossible. The following suggestions for the fabrication and care of the basic equip- ment for handling spikelets and similar organs are the outgrowth of many years of working with grass structures. The common wooden-handled dissecting needle is one of the most useful and most neglected items of the taxonomist's equipment. By culling over a collection of needles, one can select the strongest for conditioning. They should be pointed on a fine carborundum stone to remove burrs and to give the point a 45? taper. A long needle-point is not desirable, lacking the necessary strength. Final sharpening and pol- ishing should be done on the finest grade of oil stone avail- able. The Behr-Manning HF 873 Hard Arkansas Knife Blade is ideal. The tip should be circular in cross section all the way to the point. The proper sharpening of needles is greatly facilitated by the following method of handling: hold the wooden handle of the needle between the ball of the extended thumb and the second finger, which should be hooked back toward the palm of the hand. The tip of the index finger then lies on top of the tip of the needle, and presses it against the stone. With oil on the stone, slide the needle tip forward and backward over the stone. While thus sharpening the point, the handle of the needle should be rotated between the thumb and the second finger. The point produced by this manipulation will be uniformly ta- pered and as sharp as desired. Needles should not be ‘Journal Paper #J-4880 of the Iowa Agricultural and Home Eco- nomics Experiment Station, Ames, Project 1136. 95 96 Rhodora [Vol. 67 sharpened by rubbing sideways on the stone, as chatter will ruin the point. I still treasure à micro-scalpel of pre-war German manu- facture, its blade now almost worn away. Since "Ersatz" American-made products available since the war have been very unsatisfactory, we have made our own micro-scalpels by a variety of processes. The most satisfactory tools can be made from the end section of coping-saw blade (.110” wide X .020" thick). The handle is an ordinary wooden dissecting needle handle with the needle removed. The base of a two inch section snipped from the saw blade is thrust into the wooden handle. The hole may have to be enlarged with a fine drill, and the blade can be firmly set with plastic glue. If desired, the shank of the blade can be strengthened by wrapping it in thin metal from a juice can. The saw teeth can be filed or ground off and the scapel blade roughly shaped in any desired style with wire nippers, a file or coarse stone. Final sharpening should be done on a fine oil stone. The scalpel blade ordinarily should not be over 3 or 4 mm. long and its edge should be sharpened to a V-profile. Extremely thin edges should be avoided, since they will merely roll, rather than cut. Dissection of tough or rigid plant structures is facilitated by the application of a few drops of the following solution. It penetrates rapidly, is non-staining, and requires no boil- ing. It may safely be used on herbarium sheets. We dis- pense it from a miniature polyethylene squeeze-bottle, provided with a medicine-dropper tip. Dioctyl sodium sulfosuccinate? 1% distilled water 74% Methyl alcohol 25% RICHARD W. POHL IOWA STATE UNIVERSITY, AMES *Sold commercially as “Aerosol OT”. Volume 67, No. 769 including pages 1 to 96, was issued March 31, 1965. FARLOW REFERENCE LIBRARY JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL 3 STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER » Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. J Vol. 67 April-June, 1965 No. 770 CONTENTS: Studies in the Flora of Bolivia, — III. Cyperaceae, Part 1. Robert C. Foster 97 Genotypic Variation in the Phacelia strictiflora Complex George W. Gillett i 139 Interpretation of Crossing Diagrams David L. Mulcahy .... 146 II. Monograph of the Genus Elodea, Summary Harold St. John TREASURE T 155 Flowering Plants New to or Rare in Kentucky Elizabeth M. Browne and E. T. Browne, Jr. ................. 180 The Typification of Xanthocephalum (Compositae) Eë TT 4isocesQernssrierr EH 182 (Continued on Inside Cover) The Nem England Botanical Club, Jne. Botanical Museum, Oxford St., Cambridge 38, Mass. CONTENTS: — continued Two Species of Chamaesyce (Euphorbiaceae) New to the United States Derek Burch ............ eene 185 Chromosome Numbers in Penstemon (Scrophulariaceae) I: New Mitotic Counts ..Frank S. Crosswhite and Shoichi LI TN 187 Additions to the Vascular Flora of Oklahoma R. John Taylor and Constance Taylor ................. eere 191 Chamaesaracha villosa New to Texas James F. Scudday .... 194 Notes on the Flora of Coós County, New Hampshire Stuart K. Harris |... eere eee eene eene nnns ER 195 An Eastern Variety of Carex fissa (§ Multiflorae) F. J. Hermann EE 198 Ipomoea amnicola (Convolvulaceae), A South American Waif in Missouri Lloyd H. Shiner NN 200 A Revision of the North American Species of Helianthemum (Cistaceae) H. S. Daoud and Robert L. Wilbur ................ 201 Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 67 April-June, 1965 No. 770 STUDIES IN THE FLORA OF BOLIVIA, — III. CYPERACEAE, PART 1. ROBERT C. FOSTER This treatment of the Bolivian Cyperaceae is complete except for the genus Cyperus, which will constitute Part 2 of the family treatment. Since it is floristic, rather than monographic, synonymy has mostly been limited to names, some based on misidentifications, which were included in my Catalogue of the Ferns and Flowering Plants of Bolivia (Contrib. Gray Herb. no. 184 [1958]), but which now seem properly relegated to synonymy. In a few instances, I have not seen Bolivian material of species, but after the Bolivian specimens cited I have placed in parentheses the name of a previous worker who had seen them. A generic description is given whenever two or more species of a genus are treated. When only one species of a genus occurs in Bolivia, a combined generico-specific de- scription is given, to prevent unnecessary repetition. It should be emphasized that generic descriptions are based solely on the Bolivian representatives of a genus. For the most part, the treatment is based on the col- lections of the Gray Herbarium (GH). I have, however, received supplementary material from the New York Botan- ical Garden (NY), the United States National Herbarium (us) and the Missouri Botanical Garden (MO). I am much indebted to the curators of these institutions for making their specimens available. Above all, I am most grateful to Dr. Lyman B. Smith for his patience in answering questions and his kindness in searching for material which would be helpful to me. 97 98 Rhodora [Vol. 67 KEY TO GENERA a. Flowers unisexual. b. Rachilla of pistillate flowers prolonged and conspicuous. ....... 14. Uncinia. TTT LOL b. Rachilla neither prolonged nor conspicuous. c. Achene covered by a puberulent membrane or by a perigynium. d. Achene enclosed in a puberulent membrane; staminate flower a single stamen. .............. ener 12. Calyptrocarya. d. Achene enclosed in a perigynium; staminate flower of 3 STAMENS. eeeeeesese otn onm eo one ntt oa tos rtto nm— — — 15. Carez. c. Achene not covered by a membrane nor by a perigynium. ........ 13. Scleria. a. Flowers perfect (at least some “of them). e. Leaf-blades absent. ............... eene nennen 7. Eleocharis. e. Leaf-blades present (sometimes much reduced). f. Style-base persistent on the achene. g. Style-base a tubercle, ..................-. EE 9. Bulbostylis. g. Style-base a beak. h. Perianth present (of scales or bristles). . Achene surrounded by a transparent scale or utricle. "—————————— 3. Ascolepis. i. Achene surrounded by scales or bristles. j. Perianth of 3 flattened scales alternating with 3 bristles (or the bristles absent). ................ 5, Fuirena. j. Perianth of bristles only (flattened scales absent). :n—n—— nn, 11. Rhynchospora. h. Perianth absent. ............ "E 11. Rhynchospora. f. Style-base not persistent on the achene. k. Perianth-bristles present. ............ een 4. Scirpus. k. Perianth-bristles absent. l. Inner scales present in flowers. m. Inner scales 2, like a 2-valved utricle. .... 1. Lipocarpha. m. Inner scale 1, small, between achene and axis of spikelet. |... eere rere erronee tento porta ertt 2. Hemicarpha. l Inner scales absent in flowers. n. Spikelets mostly many-flowered. o. Spikelet-seales 2-ranked, conduplicate. .... 6. Cyperus. o. Spikelet-scales spirally arranged. p. Style-base thickened or dilated. ........ 8. Fimbristylis. p. Style-base not thickened. ........................ 4. Scirpus. n. Spikelets mostly 1-2 (-few)-flowered. ........ 10. Cladium. 1. LIPOCARPHA R. Br. Lipocarpha Sellowiana Kunth, Enum. Pl. 2: 267 (1837). Caespitose perennial to 8 dm. tall. Culms slender, rigid, triangular. Leaves few, basal, shorter than the culms, basal sheaths often reddish or purplish, 1965] Flora of Bolivia — Foster 99 blades subterete, canaliculate, the apex obtuse. Involucre of 2-3 foliar bracts, the lowermost to 12 cm. long, the others much shorter. Inflo- rescence of 2 to several sessile, ovoid or subglobose spikes forming a dense capitulum. Glumes imbricate, to 2 mm. long, obovate to cuneate with a broad terminal acumen, many-nerved, striate, subtending axillary perfect flowers of 1-2 stamens and trifid style, these sur- rounded by a 2-valved utricle with one scale adaxial, the other abaxial, the 2 parts more or less transparent, membranous, many-nerved, elliptic-ovate and more or less concealed by the glumes. Achene to 1.8 mm. long, basally attenuate, obscurely trigonous. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1902 (GH), 5140 (GH), 6895 bis (GH). 2. HEMICARPHA Nees & Arn. Hemicarpha micrantha (Vahl) Pax in Engl. & Prantl, Nat. Pflanzen- fam. 2(2): 105 (1887). Scirpus micranthus of the Catalogue. Dwarf annual with dark fibrous roots. Culms to 9 cm. high, often much shorter, capillary, smooth. Leaves much shorter than culms, capillary, subterete, sheath-orifice oblique. Inflorescence terminal, pseudolateral, of 1-3 spikelets, involucral bract much exceeding spikelets and appear- ing as a continuation of the culm. Spikelets ovoid to oblong-ovoid, to 2 mm. long; scales brown with light streaks, oblong-ovate, to 1 mm. long, but usually not much exceeding the achene, not carinate; a small translucent scale between the achene and spikelet-axis; perianth- bristles 0; stamens 1; style bifid. Achene narrowly obovoid, briefly apiculate at the abruptly obtuse apex, about 0.5 mm. long, pale brown until full maturity, then dark brown, minutely and evenly papillate in longitudinal rows, style-base not persistent as a tubercle. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1919 (GH). 3. ASCOLEPIS Nees Ascolepis brasiliensis (Kunth) Benth. ex C. B. Clarke in Dur. & Schinz, Conspect. Fl. Afr. 5: 651 (1894). Perennial, from a more or less horizontal slender rhizome, to 6 dm. tall. Culms slender to filiform. Leaves basal, few in a tuft, shorter than the culms, filiform, basally sheathing. Involucre of 1-3 unequal bracts, the longest to 9 cm. long. Inflorescence capitulate, of 1-3 dense, ovoid to globose sessile spikes to 8 mm. long. Spikelets spirally inserted on the rachis of the spike, each consisting of a long-triangular or long-deltoid several-nerved glume to 2 mm. long and less than 1 mm. wide at the base, subtending a flattened, oval, several-nerved scale or utricle to 2.5 mm. long, the apex abruptly acuminate, the wide margins hyaline, this containing the perfect flower of 1-2 stamens and bifid style, the style-base persistent on the oblong, biconvex, basally attenuate achene, the achene-coat shining brown, finely reticulate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6727 (GH). 100 Rhodora [Vol. 67 4. SCIRPUS L. Annual or perennial, in wet or damp habitats. Plants dwarf to very tall, the culms filiform to broad-trigonous. Leaves with basal sheaths, the blades mostly short and narrow, or greatly reduced, Inflorescence terminal, reduced to 1-many spikelets, pseudolateral in some species. Spikelets few- to several-flowered, the lowest scales empty, flowers perfect, perianth-bristles present or absent; stamens 2-3; style bifid or trifid. Achene plano-convex or trigonous, smooth or somewhat reticulate or scrobiculate. a. Inflorescence not pseudolateral. b. Perianth-bristles 0. c. Achene contracted apically into a short neck. d. Culms to 2 dm. high; achene trigonous. ........ 2. S. cernuus. d. Culms to 3 em. high; achene plano-convex. .. 4. S. boliviensis. c. Achene not contracted apically into a neck, but abruptly rounded-obtuse. een "n" EE b. S. atacamensis. b. Perianth-bristles present. e. Inflorescence sessile; leaves in a rosette; achene pale. ............ :——————— 8. S. acaulis. e. Inflorescence not sessile; leaves not in a rosette; achene dark. — — ————————— 9. S. deserticola. a. Inflorescence pseudolateral. f. Culms to 2.5 dm. tall, 0.5-1 mm. wide; perianth-bristles 0. g. Achene pale brown or stramineous, plano-convex. .. 1. S. rigidus. g. Achene shiny black or deep brownish-black, trigonous. ............ "—-———————— — 3. S. inundatus. f. Culms 0.5-2 m. tall, 2-10 mm. wide; perianth-bristles present. h. Leaf-blades present; perianth-bristles not plumose. i. Inflorescence a single capitulum; spikelet-scales bilobed or emarginate. EEN — TERR 6. S. americanus. i. Inflorescence anthelate; spikelet-scales not bilobed nor emarginate. ........ eene eee nennen nennen nennen enne 7. S. asper. h. Leaf-blades rudimentary or absent; perianth-bristles plumose- ciliate. EN 10. S. californicus. 1. Scirpus rigidus Boeckl. in Linnaea, 36: 492. (1870). Stiff, rigid perennial, the fasciculate stems to 2.5 dm. tall, more or less angular- terete, striate, glabrous, smooth or scabrid above. Leaves shorter than the culms, mostly less than 1 mm. wide, glabrous, the margins scabri- dulous, apex obtuse, sheath-orifice oblique and attenuate. Inflorescence usually a solitary pseudolateral spikelet subtended by 2 bracts, the lower foliar, appearing as a continuation of the culm, somewhat exceeding the spikelet. Spikelet to 1 cm. long; scales oval, obtuse, the lowermost broadly carinate and muticous, yellowish or stramineous below and warm brown above; bristles 0; stamens 3; style deeply trifid, not enlarged at the base. Achene to 2.3 mm. long, obovoid, plano-convex (or subtrigonous), apex abruptly rounded-obtuse, shiny 1965] Flora of Bolivia — Foster 101 black or brownish-black. LA PAZ: LARECAJA: Caypichuru, 3500- 4200 m., Mandon 1412 (GH). 2. S. cernuus Vahl, Enum. 2: 245 (1806). Dwarf caespitose annual. Culms to 2 dm. high, but usually shorter, filiform, striate-sulcate, smooth. Leaves mostly shorter than the culms, filiform; sheath- orifice oblique, unilaterally produced as a long, broad, obtuse apiculus. Inflorescence of 1(-3) terminal pseudolateral spikelets, subtended by 2 bracts, the lower appearing as a continuation of the culm, hardly or only somewhat exceeding the spikelet. Spikelets to 1 cm. long, usually shorter; scales carinate, the midrib excurrent as a very short blunt apiculus, the midrib green, margins bright reddish-chestnut; bristles 0; stamens (1-) 3; style trifid. Achene obovoid, trigonous, obtuse, with a short apiculus nearly 1 mm. long, obscurely to plainly reticulate. LA PAZ: LARECAJA: near Millipaya, 3200-3700 m., Mandon 1409 (GH); MURILLO: near La Paz, 3300 m., Bang 71 (GH). 3. S. inundatus (R. Br.) Spreng. Syst. Veg. 1: 207 (1825). Perennial from a filiform, branched, spreading rhizome. Culms to 2.5 dm. tall, setaceous. Leaves mostly shorter than the culms, to 1 mm. wide, flat; sheath-orifice oblique and the sheath prolonged unilaterally. Inflo- rescence pseudolateral, of 2-several spikelets aggregated in a capitu- lum, subtended by 2 bracts, the lower erect, exceeding the head and appearing to be a continuation of the culm; spikelets ovoid, 4-12 mm. long; scales ovate, obtuse or mucronulate, somewhat carinate, the keel green, the borders pale or hyaline; bristles 0; stamens 1-2(-3); style trifid. Achene ellipsoidal, trigonous, the apex acute, to 0.6 mm. long, finely or obscurely reticulate, pale brown or stramineous. LA PAZ: LARECAJA: Cochipata, etc., 3000-3800 m., Mandon 1406 (GH); near Acouma, Mandon 1407 (GH). 4. S. boliviensis M. Barros in Darwiniana, 11: 764 (1959). Dwarf perennial from an erect branching rhizome. Culms to 3 cm. high, obtusely trigonous. Leaves few (3-5), shorter than the culms, the sheaths loose, 6-7 mm. long, the blades 4-5 mm. long. Inflorescence a single terminal ebracteate spikelet about 3 mm. in diameter; scales broadly ovate, obtuse, fuscous, at least the lowermost mucronulate; bristles 0; stamens 2; style bifid or trifid. Achene about 1.2 mm. long, suborbicular, the apex abruptly obtuse and contracted into a short neck, plano-convex, slightly reticulate, pale-castaneous or fuscous. PoTosí: CERCADO: Potosí, Petersen & Hjerting 1043b (type-number; not seen). As no material of this species has been available, this description has been based on the original description and figure. 5. S. atacamensis (Phil. Boeckl. in Linnaea, 36: 482 (1870). Perennial, caespitose, from a descending rhizome. Culms to 6 cm. 102 Rhodora [Vol. 67 high. Leaves shorter than the culms, rigid, canaliculate, the margins scabrid on the upper portion. Inflorescence usually a single, terminal, few-flowered spikelet about 6-7 mm. long, the subtending bract not foliar but resembling a scale; scales elliptic-ovate, obtuse, stramineous with brown striations; bristles 0; stamens 3; style trifid. Achene to 1.5 mm. long, plano-convex, obovoid, olivaceous to brownish, shining, the apex abruptly rounded-obtuse. Without data: Bang 1882 (GH). The specimen cited resembles S. rigidus vegetatively, but the achene is that of S. atacamensis. 6. S. americanus var. polyphyllus (Boeckl.) Beetle in Amer. Journ. Bot. 30: 399 (1943). S. americanus subsp. polyphyllus (Boeckl.) T. Koyama in Canad. Journ. Bot. 41: 1118 (1963). Perennial from a rather stout rhizome. Culms to 8 dm. tall, glaucous, acutely trigonous, the sides flat. Leaves few, mostly sheathing, the produced blades seldom equaling the inflorescence, basally flattened, the apex trigonous, pungent; sheath-orifice oblique. Inflorescence pseudolateral, a capit- ulum of 1-6 spikelets, subtended by a bract much exceeding the capitulum and appearing as a continuation of the culm. Scales chestnut-brown, the midrib pale, bilobed with a central mucro equal to or shorter than the lobes; bristles 3-6, exceeding the achene, barbs very large; stamens 3; style bifid (occasionally trifid). Achene obovoid, pale to dark brown, to 2.3 mm. long, plano-convex, apically prolonged into a short neck. COCHABAMBA: CERCADO: near Cocha- bamba, Bang 997 (GH). PoTosí: CHICHAS: road from Quechisla to Chorolque, 3600 m., Cárdenas 36 (GH). 7. S. asper Presl, Rel. Haenk. 1: 194 (1830). Perennial from a rather woody ascending rootstock. Culms to 8 dm. tall, trigonous, the angles scabrid on the upper portion. Leaf-blades long, to 1 cm. wide, often glaucous, the margins scabrid; sheath-orifice truncate to shortly oblique, the margin finely ciliolate. Inflorescence terminal, compound- anthelate, of several to many capitula, subtended by 2-3 foliar in- volucral bracts, at least the lowermost exceeding the inflorescence and appearing as a continuation of the culm. Spikelets fasciculate (occasionally solitary), oblong, to 1.2 em, long; scales ovate, broadly carinate, mucronate, reddish-brown, the keel green, the margin ciliate, especially at the apex; bristles 6, of varying lengths, the longest exceeding the achene; stamens 3; style trifid. Achene ovoid to obovoid, trigonous, to 1.2 mm. long, the apex prolonged into a short neck, smooth, pale (almost white) to brownish. LA PAZ: MURILLO: La Paz, 3460 m., Buchtien 529 (GH); Cotana, 2450 m., Buchtien 3141 (GH); Capi, Bang 765 (GH). 8. S. acaulis Boeckl. in Linnaea, 36: 494 (1870). Acaulescent perennial from a slender creeping rhizome. Leaves numerous in a flattened rosette, the basal sheath large and much dilated, the apex 1965] Flora of Bolivia — Foster 103 more or less obtuse, canaliculate, to 2.5 cm. long, narrow. Inflorescence a sessile capitulum of several to many spikelets, about 7 mm. long. Spikelets oblong, many-flowered (about 8); scales densely aggregated, to 6 mm. long, lance-elliptic, the back green, the sides brownish, apex acute; bristles 5-6, whitish; stamens 3; style long, trifid. Achene obovoid, trigonous, to 1.5 mm. long, the apex acuminate, pale in color, smooth. LA Paz: MUÑECAS: cordillera above Carabuco, 4200 m., Buchtien 6388 (vus). 9. S. deserticola Phil. Fl. Atac. 53 (1860). Dwarf, densely caespitose perennial from creeping, scaly, slender, much-branched rhizomes. Culms to 10 cm. tall, usually shorter. Leaves very short, much shorter than the culms, the membranous sheathing base somewhat enlarged, blades to 2 cm. long, scabrid on the margins. Inflorescence a solitary terminal (not pseudolateral) capitulum about 1 cm. long, of 2-3 (rarely 1) spikelets, the involucral bracts shorter than the capitulum. Spikelets ovoid-oblong; scales ovate or lance-ovate, the margins pale brown, the broad midrib stramineous to greenish, somewhat excurrent; bristles 3-6, the longest exceeding the achene; stamens 3; style trifid. Achene ellipsoid, trigonous, to 1.5 mm. long, brown, the apex prolonged almost like an apiculus, the surface finely reticulate-scrobiculate. LA PAZ: LARECAJA: Lacatia, etc., Mandon 1417 (GH). PoTosi: CHICHAS: near Chorolque, 3500 m., Cárdenas 62 (GH). 10. S. californicus (C. A. Mey.) Steud. Nomencl. (ed. 2) 2: 538 (1841). S. riparius of the Catalogue. Perennial from a thick hori- zontal rhizome. Culms to 1.8 m. high, obtusely trigonous, to 1 cm. wide, smooth. Leaves reduced to thin membranous sheaths, the upper- most with rudimentary blades. Inflorescence a large, terminal, compound anthela, subtended by a single large involucral bract to 5 cm. long, appearing as a continuation of the culm, the inflorescence thus pseudolateral. Spikelets solitary at ends of secondary rays, to 1.8 em. long; scales broadly ovate to obovate, to 4 mm. long, the apex entire or emarginate, ciliate, long-mucronate, brown or reddish-brown, the midrib and mucro green; bristles 3, plumose-ciliate, equaling the achene; stamens 3; style bifid. Achene to 2.25 mm. long, obovoid, plano-convex, the apex produced as a short apiculus, grayish to brownish, finely scrobiculate. TARIJA: ARCE: Bermejo, 1400 m., Fiebrig 2311 (GH). 5. FUIRENA Rottb. Fuirena umbellata Rottb. Descr. & Icon. 70, t. 19, fig. 3 (1778). Perennial from a creeping rhizome. Culms to 8 dm. tall. Leaves cauline, basally long-sheathing, the linear-lanceolate blades to 18 cm. long and 1.2 cm. wide, acute, scabrid above and beneath, the ligule puberulent or shortly pubescent. Inflorescence of terminal and axillary corymbs. Spikelets few- to many-flowered, on villous peduncles and 104 Rhodora [Vol. 67 pedicels, oblong-ovoid, acute, in groups of 3-15, to 7 mm. long. Glumes obovate, red-brown, with 3 prominent green dorsal nerves, the upper half pubescent, 2 mm. long, terminated by a short, thick, straight mucro originating shortly below the apex. Perianth of 3 membranous, flattened, stipitate, ovate, aristate, 3-nerved scales, alternating with 3 bristles (or these absent), surrounding 8 stamens and an obovoid trigonous achene crowned with the persistent base of the trifid style. LA Paz: CAUPOLICÁN: Apolo, R. S. Williams 917 (GH); Ixiamas, 330 m., Cárdenas (M. E.)* 1950 (GH). SANTA CRUZ: SARA: Buena- vista, 500 m., Steinbach 5217 (GH). 7. ELEOCHARIS R. Br. See Svenson in Rhodora, 31: 121-135, 151-163, 167-191, 199-219, 224-242 (1929); 34: 193-2083, 215-227 (1932) ; 35: 377-389 (1934) ; 39: 210-231, 236-273 (19377) ; 41: 1-19, 43- 77, 90-110 (1939). Annual or perennial leafless aquatics or semi-aquatics. Culms septate or not septate, angled or terete, with bladeless basal sheaths. Inflorescence a single terminal spikelet, few- to many-flowered, the lowest scales empty, the central scales with perfect flowers, the upper- most scales staminate or sterile. Perianth present as hypogynous bristles (rarely absent), the bristles retrorsely barbed or smooth. Stamens 1-3. Style bifid or trifid, glabrous, the bulbous thickened base persistent on the achene as a tubercle. Achene lenticular, biconvex or trigonous. a. Culms septate. b. Culms 1-2 mm. wide; spikelet much wider than the culms. ........ "——————————— 10. E. nodulosa. b. Culms 5-10 mm. wide; spikelet hardly or not at all wider than the culm. c. Achene 2 mm. long, longitudinally ribbed and pitted, the cells rectangular; style-base broadly pyramidal. .... 2. E. interstincta. c. Achene 1.5 mm. long, scrobiculate; style-base lanceolate, half the length of the achene. EN 11. E. elegans. a. Culms not septate. d. Sheath oblique at orifice. e. Achene apically narrowed to a neck. f. Neck of achene broadened upward. ................ 1. E. fistulosa. f. Neck not broadened upward. ...................ssss 3. E. radicans. e. Achene lacking an apical neck. g. Achene black or deep brownish-black. ........ 6. E. geniculata. *The initials M. E. refer to specimens collected on the Mulford Bio- logieal Exploration. The number-series of these collections duplicate other number-series of the same collectors. 1965] Flora of Bolivia — Foster 105 g. Achene white, greenish or brownish. h. Achene obovoid, greenish to brownish, 1.5 mm. long. ........ mr Dp Puce MO gee ne eRe ee e 9. E. albibracteata. h. Achene ovoid, white to brownish, 1 mm. long. i. Scales lance-ovate, acute, not emarginate at apex. ....... PIG ADOBE NL ENIRO SET ea Tae LM 12. E. minima. i. Seales mostly oblong, obtuse, at least some emarginate e ese RN IBN IBEEH 13. E. filiculmis. d. Sheath truncate at orifice. j. Sheath bifid at apex. E Achene shiny black. ................. eee 4. E. maculosa. k. Achene greenish-yellow to olivaceous, never black. l. Achene olivaceous, with a short apical neck. ...................... e ANUS 5. E. Sellowiana. l. Achene greenish-yellow, without an apical neck, ................ E 8. E. crinalis. J- Sheath not bifid at apex; nnana a. 7. E. Dombeyana. 1. Eleocharis fistulosa (Poir.) Schult. Mant. 2: 89 (1824). Culms not septate, acutely trigonous, to 6 dm. high, the coarse roots reddish- brown; sheaths thin, membranous, loose, the orifice oblique, acute, pale reddish-brown. Spikelets to 3 (-3.5) em. long, as wide as the culm, or sometimes a little wider; scales to 4 mm. long, broadly ovate, subacute or obtuse, firm, conspicuously striate, stramineous to gray, the very narrow, thin, white margin erose; stamens 3; style trifid. Achene about 2 mm. long, obovoid, greenish to light brown, with about 20 rows of deep quadrangular to oblong-hexagonal cells, apex narrowed to a neck and broadened upward to the style-base; bristles 6, exceeding the achene. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7444 (GH). 2. E. interstincta (Vahl) Roem. & Schult. Syst. Veg. 2: 149 (1817). Culms septate, terete, to 10 dm. high and 5 mm. wide, from a short thick caudex with coarse red-brown or light brown roots; sheaths membranous, the orifice oblique and acute. Spikelet to 4 cm. long, more or less cylindrical, as wide as or a little wider than the culm; scales oblong, obtuse to subacute, striate, stramineous to gray, with a rather broad, thin, white-hyaline, erosulous margin; stamens 3; style bifid or trifid. Achene 2 mm. long, yellow to gray, longitudinally ribbed and pitted, the cells transversely rectangular and prominent, narrowed to a marked apical annulus (but not narrowed into a neck), the style-base broadly pyramidal, dark brown; bristles 6, exceeding the achene, broad. BENI: YACUMA: near Lake Rogagua, 330 m., Rusby (M. E.) 1422 (GH). SANTA CRUZ: SARA: Río Curichi, 450 m., Steinbach 1917 (GH); Buenavista, 450-500 m., Steinbach 7445 (GH), 7043bis (GH), 5216 (GH). 3. E. radicans (Poir.) Kunth, Enum. Pl. 2: 142 (1837). E. exigua and E. costulata of the Catalogue. Perennial from filiform creeping 106 Rhodora [Vol. 67 rootstocks. Culms to 1 dm. tall, filiform or capillary, not septate, often recurved, somewhat angular; sheath greenish-scarious, the orifice oblique and acute. Spikelet to 3 mm. long, wider than the culm, scales greenish, often with a broad purple margin, striate, lance- oblong to oblanceolate, obtuse to subacute; stamens 2; style trifid. Achene 1 mm. long, obovoid, yellowish-green to nearly white, obscurely trigonous with about 15 elevated longitudinal ribs, the inter-costal areas about 40-trabeculate, the cells narrowly rectangular, the apex constricted into a neck, but not dilated upward into the dark-green acute style-base; bristles 2 (or 0), white. LA PAZ: LARECAJA: Mapiri, Bang 1503 (GH); between Cochipata and Millipaya, 3100 m., Mandon 1413 (GH). 4. E. maculosa (Vahl) Roem. & Schult. Syst. Veg. 2: 154 (1817). Perennial, with long, slender, scaly, chestnut-brown stolons. Culms to 3.5 dm. long, not septate, more or less quadrangular and canaliculate, smooth, rigid and erect; uppermost sheaths scarious at apex, enlarged, rugose, the orifice truncate, bifid. Spikelet to 1.2 em. long, much broader than the culm, lanceolate to ovoid; scales broadly ovate, obtuse, firm-textured, shining, purplish-brown, the scarious margin erosulous; stamens 3; style bifid. Achene to 1 mm. long, obovoid, shiny black, the surface minutely striatulate, narrowed apically to a very short neck, this not enlarged below the style-base; style-base dilated at base, half the achene-width, light-brown, the beak subulate; bristles 7-8, unequal, at least some equaling the achene, teeth small, numerous. LA PAZ: CAUPOLICAN: Apolo, R. S. Williams 914 (Sven- son). 5. E. Sellowiana Kunth, Enum. Pl. 2: 149 (1837). E. ocreata var. flaccida of the Catalogue. Annual, with rather pale slender roots. Culms not septate, to 1.5 dm. tall, or occasionally to 3.5 dm., to 2 mm. wide, striate, slightly constricted below the spikelet; sheaths hyaline at the truncate unilaterally bifid apex. Spikelet to 1 cm. long, wider than the culm, ellipsoid; scales oblong, obtuse, stramineous (or some- what pale yellow-green), with a narrow brown stripe on each side of the scarcely keeled midrib; stamens 3; style bifid. Achene to 1 mm. long, obovoid, lenticular, olivaceous, shiny, puncticulate-striate, the striae sometimes dark or blackish, narrowed abruptly at the apex into a very short neck, style-base shortly conic, yellow-green, less than half the width of the achene; bristles 7-8, white, shorter than the achene. LA Paz: CAUPOLICAN: Apolo, R. S. Williams 909 (Svenson). SANTA CRUZ: SARA: Curiche de Piedritas, 450 m., Steinbach 2870 (GH). 6. E. geniculata (L). Roem. & Schult. Syst. Veg. 2: 150 (1817). E. capitata and E. geniculata of the Catalogue. Annual (occasionally stoloniferous ?). Culms to 4 dm. high, not septate, more or less terete, sulcate; sheaths basally brown, stramineous above, the orifice oblique and subacute. Spikelets to 0.8 cm. long, ovoid to subglobose, much 1965] Flora of Bolivia — Foster 107 wider than the culms; scales suborbicular to ovate, obtuse, yellow to pale brown with a very narrow, poorly defined, hyaline margin; stamens 2-3; style bifid. Achene about 1 mm. long, obovoid, more or less biconvex or obscurely trigonous, not narrowed into an apical neck, shiny, deep brownish-black or black, the style-base mostly depressed, whitish to brownish; bristles 6-8 (or 0), brown, broad-based, the barbs conspicuous. BENI: YACUMA: near Lake Rogagua, Mulford Explo- ration 2519% (GH). LA PAZ: SUR YUNGAS: cataracts of the Rio Bopi, 1000 m., Rusby (M. E.) 738 (GH). SANTA CRUZ: SARA: Rio Perdix, 450 m., Steinbach 7454 (GH). 7. E. Dombeyana Kunth, Enum. Pl. 2: 145 (1837). Perennial from slender, scaly, red-brown rhizomes. Culms to 3 dm. tall, not septate, subterete, sulcate; sheaths basally deep brown, stramineous at the truncate apiculate orifice. Spikelets to 1 cm. long, ovoid to lance-linear, much wider than the culms; scales elliptic to ovate, obtuse, brown, the midrib yellow or green, the margin hyaline (at least on the lowest scales); stamens 3; style trifid. Achene to 1.3 mm. long, obovoid, trigonous, not narrowed to an apical neck, shiny, yellow or brown, the surface faintly reticulate, the style-base elongate, acute (mucroni- form); bristles 4, brown, longer or shorter than the achene. LA PAZ: MURILLO: La Paz, Buchtien 149 (GH), 8824 (GH), Bang 144 (GH). COCHABAMBA: CERCADO: near Cochabamba, Bang 996 (GH). 8. E. crinalis (Griseb.) C. B. Clarke in Kew Bull. Add. Ser. 8: 23 (1908). E. boliviana and E. Brehmeriana of the Catalogue. Perennial. Culms filiform, in remote clumps from a long, slender, scaly rhizome, to 2 dm. high (rarely higher), not septate, sulcate, angled; sheaths reddish or reddish-brown at the base, the apex darker or lighter, orifice truncate and briefly bifid. Spikelets to 6 mm. long, ovate to elliptie, much broader than the culms; scales ovate to lanceolate, mostly obtuse to subacute, not (or hardly) carinate, deep chestnut- brown to reddish, the midrib paler; stamens 3; style trifid. Achene to 1 mm. long, greenish-yellow, ovoid to suborbicular, obscurely trigonous, the 3 angles thickened, longitudinally striatulate, faintly reticulate, style-base acuminate-deltoid, brownish; bristles pale-brown, slender, about equaling the achene. LA PAZ: MURILLO: Cotana, 2000-2900 m., Mandon 1416 in part (GH; type-number of E. Brehme- riana), 2450 m., Buchtien 3143 (GH); La Paz, 3750 m., Buchtien 4482 (GH; type-number of E. boliviana). TARIJA: ARCE: Padcaya, 2100 m., Fiebrig 2524 (GH). 9. E. albibracteata Nees & Meyen ex Kunth, Enum. Pl. 2: 143 (1837). E. nubigena of the Catalogue. E. albibracteata var. nubigena (C. B. Clarke) M. Barros in Gen. & Sp. Pl. Argent. 4(1): 179 (1947). Perennial from a rather woody spreading rootstock. Culms filiform, mostly less than 1 dm. long; sheath-orifice oblique, acute. ‘Spikelets to 6 mm. long, wider than the culms; scales ovate, dark brown, the 108 Rhodora [Vol. 67 margin hyaline, entire, the midrib green, obtuse to acute; stamens 3; style trifid. Achene to 1.5 mm. long, obovoid, trigonous, the angles thickened and broad, greenish to brownish, faintly striatulate or reticulate, style-base pyramidal to conic, with a basal ridge at junction with the achene; bristles 3-4, slender, exceeding the achene, occasional- ly shorter. LA PAZ: LARECAJA: Cochipata, etc., 3100-4100 m., Mandon 1414 in part (Gray Herbarium sheet mixed with a very dwarf Carex) ; MURILLO: La Paz, 3300 m., Bang Tla (GH). The variety nubigena seems to differ from the species only in the orifice of the sheath, and even this character seems somewhat variable. 10. E. nodulosa (Roth) Schult. Mant. 2: 87 (1824). Perennial from a creeping rhizome, Culms to 8 dm. tall, terete, septate, the septations approximate; sheaths long, stramineous, the orifice truncate, uni- laterally apiculate. Spikelets to 2.7 cm. long, lance-oblong, wider than the culms; scales suborbicular to ovate, acute to obtuse, brown, the midrib often lighter in color, with a broad hyaline margin and apex; stamens 2; style bifid or trifid. Achene 1 mm. long, obovoid, biconvex, yellowish, brownish or olivaceous, regularly scrobiculate, the brown style-base flattened-deltoid, with a slight elevation at junction with achene; bristles equaling or somewhat exceeding the achene, red- brown. LA PAZ: CAUPOLICAN: Apolo, R. S. Williams 911 (GH); NOR YUNGAS: Hacienda El Choro, Coripata, Buchtien 8065 (NY). COCHA- BAMBA: TOTORA: Pocona, 2500 m., Steinbach 8710 (GH). SANTA CRUZ: SARA: Buanavista, 500 m., Steinbach 5298 (GH). ll. E. elegans (HBK.) Roem. & Schult. Syst. Veg. 2: 150 (1817). E. geniculata of the Catalogue. Coarse perennial from a creeping rootstock. Culms septate, terete, to 1.5 m, high, to 1 cm. wide; sheaths reddish or pale, the orifice truncate and unilaterally briefly apiculate. Spikelets to 3 em. long, subglobose to lanceolate to cylindrical, about as wide as the culm or a little wider; scales oblong to obovate, 2 mm. long, obtuse, not carinate, the midportion brown, the broad scarious margin pale brown to whitish; stamens 3; style bifid or trifid. Achene obovoid, 1.5 mm. long, biconvex or obscurely trigonous, the apex narrowed into a short neck, yellow to brown, more or less regularly scrobiculate, style-base dark brown, lanceolate, half the length of the achene; bristles exceeding the achene, dark brown. BENI: YACUMA: Lake Rogagua, Rusby 1602 (Svenson). 12. E. minima Kunth, Enum. Pl. 2: 139 (1837). Matted caespitose plants with whitish fibrous roots. Culms to 7 cm. tall, capillary, angular-sulcate, sheaths oblique at the orifice. Spikelets to 4 mm. long, much wider than the culms; scales lance-ovate, acute, dark brown, the midrib green, margin broad, hyaline, white; stamens 3; style trifid. Achene to 1 mm. long, ovoid, trigonous, apically narrowed 1965] Flora of Bolivia — Foster 109 into a very brief neck, subreticulate to striatulate, whitish to olivaceous to brown, style-base short-pyramidal, gray or brown; bristles mostly whitish, not equaling the achene, obscurely barbed. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 2208 (GH), 5499 (GH). 13. E. filiculmis Kunth, Enum. Pl. 2: 144 (1837). E. sulcata of the Catalogue. Perennial from an ascending caudex. Culms to 4 dm. tall, not septate, flattened, sulcate, slender, sometimes to 1 mm. wide; sheath purple-brown to stramineous, the orifice oblique. Spikelets to 1 em. long, ovoid to cylindric, much wider than the culms; scales more or less oblong, obtuse, often emarginate, stramineous to rufescent, the keel lighter, the broad margin scarious; stamens 3; style trifid. Achene 1 mm. long, ovoid, trigonous, shining white, subreticulate or striatulate, style-base pyramidal, almost as wide as the achene, the basal margin overhanging the achene; bristles about 6, white, nearly equaling the achene. COCHABAMBA: MIZQUE: near Vilavila, 2500 m., Eyerdam 24970 (GH). SANTA CRUZ: SARA: Dolores, 450 m., Steinbach 1900 (GH); Buenavista, 450-500 m., Steinbach 5340 (GH), 6946 (GH). 8. FIMBRISTYLIS Vahl Annual or perennial. Leaves filiform or linear. Inflorescence anthelate, with few to many heads, or reduced to a single terminal head. Spikelets several- to many-flowered, terete or ellipsoid (ovoid in one case), the glumes all, or mostly, floriferous, imbricate, several- ranked. Flowers perfect; perianth absent; stamens 3; style dilated at the base and abruptly constricted above the achene, bifid or trifid, not persistent on the achene; achene basally attenuate, apically short- apiculate, biconvex or trigonous, regularly ribbed and cancellate or tuberculate. Inflorescence anthelate, with several to many spikelets; achenes not tuberculate. Achenes 1 mm. or more in length, strongly ribbed and cancellate. A NE EUER EE 1. F. annua. Achenes less than 0.5 mm. long, not cancellate, the ribbing obscure. 2. F. aestivalis. Inflorescence reduced to a single head; achenes tuberculate, not (25 5 5 E 3. F. monostachya. 1. Fimbristylis annua (All) Roem. & Schult. Syst. Veg. 2: 95 (1817). Annual, to 8 dm. tall, glabrous, or in some forms the leaf- sheaths and involucral bracts pubescent. Inflorescence anthelate, involucrate, the lower bract as long as or much exceeding the anthela. Spikelets to 9 mm. long, elliptic, acute, the ovate glumes apiculate, not conspicuously keeled. Style bifid, ciliate on at least the upper portion. Achene 1 mm. long, or a little more, obovoid, biconvex, cancellate, with about 6 prominent longitudinal ribs on -each face, the intercostal pitting regular and pronounced. YUNGAS: Bang 530 (GH). BENI 110 Rhodora [Vol. 677 (2): Río Beni, Rusby 71 (GH). LA Paz: LARECAJA: San Carlos, Mapiri, 600 m., Buchtien 304 (GH); NOR YUNGAS: Coripata, 1300 m., Buchtien 8066 (NY). SANTA CRUZ: SARA: Buenavista, 450 m., Stein- bach 1035 (GH), 1071 (GH), 5141 (GH), 5337 (GH), 5466 (NY). BENI: YACUMA: Hacienda Rosario, near Lake Rogagua, 330 m., Rusby (M. E.) 1655 (GH). 2. F. aestivalis (Retz.) Vahl, Enum. 2: 288 (1806). Annual, with fibrous roots. Culms to 2.5 dm. high, filiform. Leaves filiform or very narrowly linear, mostly shorter than the anthelate inflorescence. Involucre of 2 bracts, the lower equaling or exceeding the inflorescence, the other very short. Glumes rather narrowly ovate, 1-1.5 mm. long, carinate, the keel green and excurrent as a short tooth. Style bifid. Achene less than 0.5 mm. long, pyriform, lenticular or obscurely trigonous, evenly papillose. BENI (?): Río Beni, Rusby 287 (GH). Note: the typical form of this species is somewhat pubes- cent or pilose. This glabrous specimen was determined by H. K. Svenson. 3. F. monostachya (L.) Hassk. Pl. Jav. Rar. 61 (1848). Perennial, caespitose, or with a short rhizome, to 3 dm. tall. Leaves flat or canaliculate, shorter than the culms, acute, the margins scabrid. Inflorescence reduced to a single ovoid head (rarely 2), to 1.5 cm. long. Involuere of a single reduced bract, shorter than the head. Lowermost glumes sterile and distichous, the fertile upper glumes spirally inserted on the axis; glumes to 5-6 mm. long, naviculate, carinate, the prominent green keel excurrent as a stout mucro. Style trifid, triangular, the angles densely pectinate-ciliate from base to top. Achene to 2.5 mm. long, somewhat stipitate, pyriform-globose or trigonous-globose, white or yellowish, tuberculate. BENI: YACUMA: near Lake Rogagua, 330 m., Cárdenas (M. E.) 1395 (GH). LA Paz: NOR YUNGAS: Milluguaya, 1900 m., Buchtien 4177 (GH). TARIJA: ARCE: Padcaya, 2100 m., Fiebrig 2553 (GH). 9. BULBOSTYLIS Kunth Annual or perennial, the slender, often filiform, culms usually caespitose, leafless, glabrous. Leaves filiform, the sheaths ciliate, especially at the orifice, cilia sometimes absent. Inflorescence mono- cephalous or anthelate, or the spikelets solitary or fasciculate, the scales often puberulent; flowers perfect. Perianth-bristles 0; stamens 2-3; style glabrous, trifid, the base persistent as a tubercle on the achene. Achene trigonous or compressed, smooth, puncticulate or undulate-rugose. a. Inflorescence monocephalous. b. Inflorescences terminal and basal. ............................ 1. B. Funckü. 1965] Flora of Bolivia — Foster 111 b. Inflorescences terminal only. c. Inflorescence a single obconic spikelet. ................ 2. B. paradoxa. c. Inflorescence not a single spikelet. d. Achene puncticulate but not rugulose. .. 3. B. sphaerocephala. d. Achene transversely rugulose. ........................ 4. B. juncoides. a. Inflorescence anthelate. e. Leaves and sheaths densely puberulent. ................ 6. B. hirtella. e. Leaves and sheaths glabrous, but scabrid on the margins. f. Achene puncticulate. g. Rhizome present, very short; cilia at sheath-orifice brown. c Kc EE 8. B. junciformis. g. Rhizome absent; cilia at sheath-orifice white, curling. ........ BN IU AU nh opener oye E 5. B. papillosa. f. Achene transversely rugulose (occasionally quite smooth). h. Spikelets solitary. ................. eere eene 7. B. capillaris. h. Spikelets fascicled (3-5). .... 4. B. juncoides var. ampliceps. 1. Bulbostylis Funckii (Steud.) C. B. Clarke in Kew Bull. Add. Ser. 8: 26 (1908). Annual (?). Culms to 6 cm. high, capillary. Leaves filiform-capillary, shorter than culms, occasionally exceeding the inflorescence; sheath white-ciliate at the orifice. Inflorescence of 2 kinds: terminal solitary spikelets, and basal spikelets. Terminal inflorescence to 4 mm. long, the subtending bract small, usually shorter than the spikelet; scales ovate, somewhat carinate and shortly mucronate; achene less than 1 mm. long, trigonous, transversely rugulose, yellowish-white to very pale brown, the style-base a small flattened brown disc with a minute central apiculus, Basal inflo- rescence: several flowers enclosed in the bases of modified, rather hyaline leaf-sheaths; achenes about 1.5 mm. long, more markedly rugulose than the terminal achenes. LA PAZ: LARECAJA: Cerro del Iminapi, 2700 m., Mandon 1410 (GH). At first glance this extraordinary little plant may appear to be intermixed with Jsoétes, for the basal achene-groups bear a resemblance to the megasporangia of Isoëtes. 2. B. paradoxa (Spreng.) Lindm. in Bih. Sv. Vet.-Akad. Handl. 26, Afd. 3, no. 9: 17 (1900). Small, densely caespitose perennial from a short woody rhizome. Culms to 1.5 dm. high, but usually shorter. Leaves shorter than the culms, or sometimes nearly equaling them, narrow, obtuse, canaliculate, the base somewhat enlarged, sheath papery, open, densely long-ciliate with white curling hairs. Inflo- rescence a solitary, terminal, obconic spikelet to 1 cm. long, subtended by several flattened, narrow, long-acuminate bracts, these equaling or shorter than the spikelet; scales lanceolate, the thick green midrib long-excurrent, the basal margins thin, brownish, densely long-ciliate; stamens 2. Achene to 2 mm. long, obovoid, trigonous, the thickened obtuse angles pale, finely rugulose, pale to darkish brown, rather 112 Rhodora [Vol. 67 shining. LA PAZ: CAUPOLICÁN: Altunkama, 2600 m., Cárdenas 5679 (US). 3. B. sphaerocephala (Boeckl.) C. B. Clarke in Bull. Herb. Boiss. (ser. 2) 3: 1018 (1903). Caespitose perennial from a very short rhizome. Culms to 6 dm. high, rounded below, quadrangular above. Leaves filiform, flexuose, mostly shorter than the inflorescence, but occasionally longer, puberulent, ciliate; sheaths sparsely to copiously long-ciliate or even fimbriate at the orifice. Inflorescence a single more or less globose head of several to numerous spikelets, the heads to 1.2 cm. wide, subtended by 3 conspicuous puberulent involucral bracts (reflexed at maturity), the lowermost to 3 em. long. Spikelets oblong, acute, to 4-5 mm. long, with 6-7 flowers; scales orbicular to oval, reddish-brown, ciliolate, mucronulate, to 2.5 mm. long; stamens 3. Achene whitish to brownish, obovoid, trigonous, to 0.7 mm. long, finely puncticulate, style-base flattened, with a dark center. SANTA CRUZ: SARA: Buenavista, Steinbach 6805 (Barros). 3a. B. sphaerocephala var. macrocephala Kükenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2) 3: 186 (1931). Taller, with a larger head of spikelets. Achene to 0.9 mm. long, grayish, the style-base dark. SANTA CRUZ: SARA: Buenavista, 400 m., Steinbach 6806 in part (as to Gray Herbarium sheet). 4. B. juncoides (Vahl) Kükenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2) 3: 185 (1931). Annual (?). Culms to 8 dm. tall, caespitose, obscurely quadrangular, scabrid. Leaves filiform-setaceous, scabrid on the veins and margins, shorter than the culms. Inflorescence a single terminal head of relatively few spikelets (up to 8, rarely more), to 1 cm. in diameter but usually narrower, subtended by 3 involucral bracts, the longest exceeding the head, puberulent, the base enlarged, long-ciliate. Spikelets oblong-ovate, to 5 mm. long, with 6-8 flowers; scales dark chestnut-brown, puberulent (at least the lower ones), ciliate, mucronate; stamens 2-3. Achene obovoid, 1 mm. long, trigonous, finely undulate-rugulose, light brown or grayish, the style- base the same color as the achene. LA PAZ: LARECAJA: near Sorata, 2900 m., Mandon 1408 (GH). 4a. B. juncoides var. ampliceps Kükenth. ex Osten in Anal. Mus. Hist. Nat. Montevideo (ser. 2) 3: 188 (1931). B. sphaerolepis of the Catalogue. Mostly smaller than the species, the stems smooth, the inflorescence a contracted anthela of 3-5 spikelets sessile and 2 fas- cicles of 3-5 spikelets terminal on the rays, the involucral bracts mostly glabrous. Spikelets 4-6 mm. long; scales asymmetrical, cilio- late near the apex. Achene pale bluish-gray to whitish. La PAZ: NOR YUNGAS: Milluguaya, 1100 m., Buchtien 4178 (GH); Polo-Polo, 1100 m., Buchtien 3649 (GH); SUR YUNGAS: Sirupaya, 2100 m., Buchtien 401 (GH). YUNGAS: Rusby 67 (GH), Bang 306 (GH). SANTA CRUZ: SARA: Buenavista, Steinbach 3544 (GH). 1965] Flora of Bolivia — Foster 113 5. B. papillosa Kükenth. in Fedde, Rep. Sp. Nov. 23: 198 (1926). Annual, caespitose. Culms to 8 dm. tall, basally terete, apically obscurely trigonous, smooth. Leaves very narrowly linear to filiform, to 1.5 mm. wide, shorter than the culms, the margins scabrid; sheath- orifice very long-ciliate with white curling hairs. Inflorescence a terminal anthela with up to 9 rays, subtended by 3-4 involucral bracts mostly shorter than the rays. Spikelets lanceolate, to 6 mm. long, sessile, in dense fascicles of 5-8; scales lance-ovate, dark reddish- brown, mucronate, glabrous or puberulent, to 2 mm. long; stamens 3. Achene oblong-obovoid, 1 mm. long, trigonous, grayish-white, minutely papillate-puncticulate. LA PAZ: SUR YUNGAS: Sirupaya, 2200 m., Buchtien 6929 (GH). SANTA CRUZ: CHIQUITOS: El Poston, 506 m., Cárdenas 4500 (Us). 6. B. hirtella (Schrad. ex Schult.) Urb. Symbol. Antill 2: 166 (1900). Annual. Culms to 4.5 dm. tall, filiform, scabridulous. Leaves much shorter than the culms, filiform-setaceous, shortly pubescent, the Sheaths with copious long white ciliation at the orifices. Inflorescence a terminal compound anthela, somewhat compressed, with 4-6 unequal spreading rays, subtended by 2-3 setaceous involucral bracts, the longest exceeding the anthela. Spikelets to 3-4 mm. long, lance-ovate; scales dark red-brown, mucronate or mucronulate, ciliolate, to 1.2 mm. long; stamens 3. Achene to 0.8 mm. long, oblong-obovoid, trigonous, finely puncticulate or somewhat transversely rugulose, the style-base more or less globose. SANTA CRUZ: CORDILLERA: Puerto Suárez, Etchichury (Barros). 7. B. capillaris (L.) C. B. Clarke in Hook. Fl. Brit. Ind. 6: 652 (1893). Annual, caespitose. Culms to 3.5 dm, high, filiform, glabrous or scabrid. Leaves much shorter than the culms, filiform-setaceous, the margins scabrid, sheath-orifice long-ciliate. Inflorescence anthelate, with 2-5 principal rays, subtended by 2 foliar bracts, the longer usually much exceeding the inflorescence. Spikelets solitary, oblong- ovoid, to 6 mm. long, 6-15-flowered; scales lance-ovate, the upper margins ciliolate, the keel prominent, strongly serrate-scabrid, acute or mucronulate, to 2 mm. Jong, blackish-brown, finely puberulent; stamens 2 (-3). Achene to 0.8 mm. long, trigonous, obovoid to cordi- form, pale brown, slightly or not at all transversely rugulose, the style-base dark. SANTA CRUZ: SARA: Buenavista, Steinbach 1913 (GH), 5186 (NY). . Ta. B. capillaris var. tenuifolia (Rudge) C. B. Clarke in Urb. Symbol. Antill. 2: 89 (1900). B. tenuifolia of the Catalogue. Plants to 4.5 dm. tall. Involucral bracts shorter than inflorescence. Spikelets narrower (about 1 mm. wide). Achene to 0.9 mm. long, smooth or somewhat transversely rugulose. LA PAZ: LARECAJA: San Carlos, Mapiri, 950 m., Buchtien 324 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 5201 (Ny, Us), 6804 (GH), 6989 (GH). 114 Rhodora [Vol. 67 8. B. junciformis (HBK.) C. B. Clarke in Trans. Linn. Soc. (ser. 2) 4: 512 (1895). Caespitose, with a very short rhizome. Culms to 8 dm. tall, glabrous. Leaves much shorter than the culms, filiform, smooth or the margins slightly scabridulous, sheath-orifice ciliate with brown hairs. Inflorescence anthelate, compound but condensed, the involucral bracts usually shorter than the inflorescence, not ciliate or only slightly so at the base. Spikelets fasciculate, to 8 mm. long, oblong. Scales ovate, dark brown, minutely puberulent, the midrib somewhat ex- current. Achene obovoid to cuneate, not rugulose but finely puncticu- late, pale brown to white, the style-base dark. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1903 (GH), 5185 (GH). Bulbostylis boliviana Palla in Oesterr. Bot. Zeitschr. 59: 191 (1908). I have seen no material of this species, and the original description does not enable me to place it. Conse- quently, it has been omitted from the key and from the formal treatment of the genus. 10. CLADIUM P. Br. Cladium jamaicense Crantz, Inst. 1: 362 (1766). Perennial, to 1-3 m. tall. Leaves long, to 1 cm. wide, flat, the margins and the midrib beneath scabridulous. Involucre of several bracts, the longest shorter than the inflorescence. Inflorescence to 9 dm. long, paniculate, the numerous spikelets small, red-brown, acute, fasciculate at the ends of the rays. Perianth absent; lowest glumes short, sterile; 1 or 2 above subtending staminate flowers, the terminal flower perfect, its glume to 5 mm. long, acute. Style bifid (or trifid) ; achene obovoid, the base truncate, apical tubercle not present. SANTA CRUZ: SARA: Cerro de Amboró, 1000 m., Steinbach 3007 (GH). 11. RHYNCHOSPORA Vahl (nomen conservandum) (including Dichromena and Pleurostachys) See Kükenthal in Bot. Jahrb. 74: 375-509 (1949); 75: 90-195 (1950), 2773-314 (1951), 451-484 (1952). Perennial from rhizomes. Culms mostly triangular and leaf- bearing. Leaves long-sheathing, flat, sometimes subplicate. Inflo- rescence paniculate, composed of smaller panicles or corymbs, or often reduced to 1-3 rather dense glomerulate heads. Spikelets with the lowermost scales empty, the middle scales subtending perfect flowers, the uppermost bearing staminate or sterile flowers. Perianth present as scabridulous or smooth hypogynous bristles, or absent; stamens 2-3; style long, shortly or deeply bifid, the base persistent on the achene. Achene orbicular, obovoid, ovoid or oblong, biconvex, trans- versely rugulose, or finely cancellate, or scrobiculate, or smooth, usually shining. 1965] Flora of Bolivia — Foster 115 a. Culms monocephalous. b. Bracts foliar, long, much exceeding the heads. c. Style-base bluntly triangular, not basally lobed. ....................... eee: mete a E A TII EE 22. R. nervosa. c. Style-base flattened, 2-lobed basally, the lobes decurrent on the EUR MUS EE 23. R. radicans. b. Bracts not foliar, short, not equaling the heads. d. Bristles 3, not at all plumose. ...................... 12. R. hirta. d. Bristles 2-5, plumose on lower half. ..................-. 11. R. globosa. a. Culms not monocephalous. e. Bristles present. f. Bristles not plumose. g. Achene pitted, scrobiculate or cancellate, but not transversely rugulose. h. Achene pitted or scrobiculate. i. Spikelet many-flowered. j. Leaf-blade to 2 cm. wide, the base petiolately at- tenuate. .......— eno oenenroos tia Dr DA 1. R. umbraticola var. Kuntzei. j. Leaf-blade filiform, to 1 mm. wide, the base not attenuate. ......«... 2. eee e 20. R. confinis. i. Spikelet 2-3-flowered. ............................ 6. R. corymbosa. h. Achene cancellate. k. Spikelet 5-6-flowered. ...................—......- 5. R. polyphylla. k. Spikelet 2-3-flowered. l. Rachilla strongly curved between spikelets. ................... 4. R. aristata. l. Rachilla straight or nearly so, but not strongly curved between spikelets. m. Achene to 1.5 mm. long, very minutely cancellate. e ols 2. R. macrochaeta. m. Achene to 2 mm. long, obviously cancellate. ........ 3. R. Schiedeana. TTT AAA g. Achene transversely rugulose. n. Inflorescence of a few dense heads; style only slightly bifid. .........11. a eeeceeees cos ecs cesa EE 7. R. cyperoides. n. Inflorescence of distant corymbs; style deeply bifid. o. Style-base conic-subulate, as long and broad as the Greg eode tea "SACO ER 13. R. Marisculus. o. Style-base conic, as wide as the achene but only half as long. ...—....... erronee oen ERE ERREUR ee UR 14. R. glauca. f. Bristles plumose, at least in part. p. Inflorescence of 5-6 lax distant corymbs. .... 24. R. millegrana. p. Inflorescence of 7-12 distant pyramidal panicles, ...................- desesedecscaascoccasessscsccsovevsosecossecscasceess nadari DR ERE E he 25. R. puberula. e. Bristles O. q. Achene horned at shoulders, appearing tridentate. ..................- 116 Rhodora [Vol. 67 :———— ——— — 21. R. setacea. q. Achene not horned at shoulders, not appearing tridentate. r. Achene gray. s. Achene hardly or not at all rugulose. ............ 9. R. andina. s. Achene obviously rugulose. ........................ 19. R. emaciata. r. Achene brownish to blackish. t. Achene not transversely rugulose, but cancellate or scrobi- culate. u. Spikelets fasciculate; achene scrobiculate; style-base depressed, much shorter and narrower than achene. jevvescepacessedssasdssqaresessevssanvsenessesonecssssocauneuoossesssee 20. R. confinis. u. Spikelets solitary; achene cancellate; style-base conic- subulate, longer than and nearly as wide as achene. n———— eese LO, R. boliviensis. t. Achene transversely rugulose. v. Leaf-blades to 1.4 cm. wide; panicle of relatively few dense heads. ................. esee eene 8. R. exaltata. v. Leaf-blades to 2-3 mm. wide; inflorescence not of dense heads. w. Spikelets solitary or paired at ends of peduncles. KEREN 16. R. velutina var. Sellowiana. w. Spikelets fasciculate. x. Lowest scales (3-4) more or less long-apiculate, the apiculus hirtellous. ........................ s. 15. R. robusta. x. Lowest scales not long-apiculate, not hirtellous. y. Inflorescence of 2-3 remote lax corymbs. ................ RM 18. R. tenuis. y. Inflorescence of 2-3 remote lax pyramidal panicles. ........... sse 17. R. luzuliformis. 1. Rhynchospora umbraticola var. Kuntzei (C. B. Clarke) Kükenth. in Bot. Jahrb. 74: 391 (1949). Dichromena Kuntzei of the Catalogue. Culms to 4 dm. tall. Leaves mostly basal, the blades to 3 dm. long (occasionally to 5 dm.) and 2 em. wide, glabrous, the bases subpetio- lately attenuate. Inflorescences axillary and terminal, corymbose- paniculate; bracts foliar, usually exceeding the inflorescences. Spikelets solitary, oblong-elliptic, to 7 mm. long, with numerous (12-20) flowers; scales light brown, ovate, mucronate; bristles 6, antrorsely scabridulous, exceeding the achene; stamens 3; style shortly bifid, the style-base as long and broad as the achene. Achene to 3 mm. long, obovate, shining, brown, biconvex, densely and evenly scrobicu- late. SANTA CRUZ: CERCADO: Cerro Amboró, Steinbach 3459 (GH); SARA: Rio Yapacani, O. Kuntze (type; photo, GH); Buenavista, 450 m., Steinbach 2878 (GH). BENI: YACUMA: Rurrenabaque, Cárdenas (M. E.) 1586 (GH, NY), White (M. E.) 851 (GH, Us). 2. R. macrochaeta Steud. ex Boeckl. in Linnaea, 38: 632 (1874). 1965] Flora of Bolivia — Foster 117 Culms to nearly 1 m. tall, with up to 3 cauline leaves. Leaves shorter than culms, to 6 mm. wide, glabrous, the revolute margins scabrid. Inflorescence of terminal and axillary, short, pyramidal or sub- pyramidal panicles, the terminal panicle larger and denser than the laterals; bracts foliar, exceeding the panicles. Spikelets linear to narrowly lanceolate, to 6-7 mm. long, solitary or paired, few-flowered (2-3) ; scales lance-oblong, membranous, brown or red-brown; bristles 4-6, antrorsely scabrid, slightly exceeding the achene; stamens 3; style apically subentire or somewhat bifid, the style-base pyramidal, rugose, as long and broad as the achene. Achene to 1.5 mm. long, brown to blackish, shining, minutely cancellate. COCHABAMBA: CERCADO: Cocha- bamba, Bang 1071 (GH); CHAPARE: Locotal, 1600m., Steinbach 9096 (GH). 2a. R. macrochaeta var. Ruiziana (Boeckl.) Kiikenth. in Bot. Jahrb. 74: 393 (1949). Components of inflorescence longer (to 9 em.), spike- lets 2-4-flowered, scales with the midrib excurrent as a long mucro, LA PAZ: NOR YUNGAS: Polo-Polo, 3200 m., Buchtien 698 (GH); LARECAJA: Ingenio del Oro, Rusby 167 (GH). SANTA CRUZ: VALLE- GRANDE: Comarapa, 2800 m., Steinbach 8340 (GH, NY); Cerro San Mateo, Comarapa, 3500 m., Steinbach 8532 (GH, NY). Rhynchospora macrochaeta var. colombiensis f. conden- sata and R. macrochaeta var. quinquespicata, listed in the Catalogue, seem hardly worthy of recognition. 3. R. Schiedeana (Schlechtd.) Kunth, Enum. Pl. 2: 300 (1837). Culms to 1.5 m. high, glabrous. Leaves basal and cauline, very long, mostly exceeding the inflorescence, acuminate, to 1 cm. wide, the margins scabrid. Inflorescence paniculate, compound, interrupted, the component panicles pyramidal or subpyramidal. Spikelets solitary or paired, lanceolate, to 6-7 mm. long, 2-3-flowered; scales 5-6, red- brown, lance-ovate, mucronate; bristles 3-4, antrorsely scabrid, shorter than the achene; stamens 3; style entire or the apex very briefly bifid, the style-base conic-subulate, blackish, rugose, as long and broad as the achene. Achene much shorter than the scales, about 2 mm. long, broadly obovoid, shiny brown, cancellate. LA PAZ: NOR YUNGAS: Unduavi, 3200 m., Buchtien 2588 (GH), 2593 (GH), SUR YUNGAS: San Felipe, Holway & Holway (GH, NY). 4. R. aristata Boeckl. in Flora, 40: 36 (1857). R. aristata var latilaminata Kükenth. in Fedde, Rep. Sp. Nov. 26: 253 (1929). Culms to 9 dm. (-1.5 m.) high, glabrous. Leaves mostly basal (about 3 cauline), equaling the inflorescence, to 8(-20) mm. wide, glabrous, scabridulous near the apex. Inflorescence a lax, open, compound panicle, the terminal portion corymbose, the laterals pyramidal, branches divaricate; bracts foliar, exceeding the panicle, bractlets setaceous, basally ciliate. Spikelets solitary, distant, the rachilla 118 Rhodora [Vol. 67 strongly curved between spikelets, 6-9 mm. long, 2-3-flowered, the lowermost flower perfect, the rest staminate or sterile; scales 6-7, brown or reddish-brown, mucronate or aristate; bristles 4-5, antrorsely scabrid, shorter or longer than the achene; stamens 3; style long, shortly bifid, the style-base ashy-gray, conic-subulate, longer and much narrower than the achene. Achene about 2 mm. long, broadly ovoid, shiny brown or stramineous, cancellate. LA PAZ: NOR YUNGAS: Polo- Polo, 1100 m., Buchtien 3651 (GH; type-number of var. latilaminata) . COCHABAMBA: CHAPARE: Locotal, 1600 m., Steinbach 9100 (GH). The variety latilaminata was recognized primarily on the basis of broader leaves (2 cm. wide), but an isotype avail- able to me has leaves rather less than 1 cm. wide. Its sepa- rateness seems dubious. 5. R. polyphylla Vahl, Enum. 2: 230 (1806). Culms to 9 dm. high, basally thickened, glabrous. Leaves increasing in size upwards on the culm, exceeding the inflorescence, to 1 cm. wide, glabrous or more or less pilose. Inflorescence a long, lax, compound, pale panicle, the 4-7 components distant, much-branched, more or less pyramidal; bracts foliar, exceeding the component panicles, Spikelets solitary, or occasionally 2-3 in a fascicle, 5-6-flowered, terete or lanceolate, 4-5 mm. long, distant on the produced rachilla; scales light-stramineous, lance-ovate, mucronate; bristles few, minute (rarely 1 or more exceeding the achene); stamens 2-3; style long, entire or slightly bifid, style-base pyramidal, greenish, as wide as the achene, but about half its length. Achene obovoid, slightly less than 1 mm. long, bi- convex, cancellate, light brown. LA Paz: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 341 (Kükenthal). 5a. R. polyphylla var. laxa (Beauv.) Pfeiff. in Fedde, Rep. Sp. Nov. 49: 78 (1940). Dichromena Mandonii of the Catalogue. Culms taller, to 1.4 m., leaves broader (to 2 cm.), component-panicles 3-4, scales rusty-brown, bristles 4, exceeding the achene. LA PAZ: LARECAJA: Sorata, Mandon 1421 (Kiikenthal). 5b. R. polyphylla var. longispiculosa Kükenth. in Bot. Jahrb. 74: 406 (1949). Components of the panicle 3, approximate, spikelets dull brown, linear-subulate, to 9-10 mm. long. SANTA CRUZ: SARA: La Perdix, 450 m., Steinbach 7993 (GH). 6. R. corymbosa (L.) Britton in Trans. N. Y. Acad. Sci. 11: 84 (1892). Culms to 1 m. high, glabrous, the cauline leaves inserted remotely. Leaves equaling or shorter than the culms, to 2 cm. wide, the midrib and margins scabrid. Inflorescence a panicle of 2-5 distant corymbs; bracts long, foliar. Spikelets many, usually fasciculate (2-5 in a fascicle), occasionally solitary, 6-8 mm. long, 2-3-flowered; scales red-brown, mucronate; bristles 6, antrorsely scabrid, exceeding the 1965] Flora of Bolivia — Foster 119 achene; stamens 3; style nearly entire, the conic base as wide and long as the achene, brownish, basally bilobed, sometimes finely puberu- lent. Achene obovoid, 2-3 mm. long, brown or chestnut-brown, the 2 broad faces with irregular depressions, densely and finely pitted. SANTA CRUZ: SARA: Buenavista, Steinbach 6674 and 7051 (Küken- thal, 5215 (GH, NY). BENI: YACUMA: Reyes, 300 m., White (M. E.) 1527 (GH, NY); near Lake Rogagua, 300 m., Rusby (M. E.) 1642 (GH, NY). 7. R. cyperoides (Swartz) Mart. in Denkschr. Akad. Wiss. Münch. 6: 149 (1820). Culms solitary, to 1 m. tall, scabrid on the angles, with 2-3 distant cauline leaves. Leaves shorter than the culms, to 7 mm. wide, the margins scabrid. Inflorescence a panicle of a few, dense, distant, globose heads (anthelate), bracts mostly exceeding the heads. Spikelets to 6 mm. long, ovate-lanceolate, many in a head; scales 7-8, mucronate, pale red-brown; bristles 6, antrorsely scabrid, 3 or 0 in male flowers; stamens 3; style slightly bifid, the base pale green, longer and much narrower than the achene, conic-subulate. Achene about 1.5 mm. long, shorter than the scale, obovoid, brown, shining, the margins slightly scabrid, finely transverse-rugulose. SANTA CRUZ: CERCADO: Santa Cruz, Steinbach 5343 (NY). 8. R. exaltata Kunth, Enum. Pl. 2: 291 (1837). R. exaltata var. ovalis of the Catalogue. Culms to 1 m. or more. Leaves as long as the culms or shorter, to 1.4 cm. wide, the margins and nerves scabrid or serrulate-scabrid. Inflorescence a much-interrupted panicle of rather dense heads; bracts exceeding the heads. Spikelets more or less stellately arranged in the heads, to 6 mm. long, narrowly lanceolate to subulate; scales stramineous or pale brown, mucronate; bristles 0; stamens 3; style with 2 linear branches, the style-base subulate, ashy- gray, shorter than the achene. Achene to 2.5 mm. long, orbiculate to ovoid, chestnut-brown, shining, transversely rugulose. LA PAZ: LARE- CAJA: San Carlos, Mapiri. 850 m., Buchtien 339 (GH). 8a. R. exaltata var. cephalophora (Nees) Kiikenth. in Bot. Jahrb. 74: 440 (1949). Inflorescence racemose, of solitary, distant spikes. Bristles present (2-3). SANTA Cruz: [prov. uncertain]: Cerro Hosana, Steinbach 3387 (Kiikenthal). 9. R. andina Kükenth. in Fedde, Rep. Sp. Nov. 53; 73 (1944). Culms to 3.5 dm. tall, glabrous, cauline leaves absent or 1, culms clothed basally in fibrous remains of old leaf-sheaths. Leaves much shorter than the culms, to 1 mm. wide, glabrous. Inflorescence a panicle of 2-4 distant corymbs, bracts shorter than or equaling the peduncles. Spikelets few, fasciculate, rarely solitary, 'to 4 mm. long, lance-oblong; scales 7-8, brown, midrib excurrent as an arista; bristles 0; stamens 3; style entire, the flattened base brownish, as broad as, but shorter than, the achene. Achene to 1 mm, long, obovoid, shining, gray, 120 Rhodora [Vol. 67 obscurely or not at all transversely rugulose. LA PAZ: LARECAJA: Hacienda Casana, 1400-1600 m., Buchtien 7159 (GH; type-number). 10. R. boliviensis C. B. Clarke in Kew Bull. Add. Ser. 8: 37 (1908). Culms to 2-3.5 dm. high. Leaves to 15 cm. long and 1-2 mm. wide, flat, glabrous. Inflorescence a lax slender panicle to 15 cm. long, spikelets solitary, pedicellate, 10-20 on a culm. Spikelets 5 mm, long, oblong-lanceolate; scales lanceolate to ovate, not mucronate, brown, the lowest 4 sterile; bristles 0; style very shortly bifid, style-bases basally inconspicuously bilobed, conic-subulate, 1.5 mm. long and nearly as wide as the achene. Achene 1 (-1.25) mm. long, obovoid, plano-convex, abruptly narrowed at the apex into an extremely short neck, brownish, cancellate. YUNGAS: 2000 m., Rusby 83 (NY; type- number). LA PAZ: NOR YUNGAS: Unduavi, 3300 m., Rusby 82 (NY), 85 (US). 11. R. globosa (HBK.) Roem. & Schult. Syst. Veg. 2: 89 (1817). R. globosa var. Loefgrenii of the Catalogue. Culms to 9 dm. high, caespitose, glabrous, cauline leaves 0. Leaves shorter than culms, to 2 mm. wide, margins scabrid, grayish-green. Inflorescence mono- cephalous, the globose head to 1-2 cm. in diameter, bracts short, scale- like, not foliar, inconspicuous, firm-textured, brownish, mucronate, the margin ciliate on the upper portion. Spikelets 6-7 mm. long, lanceolate; scales reddish-brown, the lowest short, the upper longer, acute; bristles 5 in perfect flowers, usually 2 in staminate flowers, the basal portion densely white-plumose, the apical portion antrorsely scabrid; stamens 3; style very briefly bifid, the conical base light greenish-brown, narrower and shorter than the achene. Achene about 2 mm. long, oblong-obovoid, brown, the shoulders scabrid-ciliate at the apex. BENI: YACUMA: near Lake Rogagua, 330 m., Rusby (M. E.) 1700 (GH). LA PAZ: NOR YUNGAS: Milluguaya, 1800 m., Buchtien 4154 (GH). COCHABAMBA: CERCADO: near Cochabamba, Bang 873 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach. 5351 (GH); Dolores, 450 m., Steinbach 1896 (GH); CHIQUITOS: 5 km. north of Santiago de Chiquitos, 700 m., Cutler 7020 (am). The heads of the last collection cited are described by the collector as lemon-yellow. 12. R. hirta (Nees) Boeckl. in Vidensk. Meddel. Kjoebenh. 1869: 146 (1870). Culms solitary, to 6.5 dm. long, glabrous or somewhat pilose. Leaves shorter than the culms, to 5 mm. wide, the lower portion pilose-ciliate. Inflorescence monocephalous, the hemispherical to globose head to 2 cm. in diameter; bracts numerous, imbricate, ovate, acuminate, light brown, the margins entirely long-ciliate. Spikelets to 6-7 mm. long, lanceolate; scales 6, lance-oblong, pale brown; bristles 3, unequal, 1 exceeding, 2 shorter than, the achene, antrorsely scabrid, but not plumose; stamens 3; style hardly bifid, 1965] Flora of Bolivia — Foster 121 the long-conic greenish-brown base half as long as and narrower than the achene. Achene 2 mm. long, oblong, concavo-convex, brown, finely puncticulate, apically papillose. SANTA CRUZ: eastern Velasco, O. Kuntze (Kükenthal). 13. R. Marisculus Lindl. & Nees in Mart. Fl. Bras. 2(1): 142 (1842). Rhizome occasionally shortly stoloniferous. Culms to 1.5 m. high, smooth, with 4-5 distant cauline leaves. Basal leaves long, shorter than the culms, to 7 mm. wide, the margins scabrid. Inflorescence a panicle of 2-3 distant corymbs, the bracts shorter than the peduncles. Spikelets oblong-lanceolate, to 6-7 mm. long; scales 6, reddish-brown, lanceolate, mucronate; bristles 6, exceeding the achene, antrorsely scabrid; stamens 3; style deeply bifid, the conic-subulate base ashy- gray, as long and as broad as the achene. Achene 1.5 mm. long, obovoid, rather shiny, transversely rugulose. Without data: Bang 2204 (GH). 14. R. glauca Vahl, Enum. 2: 233 (1806). Culms slender, to 7.5 dm. tall, with 3 distant cauline leaves. Leaves shorter than the culms, to 3 mm. wide, the margins scabrid. Inflorescence a small terminal panicle of 3-4 distant corymbs; bracts narrow, about equaling the corymbs. Spikelets few, more or less fasciculate, 3-4 mm. long, lanceolate; scales ovate, dark grown, shortly mucronate; bristles 6, of varying lengths but mostly equaling the achene, antrorsely scabrid; stamens 3; style deeply bifid, the conical base gray, as wide as the achene and up to half as long. Achene suborbicular to obovoid, yellowish-brown, shiny, to 2 mm. long, transversely undulate-rugose. YUNGAS: Bang 433 (GH). LA PAZ: LARECAJA: Guanai-Tipuani, Bang 1426 (Kükenthal); Hacienda Simaco, 1400 m., Buchtien 5360 (GH). 15. R. robusta (Kunth) Boeckl. in Linnaea, 37: 616 (1873). Stoloni- ferous. Culms to 1.2 m. high, glabrous, smooth. Leaves firm, to 8 mm. wide, long, glabrous or ciliate. Inflorescence of 2-3 distant terminal corymbs; bract foliar, pilose. Spikelets solitary or fasciculate, to 1 cm. long, oblong-ovate; scales ovate to ovate-oblong, the lower with the midrib prolonged as a conspicuous hirtellous apiculus; bristles 0; stamens 3; style bifid, the base campanulate to bilobed, decurrent on the achene. Achene orbicular, to 1.5 mm. long, brown, undulate- rugose. BENI: YACUMA: near Lake Rogagua, 300 m., Rusby (M. E.) 1425 (GH, NY), 1442 (Ny, US). SANTA CRUZ: SARA: Río Cuchi, 450 m., Steinbach 6619 (GH). This species was not ascribed to Bolivia by Kükenthal. Although the material cited is young, there seem to be no perianth-bristles present. The hirtellous apiculus of the lowest scales is quite obvious, and I cannot attribute the specimens to any other species known from Bolivia. 122 Rhodora [Vol. 67 16. R. velutina var. Sellowiana (Kunth) Boeckl. in Linnaea, 37: 614 (1873). Culms to 9 dm. high, with 3-5 cauline leaves. Leaves shorter than the inflorescence, glabrous, 2-3 mm. wide, margins scabrid. Inflorescence of 1 terminal and 2 lateral corymbs; bracts 3, foliar. Spikelets solitary or paired at ends of peduncles, oblong-cylindrical, 5-6 mm. long; scales lance-ovate, the uppermost mucronate, brown or reddish-brown; bristles 0; stamens 3; style bifid, the campanulate base broadly bilobed at its base. Achene obovoid to orbicular, 2 mm. long, margined, chestnut-brown, shiny, transversely rugose. SANTA CRUZ: SARA: Buenavista, Steinbach (Kükenthal). 17. R. luzuliformis Boeckl. in Linnaea, 37: 632 (1873). Rhizome elongate, producing scaly stolons. Culms to 3 dm. high, glabrous, not scabrid, with 2-3 distant cauline leaves. Leaves equaling the culms, to 2 mm. wide, the margins scabrid. Inflorescence a compound panicle of 2-3 remote, smaller, more or less pyramidal panicles; bracts long, linear, foliar, much exceeding the component panicles. Spikelets somewhat fasciculate, 3 mm. long, oblong-ovate, reddish or brown; scales densely imbricate, ovate to oblong-ovate, mucronate; bristles 0; stamens 3; style bifid, the base pyramidal, gray-green, decurrent on the shoulders of the achene, as broad and half as long as the achene. Achene subglobose, lenticular or biconvex, yellow-brown, transversely undulate-rugose. TARIJA: ARCE: Camacho, 2500 m., Fiebrig 2575 (GH). 18. R. tenuis Link in Spreng. Schrad. & Link, Jahrb. 1(3): 76 (1820). Rhizome very short or lacking. Culms to 4 dm. high, filiform, smooth and glabrous, with 1-2 cauline leaves. Leaves shorter than the culms, to 2 mm. wide, glabrous. Inflorescence of 2-3 lax remote corymbs; bracts almost filiform. Spikelets usually fasciculate (2-3), 4-5 mm. long; scales 5-6, red-brown or stramineous, ovate to ovate- lanceolate, mucronate; bristles 0; stamens 3; style bifid, the base flattened-pyramidal, gray to blackish, its base bilobed, decurrent on the angles of the achene, half as long as the achene. Achene 1 mm. long, orbicular to obcvoid, reddish-brown to blackish, conspicuously transverse-rugose. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5056 (GH, NY), 5060 (GH, NY). 19. R. cmaciata (Nees) Boeckl. in Vidensk. Meddel. Kioebenh. 1869: 149 (1870). R. tenuis var. emaciata of the Catalogue. Culms to 6 dm. tall, trigonous, smooth and glabrous. Leaves very much shorter than the culms, filiform, canaliculate. Inflorescence terminal, of 1-2 remote lax corymbs; bracts long, bracteoles long-setaceous. Spikelets solitary, pedicellate, lance-oblong, 5-6 mm. long; scales 5-6, brownish with hyaline margins, aristate; bristles 0; stamens 3; style deeply bifid, the depressed-pyramidal base about one-third the length of the achene and much narrower, stramineous to castaneous, its base almost entire 1965] Flora of Bolivia — Foster 123 and not decurrent on the achene. Achene 1 mm. long, orbicular, lead- gray, undulate-rugose. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1899 (GH), 6926 (GH). 20. R. confinis (Nees) C. B. Clarke in Kew Bull. Add. Ser. 8: 40, 119 (1908). Rhizome long. Culms solitary, to 5 dm. tall, smooth, with remote cauline leaves to 1 mm. wide, shorter than the culms, scabrid. Inflorescence 2-3 dense long-pedunculate corymbs; bracts setaceous, exceeding the corymbs. Spikelets rather densely fasciculate, to 8 mm. long, many-flowered; scales oblong-lanceolate, aristate, reddish-brown; bristles usually 0, occasionally 3; stamens 3; style deeply bifid, the blackish base depressed-campanulate, narrower than the achene. Achene 1 mm. long, broadly obovoid, pale to chestnut-brown, finely scrobiculate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6924 (GH), 7091 bis (GH). 21. R. setacea (Berg.) Boeckl. in Vidensk. Meddel. Kjoebenh. 1869: 159 (1870). R. tenerrima of Catalogue. Caespitose, the bases of tufts somewhat enlarged, rhizome absent. Culms to 3.5 dm. high, filiform, smooth. Leaves mostly longer than culms, sometimes shorter, filiform, glabrous. Inflorescence of 2-4 axillary and terminal, somewhat lax, distant corymbs with 1-6 spikelets in each; bracts setaceous, exceeding the corymbs. Spikelets in small fascicles or solitary, 4-5 mm. long, lanceolate to terete; scales 7-8, ovate to lanceolate, firm, stramineous to reddish, with the midrib prolonged as an arista; bristles 0; stamens 2; style bifid, the base broad, flat, elongated as 2 horns at the shoulders of the achene, the achene thus appearing tridentate. Achene 1 mm. long, orbicular, brown, transversely undulate-rugose. SANTA CRUZ: SARA: Buenavista, Steinbach 1901 (GH). 22. R. nervosa (Vahl) Boeckl. in Vidensk. Meddel. Kjoebenh. 1869: 143 (1870). Dichromena ciliata of the Catalogue. Rhizome short, often developing woody stolons. Culms to 7 dm. high, usually shorter, usually glabrous. Leaves as long as the culms or shorter, 1-2 mm. wide, basally sheathing, the old sheaths reduced to a basal collar of fibers, blades scabrid-ciliate. Inflorescence a single terminal head of 3-5 spikelets 6-8 mm. long; involucre of 4-7 unequal bracts, the bracts pubescent on the abaxial side and at least the basal portion ciliate. Seales oblong-ovate to lance-ovate, the midrib thickened, glabrous, white; bristles 0; stamens 3; style deeply bifid, the style-base bluntly pyramidal. Achene to 1.5 mm. long, orbicular to obovoid, or more or less lenticular, dark brown, transversely rugulose. YUNGAS: Bang 251 (GH). BENI: CERCADO MOJOS: Trinidad-Misiones Guarayos, 250 m., Werdermann 2487 (MO); YACUMA: Reyes, White (M. E.) 1537 (GH). LA Paz: NOR YUNGAS: Milluguaya, 1900 m., Buchtien 396 (GH); Polo-Polo, 1100 m., Buchtien (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 3277 (GH), 6725 (GH). 124 Rhodora [Vol. 67 The binomial Rhynchospora ciliata (Vahl) Kükenth. J (based on Dichromena ciliata Vahl) is a later homonym of R. ciliata Vahl and therefore cannot be used for this species. 22a. R. nervosa var. Jelskiana (Boeckl.) Kükenth. in Bot. Jahrb. 75: 298 (1952). Differs in having wider leaves (3-4 mm.), the culms often pilose, and spikelets to 12 mm. long. SANTA CRUZ: SARA: Río Surutü, Steinbach 6841 bis (Kükenthal). 24. R. millegrana (Nees) R. C. Foster, comb. nov. Nemochloa millegrana Nees in Mart. Fl. Bras. 2(1) : 148 (1842). Rhizome slender, scaly, horizontal. Culms to 1.2 m. high, smooth. Leaves long, but usually shorter than the culms, 5-7 mm. wide, the margins and nerves scabrid. Inflorescence of terminal and lateral distant lax corymbs; bracteoles narrow, setaceous. Spikelets to 2 mm. long, solitary or 2-3 approximate; scales broadly ovate, often emarginate, coppery or pale red-brown; bristles 4-5, plumose; stamens 3; style deeply bifid, the base conic-pyramidal, ashy gray, less than half the length of the achene. Achene to 1.5 mm. long, orbicular-obovoid, shiny, deep chest- nut-brown, granulate-rugulose. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 332 (GH); San Antonio, Mapiri, 850 m., Buchtien 1233 (GH). This is the variant which has been distinguished as Pleurostachys minuscula Kükenth. in Fedde, Rep. Sp. Nov. 26: 254 (1929). According to the Index Kewensis, the binomial Rhyncho- spora millegrana has been published before, being attributed to Schrad. ex Steud. Syn. Pl. Cyp. 139 (1855). Examination of this citation shows that the binomial was published in synonymy, so that the name cannot serve as a bar to the proper transfer of the basionym. 25. R. puberula (Boeckl.) L. B. Smith in Phytologia, 1: 82 (1934). Pleurostachys puberula var. Buchtienii of the Catalogue. Rhizome short, thick. Culms to 1.2 m. tall, thick. Leaves about equaling the culms, to 2 cm. wide, pilose or glabrate beneath, the sheaths pube- rulent. Inflorescence of 7-12 distant pyramidal panicles; bracts exceeding the panicles. Spikelets more or less globose, to 1.5 mm. long, in dense spikes at ends of branchlets; scales broadly ovate, reddish- brown; bristles 4-5, plumose, at least in part; stamens 3; style-base conic-pyramidal, much shorter than the achene. Achene 1 mm. long, obovoid, shiny chestnut-brown, densely granulate-rugose. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 336 and 337 (Küken- thal). 1965] Flora of Bolivia — Foster 125 12. CALYPTROCARYA Nees Calyptrocarya glomerulata (Brongn.) Urb. Symbol. Antill. 2: 169 (1900). Becquerelia glomerulata and Calyptrocarya fragifera of the Catalogue. Perennial from a slender woody rhizome, to 3.5 dm. high, the entire plant glabrous. Leaves eligulate, linear, to 9 mm. wide, usually exceeding the inflorescence. Inflorescence of small terminal or axillary corymbose panicles, the individual subglobose heads to 3 mm. long, the flowers unisexual. Female flowers: 2 distichous basal glumes or scales, many-nerved, the mid-nerve very prominent; achene lentic- ular to biconvex, apiculate, covered by a puberulent membrane, about 1 mm. long; style bifid. Male spikelets 2-4 at base of pistillate flowers, each of 1-4 flowers reduced to single stamens. SANTA CRUZ: ICHILO: Río Vibora, 350 m., Steinbach 7579 (GH). 13. SCLERIA Berg. See Core in Brittonia, 2: 1-105 (1936). Perennial, rhizomatous or stoloniferous (1 species sometimes annual, with fibrous roots). Culms mostly triangular. Leaf-sheaths often trialate, ligulate; blades sometimes plicate. Inflorescence of distant glomerulate heads, or densely spicate, or obviously paniculate, or, in no. 1, of dense heads subsessile in the leaf-axils. Flowers uni- sexual, perianth absent. Pistillate spikelets 1-flowered, often in the axil of the lowest sterile scale of a staminate spikelet. Staminate spikelets few-to many-flowered. Hypogynium obscure or absent in a few species, but present in most. Achenes hard, bony, often smooth and shining, or reticulate, rugose, verrucose or tuberculate. a. Inflorescence of dense spikelet-heads subsessile in leaf-axils. ........ wc ME o ec 1. S. amphigaea. a. Inflorescence not of dense spikelet-heads subsessile in leaf-axils. b. Achenes smooth and glabrous. c. Hypogynium obscure or absent. d. Inflorescence simple, the spikelets in distant glomerulate heads. e. Scales hirsute. .......cccccccccecccecccceceecceeeeeeecaeeeneeeenees 2. S. hirtella. e. Scales glabrous. e 9. S. leptostachya. d. Inflorescence branched. f. Achenes 2-2.5 mm. long. ................... 4. S. lithosperma. f. Achenes 1.5-2 mm. long. .................... s eeeeeee 5. S. castanea. c. Hypogynium obviously present. g. Margin of hypogynium entire or undulate, not 3-lobed. h. Achenes 3.5-6 mm. long. ................... 7. S. macrophylla. h. Achenes 3 mm. or less in length. i. Hypogynium long-ciliate on margin. ............ 9. S. mitis. i. Hypogynium short-ciliolate on margin. j. Largest achenes 2 mm. long. ............ 8. S. microcarpa. 126 Rhodora [Vol. 67 j. Largest achenes 3 mm. long. sesser 10. S. obtusa. g. Margin of hypogynium 3-lobed. k. Margin obscurely 3-lobed. ..................... 19. S. vaginata. k. Margin obviously 3-lobed. l. Lobes fimbriate. m. Inflorescence and achenes purplish. ................. eese BEEN 12. S. arundinacea. m. Inflorescence brown; achenes white. .. 13. S. latifolia. l. Lobes entire. n. Inflorescence purple or brownish-purple. ...................- NEM 20. S. melaleuca. n. Inflorescence green or yellow-brown. .... 21. S. pterota. b. Achenes not smooth and glabrous. o. Hypogynium obscure or none. p. Inflorescence of few, distant, glomerulate heads. .................... ususuueweuvssssessavewusrseseassseengsaasenssscssse+ssssssanesesuaesssstese 3. S. leptostachya. p. Inflorescence dense, spicate. ...............eee 6. S. composita. o. Hypogynium obviously present. q. Hypogynium subentire, the margin undulate. .. 22, S. secans. q. Hypogynium 38-lobed. r. Plant scandent. ..................eeeeeeeeenee 17. S. tenacissima. r. Plant not scandent. s. Achenes reticulate. t. Achenes faintly reticulate, sparsely puberulent, 2-3 mm. long. ...........eeeeeeeeeeeeeene nere 18. S. lagoénsis. t. Achenes reticulate, the ridges pubescent, 2 mm. long. BEN UO I 16. S. setacea. S. Achenes rugose, verrucose or tuberculate. u. Staminate and pistillate spikelets in separate parts of inflorescence. ......cceccecceeceeesccseeseeeceeeenseceeees 15. S. bracteata. u. Staminate and pistillate spikelets not in separate parts of inflorescence. v. Achenes minutely verrucose, .... 11. S. eyperinoides. v. Achenes rugose-tuberculate, ........ 14. S. panicoides. 1. Scleria amphigaea Raymond in Nat. Canad. 91: 132 (1964). Diplacrum longifolium of the Catalogue. Perennial, with creeping sealy stolons, plants to 0.5 m. high. Stems leafy, leaves linear, to 5 mm. wide, glabrous, bearing dense, globular, subsessile heads of spikelets in the axils. Trifid style and achene subtended by 2 broad scales to 4-5 mm. long, flanked by 2 staminate spikelets, the staminate flowers monandrous, usually 6 to a spikelet, subtended by 2 narrow scales. Fruit suborbicular, shining white or ivory, longitudinally ridged or irregularly conspicuously reticulate. SANTA CRUZ: SARA: Cucichisito, 450 m., Steinbach 7014 (GH). 2. S. hirtella Swartz, Prodr. 19 (1788). Perennial, with a long slender rhizome. Culms to 6.5 dm. high, sharply triangular, slender, 1965] Flora of Bolivia — Foster 127 glabrous, sometimes pubescent near the apex. Leaves to 2 dm. long and 5 mm. wide, pubescent to glabrous; sheaths pilose, at least in a vertical line below the orifice, ligule usually absent (minute, if present). Inflorescence terminal, simple, interrupted, the spikelets aggregated in small, sessile, remote, glomerulate heads, the rachis sparsely to densely pilose; bractlets not exceeding the glomerule, long- pilose or hirsute. Staminate scales narrow, brown, dorsally hirsute. Pistillate scales cuspidate, brown, broader than staminate scales. Spikelets 4-5 mm. long, brown, red-brown, or almost black; hypogy- nium absent; achene subglobose-trigonous, smooth, white, shining, apiculate, the base not or only slightly excavated. LA PAZ: LARECAJA: Hacienda Casana, 1400 m., Buchtien 7165 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1802 (GH), 5327 (GH), 6861 (GH). 3. S. leptostachya Kunth, Enum. Pl. 2: 354 (1837). Perennial, with a long rhizome. Culms to 6 dm. high, filiform. Leaves to 3 dm. long, 1 mm. wide, blades glabrous; sheaths more or less pilose, ligule absent. Inflorescence terminal, interrupted, simple (sometimes few-branched), the spikelets aggregated in small heads; rachis ciliolate on the angles, otherwise glabrous; bractlets basally dilated, setaceous, glabrous. Spikelets to 5 mm. long, linear-oblong, obtuse. Staminate and pistillate scales reddish-brown, acuminate, membranous, glabrous, oblong, the pistillate much exceeding the achenes; hypogynium absent; achene short-stipitate, subglobose, white or dirty-white, transversely rugulose or nearly smooth, 1 mm. long. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5300 (GH). 4. S. lithosperma (L.) Swartz, Prodr. 18 (1788). Perennial, with short, nodulose rhizomes. Culms filiform, triangular, glabrous above, the angles often scabrid-ciliate below, to 6 dm. tall. Leaves to 2 dm. long and 3 mm. wide, margins scabrid; sheaths pilose to glabrous, ligule triangular, short, pilose. Inflorescences terminal and axillary, simple or branched, interrupted, the few spicate heads remote, few- flowered. Spikelets to 4 mm. long, bracts filiform, much exceeding the heads, bractlets dark brown; scales dark brown, ovate-lanceolate; hypogynium absent; achene to 2.5 mm. long, white, shining, smooth, oblong, basally attenuate, trigonous, the apex more or less umbonate. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5203 (GH). As noted by Core, this collection is distinctive in that there are 2 narrow red-brown lobes at 2 corners of the base of the achene. These lobes may be entire or apically bifid, ciliolate or glabrous. 5. S. castanea Core in Brittonia, 1: 239 (1934). S. pleostachya of the Catalogue. Perennial, the rhizome thick. Culms to 8 dm. tall, acutely triangular, the angles scabridulous. Leaves to 4.5 dm. long and 3 mm. wide, somewhat short-pilose along the margins and the 128 Rhodora [Vol. 67 midrib beneath; sheaths reddish-purple, pubescent, the ligule shortly pilose, obtusely triangular. Inflorescence a terminal panicle, few- branched. Bracts setaceous. Spikelets to 7 mm. long, oblong or linear-oblong, densely clustered. Staminate scales red-brown, acumi- nate. Pistillate scales red-brown, lance-ovate; hypogynium absent; achene to 2 mm. long, oblong-ovoid, obscurely trigonous, apiculate, shining white and smooth, the triangular base narrowed. YUNGAS: Bang 203 (GH). COCHABAMBA: CHAPARE: Locotal, 1700 m., Steinbach 9068 (GH; type-number). SANTA CRUZ: SARA: Cerro Hosana, Stein- bach 3421 (GH). 6. S. composita (Nees) Boeckl. in Linnaea, 38: 444 (1874). Perennial from a slender rhizome. Culms to 6 dm. tall, slender, triangular. Leaves to 2 dm. long and 2-5 mm. wide, pilose beneath and sparsely pilose to glabrous above; sheaths glabrous to densely ciliate, ligule rounded, ovate, pilose. Inflorescence terminal, densely spicate, the rachis sparsely to densely pilose; bracts setaceous (if present) ; bractlets setaceous, hairy, mostly not exceeding the spikelets. Spikelets brown, 2-3 mm. long; staminate scales lanceolate, to 4 mm. long; pistillate scales with pilose keel; hypogynium none, or almost none; achene obscurely trigonous, white or dirty-white, irregularly reticulate, mucronate, 1 mm. long. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6891 (GH). 7. S. macrophylla Presl, Rel. Haenk. 1: 200 (1830). S. paludosa of the Catalogue. Perennial, with thick rhizomes. Culms 1-3 m. tall, acutely triangular, the angles scabridulous, thick. Leaves to 4 dm. long and 4.5 em. wide, the angles and margins scabridulous; sheaths trialate, the wings scabridulous, ligule ovate, obtuse, short. Inflo- rescence an axillary and terminal panicle with erect branches; bract leaf-like, bractlets setaceous, basally pubescent. Staminate spikelets to 4 mm. long, the scales lanceolate, acuminate-acute, puberulent. Pistillate scales broad, more or less puberulent, ciliate; hypogynium conspicuous, undulate, the margin ciliate; achene to 6 mm. long, subglobose, but very obscurely subtrigonous, white or dirty-white to pale brown, smooth, shining, the puberulent style-base persistent. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1465 (GH), 8021 (GH). 8. S. microcarpa Nees ex Kunth, Enum. Pl. 2: 341 (1837). Peren- nial from elongate rhizomes. Culms to 2 m. high, acutely triangular, glabrous. Leaves to 4 dm. long and 1 cm. wide, glabrous; sheaths trialate, glabrous, the lanceolate glabrous ligule to 1 cm. long. Inflo- rescence of terminal and axillary panicles, the branches erect, spikelets somewhat remote; bract foliar, bractlets very small, more or less subulate. Staminate spikelets oblong-ovate, with obtuse, ovate, straw- colored scales. Pistillate scales suborbicular, straw-colored; hypo- gynium undulate and densely ciliate; achene ovoid, to 2 mm. long, 1965] Flora of Bolivia — Foster 129 smooth, white to pale buff, the style-base somewhat persistent. SANTA CRUZ: ICHILO: Río Vibora, 350 m., Steinbach 7564 (GH). In its original use, in Linnaea, 9: 302 (1834), this binomial was a nomen nudum. Kunth’s validation of the name in 1837 is the earliest known to me. 9. S. mitis Berg. in Vet. Akad. Handl. Stockh. 27: 145, t. 5 (1765). Perennial, from moderately thick rhizomes. Culms to 3 m. tall, sharply triangular, smooth, rather thick. Leaves to 6 dm. long and 2.5 cm. wide, subplicate, the margins and nerves on the upper surface scabrid; sheaths trialate, glabrous, the lanceolate to long-deltoid ligule glabrous, to 3 cm. long (usually shorter). Inflorescence paniculate, lax, the branches erect; bract foliar, bractlets setaceous, short. Spike- lets about 2 mm. long, ovoid-elliptic; pistillate scales orbicular, shortly acuminate; hypogynium with an undulate ciliate margin, the hairs dense, long, red-brown; achene ellipsoid-conic, 2-8 mm. long, white, smooth, the black style-base persistent. SANTA CRUZ: SARA: Buena- vista, 500 m., Steinbach 2866 (GH). 10. S. obtusa Core in Brittonia, 1: 240 (1934). Probably perennial (rhizomes lacking in material seen). Culms to 7.5 dm. high, slender, triangular, slightly scabridulous. Leaves to 2 dm. long and 1 cm. wide, scabridulous, especially on the upper surface; sheaths trialate (some- times narrowly so), ligule narrow, 2-4 mm. long. Inflorescence a terminal and axillary few-branched panicle, the branches erect; bract long, slender. Spikelets to 2 mm. long, the pistillate scales brown, suborbicular, abruptly short-acuminate or apiculate; hypogynium brown or purplish-brown, the undulate margin ciliate; achene 2-3 mm. long, oblong-globose, the apex somewhat tapering, obtuse, shining white, smooth. BENI: YACUMA: Hacienda Rosario, 300 m., White (M. E.) 1224 (type-number; Gray Herbarium sheet mixed with Scleria hirtella). 11. S. cyperinoides C. B. Clarke in Kew Bull. Add. Ser. 8: 61 (1908). Perennial from a rather thick rhizome. Culms to 1 m. high, sharply triangular, the angles scabridulous. Leaves to 3 dm. long and 1.6 cm. wide, the margins and prominent veins scabrid beneath; sheaths narrowly trialate, wing-margins scabrid, ligule ovate, conspicuous, membranous, to 1.5 cm. long. Inflorescence a dense terminal panicle, the rachis sparsely pilose to puberulent; bract long, foliar, bractlets subulate or setaceous. Staminate spikelets subsessile, numerous, the acuminate scales lanceolate, purplish. Pistillate scales lance-ovate. purplish; hypogynium 3-lobed, the lobes deeply dissected into several teeth; achene to 3 mm. long, shorter than the pistillate scales, verru- cose, puberulent, especially near the apex, ovoid, subtrigonous (the angles rounded), usually purplish. SANTA CRUZ: ICHILO: Rio Vibora, 350 m., Steinbach 7575 (GH). 130 Rhodora [Vol. 67 12. S. arundinacea Kunth, Enum. Fl. 2: 347 (1837). Perennial from a short thick rhizome. Culms to 2 m. tall, triangular, the angles scabridulous. Leaves few, to 5 dm. long and sometimes to 5 cm. wide, glabrous, plicate or subplicate, abruptly narrowed to an acuminate tip; sheaths broadly trialate, inflated, pubescent, especially near the ovate ligule. Inflorescence an axillary and terminal panicle, purple, the rachis scabrid; bract foliar, to 2.5 dm. long, bractlets setaceous, to 2 cm. long. Staminate spikelets to 3 mm. long, lanceolate. Pistillate scales ovate, purplish, midrib green; hypogynium 3-lobed, lobes fimbriate, fimbriae to 1 mm. long, purple; achene to 3 mm. long, depressed-globose, shining, smooth, purplish to black (portions shielded by scales often whitish). La PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 338 (NY). 18. S. latifolia Swartz, Prodr. 18 (1788). Perennial from a thick rhizome. Culms to 1 m. tall, triangular, short-pilose (at least on the upper portion). Leaves to 6 dm. long and 5 cm. wide, abruptly acuminate at the apex, the margins and principal nerves scabridulous on lower surface; sheaths trialate, puberulent to shortly pubescent, the deltoid-ovate ligule to 8 mm. long, sometimes pubescent. Inflo- rescence an axillary and terminal panicle, brownish, the peduncles pubescent; bract to 1.5 dm. long, foliar, bractlets filiform. Staminate spikelets lanceolate, to 4 mm. long, short-pedicellate, Pistillate spike- lets with membranous, brown, puberulent, ovate scales; hypogynium 3-lobed, the lobes deeply toothed or fimbriate; achene depressed- globose, to 3 mm. long, shorter than the pistillate scales, smooth, white to purple. LA PAZ: LARECAJA: San Carlos, Mapiri, Buchtien 1227 (US); Hacienda Casana, Buchtien 7161 (Us). The only marked difference between S. latifolia and S. arundinacea seems to be in the color of the inflorescence and achene. This hardly seems to warrant retention of two "species", especially as the two are sympatric. The range of S. latifolia is apparently entirely included in that of S. arundinacea. 14. S. panicoides Kunth, Enum. Pl. 2: 348 (1837). Perennial from a thick rhizome. Culms to 2 m. high, triangular, sparsely pilose, the angles scabrid. Leaves to 3 dm. long and 5 cm. wide, sparsely pubescent; sheaths trialate, pubescent, the short ovate ligule densely pubescent to pilose. Inflorescence of axillary and terminal purplish panicles, the rachis and branches pubescent to puberulent; bract foliar, bractlets filiform, long. Staminate spikelets to 4 mm. long, ellipsoid-oblong, the scales purplish. Pistillate scales suborbicular, pubescent, acuminate; hypogynium 3-lobed, laciniate, purple; achene depressed-globose, to 4 mm. long, whitish, shining, rugose-tuberculate, the tubereles puberulent or glabrous. Without data: Bang 2350 (GH). 1965] Flora of Bolivia — Foster 131 15. S. bracteata Cav. Ic. 5: 34, t. 457 (1799). S. bracteata f. simpli- cior Kükenth. in Fedde, Rep. Sp. Nov. 26: 253 (1929). Perennial from a thick rhizome, Culms to 3 m. long, triangular, scandent or subscandent. Leaves to 4.5 dm. long and 1.8 em. wide, pubescent on both surfaces, tapering to a long-caudate tip; sheaths shortly villous, often purple-tinged, the ovate ligule to 4 mm. long, pubescent. Inflo- rescence a terminal and axillary panicle, with the staminate and pistillate portions separate. Pistillate inflorescence: bracts to 7 cm. long, filiform, scabrid; spikelets few-flowered, lance-ovate, cuspidate, purple; hypogynium 3-lobed, the margin purple, entire or dentate; achene subglobose, to 3 mm. long, white or dirty-white, subtuberculate to verrucose, the tubercles puberulent. yYUNGAS: Bang 262 (GH). LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5102 (GH), 5107 (GH); Hacienda Casana, 1400 m., Buchtien 7164 (GH); NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 3652 (GH); Hacienda El Choro, Buchtien 8067 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7038 (GH). 16. S. setacea Poir. Encycl. 7: 4 (1806). Annual or perennial, with fibrous roots or a short rhizome. Culms to 8 dm. high, diffuse, triangu- lar. Leaves long, linear, 1-4 mm. wide, glabrous or sparsely pilose; sheaths glabrous, sometimes alate, the short ligule rounded. Inflo- rescence of terminal and axillary panicles, the lateral peduncles very slender, drooping; bracts linear or lance-linear, mostly exceeding the culms, bractlets linear to subulate. Spikelets 2-4 mm. long. Pistillate scales lance-ovate, acuminate, straw-colored to purplish; hypogynium prominently 3-lobed; achene globose to ellipsoid, 2 mm. long, reticulate, the ridges pubescent, dirty-white. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5425 (GH). 17. S. tenacissima (Nees) Steud. Syn. Pl. Cyp. 175 (1855). Base not seen, plant scandent. Culms to 3 dm. tall, clustered at nodes of the principal axes, triangular, the angles scabrid. Leaves to 3 dm. long and 4 mm. wide, pilose on both surfaces; sheaths pubescent to glabrous, angled, the angles scabrid, the rounded short ligule pilose. Panicles few (1-3) at ends of branches, rachis pilose; bracts foliar, bractlets setaceous. Spikelets to 4 mm. long, 'the scales lanceolate, pubescent; hypogynium somewhat 3-lobed; achene globose, to 3 mm. long, white, reticulate-tuberculate. LA PAZ: LARECAJA: San Carlos, Buchtien 1235 (us). COCHABAMBA: CHAPARE: Puerto Polonia, Cárdenas & Cutler 7348 (GH). 18. S. lagoénsis Boeckl. in Vidensk. Meddel. Kjoebenh. 1869: 151 (1870). Perennial from a rather thick rhizome. Culms to 7 dm. high, triangular, the angles scabrid. Leaves to 3 dm. long and 7 mm. wide, the margins and nerves scabrid; sheaths puberulent, the ovate ligule short, pilose on the margins. Inflorescence a terminal and axillary panicle, spikelets few in a cluster; bract foliar, bractlets subulate, 132 Rhodora [Vol. 67 long-attenuate. Staminate spikelets oblong, the scales lanceolate. Pistillate scales lance-ovate, acute to mucronate, brownish, exceeding the achenes; hypogynium 3-lobed, the lobes small, acute; achene 2-3 mm. long, ovoid, white, subreticulate and very sparsely puberulent. LA Paz: NOR YUNGAS: Milliguaya, Buchtien 4283 (Us). 19. S. vaginata Steud. Syn. Pl. Cyp. 179 (1855). Base unknown, but the plant probably perennial. Culms to 1 m. tall, acutely triangu- lar, the angles scabrid. Leaves to 6 dm. long and 1 cm. wide, exceed- ing the inflorescence, glabrous, the nerves of lower surface scabrid; sheaths glabrous, broadly trialate, the wings scabrid, the obtusely rounded glabrous ligule to 1 cm. long. Inflorescence a much-branched purplish panicle; bracts foliar, bractlets subulate. Staminate spikelets to 5 mm. long, few-flowered, scales lance-oblong, acuminate. Pistillate spikelets lance-ovate, purplish, acuminate, the scales exceeding the achenes; hypogynium small, obscurely 3-lobed; achene 3-4 mm. long, ovoid, shining, smooth, white (often tinged with purple). La Paz: LARECAJA: San Carlos, Mapiri, 750 m., Buchtien 1284 (GH). 20. S. melaleuca Reichb. ex Schlechtd. & Cham. in Linnaea, 6: 29 (1831). Perennial from a rhizome. Culms to 9 dm. high, usually glabrous, narrowly trialate, the wings scabrous, the lance-ovate ligule to 1 em. long, glabrous or pubescent, Inflorescence paniculate, terminal and axillary, purple or brownish-purple; bract foliar, bractlets subulate, ciliate. Staminate scales reddish-brown, lanceolate. Pistillate scales purple, suborbicular, acuminate; hypogynium 3-lobed, the lobes rounded, brown; achene subglobose, 2 mm. long, smooth and shining, purplish, occasionally with short pubescence near the base. LA Paz: LARECAJA: San Carlos, Mapiri, Buchtien 333 (NY). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6950 bis (GH). This is probably better treated as a variety of the next species, S. pterota. There seems to be almost no distinguish- ing mark aside from the color of the inflorescence. Since the latest monographer of the genus has retained them as Species, as was also true of S. arundinacea and S. latifolia, I have followed his example, reluctantly. 21. S. pterota Presl in Isis, 21: 268 (1826). Perennial from a thick rhizome, Culms to 3 m. tall, triangular, the angles scabrid, glabrous. Leaves to 4.5 dm. long and 1.5 cm. wide, the margins scabrid; sheaths narrowly trialate, the triangular ligule pilose, to 1 cm. long. Inflo- rescence of axillary and terminal panicles, greenish or yellowish-brown (sometimes purplish) ; bract foliar, large, bractlets filiform. Staminate scales lanceolate, acuminate, greenish-brown or tinged with purple. Pistillate scales orbicular, acuminate, stramineous; hypogynium 3- lobed, the lobes rounded, glabrous or ciliate ; achene to 2.5 mm. long, 1965] Flora of Bolivia — Foster 133 globose or depressed-globose, smooth, white or yellowish-brown. LA PAZ: LARECAJA: Mapiri, Rusby 79 (NY). 22. S. secans (L.) Urb. Symbol. Antill. 2: 169 (1900). S. reflexa of the Catalogue. Perennial from a rhizome. Plants scandent, the culms to 10 m. long, triangular, the angles scabrid. Leaves long, linear, to 5 mm. wide, usually basally pubescent on the upper surface and pilose beneath; sheaths scabrid, apically pubescent, the short ligule ovate, usually with an apical scarious appendage. Inflorescence terminal and axillary; bracts foliar, bractlets filiform-subulate, pubescent. Pistillate scales ovate, glabrous, acute, dark in color; hypogynium subentire, the margin undulate; achene 2-4 mm. long, globose, shining, white or dirty-white, usually minutely puberulent. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 329 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5308 (GH). 14. UNCINIA Pers. Stoloniferous perennials, the leaves mostly basal. Inflorescence a terminal spike, flowers unisexual; female flowers below the apical male flowers on the spike. Female flowers with a subtending scale, a 2-parted utricle, the achene with a trifid style, the rachilla of the spikelet long-exserted and apically hooked; achene oblong, obscurely trigonous. Male flowers with a smaller scale subtending 3 stamens, filaments broad. Inflorescence narrowly cylindrical; rachilla markedly divaricate at maturity; utricles elliptic. ............ e 1. U. hamata. Inflorescence clavate, broadest near the apex; rachilla hardly divari- cate; utricles oblong. ............. nnn 2. U. phleoides. 1. Uncinia hamata (Swartz) Urb. Symbol. Antill. 2: 169 (1900). Plants to 7 dm. tall, the leaves exceeding the stem. Inflorescence to 2 dm. long. Female flowers: glume oblong-ovate, obtuse, ciliolate at the apex; utricles to 5-6 mm. long, exceeding the glumes, the margins coarsely hispid-ciliate, at least on the upper half; exserted portion of rachilla at least as long as the utricle, usually about 7 mm. long; achene oblong-obovate, to 4 mm. long. Male flowers: glumes shorter, abruptly obtuse. LA PAZ: NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 3653 (GH). 2. U. phleoides (Cav.) Pers. Synops. Pl. 2: 534 (1806). Plants to 7.5 dm. tall, leaves to 12 mm. wide. Inflorescence to 15 cm. long. Female flowers: glume oblong, obtusely rounded, finely ciliolate in the upper portion; utricles to 5-6 mm. long, much exceeding the glumes, densely hispid on upper portion; exserted portion of rachilla much shorter than utricle in length; achene narrow, oblong, to 5 mm. long. Male flowers: glumes much shorter and proportionately broader. LA PAZ: LARECAJA: Lancha de Cochipata, 3300-3700 m., Mandon 1424 (GH). 134 Rhodora [Vol. 67 15. canEX L. Perennial from a rather slender scaly rhizome. Culms mostly trigonous, the angles often scabrid or scabridulous, often clothed at the base in the fibrous remains of old leaves. Leaves mostly flat, occasionally reduced to sheaths. Inflorescence of solitary spikes, or spikes aggregated in dense capitula, or obviously paniculate, Flowers unisexual, the staminate flowers consisting of a scale and 3 stamens, borne at the base (gynaecandrous) or apex (androgynous) of a spike (in 1 species, staminate and pistillate flowers borne in separate spikes); pistillate flowers consisting of a scale subtending a special- ized sae (perigynium) containing an achene; achene flattened, bi- convex (lenticular) or trigonous; stigmas 2 or 3. a. Pistillate and staminate flowers in separate spikes on the same plant. REM 5. C. decidua var. Brehmeri. a. Pistillate and staminate flowers in the same Spike. b. Staminate flowers at base of spike (gynaecandrous). c. Achene trigonous; stigmas 3. 9. C. boliviensis. c. Achene not trigonous; stigmas 2. d. Perigynia pale green, plano-convex; beak long; style-base thickened. oo... eeeesescssssescssesscssceseesseseessecseessesesse 3. C. Bonplandii. d. Perigynia olivaceous to brown, concavo-convex; beak short; style-base not thickened. ccc 2. C. pinetorum. b. Staminate flowers at apex of spike (androgynous). e. Achene trigonous; stigmas 3. f. Achene not constricted in the middle. g. Perigynia pubescent to glabrate. ............ 10. C. phalaroides. g. Perigynia glabrous. ........... ees 4. C. cladostachya var. maxima. f. Achene constricted in the middle. ............... 8. C. Jamesonii var. subfulva. e. Achene not trigonous; stigmas 2. h. Achene biconvex and torulose. .............. es 7. C. fecunda. h. Achene not biconvex, not torulose. i. Perigynia green, subalate, ampullaceous, beak long. ........ ————————HÉÉaÁá AMNEM 1. C. nebularum. i. Perigynia deep chestnut or purplish-brown, not alate, not ampullaceous, beak short. ................ 6. C. pichinchensis. l. Carex nebularum Phil. in Anal. Univ. Chile, 93: 492 (1896). Plants to 2 dm. high, the culms trigonous, scabridulous above. Leaves shorter or longer than the culms, to 4 mm. wide, the midrib prominent, at least the margins scabridulous. Inflorescence a dense, terminal, ellipsoid to ovoid, ebracteate capitulum of numerous androgynous (staminate flowers at apex) spikes. Staminate flowers: scales narrow- 1965] Flora of Bolivia — Foster 135 ly lanceolate, to 4 mm. long, acute to somewhat mucronate, the broad margins hyaline or brown-hyaline, the rather prominent midrib brown. Pistillate flowers: scales lanceolate, broader, to 6 mm. long, acute, the margins broadly hyaline, the center pale to dark brown, midrib green; perigynium equaling or shorter than the scale, to 4 mm. long, plano-convex, the over-all shape ampullaceous, subalate, the long beak and upper wings scabrid-serratulate, apex of beak deeply bidentate; stigmas 2; achene small, flattened, pale brown. LA PAZ: MURILLO: La Paz, 3800 m., Buchtien 9245 (GH; det. Kükenthal). 2. C. pinetorum Liebm. in Vidensk. Selsk. Skr. Kjoebenh. (ser. 2) 5: 263 (1851). C. Mandoniana of the Catalogue. Culms mostly to 1 dm. tall, rarely to 3.5 dm., trigonous, the angles smooth or occasionally scabridulous. Leaves mostly shorter than the culms, to 2 mm. wide, scabridulous beneath. Inflorescence a congested ovoid capitulum of 3-5 gynaecandrous (staminate flowers at base) spikes, the bracts and bracteoles mostly setaceous. Staminate flowers: scales ovate, obtuse, brown with a green keel, the apex hyaline. Pistillate flowers: scales ovate, acute, chestnut-brown with a green keel; perigynium to 4 mm. long, slightly exceeding the scale, ovoid, concavo-convex, nerves sub- conspicuous, narrowly alate, the margin ciliate-scabrid, olivaceous, becoming brown, the beak medium long, reddish-brown, bidentate; stigmas 2; achene shining brown, ovoid, style-base not thickened. LA PAZ: LARECAJA: Lancha de Cochipata, 3100-4300 m., Mandon 1429 in part (GH; type-number of C. Mandoniana). 3. C. Bonplandii Kunth, Enum. Pl. 2: 380 (1837). Culms 3 dm. tall, trigonous, the angles scabrid. Leaves shorter or longer than the culms, to 2.5 mm. wide. Inflorescence of 4-10 densely flowered, ovate, ap- proximate, gynaecandrous (staminate flowers at base) spikes, mostly ebracteate. Staminate flowers: scales ovate, obtuse or subobtuse, chestnut-brown, the keel green, hyaline margin broad. Pistillate flowers: scales lance-ovate, subacute, chestnut-brown, the keel green; perigynium to 3-3.5 mm. long, exceeding the scale, plano-convex, ovoid, nerves few and inconspicuous, pale green, subalate, the margins scabrid from the middle upward, beak long, reddish, bidentate; stigmas 2; achene small, ovoid, brown, style-base thickened. LA Paz: LARECAJA: Lancha de Cochipata, Mandon 1429 in part (GH); MURIL- Lo: near La Paz, 3300 m., Bang 62 (GH); 3700 m., Buchtien 9244 (Gu; inflorescence like C. pinetorum but technical details of C. Bonplandii); NOR YUNGAS: Unduavi, 3200 m., Buchtien 6405 (vs). 4. C. cladostachya var. maxima Kükenth. in Engler, Pflanzenr. IV, 20(38): 268 (1909). Culms to 5 dm. high. Leaves longer or shorter than the culms, to 1 cm. wide, scabrid above and beneath. Inflo- rescence paniculate or sometimes apparently racemose, the spikes short-pedicellate; basal bract foliar, exceeding the inflorescence, bracteoles setaceous, equaling or shorter than the spikes. Spikes ovoid, 136 Rhodora [Vol. 67 solitary (occasionally paired), to 1 cm. long, androgynous (staminate flowers at apex). Staminate flowers: scales ovate, about 2 mm. long, the central portion thin, with 3 prominent ribs, light greenish-brown, the broad margins hyaline, apex slightly emarginate, midrib excurrent as a short scabridulous mucro. Pistillate flowers: scales slightly broader, thinner-textured, paler, midrib green, very prominent, mucro longer and more scabrid; perigynium about 2-3 mm. long, exceeding the scale, trigonous, ovoid-ellipsoid, the sides thickened, 3-5 nerves prominent, thin-textured, pale green, gradually contracted into the bidentate beak; achene ellipsoid, trigonous, warm brown, the angles stramineous, the sides densely and evenly papillate; stigmas 3. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5108 (GH); MURILLO: near La Paz, Rusby 74 (NY). Without data: Bang 2210 (GH; type-number). 5. C. decidua var. Brehmeri (Boeckl. Kükenth. in Engler, Pflan- zenr. IV, 20 (38): 307 (1909). C. Goodenowii var. stolonifera of the Catalogue. Plant to 1 dm. high, culms shorter than or about equaling the leaves. Leaves to 3 mm. wide, scabridulous beneath. Inflorescence contracted, of 4-7 spikes, the uppermost staminate, the lower pistillate, bracts small but foliar, mostly exceeding the inflorescence. Staminate flowers: scales to 2.5 mm. long, cuneate to spatulate, obtuse, brown, the midrib and central portion green. Pistillate flowers: scales to 2 mm. long, ovate to obovate, subacute, warm deep brown with a narrow, hyaline, white margin; perigynium to 2 mm, long, not, or only slightly, exceeding the scale, plano-convex, few-nerved, ovate to lance-ovate, papillose, abruptly contracted into a very short entire beak; stigmas 2; achene nearly as large as the perigynium, flattened, finely scrobicu- late, pale brown, the style-base about 0.5 mm. long. LA PAZ: LARECAJA: Cochipata, 3200 m., Mandon 1427 (GH; type-number of C. Brehmeri). 6. C. pichinchensis HBK. Nov, Gen. & Spec. 1: 233 (1815). Culms to 4.5 dm. high, acutely trigonous, the angles scabridulous. Leaves numerous, as long as the culms, to 7-8 mm. wide, glaucous, not scabri- dulous, finely nerved, the midrib usually prominent. Inflorescence a terminal panicle of several to many dense, more or less oblong, androgynous (staminate flowers at apex) spikes, the basal bract long, foliar, sometimes exceeding the inflorescence. Staminate flowers: scales narrowly ovate, to 3 mm. long, acute, deep chestnut-brown with a pale midrib. Pistillate flowers: scales ovate, acute to acuminate, to 3.5-4 mm. long, deep chestnut-brown to blackish-brown; perigynium nearly as long as the scale, ovoid, plano-convex, deep chestnut-brown or purplish-brown, abruptly contracted into a short bidentate beak; stigmas 2; achene oblong to obovoid, pale brown to stramineous, finely pitted, plano-convex. L4 PAZ: LARECAJA: Ticonguaya, 3300-3800 m., Mandon 1426 (Gn); Ingenio del Oro, 3300 m., Rusby 170 (GH); NOR YUNGAS: Unduavi, 3600 m., Buchtien 2596 (GH). SANTA CRUZ: VALLE- GRANDE: Comarapa, 2800 m., Steinbach. 8339 (NY). 1965] Flora of Bolivia — Foster 137 6a. C. pichinchensis var. obtusisquamea R. Gross in Notizbl. 14: 198 (1938). Inflorescence small; scales obtuse or subobtuse, not acute to acuminate. COCHABAMBA: AYOPAYA: Sailapata, 3000 m., Cardenas 3204 (type-number; not seen). 7. C. fecunda Steud. Syn. Pl. Cyp. 194 (1855). Culms to 1 m. high, acutely trigonous, the angles scabridulous. Leaves mostly as long as the culms, to 8 mm. wide, scabridulous. Inflorescence paniculate, the lower bracts foliar, the bracteoles setaceous. Spikes numerous, many- flowered, cylindric, androgynous (staminate flowers at apex). Stami- nate flowers: scales lance-ovate, acuminate to shortly mucronate, reddish-brown, the midsection green, about 3 mm. long. Pistillate flowers: scales like the staminate scales; perigynium a little shorter than the scale, ovoid, plano-convex, the marginal nerves thickened (almost subalate), contracted rather abruptly into a short bidentate beak, the beak-margins sometimes serrate-scabrid; stigmas 2; achene biconvex, pale to medium brown, once or twice contracted near the middle, hence somewhat torulose. LA PAZ: LARECAJA: Cochipata, 3200 m., Mandon 1425 (GH). SANTA CRUZ: VALLEGRANDE: Cerro San Mateo, Comarapa, 3000 m., Steinbach 8390 (NY). 8. C. Jamesonii var. subfulva Kükenth. in Fedde, Rep. Sp. Nov. 8: 7 (1910). Culms to 1 m. tall, but usually shorter, strongly angled, the angles scabrid. Leaves equaling the culms, to 1 cm. wide, with 2 prominent nerves on upper surface. Spikes in a panicle, androgynous (staminate flowers at apex). Scales fulvous, lanceolate, to 4 mm. long, prominently 3-nerved, the apex-margins scabrid; perigynium about equaling the scale, outwardly curved, narrowly ovoid or lanceolate, subtrigonous, nerves inconspicuous to obsolete; stigmas 3; achene trigonous, ecnstricted (subtorulose) in the middle, the style-base not thickened. Without data: Bang 2376 (GH; type-number). 9. C. boliviensis v. Heurck & Muell. Arg. in v. Heurck, Obs. Bot. 32 (1870). Culms to 4.5 dm. high, very slender, decumbent, the upper portion scabrid on the angles. Leaves entirely basal, very much Shorter than the culms, to 2-3 mm. wide. Inflorescence of 2-3 approxi- mate spikes, the uppermost gynaecandrous (staminate flowers at base), the others pistillate, bracts short, hardly foliar, the lowermost slightly longer than the lowest spike. Spikes to 1.5 cm. long, oblong- ovoid. Staminate flowers: scales to 3 mm. long, ovate to obovate, brown, the green midrib prominent but not excurrent, the apex somewhat hyaline at the margin. Pistillate flowers: scales to 3 mm. long, broadly ovate, thin, brown, the central portion green and 3- nerved, with a short blunt mucro, or the mucro absent; perigynium exceeding the scale, lance-ovoid, with 3-5 prominent ribs dorsally and about 5 less prominent ribs ventrally, apically shortly attenuate into the short entire or subentire beak; achene obovoid, sharply trigonous, finely and evenly papillate; stigmas 3, the lower portion densely 138 Rhodora [Vol. 67 glandular-pilose. LA Paz: LARECAJA: near Lacatia, 3200-3300 m., Mandon 1428 (GH; type-number). 10. C. phalaroides Kunth, Enum. Pl. 2: 482 (1837). Culms to 2 dm. tall, obtusely trigonous, smooth, the bases mostly clothed in fibrous remains of old leaves. Leaves exceeding the culms, to 2-3 mm. wide, smooth or scabridulous on the margins and more prominent veins. Inflorescence of 4-6 distant, androgynous (staminate flowers at apex), dense, subglobose spikes to 1 cm. long; bract foliar, exceeding the inflorescence. Staminate flowers: scales narrowly elliptic-ovate, the margins whitish, central portion yellowish-green, the midrib dark green, with a thick scabrid arista less than half the length of the scale. Pistillate flowers: scales similar but much broader, the arista usually longer than the body of the scale; perigynium coriaceous, obovoid-oblong, trigonous, 3-4 mm. long, yellowish-green, densely short-pubescent (the hairs white, thick, curled), 2 lateral nerves prominent, basally long-attenuate or stipitate, the apex abruptly contracted into a short entire beak; achene obovoid, trigonous, the somewhat thickened style-base about half the length of the achene; stigmas 3. TARIJA: MENDEZ: Tucumilla, Fiebrig 2639 (Barros). LA PAZ: MURILLO: La Paz, 3800 m., Buchtien 6407 (US). 10a. C. phalaroides var. moesta (Kunth) Kükenth. in Verhandl. Bot. Ver. Brandenb. 47: 208 (1905). Taller (to 3 dm.), leaves narrower (1-2 mm.). Spikes oblong-clavate, distant, androgynous. Perigynium shorter, the base shortly or hardly at all stipitate, the beak shorter and its apex shortly notched or emarginate. Without data: Bang 1870 (GH). LA Paz: LARECAJA: Cerro del Iminapi, 2650-3950 m., Mandon 1430 (GH). GRAY HERBARIUM, HARVARD UNIVERSITY GENOTYPIC VARIATION IN THE PHACELIA STRICTIFLORA COMPLEX" GEORGE W. GILLETT The Phacelia strictiflora complex occurs from the Texas Gulf Coast to peripheral upland habitats of northeast Mexi- co, the Edwards Plateau, Oklahoma, Arkansas, Louisiana, Mississippi, and eastern Alabama. It embraces populations nominally included under P. strictiflora (Engelm. & Gray) A. Gray; and P. patuliflora (Engelm. & Gray) A. Gray; also a large number of intermediate populations. The two species of this complex were included in a recent biosystem- atic study of the Cosmanthus phacelias, in which a crossing program was completed with eight additional Cosmanthus phacelias (Gillett, in press). Constance (1949) ecognized four varieties of P. strictiflora and two varieties of P. patuliflora. Material grown in this study would corre- spond more nearly to P. strictiflora var. lundelliana Con- stance and to P. patuliflora var. patuliflora. Recognition of the P. strictiflora complex as a distinct evolutionary line seems justified by genetic and morphological evidence presented earlier (op. cit.). The current objective is to present evidence that has an important bearing on the evolutionary relationship of the two species of the complex. Both species of this complex are self-compatible out- crossers with nine pairs of chromosomes. Populations of Phacelia patuliflora occur on lowland habitats, very often on sandy alluvium near sea level along the coast, extending inland to central Texas. Its penetration inland centers along alluvial lowlands between the Colorado and Nueces Rivers. Its coastal distribution extends from the vicinity of Houston to the mouth of the Rio Grande, the southern populations showing considerable gradation to intermediate forms. On the other hand, P. strictiflora is a species of upland habitats and occurs over a wide expanse of territory from northeast Mexico to Oklahoma and eastern Alabama. It is highly variable and very often occurs as a weed in fallow fields, roadsides, and disturbed areas. 'Aided by grants from the National Science Foundation. 139 140 Rhodora [Vol. 67 O — m WwW Bw Fig. 1. Habit; pedicel length; and size and orientation of non- glandular hairs in: (A) Phacelia patuliflora (Gillett 1216 UC) ; and (B) P. strictiflora (Gillett 1257 uc). 1965] Phacelia strictiflora — Gillett 141 A useful comparison of Phacelia strictiflora and P. patuli- flora, is given in the following table: Phacelia strictiflora Phacelia patuliflora Corolla with crenulate to entire Corolla with entire margin. margin. Calyx accrescent. Calyx non-accrescent. Colleters small; 0.1 - 0.2 mm long. Colleters of two size classes: small, 0.1 - 0.2 mm; and large, 0.3 - 0.5 mm long. Habit virgate to ascending. Habit reclining to ascending.* Non-glandular hairs fine, ascend- Non-glandular hairs coarse, ing. spreading.* Pedicel length 0.5 - 1.5 em. Pedicel length 1.5 - 2.0 cm.* *Clinal variation (determined by the study of herbarium specimens). The variability of the last four characters in the above table was determined by direct measurement or by compari- son. Voucher specimens of each species were prepared from plants grown together in the greenhouse and were used for making comparisons with herbarium specimens. Line drawings of these vouchers are shown in Fig. 1. The glandular hairs (colleters) of Phacelia strictiflora and P. patuliflora are of two size classes that show no inter- gradations (Fig. 2). The larger glandular hairs are quite distinct and easily recognized on dried material, but the 5 : M gp feu L.! l^ a O Fig. 2. Glandular hairs (colleters) of the Phacelia strictiflora complex. (A) small glandular hairs of P. strictiflora and southern populations of P. patuliflora (Gillett 1257 uc); (B) large glandular hairs of northern populations of P. patuliflora (Constance and Cory 3247 UC). 142 Rhodora [Vol. 67 smaller become shrunken upon drying, presenting a dis- torted appearance. It was, therefore, quite easy to score herbarium specimens for these. The larger glandular hair is apparently restricted to the northern and interior popu- lations of P. patuliflora. The three additional characters scored in this complex were: (1) the length of the lower pedicels of the inflo- 102 100° 98 96 94" 92° Se | O 25 50 op wën A | o zs $0 100 omnes || EXTREME rom OF PHACELA PATULFLORA D | EXTACME FORM Of PRACELIA STRCTIFLORA [e] BEES d 20x "a IN THE PHACELIA STRICTIFLORA COMPLEX SCORING SYMBOLS PEOICEL LENGTH NOM-GLARDULAR NOIRS — BLAMDULAR NAIRA MAAIL CN 2.0 é COARSE, SPREADING d RECLINING A T D COARSE Q COARSE, SPREADING ó * riw, Mou. Ò Asctams. Ò (0 rat O os rat, ASCENDING O) E.A o Fig. 3. Geographical distribution and variation in Phacelia stricti- flora and P. patuliflora. 1965] Phacelia strictiflora — Gillett 143 rescence (by direct measurement); (2) the cauline pu- bescence of non-glandular hairs (by comparison) ; and (3) habit (by comparison). Each of these three characters is metrical with a continuous spectrum of expression between the extremes portrayed in Fig. 1. The classes scored were the two extreme expressions, each with a representation of near-extremes, and an intermediate class that included an arbitrarily designated central expression along with inter- mediates on either side. In scoring the variation in pedicel length, two intermediate classes were designated, so that four classes were recorded for this character. The scoring for these metrical characters was admittedly unsophisti- cated, but it permitted a graphic portrayal of variability in each character over the entire range of the complex, so that morphological variation may be related to ecological differ- ences. (Fig. 3). The presence of both qualitative (glandular hairs) and quantitative characters in this material probably reflects the genetic make up of many wild plants. This genetic versatility may be even greater, for the metrical or quanti- tative characters could represent more than one of the four basic types of polymeric gene systems cited by Clausen & Hiesey (1958). The four characters portrayed in Fig. 3 are correlated, so that the gene frequencies for the extreme expression of Phacelia strictiflora are relatively higher in the populations in Oklahoma and are progressively lower in those of central Texas, portraying generalized genoclines. On the other hand, the gene frequencies for the extreme expression of P. patuliflora are relatively higher in populations from central Texas (Burnet County) to the Gulf Coast and are progressively lower south to the Rio Grande. It is notable that the intermediate expression of all three metrical characters is found in widely separated populations of the southern Gulf Coast, the Edwards Plateau (about 300 miles northwest), and central Texas. The clinal variation of these characters suggests that the past migrational histories of P. patuliflora, and P. strictiflora have involved contact and introgressive hybridization along the Rio Grande, or along 144 Rhodora [Vol. 67 the northern distributional limit of P. patuliflora, or possi- bly in both regions. It was not possible to confirm experimentally the abundant evidence of natural hybridization between Phacelia stricti- flora and P. patuliflora. In the first cultures of experimental hybrids grown only two flowers of each species were cross- pollinated, with no seed resulting. In later crosses with cultures grown at Turku, Finland, seven flowers of P. patuliflora were pollinated with P. strictiflora pollen. The P. patuliflora stigmas were examined on the stage of a 30X stereoscopic microscope and thus verified to be clean and free of "selfed" pollen before cross-pollination. The ovules of these seven cross-pollinated flowers were analyzed be- tween 25 and 30 days after pollination. Of 69 ovules examined, only five had normal endosperm and apparently normal embryos. The remaining 64 ovules had no visible endosperm, but 14 of these had deformed embryos. This would suggest that the sterility barrier involves failure of endosperm development and incompatibility between the embryo and surrounding tissue as discussed by Stebbins (1958) and reported in PRIMULA by Valentine (1952, 1955). In the latter work, Valentine was able to secure hybrids from over 4000 seeds obtained from his crosses. In the present material of P. strictiflora and P. patuliflora, com- patibility ranges from zero to apparently normal endosperm and embryos in a very limited population of only 69 ovules. It seems most likely, therefore, that a larger number of cross-pollinations might have produced hybrids. It is also possible that additional reciprocal crosses to the P. stricti- flora seed parent might have produced hybrids. Therefore, the incompatibility shown by these experiments appears to be relative and by no means absolute. The taxonomic conclusions of this study provide little satisfaction in terms of readily identifiable species or varieties for the evolutionary dynamics involved here work against, rather than for, discrete taxa. Species limits are established by morphological and often geographical refer- ence points. In P. strictiflora and P. patuliflora a broad expanse of territory contains innumerable intermediates, 1965] Phacelia strictiflora — Gillett 145 each of which is a logical reference point for circumscribing the limit of either species. It is impossible to communicate a species or subspecies limit that can be unerringly con- firmed by subsequent workers because a given intermediate population could be ascribed to P. patuliflora or to P. stricti- flora with equal justification. Combining eco-geographical and morphological criteria, I suggest the reasonable compro- mise of including in P. strictiflora those populations of upland habitats from northeastern Mexico through the Edwards Plateau, and including those populations north of central (Burnet County) Texas. On the other hand, it seems reasonable to circumscribe within P. patuliflora the coastal populations and those contiguous populations extend- ing inland to and including Burnet County, Texas. This is an admittedly imperfect designation because it leaves popu- lations of P. strictiflora within the area ascribed to P. patuliflora, as clearly shown on the map of Fig. 3. ACKNOWLEDGEMENTS This research was carried out at the Department of Botany, University of Turku, Finland, under a Fulbright Lectureship. The space and facilities provided by Professor Antero Vaarama and his staff at Turku are gratefully acknowledged. The line drawings are by Miss Nina Tarén. DEPARTMENT OF BOTANY, UNIVERSITY OF HAWAII LITERATURE CITED CLAUSEN, J. and W. M. HiEsEY. 1958. Experimental studies on the nature of species. IV. Genetic structure of ecological races. Carnegie Inst. Wash. Pub. 615. CONSTANCE, L. 1949. A revision of Phacelia subgenus Cosmanthus (Hydrophyllaceae). Contr. Gray Herb. 168: 1-48. GILLETT, G. W. In Press. Genetic barriers in the Cosmanthus phacelias (Hydrophyllaceae). Rhodora 66: 359-368. STEBBINS, G. L. 1958. The inviability, weakness and sterility of interspecific hybrids. Advances in Genetics 9: 147-215. VALENTINE, D. H. 1952. Studies in British Primulas. III. Hybridi- zation between Primula elatior (L.) Hill and P. veris L. New Phytologist 50: 383-398. . 1955. Studies in British Primulas. IV. Hybridi- zation between Primula vulgaris Huds. and P. veris L. New Phytologist 54: 70-80. INTERPRETATION OF CROSSING DIAGRAMS DAVID L. MULCAHY! Since the taxonomic value of experimentally performed interspecific crosses was so ably demonstrated by the work of Clausen, Keck, and Hiesey (1941), most biosystematic monographs have included one or more of the now classic crossing diagrams. Those first authors included data on the fertility of the F, and the F, generations of crosses between taxa. Similar data, coupled with other information, have been used by many authors in the determination of taxo- nomic affinities, the assumption being that the closer the relationship between taxa, the greater is the probability that they will produce a fertile hybrid. This assumption is generally valid and, when coupled with data from morpho- logical studies, constitutes the basis for a valuable method of analysis. When, however, the conclusions based on crossing diagrams are not substantiated by other studies, gross misinterpretations may result. In a recent study of six species within Oxalis section CORNICULATAE (Oxalidaceae), a series of interspecific crosses was made in an attempt to determine the taxonomic relations between these species. The results of these crosses are shown in Fig. 1. Oxalis corniculata L., O. dillenii Jacq., and O. stricta L. are fully interfertile and also receptive to pollen from O. priceae Small, O. suksdorfii Trelease, and O. grandis Small but the reciprocal crosses fail. Oxalis priceae, O. suksdorfii, and O. grandis are intersterile. According to the general interpretation of crossing diagrams, since Oxalis corniculata, O. dillenii, and O. stricta are interfertile, they should be closely interrelated. Similar- ly, because O. priceae, O. suksdorfii, and O. grandis are not interfertile, they should be distantly interrelated. "The author wishes to express his appreciation to Dr. R. B. Channell, Vanderbilt University, under whose guidance this work was conducted. 146 1965] Crossing Diagrams — Mulcahy 147 O. corniculata «e MERE d O. Priceae y e 2 | Hal Kë 3 | E o Kal 2 EX Ll bb p EI : * a - o "ee $ së CO e * D Ze e, . a Se Ze s 6 See, "e C See Kä * E) Ze, * * - Se Ze a p + *, ry La a * Se ta, o * Se "d 8 EI * E: D * D Ki 4 e Figure 1. Diagram of crosses between six species of Oxalis. Arrows indicate the direction of pollen transfer. Solid lines indicate that seeds were produced. Broken lines indicate that no seeds were produced. 148 Rhodora [Vol. 67 Morphological evidence indicates that Oxalis stricta and O. grandis are closely interrelated while the remaining four species form another natural group. Eiten (1963) refers to these two groups as subsection STRICTAE and subsection CORNICULATAE, respectively. He, however, places O. suks- dorfii in subsection STRICTAE. A combination of morpho- logical data and the results of interspecific crosses, interpreted in the usual way, indicates the phylogenetic sequence represented in figure 2. Other data must be considered however. The species Oxalis corniculata, O. dillenii, and O. stricta are homostylous and predominantly inbreeding. The re- maining three species are heterostylous and mostly out- breeding. In Fig. 2, furthermore, the homostylous species are represented as ancestral to the heterostylous ones, but several facts indicate that, generally, and in Ovzalis in particular, heterostyly represents the ancestral condition. These facts are: 1. Homostylous species are inbreeding. Inbreeding species are considered to be derived (Stebbins, 1957). Therefore, homostyly represents the derived condition. (See also Baker, 1959.) 2. A vast majority of Oxalis species in South Africa are heterostylous (Salter, 1944), as are most South American species (Eiten, 1963). Such a preponderance of heterostyly, O.STRICTA O.CORNICULATA O.DILLENII O. GRANDIS O.SUKSDORFII OPRICEAE Figure 2. The taxonomic relations suggested by a strict interpre- tation of data derived from interspecific crosses. 1965] Crossing Diagrams — Mulcahy 149 especially in these, the generic centers of distribution, strongly indicates the ancestral position of heterostyly within the genus. 3. Biophytum (Oxalidaceae), closely related to Oxalis, is also heterostylous. Since tristyly is known for only three angiospermous families, its evolution must be considered to be a relatively rare phenomenon and its occurrence in two genera of the same family probably indicates a single and early origin of tristyly within the family. If heterostyly represents the ancestral condition in the Oxalidaceae, it must certainly be considered ancestral in Oxalis. These considerations contradict the conclusions based on the interspecific crosses represented in Fig. 1 and a revised phylogenetic sequence (Fig. 3) has been proposed. This revised sequence can be supported by morphological and cytological evidence. Although the homostylous species are interfertile, it is unlikely that they are ancestral to the heterostylous ones. This indicates that experimentally performed interspecific OGRANDIS O.SUKSDORFII4-—J9 0.PRICEAE O. DILLENII Y O.STRICTA 0. CORNICULATA Figure 3. Revised taxonomic relations suggested by a consideration of the reproductive biology of the six species. 150 Rhodora [Vol. 67 crosses cannot be relied upon as an unfailing guide in biosystematic studies. Factors other than taxonomic affini- ties must be capable of influencing the outcome of such crosses. One aspect of the relationships between inbreeding (self compatible) species and outbreeding (self incompatible) species was reviewed by Lewis and Crowe in 1958. Those authors, concerned primarily with pollen tube growth or inhibition, performed many crosses between species, genera, and even families. Combining their data with that already present in the literature, they formulated the following rule: “In the four possible combinations in crossing self- incompatible (SI) and self-compatible (SC) pollentube growth is uninhibited except in the cross SI style X SC pollen, where it is inhibited similarly to SI polli- nation." Lewis and Crowe present data which indicate that the above rule applies to representatives of gametophytic and sporophytic systems of incompatibility, as well as to heter- omorphic systems, although only distylous species are specifically mentioned. Homostyled species generally are self fertile and hetero- Styled ones are not; thus, the results of the present study appear to be largely in conformity with the Lewis and Crowe rule although some crosses were not performed. The homostyled species are interfertile. Furthermore, these three species are receptive to pollen from the heterostyled species, and the reciprocal crosses, pollen from homostyles on the stigmas of heterostyles, fail. The primary indication of this unilateral incompatibility is that it reveals the profound and overriding influence that differences in breed- ing systems have on the outcome of interspecific crosses. Oxalis corniculata cannot be closely related to O. priceae without O. priceae being closely related to O. corniculata. Yet this is what would be indicated by a strict interpretation of crosses between these two species. It is necessary, there- fore, to consider the details of these breeding systems before the crossing diagram can be understood. 1965] Crossing Diagrams — Mulcahy 151 Though heterostylous, Oxalis priceae is self compatible; yet, like a self incompatible species, it cannot be fertilized by self compatible, homostylous species. Thus, it appears to violate the Lewis and Crowe postulate. This anomalous behavior, however, can be explained by considering that several events are involved in the degeneration of self incompatibility. Lewis (1951) indicates that a well established system of self compatibility is the result of three mutational steps. The first step results in pollen which will grow in the style of the flower which produced it, as well as in any other SI style. The properties of the style are, however, not affected by this first mutation. A second mutation destroys the activity of the style so that no pollen is inhibited on it. A final mutation is postulated ‘‘which leaves the pollen un- protected from the inhibitory effect of the SI style." If it can be assumed that Oxalis priceae, like several homomorphic examples listed by Lewis and Crowe (1958), has passed through only the first step in the evolution of self compatibility, its behavior in interspecific crosses be- comes clear. The properties of the style are unaffected and, thus, growth of pollen from homostyles is inhibited, even though the species is self fertile. This assumption implies, however, that the nature of incompatibility in tristylous species is fundamentally the same as that found in non- tristylous groups of the plant kingdom. Whether or not this assumption is valid cannot be decided until further investi- gations have been performed. According to Lewis and Crowe (1951), heterostylous species are expected to be interfertile, as are homostylous ones. The homostylous species are indeed interfertile but the heterostylous ones are not. A possible explanation for this behavior may be that, since these are interspecific crosses, failure to set seeds may be due to genetic difference between the heterostyled species. Under the proposed system of taxonomic relationship (Fig. 3), however, the genetic differences between the homostylous species Oxalis corniculata and O. stricta are expected to be even greater 152 Rhodora [Vol. 67 than those between the heterostylous species. This apparent conflict may be resolved by a consideration of inbreeding species in general. The proximity of anthers to stigmas in homostylous species, coupled with self compatibility, promotes a high degree of inbreeding in homostylous species. This arrange- ment contributes to the success of homostylous species as weeds. A single individual (Baker, 1959) can, in a few seasons, give rise to a large population and thus colonize any suitable niche which may have become available. In- breeding leads to a loss of genetic variability and evolution- ary potential but Stebbins (1957) points out that inbreeding species occasionally hybridize with other taxa, resulting in greatly increased heterozygosity and the production of many new gene combinations. This is followed by a period of intensive selection. Surviving biotypes may give rise to new inbreeding races. This concept implies that introgres- sion is an important source of genetic variability for in- breeding species. Within Oxalis section CORNICULATAE, the homostylous species exhibit great morphological variability, often in the direction of the heterostylous species (Eiten, 1963). This indicates that within these inbreeding species, as in others discussed by Stebbins (1957), introgression is an important source of genetic variation. The selective advantage of introgression will favor, in homostylous species, the se- lection of genes or genotypes which are, in the words of Mayr (1954), “good mixers” i.e., those which will function in a wide variety of genetic environments. Thus, homosty- lous species are expected to be more likely to participate successfully in interspecific crosses than are heterostylous species. This concept may apply to many inbreeding species and thus must be considered in the interpretation of experi- mentally performed interspecific crosses. In view of the above discussion, the crossing diagram (Fig. 1) can obviously be interpreted in two ways. The first possibility, based solely on the crossing diagram and morphological data, assigns to the homostylous species, Oxalis corniculata and O. stricta, a position ancestral to the 1965] Crossing Diagrams — Mulcahy 153 heterostylous species. This necessitates a rejection, in this case, of the rule formulated by Lewis and Crowe, as well as information which indicates the derived nature of homo- styly. The second interpretation is that, although often of considerable value in determining taxonomic relationships, the crossing diagram, because of the differences in the breeding systems between the species involved, provides little useful information in this particular example. The results of interspecific crosses in this instance are de- termined largely by reproductive characteristies of the species involved. The morphological variability of the homostylous species indicates, not that they are ancestral to the heterostylous species, but rather that they have undergone extensive introgression from the heterostylous species. When interspecific crosses are incorporated into bio- systematic studies, it is essential to know which of the taxa involved are self fertile and which are self sterile. Otherwise the several factors which influence the compatibility re- actions may produce very deceptive results. Furthermore, crosses should be carried through the F, and F, generation. This gives a better indication of the genetic similarity between taxa but even these results can be modified by selective forces favoring genetic compatibility or genotypic flexibility. If, because of the nature of the species, further generations cannot be obtained, caution must be exercised during interpretation of the data. DEPARTMENT OF BOTANY, UNIVERSITY OF GEORGIA, ATHENS LITERATURE CITED BAKER, H. G. 1959. The contributions of autecological and geneco- logical studies to our knowledge of the past migrations of plants. American Naturalist 93:255-272. CLAUSEN, J., D. D. KECK, and W. M. HiESEY. 1941. Experimental taxonomy. Carnegie Institute of Washington, Yearbook, No. 40:160-170. EITEN, G. 1959. Taxonomy and Regional variation of Oxalis section CORNICULATAE. Thesis, Columbia University. 154 Rhodora [Vol. 67 —— ——————. 1963 Taxonomy and regional variation of Oxalis section CORNICULATAE I, American Midland Naturalist 69:275-300, Lewis, D. 1951. Structure of the incompatibility gene III — Types of spontaneous and induced mutation. Heredity 5:399-414. Lewis, D. and LESLIE K. Crowe, 1958. Unilateral interspecific incompatibility in flowering plants. Heredity 12:233-256. Mayr, ERNST. 1954. Change of Genetic Environment and Evolution, In, Evolution as a process, edited by J. Huxley, MacMillan Co., New York. SALTER, T. M. 1944. The genus Ozalis in South Africa, a taxonomic revision. Journal of South African Botany Suppl. Vol. 1. STEBBINS, G. L. 1957. Self fertilization and population variability in the higher plants. American Naturalist 91:337-354. II. MONOGRAPH OF THE GENUS ELODEA, SUMMARY. CONTINUED FROM P. 35 HAROLD ST. JOHN Resurrection of the Family Elodeaceae. Key to genera. Egeria, Elodea. Key to subgenera, sections and species. List of Elodea species. Names and synonyms in Elodea. Index to exsiccatae of Elodea. ELODEACEAE Dumortier Analyse des familles des plantes (1829) 54. Submerged perennial aquatics, growing from rootstocks or stolons. Leaves opposite or mostly whorled. Flowers perfect or dioecious, enclosed at first by a spathe; hypanthium in perfect and in pistillate flowers well developed, an elongating thread-like tube; perianth regular; sepals 3; petals 3 (or 0) ; stamens 3 in perfect flowers, or as 3 staminodia in pistillate flowers, or reduced to 1-2; style equaling or exceeding the hypanthium; stigmas 3, and 2-lobed (or entire) ; ovary superior, 1-celled, with 3 parietal placentae; ovules orthotropous or anatropous, several to numerous; capsule ellipsoid, indehiscent or tardily dehiscent; seeds ellipsoid. Staminate flowers of most dioecious species exserted from the spathe by the elongating, slender tubular base of the hypanthium; stamens 9, mostly in 2 series, the 6 outer ones with short filaments, radiating in a lower ring, the 3 inner ones raised on their fused filaments. In E. Nuttallii the staminate flowers sessile, becoming detached and floating to the surface of the water. Lectotype: Elodea Rich. in Michx., Fl. Bor.-Am. 1 (1803) 20. Also in this family is the genus Egeria Planchon. Dumortier based his new family Elodeaceae on the genera Elodea, Anacharis, and Hydrilla. Anacharis is a taxonomic synonym of Elodea. Hydrilla is superfically very similar, but the pistillate flowers are epigynous, so it is here restored to the Hydrocharitaceae. That leaves of the original ones in the Elodeaceae only the genus Elodea, and it is here designated as the lectotype of the family. 155 156 Rhodora [Vol. 67 Though very different in vegetative characters the genus Vallisneria has flowers similar to those of Elodea. However, its staminate spathes have many flowers and the flowers have 1-3 stamens; the pistillate flowers have the ovary and fruit truly inferior. Consequentlv it must remain in the Hydrocharitaceae. There is a publication by Moldenke (1940: 354) which credits St. John with the Elodeaceae. This publication was not authorized by the present writer, and at that time he did not classify Egeria or Elodea as in the Elodeaceae, but rather in the Hydrocharitaceae. NATURAL KEY TO GENERA. Staminate spathes 1-flowered, with a slender stalk-like base, or if sessile, subglobose and containing a sessile staminate flower that is liberated at anthesis, or if with an urceolate spathe the flowers perfect; staminate flowers with 9 stamens, the 3 central ones raised on a common stalk of connate filaments, or rarely the stamens 9 and equal; anthers with 2 locules explosively dehiscing at the axial side of the connective into one widespreading valve (septifragal) which then simulates a heavy ribbed petal; or the flowers perfect, with 3 stamens; filaments smooth, non-glandular, much shorter than the anthers in the staminate flowers; petals smaller than or but little larger than the sepals; nectary lacking; pistillate spathe evenly and shallowly bifid at apex; pistillate flowers with 3 (or 4) entire or bifid or bipartite stigmas; flowers water-pollinated after the ex- plosion of the anthers. Elodea. Staminate spathes 2-4-flowered, sessile, funnelform or ellipsoid; stamens 9 (-10), distinet; filaments papillese glandular above, at least thrice as long as the anthers; anthers each with 2 locules dehiscing by longitudinal lateral sutures into 2 equal valves (loculieidal); petals about thrice larger than the sepals; in the center of the flower a dark nectary; pistillate spathe split halfway down one side; the 3 stigmas 2-3-parted; flowers insect-pollinated. Egeria. EGERIA Planch. Ann. Sci. Nat. Bot. III, 11: 79-81. 1849. St. John in Darwiniana 12: 293-307. 1961. Egeria densa Planch., Ann Sci. Nat. Bot. III, 11 (1849) 80. Elodea densa (Planch.) Caspary, Monatsber. Kgl. Preuss. Akad. Wissensch. 1857 (1857) 48; also in Pringsheim’s Jahrb. Wissensch, Bot 1 (1858) 475 (and 99 in repr.) ; Elodea canadensis Rich. in Michx. var. gigantea Hort. in Bailey, L. H., Standard Cyclop. Hort. 2 (1914) 111; Elodea 1965] Elodea — St. John 157 gigantea Santos, Bot. Gaz. 75 (1923) 44, 50; Elodea densa (Planch.) Casp. var. longifolia Hort. in Parev's Blumengártnerei, ed. by C. Bonstedt (1931) 110; Anacharis densa (Planch.) Victorin, Contrib. Lab. Bot, Univ. Montréal 18 (1931) 41; Philotria densa (Planch.) Small, Man. Southeastern Flora (1933) 28; see St. John, in Darwin- iana 12 (1961) 297-298, 300-302, fig. 1, a-e, 2. Description of all specimens examined: Submersed aquatic; stem stout, 2-8 mm. in diameter; lower leaves in whorls of 3, and 3-7 mm. long, 1.5-2 mm. wide, ovate or lance-ovate; middle and upper leaves in whorls of 4 or 5, and 12-40 mm. long, 1.7-5 mm. broad, oblong to broadly linear, acute, serrulate; nodes close together, but the leaves mostly divergent; flowers dioecious; staminate spathes borne in the upper axils, 11-12 (-14) mm. long, 3.5 mm. broad, elliptical, cleft down one side, the apex bifid with 2 short blunt teeth; staminate flowers 2-4 from each spathe, long stipitate, raised above the water by the slender, thread-like, elongating hypanthium which is 3-6 cm. long; sepals 3-4 mm. long, elliptic-oblong, boat-shaped, dark striate; petals 9.5-11 mm. long, 3.5-9 mm. wide, obovate to suborbicular, subacute at tip, cuneate at base, white, erect; stamens 9 (-10), distinct, 2-2.5 mm. long, the central ones not connate; filaments heavy, clavate, flattened, glandular papillose above; anthers 0.5-1.2 mm. long, unequal; nectary central, green, 3-lobed, secreting abundant nectar; pistillate plants unknown to the writer, not represented in any of the numerous herbaria examined, including that of L. Hauman-Merck in Brussels, but described in detail by him while in Buenos Aires, and the following description is drawn from a translation of his article in Rec. Inst. L. Errera 9 (1913 = 1912) 34-35; pistillate spathe 10-11 mm. long, entire or slightly cleft along the median line, 1-flowered; calvx like the above; corolla smaller, the petals 8 mm. long, 8 mm. wide, less regularly plaited; the 3 staminodia 2.5 mm. long, filiform, sharp, vellow, covered with papillae, alternating with the branches of the style; style white, trifid to the base, with filiform lobes 3 mm. long, deeply 2-3-cleft and covered with papillae for their whole length; at the base of each lobe towards the cutside, a little green nectary; ovary surmounted by a thread-like stalk 2-6 cm. long, 1 mm. in diameter; ovary cavity small, 1-celled, producing a swelling at the base of the neck; ovules 3-6, erect; fruit 7-8 mm. long, 3 mm. in diameter, cbscurely trigonous, attenuate at the summit and sur- mounted by a vestige of the neck of the ovary; pericarp membranous, transparent; seeds 7-8 mm. long, (rare), sessile, fusiform, completely filling the locule owing to their swollen, mucilaginous seed coat and papillae, surmounted by a slender 2 mm. filament. Holotype: “in ditione Platensi, prope Bonariam; Tweedie in herb, Hook." (= Buenos Aires, Argentina), (K)! Type examined! Range: Southern Brasil, Paraguay, Uruguay, and northern Argentina. Egeria densa is universally cultivated in biological labo- 158 Rhodora [Vol. 67 ratories, and frequently grown in aquaria and water cultures. It has escaped from cultivation and has established itself in the waters of Massachusetts, New York, New Jersey, Pennsylvania, Maryland, Virginia, North Carolina, South Carolina, Georgia, Florida, Kentucky, Alabama, Louisiana, Texas, Nebraska, Arkansas, Missouri, Arizona, Oregon, and California. It is also established in Germany, France, Italy, Japan. ELODEA L. C. Richard in Michaux Perennial submerged aquatic herbs in fresh (or rarely brackish) water, rooting in the (usually muddy) bottom or free drifting when broken loose; roots unbranched, smooth, with a root cap; stems slender, simple or sparsely dichotomously branched; lowest leaves alternate, opposite, or in whorls of 3, and reduced; middle and upper leaves opposite or in whorls of 3-7, linear to oblong, acute or obtuse, sharply serrulate, 1l-nerved; stipules minute, evanescent; flowers hypogynous, and in a few species perfect, with 3 alternate stamens and 3 stigmas; flowers of most species dioecious and raised to the surface by a slender hypanthium enclosing the style; sepals 3, elliptic, chartaceous; petals 3 (rarely 0), membranous, white to purple, elliptie to linear; staminate flowers with 9 stamens, usually 6 in a lower, outer ring, and the inner 3 raised on a fused filament column; anthers bilocular, explosively septifragal; pollen exine spiny; pistillate flowers with 3 alternate staminodia; common style running up through and slightly exceeding the hypanthium tube; style branches 3; stigmas 3, bifid (or entire), exceeding the perianth; ovary unilocular with 3 parietal placentae and/or a basal placenta; ovules several, orthotro- pous or anatropous; fruit capsular; seeds cylindric to fusiform, glabrous, papillose, hirsutulous, or mucilaginous. The writer has been doing research on Elodea since 1919, has studied it in the field and in many of the major and minor herbaria of the world. He has seen the holotypes of all previously described species and minor taxa. Like most other flowering plants, the species of Elodea have their most constant and best diagnostic characters in the structure of their flowers. These are tiny, so in our illustrations they are shown five times magnified. For the benefit of curators of herbaria and for students of local floras, the distribution of each species is mapped, and all the collections that have been studied, are cited, but this is done as briefly as possible. Such a detailed presentation is valuable, as the identification of collections of Elodea has 1965] Elodea — St. John 159 proven so difficult that the individual botanist has usually felt uncertain of his determinations. The natural ranges of the species of Elodea are of inter- est. In North America only E. canadensis spans the conti- nent, from Quebec to British Columbia, south to Alabama, and California. This species is also the one which, when introduced to Ireland in 1836, soon ran rampant over Europe, and now is also established in Asia, Africa, and Australia. Elodea Nuttallii has the next largest range, being abundant in the lowlands from Maine to North Carolina, and occurring westward to Missouri, and also in Idaho (where it is probably introduced). The other North Ameri- can species are more restricted in occurrence, and several are known only from single localities. In South America, too, there is one widespread species, Elodea granatensis Humb. & Bonpl. It occurs from Colombia eastward to French Guiana, and southward to Argentina. The other species of that continent have much smaller ranges. Between the northern border of Mexico and the southern border of Panama there are no known collections of indige- nous Elodea. The genus does not occur naturally in any part of the West Indies or Central America. The North American species are all of temperate regions, but several of the South American ones are of the tropics, so the absence in Central America cannot be due solely to the climate, nor does it seem to be due to lack of collecting. The writer can offer no explanation of this broad gap between the areas occupied by the 9 North American and the 8 South American species. Many botanists have assisted the writer during his pro- longed investigation. They are too many to thank individu- ally, but one of them must be singled out. Dr. J. E. Dandy, Keeper of Botany, Natural History Museum, London, has contributed much from his knowledge of aquatics and their nomenclature. GENERAL MORPHOLOGY — The members of the genus Elodea are submerged aquatics, rooting in the bottom, or free floating (when broken loose). Their growth is erect until they reach the surface of the water. 160 Rhodora [Vol. 67 The roots are slender, white or pale, and unbranched. They have a root cap, but no root hairs. Adventitious roots are freely produced from nodes of the stem, and most abundantly so on detached, drifting branches. The stems are erect, rather weak and brittle, cylindric, simple or dichotomously branched. The lowest leaves are alternate, opposite, or in whorls of three. The middle and upper leaves are opposite or in whorls of 3-7. They are from linear to oblong, acute or obtuse, and sharply serrulate. Their venation consists of a single midrib, and the thin, bright or pale green blades are translucent. Axillary scales or stipules, entire or fimbriate, are present and visible on fresh specimens. They are minute and evanescent, and on dried specimens are invisible. The flowers are solitary, axillary, and in bud are enclosed in a membranous, globose or urceolate spathe which opens at the bifid apex. Hypanthium (present in most species) filiform, elongating and raising the bud to the surface where it expands. Flowers perfect and with only 3 stamens in a few species, or as in most species, dioecious. Sepals 3, elliptic, concave, firm, chartaceous, in part greenish and in many streaked with black or purple. Petals 3, membranous, white or bluish or the outer part purple, elliptic to spatulate or linear. Staminate flowers with 9 erect stamens, of which 6 are in an outer, lower ring, and the inner 3 are mostly elevated on a smooth, fused filament column, or rarely all of equal length and attachment. Anthers with 2 locules, these septifragal, explosively dehiscent at the axial side of the connective, becoming flat and resembling heavy ribbed petals. Pistillate flowers with similar hypanthium, sepals and petals. Staminodia 3 and diverse in the different species, being a mere subulate filament or a filament with more or less abortive, sterile anther remnants. Ovary 1-celled; ovules several, borne on parietal or basal placentae. Style filiform, exceeding the hypanthium which surrounds it, the apex 3-lobed. Stigmas 3, ligulate or spatulate, deeply bifid (or entire), reddish papillose on the upper receptive surface. Fruit capsular; seeds cylindric to fusiform, glabrous, papil- lose, hirsutulous, or mucilaginous. 1965] Elodea — St. John 161 POLLEN GRAINS — The pollen grains are nonaperturate, 94-125 „ long, and the surface bears thin, sharp spinules, separate or on a connecting reticulum. The grains become separate or remain attached in tetrads. Apparently only the following species have had their pollen described: E. calli- trichoides, E. canadensis, E. Ernstae, and E. Matthewsü (Erdtman 1943: 62, pl. 2, fig. 22; 1952: 207-208). CHROMOSOMES — The 2X number of chromosomes is 16 in Elodea callitrichoides, and 24 in E. canadensis. The related genus Egeria has in its species E. densa 48 chromo- somes (Darlington & Janaki Ammal, 1945: 276). The record of the number 48 for Elodea canadensis is evidently due to examination of a misidentified specimen of Egeria densa. POLLINATION — Nearly all of the species of Elodea are dioecious. Their flowers grow upward, raised by the elongating hypanthium, till they reach the surface of the water. There the flowers expand, the three sepals spread outward and reflex slightly, and their waxy outer surfaces repel water and serve as floats to uphold the flower. In tlie pistillate flowers the longest, heaviest parts are the three bifid stigmas which project beyond the perianth parts, overweight it, causing it to partly flop over on one side. Thus the two nearest stigmas arch over and their tips rest on the water surface. Due to their weight and unwettable nature, they push down and cause a depression of the surface film of the water. The staminate flowers, in all the dioecious species but one (E. Nuttallii), in growth are raised to the water surface by an elongating, thread-like hypanthium, just as are the pistillate ones. The anther dehiscence is by an explosion which scatters the pollen grains on the nearby water surface. The outer coat of the pollen grains is covered with spines which entrap air, hold back the water, and give buoyancy to the grains. They float and bob along on the surface of the water. When and if they drift to a pistillate flower, the grains slide down into the depression of the water surface around a stigma, contact its sticky papillose 162 Rhodora [Vol. 67 surface, and are in position to accomplish pollination and fertilization. This ingenious method of pollination is effective, and it does work. Fertilization does occur and seeds are formed, but rarely. It is quite clear that the two sexes of the species are not equal in number. In the more local and rarer species, mere chance has determined whether the staminate or the pistillate was collected and hence is the only sex known. Among the other species of wide distribution, in which both sexes are known, there is still an evident disparity. In collections the pistillate plants are many times more abundant than the staminate. It appears that seed forma- tion is rare. DISPERSAL — Ripe seeds, freed in the water of the lake or stream, may be carried some distance by moving water and allow the establishment of a new colony. From the rarity of seed production, this natural dissemination seems to be a minor method of dispersal of a species. The branches of Elodea plants are slender and rather brittle. They bend and sway with water currents, but are easily broken. Detached branches, still submerged, drift away. Unless stranded and dessicated they retain life and continue growth. Adventitious roots are produced at the nodes, and by them the loose branch may again be rooted in the bottom of the body of water. It seems very clear that this vegetative propagation by fragmentation is the princi- pal method of dispersal of the species, Species of Elodea may also be dispersed by water birds. Stems removed from the water and exposed to the air for 23 hours, and then replaced in water have recovered in one day. Rootstocks of E. canadensis have been found in the nests of the common tern on islands on the coast of Sweden (Ridley, 1930: 537-538). Such deliberate or accidental transport by birds is apparently the principal means of spread of the plant from one water system to another. That these methods are effective, has been evidenced by the phenomenal spread of the pistillate plant of E. canadensis throughout Europe. DISTRIBUTION — The genus Elodea contains at present 17 1965] Elodea — St. John 163 species, 9 of which occur in North America, and 8 in South America. In North America the indigenous species occur across southern Canada from Quebec to British Columbia, and southward in the United States to Alabama, Mississippi, New Mexico, Arizona, and California. The published records of occurrence in Texas are unsubstantiated. In South America native species are known from Colom- bia southward to Peru, at both low and high altitudes. They also occur from Colombia eastward to French Guiana, and south to Argentina. In the West Indies there are no records of Elodea. In Central America, the only records are in Mexico, of Elodea canadensis and Egeria densa, recently established at a few localities near Mexico City. They are doubtless adventives, escaped from culture in gold fish bowls or tanks. Good habitats are numerous both in the temperate and in the tropical sections of the region. Having been created in one of the Americas, the genus must have spread to the other one across Central America. Its absence in Central America now seems to be a fact and not a mere apparent absence due to lack of collecting. Panama, Costa Rica, Guatemala, and Mexico have been explored too long and too thoroughly for that hypothesis to receive any credence by the writer. PHYLOGENY — Most of the species of Elodea are dioecious or functionally dioecious. However, in their pistillate flowers they all have staminodia, three in number, alternate with the petals. These staminodia are abortive in various degrees. The extreme in reduction is to a slender, subulate structure, representing solely a filament. Other species have an expanded apex, bidentate or retuse, representing an abortive remnant of an anther. These diverse sorts of staminodia evidence an evolutionary trend towards loss of stamens in the flowers. It would not be reasonable to postu- late evolution in the other direction, — that these staminodia were stages in the development of perfect stamens. Hence, it seems clear that in this genus the more primitive type of flower is the bisexual or perfect one. These more primitive 164 Rhodora [Vol. 67 Species are the South American E. granatensis and the North American E. Brandegeae, and E. Schweinitzii. The second and third of these are rare and very local, but the first is common and with a very broad range. Among these three the writer cannot single out one as the most primitive species. NATURAL KEY TO THE SUBGENERA, SECTIONS AND SPECIES OF ELODEA Flowers perfect. subgenus Apalanthe. Stigmas entire, included, equaling the filaments; median and upper leaves in whorls of 3. 2. E. Brandegeae. Stigmas exserted, Stigmas 3, bifid, equaling the anthers; middle and upper leaves in whorls of 7 (or 5). 8 (part 1), 13 (part 3). E. granatensis. Stigmas 4 (or 3), usually entire; middle and upper leaves in whorls of 3. 17. E. Schweinitzü. Flowers dioecious. subgenus Elodea. Staminate flowers sessile, at anthesis liberated, floating to and expanding on the surface of the water. section Natator, with a single species. 7 (part 1), 18 (part 4). E. Nuttallü. Staminate flowers protruded from the spathe by the elongating, filiform base of the hypanthium, not liberated. ` section Elodea. Section ELODEA Dioecious species with both sexes known. Stigmas entire. Staminodia 1.5 mm. long, obtuse; pistillate spathe 3-7 em. long, the apical teeth divergent; staminate petals 0.6 mm. broad, linear. 5 (part 1), 21 (part 4). E. longivaginata. Staminodia 0.8 mm. long, bifid; pistillate spathe 1.8 em. long, the apical teeth erect; staminate petals 1.5 mm. wide, oblanceolate; pistillate sepals oblong-elliptic. 12. E. titicacana, Stigmas bifid. Stigmas bifid for half or more than half of their length. Anthers 4 mm. long; middle and upper leaves in whorls of 3-4. 11. E. Potamogeton. Anthers 3.5 mm. or less in length. Middle and upper leaves opposite or occasionally some of them in whorls of 3. 14. E. callitrichoides. Middle and upper leaves in whorls of 3-4. Pistillate petals wanting; staminate petals 0.4-0.6 mm. wide, linear, acute; middle and upper leaves in whorls of 3-4. 16. E. Richardi. Pistillate petals present; staminate petals 1.1-2 mm. wide, somewhat dilated, obtuse; middle and upper leaves in whorls of 3. 1965] Elodea — St. John 165 Pistillate sepals oblong-deltoid; stigmas 5-5.5 mm. long; staminate petals 1.1 mm. wide, slightly dilated. 15. E. Ernstae. Pistillate sepals oblong to elliptic; stigmas 3.5-9.8 mm. long; staminate petals spatulate. Pistillate sepals 3 mm. long; staminate sepals ovate; staminate petals 1.5 mm. wide. 10. E. peruviensis. Pistillate sepals 2.4-2.6 mm. long; staminate sepals elliptic; staminate petals 1.8-2 mm. wide. 9. E. Matthewsii. Stigmas bifid for a third or less than a third of their length. Pistillate sepals 2-2.2 mm. long; staminate petals slightly dilated near the apex; middle and upper leaves in whorls of 3. 3 (part 1), 19 (part 4). E. canadensis. Pistillate sepals 1.4 mm. long; staminate petals linear; middle and upper leaves opposite or occasionally in whorls of 3 at a few of the upper nodes. 1. E. bifoliata. Dioecious species of which only one sex is known. Leaves 0.5-1.5 mm. broad, the median and upper ones in whorls of 3. Leaves less than 1 mm. broad; pistillate flowers unknown; staminate petals 0.5 mm. wide; stamens equal, all attached to summit of hypanthium. 20. E. linearis. Leaves 0.5-2 mm. broad; pistillate flowers known; petals 1.1 mm. wide. Staminodia obdeltoid, petaloid throughout; middle and upper leaves 0.5-1.5 mm. wide, flaccid. 4. E. columbiana. Staminodia subulate at base, the tip petaloid, elliptic; middle and upper leaves 1-2 mm. wide, thin but not flaccid. 6. E. nevadensis. LIST OF ELODEA SPECIES. Monograph, part 1. Res. Stud. Wash. State Univ. 30: 19-44. 1962. . bifoliata St. John . Brandegeae St. John . canadensis Rich. in Michx. . columbiana St. John . longivaginata St. John . nevadensis St. John . Nuttallii (Planch.) St. John ograph, part 2. Caldasia 9: 95-113, 1964. S 300 4 ctn S Fe eee ee Mo 8. E. granatensis Humb. & Bonpl. 9. E. Matthewsii (Planch.) St. John 10. E. peruviensis, St. John 11. E. Potamogeton (Bert.) Espinosa 12. E. titicacana, St. John 166 Rhodora [Vol. 67 Monograph, part 3. Darwiniana 12: 639-652. 1963. 13. (8.) E. granatensis Humb. & Bonpl. 14. E. callitrichoides (Rich.) Casp. 15. E. Ernstae St. John 16. E. Richardii St. John Monograph, part 4. 17. E. Schweinitzii (Planch.) Casp. 18. (7.) E. Nuttallii (Planch.) St. John 19. (3.) E. canadensis Rich. in Michx. 20. E. linearis (Rydb.) St. John 21. (5.) E. longivaginata St. John NAMES AND SYNONYMS IN ELODEA, Synonyms are in italie and accepted names in roman type. The names published in Elodea Juss. (an orthographic variant of Elodes Adans.), and like it in the Hypericaceae, are omitted here. Anacharis A. Alsinastrum Babington, Ann. & Mag. Nat. Hist. II, 1: 83-84, 86, 1848; Ann. Sci. Nat. Bot. III, 11: 74, 1849 = Elodea canadensis. A. callitrichoides Rich., Mém. Inst. de France 12(2): 7-8, 75, pl. 2 bottom, 1811 = 1814 = Elodea callitrichoides. A. canadensis Planch., Ann. & Mag. Nat. Hist. II, 1: 86, 1848; Ann. Sci. Nat. Bot. II, 11: 75, 1849, independent species, not based on Elodea canadensis Rich. in Michx., but = Elodea canadensis. var. latifolia (Casp.) Sanio, Verhandl. Bot. Verein Prov. Brandenburg 32: 121, 1890 — Elodea canadensis. var. latifolia (Casp.) Sanio, var. repens Sanio, Verhandl. Bot. Verein Brandenburg 32: 121, 1890, invalid, a variety under a variety. var. Planchonii (Casp.) Victorin, Contrib. Lab. Bot, Univ. Montréal 18: 40, 1931 — Elodea canadensis. A. chilensis Planch., Ann. & Mag. Nat. Hist. II, 1: RF, 1848, Ann. Sci. Nat. Bot. III, 11: 75, 1849 — Elodea Potamogeton. A. densa (Planch.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 41, 1931 = Egeria densa. A. Hilariana Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 49, 1857 (in repr. p. 13); Pringsheim's Jahrb. 1: 476, 1858, published in the synonymy of Elodea Naias — Egeria Naias. A. linearis (Rydb.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 41, 1931 = Elodea linearis. A. Matthewsii Planch., Ann. & Mag. Nat. Hist. II, 1: 86, 1848 = Elodea Matthewsii. A. Naias (Planch.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 42, 1931. The authorities were published as “(Casp.) Victorin.” = Egeria Naias. A. Nuttallii Planch., Ann. & Mag. Nat. Hist. II, 1: 86, 1848; Ann. Sci. Nat. Bot. III, 11: 74, 1849 — Elodea Nuttallii. 1965] Elodea — St. John 167 A. A p. occidentalis Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 40, 1931, based on Serpicula occidentalis Pursh, a superfluous name which cannot be transferred — Elodea Nuttallii. . Planchonii (Casp.) Rydb., Fl. Prairies & Plains Cent. N. Am. 57, 1932 — Elodea canadensis. . Planchonii (Casp.) M. E. Peck, Man. Higher Pl. Ore. 76, 1941 — Elodea canadensis. . pomeranica (Reichenb.) Peterm., Deutschl. Fl. 530, t. 82, fig. 650 a, b, 1849. The description and the leafy shoot, fig. 650 = Hydril- la verticillata (L. f.) Royle. The staminate and pistillate flowers shown in fig. 650 — Elodea canadensis. . Potamogeton (Bert.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 41, 1931 — Elodea Potamogeton. . Tripteris (L.) Steud., Nom. Bot., ed. 2, 1: 82, 360, 1840. This was an obvious error, not a combination intended by the author, as is confirmed under Chrysostemma on page 360. = Coreopteris Tripteris L. A palanthe . granatensis (Humb. & Bonpl.) Planch., Ann. & Mag. Nat. Hist. II, 1: 87, 1848; Ann. Sci. Nat. Bot. III, 11: 76, 1849 — Elodea granatensis. guyannensis (Rich.) Planch., Ann. & Mag. Nat. Hist. II, 1: 87, 1848; Ann. Sci. Nat. Bot. III, 11: 76, 1849 = Elodea granatensis. . Schweinitzii Planch., Ann. E Mag. Nat. Hist. II, 1: 87, 1848; Ann. Sci. Nat. Bot. III, 11: 76, 1849 — Elodea Schweinitzii. Babingtonia . pestifera. Syme in Sowerby, English Bot. ed. 3, 9: 83, 1869, published in synonymy = Elodea canadensis. Diplandra . Potamogeton Bert., Mercurio Chileno 13: 612, 1829 = Elodea Potamogeton. Elodea . bifoliata St. John, Res. Stud., Washington State Univ. 30: 23-24, figs. 3, b, c, 4, a, b, 5, 1962. . Brandegeae St. John, Res. Stud. Wash. State Univ. 30: 25-26, Ses, 2, c, d, 4, m, n, 5, 1962. . callitriehoides (Rich.) Casp. Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 47-48, 1857 (in reprint pp. 11-12). The combi- nation was made by Caspary, but with a question mark. Yet, it was a valid publication. var. ernstae Erdtman, Pollen Morphology and Plant Taxonomy, Angiosperms 207-208, 1952. The variety lacks a diagnosis, and is invalid — Elodea Ernstae. . canadensis Rich. in Michx., Fl. Bor.-Am. 1: 20, 1803. var. B angustifolia Aschers. & Graebn., Syn. Mitteleur. Fl. 1: 403, 1897, nomen nudum. 168 Rhodora [Vol. 67 var. angustifolia (Britton ex Rydb.) Farw., Am. Midl. Nat. 10:203, 1927 — E. Nuttallii. var. gigantea Hort., Bailey, Stand. Cyclop. Hort. 2: 1,111, 1914 — Egeria densa. var. latifolia (Casp.) Sanio var. repens Sanio, Verhandl. Bot. Vereins Prov. Brandenb. 32: 121, 1891 = 1890, a variety under a variety, and hence invalid. var. latifolia Aschers. & Graebn., and forma latifolia Aschers. & Graebn., Synops. Mitteleur. Fl. 1: 403, 1897 — E. cana- densis. var. Planchonii (Casp.) Farw., Am. Midl. Nat. 10: 203, 1927 = E. canadensis. . capensis M. Ernst, Ber. Schweiz. Bot. Ges. 55: 35, 1945, published in synonymy. . chilensis (Planch.) Casp., Monatsber. Kgl. Preuss. Akad. Wis- sensch. 1857: 47, 1857 (in repr. p. 11) — E. Potamogeton. . chinensis Casp. ex. Ind. Kew. Suppl. 1: 152, 1901-1906, misprint for E. chilensis. . columbiana St. John, Res. Stud., Wash. State Univ. 30: 37-38, figs. 3, d, 4, g, h, 5, 1962. . crispa Hort. ex Henkel, Haupt-Katalog 26, 1908 = Lagarosiphon major (Ridl.) Moss. - densa (Planch.) Casp. Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 49, 1857 (in repr. p. 13). The combination was made with a question mark, but was a valid one = Egeria densa. var. crispa Hort. ex Wehrh., Gartenstauden 1: 10, 1929, published in Synonymy = Hydrilla verticillata Royle var. crispa Casp. var. longifolia Hort. ex Bonstedt in Parey's Blumengaertnerei 1: 110, 1930, — Egeria densa. . Ernstae St. John, Darwiniana 12: 644, 646, 648, fies. 1, a, b, d, e, 3, 1963. . gigantea Santos, Bot. Gaz. 75: 44, 50, 1923 = Egeria densa. . granatensis Humb. & Bonpl., Pl. Aequin. 2: 150, pl. 128, 1813. . guyannensis Rich., Mém. Inst. France 12(2) : 4, 75, pl. 1, 1811 = 1814 — E. granatensis. var. dicranoides Casp., Monatsber. Kgl. Preuss. Akad. Wis- sensch. 1857: 48, 1857 (in repr. p. 12), and in Pringsheim’s Jahrb. 1: 502, 1858 — E. granatensis. f. longifolia Chodat & Hassler, Bull. Herb. Boiss. II, 3: 1,033, 1903 — Egeria Naias. E. ioensis Wylie, Nat. Hist. Bul, State Univ. Iowa 6(4): 48-50, pl. 1-2 1913 — E. canadensis. E. iowensis Wylie (as Iowensis), Proc. Iowa Acad. Sci. 17: 82, 1910, momen provisorium, — E. canadensis. E. Kochii Herter, Revista Sudamer. Bot. 6: 134, fig. 2, 1940 = E. Naias. &y RE mH tot N mri m 1965] Elodea — St. John 169 E fy bj latifolia Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 46, 1857 (in repr. p. 11), published as a doubtful species, based upon sterile material — E. canadensis. . linearis (Rydb.) St. John, made above in this paper. . longivaginata St. John, Res. Studies, Wash. State Univ. 30: 38-40, figs. 2, e-j, 4, c, d, 5, 1962. . Matthewsii (Planch.) St. John ex Erdtman (as Plach.), Introd. Pollen Analysis 62, pl. II, fig. 22, 1943. It has the measurements of a pollen grain, and an outline drawing of one grain. It lacks the reference to the basionym, and the publication was not authorized by St. John. Erdtman apparently saw the binomial written upon an herbarium sheet. . Matthewsii (Planch.) St. John, Darwiniana 12: 307, 1961; Caldasia 9: 101, 108, figs. 4, a, b, 5, a, b, 8, 1964. E. minor (Small) Farwell, Rept. Mich. Acad. Sci. 17: 181, 1916 — e So & PR y ba pm bM a E. Nuttallii. . Naias (Planch.) Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 49, 1857 (in repr. p. 13-14) ; and altered to E. Najas in Pringsheim's Jahrb. 1: 476-477, 503, 1858. — Egeria Naias. . nevadensis St. John, Res. Stud., Wash. State Univ. 30: 41-43, figs. 8, a, 4, i, j, 5, 1962. . Nuttallii (Planch.) St. John, Rhodora 22: 29, 1920, sensu basionym, Anacharis Nuttallii Planch., non sensu St. John (1920). . oblongifolia Michx. ex Casp., Pringsheim's Jahrb. 1: 462, 1858, published in synonymy = E. canadensis. . occidentalis St. John, Rhodora 22: 27-29, 1920. This was ostensibly a combination based upon Serpicula occidentalis Pursh, excluding his synonym E. canadensis Michx. The epithet canadensis was available, and Pursh should have adopted it, so his new epithet occidentalis is illegitimate and cannot be transferred. St. John also cited in synonymy Philotria minor Small. The epithet minor was available and should have been adopted, so St. John’s epithet is also illegitimate = Elodea Nuttallii. . orinocensis Rich. (as Orinocensis), Mém. Inst. de France 12(2): 75, 1811 — 1814, — E. granatensis. . paraguayensis Herter, Revista Sudamer. Bot. 6: 134, fig. 3, 1940 — Egeria Naias. . peruviensis St. John, Caldasia 9: 103, 105, 107, figs. 3, a, b, 5, g. h, 6, 8, 1964. . Planchonii Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 47, 1857; and Pringsheim's Jahrb. 1: 468-469, 500-501, 1858 — E. canadensis. This was based upon Anacharis canadensis Planch. and doubtfully on A. canadensis Chatin. . Potamogeton (Bert.) Espinosa, Rev. Chil. Hist. Nat. 31: 150-155, fig. 10, 10(2), 1928. . Richardii St. John, Darwiniana 12: 649, 651, figs. 1, f, g, 2, a, b, 3, 1963. 170 Rhodora [Vol. 67 E. Sehweinitzii (Planch.) Casp., Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 46, 1857 (in repr. p. 10-11) ; Pringsheim's Jahrb. 1: 468, 500, 1858, published with a question mark as a doubtful species, yet it is valid. E. titicacana St. John, Caldasia 9: 111-112, figs. 3, d, e, 5, c, d, 8, 1964. 7. verticillata (L. f.) F. Muell, Key Syst. Vict. Pl. 1: 423, 1888 = Hydrilla verticillata (L. f.) Royle. Hapalanthe H. Sehweinitzii Planch, ex Peterm, Deutschl. Fl. 530, t. 82, fig. 650 e, 1849. The generic name is merely an orthographic variant of 'Apalanthe, and the species = Elodea Schweinitzii. Luchia L. Berteroniana Steud., Nom. Bot. ed. 2, 2: 75, 1841, nomen nudum — Elodea Potamogeton. Philotria P. angustifolia Britton ex Rydb., Fl. Colo. 15, 1906 — Elodea Nuttal- lii. Rydberg in Bull. Torrey Bot. Club. 35: 460, 1908 stated that he applied the name to a different species, that is to Philotria Planchonii (Casp.) Rydb. — Elodea canadensis. P. densa (Planch.) Small, Man. S. E. Fl. 28, 1,503, 1933 = Egeria densa. P. Canadensis (Rich. in Michx.) Britton, Science II, 2: 5, 1895 — Elodea canadensis. P. granatensis (Humb. & Bonpl.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 42, 1931 — Elodea granatensis. P. guyannensis (Rich.) Victorin (as guyanensis), Contrib, Lab. Bot., Univ. Montréal 18: 42, 1931 — Elodea granatensis. P. Iowensis Wylie, Proc. Iowa Acad. Sci. 17: 82, 1910, published as a synonym of Elodea iowensis, a provisional name = Elodea canadensis. P. Iowensis Wylie, Science, n. s. 33: 263, 1911 — Elodea canadensis. P. linearis Rydb., Bull. Torrey Bot. Club 35: 464, 1908 = Elodea linearis. P. minor Small, Fl. S. E. U. S. 47, 1903, published as (Engelm.) Small, but Engelmann's Udora verticillata var. minor was not validly published — Elodea Nuttallii. P. Nuttallii (Planch.) Rydb., Bull. Torrey Bot. Club 35: 461-462, 465, 1908, a provisional name, and hence invalid — Elodea Nuttallii. P. Nuttallii (Planch.) Rydb. ex Britton & Brown, ILL. Fl. N. E. U. S., ed. 2, 1: 105, 1913 ~ Elodea Nuttallii. P. occidentalis House, Bul. N. Y. State Mus. 243-244: 55, 1923, based upon Serpicula occidentalis Pursh, a superfluous name which cannot be transferred = Elodea Nuttallii. P. orinocensis (Rich.) Victorin, Contrib. Lab. Bot., Univ. Montréal 18: 42, 1931 — Elcdea granatensis. P. Planchonii (Casp.) Rydb., Bull. Torrey Bot. Club 35: 462-463, 1908 — Elodea canadensis. 1965] Elodea — St. John 171 Serpicula S. canadensis (Rich. in Michx.) Eaton, Man. ed. 5, 390, 1829. In large part and as to basionym = Elodea canadensis. S. occidentalis Pursh, Fl. Am. Sept. 1: 33, 1814. A superfluous name substituted for the available Elodea canadensis Rich. in Michx. In part, Elodea Nuttallii, and illegitimately adopted for that by St. John (1920). S. verticillata L. f. var. B angustifolia Muhl., Cat. Pl. Am. Sept. 84, 1815, nomen nudum = Elodea Nuttallii. Udora U. brasiliensis Mart., Fl. Brasil. 3(1): 99-100, 1847, a superfluous name, as he included as a synonym the valid and available Elodea granatensis Humb. & Bonpl. U. canadensis (Rich. in Michx.) Nutt., Gen. N. Am. PI. 2: 242, 1818, a superfluous name, since his monotypic new genus and species included the earlier and available Elodea canadensis Rich. in Michx. U. granatensis (Humb. & Bonpl.) Spreng., in Linnaeus’ Syst. Veg., ed. 16, 4(2) : 25, 1827 — Elodea granatensis. U. guyannensis (Rich.) Steud., Nom. Bot., ed. 2, 2: 727, 1841, (as guianensis) — Elodea granatensis. U. lithuanica Andrz. ex Bess., Flora, Beibl. 1832(2): 13, 1832 (er- roneously as Hydora lithuanica) = Hydrilla verticillata (L. f.) Royle. U. occidentalis Koch, Syn. Fl. Germ. & Helvet. 669, 1837, based upon Serpicula occidentalis Pursh, and therefore, as to type — Elodea canadensis. U. orinocensis (Rich.) Spreng., Syst. Veg. of Linnaeus, ed. 16 by Spreng., 4(2): 25, 1827 = Elodea granatensis. U. pomeranica Reichenb., Ic. Fl. Germ. Helvet. 7: 31, 1845 = Hydrilla verticillata (L. f.) Royle U. surinamensis Miq. ex Schomburgh, Reisen Brit.-Guiana 3: 900, 1848, a dubious species, with inadequate diagnosis. He said only that it was a perennial herb that bloomed all year. U. verticillata Spreng. in Linnaeus’ Syst. Veg., ed. 16 by Spreng., 1: 170-171, 1825. As to basionym = Hydrilla verticillata (L. f.) Royle, but, in part, is Elodea canadensis. minor Engelm. ex Caspary Monatsber. Kgl. Preuss. Akad. Wissensch. 1857: 46, 1857 (in repr. p. 10); Pringsheim’s Jahrb. 1: 465, 500, 1858, published in synonymy = Elodea Nuttallii. INDEX TO EXSICCATAE OF ELODEA The collector’s name, with initials, is followed by the collection number, or, if there is none, by s.n. (for sine numero), and then a number, or numbers in parentheses. The latter is the number or numbers of the species as described in this monograph. Species which 172 Rhodora [Vol. 67 occur in two of the geographic regions covered by different parts of this monograph will have two parenthetical reference numbers. The species names and numbers are given previously in a list in this paper. Abrams, L. R. 9719 (3), (19). Adams, J. W. 76, 99, 263, 264, 294 (7), (18). Alexander, C. P Anderson, W. A. s.n. (3), (19) Asplund, E. 72, 2275, 2595 (9); 2815 (11); 3449 (9). Austin, R. M. 573, 1179, s.n. (3), (19). Babcock, H. H. s.n. (7), (18). Bailev, W. s.n. (7), (18). Baker, M. S. s.n. (3), (19). Baldwin, W. K. W. & Breitung 3515, 3898, 4062 (3), (19). Ballard, C. A. s.n. (3), (19). Bang, M. 165 (12). Bardell, E. M. s.n. (3), (19). Bartram, E. B. s.n. (3), (19). Barker, H. C. s.n. (7), (18). Bassett, G. W. s.n. (3), (19). Battey, T. J. s.n. (7), (18). Beal, E. O. 3737, 3816, 3956, 4114, 5690, 5835, 6006 (7), (18). Bean, R. C, s.n. (3), (19). Bebb, M. S. s.n. (3), (19). Bebb, R. 1531 (3), (19). Bechtel, A. R. 13364 (3), (19). Bernard, J.-P. 58/284 (3), (19). Bertero, C. G. 225 (11). Beyrich, H. K. s.n. (3), (19). Bicknell, E. P. 174 (3), (19) ; 176 (7), (18) ; 177 (3), (19) ; sm (7), (18). Bigelow, s.n. (3), (19). Biltmore Herb. 1-897 (3), (19). Blake, J. 2210 (3), (19); sm. (7), (18). Blake, S. F. 2210 (3), (19). Blewett, A. E. 573, 586, 609, 1680, 1980 (7), (18) ; 1981 (3), (19). Blewett, A. E. & E. B. Harger 6631 (7), (18). Boetcher, F. L. J. 2543 (7), (18). Boice, C. A. s.n. (7), (18). Bonpland, A. 1757, s.n. (8), (13). Boott, W. s.n. (7), (18). Boott, Herb. s.n. (3), (19). Brade, A. C. 5795, 6409 (8), (13). Brainerd, E. s.n. (3), (19). Brandegee, K. s.m. (2). Brendel, F. s.m. (7), (18). Britton, N. L. s.n. (3), (19). Brown, S. s.n. (7), (18). Buckley Herb. s.n. (3), (19). Buell, M. F. 1650, 1651 (3), (19). Bünzli, G. H. s.n. (8), (13). Burgess, 2208 (3), (19). Burnham, S. H. 53 (3), (19). Burnett, H. S. sm. (3), (19). Bush, B. F. sm. (7), (18); sm. (3), (19). Butler, G. D. 27 (3), (19). Calder, J. A. & D. B. O. Saville 11838 (3), (19). Campbell, J. E. s.n. (3), (19). Canby, W. M. s.n. (7), (18). Cannon, E. s.n. (7), (18). Carson, J. s.n. (7), (18). Carter, J. J. s.n. (T), (18). Castle, H. & W. R. Dann 114 (7), (18). Chamberlain, E. B. 774 (7), (18). Chamberlain, E. B. & C. H. Knowlton s.m. (3), (19). Chandler, B. F. s.n. (3), (19). Chapman, s.m. (7), (18). Chapman Herb. s.n. (3), (19). Chase, A. 1711 (3), (19). Churchill, J. R. s.n. (3), (19). Clark, H. T. s.n. (3), (19). Clark, H. W. 1 (3), (19). Clausen, R. T. 2386 (3), (19). Cleghorn, s.n. (3), (19). Clinton, G. W. s.n. (3), (19). Clothier, G. L. 1052 (7), (18). Collins, F. S. 2445, 3639, s.n. (7), (18). Collins, J. F. s.n. (7), (18). 1965] Colorado State College 2421 (5), (21). Commerson, P. s.n. (14). Commons, A. sm. (7), (18), sm. (3), (19). Congdon, J. W. s.n. (3), (19). Cook, Fannye A. s.n. (7), (18). Cook, O. F. & G. E. Gilbert 195 (9). Coombs, A. L. s.n. (3), (19). Coville, F. V. 120, s.n. (3), (19). Coville, F. V. & J. B. Leiberg 318 (3), (19). Cowen, J. H s.n. (5), (21); s.n. (7), (18). Cowles, s.n. (7), (18). Crandall, C. S. 2423, 2528, 6720 (5), (21). Crandall C. S. & J. H. Cowen 6720 (5), (21). Cratty, R. I. s.n. (3), (19). Crawford, J. s.n. (7), (18). Cuming, H. 636 (11). Curtiss, A. H. s.n. (3), (19). Cushman, H. & M. Wood s.n. (7), (18). D. R. sn. (3), (19). Daguerre, 440 (15). Darlington, W. s.n. (3), (19). Davidson, R. A. 4409 (7), (18). Davis, H. A. 205 (3), (19). Deam, C. C. 11765 (3), (19); 14458 (7), (18); 20245 (3), (19); 20721, 22982 (7), (18); 25520, 25599, 29224, 30087, 42177, s.n. (3), (19). Deane, W. s.n. (7), (18). Deane, W., E. & C. E. Faxon s.n. (7), (18). De Selm, A. W. 717 (3), (19). Dewart, F. s.n. (7), (18). Dodge, C. K. s.n. (3), (19). D'Orbigny, A. s.n. (11). Drake & Dickson s.n. (4). Dreisbach, R. R. 1598 (7), (18). Drummond, T. s.n. (3), (19). Dryander, E. 2618 (8), (13). Dudley, W. R. s.n. (3), (19). Agric. Elodea — St. John 173 Dupee, D. s.n. (3), (19). Durand, E. s.m. (7), (18); sm. (3), (19). Dutton, D. L. s.n. (3), (19). Dutton, D. L. & C. W. Dodge s.n. (3), (19). Eames, A. J. 13362 (3), (19). Eames, A. J. & K. M. Wiegand 13351 (1), (18) ; 13353, 13353a, 13356 (3), (19). Eames, E. H. 5265, (19) ; s.n. (7), (18). Earle, E. C. 4281 (7), (18). Eastwood, A. 1268 (3), (19). Eaton, D. C. s.n. (7), (18). Eby, A. E sm. (3), (19). Eggert, H. s.n. (7), (18). Eggleston, W. W. 2085 (7), (18); s.n. (3), (19). Eggleston, W. W. & A. J. Grout s.n. (3), (19). Ehlers, J. H. 5902, 6256, 6280 (3), (19). Elliot, S. 340 (11). Emig. W. H. 218 (7), (18). Engelmann, G. 2, s.n. (7), (18); sn. (3), (19). Enneander, D. s.n. (7), (18). Epling, C. C. & Houck 10053 (7), (18). Espinosa, B., M. R. s.n. (11). Evermann, B. W. 1070, s.n. (3), (19). Eyerdam, W. J. 1221, 6627, s.n. (3), (19). Eyles, D. E. & M. S. 318, 338 (7), (18). Fasset, N. C. 14, 94 (7), (18); 5049, 5328, 5691 (3), (19); pert. 797 (T) (18); 7878, 7379, 7380, 7381, s.n. (3), (19). Fassett, N. C. & L. R. Wilson 5050, 5051 (3), (19). Fendler, A. s.n. (7), (18). Ferguson, W. C. 3405 (3), (19); 4147, 4414, 4832, 7843 (7), (18); 7879 (3), (19); 8072, 8154, s.n. (7), (18). 8457 (3), 174 Fernald, M. L. 2750 (7), (18). Fernald, M. L., F. Hunnewell & B. Long s.n. (7), (18). Fernald, M. L. & B. Long 12748 (7), (18). Fernald, M. L., B. Long & Smart 5592 (7), (18). Fernald, M. L. & H. K. Svenson 758 (7), (18). Fernald, M. L. & K. M. Wiegand 14548 (3), (19). Fernald, M. L., K. M. Wiegand & A. J. Eames 14123, 14124 (7), (18); 14125, 14126, 14127, 14128 (3), (19). Ferreyra, R. 5206, 5243 (11). Fink, B. 552 (3), (19) ; s.n. (7), (18). Fitzpatrick, T. J. s.n. (7), (18). Focke, H. C. 602 (8), (13). Fogg, J. M., Jr. s.n. (7), (18). Forbes, F. F. s.m. (7), (18). Frank, s.n. (3), (19). Fretz, C. D. s.n. (7), (18); s.m. (3), (19). Furbish, K. s.n. (7), (18). Gall, E. 503 (4); 1079, s.n. (7), (18). Garton, 1676 (3), (19). Gates, F. C. 632, 913a (7), (18); 1758.4, 14132 (3), (19). Gates, F. C. & M. T. 10592 (3), (19). Gattinger, A. s.n. (20). Gay, C. 166, s.m. (11); s.n. (14). Gehrt, A. 3333 (8), (13). Germain, see Rolland-Germain Gillman, H. s.n. (7), (18); sm. (3), (19). Githens, T. S. s.n. (7), (18). Glasson, M. s.n. (3), (19). Glaziou, A. 13290, 14355 22230 (8), (13). Glaziou, A. & Letegrier 22229 (8), (13). Glück, H. 7, 2 (15); s.n. (18). Godfrey, R. K. 62102 (3), (19). bis, (7), Rhodora [ Vol. 67 Goessl, C. (19). Goldie, J. s.n. (7), (18). Gooding, L. N. 597 (3), (19). Gorman, M. W. s.n. (3), (19). Gray, A. 4606, sm. (7), (18); s.m. (3), (19). Greene, A. s.n. (7), (18). Greeman, J. M. s.n. (3), (19). Greville, s.n. (3), (19). Griffith, R. E. 260, s.n. (7), (18). Grove, J. H. 490 (7), (18). Gurney, s.n. (7), (18). Guthnik Herb. s.n. (3), (19). Hale, T. J. s.n. (7), (18). Hall, E. 503 (4); 1079, s.n. (7), (18). Hall, J. s.n. (7), (18). Hanson, H. C. & E. E. A892 (1). Harger, E. B. s.n. (7), (18). Harger, E. B. & C. A. Weatherby s.n. (3), (19). Harper, E. T. & S. A. sn. (19). Harper, R. M. 3287 (3), (19). Harrington, W. H. s.n. (3), (19). Harshberger, J. W. sm. (7), (18). Harvey, J. L. s.n. (7), (18). Hasse, H. E. 2531 (3), (19) ; s.n. (7), (18). Hauman, L. 14, 10989, 10990, s.n. (14). Hayden, F. V. 1553, 1554, s.m. (3), (19). Henderson, L. F. 934 (4). Henderson, N. C. s.n. (3), (19). Hennecart, J. s.n. (3), (19). Hermann, F. J. 7642 (3), (19). Herrera, F. L. 217, 427 (9); 537 (10); 662 (9); 2082 (11). Heuser, P. s.n. (7), (18). Hicken, C. M. 872 (15); (14) 31574 (15). Hitchcock, A. S. 16532 (8), (18). Hitchcock, C. L. & Muhlick 13712 (3), (19). 6907, 7471, 8785 (8), (3), $73 1965] Hiteheock, R. & A. R. Bechtel 11218 (3), (19). Hodgdon, A. R. 5889, 5893 (7), (18). Hodgdon, A. R. & E. Hooghkirk 5890 (7), (18). Hodgdon, A. R. & F. Steele 70048 (7), (18). Hoehne, F. C. 19260, 20549, s.n. (8), (13). Hoffman, Ra: (9); (19): Holcomb; L sn: O (18) Hollick Av sn et Holton, I. F. 138 (8), (13) ; sm. (3), (19). Holzinger, J. M. s.n. ams E CLO): Hope, T. s.m. (3), (19). (7), (18); Hotchkiss, N. 4933, 6348 (5), (21); 6349 (3), (19). House, H. D. 6561, 8223 (3), (19). Howell, J. s.n. (4). Hueske, E. E. s.n. (3), (19). Jahn, A. 167 (7), (18). James, J. E s.m. (3), (19). James, L. E. 1711 (3), (19). James T. P. sm. (O (L8). Jennman, G. S. 4791 (8), (13). Jennings, O. E. s.m. (3), (19). Johannsen, P. L. 652 (3), (19). Jones, M. E. 6018 (3), (19). Jórgensen, P. 2535 (16). Jussieu, J. s.n. (8), (13). Kearney, T. H., Jr. 1697 (18). Keller, I. A. s.n. (7), (18). Kellogg, J. H. s.n. (7), (18). Kennedy, G. G. s.n. (7), (18); s.n. (3), (19). Keyes, H. H. s.n. (7), (18). Kidder, N. T. s.n. (7), (18). Kiener, R. 15537, 17514 (19); 17601 (7), (18). Kilip, E. P. 1063, 12533 (8), (19); 32656 (7), (18); 37595 (8), (13). Kingman, C. C. s.n. (7), (18). (7), (3), Elodea — St. John 175 Kirk, M. 679 (3), (19). Kleeberger, G. R. s.n. (7), (18). Knight, E. G. s.n. (7), (18). Knowlton, C. s.n. (3), (19). Kral, R. 13971 (3), (19). Krause, Àj s.n. (T): CIS). Krautter, L. s.n. (3), (19). Kriebel, R. M. 3479 (3), (19). Krochmal, S. B. 1227 (7), (18). Krotkov, P. V. 8649 (3), (19). Bansin OC E, JD 1762 (8), (19). Lapham, I. A. sm. (7), (18); Seed) C19). Lawrence, G. H. M. 831 (19). Lechler, W. 3144 (9). Leconte, s.n. (7), (18). Lee, C. E. s.n. (3), (19). Leggett, W. H. s.m. (7), (18). Lehman, F. C. 2287 (8), (13). Leiberg, J. B. 9 (3), (19). Leprieur, 26, 138, s.n. (8), (13). Lesquereux, L. s.m. (7), (18). Lippincott, C. D. s.n. (7), (18). Locke, S. B. 4 (3), (19). (3), Loefgren, & Edwall 2562 (8), (13). Long, B. 4595, 6036, s.n. (7), (18); sn, (3), (19). Long, C. A. E. 906, s.n. (3), (19). Looser, G. 3372 (11). Louis-Marie, Pere s.n. (3), (19). Luetzelburg, P. von s.n. (8), (13). Lunell, J. s.n. (3), (19). Mabbott, D. C. 321 (3), (19). McAtee, W. L. 2281 (3), (19); 2311, 2339 (7), (18) ; 3424 (3), (19). MacElwee, A. s.n. (7), (18). Mackenzie, K. K. 1505, 4731, 5095, 6833 (7), (18) ; 7233, 7258 (3), (19); 7352 (7), (18); 7732, sm. (3), (19) ; s.n. (7), (18). MacMillan, C. & E. P. Sheldon 572 (8), (19); 3334 (3), (19). Macoun, J. 13724, (3), (19); 176 13725 (5), (21); 27001, 61003, 68806, 82331, 85553, sm. (8), (19). Macoun, J. M. 27002 (3), (19). Mahoney, K. L. s.n. (7), (18). Mandon, G. 1461 (11). Martindale, I. C. s.n. (7), (18). Martius, K. F. P. von 453, sm. (8), (13). Martyn, E. B. 425 (8), (13). Massey, A. W. R. s.n. (3), (19). Matthews, A. 581 (9). Maxon, W. R. 6329 (7), (18). Mayall, & Cormack, s.n. (3), (19). Mearns, E. A. 804 (3), (19). Melchior, H. sm. (7), (18). Mendonga, R. 688 (8), (18). Meredith, H. B. s.n. (7), (18); s.n. (3), (19). Merrill, E. D. 508, 963, s.n. (7), (18). Metcalf, F. P. 358, 569 (5), (21); 924, 929, 1089, 1232 (7), (18); 2222, 2364 (8), (19). Meyer, F. G. s.n. (3), (19). Michaux, A. sm. (3), (19). Mitchill, M. s.n. (7), (18). Moldenke, H. N. 6765 (7), (18). Moore, J. W. 23682 (7), (18). Morong, T. s.m. (7), (18); sm. (3), (19). Morris, E. L. s.n. (7), (18). Moseley, E. L. s.n. (3), (19). Mosén, H. 3479 (8), (13). Moser, C. J. s.n. (17); an (3), (19). Mühlenberg, H. s.n. (3), (19). Muenscher, W. C. 3614, 13352 (7), (18) ; 13354, 13363, 14547, 19584 (3), (19). Muenscher, W. C. & A. R. Bechtel 50, 51 (3), (19). Muenscher, W. C. & P. R. Burk- holder 16366, 16367 16368, 16369, 16370 (8), (19). Muenscher, W. C. & O. F. Curtis 5570 (7), (18). Rhodora [Vol. 67 Muenscher, W. C., K. M. Wie- gand & A. H. Wright 15197, 15198 (8), (19). Mumbauer, J. R. 213 (7), (18). Murrill, W. A. s.n. (3), (19). Nash, G. V. 1 (3), (19). Nelson, A. 1429 (3), (19). Nicollet, I. N. 415 (3), (19). Northrup, J. I. & G. R. s.an. (3), (19). Nuderlein, G. s.n. (8), (13). Nuttall, T. sn. (4); sm. (18). O., R. C. & E. B. W. sm. (3), (19). Oberlin College s.n. (7), (18). Olney, S. T. s.n. (3), (19). Ostenfeld, C. H. 552, 554a, 584c, s.n. (3), (19). Palmer, E. J. 3765, 3766 (T), (18). Palmer, E. L. 73 (3), (19). Palmer, E. L. & A. J. Eames 72 (3), (19). Parker, s.n. (8), (13). Parodi, L. R. 11889, 11890 (14). Passarge, & Selwyn s.n. (8), (13). Pastore, F. 135 (15). Patterson, H. N. s.n. (7), (18). Pease, A. S. 2063, 2638, 29912, 36781 (7), (18). Peck, M. E. 4250 (3), (19). Pennell, F. W. 2280, 5014, 5058, 5110, 6498 (7), (18). Perrottet, G. S. 193 (8), (13). Peter, R. s.n. (3), (19) ; s.n. (7), (18). Phelps, O. P. 279 (3), (19). Philippi, R. A. 87, 682, s.n. (11). Pickering, C. s.n. (3), (19). Pieters, A. J. s.n. (3), (19). Piper, C. V. s.n. (3), (19). Porter, T. C. 91 (3), (19); sm. (7), (18); sm. (3), (19). Preston, D. 860 (3), (19). Pretz, H. W. 4183, 4784 (3), (19); 5175, 6227 (7), (18); (7), 1965] 6962 (3), (19); 7279 (18) ; 11212 (3), (19). Pur, ES os*. (D, (18). R., J. L. s.n. (3), (19). Radford, A. E. 4614, 4686 (7), (18); 4950 (3), (19); 4956 (7), (18); 5279 (3), (19); 5409, 5436, 5739 (7), (18). Radford, A. E. & Haesloop & Miller 7683 (7), (18). Raimondi, A. 9524, 10506 (9). Hedfeunrm PL Jr. 5749 (2 (19). Redfearn, P. L. & T. Stombaugh 4210 (7), (18). Reed, A. N. & Webb 929 (3), (19). Reed, E. C. s.n. (11). Remy, J. s.n. (3), (19). Rendle & Good s.». (3), (19). Rhoads, J. R. 831 (7), (18). Richard, L. C. M. s.n. (8), (13). Riedel, L. $48 (8), (13). Ripley, W. S., Jr. 16138, 17494 (3), (19). Robert, A. s.n. (8), (13). Robbins, s.n. (3), (19). Rolland-Germain, Freére 6472 693, 019) 7498 (7), (18). Rose, J. N., A. Pachano & G. Rose 22823 (9). Rossbach, G. B. 96, 97 (3), (19). Rothrock UT Toswu (05 UNT! Sw Or GLO) Rusby, H H sn. (7), (18); sm. (3), (19). Russell, C. s.n. (3), (19). Rust, H. J. 373 (7), (18). St.-Hilaire, A. de B1, 581, s.m. (7), (8), (13). St. John, H. 679, 2548a (7), (13) 8298" (3) (19) sm. (15). St. John, H. & F. A. Warren 3423 (3), (19). Sandberg, J. H. 563, s.n. (19). Sartwell, H. P. s.n. (3), (19). (3), Elodea — St. John 177 Schallert, P. O. s.m. (7), (18). Schott, A. s.n. (7), (18). Schrenk, H. von s.n. (3), (19). Schinz Herb. s.n. (3), (19). Schuette, J. H. s.n. (7), (18). Schulz, A. G. 833 (15). Scott, W. s.n. (3), (19). Schwake, W. 9298 (16). Schweinitz, J. B. de sm. s.n. (3), (19). Scoggan, H. J. 6680, 6900, 8723, 10268 (3), (19). Scovell, J. T. & Clarke 1070 (3), (19). Selby & Duvel 1321 (7), (18). Sellow, F. s.n. (8), (13). Senn, H. A. 552 (3), (19). Setchell, W. A. sm. (7), sm. (83), (19). Seymour, F. C. 257 (3), (19); 3620, 4824, 21058 (7), (18). Shear, C. L. 3746 (3), (19). Sheehan, & Krochmal 53 (8), (19). Short, C. W. sen. (7), (18). Short, Herb. sm. (3), (19). Shull, G. H. 146 (7), (18); 467 (3), (19). Small, J. K. s». (1), (18); 8n. (3), (19). Smith, B. H. s.n. (7), (18). Smith, C. E. s.n. (7), (18). Smith, H. H. 7422 (3), (19). smith: d. D. sn. (8) (19). Smith, L. B. & H. K. Svenson 825 (7), (18). Sodiro, A. 195/2 (11). Speegazzini, C. 2218 (15). Splitgerber, F. L. s.n. (8), (13). (17); (18) ; Spreadborough, W. 62633 (8), (19). Sprengel, Herb. sn (3), (19). Spruce, R. 1991 (8), (13); 5458 (9). Standley, P. C. 8274 (5), (21). Steele, E. S. s.n. (7), (18). Steele, F. L. 3374 (7), (18). Stevens, F. L. 305 (3), (19). 178 Strohm, F. H. s.». (3), (19). Stübel, A. 194c (8), (13). Sturtevant, E. L. s.n. (3), (19). Suksdorf, W. N. 87 (3), (19). Sullivant, W. T. S. s.n. (7), (18). Swezey, G. D. s.n. (7), (18). Tatnall, E. s.n. (3), (19). Taylor, B. C. s.n. (7), (18) ; s.m. (3), (19). Taylor, K. A. s.n. (7), (18). Taylor, N. 847, 1372 (7), (18). Taylor, T. M. C., Hosie et al. 1454 (3), (19). Thomson, R. 9, 177 (7), (18). Thorne, R. F. 10472, 11395 (7), (18); 14595, 17862, s.m. (3), (19). Tidestrom, I. 94 (7), (18); 7185 (3), (19); 10664 (6). Topping, D. L. s.n. (3), (19). Torrey, J. sm. (3), (19); sm. (17); s.m. (7), (18). Torrey Herb. s.n. (3), (19). Torrey & Gilman s.n. (7), (18). Tracy, S. M. & Evans 475 (6). Trelease, W. s.n. (3), (19). Troll, C. 2321, 3043 (10). Trudell, H. W. s.n. (8), (19). Tutin, T. G. s.n. (8), (13). U., E. B. s.n. (3), (19). Ule, E. 4074, 7758 (8), (18). Underwood, L. M. 3211, s.n. (3), (19). Usteri, A. s.n. (8), (13). Vail, A. M. s.n. (7), (18). Van Eseltine, G. P. & T. W. Moseley 203 (7), (18). Van Pelt, S. S. s.n. (3), (19). Van Pelt, S. S. & B. Long s.n. (7), (18). Vasey, G. R. s.n. (3), (19) ; s.n. (5), (21). Venturi, S. 175 (15). Victorin, Fr. Marie 8171, 9910, 11256, 20318 (3), (19); 20322 (7), (18); 24725, 28388, 28393 (3), (19). Rhodora [Vol. 67 Victorin, Fr. Marie & Fr. R- Germain 7310, 7410, 33905, 33996, 44974, 456381, 49140, 49147, 49306, 49361 (3), (19). Victorin, Fr. Marie, Fr. R.-Ger- main & B. Boivin 4302 (3), (19). Victorin, Fr. Marie, Fr. R.-Ger- main & Jacques 45855 (3), (19). Victorin, Fr. Marie, Fr. R.-Ger- main & Meilleur 44974 (3), (19). Victorin, Fr. Marie, Fr. R.-Ger- main & Rouleau 2017, 6534 (3), (19). Waite, M. B. s.n. (3), (19). Waldron, L. R. s.». (3), (19). Ward, L. F. 126, s.n. (7), (18); s.n. (3), (19). Ware, R. A. 3472 (7), (18). Warming, E. 1103, s.n. (8), (13). Watson, s.n. (7), (18). Weatherby, C. A. 367 (7), (18); 2700 (3), (19) ; 3373, 3591 (7), (18); 3595, 3604 (3), (19); 3277 (7), (18). Webb, R. J. s.n. (7), (18). Weddell, H. A. 3367 (8), (13). Weinland, 49 (3), (19) ; s.n. (7), (18). Westerfeld, W. F. 6357 (7), (18). Wheeler, C. F. 7663 (3), (19). Wheeler, L. 2325 (3), (19). Whitford, L. A. 231, s.an. (18). Widmann, O. s.n. (7), (18). Wied-Neuwied s.n. (17). Wiegand, K. M. 13355 (3), (19); 13358 (7), (18); 13358 in part (3), (19) ; 12359, 13360, 13361 (3), (19). Wiegand, K. M. & M. L. Fernald 14546 (3), (19). 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Club 35: 457-465. SANTOS, J. K. 1923. Differentiation among Chromosomes in Elodea. Bot. Gaz. 75: 42-59, 1 pl. 1924. Determination of Sex in Elodea. Bot. Gaz. TT: 353-376, figs. 1-8, pl. XXIII-XXVII. VICTORIN, FRÈRE MARIE-. 1931. L'Anacharis canadensis. Histoire et solution d'un imbroglio taxonomique. Contrib. Lab. Bot., Univ. Montréal, 18: 1-43, figs. 1-7. 180 Rhodora [Vol. 67 WIEGAND, K. M. & A. J. EAMES. 1926. The Flora of the Cayuga Lake Basin, New York, Vascular Plants. Mem. Agric. Exp. Sta., Cornell Univ. 92: 1-491. WYLIE, RoBERT B. 1904. The Morphology of Elodea canadensis. Bot. Gaz. 37: 1-22, pl. I-IV. This describes the floral development and the female gametophyte of genuine E. canadensis, but the stami- nate flowers described were certainly those of E. Nuttallit. Addenda Vol. 67, p. 7. At the end of the first paragraph add: capsules 4-6 mm. long, 1.3-1.8 mm. in diameter, lance-ellipsoid, 1-loculed, some- what involute on the 2 flattened sides; seeds 3.5-5 mm. long, 0.5 mm. in diameter, cylindric, brown, white puberulous. Corrigenda Vol. 67, p. 1, line 3 from below, after 1-8, change the comma to a semicolon, and delete: is in press in Mutisia; Page. 32, in the legend, for longivaginat, read: longivaginata FLOWERING PLANTS NEW TO OR RARE IN KENTUCKY! ELIZABETH M. BROWNE AND E. T. BROWNE, JR.” In an effort to catalog more completely the species of vascular plants in Kentucky and to plot their distribution, extensive field work has been done in the last three years. As a result, several species have been found in the state which were previously unknown to occur here or were first reported many years ago without having been reported since. Among these are the following: SPOROBOLUS CRYPTANDRUS (Torr.) Gray. This species was reported for the only time previously by Price (1893). Fulton Co. Kentucky Point. EMB & ETB 6051; EMB & ETB 6469; EMB & ETB 7624.5. ‘Contribution No. 3 of the Kentucky Flora Project, University of Kentucky. "The junior author wishes to express his sincere appreciation to the Faculty Research Committee, Graduate School, University of Ken- tucky, for several grants which partially defrayed field expenses and enabled him to make the trips to the herbaria, Dr. H. P. Riley has kindly reviewed the original manuscript. 1965] Kentucky Plants — Browne & Browne 181 In addition to the above-named species, others have been collected of which there is no record in the literature of their occurrence in Kentucky: ERIOCHLOA GRACILIS (Fourn.) Hitche. Fulton Co. Kentucky Point. EMB & ETB 6020, 6098; EMB & ETB 6409; EMB & ETB 7632. CHENOPODIUM PUMILIO R. Br. Trimble Co. Strawberry field near Bedford. Millard Maxey, Co. Agent, s.n. (Det. by James W. Herron and ETB.) AMARANTHUS ARENICOLA I. M. Johnst. Fulton Co. Kentucky Point. EMB & ETB 6060. TRIBULUS TERRESTRIS L. Fulton Co. Kentucky Point. EMB & ETB 6018. AMPELOPSIS ARBOREA (L.) Koehne. Fulton Co. Kentucky Point. EMB & ETB 5607; EMB & ETB 6070. JUSSIAEA LEPTOCARPA Nutt. Fulton Co. Kentucky Point. EMB & ETB 7618; Henry Co. Ox-bow W of Kentucky R. and 0.5 mi. E of dirt road to Lockport. EMB & ETB 5976. DICLIPTERA BRACHIATA (Pursh) Spreng. Fulton Co. Kentucky Point. EMB & ETB 5562; EMB & ETB 6100. HETEROTHECA LATIFOLIA Buckley. Fulton Co. Kentucky Point. EMB & ETB 5572; EMB & ETB 6067; EMB & ETB 6431; EMB & ETB 7963. No herbarium specimens of these species from Kentucky are to be found in A, GH, NY or US. Consequently, all except the first species mentioned herein are believed to be new state records. The kindness of the curators and staffs of these herbaria is gratefully acknowledged. 1234 KASTLE RD., LEXINGTON, KENTUCKY DEPARTMENT OF BOTANY, UNIVERSITY OF KENTUCKY, LEXING- TON LITERATURE CITED PRICE, SADIE F. 1893. A Flora of Warren County, Kentucky. New London, Wisconsin. THE TYPIFICATION OF XANTHOCEPHALUM (COMPOSITAE) OTTO. T. SOLBRIG The genus Xanthocephalum was described by Willdenow in 1807 (Ges. Naturf. Fr. Berlin Mag. 1:140. 1807) based on material collected and described by Humboldt. In the original description, which was very short and not diag- nostic of any species, no specific name was given. Kunth (Humboldt, Bonpland and Kunth, Nov. Gen. et Sp. Pl. 4:310. 1820) referred the species X. centauroides (without a description) to Willdenow's description, stating that he had not seen any material of it. This is the first specific epithet ascribed to Xanthocephalum, and was considered by me (Rhodora 63:151-164. 1961) as the type species, Never- theless, in the same work Kunth described the new genus Xanthocoma with one species X. humile HBK. (= Xantho- cephalum humile (HBK.) Sch. Bip. ex Hemsley), and re- described X. centauroides as Pyrethrum bonplandianum Kunth. Since I was unable to see type material there was some doubt as to the identity of X. centauroides. Further- more, there was the possibility, in case no original material was extant, that Xanthocephalum would have to be abandoned as a *nomen confusum" and have to be replaced by Xanthocoma. During the last summer I had an opportunity to visit and study at the Botanisches Museum in Berlin-Dahlem! where the Willdenow herbarium is deposited. In it a specimen (see Fig. 1) of Xanthocephalum (Willdenow no. 16523), col- lected by Humboldt is preserved. In the upper left hand corner is the annotation of Willdenow (see Schlechtendahl, Flora 15:561-567. 1832) "Syngenesia frustranea, Xantho- cephalum centauroides, Habitat in Mexico." In addition, in the lower right hand corner is stated “Humboldt W,” pre- '] want to thank the Botanical Society of America and the National Science Foundation for a travel grant. I also want to thank very particularly Dr. W. Domke, for all the help while in Berlin and for the photograph of the type of Xanthocephalum. 182 1965] Xanthocephalum — Solbrig 183 Mus. Bot. Berol. Film Nr. 7232 ON | LL m suada 6 li 2 3 fr RU | L MM DI | Plate, VN Bags). Det no. 16523, B) ail of the type specimen of X. centauroides (Willdenow 184 Rhodora [Vol. 6% sumably copied from the back of the original sheet, where Willdenow used to write the procedence of his material (see again Schlechtendahl, 1832). There is also a copy of the original diagnosis affixed after Willdenow's death by Schlechtendahl, and a later annotation by Schulz-Bipontinus, where Xanthocephalum (Pyrethrum) ` bonplandianum (HBK.) DC is equated to X. centauroides. A last annotation "Pyrethrum Bonplandianum Kunth" could not be traced to its author. It is clear, that this specimen is the type of Xanthoce- phalum centauroides Willd. ex HBK. (Nov. Gen. et Sp. 4:312. 1820), and that this is the type species of the genus as I had concluded in my previous work. Furthermore, it confirms Xanthocoma as a generic synonym provided X. humile is considered a member of Xanthocephalum. GRAY HERBARIUM, HARVARD UNIVERSITY TWO SPECIES OF CHAMAESYCE (EUPHORBIACEAE) NEW TO THE UNITED STATES Collections made in peninsular Florida for a taxonomic revision of the genus Chamaesyce in the Caribbean region have included two species not previously reported for the United States. The two, Chamaesyce mendezii (Boiss.) Millsp. and C. thymifolia (L.) Millsp., are known from Mexico and Cuba, with the range of the latter extending through the West Indies to South America. Chamaesyce mendezii resembles the commoner C. pros- trata (Ait.) Small, and C. thymifolia is very similar in appearance to C. maculata, (L.) Small (sensu most authors, not Wheeler). All four are prostrate plants with serrate leaves, pubescent stems and capsules, and cyathia borne in the axils of leaves or on leafy side shoots, but they may readily be distinguished on characters of inflorescence and seed. Capsule pubescent only along the angles. Stem short-pubescent on lines; cyathial appendages minute or obsolete; seed with deep transverse furrows often extending through the angles o sns E ge teers C. prostrata Stem long-hirsute, at least on lines; cyathial appendages prominent, pink or white; surface of seed rippled, the ridges not extending through the angles ee C. mendezit Capsule pubescent all over, sometimes sparingly so. Capsule completely exserted from cyathium when mature, widest just below the equator; styles .2-.4 mm, long, spreading; angles of seed rounded, faces with low irregular ridges; diameter of seed sub-equal except at extremities ............ eese C. maculata Capsule only partially exserted, splitting one side of cyathium when mature, widest close to base; styles .4-.6 mm. long, only rarely spreading; seed with sharp angles, ridges on faces transverse, well marked; seed tapering from base to apex .................... C. thymifolia Chamaesyce mendezii was first collected in Florida in 1957 (Brass 29046, St. Lucie County, as C. adenoptera (Bertol.) Small), and then in 1959 in the Redlands area of Dade County (Atwater M-113, as Euphorbia adenoptera). It is now widespread in disturbed areas in Dade, Monroe 185 186 Rhodora [Vol. 67 and Collier counties, and has been collected in Highlands and Polk counties. It must be considered a well established element of the flora of the southern counties, and at least an occasional weed in the central part of the state. C. mendezii: COLLIER COUNTY: Goodland, 26 Sept 1964, D. Burch 401 (GH, US); Naples, 27 Sept 1964, DB 412 (BM, F) ; Goodland, 22 Nov 1964, R. K. Godfrey, 65468a; Naples, 22 Nov 1964, RKG 65527; DADE: Redlands, 5 July 1959, Atwater M-113; S. Miami, 11 Aug 1963, DB 158; Coral Reef Dr., Miami, 11 Aug 1963, DB 160; Old Cutler Rd., Miami, 29 Nov 1963, DB 219 (Ny, US); Everglades National Park, 17 Apr 1964, D. Burch & D. B. Ward 293; Homestead, 19 Apr 1964, DB & DBW 313; Homestead, 19 Apr 1964, RKG 63361; HIGHLANDS: Lake Annie, junction FLA 17 and FLA 70, 18 Nov 1964, DB & DBW 528 (NY); MONROE: Flamingo, Everglades National Park, 17 Apr 1964 DB & DBW 288; Plantation Key, 21 Nov 1964, DB & DBW 563; Cudjoe Key, 30 Dec 1964, G. Avery s.n.; POLK: Babson Park, 27 Nov 1964, DB 580; ST. LUCIE: Nigger Jim Scrub, 26 Oct 1957, L. Brass 29046. Chamaesyce thymifolia has only been found in three counties to date, all on the west coast of Florida and in each case close to salt or brackish water. The collections were all made in 1963 or 1964, but the colonies were thriving and appeared from their extent to have been in the area for some time. C. thymifolia: CHARLOTTE COUNTY: Charlotte Harbor, 27 Sept 1964, DB 417 (BM, GH, NY, US) ; LEE: Fort Myers, 13 Aug 1963, DB 156a; Fort Myers, 27 Sept 1964, DB 413 (BM, F, FSU, GH, NY, US) ; Fort Myers, 27 Sept 1964, DB 414; PINELLAS: Dunedin, 15 Oct 1964, DB 451, Specimens representing the above collections are on de- posit in the herbarium of the Agricultural Experiment Station, University of Florida, Gainesville, and duplicates are being distributed to the institutions indicated. DEREK BURCH DEPARTMENT OF BOTANY, UNIVERSITY OF FLORIDA, GAINESVILLE CHROMOSOME NUMBERS IN PENSTEMON (SCROPHULARIACEAE) I: NEW MITOTIC COUNTS FRANK S. CROSSWHITE AND SHOICHI KAWANO Living plants of Penstemon were collected for cytological analysis by Delzie Demaree, Dr. W. G. Dore, Dr. Dennis Knight, and the authors. These were transported to the University of Wisconsin greenhouses and botanical garden, where they have been grown for two successive years. Root- tips were used exclusively. Initially, root-tips were sectioned and stained with erystal-violet, but most of the preparations were made by a new orcein technique (Kawano, unpublished). Excised root-tips were stored in 0.001-0.002 mol. 8-hydroxyquinoline aqueous solution for 4-6 hours at 17-20°C, then fixed in Os- tergren & Heneen's fixative overnight, then treated with 5% aqueous pectinase for 1-2 hours at 30-38°C, and sub- sequently stained in lactopropionic orcein (Dyer, 1963) for more than 6 hours. The root-tips were then transferred into lactopropionic orcein-1 N hydrochloric acid (1:1) in which they were allowed to remain for 30 seconds to 5 minutes. Each of the root-tips was then transferred to a glass slide, treated with a drop of propionic acid (45%)- glycerine (9:1) for 5 to 20 seconds, after which it was gently heated over an alcohol flame and the root-tip squashed. Since part II of this series, a revue of the cytology of Penstemon, is in preliminary manuscript, and will be pub- lished soon, no discussion of the taxonomic ramifications of the new counts is presented here. The new counts, together with voucher specimens, are cited below. Metaphase draw- ings are presented in Figure 1. Penstemon arkansanus Pennell. 2n = 16. (including P. multicaulis Pennell and P. wherryi Pennell) ARKANSAS. FAULKNER Co.: 2 mi. e. of Conway, Crosswhite & Crosswhite (WIS); 4-5 mi. n. of Conway, Crosswhite & Crosswhite 62-325 (WIS). FRANKLIN Co.: 7 mi. w. of Ozark, Crosswhite & Crosswhite 62-310 (WIS). GARLAND Co.: Hwy. 7, 1 mi. s. of Hamilton Lake, Crosswhite & Crosswhite 62-320 (wis). HoT SPRINGS Co: 187 188 Rhodora [Vol. 67 Big Hill Creek, 2 mi. n. of Bismark, Crosswhite & Crosswhite 62-322 (WIS). INDEPENDENCE Co.: 5 mi. s. of Batesville, Crosswhite & Kawano 63-,07 (WIS). JACKSON Co: 2 mi. s. of Denmark, Cross- white & Kawano 63-408 (wis). LOGAN Co.: 8 mi. se. of Paris, Crosswhite & Crosswhite 62-314 (WIS). PERRY Co.: 38% mi. s. of Hollis, Crosswhite & Crosswhite 62-318 (WIS). SALINE Co.: 1 mi. e. of Owensville, Crosswhite & Crosswhite 62-324 (WIS). YELL Co.: 2 mi. n. of Havana, Crosswhite & Crosswhite 62-315 (WIS). oke y LAS * us ae AS a vi HARBOURII ARKANSANUS PALLIDUS oe Mi “, GK A LAXIFLORUS PACHYPHYLLUS TEUCRIOIDES Le Le, ay M SM ts A d Le € 7722 ZLA 2 a H <$ SN Té SS s. CALYCOSUS GRACILIS. ssp. Ver (Cit WISCONSINENSIS Zo | TUBEFLORUS u FB ~é dëtt UA ae) Ze 9 "1 ; " 24 REDE eo $ >a" TRS Cros yA bw "e Á DIGITALIS Fig. 1. Camera-lucida drawings of the chromosome complements of several Penstemon taxa. 1965] Penstemon — Crosswhite & Kawano 189 Penstemon calycosus Small. 2» — 96. ILLINOIS. MARSHALL Co.: 2 mi. n. of Lacon, Crosswhite & Cross- white 62-305 (wis). SHELBY Co.: 9 mi. s. of Pana, Crosswhite & Crosswhite 62-330 (wis). WASHINGTON Co.: 2 mi. s. of Irvington, Crosswhite d Crosswhite 62-239 (wis). Penstemon digitalis Nutt. 2n — 96. ARKANSAS. PERRY Co.: 3 mi. s. of Hollis, Crosswhite & Cross- white 62-317 (wis). SHARP Co.: 1 mi. s. of Hardy, Crosswhite & Kawano 63-414 (wis). LOUISIANA. EVANGELINE PARISH: Chicot State Park, Crosswhite & Kawano 63-415 (WIS). MISSOURI. MILLER Co.: 5 mi. s.e. of Eldon, Crosswhite & Nickerson 62-306 (WIS). WISCONSIN. RicHLAND Co.: Richland Center, Crosswhite 63-412 (WiS). SAUK Co.: Lake Delton, Crosswhite & Crosswhite 63-413 (wis). ONTARIO: Jaspar, Dore 19184 (WIS). Penstemon gracilis Nutt. ssp. gracilis, 2n — 16. SOUTH DAKOTA. DEUEL Co.: 4 mi. e. of Clear Lake, Knight s.n. (WIS). PENNINGTON Co: Mt. Rushmore, Crosswhite & Cross- white 63-400 (Wis). WISCONSIN. TREMPEALEAU Co.: Trempealeau, Crosswhite 63-401 (WIS). Penstemon gracilis ssp. wisconsinensis Pennell. 2n = 16. WISCONSIN. Apams Co.: 2 mi. n. of Plainville, Crosswhite & Crosswhite 62-303 (wis) ; Friendship, Crosswhite & Crosswhite 62-304 (WIS). WAUSHARA Co.: Plainfield, Crosswhite & Crosswhite 63-402 (WIs). Penstemon grandiflorus Nutt. 2n = 16. WISCONSIN. ApAMS Co.: New Rome, Crosswhite & Crosswhite 63-416 (WIS). Penstemon harbourii A. Gray. 2n = 16. COLORADO. GUNNISON Co.: Mt. Belleview, Crosswhite 62-221X (PH, RM, SMU, WIS). Penstemon laxiflorus Pennell. 2n = 16. ALABAMA. MoBILE Co.: Creola, Crosswhite & Kawano 63-411 (wis). LOUISIANA. GRANT PARISH: 15 mi. s. of Winnfiield, Cross- white & Kawano 63-409 (WIS). EVANGELINE PARISH: 5 mi. n. of Turkey Creek, Crosswhite & Kawano 63-410 (WIS). Penstemon pachyphyllus A. Gray ssp. pachyphyllus. 2n = 16. UTAH. UINTAH Co.: 13 mi. n. of Vernal, Crosswhite & Crosswhite 62-254X (PH, RM, SMU, WIS). Penstemon pallidus Small. 2n = 16. ARKANSAS. INDEPENDENCE Co.: % mi. s. of Cave City, Cross- white & Kawano 63-405 (Wis). SHARP Co.: 5 mi. s. of Ash Flat, Crosswhite & Kawano 63-406 (wis). ILLINOIS. JACKSON Co.: Giant 190 Rhodora [Vol. 67 City State Park, Crosswhite & Kawano 63-404 (WIS). WASHINGTUN Co.: s. of Centralia, Crosswhite & Kawano 63-403 (WIS). Penstemon teucrioides Greene. 2n — 16. COLORADO. GUNNISON Co.: Almont, Crosswhite & Crosswhite 62-229X (PH, RM, SMU, WIS); Sapinero, Crosswhite & Crosswhite 62-230X (PH, RM, SMU, WIS). Penstemon tubaeflorus Nutt. 2n — 32. ARKANSAS. CRAIGHEAD Co.: 1 mi. n. of Jonesboro, Crosswhite & Crosswhite 62-328 (WIS); 5 mi. s. of Jonesboro, Crosswhite & Cross- white (WIS). CRAWFORD Co.: 4 mi. n. of Mountainberg, Crosswhite & Crosswhite 62-313 (WIS). JOHNSON Co.: Ozone, Crosswhite & Cross- white 62-309 (w1S). We wish to thank the Research Committee of the Univer- sity of Wisconsin for supplying funds for this study from the Wisconsin Alumni Research Foundation. The work by Dr. Kawano was supported in part by a grant from the National Science Foundation to Dr. Hugh H. Iltis. We are very grateful to Dr. Iltis for his interest in the project, and for critically reviewing the original manuscript. HERBARIUM, UNIVERSITY OF WISCONSIN, MADISON DEPARTMENT OF BOTANY, UNIVERSITY OF TOKYO, JAPAN ADDITIONS TO THE VASCULAR FLORA OF OKLAHOMA The vascular flora of most of the major areas of Okla- homa has been fairly well collected. There are, however, many areas within the boundary of the state that have not been thoroughly investigated. The species discussed in this paper come from two of these areas. Until recently, Bryan Co. represented an essentially uninvestigated area. Prior to 1960, only a few specimens of vascular plants had been collected from this county of approximately 900 sq. miles. Specimens from the county have added a few new taxa to the flora of the state, and have added range extensions for many eastern and a few western species, (Taylor & Taylor, 1964a, 1964b, and in press). Many unique habitats are found in Bryan County. Hillside, seepy, bog-like habitats are formed along the flanks of small valleys cut in the Woodbine Sandstone (Cretaceous in age) where porous overlying strata have been truncated. Some of these habitats are only an acre or so in size, while others are several acres in extent (Taylor & Laughlin, 1963). Another unique habitat in this area is a strip of sand dunes lying along the north side of the Red River. The more recently formed dunes contain a sparse, xeric type of vegetation composed partially of western and southwestern species. The older dunes are stabilized and a type of bottomland forest invades them. Some of these forested areas have been cleared and cultivated, then subsequently allowed to go fallow. Some western and southwestern species also occur in these old fields (Taylor & Taylor, in press). Another area in Okla- homa that as yet has not been thoroughly studied is a 48 sq. mi. area (triangular in outline with the towns of Mill Creek, Mannsville, and Ravia at the apices) lying along the eastern side of the Arbuckle Mountains. It is an area under- lain mainly by limestone strata (Tishomingo granite out- crops along the eastern side) and dissected by a number of spring-fed streams that flow year round — even in very dry years. This area, and other parts of the Arbuckle Mountains lying adjacent to it on the west, contain several eastern species. Hall (1952) and Hall and Carr (1964) discussed 191 192 Rhodora [Vol. 67 the affinities of some of this eastern element. Taylor and Taylor (in press) list a few additional eastern species for this area. The species listed below are thought to be newly reported for the state. All specimens cited were collected by John and Constance Taylor and are deposited in the Bebb Herba- rium, University of Oklahoma, Norman. We wish to express our sincere gratitude to Dr. G. J. Goodman for his advice and for placing the facilities of the Bebb Herbarium at our disposal. Dulichium arundinaceum (L.) Britt. was collected in Bryan County (No. 2332) from 4.5 mi. NE of Bennington where it was very abundant in a hillside seep. The Three- Way Sedge is a species of marshes and stream margins ranging from California N to British Columbia, E to New- foundland and S to Florida and Texas. Rhynchospora capillacea Torr. This almost hair-like, tussock forming member of the sedge family was collected (No. 2461) in the Arbuckle Mt. area of Johnston Co., 4 mi. S of the town of Mill Creek from a damp, calcareous meadow near Bee Branch. It would seem that this species has not previously been known further south and west than south- ern Missouri, where Steyermark (1963) considers it a relict. Quite possibly its occurrence in the Arbuckle Mts. of Okla- homa may be explained in the same manner. Tigridia purpurea (Herb.) Shinners. This combination follows Shinners (1964). We have this very beautiful member of the Iridaceae (formerly known under Eustylis) from two nearby locations in eastern Bryan County. No. 2220 was collected from a damp low lying area in an oak- pine forest, 5 mi. E and 2 mi. N of Bennington. Other specimens (No. 2269 and duplicates) were collected ap- proximately 1 mi. S of this location from an upland forest that fringes a hillside seep. Apparently the rather rare Pinewoods Lily has previously been known only from Ar- kansas, Louisiana, and Texas. Linum imbricatum (Raf.) Shinners has been collected from a sandy old field 3.5 mi. SW of Mead in Bryan County. This species has previously been known from as near as 1965] Oklahoma Flora — Taylor & Taylor 193 Cook and Grayson Cos. across the Red River in Texas (Rogers, 1964), and is known only from east centra] Texas. R. JOHN AND CONSTANCE TAYLOR SOUTHEASTERN STATE COLLEGE, DURANT, OKLAHOMA LITERATURE CITED HALL, Marion T. 1952. Variation and hybridization in Juniperus. Annals of the Mo. Bot. Gard. 39:1-64. , and CLAUDIA J. CARR. 1964. Differential se- lection in Juniper populations from the Baum Limestone and Trinity Sand of southern Oklahoma. Butler Univ. Bot. Stud. 14:21-40. Rocers, C. M. 1964. Yellow-flowered Linum (Linaceae) in Texas. SIDA Contr. to Botany 1(6) :328-336. SHINNERS, LLovp H. 1964. Tigridia purpurea (Herb.) Shinners comb. nov. (Iridaceae). SIDA Cont. to Botany 1(5):295. STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames. p. 306. TAYLOR, R. JOHN and HAROLD LAUGHLIN. 1964. Additions to the Herpeteofauna of Bryan County, Oklahoma. Southwestern Naturalist 9:41-43. TAYLOR, CONSTANCE and R. JOHN. 1964a. Comments on the Flora of Bryan Co., Okla. Proc. Okla. Acad. Sci. 45:6-10. . 1964b. Podophyllum peltatum f. Deamii from Bryan Co., Okla. Rhodora 66:766. TAYLOR, R. JOHN and CONSTANCE. 1965 (in press). Comments on the vascular flora of Oklahoma. Proc. Okla. Acad. Sci. 46. CHAMAESARACHA VILLOSA NEW TO TEXAS — While identifying a rather large collection of specimens from the Chinati Mountains of west Texas, (as a research participant in botany under the NSF sponsored program at Oklahoma State University) I noticed a rather unusual specimen of Chamaesaracha. This specimen (Scudday 372) proved to be Chamaesaracha villosa Rydberg, as determined by Dr. U. T. Waterfall of Oklahoma State University. The specimen was found growing in gravelly alluvium and sand beside the Marfa-Ruidosa road about two and one-half miles north of Ruidosa. It is deposited in the herbarium of Okla- homa State University. Associated species collected from the same site were Tetraclea Coulteri Gray, Physalis hederaefolia Gray var. hederaefolia, and Jatropha dioica Sesse ex Cerv. var. graminea McVaugh. This species of Chamaesaracha was described from col- lections made in Coahuila, Mexico (Rydberg, 1895). It is not included in the checklist of Texas plants (Gould, 1962). The addition of this species to the flora of Texas is hereby noted. The author is grateful to Dr. U. T. Waterfall for deter- mining the specimen of Chamaesaracha. JAMES F. SCUDDAY DEPT. OF BIOLOGY, SUL ROSS STATE COLLEGE, ALPINE, TEXAS. LITERATURE CITED GouLp, F. W., 1963 — Texas Plants. A checklist and ecological summary. Tex. Agric. Exper. Sta. MP — 585. 112 p. RYDBERG, P. A., 1895 — The North American Species of Physalis and Related Genera. Mem. of the Torrey Bot. Club. 4 (5): 368. 194 NOTES ON THE FLORA OF CO6S COUNTY, NEW HAMPSHIRE The apearance of a new flora such as Professor Pease's Flora of Northern New England. stimulates the desire to find new taxa in the area and thus renders the work obsolete as soon as possible. With this in mind, I did a limited amount of collecting in Coós County during the summer of 1964. The results are presented below together with a few changes which should be made in the Pease Flora. Equisetum fluviatile L. The following townships may be added to the six in Pease. Wentworth location, along Magalloway River, Harris 26789 (64); Errol, Bear Brook Logan, Harris 26548 (64) ; Stratford, Stratford Bog Pond, Harris 26758 (64). Lycopodium clavatum L., var. megastachyon Fern. & Bissell, forma furcatum (Luerss.) Vict. Dummer, runout field Dummer Hill, Harris 26575 (64). First collection for the County. Larix laricina (DuRoi) K. Koch. Bean Grant, between Mts. Clinton and Pleasant, Harris & Hans Stauffer 26535 (64). A small group of trees over two meters tall were found growing at the upper limit of timber on the Crawford Path. Potamogeton natans L. Errol, Bear Brook Logan, Harris 26553 (64). The fourth township to be represented. Elodea Nuttallii (Planch.) St. John. Pittsburg, First Lake, Harris 26745 (64); Wentworth Location, Magalloway River, Harris 26797 (64); Berlin, Androscoggin River, Harris 26634 (64). Adds three townships to the two listed in Pease. Vallisneria americana Michx. Berlin, Androscoggin River, Harris 26647 (64). Adds a fourth township to the representation of this species. Eragrostis multicaulis Steud. Colebrook, roadside, Harris 26732 (64). The Pease collection 29690 (42) cited under Colebrook in E. pectinacea should be moved to E. multicaulis. Phleum pratense L., forma viviparum (S. F. Gray) Louis-Marie. Milan, pasture West Milan, Harris 26610 (64). Zizania aquatica L., var. augustifolia Hitche. Wentworth Location, oxbow pond, Harris 26786 (64). The fourth township for the County. Setaria Faberi Herrm. The Pease collection cited under S. viridis, Berlin 30308 (43) should be changed to this species, a new record for the County. Eleocharis obtusa (Willd.) Schultes, var. jejuna Fern. Berlin, shore of Androscoggin River, Harris 26624 (64). The second collection from the County. Lemna minor L. Stewartstown, pool in County Farm pasture, Harris 26733 (64); Milan, pool in pasture West Milan, Harris 26614 (64). 195 196 Rhodora [Vol. 67 Previous records from three townships, all in the southern portion of the County. Iris Pseudacorus L. Milan, bank of Upper Ammonoosuc River, West Milan, Harris 26652 (64). New to the County. Silene pumilio Wulf. Sargent Purchase, Ammonoosuc Ravine Trail near upper limit of trees, Harris 26529 (64). This species native to the mountains on the boundary of Austria and Yugoslavia, and not too common there, was a most unexpected find. The small colony confined to a crack in a ledge had evidently been there for several years. Its establishment on Mount Washington cannot be accidental. Some practical joker must be attempting to spice up the mountain flora. Ceratophyllum demersum L. Errol, Bear Brook Logan, Harris 26555 (64). This collection substantiates the statement made in Pease that Provost reports that the species is abundant at this locality, the second township for the County. Ranunculus acris L., var. latisectus G. Beck. Milan, pasture West Milan, Harris 26611 (64). The second collection from the County. Rubus pensilvanicus Poir. Dummer, old field Dummer Hill, Harris 26558 (64). Known previously from three townships. Lotus corniculatus L. Colebrook, berm U. S. rte. 3, Harris 26729 (64); Berlin, roadside rte. 16, Harris 26531 (64) ; Gorham, roadside rte. 16, Harris 26530 (64). This is a new genus for the County. The species seems to be rapidly establishing itself over much of New Eng- land. Linum catharticum L. Pittsburg, berm of highway two miles north of Idlewild, Harris 26742 (64). This is apparently the first collection of the species in New Hampshire. Daphne Mezereum L. Milan, spontaneous behind Steady Camp, Cedar Pond, Harris 26806 (64). This is a new genus to the County. Mr. Steady has occupied the camp for many years and has never noticed the shrub until the past summer. Myriophyllum verticillatum L., var. pectinatum Wallr. Berlin, An- droscoggin River, Harris 26645 (64). The third township for the County. Myriophyllum tenellum Bigel. Stark, Christine Lake, Harris 26600 (64). The fourth township for the County. Hippurus vulgaris L. Stratford, Stratford Bog Pond, Harris 26768 (64). The fifth township for the County. Carum Carvi L. Low & Burbank Grant, abundant at Madison Hut, Harris 26527 (64). This is the first collection of the species from above treeline in the County. Moneses uniflora (L.) Gray. Bean Grant, coniferous woods south- west of summit of Mt. Clinton, Harris & Hans Stauffer 26534 (64). The fourth township for the County and the only collection made above 4000 feet. Lycopersicum esculentum Mill. Milan, waif on dump, West Milan, Harris 26604 (64). A new genus for the County but of little moment. 1965] Flora of Coos County — Harris 197 Verbascum Blattaria L. Pittsburg, roadside two miles north of Idlewild, Harris 26743 (64); Colebrook, berm of U. S. rte. 3, Harris 26730 (64). Known previously from three southern townships. Gratiola aurea Muhl. Stark, Christine Lake, Harris 26599 (64). Third township for County. Utricularia inflata Walt., var. minor Chapm. Errol, Bear Brook Logan, Harris 26542 (64); Berlin, Androscoggin River, Harris 26618 (64). Known previously from two townships. Centaurea Jacea L. The collection Milan, Harris & Humes 7688 (51) cited under C. nigra, var. radiata should be moved to this species. The second collection for the County. Crepis capillaris (L.) Wallr. Colebrook, berm U. S. rte. 3, Harris 26731 (64). The second township for the County. Hieracium aurantiacum L. Low & Burbank Grant, Madison Hut, Harris 26528 (64). The species is abundant about the Hut at 4800 feet and well above treeline. STUART K. HARRIS BOSTON UNIVERSITY. AN EASTERN VARIETY OF CAREX FISSA (SMULTIFLORAE) F. J. HERMANN In 1960, specimens of a puzzling sedge from Lake and Seminole counties, Florida, were received for review from Richard J. Eaton. Mr. Eaton commented that although re- peated attempts to key out the plant in Mackenzie’s mono- graph (North American Flora, Vol. 18, 1931-1935), as well as in Fernald's treatment in Gray's Manual, ed. 8, (1950), and Gleason's in the New Britton and Brown Illustrated Flora (1952), lead invariably to the widespread and poly- morphous Carex annectens Bickn. (Section Multiflorae), it was not in the least suggestive of that species in the field. My own attempts at identification produced the same result and it was not until I had seen an additional specimen (Ray, Wood, Smith & Eaton 10750), collected the following year, that another possible affinity suggested itself, namely with the rare, or at any rate highly localized, C. fissa Mack. of eastern Oklahoma. Further collections from Florida by R. J. Eaton, an extensive series obtained from several counties in the same state by R. K. Godfrey, and recent collections of C. fissa kindly made in the type locality by Professor U. T. Water- fall, were compared with the type specimen of C. fissa. The results indicated that in their fundamental character- istics the Florida plants agreed most closely with the pre- sumed Oklahoma endemic. Pronounced differences were apparent, so that nomenclatorial recognition seemed imper- ative, but without exception the diagnostic characteristics were found to be inconstant, and so the eastern plant is here proposed as a geographic variety. Carex fissa Mack. var. aristata F. J. Herm., var. nov., a varietate typica recedit ligula concava, vaginis ad apicem ventraliter Saepe minus prolongatis, squamis foemineis longiaristatis, perigyniis ventraliter paucinervatis. FLORIDA: SEMINOLE COUNTY: between roadside and moist margin of pine- Serenoa flatwood, 1 mile southeast of Oviedo, Ray, Wood, Smith & Eaton 10750, April 26, 1961 (GH-Type; NY; USFs; FSU; OKLA); near Slavia, Cooley, Eaton & Ray 7459, May 2, 1960 (GH; Us; USFS; USF). LAKE COUNTY: near Howie-in-the-Hills, Cooley & Eaton 7350, 198 1965] Carex fissa — Hermann 199 April 29, 1960 (GH; USFS; USF). WAKULLA COUNTY: Live Oak Island, R. K. Godfrey 56799, May 21, 1958 (GH; USFS; FSU) ; and 62727, May 3, 1963 (GH; FS; NY; BM; SFU; US; V); vicinity of Shell Island near St. Marks, R. K. Godfrey 62734, May 3, 1963 (GH; FS; FSU; US). Nassau COUNTY: clearing of swamp, vicinity of O'Neil, R. K. Godfrey 64096, May 23, 1964, (FSU; USFS). MARIAN CouNTY: wet ditch along Alexander Springs River, near Alexander Springs, R. K. Godfrey & R. D. Houk 62795, May 11, 1963, (SFU; USFS). TAYLOR COUNTY: low wet area near river, Econfina Landing, S. McDaniel & R. K. Godfrey 4299, May 3, 1964 (FsU; USFS). OKLAHOMA: CREEK CouNTY: ditch, valley just north of Kiefer, M. T. Waterfall 17016, June 1, 1962 (GH; OKLA; USFS) (approaching var. fissa). The following key may serve to distinguish Carex fissa var. aristata from var. fissa and from C. annectens. Plant coarse in habit; culms thick (3-6 mm. wide at base) ; leaves wide (3-5 mm.) ; spikes congested into a short, broad head (2-4.5 cm. long, 8-15 mm. wide); perigynia large (3.25-3.9 mm. long), usually green at maturity, truncate to shallowly cordate at base. Ligule convex; sheaths conspicuously prolonged ventrally at the mouth; scales acuminate to cuspidate; perigynia ventrally nerveless or occasionally 1-2 nerved .....................-. C. fissa. Ligule generally concave to V-shaped; sheaths generally less pro- longed ventrally at the mouth; pistillate scales cuspidate to long-awned; perigynia usually with 1-4 ventral nerves E E C. fissa var. aristata. Plant slender; culms not thick (2.5-4 mm. wide at base); leaves rela- tively narrow (2-4 mm. wide) ; spikes arranged in an elongated, relatively narrow inflorescence (3-8 cm. long, 5-10(12) mm. wide); perigynia small (2.3-2.9 mm. long), usually nerveless ventrally, yellowish to brownish at maturity, tapered at base C. annectens. DEET During the course of a preliminary study of his Florida collections and before asking me to make the taxonomic decision and publish my findings, Mr. Eaton had enlisted the indispensable aid of Prof. Waterfall and Dr. Godfrey in securing the many collections of plants pertinent to the problem. Both Mr. Eaton and I are deeply grateful for their invaluable cooperation, without which it is unlikely that any plausible conclusion could have been reached. FOREST SERVICE HERBARIUM, RANGE AND WILDLIFE HABITAT RESEARCH, U. S. DEPARTMENT OF AGRICULTURE, WASHINGTON, D. C. IPOMOEA AMNICOLA (CONVOLVULACEAE), A SOUTH AMERICAN WAIF IN MISSOURI. — Among some unidentified specimens of Ipomoea in the Herbarium of the Missouri Botanical Garden, received on loan through the kindness of Dr. Walter H. Lewis, was one which proved to be Ipomoea amnicola Morong, a native of Paraguay and parts of adjoining countries, well established as a weed in extreme southern Texas and adjacent Mexico (Tamaulipas). The specimen was collected by B. F. Bush (no. 9691) in waste ground at Sheffield (now part of Kansas City), Missouri, Sept. 26, 1921. In reporting the species from Texas (see Lilloa 29: 108 and 110, 1959), the late Carlos O’Donell assumed it to be native there and disjunct, with one inter- vening record in Colombia between the North American and main South American portions of its range. The earliest collection from Texas that I have seen was made in 1929; the plant occurs in disturbed habitats such as roadsides and fields and in towns; and it is steadily increasing its range. These facts all suggest an introduced species. The Missouri collection, eight years earlier than the oldest from Texas, is additional evidence of introduced rather than native status. The manner of its introduction is unknown. It behaves like another introduced Ipomoea at first taken to be native, J. heptaphylla (Roxburgh) Voigt (I. Wrightti Gray; I. pul- chella of authors, not Roth), in showing a preference for heavier soils rather than sandy ones. The latter species, native to the Old World (probably originally from India), has been in Texas for more than 100 years, but although it has been collected in 17 central and southern counties with calcareous silt and clay, it is absent from the sandy eastern counties and from most of Louisiana. It reappears locally in southern and eastern Louisiana and in western Missis- sippi, where calcareous silts are present. LLOYD H SHINNERS SOUTHERN METHODIST UNIVERSITY, DALLAS, TEXAS 200 A REVISION OF THE NORTH AMERICAN SPECIES OF HELIANTHEMUM (CISTACEAE) (CONTINUED FROM P. 82) H. S. DAoUD & ROBERT L. WILBUR SYSTEMATIC TREATMENT* HELIANTHEMUM [Tourn.] Mill, Gard. Dict. Abr. ed, 4. 1754. Cistus L., Sp. Pl. 523-529. 1753 & Gen. Pl. 234. 1754, (in part). Helianthemum sect. Lecheoides Dunal, DC. Prodr. 1: 269. 1824. Crocanthemum Spach, Ann. Sci. Nat. 2nd ser. 6: 370. 1836. Lectotype: C. carolinianum (Walt.) Spach = H. carolinianum (Walt.) Michx. Heteromeris Spach, Ann. Sci. Nat. 2nd ser. 6: 370. 1836. Lectotype: H. canadensis (L.) Spach = Helianthemum canadense (L.) Michx. Taeniostema Spach, Ann. Sci. Nat. 2nd ser. 6: 371. 1836. Type: T. micranthum Spach = H. glomeratum (Lag.) Dun. Anthelis subg. Horanthes Raf., New Fl. N. Am. 3: 30. 1836 [1838]. Horanthes (Raf.) Raf., Sylva Tell. 132. 1838. Helianthemum subg. Lecheoides (Dunal) Reiche, Nat. Pflanzenfam. III. 6: 306. 1895. Halimium sect. Spartioides Gross., Pflanzenreich 14 (IV. 193): 33. 1903. Halimium sect. Lecheoides (Dunal) Gross., Pflanzenreich 14 (IV. 193): 33. 1903. Crocanthemum sect. Spartioides (Gross.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. Crocanthemum. sect. Lecheoides (Dunal) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. Generic Description Perennial herbs or small shrubs. STEMS one to many, decumbent to erect, arising from a woody caudex or rarely from an elongate, subterranean, woody rootstock. BASAL LEAVES, when present, either in a rosette or forming a mat. CAULINE LEAVES alternate, estipulate (an axillary tuft of leaves characteristic of several species sometimes mistaken for stipules), sessile to short-petioled, gradually reduced in size above: the blade entire or rarely subdenticulate with a con- spicuous midvein and with the pinnate secondary veins varying from very obscure to prominently elevated beneath. PUBESCENCE typically present and usually abundant; trichomes predominantly stellate but *The generic description and synonyms presented here are specific- ally intended to apply only to the North American species of Helian- themum. 201 202 Rhodora [Vol. 67 occasionally intermixed with simple or glandular hairs. INFLO- RESCENCE: cymose, paniculate, or thyrsoid. FLOWERS either chasmog- amous (petals 5; stamens 10 or more, anthers (1-) 1.5-2 times longer than wide and dehiscing laterally by means of longitudinal slits) or cleistogamous (petals lacking; stamens up to 8, anthers short and about as long as wide and dehiscing by means of rupture in the wall adherent to the stigma). Flowers either isomorphie (chasmogamous or cleistogamous) or dimorphic (both chasmogamous and cleistog- amous). When dimorphic, either occurring together in the same cluster, or separately. Bracts linear, lanceolate, or somewhat spatu- late. CHASMOGAMOUS FLOWERS: SEPALS 5, the two outer ones linear, spatulate or lanceolate, mostly shorter than the ovate inner sepals. PETALS 5, yellow, fugacious. STAMENS 10-50, filaments slender, mostly exceeding the pistil in length; anthers oblong to linear, mostly (1-)1.5-2 times longer than wide and dehiscing laterally by means of longi- tudinal slits, neither cohering with one another nor adhering to the stigma at anthesis. OVARY ovoid, one-celled, glabrous or stellate- pubescent (in 2 species); style straight and erect, varying from almost wanting to 1 mm. in length (always shorter than the ovary); stigma capitate. CAPSULE loculicidally dehiscent, ovoid, ellipsoid, or ovoid-triquetrous, shorter than calyx, glabrous or stellate-pubescent on the upper half, unilocular, usually 3-valved (one species character- istically 2-valved). SEEDS 1-45(-135), mostly irregular, ovoid, some- what flattened or rarely globose, smooth, reticulate, or papillate; the smooth seeds and the ones covered with white papillae having a thin separable membrane when moistened, (and then the embryo becoming somewhat visible), while the outer testa of the reticulate or dark papillate seeds not readily separable when wetted. CLEISTOGAMOUS FLOWERS: SEPALS 5, the two outer ones linear, spatulate, or rudimentary and knob-like and then attached near the middle edge of the ovate inner sepals. PETALS lacking. STAMENS (3-)4-6(-8), filaments slender, equaling the pistil in length; anthers short and about as long as wide, mostly coherent, dehiscing by means of a rupture of the wall adherent to the stigma and often remaining attached to the stigma even in fruit. OVARY similar to that of the chasmogamous flower but smaller; its style shorter. CAPSULE ovoid to mostly ovoid-triquetrous, other characters similar to those of the chasmogamous capsule. sEEDS smaller and fewer (1-22) but otherwise resembling those of the chasmogamous fruit. KEY TO THE NORTH AMERICAN SPECIES OF HELIANTHEMUM 1. Ovary and capsule densely stellate-pubescent at least in the upper third. 2. Capsule 3-valved: plants mostly 20 cm. tall or less; inflorescence terminal and umbellate; inner sepals of largest flowers mostly 1965] Helianthemum — Daoud & Wilbur 203 4-8 mm. long and in fruit over 5.5 mm. long; seeds mostly more numerous than 10/capsule; nw. Fla. to Miss. ........ 5. H. arenicola. 2. Capsule 2-valved; plants mostly 20 cm. tall or more; inflorescence a terminal and/or lateral, cylindrical, leafy thyrse; inner sepals of the largest flowers mostly 3.5-5.0 mm. long and in fruit not longer than 5.5 mm. long; seeds mostly less numerous than 10/ capsule; peninsular Fla. .................... eene eene 6. H. Nashü. 1. Ovary and capsule glabrous throughout. 3. Pedicels and branches of the inflorescence densely beset with coarse dark red trichomes 0.3-0.6 mm. long terminated by a small glandular knob; Channel Islands of s. Calif. ....... 9. H. Greenet. 3. Pedicels and branches lacking glandular trichomes. 4. Leaves spatulate, less than 1 cm. long, broadest well above the middle; margins with ciliate trichomes half to as long as the width of the leaf; low woody shrub; Baja Calif. .. 11. H. nutans. 4. Leaves various but not all spatulate and often exceeding 1 cm. in length, usually broadest at the middle or towards the base; leaf-margins eciliate or if trichomes present these never equal- ling in length half the width of the blade; herbaceous to shrubby. 5. Median cauline leaves 2 mm. wide or less and less than 1 cm. long; Mexico or Calif. 6. Flowers isomorphic, all petaliferous; pedicels 2-12 mm. long; foliage typically densely stellate-pubescent but varying to glabrate or rarely even glabrous; seeds 4-20/ capsule; Calif., Baja Calif. and central Mexico. 7. Median cauline leaves more than 8 times as long as broad; inner sepals up to 7 mm. long; outer sepals up to 4:.5mm; long; Calf snd m Baja Calf o a ar EE 10. H. scoparium. 7. Median cauline leaves less than 8 times as long as broad; inner sepals not exceeding 4 mm. long; outer sepals 2.5 mm. or less in length; central Mexico. ............ Een t c Li 12. H. patens. 6. Flowers dimorphic, petaliferous and apetalous; pedicels of the apetalous flowers 1 mm. long or less; foliage finely but exceedingly densely pubescent (and the pubescence not apparently stellate); seeds 3 or fewer/capsule; central Mexico sias e t TEN 20. H. argenteum. 5. Median cauline leaves 2 mm. or more in width and mostly over 1 em. long. 8. Lower surface of leaf-blade with both preminently elevated secondary veins and a visible epidermis (i.e. the pubescence sparse enough so that the epidermis is not completely obscured.) 9. Herbs up to 4 dm. high with persisting basal leaves; flowers 6 or fewer borne in a loose scorpioid cyme but 204 Rhodora [Vol. 67 appearing solitary; outer sepals lanceolate, mostly more than 0.5 mm. wide (and often up to 1.5 mm.) ; seeds 50 or more/capsule and lacking a separable membrane when moistened; se. Coastal Plain of the U.S. ennen: MEMMMMMMMMMNMMMMMMMZMM M 1. H. carolinianum. 9. Shrubs up to 10 dm, high and lacking basal leaves; flowers in dense corymbose clusters terminating branches; outer sepals linear, less than 0.5 mm. wide; seeds 10 or fewer/capsule and possessing a readily separable membrane when moistened; central and w. Mexico. ........ 14. H. concolor. DEET 8. Lower surface of leaf-blade either lacking prominently elevated veins and/or the pubescence so dense that the lower epidermis is completely obscured. 10. Lower half of stem densely beset with simple, villous trichomes as well as stellate pubescence; Mexico and Central America. 11. Secondary venation conspicuously elevated beneath; seeds lacking a hygroscopic membrane when wetted. BERE 4. H. Coulteri. 11. Secondary venation obscure or at least not conspicu- ously elevated beneath; seeds with a readily separable, hygroscopic membrane when wetted. 12. Inner sepals merely stellate-pubescent and never with villous trichomes; chasmogamous and cleistog- amous flowers borne together at the end of the branches .......... HH 16. H. Pringlei. 12. Inner sepals stellate-pubescent and also bearing villous trichomes; chasmogamous and cleistoyamous flowers borne separately (the chasmogamous termi- nating the main stem and major branches and the cleistogamous borne in 1-to few-flowered, often glomerate clusters on short axillary branches). ........ KEEN 17. H. chihuahuense. 10. Lower half of stem merely stellate-pubescent to gla- brate and never partly villous. 13. Cleistogamous flowers congested in a dense, terminal, corymbiform cyme and chasmogamous flowers, if pres- ent, long-pedicellate and overtopping the cluster; outer sepals and closely associated bracts narrowly spatulate and obtusely tipped; Coastal Plain of the se. U.S. (N.C. to Miss.) ................ sese 13. H. corymbosum. 13. Cleistogamous flowers not borne in a dense, terminal, corymbiform cyme (although often in dense glomer- ules) and chasmogamous flowers, if present, not over- topping such an inflorescence, outer sepals and the 1965] Helianthemum — Daoud & Wilbur 205 closely associated bracts linear and neither spatulate nor obtusely tipped. 14. Cleistogamous flowers (or fruit) lacking and secondary veins of leaf-blade not elevated beneath; Calif. and/or Mexico. 15. Leaves mostly 8-24 times as long as wide, their margins often revolute, often exceed- ing 1.5 em. long; plants usually exceeding 2 dm. tall; Calif. and n. Baja Calif. ........... eese EEN 10. H. scoparium. 15. Leaves mostly less than 8 times as long as wide, their margins not revolute, usually less than 1 em. long and not known to exceed 1.5 em. long; plants mostly less than 2 dm. tall and not exceeding 3 dm.; central Mexico „eenn. RR SERERE RC 12. H. patens. 14. Cleistogamous flowers (or fruit) present and/or secondary veins of leaf-blade noticeably elevated beneath. 16. Secondary veins of leaf not noticeably elevated beneath; seeds encompassed by a thin mem- brane that is readily separable when wetted; se. Coastal Plain of the U.S. or Mexico. 17. External sepals of the cleistogamous flowers less than 1.2 mm. long; internal cleistoga- mous sepals less than 2 mm. long; leaf-margin usually somewhat revolute; Coastal Plain from N.C. to e, Tex., also known from central Hispaniola. ................ 19. H. rosmarinifolium. 17. External sepals of the cleistogamous flowers 1.2 mm. long or longer; internal cleistogamous sepals exceeding 2 mm. long; leaf-margin non- revolute; Central America, Mexico, or extreme s. Tex. 18. Inflorescence a loosely racemose cyme of 2-5 + chasmogamous and cleistogamous flowers borne on elongate branches from the upper quarter of the plant; pedicels in- conspicuously jointed, articulate; plants typically herbaceous and color not ashy. RM ESOS HEURE ONG EN 16. H. Pringlei. 18. Inflorescence in dense axillary and terminal glomerules with the chasmogamous flowers (when present) on elongate pedicels con- spicuously overtopping the cleistogamous clusters; pedicels not jointed; plants suffru- 206 Rhodora [Vol. 67 ticose and of an ashy color. ........................ EE 18. H. glomeratum. 16. Secondary veins of leaf noticeably elevated be- neath; seeds either membraneous or enveloped by a readily separable thin membrane when wetted. 19. Only chasmogamous flowers and/or fruit present on specimen. 20. Chasmogamous flowers solitary and term- inating the stem but later overtopped by lateral branches (or if, when immature, 2 flowers appear together, the upper surface of the upper leaves with apparently simple trichomes intermixed with the stellate pubescence) ; e. U.S. and Canada. 21. Stems (0.5-) 1.0-2.0 (-3.0) dm. high with widely divergent branches and branch- lets; pubescence of the upper surface of the midcauline leaves dense with the stel- late pubescence so closely placed that the trichome-branches overlap; the upper epi- dermis hence obscured from view; Mass. to Long Island, N.Y. ...... 2. H. dumosum. 21. Stems (0.8-)1.5-4.5(-6.5) dm. high with strongly ascending branches and branch- lets; pubescence of the upper surface of the midcauline leaves relatively sparse with the stellate pubescence spaced so that the trichome branches rarely over- lap, the upper epidermis hence readily visible; s. Canada to Ga. and w. to Mo. and Minn. ..................ss 3. H. canadense. 20. Chasmogamous flowers clustered in groups of 2 or more. 22. Mature seeds with a separable mem- brane when wetted; calyces with only uni- form short stellate puberulence; Coastal Plain from e. N.C. to n. Fla. and w. into Ark., s. Okla. and e. Tex. ................... VERSER ANRETEERESTRREEESERIERIRRRRNERENE 15. H. georgianum. 22. Mature seeds lacking a readily separa- ble membrane after wetting; calyces oft- en with elongate trichomes in addition to the short stellate puberulence; central and e. U.S. and s. Canada (Me. to n. Ga. 1965] Helianthemum — Daoud & Wilbur 207 w. to the Dakotas and Col.) or from Mex- ico and/or Central America. 23. Midcauline leaves mostly less than 3.5 times as long as wide, leaves often broadly elliptic to somewhat obovate; Mexico and Central America ................ PEE ETA EEE UST 4. H. Coulteri. 23. Midcauline leaves mostly more than 3.5 times as long as wide, leaves usual- ly oblanceolate to narrowly elliptic; central and e. U.S. and s. Canada. 24. Chasmogamous outer sepals fused with the inner sepals only near the base and the free portion more than half the length of the inner sepals, the free portion over 3.2 mm. long; stems arising from a multicipital CEU RE eee 7. H. Bicknellii. 24. Chasmogamous outer sepals fused for half to two-thirds the length of the inner sepals, the free portion less than 3.2 mm. long; stems arising from a horizontal rootstock ................ HERRERA 8. H. propinquum. 19. Cleistogamous flowers and/or fruits present on specimen (the chasmogamous may also be present). 25. Chasmogamous flowers, when present, borne on filiform pedicels greatly overtop- ping the nearly sessile, glomerate, cleistoga- mous flowers; seeds with a separable membrane when wetted. 26. Internal sepals of the cleistogamous or fruit averaging less than 1.8 mm. long; cleistogamous capsules rounded in cross-section; median cauline leaves mostly 6 times or more as long as wide; Santo Domingo and the Coastal Plain from N.C, to n. Fla. and w. to e. Tex. and Okla. .................... 19. H. rosmarinifolium. 26. Internal sepals of the cleistogamous flowers or fruit averaging over 2.2 mm. long; cleistogamous capsules 3-angled in cross-section; median cauline leaves less than 6 times as long as wide; extreme sw. 208 Rhodora [Vol. 67 Tex., Mexico, and Guatemala. .................... NEMPE 18. H. glomeratum. 25. Chasmogamous flowers lacking or, if present, not borne on filiform pedicels over- topping the subsessile, glomerate, cleistoga- mous flowers; seeds lacking a separable membrane when wetted or with such a membrane only in A. georgianum, H. glomeratum and H. rosmarinifolium. 27. Inner sepals of the cleistogamous flowers or fruits averaging less than 1.8 mm. long; the Coastal Plain from N.C. s. to n. Fla. and w. into e. Tex. and Okla. and also in the Dominican Republic ................ NER 19. H. rosmarinifolium. 27. Inner sepals of the cleistogamous flowers or fruits averaging more than 1.8 mm. long. 28. Mature seeds present on specimen. 29. Seeds with a hygroscopic, separable membrane when wetted. 30. Cleistoramous flowers (and chasmogamous also, if present) loosely arranged and each sepa- rately borne on the ends of branch- lets; seeds per cleistogamous capsule 12 or more; capsules glo- bose to ovoid; coastal N.C. to n. Fla. and w. to e. Tex. .............. BEEN 15. H. georgianum. 30. Cleistogamous flowers in dense axillary and terminal glomerules (the chasmogamous, if present, overtopping the compact, cleistog- amous clusters); seeds/cleistoga- mous capsule fewer than 12 and mostly 2-7; capsules triquetrous; the Big Bend area of Tex. s. throughout much of Mexico and Guatemala ... 18. H. glomeratum. 29. Seeds lacking a hygroscopic, sepa- rable membrane when wetted. 31. External sepals of the cleistog- amous flowers averaging over 1.2 mm. long; Mexico and Central America sosse. 4. H. Coulteri. 1965] Helianthemum — Daoud & Wilbur 209 31. External sepals averaging less than 1.2 mm. long; U.S. e. of the Rocky Mts. and Canada. 32. Seed per cleistogamous capsule more numerous than 4, seed- surface papillate. 33. Stems (0.5-)1.0-2.0(-3.0) dm. high with widely divergent branches and branchlets; pubescence of the upper sur- face of the midcauline leaves dense with the stellate pubes- cence so closely placed that the trichome-branches overlap ob- scuring the epidermis from view; mature cleistogamous capsules and their associated calyces solitary or rarely two together at the tips of the branchlets; external sepals 0.4- 1.2 mm. long; Mass. s. to Long Island, N.Y. .. 2. H. dumosum. 33. Stems (0.8-)1.5-4.5(-6.5) dm. high with strongly ascending branches and branchlets; pubescence of the upper surface of the midcauline leaves often relatively sparse with the stellate pubescence so spaced that the trichome branches rarely overlap and hence the upper epidermis readily visible; mature cleistog- amous capsules and their as- sociated calyces one to few and glomerate, terminating the short axillary branchlets; ex- ternal sepals 0.2-0.4 (-0.6) mm. long; s. Canada to Ga. and w. to Mo. and Minn. .....:............. uc MIR RUM 3. H. canadense. 32. Seeds per cleistogamous capsule 1-3, seed-surface reticulate. 34. Stems usually clustered and upright, arising from an erect caudex; free portion of the 210 Rhodora [Vol. 67 outer sepals of the cleistoga- mous flowers linear, (0.3-)0.6- 1.2(-1.8) mm. long, about 3-5 times longer than wide; cleis- togamous capsule sharply 3- angled in cross-section ............ REM 7. H. Bicknellii. 34. Stems scattered, arising from a horizontal, elongate root- stock; free portion of the cleis- togamous outer sepals rudi- mentary and knob-like, 0.2-0.5 mm. long, 1-2 times longer than wide; cleistogamous cap- sules somewhat rounded in CroSS-Section ............eeeeeeeeeeennne BEE 8. H. propinquum. 28. Mature seeds lacking on specimen. 35. Pedicels of cleistogamous flowers over 0.5 mm, long; flowers loosely arranged at the ends of the branches or at least not in few- to many- flowered glomerules. 36. Outer sepals of the cleistogamous flowers 0.3-0.7 mm. wide; Mexico and Central America .............. ee HERR 4. H. Coulteri. 36. Outer sepals of the cleistogamous flowers about 0.3 mm. wide; Coast- al Plain from N.C. to n. Fla. and w. into e. Tex. .. 15. H. georgianum. 35. Pedicels of the cleistogamous flowers less than 0.5 mm. long, sessile or nearly so; flowers often borne in axillary clusters of 2 to many. 37. Leaves usually canescent and equally pubescent on both surfaces and hence of similar color; ex- ternal sepals of cleistogamous flowers usually 1 mm. long or long- er; extreme s, Tex. and in Mexico and Guatemala. ............ eene MEN 18. H. glomeratum. 37. Leaves neither canescent nor equally pubescent on both surfaces and usually of dissimilar color; ex- ternal sepals of the cleistogamous 196 5 ] Helianthemum — Daoud & Wilbur 211 flowers usually less than 1 mm. long (or longer in H. Bicknellit) ; U.S. and Canada e. of the Rocky Mts. 38. Cleistogamous flowers few in number and not sharply differ- entiated from the chasmogamous, borne solitary or rarely two to- gether at ends of branchlets; plants diffusely branched, 1-2 (-3) dm. high; e. Mass. s. to Long Island, N.Y. ...... 2. H. dumosum. 38. Cleistogamous flowers typically numerous and sharply differ- entiated from the chasmogamous, often borne in axillary glome- rules; plants erect, mostly 2-5 (-6.5) dm. high; e. N. Am. 39. External sepals of the cleis- togamous flowers 0.6 mm. long or more; capsule sharply tri- angular in cross-section .......... isis EEE 7. H. Bicknellii. 39. External sepals of the cleis- togamous flowers less than 0.6 mm. long; capsules weakly 3- angled or rounded in cross- section. 40. Stems arising from a horizontal rootstock, mostly less than 8 dm. tall; seeds 1-2(-3) per cleistogamous capsule; cleistogamous flow- ers few-numerous in each cluster .... 8. H. propinquum. 40. Stems arising from an erect caudex, mostly more than 2.5 dm. tall; seeds 5 or more/cleistogamous cap- sule; cleistogamous flowers one-few in each cluster ........ MORES 3. H. canadense. 1. Helianthemum carolinianum (Walt.) Michx. Cistus carolinianus Walt., Fl. Car. 152. 1788. Walter's Herb. (BM), Photo: (GH!). Probably from eastern S. Carolina. 212 Rhodora [Vol. 67 Helianthemum carolinianum (Walt.) Michx., Fl. Bor.-Am. 1: 307. 1803. Crocanthemum carolinianum (Walt.) Spach, Ann. Sci. Nat. 2nd ser. 6: 870. 1836. Halimium carolinianum (Walt) Gross., Pflanzenreich 14 (IV. 193) : 44. 1903. Heteromeris caroliniana (Walt.) Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 171. 1921. (No basionym.) Perennial herb (4)10-30(38) cm. tall with one to several stems arising from a caudex or rarely from subterranean rootstock. STEMS ascending to erect, covered with spreading, white, stellate pubescence up to 2.5 mm. long. BASAL LEAVES in a rosette; blade (3)10-35 (60) mm. long, (3)5-18(28) mm. wide, otherwise similar to the cauline leaves. CAULINE LEAVES few and remote; petiole 1-4 mm. long; blade (8)18-36(55) mm. long, (3)5-16(26) mm. wide, spatulate to obovate to elliptic or even elliptic-lanceolate near the apex, green on both sides but turning brown upon drying, sparsely pubescent above (the trichomes up to 1 mm. long), sparsely and more shortly stellate- pubescent beneath; midvein and secondary veins prominent beneath; base of blade attenuate to cuneate, apex obtuse to acute; margin sub- dentieulate, non-revolute. Inflorescence: a few-flowered, scorpioid cyme. Flowers mostly isomorphic (chasmogamous), very rarely dimorphic (chasmogamous and cleistogamous); the cleistogamous flowers, when present, one or two near the apex; bracts 2.5-6 mm. long, 0.5-1 mm. wide, lanceolate. Pedicel and calyx covered with spreading, white, pilose or 2-several-branched stellate pubescence (up to 1.5 mm. long) resembling the pubescence of the stem but often less-branched. CHASMOGAMOUS FLOWERS: pedicels 4-15(24) mm. long, slender, as- cending or sometimes curving upwards. OUTER SEPALS (free portion) (2.5)4-5.5(7.5) mm. long, (0.4)0.8-1.2(1.5) mm, wide, lanceolate, acute; INNER SEPALS (6)7.5-12(14) mm. long, (3)4-5.5(6.3) mm, wide, ovate, acuminate. COROLLA yellow, petals 8-18 mm. long, 8-16 mm. wide, broadly spatulate. STAMENS 20-35(50). PISTIL 1.9-2.7 mm. long; ovary 1.5-2.3 mm. long, 1.3-2.2 mm. in diameter, ovoid, glabrous; style 0.1-0.4 mm. long; stigma c. 1 mm. wide, capitate, parillate. FRUITING PEDICELS up to 28 mm. long; fruiting calyx 8.5-12(16.5) mm. long, 6-9 (11) mm. in diameter. OUTER SEPALS (free portion) 4-7(9) mm. long, lanceolate, up to 28 mm. long; fruiting calyx 8.5-12(16.5) mm. long, 6-9(11) mm. acute; INNER SEPALS 8.5-12(16.5) mm. long, 1.5-3-times longer than the outer sepals, 5.4-7.5 (9) mm. wide, ovate, acuminate. CAPSULE 6-9 (10.5) mm. long, 4.5-9(10) mm. in diameter, ovoid, glabrous, 3-valved (very few capsules having 4 or even 5 valves); each valve 3.8-7 mm. wide, ovate, acute to subacute. SEEDS 80-92(135), ovoid, reddish to dark brown, papillate, CLEISTOGAMOUS FLOWERS: lacking in about 99 per cent of the speci- mens examined; when present, pedicels 0.4-3.8 mm. long. OUTER 1965] Helianthemum — Daoud & Wilbur 213 SEPALS (free portion) 1.6-2.8 mm. long, lanceolate; INNER SEPALS 3-4.5 mm. long, c. 1.6 mm. wide. STAMENS 4-6. PISTIL c. 1.5 mm. long; ovary c. 1.2 mm. long, c. 0.7 mm. in diameter, style c. 0.2 mm, long; stigma c. 0.4 mm. wide. CLEISTOGAMOUS FRUITS not seen. FLOWERING: March-May. HABITAT: dry pine barrens, sandy open woodlands and fields. DISTRIBUTION: along the Coastal Plain from southeastern North Carolina into central Florida and westward into eastern Texas and southern Arkansas. (Map 1.) Grosser (Pflanzenreich 14 (IV. 193) : 45. 1908.) first de- scribed this species as possessing both chasmogamous and cleistogamous flowers. He was followed by Barnhart (in Small, Man. SE. Fl. 879. 1933.) who described it as having *petaliferous flowers with 25-35 stamens and 80 ovules, the apetalous with 5 stamens and 50-60 ovules." Usually this species has only chasmogamous flowers. However, a score or so specimens from Florida amounting to less than 1 per cent of the collections examined had one or two cleistoga- mous flowers borne near the apex late in the growing season. No cleistogamous fruits were observed. Spach (Ann. Sci. Nat. 2nd ser. 6: 370. 1836.) included H. carolinianum and H. brasiliense in his generic segregate, Crocanthemum. The basic criterion for this segregation was that both species have only petaliferous flowers. However, H. carolinianum very rarely has cleistogamous flowers while H. brasiliense typically has them. Therefore the basic charac- teristic employed by Spach to justify this segregation fails to hold. In addition, the two species do not form a particwar- ly closely related group. For example, the seeds of H. carolinianum are papillate and without a separable mem- brane, while the seeds of H. brasiliense are smooth and possess a separable membrane. REPRESENTATIVE SPECIMENS: NoRTH CAROLINA: Robeson Co., 1.7 mi. n. of Lumberton, Ahles 23716 (DUKE, NCU); Wilson Co., 1.4 mi. sw. of Black Creek, Radford 35719 (DUKE, NCU), SOUTH CARO- LINA: Hampton Co., 3.1 mi. nw. of Yemassee, Ahles 12432 (DUKE, UNC) ; Horry Co., 2 mi. s. of Myrtle Beach, Weatherby & Griscom 16585 (GH, NY). GEORGIA: Screven Co., about 1 mi, n. of Sylvania, Harper 2081 (GH, MO, NY, US); Sumter Co., 1.5 mi. se. of Flintside, Moore & Lawrence 671 (GA, SMU, UC). FLORIDA: Alachua Co., Curtiss 225 (CU, F, KANU, NY, PH, UARK, US); Clay Co., Hibernia, March 1869, Canby 214 Rhodora [Vol. 67 (F, MO, NY, PH, US); Duval Co., near Jacksonville, Curtiss 5830 (FLAS, GH, ILL, ISC, MIN, NCU, NY, SMU, UC, US). ALABAMA: Covington Co., 13 mi. sw. of Andalusia, Duncan & Hardin 14970 (GA); Mobile Co.: Mobile, Benke 3370 (F). MISSISSIPPI: Harrison Co., Biloxi, Tracy 5142 (IND, MIN, MSC, NY, 08) ; Wayne Co., 5 mi. s. of Buskatunna, Shinners 27096 (SMU). ARKANSAS: Hot Springs Co., near Malvern, Palmer 29689 (MO, UARK); Ouachita Co., Camden, Demaree 16783 (SMU). LOUISIANA: Natchitoches Par., Natchitoches, Palmer 7703 (NY, MO, US); Rapides Par., vicinity of Alexandria, Ball 614 (GH, MO, NY, US); Vernon Par., 1 mi. s. of Mayo, McVaugh & Harvill 8474 (GH, MICH). TEXAS: Harris Co., near Lindale n. of Houston, Tharp & Barkley 171006 (COLO, CU, DUKE, FLAS, GH, ILL, ISC, MIN, OKL, PENN, TEX, US) ; Jasper Co., 3 mi. ne. of Evadale, Cory 52727 (GH, SMU). 2. Helianthemum dumosum (Bickn.) Fern. Crocanthemum dumosum Bickn., Bull. Torrey Club. 40: 613. 1913. Type: Bicknell, 21 Sept. 1899 (Nv!). Nantucket Island, Massa- chusetts. Helianthemum dumosum (Bickn.) Fern., Rhodora 19: 60. 1917. Cespitose, perennial herb, (5)10-20(30) em. tall, arising from a multicipital caudex. STEMS numerous, forming loosely ascending to de- pressed mounds, the pubescence pale and stellate, sometimes intermixed with minute, red, glandular hairs; unbranched at first flowering but soon becoming much branched with stiffly divergent to ascending branches. CAULINE LEAVES: petiole 1-2 mm. long; blade (7)12-17(26) mm. long, (2)3-5(10) mm. wide, elliptic to rarely oblanceolate; the blade’s upper surface yellowish-green and densely stellate-pubescent but also sparsely intermixed (especially when young) with simple pilose hairs (0.5-1.0 mm. long) and occasionally with minute, red, glandular hairs; lower surface hoary stellate-tomentose; midvein and secondary veins prominent beneath; base of blade cuneate, apex acute; margin entire, mostly non-revolute. FLOWERS: dimorphic (chasmog- amous and cleistogamous); maturing at different times during the growing season and at different positions on the plant. At first anthesis with terminal or subterminal, solitary (but sometimes at first appear- ing paired) chasmogamous flowers (but subsequent development leav- ing it in the angle of the bifurcated stem, or in the axil of the branch when one of the two branches fails to develop) ; mostly succeeded by few chasmogamous flowers intermediate in size of calyx and number of stamens between the early ones and the late cleistogamous flowers. Pedicel and calyx of the chasmogamous flowers stellate-pubescent inter- mixed with apparently simple, pilose trichomes (1.0-1.5 mm. long) and with minute, red glandular hairs. Cleistogamous flowers solitary (rare- ly two) at tips and forks of the leafy branchlets. Pedicels and calyces of cleistogamous flowers covered with stellate pubescence and simple pilose hairs (c. 1 mm. long) intermixed with red, glandular hairs. 1965] Helianthemum — Daoud & Wilbur 215 CHASMOGAMOUS FLOWERS: pedicels 2.4-6.0 mm. long, mostly shorter than the calyx. OUTER SEPALS (free portion) 2-6 mm, long, 0.8-3.0 mm. wide, lanceolate to narrowly triangular, acuminate to acute, varying in extent of fusion with the edge of inner sepals from near the base to the middle of the inner sepals; INNER SEPALS 7-10 mm. long, 4.0-6.5 mm. wide, ovate, acute to acuminate. PETALS 5-15 mm. long, 3.6-13.0 mm. wide, yellow, obovate. STAMENS 12-36. PISTIL 2.4-3.5 mm, long; ovary 1.8-2.4 mm. long, 1.4-1.8 mm. in diameter, ovoid, glabrous; style 0.6-1.0 mm. long; stigma 1.2-2.0 mm. wide, capitate. FRUITING PEDICELS 3-9 mm. long. FRUITING CALYX 7-11 mm. long, 3.5-6.0 mm. in diameter, ovoid. OUTER SEPALS (free portion) 2.4-7.2 mm. long, 0.8- 3.0 mm. wide; INNER SEPALS 7-11 mm. long, 4.0-6.5 mm, wide. CAPSULE 4-7 mm. long, 2.5-4.0 mm. in diameter, ovoid, glabrous, 3-valved, each 22.3.8 mm. wide, ovate, acute. SEEDS 16-53, ovoid to inequilateral, dark brown, papillate, without separable membrane. CLEISTOGAMOUS FLOWERS: very few on the plant and formed late in the growing season, solitary (or rarely two), subsessile. FRUITING PEDICEL 1-3 mm. long, OUTER SEPALS (free portion) 0.4-1.2 mm. long, 0.3-0.6 mm. wide, rudimentary and knob-like to triangular and acute; INNER SEPALS 3-6 mm. long, 2.4-3.2 mm. wide, ovate, acute. STAMENS 5(8). CAPSULE 3.2-4.5 mm. long, 2.4-3.2 mm. in diameter, ovoid- triquetrous, glabrous, 3-valved; each valve 2.4-3.2 mm. wide, ovate- elliptic, acute. SEEDS 8-14, similar to the seeds of the chasmogamous flowers. FLOWERING: chasmogamous flowers late May-June, cleis- togamous flowers July-September. HABITAT: dry sandy bar- rens and open wcods. DISTRIBUTION: eastern Massachusetts southward to Long Island, New York. (Map 9). Helianthemum dumosum is very limited in distribution and consequently relatively little collected. It is apparently closely related to H. canadense and is often confused with it. The similarity and the differences between the two species are presented in the discussion of H. canadense. In addition, H. dumosum has leaves often smaller and more crowded on the divergent to ascending branches and branchlets and its chasmogamous outer sepals are relatively broader. Accord- ing to Bicknell (Bull. Torrey Club 40: 614. 1913.), it blooms earlier than H. canadense when both are growing in the same locality. Usually, H. dumosum may be clearly distinguished by the above mentioned characters and those indicated under H. canadense. However, it is sometimes difficult to refer a specimen to one species or the other, especially if it was 216 Rhodora [Vol. 67 collected in the middle of the growing season when H. dumo- sum becomes somewhat more erect and approaches H. cana- dense in habit. A few perplexing specimens have been col- lected on Long Island. The temptation to treat H. dumosum as a subspecific variant of H. canadense was deemed unwar- ranted largely because the two when growing together, as they often do within the limited range of H. dumosum, main- tain their distinguishing features, which implies some sort of effective isolating mechanism. Bicknell (Bull. Torrey Club 40: 614. 1913.) in his original description of this spe- cies concluded that “C. dumosum is evidently a strongly established derivative of C. canadense even if it be not yet wholly disconnected from that species.” REPRESENTATIVE SPECIMENS: MASSACHUSETTS: Barnstable Co.: Harwich, Fernald 383 (DS, DUKE, F, GA, IA, IND, ISC, KANS, MICH, MIN, MO, NCSC, NO, NY, OKL, PENN, PH, SMU, TENN, TEX, UARK, UC, UMO, US, WIS, WVA); Bristol Co.: Nonquitt, 6 June 1889, Sturtevant (MIN); Dukes Co.: near Terry's Pond, West Tisbury, Seymour 1278 (DUKE, GH, NY, US); e. end of Island, Nashawena, Fogg 3557 (MIN, PENN) ; Middlesex Co.: southerly slopes of Arlington Heights, Fernald, Long & St. John 9945 (PH); Nantucket Co.: Nantucket Island, Bick- nell 5893 (NY, PH); Plymouth Co.: e. of Plymouth, Cape Cod, Schus- ter A-3964 (DUKE). RHODE ISLAND: Newport Co.: Grace Point, Block Island, Fernald, Long & Torrey 9941 (GH, ILL, PH). CONNECTICUT: New London Co.: Groton, 14 July 1929, Janssan (ISC). NEW YORK: Suffolk Co.: Montauk, Long Island, Ferguson 1 (NY); Nassau Co.: Hempstead Plains, Long Island, 30 May 1906, Bicknell (GH, NY). (To BE CONTINUED) Volume 67, No. 770 including pages 97-216, was issued June 30, 1965. w REFERENCE LIBRARY 73 R47 FARLO OCT 1 1960 Hovdova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL d STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER » Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. Vol. 67 July-September, 1965 No. 771 CONTENTS: Taxonomy of Haplopappus, Section Isopappus (Compositae) BB OMh eioten nan aid a a e 217 A New Species of Senecio from Costa Rica R. M. King .... 239 Soil Algae of Forty Ponds Under Construction at Ithaca, New York Grace D. Lanciani and John M. Kingsbury .... 242 A Revision of the North American Species of Helianthemum (Cistaceae) concluded H. S. Daoud and Robert L. Wilbur 255 The Extended Distribution of Eragrostis tracyi from Sanibel Island, Florida Olga Lakela ...................... eere ennt 312 The New England Botanical Club, Jue. Botanical Museum, Oxford St., Cambridge 38, Mass. RHODORA.—A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price, $6.00 per year, net, postpaid, in funds payable at par in United States currency in Boston; single copies (if available) $1.80. Back vol. umes 1-58, with a few incomplete, can be supplied at $5.00 per volume, Volume 59— available at $6.00. Somewhat reduced rates for complete sets can be obtained upon application. Scientific papers and notes, relating directly or indirectly to the plants of North America, will be considered by the editorial com- mittee for publication. Articles concerned with systematic botany and cytotaxonomy in their broader implications are equally accept- able. All manuscripts should be double-spaced throughout, Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Albion R. Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire. Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to Albert F. Hill, Botanical Museum, Oxford Street, Cambridge 38, Mass. Second Class Postage Paid at Boston, Mass. Manufactured by THE LEXINGTON PRESS, INC. Lexington, Mass. Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 67 July-September, 1965 No. 771 TAXONOMY OF HAPLOPAPPUS, SECTION ISOPAPPUS (COMPOSITAE). ? EDWIN B. SMITH Haplopappus divaricatus (Nutt.) Gray is a wide-ranging, herbaceous annual which is found in sandy soil throughout most of the Coastal Plain of the southern and southeastern United States. Because of differing chromosome numbers reported for H. divaricatus (n — 4, Turner & Ellison, 1960, as Croptilon divaricatum Raf.; n —5, Jackson, 1959), a study of the taxon was undertaken in 1961. An aneuploid chromosome series was discovered in which the different chromosome numbers were found in morphologically recog- nizable entities (Smith, 1964). These are designated Hap- lopappus divaricatus (n — 4), H. rigidifolius (n — 5), H. validus subsp. validus (n — 5), H. validus subsp. torreyi (n — 6), and H. validus subsp. graniticus (n — T). Formal taxonomic recognition of the new taxon and new combina- tions is given in a later section. A metaphase I configuration of H. rigidifolius is shown in Fig. 1. Configurations of the other taxa at M, have been presented earlier (Smith, 1964). Haplopappus divaricatus and relatives have been recog- "This study was supported in part by an NSF Summer Fellowship, an NSF Cooperative Graduate Fellowship, and a Sigma Xi-RESA Grant-in-Aid of Research. zt am indebted to Dr. R. L. McGregor and Dr. R. C. Jackson (U. of Kansas, Lawrence, Kansas) for their generous assistance in this study and their valuable suggestions concerning the manuscript. I thank Dr. B. L. Turner (U. of Texas, Austin, Texas) for critical review of the preliminary manuscript. 217 218 Rhodora [Vol. 67 Km |, leg Figure 1. Meiotic metaphase I of Haplopappus rigidifolius. The ver- tical reference line = about 4 microns. Inked-in tracing from a photo- graph. nized in several different genera, including Inula, Chrysop- sis, Croptilon, Diplopappus, Isopappus, and Aster, as indicated in the synoymy given later. Recently, however, this group has been treated as either section Isopappus of Haplo- pappus (Hall, 1928) or as the separate genus Croptilon (Shinners, 1951). Hall’s treatment included H. divaricatus (Nutt.) Gray and H. occidentalis Hall. The latter has since been recognized as Benitoa occidentalis (Hall) Keck (Keck, 1956). Blake (1932) referred a new species from Mexico, Haplopappus bartlettii (as “Aplopappus’’), to section Iso- pappus but this entity was later transferred to Heterotheca by Johnston (1957). I have examined the holotype of H. bartlettii (Bartlett 10046, MICH), plus a second specimen (Muller 2854, MICH), and found that the entity differs in numerous details from other section /sopappus taxa. Its general appearance is characteristic Heterotheca and I would concur with Johnston's placement of it there, at least until considerably more information (biosystematic, cyto- genetic) on it becomes available. Shinners' treatment included the single species Croptilon divaricatum (Nutt.) Raf., with varieties divaricatum, hir- tellum (Shinners) Shinners, and hookerianum (T. & G.) Shinners. Although my own work has been at the specific and infra- specific level, I see no reason to treat the section as a separ- ate genus and, for the present at least, I prefer to follow Hall (1928) in this regard. 1965] Haplopappus — Smith 219 METHODS AND MATERIALS Field studies were made throughout the range of Haplo- pappus divaricatus, including Kansas, Oklahoma, Arkansas, Texas, Lousiana, Mississippi, Alabama, Florida, Georgia, South Carolina, and North Carolina. Live transplants and seeds were collected for hybridization studies, buds were fixed in a modified Carnoy's solution (95% ethanol, chloro- form, propionic acid; 2:1:1) for cytological analysis, and herbarium specimens were taken for later examination and measurement. Meiotic chromosome counts were made from anther squashes in propio-carmine. Mitotic counts were made from root tip squashes in aceto-orceine, following a 3-5 minute hydrolysis in 15% HCl. Hybridizations between pairs of the taxa were attempted in all combinations. The technique used in the crosses was the same as described by Jackson (1962), except the crossing of two particular heads was made only once and many dif- ferent heads were used. All taxa in the section are self- incompatible. The F,, F., and backcross progeny were grown in the greenhouse. Pollen stainability of each plant was sampled by staining 20-30 minutes in Aniline Blue (0.1%) in lacto- phenol and counting a minimum of 300 grains per sample. Those grains staining deep blue were counted as stainable; grains not evenly stained were counted as non-stainable. Later, an apparently equally accurate and much more rapid stain was used: Aniline Blue (water soluble, c. 0.2%) in 90% propionic acid (propionic acid: water, 9:1). With this stain, counting can begin within 60 seconds after prep- aration of the slide. RESULTS Table 1 contains a synopsis of the major quantitative characters by which the taxa differ. Formal and more com- plete descriptions of the taxa are given in a later section. Table 2 shows chromosome counts from field collections of the different taxa made during 1962-64. Voucher specimens for at least one count of each taxon are in the University of Kansas Herbarium. Generally 1-2 (-14) counts were 220 Rhodora [Vol. 67 Figure 2. Known distribution by county of the taxa in section Iso- pappus. Circles = . Haplopappus divaricatus (n = 4); squares = H. rigidifolius (n — 5); parabolas — H. validus subsp. validus (n — 5); triangles = H. validus subsp. torreyi (n = 6); rectagles = H. val- idus subsp. graniticus (n = 7). made per population. Additional counts from first and sec- ond generation plants grown in the greenhouse from random seed collections of wild populations would make the total number of counts about double the number shown in Table €) The count under subsp. graniticus marked with an asterisk is the only evidence of euploidy found in the field in the section. A single plant of this makeup was found in 1962 and previously reported as “morphologically n = 7 with anomalous pairing” (Smith, 1964). Reexamination of permanent slides of meiotic material from the plant show it is a simple triploid. Supernumerary chrosomes occur in a low frequency in subspecies of Haplopappus validus. Their absence in H. rigidifolius (Table 2) may be due to insufficient sampling, 1965] Haplopappus — Smith 221 while in H. divaricatus they appear to be absent or rare. The supernumerary chromosomes are small and near-meta- centrie, presumably consisting only of a centromere and adjacent chromatin. Figure 2 is a distribution map by county of the five taxa. Most of these locations were visited by the author. However, numerous locations were obtained from specimens from various herbaria, particularly the excellent collection of the University of North Carolina. Table 1. Synopsis of key quantitative characters of taxa in the section Isopappus. The first number is the range, the second (under- lined) is the arithmetic mean, the third is the standard deviation. Based on measurements from 25-40 individuals of each taxon. divar. = H. divaricatus; rigid. = H. rigidifolius; validus = H. validus subsp. validus; torreyi = H. validus subsp. torreyi; granit. = H. validus subsp. graniticus. Character divar. rigid. validus torreyi granit. Hgt. to 1st 30-97 6-97 20-81 19-88 16-51.5 head (cm) 59:7 51.8 49.7 46.7 31.8 18.8 22.8 14.3 13.8 8.8 Number of 5-11 5-22 13-29 10-17 10-21 ray flowers 8.1 14.6 19.7 14.4 16.6 1.4 4.8 1.0 1.8 3.2 Number of 9-26 6-60 33-108 23-51 20-79 disc 15.8 28.2 59.2 $8.9 45.9 flowers 3.9 15.7 15.9 ES, 16.9 Ligule 3.9-6.0 4.0-7.4 6.0-12.0 5.7-11.0 6.3-14.0 length. (mm) 5.29 5.21 9.16 8.30 9.96 0.59 0.96 1.42 1.28 2.04 Ligule 1.9-3.2 1.7-2.6 1.8-3.1 24:89 2.7-5.0 width (mm) 2.39 208 2.48 2.87 3.86 0.34 0.28 0.31 0.25 0.58 Involucre 5.0-7.0 4.5-8.0 5.5-8.0 5.0-9.5 5.5-10.0 height (mm) 6.08 6.04 7.01 7.93 7.76 0.53 0.99 0.59 0.84 1.28 Receptacle 2 5-4.0 1.8-5.0 4.0-7.0 3.5-6.0 3.9-6.0 diameter (mm) | 3.01 3.54 5.95 4.83 4.89 0.39 0.95 0.72 0.40 0.69 222 Rhodora [Vol. 67 Table 2. Chromosome counts made from field collections of the dif- ferent taxa during 1962-64. Designations for the taxa follow the scheme of Table 1. su — supernumerary univalent; sp — supernumer- ary pair; III — trivalent, II — bivalent, I — univalent. No. locations No. plants Chromosome number Taxon sampled counted divar. 30 36 n=4 rigid. 5 9 n=5 validus 41 127 n=5 2 2 n = 5 + sp; 2n = 12 1 1 2n = 11 torreyt 18 35 n= 1 1 n — 6 + su; 2n = 13 granit. 16 69 n=T7 2 3 n =T tsu; 2n = 15 1 1 n =T + sp; 2n — 16 *1 1 2n = 21; 3III, 5II, 2I, etc. The results of attempted hybridization between pairs of the taxa are shown in Figure 3. F, hybrids were generally vigorous and intermediate in most characters. They flow- ered somewhat later than the average time of the parental taxa. Minor aberrations, such as chlorotic spotting and striping, fused leaves, and abortion of the first few heads occurred at a low frequency in the F, hybrids. One voucher specimen of each successful F, hybrid conbination (Fig. 3) is in the University of Kansas Herbarium. Only a few F, plants were produced from most of the different Fis, despite numerous crosses. Table 3 gives the results of pollen stainability tests of the F». Essentially all crosses involving Haplapappus rigidifolius failed completely Table 3. Pollen stainability of F: progeny. Designations for taxa fol- low the scheme of Table 1. F: No. plants Pollen stainability (%) Range Average divar. X validus 20 | . 119-993 | | 693 ` divar. X torreyi 15 5.3-92.2 33.6 divar. X granit. 1 — 95.6 validus X granit, 6 39.8-70.3 54.6 torreyi X validus 3 42.4-63.7 53.0 torreyi X granit. 29 32.7-99.7 76.4 1965] Haplopappus — Smith 223 Haplopappus divaricatus PN N ~ EN. IN Haplopappus validus Haplo. (edd? subsp. graniticus 1.8% (3) Ppeppus vigidifolius n=?7 D. Haplopappus validus subsp. validus Haplopappus validus subsp. torreyi n=6 n=5 Figure 3. Crossing polygon, showing avenues for gene exchange among taxa in the section Jsopappus, based on average pollen staina- bility of F, hybrids. The width of the avenues connecting the taxa is proportional to the average pollen stainability of the F, hybrids. Average pollen stainability is given, with the number of individuals tested following in parentheses. Arrows on the avenues point toward the pistillate parent in crosses in which successful Eis were obtained in one direction only. F; seed was obtained in the cross subsp. validus (pistillate) X subsp. torreyi (staminate), but all resultant progeny died in the seedling stage. The dotted line indicates that the attempted hybridization failed in both directions. in the F.. One cross, H. validus subsp. torreyi X H. rigidi- folius, is represented in the F, by a mere two seedlings presently growing in the greenhouse. A small number of backcross progeny were obtained. However, since the number of these was low and their aver- 224 Rhodora [Vol. 67 age pollen stainability did not differ significantly from that of the F,, they are not included in Table 3. Benitoa occidentalis would not hybridize with either H. validus subsp. validus or subsp. graniticus. This evidence while negative, lends genetic support to Keck's (1956) rec- ognition on morphological grounds of H. occidentalis as the monotypic genus Benitoa. DISCUSSION The synopsis of quantitative characters separating the taxa in section /sopappus (Table 1) shows that there is considerable overlap for these characters. It is pertinent here to point out that, for the most part, plants from which the measurements were taken were selected to show the range of morphological variation found in the field and that the characters listed in Table 1 are influenced more or less strongly by environmental conditions. Available moisture influences the number of disc and ray flowers and thus head size, the width of the leaves, the amount of serration of the leaves, and the height of the plant. Thus, under drought conditions, the plants tend to be shorter, with fewer ray and disc flowers (hence, smaller heads), and narrower leaves which tend toward entire margins or serrulation apically. A similar effect is seen in late season forms with regard to head size. Collections of H. validus subsp. validus made after about mid-September and of H. rigidifolius after about mid-November tend to have much smaller heads (with reduced numbers of disc and ray flowers). Neverthe- less, if collections are made during normal peak blooming period from populations under more or less normal environ- mental conditions, the overlap is relatively small. The quan- titative characters (Table 1) combined with the qualitative characters of peduncular pubescence are generally sufficient to distinguish the taxa with ease. Haplopappus rigidifolius is sharply delimited from the other taxa in the section by its lack of glandular stipitate pubescence on the peduncles. H. divaricatus is likewise easily distinguished in the vast majority of cases from H. 1965] Haplopappus — Smith 225 validus by its small heads. It is only in H. validus that the overlap becomes a problem, and that is precisely one of the reasons for placement of the entities validus, torreyi, and graniticus at the subspecific level. These three entities over- lap in morphology but have different means, are separated at least partially by sterility barriers, and are mostly allo- patric in their distributions (Fig. 2). Where they overlap geographically, or closely aproach one another, there is evi- dence of a low frequency of hybridization (at least between validus and torreyi). The distributions of the taxa overlap somewhat in south- ern Texas (Fig. 2). This sympatry, however, is more appar- ent than real. All of the taxa generally occur in small, localized populations, covering several square acres to sev- eral square miles. Their distribution is generally not con- tinuous. Thus, although the distribution of Haplopappus divaricatus, H. validus subsp. torreyi, and H. rigidifolius appears to be sympatric in Bastrop and Lee counties, Texas (Fig. 2), they are allopatric for the most part. In Lee County, divaricatus and rigidifolius are sympatric at Tangle- wood, but subsp. torreyi was found about 7, 17, and 23 miles south of Tanglewood, with the areas between apparently devoid of the three taxa. H. divaricatus was found again about 4 miles southeast of the extreme southern population of subsp. torreyi in the county. A similar situation prevailed in Bastrop County, where divaricatus was found about 10 miles northeast of Bastrop, subsp. torreyi about 2 miles southeast and rigidifolius about 5 miles southeast of Bastrop. While sympatry of two or more of the taxa appears to be indicated in 10 counties in Texas (Fig. 2), it was actually observed in only a few cases, and in these cases the sympatry involved only H. rigidifolius and either H. divaricatus or H. validus subsp. torreyi. Where two or more taxa were found to be sympatric, such as at Tanglewood, Texas (H. divaricatus & H. rigidi- folius) and at Palmetto State Park in Gonzales County, Texas (H. rigidifolius & H. validus subsp. torreyi), an at- tempt was made to locate hybrids between the taxa, but none 226 Rhodora [Vol. 67 were found. However, there is an area in northern Texas, discussed later, in which it appears likely that subspecies validus and torreyi have hybridized. Artificial hybridizations between the taxa were success- ful in 9 of the 10 possible combinations. Reciprocal crosses were lumped and considered the same when it became appar- ent that no significant differences in morphology or sterility could be detected between them. This was true of all crosses except those involving Haplopappus rigidifolius as the pis- tillate parent and in H. validus subsp. validus (pistillate) X subsp. torreyi (staminate) (see Fig. 3). Probably the failure of the F, seedlings of the validus X torreyi cross and the few successful hybrids obtained in the reciprocal cross (torreyi X validus) was due to minor genetic differ- ences accidentally introduced in the small size of the breed- ing stock used. Failure of crosses using H. rigidifolius as the pistillate parent, however, was consistent and presum- ably reflects a reciprocal difference, perhaps cytoplasmic in action. The relatively small number of hybridizations at- tempted (total of c. 40 heads crossed in each case) and the high sterility barrier, resulting in very few Fis in the crosses that succeeded, do not allow any definite conclusions to be made. Assuming that pollen stainability can be taken as an index of fertility, there is no great sterility barrier among the subspecies of Haplopappus validus (Fig. 3). While subsp. graniticus and validus would appear to be able to exchange genes directly only with difficulty, there is an avenue for gene exchange through the intermediate subsp. torreyi, at least in the direction of validus to graniticus. These three taxa hence form a natural group. The remaining two taxa, Haplopappus rigidifolius and H. divaricatus, are strongly isolated from the H. validus group, rigidifolius being more strongly isolated than divari- catus. Also, the isolation of these two taxa is phyletically in differing directions. The only hybridization of the 10 pos- sible combinations among the taxa which failed was that between rigidifolius and divaricatus. It should be empha- 1965] Haplopappus — Smith 221 sized that H. rigidifolius X H. divaricatus (and a few reciprocal) pollinations were performed many times in 1962, 1963, and 1964. The few achenes that did develop produced 2n — 10 seedlings without exception. Whether these resulted from parthenogenesis or self-fertilization of H. rigidifolius is not known, but the latter is presumably the cause. “Self- ing" is known in other self-incompatible Haplopappus taxa when stimulated by pollen from a related entity (Jackson, 1962). Haplopappus rigidifolius is essentially totally isolated from all other members of the section, and is morphologi- cally and cytologically the most distinct member of the section. It is not surprising that rigidifolius can occur sympatrically with H. divaricatus and H. validus subsp. torreyi with few or no hybridizations occurring. In comparing the average pollen stainability of the F: hybrid combinations (Table 3) with the F, (Fig. 3), it can be seen that the average pollen stainability increases in the F,. Indeed, a few F, plants in each category have regained nearly complete fertility. Most of these were, as far as chromosome number is concerned, reconstituted par- ental types. It is apparent that a minor amount of gene flow between all the taxa, including Haplopappus rigidi- folius, is possible. But when one considers that (a) the taxa are mostly allopatric, (b) the reproductive effectiveness of the F, hybrids is rather sharply reduced (based on the low pollen stainability of the F, and the low number of F, obtained), and (c) the average fertility of the F, is con- siderably below normal, it would appear that the amount of such gene flow is, at most, minor. It would seem quite likely that competition from normal individuals in the popu- lation would preclude the survival of most of the progeny resulting from the occasional hybridizations that might occur. The H. validus group may be exceptional in this respect. In the field, hybrids would be much more likely to back- cross with one of the parents than cross with each other to produce an F,. However, the few backcross progeny obtained 228 Rhodora [Vol. 67 indicate that average fertility in backcrosses would also be rather low. There appear to be three natural groups among the taxa of section Isopappus: H. validus, H. divaricatus, and H. rigidi- folius. Based on morphological differences, the geographical distributions, chromosome number differences, difficulty in obtaining F, and F, hybrids, and high sterility in the F; and F., appropriate names are formally proposed for the taxa. SECTIONAL DESCRIPTION OF ISOPAPPUS Mostly erect annual to weak perennial herbs, with alternate, spatu- late, lanceolate, oblanceolate, to linear leaves, height to first head mostly about 25-65 cm. Leaves setose-ciliate basally and glandular stipitate on the margins up to about 1/3-1/2 their length, or, in one species (rigidifolius), not glandular stipitate but ciliate nearly over entire margin and more or less on the flat of the blade. Margins of median leaves irregularly serrate or serruate apically, or as much as the upper 1/3-3/5 serrate, rarely entire (but commonly entire in rigidifolius). Stems striate, more or less glabrate basally. Peduncles with hispid or glandular stipitate, or both hispid and glandular stipitate pubescence; in one taxon (rigidifolius) sometimes with hir- tellous, a low crisp-hairy pubescence, or even glabrous. Inflorescence a panicle, with heads of bright yellow or orangish-yellow flowers. Heads with fertile dise and ray flowers, Receptacle 1.8-7.0 mm. wide. Involueres turbinate or, infrequently, hemispherical or cylindric- turbinate. Phyllaries several-many, imbricated in several (c. 3) series, linear-attenuate to lanceolate, greenish, with scarious margins, strong- ly reflexed in age, with short glandular stipitate, hirtellous, glandular, or hispid and glandular stipitate pubescence on the back. Ray flowers 5-29, pistillate, with conspicuous ligules (ligules 3.9-14.0 mm long, 1.7-5.0 mm wide). Disc flowers 6-108, perfect, the corolla gradually enlarged above the tube. Achenes subulate, about 2.0-3.2 mm long, serecious-canescent, stramineous to brown cr reddish-brown. Pappus a single series of equal (or nearly equal) capillary bristles, more or less ferruginous. In the following key, and the descriptions which follow it, peduncle pubescence is based on the first 2.5 cm below the heads. Hispid pubescence at the base of bracts should be ignored. Floral measure- ments are based on fresh heads and on boiled heads from herbarium specimens. KEY TO TAXA IN HAPLOPAPPUS, SECTION ISOPAPPUS A. Peduncles hirtellous to hispid, with or without a lower, crisp- hairy pubescence intermixed, rarely glabrous, sometimes slightly 1965] Haplopappus — Smith 229 glutinous; plants leafy to summit, decumbent to erect; cauline leaves spatulate to lanceolate, entire (sometimes serrulate apical- IA DM ue eU Mene nene depre iecte nt 1. H. rigidifolius. A. Peduncles obviously or densely glandular-stipitate pubescent, with or without hispid pubescence; plants not leafy to summit (leaves decreasing in size up the stem to small bracts above), erect; cau- line leaves lanceolate to linear, serrate to serrulate apically (rare- IV A23 001.13 BEER B. B. Number of ray flowers 6-9 (5-11); receptacle diameter about SAL ic E CIT OT S MET NRW PEN S 2. H. divaricatus. B. Number of ray flowers 13-21 (10-29); receptacle diameter 4.7- 5.4 mm (8.5-1.0) e a e e a A Eene eet C. C. Peduncles with long-stiped, glandular-stipitate pubescence (longest hairs c. 0.4-0.5 mm, including gland), often slightly hispid just below the heads; ligule width 2.0-2.8 mm (1.8- 3.1); taxon of northern Texas and northward to central Kansas e NEM 3a. H. validus subsp. validus. C. Peduncles with short-stiped, glandular-stipitate pubescence (longest hairs c. 0.2-0.3 mm, including gland), never hispid; ligule width 2.6-4.2 mm (2.4-5.0); taxa of central and EUNDO RS, epiesptcenpncurcatsnnntiad san uitae siben OE NEUE D. D. Ligule width 2.6-3.2 mm (2.4-3.3); occurs on sand and gravel of Coastal Plain from near Austin south to near the Gulf Coast (one exceptional population occurs near Carrizo Springs in Dimmit Co.) ...... 3b. H. validus subsp. torreyi. D. Ligule width 3.2-4.2 mm (2.7-5.0); occurs exclusively in granite outcrop areas in and near the Central Mineral Region of Texas, rooted in cracks in the granite and in shallow sand deposits on the granite (one exceptional population occurs in coarse sand from c. 4-9 mi. se. of La Grange in Fayette Co.) .. 3c. H. validus subsp. graniticus. 1. Haplopappus rigidifolius Smith spec. nov. Isopappus divaricatus (Nutt.) T.&G. var. hirtellus Shinners, Field & Lab. 18:157. 1950. (T: 10 mi. s. of Falfurrias, Brooks Co., Texas, C. L. € A. A. Lundell 10813, SMU!) Croptilon divaricatum (Nutt.) Raf. var. hirtellum (Shinners) Shin- ners, Field & Lab. 19:134. 1951. Annual to weak perennial herb, decumbent to sprawling, or erect; stem highly variable in length, striate, strongly branched, with the basal branches often equalling or even exceeding the main axis in diameter; leaves rather rigid (hence: rigidifolius), alternate, spatu- late to THAM entire (occasionally serrulate apically), ciliate near- ly over entire margin and often on the flat surface of the blade, about 2.5-4.5 (2.0-5.8) cm long and 0.2-0.5 (0.2-0.8) cm wide; peduncles hirtellous to hispid, often with a lower crisp-hairy pubescence inter- 230 Rhodora [Vol. 67 mixed, sometimes slightly glutinous, rarely glabrous; heads several in a somewhat closed or compact panicle; receptacle diameter 2.9-4.0 (1.8-5.0) mm; phyllaries narrowly lanceolate, acuminate, c. 4.5-7.0 mm long and 1.0-1.6 mm wide; rays 9-21 (5-22); ligules 4.5-6.3 (4.0-7.4) mm long and 2.0-2.5 (1.7-2.6) mm wide; disc corollas 25-45 (6-60); achenes subulate, serecious-canescent, 2.0-2.4 mm long, stramineous to pale reddish-brown, sometimes mottled. Chromosome number n = 5. Blooms September-November and later. Distinguished easily from other taxa in the section by its lack of glandular stipitate pubescence on the peduncles. Known from 23 counties in southern Texas (Fig. 2). A new epithet is required for this species under Haplo- pappus, since the name H. hirtellus had previously been used for a species from Chile (Hall, 1928). Haplopappus rigidifolius is highly variable, especially with regard to number of disc and ray flowers, as well as general size. Collections range from strictly decumbent, weak peren- nials with stems less than 10 cm long and small heads (Ar- ansas and Victoria counties, Texas) to erect annuals with stems as much as 70 cm or more long and large heads (Live Oak and Refugio counties, Texas). There is some correla- tion of the above characteristies with distance from the ocean, but the variation is not strictly clinal. The coastal, decumbent population might be considered a seacoast eco- type, as in Succisa pratensis, Matricaria inodora (Stebbins, 1950), Clarkia prostrata, or C. davyi (Lewis & Lewis, 1955). Taxonomic recognition of the variation does not, however, appear justifiable, since there is considerable intergradation and, even in the same population, some heterogeneity for the characters mentioned. Representative Specimens: TEXAS: ARANSAS Co., Goose Island State Park, Smith 91, KANU, Johnston 53.280.174, TEX; AUSTIN Co, 4.8 mi. s. of Kenney, Smith 259, KANU; BASTROP Co., 5.1 mi. se. of Bastrop, Smith 553, KANU; BEE Co., c. 21.2 mi. w. of junction of 202 & US 183, Smith 635, KANU, SMU, GA, FSU, & NCU; BURLESON Co., 1.1 mi. s. of Somerville, Smith 260, KANU; COLORADO Co., 3 mi. s. of junction of 71 & US 90A, Smith 257, KANU; GONZALES Co., near Palmetto State Park, Smith 243, KANU; KARNES Co., 2.5 mi. ne. of Kenedy, Johnson 1354, SMU, TEX; KENEDY Co., 0.5 mi. n. of Sarita, Smith 92, KANU, 5 mi. n. of Armstrong, Turner 4474, TEX; KLEBERG Co., King Ranch, Johnston 54416, TEX; LAVACA Co., 13 mi. n. of Hallettsville, Shinners 1965] Haplopappus — Smith 231 28701, SMU; LEE Co., at Tanglewood, Smith 236, KANU; MEDINA Co., near Devine, Barkley 13947, FSU, TEX; NUECES Co., 1 mi. s. of Flour Bluff, Jones 745, SMU; REFUGIO Co., c. 4 mi. w. of junction of 202 & US 183, Smith 638, KANU & TEX; VICTORIA Co., 2.0 mi. sw. of Inez, Smith 252, KANU, 8.3 mi. sw. of Victoria, Shinners 25235, SMU; WHAR- TON Co., 3.7 mi. s. of Nada, Smith 256, KANU; WILLACY Co., 4 mi. w. of Redfish Bay, Johnston 54179, TEX. Also known to occur on Padre Island. 2. Haplopappus (Isopappus) divaricatus (Nutt.) Gray, Explor. Rail- road Route Miss. River to Pacific 4:99, 1856. (as “Aplopap- pus"). Inula divaricata Nuttall, Gen. 2:152. 1818. (T: vicinity of Savan- nah in Georgia, perhaps PH) Chrysopsis divaricata (Nutt.) Elliot, Bot. Sketch 2:338. 1824. Croptilon divaricatum (Nutt.) Raf., Fl. Tellur. 2:47. 1836. Diplopappus divaricatus (Nutt.) Hook., Comp. Bot. Mag. 1:97. 1836. Isopappus divaricatus (Nutt.) T.& G., Fl. N. A. 2:239. 1842. Aster divaricatus (Nutt.) Kuntze, Rev. Gen. 1:318. 1891. Annual herb, erect, virgate before fully in bloom; stem 4.0-7.0 (2.0-11.5) dm high, striate, glabrate basally, glandular stipitate at the apex, more or less ferruginous; leaves alternate, basal ones oblan- ceolate to spatulate, entire to irregularly serrate, setose-ciliate at the base, 9-12 (7-14) cm long, 1.5-2.0 em wide; median leaves lanceolate, irregularly serrate, occasionally one or more doubly serrate, setose- ciliate at the base, glandular stipitate on the margins, 3-5 (2.8-8.0) em long, 0.2-0.4 (0.15-1.5) em wide; peduncles with mixed glandular stipitate and hispid pubescence, infrequently to rarely not hispid; heads many in an open panicle; receptacle diameter 2.8-3.4 (2.5-4.0) mm; phyllaries lanceolate, the largest c. 1 X 5 mm, short glandular stipitate and sparsely hispid on the back; rays 6-9 (5-11); ligules 4.5-6.0 (3.9-6.0) mm long and 2.0-2.6 (1.9-3.2) mm wide; disc corollas 14-22 (9-26); achenes c. 2 mm long, subulate, somewhat sparsely serecious-canescent, stramineous to reddish-brown. Chromosome num- ber n = 4. Blooms late August-October. Distinguished from H. rigidifolius by its glandular-stipitate pubescence on the ped- uncles, and from other members of the section by its small heads and low number of ray and disc flowers. Known from 266 counties in 11 states (Fig. 2). A casual weed in most areas of occurrence, Haplopappus divaricatus becomes very abundant in sandy areas in Geor- gia and South Carolina. The distribution of H. divaricatus 292 Rhodora [Vol. 67 is apparently divided into an eastern and a western popula- tion by the broad band of river alluvium in the southern Mississippi River valley (see Baker, 1936; Plate 4). H. divaricatus, as well as other taxa in the section, appear to be poor competitors on soils high in clay or low in sand. The infrequent to rare form with peduncles glandular- stipitate pubescent only (no hispid pubescence) appears to be localized in a few areas at the western and southwestern edge of the distribution and does not appear to be worthy of taxonomic recognition. Haplopappus divaricatus specimens lacking hispid pubescence on the peduncles are sometimes similar to depauperate specimens of H. validus subsp. tor- reyi. Representative Specimens: ALABAMA: BUTLER Co., 6.1 mi. ne, of junction of US 31 & Ala. 10, Smith 538, KANU; CHAMBERS Co, 0.1 mi. sw. of junction of road 147 & US 431, Smith 479, KANU; HENRY Co., 8.9 mi. s. of Abbeville, Smith 537, KANU; LEE Co., 10.8 mi. w. of junc- tion of 40 & 29 in Auburn, Smith 478, KANU; WASHINGTON Co., c. 0.5 mi. s. of Choctaw Co. line on US 17, Smith 539, KANU. ARKANSAS: CLEVELAND Co., 1.6 mi. w. of Kingsland, Smith 457, KANU; Dallas Co., just e. of Manning, Smith 456, KANU; LITTLE RIVER Co. 2 mi. n. & 2.6 mi. w. of Winthrop, Smith 128, KANU. FLORIDA: ALACHUA Co. just sw. of Archer, Smith 529, KANU; GILCHRIST Co., just outside of Wilcox, Smith 527, KANU; JACKSON Co., at Marianna Caverns State Park, Mitchell 897, FSU; LAKE Co. 16.3 mi. ssw. of Leesburg, Smith 533, KANU; LEON Co., vicinity of Silver Lake, Godfrey 60356, FSU. GEORGIA: Bartow Co., 2.1 mi. s. 52° w. of Allatoona Dam, Duncan 8887, GA; CLARKE Co., vacant lot in Athens, Cronquist 4132, GA; DOUGHERTY Co., R.R. yards in Albany, Thorne 5856, GA; EMANUEL Co., 9 mi. e. of Adrian, Wilbur 2923, GA; GREENE Co., 3.7 mi. se. of Siloam, Smith 492, KANU. LOUISIANA: BIENVILLE Co., 2.2 mi. e. of Hagewood, Smith 543, KANU; DESOTO Co., 3.6 mi. nw. of Keatchie, Smith 274, KANU; NACHITOCHES Co., 2.1 mi. e. of Hage- wood, Smith 544, KANU, SMU, & TEX; OUACHITA Co., 1 mi. e. of Cal- houn exit on Interstate 20, Smith 542, KANU; SABINE Co., 3.9 mi. s. of Zwolle, Smith 273, KANU. MISSISSIPPI: JASPER Co., 2.7 mi. w. of junction of 504 & 508, Smith 462, KANU; LAUDERDALE Co., near Meri- dian, Smith 464, KANU; NEWTON Co., dry ridge thickets, Demaree 36155, FSU, 4.3 mi. nw. of junction of Miss. 504 & 508, Smith. 460, KANU. NORTH CAROLINA: ANSON Co., 2.6 mi. se. of Morvin, Smith 510, KANU; BLADEN Co, s. of Elizabethtown, Smith 513, KANU; HARNETT Co., c. 4 mi. w. of Mamers, Laing 251, GA; LEE Co., 7.5 e. of Sanford, Smith 512, KANU; MONTGOMERY Co., 1.3 mi. n. ‘of Can. 1965] Haplopappus — Smith 233 dor, Smith 511, KANU. OKLAHOMA: McCunRTAIN Co., 0.5 mi. n. of Tom, Waterfall 10496, TEX; PUSHMATAHA Co., 7 mi. w. of Antlers, Waterfall 11147, TEX. SOUTH CAROLINA: AIKEN Co., 1.1 mi. n. of Aiken, Smith 501, KANU; CHARLESTON Co., 1.9 mi. e. of junction of S.C. 174 & US 17, Smith 521, KANU; FAIRFIELD Co., c. 5 mi. w. of Rockton, Smith 507, KANU; GREENWOOD Co., 0.2 mi. e. of Abbeville Co. line on S.C. 72, Smith 4,99, KANU; KERSHAW Co., 6.2 mi. w. of junction of S.C. 34 & US 1, Smith 508, KANU; LEXINGTON Co., 1.1 mi. n. of Aiken Co. line on S.C, 215, Smith 502, KANU. TEXAS: ANDERSON CoO., 6 mi. nw. of Tennessee Colony, Marsh 283, TEX; BASTROP Co., 8.2 mi. sw. of junction of 21 & US 290, Smith 554, KANU; FANNIN Co., 1.4 mi. e. of Telephone, Smith 286, KANU; FREESTONE Co., 12 mi. ese. of Fairfield, Turner 4438, TEX; LEE Co. at Tanglewood, Smith 237, KANU; LEON Co., 5 mi. e. of Buffalo, Gould 7276, TEX; LIMESTONE CO., 5.5. mi. e. of Kosse, Shinners 30574, SMU; MILAM Co., 6.1 mi. se. of junction of 36 & US 79, Smith 558, KANU; Morris Co., 2 mi. se. of Daingerfield, Whitehouse 17656, SMU; NACOGDOCHES Co. Cushing, Tharp & Brown 53-8, TEX; RAINS Co., 5 mi. s. of Emory, Smith 562, KANU; ROBERTSON Co., 0.5 mi. sw. of Hearne, Smith 560, KANU, near New Baden, Tharp & Barkley 13978, TEX; TriTuS Co. 1.6 mi. ese. of Mt. Pleasant, Shinners 16079, SMU; TRINITY Co., 1.9 mi. nw. of Groveton, Smith 266, KANU; UPSHUR Co., 1.5 mi. e. of Big Sandy, King 2179, TEX; VAN ZANDT Co., 5.7 mi. se. of Willis Point, Shinners 8469, SMU. 3a. Haplopappus validus (Rydb.) Cory Rhodora 38:406. 1936. (as “Aplopappus”) subsp. validus Isopappus validus Rydberg, Brittonia 1:100-101. 1951. (T: 6 mi. s. of Ellinwood, Barton Co., Kansas, P. A. Rydberg & R. Imler 1309, NY. Paratypes: Rydberg & Imler 616 & 721 KANU). Annual (facultative winter annual) herb, erect; stem 4.5-6.5 (3.0- 7.5) dm high, striate, glabrate basally, glandular stipitate above; leaves alternate, basal ones oblanceolate to spatulate, entire to irregu- larly serrate, setose-ciliate at the base, 7-9 (6-10) em long, 1.9-2.3 (1.2- 3.0) em wide; median leaves lanceolate, irregularly serrate, setose- ciliate at the base, glandular stipitate on the margins, 4.5-6.5 (3.0-8.0) em long, mostly 0.3-1.2 em wide; peduncles with glandular stipitate pubescence, the stipes rather long (0.4-0.5 mm, including gland), often sparsely hispid, especially just below heads; heads several in a somewhat rigid panicle; receptacle diameter 4.7-6.0 (4.0-7.0) mm; phyllaries lanceolate, the largest about 1.1 X 6.8 mm, short glandular stipitate, and sometimes sparsely hispid, on the back; rays 16-23 (13-29) ; ligules 7-10 (6-12) mm long and 2.0-2.8 (1.8-3.1) mm wide; dise corcllas 55-70 (33-108); achenes 2.2-2.6 (2.0-3.0) mm long; subulate, serecious-canescent, stramineous to reddish-brown. Chromo- some number n = 5. 234 Rhodora [Vol. 67 Blooms July-early October. Distinguished from Haplo- pappus validus subsp. torreyi and subsp. graniticus by its narrower ligules, from H. rigidifolius by its glandular- stipitate pubescence on the peduncles, and from H. divarica- tus by its larger heads. Known from 60 counties in Kansas, Oklahoma, and nothern Texas (Fig. 2). This subspecies is mostly limited to relatively small popu- lations in sandy areas along rivers and fence rows, except in the stationary sand dunes in the Big Bend area of Kansas where it assumes great abundance. Marked seedlings have been observed to withstand pro- longed subfreezing temperatures and grow to maturity the following summer. Seedlings of the outer taxa in the section are rapidly killed by the first severe frost. Populations of subsp. validus in nothern Texas, east of the Panhandle, are almost uniformly atypical for the taxon. This is especially true of Parker, Johnson, Tarrant, Dallas, Wise, Denton, McLennan, and Callahan counties. They nearly all lack hispid pubescence on the peduncles and have smaller heads with fewer flowers than usual. It is believed that most populations in these counties are carrying one or more subsp. torreyi chromosomes, or at the very least, have picked up subsp. torreyi genes through hybridization. Num- erous somatic chromosome counts from these populations were nearly all 2n — 10. However, one 2n — 11 and a few 2n — 12 counts were made. One count of 2n — 10 showed a heteromorphic pair, one homologue of which could have been from subsp. torreyi. It is hoped that further study, including genetic analyses, will confirm the hybrid nature of certain individuals in these populations. Representative Specimens: KANSAS: DICKINSON Co., 2.9 mi. w. of Abilene, Smith 122, SMU; ELLSWORTH Co., 0.5 mi. e. RE ferra Cotta, Smith 121, KANU; KIowA Co., 3 mi. w. of Greensburgh, MeGregor 4041, KANU; MCPHERSON Co., McPherson County State Park, Smith 120, TEX; RENO Co., 10 mi. w. of Hutchinson, Wagenknecht 3124, KANU; RICE Co., 8.8 mi. e. of Sterling, Smith 118, KANU. OKLA- HOMA: CADDO Co., 5.9 mi. e. of junction of US 281 E Okl. 9, Smith 605, KANU; CANADIAN Co., 1.4 mi. n. of El Reno, Smith 220, KANU; GRANT Co., 1.7 mi. n. of junction of US 60 & US 81, Smith 218, KANU; JEFFERSON Co., 0.2 mi. n. of Red River off US 81, Smith 227, KANU; 1965] Haplopappus — Smith 285 MAJOR Co., 1 mi. e. & 1 mi. s. of Cleo Springs, Richards 1093, KANU, 1 mi. ne. of Orienta, Waterfall 10354, TEX. TEXAS: CALLAHAN CoO., 15 mi. se. of Abilene, Henderson 62-1182, FSU (rather atypical), Hen- derson 63-1772, TEX; GRAYSON Co, 4 mi. ese. of Gainsville, Smith 564, KANU; HEMPHILL Co., 7 mi. ne. of Canadian, Rowell 4259, TEX; Hoop Co., 5.5 mi. ne. of Granbury, Shinners 10330, SMU; LIPSCOMB Co., 3.5 mi. sw. of Higgens, Cory 50288, SMU; MCLENNAN Co., n. of Gholson, Smith 928, TEX; MONTAGUE Co., 7.6 mi. e. of junction of US 82 & US 81, Smith 565, KANU; SOMERVELL Co., 2.8 mi. ne. of Glen Rose, Shinners 13812, SMU. 3b. Haplopappus validus (Rydb.) Cory subsp. torreyi Smith, subspec. nov. Isopappus hookerianus T. & G., Fl. N. A. 2:239. 1842. (T: near Gonzales, Gonzales Co., Texas, Drummond 184, K, photo- graph!) not Haplopappus hookerianus DC., 1838. Aster hookerianus (T. & G.) Kuntze, Revis. Gen. 1:318. 1891. Croptilon hookerianum (T. & G.) House, N.Y. State Mus. Bul. 223-234:61. 1921. Isopappus divaricatus (Nutt.) T. & G. var. hookerianus (T. & G.) Shinners, Field & Lab. 18:157. 1950. Croptilon divaricatum (Nutt) Raf. var. hookerianum (T. & G.) Shinners, Field & Lab. 19:134. 1951. Haplopappus divaricatus (Nutt.) Gray var. hookerianus (T. & G.) Waterfall, Rhodora 62:321. 1960. Annual herb, erect; stem c. 3-6 dm high, striate, glabrate basally, glandular stipitate above; leaves alternate, basal ones oblanceolate to spatulate, entire to irregularly serrate, setose-ciliate at the base, 8-10 (6-12) em long and 1.5-2.0 (1.2-2.5) em wide; median leaves lanceolate to linear, irregularly serrate, setose-ciliate at the base, glandular stipitate on the margins, 4-8 (3.2-10.0) cm long and 0.3-0.8 (0.2-1.1) em wide; peduncles not hispid, densely glandular stipitate, the stipes short (longest c. 0.2-0.9 mm, including gland); heads several in a spreading panicle, the peduncles sometimes reclinate; receptacle diameter 4.5-5.1 (3.5-6.0) mm; phyllaries lanceolate, the largest c. 1 X 6 mm, short glandular stipitate on the back, not hispid; rays 12-16 (10-17); ligules 7.0-9.2 (5.7-11.0) mm long and 2.6-3.2 (2.4-3.3) mm wide; disc corollas 33-47 (23-51); achenes c. 2.2 mm long, subulate, densely serecious-canescent, stramineous to pale brown. Chromosome number n = 6. Blooms August-October. Distinguished from Haplopap- pus rigidifolius by its glandular-stipitate pubescence on the peduncles; from H. divaricatus and H. validus subsp. validus by its lack of hispid pubescence on the peduncles or phyl- laries and its wider ligules; end from H. validus subsp. 236 Rhodora [Vol. 67 graniticus by its narrower ligules and its ecology (never occurs in granite outcrop areas). Known from 15 counties in southern Texas (Fig. 2). This subspecies is noteworthy for the paucity of individ- uals in most observed populations. Except for two popula- tions in Refugio Co., Texas, it has not been observed to approach the abundance of other taxa in the section. Representative Specimens: TEXAS: ARANSAS Co, Rockport, Tharp 42-81, TEX; BASTROP Co., 1.5 mi. se. of Bastrop, Smith 79, GA; CALD- WELL Co., 4.8 mi. se. of Prairie Lea, Smith 244, KANU; GOLIAD CoO., 9.5 mi. s. of Goliad, Shinners 25210, SMU; GONZALES Co., near Palmetto State Park road 11 exit, Smith 242, KANU, Gonzales, Tharp (no num- ber), TEX; KARNES Co., 1.5 mi. e. of Harmony School near Ecleto Creek Crossing, Johnson 996, SMU, TEX; LEE Co., 2.3 mi. n. of Giddings, Smith 240, KANU, 0.8 mi. w. of Manheim, Smith 557, TEX; REFUGIO Co., 3.4 mi. w. of junction of 202 & US 183, Smith 639, KANU & NCU, 3 mi. e. of Woodsboro, Jones 725, SMU; VICTORIA Co., 2.8 mi. se. of Raisin Smith 251, SMU. 3c. Haplopappus validus (Rydb.) Cory subsp. graniticus Smith subspec. nov. (T: 1.9 mi. e. of Streeter, Mason Co., Texas, E. B. Smith 624, KANU. Isotype: SMU & NCU) Herba annua, 3-6 dm alta; caulibus unicis, erectis, striatis, subtus in maturitate tomentosis vel glabratis et saepe ramosis, insuper stipito- glandulosis; foliis alternis, linearibus vel lanceolatis, serratis, subtus ciliatis, ad 8 em longis et 1 cm latis, margininibus stipito-glandulosis; capitulis pluribus, paniculatis; disco dia. 4.5-5.5 (3.9-6.0) mm; phyl- lariis anguste lanceolatis, acuminatus, ca. 7.5 mm longis et 1.5 mm latis, dorsis sparse stipito-glandulosis; radiis (10-) 13-21; ligulis 8-12 (6.3-14.0) mm longis et 3.2-4.2 (2.7-5.0) mm latis; disci corollis 35-65 (20-79), 5.0-6.5 mm longis; achaeniis subulatis, pubescentis, 2.3-9.2 mm longis; pappis 2.5-4.0 mm longis. Annual herb, 3-6 dm tall; stem solitary, erect, striate, tomentose to glabrate below in maturity and often basally branched, glandular stipitate above (peduncles); leaves alternate, linear to lanceolate, serrate, ciliate basally, up to 8 cm long and 1 cm wide, glandular stipitate on the margins; heads several in an open panicle; disc dia. 4.5-5.5 (3.9-6.0) mm; phyllaries narrowly lanceolate, acuminate, c. 7.5 mm long and 1.5 mm wide, sparsely glandular stipitate on the back; rays (10-) 13-21; ligules 8-12 (6.3-14.0) mm long and mostly 3.2-4.2 (2.7-5.0) mm wide; dise corollas 35-65 (20-79), 5.0-6.5 mm long; achenes subulate, pubescent (serecious-canescent), 2.3-3.2 mm long; pappus a single row of equal bristles, 2.5-4.0 mm long. Chromo- Some number » = 7. Blooms August-October. Distinguished from Haplopap- 1965] Haplopappus — Smith 291 pus rigidifolius by having glandular-stipitate pubescence on the peduncles; from H. divaricatus and H. validus subsp. validus by its lack of hispid pubescence on the peduncles or phyllaries and its wider ligules; and from H. validus subsp. torreyi by its wider ligules and ecology (occurs almost ex- clusively in granite outerop areas). Known from 5 counties in central Texas (Fig. 2). The flowers of subsp. graniticus are a deeper yellow than those of other members of the section. However, the differ- ence is not very noticeable until specimens are compared side by side. Except for the Fayette Co. population mentioned in the key, subsp. graniticus is limited exclusively to granite out- crop areas in and near the Central Mineral Region of Texas (see Tharp, 1952) or granitic Central Basin (see Gould, 1962). The Fayette Co. population grows in coarse sand and resembles subsp. torreyi. Subspecies torreyi and graniticus overlap in morphology, but the overlap is greater in this pop- ulation. While some members of the population key easily to subsp. graniticus, others key to subsp. torreyi. Cytological examination of numerous individuals from the population showed them all to be 2n = 14 or n= 7. Therefore, the whole population is presumed to be subsp. graniticus. The fact that this population could not be placed, with certainty, in either subsp. graniticus or torreyi without cytological examination is one of the main reasons for placement of graniticus and torreyi at the subspecific level. Representative Specimens: TEXAS: BURNET Co, Inks Lake State Park, 1.8 mi. s. of Park Headquarters, Smith 76, KANU; FAYETTE CoO., 8.6 mi. se. of La Grange, Smith 649, KANU (somewhat atypical); LLANO Co. Enchanted Rock, Whitehouse 10303, SMU; MASON Co, 2 mi. s. of Air (also called Camp Air), Smith 622, KANU, GA, TEX, & FSU, and 3 !4 mi, w. of Fredonia, Cory 43052, SMU, Cory 43051, TEX. DEPARTMENT OF BOTANY, RUTGERS THE STATE UNIVERSITY, NEWARK, NEW JERSEY LITERATURE CITED Baker, O. E. 1936. Atlas of American Agriculture. U. S. Govt. Printing Office, Washington, D.C. 238 Rhodora [Vol. 67 BLAKE, S. F. 1932. New Central American Asteraceae collected by H. H. Bartlett. Jour. Wash. Acad. Sci. 22:379-386. GOULD, F. W. 1962. Texas Plants—a checklist and ecological sum- mary. Agric. & Mechan. College of Texas, College Station, Texas. HALL, H. M. 1928. The genus Haplopappus. Carnegie Inst. Wash. Pub. 389. JACKSON, R. C. 1959. in Documented chromosome numbers of plants. Madrofio 15:52. 1962. Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. Jour. Bot. 49:119-182. JOHNSTON, M. C. 1957. in Notes. The Southwestern Naturalist 2:172. Keck, D. D. 1956. Benitoa, a new genus of Compositae from Califor- nia. Leaf. West. Bot. 8:25-28. LEwis, H. & M. E. 1955. The genus Clarkia. U. Cal. Pub. Bot. 20:241-392. SHINNERS, L. H. 1951. Notes on Texas Compositae VIII. Field & Lab. 19:134. SMITH, E. B. 1964. Chromosome numbers in races of Haplopappus divaricatus. Rhodora 66:63-66. STEBBINS, G. L. 1950. Variation and Evolution in Plants. Columbia U. Press, N.Y. THARP, B. C. 1952. Texas Range Grasses. U. Tex. Press, Austin, Texas. TURNER, B. L. & W. L. ELLISON 1960. Chromosome numbers in the Compositate 1... . The Texas Jour. Sci. 12:147. A NEW SPECIES OF SENECIO FROM COSTA RICA R. M. KING While collecting cytological material of Compositae in Costa Rica, during the summer of 1962, the author made a collection of Senecio which is apparently undescribed. Senecio costaricensis King sp. nov. Herba erecta 1.2-2 m alta, sine ramis. Folia alterna, pinnatifida usque ad 23 cm longa, usque ad 7 em lata. Inflorescentiae laxe pani- culatae corymbosae ca. 22 cm altae, 26 cm latae, bracteis lanceolatis pinnatifidis usque ad 8 cm longis, 4 cm latis. Capitula radiata, multi- flora. Involucri bracteae ca. 12, lanceolatae uniseriatae, Flores radii ca. 5, corolla flavescens, ligulata, tubo ca. 3.8 mm longo, ca. 0.3 mm diam., limbo ca. 4.5 mm longo, 1.3 mm. lato. Flores disci ca. 15, per- fecti, fertiles, regulares. Corolla hypocrateriformis 5-lobata, flava, 4 mm longa. Achaenia fulva, attenuata, ca. 1.2 mm longa, basi ca. 0.25 mm diam., apice 0.5 mm diam. Coarse herb 1.2-2 meters tall; stem erect, green, chaffy, 7-8 mm in diameter, unbranched, striate; leaves alternate, sparingly short pubescent and dark green above, subglabrous and lighter green beneath, laciniately pinnatifid, sinuate dentate, up to 23 cm long, 7 em wide; inflorescence a loose paniculate corymb, terminal, ca. 22 cm high and 26 em wide; bracts laciniately pinnatifid, up to 8 cm long, 4 cm wide, sparingly short pubescent and darker green above, sub- glabrous and lighter green beneath; pedicels sparingly short-pubes- cent, many headed; phyllaries ca. 12, lanceolate, in one series, apex short-acute, sparingly short pubescent, dark brown, up to 4 mm high, ca. 1.5 mm broad; heads radiate, campanulate; ray florets 5-7, yellow, pistillate, tube ca. 3.3 mm long, ca. 0.3 mm in diameter, limb up to 4.5 mm long and 1.3 mm wide; scattered unicellular glands on outer surface of corolla tube up to 200 & long; style branches 2, divergent, ca. 0.8 mm long, 200 uw in diameter, style tube 2.6-2.8 mm long, ca. 240 u wide; stamens absent; disc florets 12-18, perfect, fertile, regular, 5 lobed, yellow; corolla salverform, up to 4 mm long; stamens 5; an- thers ca. 1.5 mm long, ca. 100 u wide; filaments 50-60 u wide, 1.0-1.2 mm long, differentiated above into a prominent collar ca. 400 4 long, 80-100 u wide, deeply channelled and with a broadly expanded cordate base, cells of collar regularly sub-quadrate (12 X 25 u), smooth; achenes dark brown, tapered, ca. 1.2 mm long, base ca. 0.25 mm in diameter, apex 0.5 mm in diameter, ribs uneven, 7-9 (8), pappus bristles 35-40, gray-white, ca. 4.0 long, serrate; pollen spherical, spicu- late, ca. 20 u in diameter. Chromosome number, n — 20 (Turner and King 1964). 239 240 Rhodora [ Vol. 67 PLANTS OF COSTA RICA Province af Cartage Senecio sp. now, abundant, flowers yellow, steep mountain slopes of Corro de La Werte, Elevation oa, 3OOOmeLerá, ‘Chromosome comet determined from, metic material aca — T0 Roben Merrill King. ae — 3395 T5 Auget MM U Eege ed thee Lmgerbzzgecg $i PL Tumer and A. M. Powell 1962 Tustitwio Interamericano De Crvmcun Agricolas 1 Deka Plate 1312. Senecio costaricensis. (Holotype: King 5392) Habit ca. X 1/3. 1965] Senecio — King 241 Holotype in the U.S. National Herbarium R.M. King 5392 (2367999). Abundant, flowers yellow, steep mountain slopes of Cerro de la Muerte, Province of Cartago, Costa Rica. Elevation ca. 3000 meters. August 24, 1962. Isotypes: (MICH, TEX, and UC). Paratypes: J. Cuatrecasas and J. Léon 26551. Cordillera de Talamanca, Macizo del Buena Vista, Province of Cartago, Costa Rica. Elevation 3300- 3400 meters. Nov. 8, 1961 (US). Arthur Cronquist and A. F. Munos 8842. Sphagnum bog along the Pan-American highway in the mountains south of Cartago, Province of Cartago, Costa Rica. Eleva- tion about 2600 meters. Nov. 10, 1960 (NY). Senecio costaricensis has the characters of the section Eremophili Greenman (1915) and shows considerable resemblance to S. townsendii Greenman. It differs from all members of the section in its generally larger habit, larger leaves, smaller bracts, larger inflorescences and fewer flowered heads. Grateful acknowledgement is made to Dr. J. Cuatrecasas who first recognized this material 2s a new species and ad- vised me during the course of my study. Special thanks also are due Dr. Harold Robinson for his very valuable assist- ance. DEPARTMENT OF BIOLOGY CATONSVILLE COMMUNITY COLLEGE CATONSVILLE, MARYLAND REFERENCES GREENMAN, J. M. 1903. Monographie der nord-und centralamerikanis- chen Arten der Gattung, Senecio. Engl. Bot. Jahrb. 32 (19) ; 1-33. GREENMAN, J. M. 1915. Monograph of the North and Central Ameri- can Species of the Genus Senecio — Part II. Ann. Mo. Bot. Gard. 11: 573-626. TURNER, B. L. and R. M. ke 1964, Chromosome Numbers in the Compositae. VIII Mexican and Central American Species. The Southwestern Naturalist. 9: 27-39. SOIL ALGAE OF FORTY PONDS UNDER CONSTRUCTION AT ITHACA, NEW YORK! GRACE D. LANCIANI? AND JoHN M. KINGSBURY To allow experimental study of basic ecology of aquatic weeds and their control by herbicides, 92 ponds have been constructed at two locations in the Ithaca area. The purpose of this study was to determine the algal flora of the soil from which one group of 40 ponds was constructed. Such algae represent a major source of introduction for the algal popu- lations that are expected to appear in the ponds when they are completed and filled. Construction of the set of ponds used in this study was begun in July, 1962 and completed in November, 1963. Each pond is 66 feet square in the bottom dimension and 124 feet square in the top dimension when measured from center dike to center dike. The area covered by one pond is one-third of an acre at the top and one-tenth of an acre at the bottom. Proposed depths of the ponds when filled range from five to eight feet. The ponds were constructed from abandoned farm lands. The soil, Volusia silt loam, had not been recently dis- turbed when construction was begun. The group of ponds under study is arranged in two rows of ten ponds each on each side of a supply canal; a second canal runs perpendicular to the first canal at the end region of the ponds (Fig. 1). Alternate ponds and alternate banks of ponds with respect to the supply canal were sampled just after they had been given final shape. Three samples were taken from the soil at each pond: the top sample was obtained from the upper margin of the pond, the middle sample from approximately three to five feet down the slope of the pond, and the bottom sample from the floor of the pond. In each case, the top three ‘Cooperative Northeast Regional Experiment Station Project, CRF-1, The Development of Principles and Practices for the control of Submerged Aquatic Vegetation. "Based on a thesis submitted to Cornell University in partial fulfill- ment of the requirements for the degree of Master of Science. 242 1965] Soil Algae — Lanciani and Kingsbury 248 Plate 1313 Figure 1. Ponds at time of sampling. to six inches of soil was sampled in a way that yielded well- mixed 10 gm samples. Twenty ponds were sampled initially, 25 in all. Half of the initial samples were examined imme- diately and the other half refrigerated. Because the latter froze accidentally, new samples were collected two months later from the same locations. Five media were used in culturing the samples: Modified Bristol's Solution (Bold, 1949) with trace elements (Chan- tanachat and Bold, 1962); Knop’s Solution (Bold, 1942) ; Kratz and Myer's (1955) Medium D; Modified Detmer's Solution (Bold, 1942) ; Soil Extract Solution (Bold, 1942). Culture vessels were 50 ml Delong culture flasks and 18 X 150 mm test tubes. Both flasks and tubes were capped with stainless steel closures (Bellco). Flasks contained 20 ml of media. Inocula consisted of one gram portions of soil. Original isolations and subsequent transfers were placed in a culture room under standard conditions of twenty-four hour ilumination at 250-450 foot candles (warm white fluorescent) on one shelf and 575-675 foot candles on an- other and temperature of 20°C. + 2°C. 244 Rhodora [Vol. 67 To facilitate identification, various techniques were em- ployed for separating the original populations into subcul- tures containing fewer species each. By these procedures, debris was removed and cultures were obtained in which one or few algal species were isolated. No attempt was made to produce cultures free of bacteria. Flasks were checked daily for presence of algal growth. Typically, the meniscus showed green color within one to two weeks. Flasks were subcultured by the methods of Bold (1942), Pringsheim (1946; 1950), Deason and Bold (1960), Chantanachat and Bold (1962), and Mattox and Bold (1962). These methods include micromanipulation with drawn-out Pasteur capillary pipettes, streaking on agar, and inoculation into fresh media. Transfers, using the above methods, were made into Bristol’s liquid medium and on the same solidified with 1.5 per cent agar; into the original medium; and into soil water tubes (Pringsheim, 1950) made with tap water and sterilized by repeated steaming. A method used by Warcup (1950) for isolating soil fungi was also employed for isolation of soil algae. This method involves distribution of soil particles throughout a thin layer of solid nutrient mediuin. Small amounts of soil were used as inocula directly and in dilutions (with water) of 1:25, 1:125 and 1:625. A method involving dispersal of soil particles through liquid nutrient medium, proposed by Willson and Forest (1957), was also employed. Transfers were made periodically to keep the algae in the best possible condition. Colonies from petri dishes were transferred to Bristol’s liquid and 1.5 per cent agar media. Isolations were made from flasks and Willson-Forest Petri dishes by means of capillary pipettes or glass needles. Fresh mounts in distilled water were used predominantly in identification of the algae. India ink and iodine were used as necessary. Aceto-carmine preparations (Deason and Bold, 1960) proved less satisfactory than the freshmount method. Algae were identified to species using standard taxonomic references. For the majority of identifications, G. W. Pres- cott's Algae of the Western Great Lakes Area (1962) served 1965] Soil Algae — Lanciani and Kingsbury 245 as the primary source. Geitler's Cyanophyceae (1932) and Collins! Green. Algae of North America (1928) were also used in particular instances. In addition, University of Texas Phycological Studies, I-III (Deason and Bold, 1960; Chantanachat and Bold, 1962; Mattox and Bold, 1962) proved useful because of their parallel nature. Identification of species of Bacillariophyceae and certain Chlorococcales was not attempted; nevertheless, an attempt was made to estimate their numbers. Patrick’s key to genera of Bacillariophyceae (1959) was helpful in this connection, and the following technique was used to aid in estimating the number of Chlorococcales. Six flasks were chosen at random from a group of fifty flasks containing putative Chlorococcales, as determined by microscopic examination. Five successive 1 to 5 dilutions were made in sterile water from cells thrown down by cen- trifuging the original culture. These were plated out on 1.5 percent Bristol’s agar in twenty-five Petri dishes. When growth was evident, fifty well-separated colonies were se- lected at random from the twenty-five Petri dishes and inoculated into Bristol’s liquid medium. The tubes were re- turned to the culture room and examined approximately three to four weeks later when good growth was apparent. Criteria used for determining the presence of differing entities among these isolates included presence or absence of green meniscus in test tube; presence and amount of sludge; presence or absence of flagellated entities in micro- scopic examination; and morphology of adult cells, includ- ing thickness of cell wall, number of pyrenoids, and shapes of chloroplasts (Deason and Bold, 1960; Chantanachat and Bold, 1962; Mattox and Bold, 1962; Herndon, 1958; and Starr, 1955). RESULTS Seventy-two species of algae have been identified from the soil samples. Divisions represented include Cyanophyta - 43 species, Chlorophyta - 29 species, and Chrysophyta ( Bacil- lariophyceae), species not identified. A taxonomic list of species is presented in Table I. 246 Rhodora [ Vol. 67 Data were examined for possible correlations of differ- ences among the species of algae found in the top, middle, and lower levels of the pond walls and among their distribu- tions horizontally from pond to pond. Vertical distribution of representative algae appearing frequently, relatively fre- quently, and rarely is given in Table II. Considerable range in frequency of occurrence was found in both distributions, but less definite patterns were noted in the horizontal dis- tribution. Among the Chlorococcales two members were identified as Chlorococcum and Chlorella, and two of the latter were keyed out further to species. Other genera and species could not be ascertained with security with the criteria available. Members of the Chlorococcales, particularly species of Chlorococcum, appeared in every soil sample analyzed. Dia- toms appeared in thirty-eight out of forty-five samples inoculated into the Delong culture flasks. Of particular interest were four species of algae described from Texas soils (Deason and Bold, 1960; Chantanachat and Bold, 1962; Mattox and Bold, 1962) : Hormidium flaccidum Plate 1314 Figure 2. Hormidium flaccidum. One-month-old filaments. 1965] Soil Algae — Lanciani and Kingsbury 247 Plate 1315 Figure 3. Ulothrix belkae. Germling. Plate 1316 Figure 4. Ulothrix belkae. Two-month-old filaments. 248 Rhodora [Vol. 67 (Fig. 2), Ulothrix belkae (Fig. 3, 4), Chlamydomonas aggregata, and Chlamydomonas typica. Chlamydomonas appeared frequently in the samples in both vertical and horizontal distributions but could not always be keyed out to species. One species of Chlorosarcinopsis was recovered from the soil samples. On the basis of descriptions by Herndon (1958), it was identified as Chlorosarcinopsis eremi. DISCUSSION The five media used in this experiment proved to be se- lective to some degree. Bristol's solution was least selective, and for the most part, the algae grew well in this medium. For this reason, Bristol's solution was used predominatly as the medium for transfer. Knop's solution appeared se- lective for certain green algae, specifically Chlorococcales, although some blue-green algae were also isolated from this medium. Kratz and Myer's medium yielded an abundance of blue-green algae, with a large number of Chlorococcales also. In general, the soil-extract solution produced poor growth and appeared to support only a few diatoms, Chloro- coccales, and blue-greens, the latter being mostly in the state of fragmented filaments. Detmer's medium appeared to select for diatoms and blue-greens. Members of the Division Cyanophyta predominated in the samples. Of seventy-two species identified, forty-three species were blue-green algae. Thirty-five of the species of Cyanophyta were filamentous in structure; seven were coccoid (unicellular or colonial) ; and one was a species of Spirulina. Of the filamentous blue-greens, twelve were species of Oscillatoria, and six each were species of Ama- baena and Nostoc. Other genera were represented by fewer species. Blue-green algae grew well in Kratz and Myer's Medium D, pH 8.0. The pH of the soil samples also was slightly basic (7.4). Diatoms were found in high frequency in the soil samples which also contained a large number of blue-green algae. These results differ from those of Bristol (1920) who re- 1965] Soil Algae — Lanciani and Kingsbury 249 ported that soils rich in blue-green algae usually contained only few species of diatoms, and vice-versa. The large amount of Chlorococcum reported in this study agrees with observations of Smith (1944) who likewise found an abundance of Chlorococcum (hwumicolum) in four different soil types analyzed in a study of soils from Florida. Another genus frequently encountered in this study was Chlamydomonas. Lund (1947) also reports the common occurrence of species of Chlamydomonas in soil, presenting ten new species in his survey. The taxonomy of Chlamy- domonas is complicated by the very large number of de- scribed species and the frequent appearance of new species descriptions. Isolation methods were not equally successful. Isolation by streaking out on 1.5 percent Bristol’s agar and by means of micromanipulation with drawn-out Pasteur capillary pipettes proved to be most often satisfactory. The Willson- Forest plating method proved excellent for the isolation and growth of blue-greens. An outstanding advantage of this method is the fact that filamentous forms could be trans- ferred from dried-out plates and, in fresh media, growth was generally good. The modified serial-dilution and plating technique was satisfactory. Well-isolated colonies on the Petri dishes frequently yielded uni-algal cultures. The Warcup method, although it eliminated the need for such serial-dilution techniques as those proposed by Bold (1942), Skinner (1932), and Bristol-Roach (1962), is limited to the use of extremely small inocula, and the plates are prone to rapid contamination with bacteria and fungi. Several conclusions may be drawn regarding the fre- quency of occurrence and distribution of species among the samples taken at different levels from the ponds. In general, most species were found relatively frequently and were equally distributed in the three levels. More than 75 percent of the species were found in all three levels. However, some- what fewer isolations could be obtained from lower levels than from higher. One hundred seventy-nine species isola- tions were obtained from soil samples taken at the top level; 250 Rhodora [Vol. 67 169 at the middle level; and 141 at the lower level. Some exceptions to the generality that most species were found in all three levels are noted in Table II. It shows that some species occurred with equal frequency in the top two levels but with lower frequency in the bottom level. On the other hand, some species appeared with equal frequency in the top two levels yet at a higher frequency in the bottom level. Few species occurred exclusively in the top, middle, or lower level. Because the ponds were still under construction at the time of sampling and were subject to frequent mechanical disturbance, it is assumed that the populations were not well established. However, some differences in habitat among the three levels were clear. The lower level of the ponds was subject to more mixing and also to moistening with rain water which remained in the new pond bottoms for long periods of time. This alone could account for the differ- ence in frequency of occurrence of species as compared to the top two levels. Reasons for the appearance of a few species one or few times may be that occasional air-borne forms were involved, that the media were not always indu- cive to good growth of certain species, or that occasional resistant forms became vegetative. The ability to persist in dormant condition under periods inimical to growth was probably a factor in determining the types of algae found and the period in culture before they appeared. Growth of members of the Chlorophyta was rapid, with visible green color appearing in one to two weeks. Likewise, when transfers to fresh media were made, zoosporulation usually occurred within a few days to a week. Certain species, however, did not appear for several weeks to several months, probably because they required the prior germination of resistant structures. Blue-green algae, in general, appeared later than the majority of green algae but were present in most samples eventually and remained viable longer. The samples that were accidentally frozen while being stored were used in the Warcup and Willson-Forest isolation 1965] Soil Algae — Lanciani and Kingsbury 251 techniques. In general, blue-greens were the predominant forms isolated. The species of blue-greens which were iso- lated from the frozen samples compared closely with those recovered from non-frozen samples. Smith (1951) reported that the bulk of species of soil algae are generally made up of Myxophyceae (blue-green algae) and Bacillariophyceae, with Chlorophyceae third, and Xanthophyceae and Euglenophyceae contributing a much smaller number. Blue-green algae predominated in this study, but members of the Chlorophyceae and Bacillario- phyceae appeared with almost equal frequency, while no species of Xanthophyceae or Euglenophyceae were found, perhaps because the media did not select for them. DEPARTMENT OF BOTANY CORNELL UNIVERSITY ITHACA, NEW YORK TABLE I TAXONOMIC LIST OF ALGAE IDENTIFIED CHLOROPHYTA CHLOROCOCCALES Gloeocystis vesiculosa Naegeli CHARACIACEAE Palmella mucosa Kützing Characium obtusum A. Braun ULOTRICHALES CHLOROCOCCACEAE ULOTRICHACEAE Chlorococcum spp. Hormidium flaccidum A. Braun COELASTRACEAE Stichococcus bacillaris Naegeli Coelastrum microporum Naegeli Ulothrix belkae Mattox and Bold OOCYSTACEAE Ulothrix subtilissima Rabenhorst Chlorella ellipsoidea Gerneck MICROSPORACEAE Chlorella vulgaris Beyerinck Microspora stagnorum (Kützing) Oocystis crassa Wittrock Lagerheim Oocystis parva West and West CHAETOPHORACEAE Trochiscia reticularis (Reinsch) Protoderma viride Kützing Hansgirg ULVALES PROTOSIPHON ACEAE SCHIZOMERIDACEAE Protosiphon botryoides (Kützing) Chlorosarcinopsis eremi Herndon Klebs VOLVOCALES SCENEDESM ACEAE CHLAMYDOMONADACEAE 2 52 Scenedesmus hystrix Lagerheim Scenedesmus obliquus (Turpin) Kützing Scenedesmus quadricauda (Tur- pin) de Brebisson TETRASPORALES PALMELLACEAE Gloeocystis ampla (Kützing) Lagerheim Gloeocystis gigas (Kützing) Lagerheim Rhodora [Vol. 67 Chlamydomonas aggregata Dea- son and Bold Chlamydomonas snowii Printz Chlamydomonas typica Deason and Bold Sphaerellopsis fluviatilis Pascher VOLVOCACEAE Gonium pectorale Mueller ZYGNEMATALES DESMIDIACEAE Cosmarium spp. CYANOPHYTA CHROOCOCCALES CHROOCOCCACEAE Aphanocapsa elachista West and West Chroocoecus minor (Kützing) Naegeli Dactylococcopsis smithii Chodat and Chodat Synechococcus aeruginosa Naegeli Synechocystis aquatilis Sauvageau Synechocystis crassa Woronichin Synechocystis sallensis Skuja HORMOGONALES NOSTOCACEAE Anabaena affinis Lemmermann Anabaena cylindrica Lemmer- mann Anabaena flos-aquae (Lyngbye) de Brebisson Anabaena oscillaroides Bory Anabaena subcylindrica Borge Anabaena torulosa (Carmichael) Lagerheim Anabaenopsis elenkinii Miller Cylindrospermum catenatum Ralfs Cylindrospermum marchicum Lemmermann Nodularia harveyana Thuret Nodularia spumigena Mertens Nostoc commune Vaucher Nostoc linckia (Roth) Bornet and Thuret Nostoc muscorum C. A. Agardh Nostoc paludosum Kützing Nostoc sphaericum Vaucher Nostoc verrucosum Vaucher OSCILLATORIACEAE Lyngbya limnetica Lemmermann Lyngbya martensiana Meneghini Lyngbya versicolor (Wartmann) Gomont Oscillatoria acutissima Kufferath Oscillatoria agardhii Gomont Oscillatoria amoena (Kützing) Gomont Oscillatoria amphibia C. A. Agardh Oscillatoria angusta Koppe Oscillatoria angustissima West and West Oscillatoria limnetica Lemmer- mann Oscillatoria rubescens De Candolle Oscillatoria splendida Greville Oscillatoria subbrevis Schmidle Oscillatoria tenuis C. A. Agardh Oscillatoria terebriformis C. A. Agardh Phormidium minnesotense (Tilden) Drouet Schizothrix calcicola (Agardh) Gomont Spirulina major Kützing RIVULARIACEAE Calothrix stagnalis Gomont 1965] Soil Algae — Lanciani and Kingsbury 253 TABLE II Vertical distribution of representative algae found frequently, less frequently, and rarely in soil samples. FREQUENCY OF OCCURRENCE NUMBER OF OCCURRENCES TOP MIDDLE LOWER FREQUENT Oscillatoria limnetica 13 13 13 Synechocystis aquatilis 14 14 13 Chlamydomonas spp. dt 9 6 Oscillatoria angusta 8 8 12 LESS FREQUENT Scenedesmus quadricauda 5 4 2 Anabaena cylindrica 4 4 5 Nostoc commune 4 3 4 Oscillatoria agardhii 2 2 6 RARE Spirulina major 0 0 1 Anabaena cscillaroides 0 0 1 Phormidium minnesotense 1 0 0 Anabaenopsis elenkinii 1 0 0 Oscillatoria splendida 0 1 0 Anabaena flos-aquae 0 1 0 LITERATURE CITED Bon H. C. 1942. The cultivation of algae. Bot. Rev. 8: 70-138. 1949. The morphology of Chlamydomonas chlamydog- ama. sp. nov. Bull. Torrey Bot. Club 76: 101-108. BRISTOL, B. M. 1920. On the alga-flora cf some dessicated English soils: an important factor in soil biology. Ann. Bot. 34: 35-80. BRisTOL-ROACH, M. 1926. On the relation of certain soil algae to some soluble carbon compounds. Ann. Bot. 40: 149-201. CHANTANACHAT, S., and H. C. Bon 1962. Phycological Studies II. Some algae from arid soils, Univ. of Texas. Publ. No. 6218, 75 p. CoLLINS, F. C. 1928. Green Algae of North America. G. E. Stechart and Co., New York. 400 p. DEASON, T. R., and H. C. Bon 1960. Phycological Studies I. Ex- ploratory studies of Texas soil algae. Univ. of Texas. Publ. No. 60225/12:p: GEITLER, L. 1932. Cyanophyceae. Vol. 14. In: L. RABENHORST, ed. Kryptogamen-Flora. Akad. Verlags., Leipzig. 1196 p. HERNDON, W. 1958. Studies on Chlorosphaeracean algae from soil. Am. J. Bot. 45: 298-308. 254 Rhodora [Vol. 67 Kratz, W. A., and J. Myer, 1955. Nutrition and growth of several blue-green algae. Am. J. Bot. 42: 282-287. LUND, J. W. 1947. Observations on soil algae III. Species of Chla- mydomonas Ehr. in relation to variability within the genus. New Phytol, 46: 185-194. Marrox K., and H. C. Bon 1962. Phycological Studies III. The taxonomy of certain Ulotrichacean algae. Univ. of Texas. Publ. No 6222, 66 p. PaTRICK, R. 1959. Bacillariophyceae. In: W. T. EDMONDSON, ed. Ward and Whipple's Fresh-water Biology, p. 171-189. John Wiley and Sons, Inc., New York. PRESCOTT, G. W. 1962. Algae of the Western Great Lakes Area. rev. ed., Wm. C. Brown, Co., Dubuque, Iowa. 977 p. PRINGSHEIM, E. G. 1946. Pure Cultures of Algae. - Their Preparation , and Maintenance. Univ. Press, Cambridge. 119 p. 1950. The eultivation of algae. Endeavour 9: 138-143. SKINNER, C. B. 1932. Isolation in pure culture of green algae from soil by a simple technique. Plant Physiol. 7: 533-537. SMITH, F. B. 1944. The occurrence and distribution of algae in soils. Proc. Florida Acad. Sci. 7: 44-49. STARR, R. C. 1955. A comparative study of Chlorococcum meneghini and other spherical, zoospore-producing genera of the Chlorococ- cales, Indiana Univ. Sci. Ser. Publ. No. 20, 111 p. TIFFANY, L. H. 1951. Ecology of fresh-water algae. In: G. M. SMITH, ed. Manual of Phycology, p. 293-311. Chronica Botanica Co., Waltham, Mass. WaRCUP, J. H. 1950. The soil plate method for isolation of fungi from soil. Nature 166: 117-118. WILLSON, D., AND H. S. Forest, 1957. An explanatory study on soil algae. Ecology 38: 309-313. A REVISION OF THE NORTH AMERICAN SPECIES OF HELIANTHEMUM (CISTACEAE) (CONTINUED FROM P. 216) H. S. DAouD & ROBERT L. WILBUR 3. Helianthemum canadense (L.) Michx. Lechea major L., Sp. Pl. 90. 1753. Type: Clayton (in the British Museum), not seen. Photo (GH!). “Habitat in Canadae aridis." but actually from Point Comfort, southeastern Virginia. Cistus canadensis L., Sp. Pl. 526. 1753. Type: Kalm (in the Linnaean Herbarium), not seen. Photo (GH!). “Habitat in Canada.” Helianthemum canadense (L.) Michx., Fl. Bor.-Am. 1: 308. 1803. As to basionym only; Michaux's specimens were H. Bicknellii accord- ing to Fernald's annotated copy of Michaux's Flora at the Gray Herbarium. Helianthemum ramuliflorum Michx., Fl. Bor.-Am. 1: 307. 18083. Type: Michaux. (P, not seen). “In Georgia et Carolina." Fernald's notes in the interleaved copy confirm the long-standing suspicion that this species is a synonym of H. canadense. Heteromeris canadensis (L.) Spach, Ann. Sci. Nat. 2nd ser. 6: 370. 1836. Heteromeris Michauxii Spach, Comp. Bot. Mag. 2: 291. 1837. Nom. illegit., H. ramuliflorum Michx. cited in synonymy. Helianthemum majus (L.) BSP., Prel. Cat. N.Y. 6. 1888. Halimium canadense (L.) Gross, Pflanzenreich 14(IV. 193): 51. 1903. Crocanthemum canadense (L.) Britt. & Brown, Ill. Fl. ed. 2. 2: 540. 1913. Heteromeris major (L.) Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 169. 1921. (No basionym and probably not the species meant by Ponzo.) Helianthemum canadense (L.) Michx. var. sabulonum Fern., Rhodora 43: 615. 1941. Type: Fernald & Long 4044 (GH!) "sand dunes south of False Cape, Princess Anne County, Virginia." Perennial herb, (6)15-45(65) cm. tall with few to many stems arising from a multicipital caudex. STEMS ascending to erect, stellate- pubescent but occasionally becoming glabrate, mostly unbranched at first anthesis, later producing numerous ascending floriferous branch- es. CAULINE LEAVES: petiole 1-2(3) mm. long; blade (7)18-30(38) mm. long, (3)5-7(10) mm. wide; oblance-elliptic to narrowly elliptie near the top of the stem; upper surface green and somewhat lustrous, sparsely stellate-pubescent and mostly intermixed with simple pilose 255 256 Rhodora [Vol. 67 hairs 0.5-1.0 mm. long, lower surface hoary and stellate-tomentose; midvein elevated and secondary veins prominent beneath; base of blade attenuate to cuneate, apex rarely obtuse, subacute to acute; margin entire, non-revolute at first anthesis, later the upper leaves become slightly revolute. FLOWERS: dimorphic (chasmogamous and cleistogamous; maturing at different times during the growing season and occupying different positions on the plant). At first anthesis with a terminal (or subterminal), solitary, chasmogamous flower (but subsequent development of the plant leaves it in the angle of the ‘bifurcated stem or sometimes one of the branches fails to develop and the flower appears in the axil of the branch, or even more occasionally a second or a third chasmogamous flower borne some distance above the first). Pedicel and calyx of the casmogamous flower stellate- pubescent intermixed with apparently simple pilose hairs 0.5-1.2 mm. long and occasionally with very few red glandular hairs. Cleistoga- mous flowers much smaller, sessile to subsessile, (1)-few in glomerules terminating the ascending branches and also in 1-few glomerate clus- ters terminating the short branchlets in the leaf-axil. Pedicel and calyx of the cleistogamous flowers stellate-tomentose. CHASMOGAMOUS FLOWERS: pedicels (1.5)4.0-9.5 (17) mm. long. OUTER SEPALS (free portion) (1.2)3-5(7) mm. long, 0.5-1.0(1.2) mm. wide lanceolate or nearly so, acute; INNER SEPALS (5)6-9(10) mm. long, (3)3.5-5.0(5.6) mm. wide, ovate, acute. PETALS 8-15 mm. long, 6-14 mm. wide, yellow, obovate. STAMENS 20-35. PISTIL 1.8-3.0 mm. long; ovary 1.5-2.4 mm. long, 1-2 mm. in diameter, ovoid, glabrous; style 0.3-0.5 mm. long; stigma 0.6-1.2 mm. wide, capitate. FRUITING PEDICELS (1.5)7-12(18) mm. long. FRUITING CALYX (5)7.0-9.6 (11.5) mm. long; (4.5)6-8(9) mm. in diameter, ovoid. OUTER SEPALS (free portion) (1.2)3.6-6.0(7) mm. long, (0.5)0.7-1.4(1.7) mm. wide, lanceolate or nearly so, acute; INNER SEPALS (5.0)7.0-9.6(11.5) mm. long, (3)5-7(7.7) mm. wide, ovate, acute. CAPSULE (4)6-7(8.5) mm. long, 4-7 mm. in diameter, ovoid, somewhat angled, glabrous, 3-valved (few capsules seen with 4 or 5 valves); each valve 3.5-6.6 mm. wide, ovate, acute. Funiculi and placentae persisting after the separation of valves and seeds. SEEDS 35-46, ovoid, to inequilateral, dark brown, papillate, lacking a separable membrane. CLEISTOGAMOUS FLOWERS: either sessile or on pedicels up to 0.5 mm. long. FRUITING PEDICELS (0.5)1-2(4) mm. long. FRUITING CALYX (2.4) 2.8-3.5 (4.3) mm. in diameter, subglobose to globose. OUTER SEPALS (free portion) 0.2-0.4(0.6) mm. long, 0.2-0.4 mm. wide, rudimentary, knob- like, and attached to the middle edge of the inner sepals; INNER SEPALS (2.4) 2.8-3.5(4.3) mm. long, (2.4)2.8-3.4(3.8) mm. wide, obliquely obo- vate, concave, imbricate. STAMENS 3(5). CAPSULE (2)2.3-3.0(3.8) mm. long, (2)2.5-3.0(4) mm. in diameter, ovoid-triquetrous, somewhat depressed, glabrous, 3-valved (few capsules seen with 4 valves) ; each valve (2)2.5-3.0(4) mm. wide, ovate-elliptic, slightly concave, some- 1965] Helianthemum — Wilbur 257 times laterally folded at maturity. SEEDS 5-9(12), similar to the seeds of the chasmogamous flowers. FLOWERING: Chasmogamous flowers, late March-July; cleistogamous flowers, May-September. HABITAT: sandy flats, dunes, “barrens”, open dry woodlands, prairies, and rocky wooded slopes. DISTRIBUTION: scattered north as far as southern Nova Scotia; and from southern Maine south to southern Georgia and westward into eastern Minnesota, Iowa, and Missouri. (Map 4.) Helianthemum canadense was the first New World species of Helianthemum to be described and for a very long time was thought to include as well the three other northeastern species, H. dumosum, H. Bicknellii, and H. propinquum. Even after their recognition as distinct entities, it has often been confused with them. Helianthemum canadense appears much more closely related to H. dumosum than to H. Bick- nellii or H. propinquum. The major characteristics shared by H. canadense and H. dumosum which distinguish them from H. Bicknellii and H. propinquum are compared below. H. canadense & H. dumosum H. Bicknellii & H. propinquum 1. Early chasmogamous flowers 1. Early chasmogamous flowers typically solitary, terminal or 2-18, rarely overtopped by the subterminal, soon overtopped lateral branches. by the lateral branches. 2. Upper surface of cauline 2. Upper surface of cauline leaves stellate-pubescent and leaves stellate-pubescent only. mostly sparsely intermixed with simple, pilose hairs 0.5- 1.0 mm. long (best observed on younger leaves). 3. Cleistogamous capsule (2.0) 3. Cleistogamous capsule 1.5-2.0 2.4-3.5(4.0) mm. in diameter (2.4) mm. in diameter each each with 5-14 papillate seeds. with 1-2(3) reticulate seeds, It is not surprising to find that H. dumosum was the last species to be separated from H. canadense. Both species have fairly uniform floral morphology (especially in the chasmog- amous flowers), similar seeds, and generally similar pubescence. The major distinctions between H. canadense and H. dumosum are as follows: 258 H. canadense . Stems ascending to erect at first anthesis, and later in the growing season up to 65 cm. tall, with ascending branches and branchlets. . Upper surface of cauline leaves somewhat lustrous and less densely stellate-pubescent. . Cleistogamous flowers (1)-few in glomerules at the top of the ascending branches and on the short axillary branchlets. Rhodora [| Vol. 67 H. dumosum 1. Stems relatively compact, loose- ly ascendent or divergent, and the plants often bushy at first anthesis, and later in the grow- ing season up to 30 cm. tall. . Upper surface of cauline leaves non-lustrous and densely stel- late-pubescent. 3. Cleistogamous flowers solitary, rarely two at the forks and tips of the divergent to ascend- ing branchlets. Helianthemum canadense and the other northeastern species, H. dumosum, H. Bicknellii, and H. propinquum, vary greatly in their morphological appearance during the period from first anthesis to maturity. These extreme variations cause considerable difficulty in identification. Hooker (Fl. Bor.-Am. 1: 72. 1830.) pointed out the remark- able dissimilarity between the chasmogamous and cleistoga- mous phases of this species which had led previous workers, including Linnaeus, to describe the latter phase as a Lechea. It is to be remembered though that part of the great diversi- ty that so impressed the elder Hooker was due to the in- clusion of all four northeastern or widespread eastern species within the concept of H. canadense. Linnaeus included this species twice, once in its petalife- rous phase as Cistus canadensis and again in its cleistoga- mous state as Lechea major. Other than the remarkable floral differences, this species is morphologically rather uniform throughout its range. A few specimens, representing not more than one percent of the total number of collections examined, possessed stems branching from near the base and with the tips of the branches crowded with cleistogamous flowers. Some of these fruiting specimens had relatively longer pedicels and slightly larger capsules. Their elliptic-obovate leaves were also more crowded. Fernald (Rhodora 43: 615. 1941.) designated some specimens with these features as the new 1965] Helianthemum — Wilbur 259 variety sabulonum. Description of Fernald's variety as it appeared in Gray's Manual is quoted below in full: Var. sabulonum ‘Fern. (of sands).— Stems few, decumbent or loosely ascending; leaves oblong-elliptic, often canescent above; cleistogamous flowers mostly long-pedicelled, in loose corymbs termi- nating upper branches, uniform, in maturity 4-5 mm. in diameter. — Dunes and open sands, local, se. Mass. and Oneida L., N.Y., to se. Va. However, the four herbarium sheets designated by Fernald as “var. sabulonum" show plants with 2-15, ascend- ing to erect stems; elliptic-obovate leaves; with cleistoga- mous flowers borne in glomerules or rarely even in raceme-like cymes which when fruiting possess pedicels 1-4 mm. long and with capsules ranging from 2-4 mm. in diamc- ter. In addition to the localities cited in the original de- scription by Fernald, specimens with similar characteristics from the following localities have been examined: St. Joseph Co., Ind., Deam 34817 (IND); Concord, Middlesex Co., Mass, Bartram (PH); Manistee Co., Mich., Schumacher | (MICH) ; Atlantic Co., N. J., Fogg 13726 (PENN); Burlington Co., N. J., Fogg 5671 (PENN); Burlington Co., N. J., Fender 926 (PENN) ; Cumberland Co., N. J., Dreisbach (PH); Lancaster Co., Pa., Tanger 4020 (PENN); Nansemond Co., Va., Heller (MIN); Isle of Wight Co., Va., Heller (MIN); Dane Co., Wis., Davis (PH); Walworth Co., Wis., Wadmond 3133 (MIN). Therefore, it seems that such variation occurs sporadically throughout the range of the species and that it does not represent an entity worthy of formal recognition. REPRESENTATIVE SPECIMENS: CANADA: NOVA SCOTIA: Queens Co., Greenfield, Weatherby & Weatherby 7120 (GH, US). ONTARIO: Comté de Carleton, Baie Constance, Marie-Victorin, Rolland- Germain & Rouleau 6296 (FSU, MO, SMU, USF). QUEBEC: Comté de Pontiac, Ile Calumet, Marie-Victorin, Rolland-Germain & Rouleau 58045 (GH). UNITED STATES: MAINE: Cumberland Co., Standish, Fernald & Long 14109 (NY, PH). NEW HAMPSHIRE: Carroll Co., Conway, Johnson 127 (ILL, UC). VERMONT: Rutland Co., Rutland, Eggleston 1047 (GH). MASSACHUSETTS: Barnstable Co., Harwich, Fernald 378 (COLO, DS, DUKE, F, GA, GH, IA, IND, ISC, KANU, MICH, MIN, MO, MSC, NCSC, NCU, NO, NY, OKL, PENN, SMU, TENN, UARK, UC, UMO, US, WIS, WVA); Fernald 379 (DS, DUKE, F, GA, GH, IA, IND, ISC, MICH, MIN, NCSC, NO, NY, OKL, PENN, SMU, TENN, TEX, UARK, UC, UMO, US, WIS, WVA). RHODE ISLAND: Providence Co., Providence, June 1874, Bennett (CU). CONNECTICUT: Windham Co., Thompson, Weatherby 260 Rhodora [Vol. 67 4926 (NCSC). NEW YORK: Queens Co., Peat Bog near Parsons Blv. & Union Turnpike, Monachino 142 (CHRB, CU, DUKE, ISC, OKL, SMU). PENNSYLVANIA: Bucks Co., Buckingham Mountain, ne. of Bucking- ham Valley, Long 66801 (PH). NEW JERSEY: Cape May Co., Anglesea, Fender 3242 (DUKE, PENN). MARYLAND: Washington Co., Warm Spring Ridge, Shreve & Jones 824 (US). DELAWARE: Sussex Co., 1 mi. e. of Lewes, McVaugh 6580 (F, GH, NY). DISTRICT OF COLUMBIA: Near Takoma Park, 27 May 1900, Steele (US). VIRGINIA: Isle of Wight Co., near Franklin, Heller 917 (CU, DS, F, GH, MIN, NY, PENN). NORTH CAROLINA: Hoke Co., 6.5 mi. nw. of Raeford, Fox & Beaman 4657 (NCSC, GH, NY, PH). SOUTH CAROLINA: Marlboro Co., 10.5 mi. n. of Bennettsville, Radford 12686 (NCU). GEORGIA: Burke Co., 1 mi. w. of Shell Bluff, Pyron & McVaugh 2460 (GH). WEST VIRGINIA: Hampshire Co., Loom, Core 3195 (SMU). KENTUCKY: Harlan Co., Pine Mountain, Kearney 220 (F, GH, ISC, MIN, MO, MSC, NY, US). OHIO: Lucas Co., 7 mi. nw. of Maumee, Kriebel 9128 (PUR). MICHIGAN: Cheboygan Co, near the Biological Station, Gleason & Gleason 28 (GH, IND, ISC, NY, SMU, WVA). INDIANA: Jasper Co., 2.4 mi. s. of Demotte, Friesner 17503 (GH, NY, UC, WVA). ILLINOIS: LaSalle Co., Starved Rock, Greenman, Lansing & Dixon 84 (F, GH, IA, NY, UC). WISCONSIN: Dane Co., about 4 mi. sw. of Madison, Shinners 1261 (UC). MINNESOTA: Washington Co., 2 mi. s. of Afton, Moore & Moore 13546 (GH, ISC, MIN, SMU). IOWA: Winneshiek Co., 1 mi. ne. of Hasper Twp., David- son & Thorne 11161 (IA). MISSOURI: Maries Co., 3-4 mi. sw. of Belle, Steyermark 27590 (F). 4. Helianthemum Coulteri S. Wats. Helianthemum arenicola sensu Hemsl., Biol. Centr.-Amer. Bot. 1:47. 1879. Coulter 743 (KEW) but not of Chapman, Fl. S. U.S. 35. 1860. Helianthemum Coulteri S. Wats., Proc. Am. Acad. 17: 323. 1882. Type: Coulter 743 (GH!). Zimapán, Hidalgo, Mexico. Cistus Virginianus Sessé & Moc., Fl. Mex. 130. 1894. Type: MA loaned to F! “in Pratis Sancti Antoni Oppid; jurisdiccionis Ixtlahuacae." Halimium Coulteri (S. Wats.) Gross. Pflanzenreich 14 (IV. 193): 46. 1903. Halimium Berlandieri Briq., Ann. Conser. Jard. Genève 10: 99: 1907. Berlandier 332. (G!) Between Tampico and Real del Monte, Mexi- co. Crocanthemum Berlandieri (Briq.) Janchen, Nat. Pflanzenfam. 2 ed. 21: 305. 1925. Perennial herb or somewhat suffruticose, 4-38 cm. tall with few to many stems arising from a caudex or occasionally a subterranean root- stock. STEMS decumbent to erect, shortly stellate-pubescent or oc- casionally appearing villous with simple and branched trichomes up to 1965] Helianthemum — Wilbur 261 1 mm. long. BASAL LEAVES mostly lacking, when present 6-24 mm. long, 2-7 mm. wide, obovate-spatulate. CAULINE LEAVES: petiole 0.6- 2.0 mm. long; blade 7-45 mm. long, 2-15 mm. wide; obovate-elliptic or merely elliptic, the upper surface dark green to grayish and stellate- pubescent (sometimes sparsely intermixed with simple or stellate- pilose hairs up to 1.2 mm. long) and hoary stellate-tomentose beneath; midvein and secondary veins conspicuous and elevated beneath; base of blade attenuate to cuneate, apex obtuse to acute; margin entire, non-revolute. FLOWERS: dimorphic (chasmogamous and cleistoga- mous), 3-22 in number borne in a congested cyme or loosely arranged in a racemose cyme. Number of cleistogamous flowers varying in each cyme from none (in which case probably not yet formed) to eight. Bracts 1-6 mm. long, 0.2-0.8 mm. wide, linear to lanceolate. Pedicel and calyx stellate-pubescent, sometimes sparsely to densely intermixed with simple hairs up to 1 mm. long, and also with red glandular hairs occasionally present on the calyx. CHASMOGAMOUS FLOWERS: pedicels 2.0-12 mm. long, mostly shorter than calyx. OUTER SEPALS (free portion) 2.0-4.2 mm. long, 0.4-1.0 mm. wide, linear to lanceolate; INNER SEPALS 4.0-6.5 mm. long, 3-5 mm. wide, ovate, acuminate to mucronate. PETALS 3.4-10.0 mm. long, 3.2- 9.0 mm. wide, obovate, “bright yellow." STAMENS 16-30. PISTIL 1.2-3.2 mm. long; ovary 1.0-2.4 mm. long, 0.8-2.0 mm. in diameter, ovoid, glabrous; style almost wanting or up to 0.3 mm. long; stigma 0.8-1.2 mm. wide, capitate. FRUITING PEDICELS 2-12 (17) mm. long. Fruiting calyx 5-12 mm. long, 3-7 mm. in diameter, ovoid. OUTER SEPALS (free portion) 2.5-5.0 mm. long, 0.4-1.0 mm. wide, linear to lanceolate; INNER SEPALS 5-12 mm. long, 3-7 mm. wide, ovate, acuminate to mucronate. CAPSULE 3.5-7.5 mm. long, 3-6 mm. in diameter, ovoid- triquetrous, glabrous, 3-valved; each valve 3-6 mm. wide, ovate, acute, slightly concave. SEEDS 16-56, inequilateral, papillate or reticulate but then on the edges papillate, lacking a separable membrane. CLEISTOGAMOUS FLOWERS: pedicels 1-3 mm. long, usually shorter than calyx. OUTER SEPALS (free portion) 1.2-3.0 mm. long, 0.3-0.7 mm. wide, linear; INNER SEPALS 3.0-4.5 mm. long, 2.3-2.8 mm. wide, ovate, acute. STAMENS 5(8). PISTIL 1.6-2.0 mm. long; ovary ovoid, glabrous. FRUITING PEDICELS 2-6 mm. long. FRUITING CALYX 4-7 mm. long, 3-4 mm. in diameter, ovoid. OUTER SEPALS (free portion) 2.0-4.2 mm. long, 0.3-0.7 mm. wide, linear; INNER SEPALS 4-7 mm. long, 3-4 mm. wide, ovate, acute. CAPSULE 3.0-4.5 mm. long, 2.8-4.2 mm. in diameter, ovoid- triquetrous, glabrous, 3-valved; each valve 2.8-4.2 mm. wide, ovate, acute. SEEDS 8-22, similar to the seeds of the chasmogamous flowers. FLOWERING: March-December. HABITAT: grassy knolls, pine forest openings, dry barren open forested hilltops and moist thickets. DISTRIBUTION: southern Nuevo Léon to 262 Rhodora [Vol. 67 central Veracruz in Mexico and also collected in Guatemala, Honduras and Nicaragua, (Map 12.) The characteristic features of this species are the hoary lower surface of the leaf-blade with conspicuously elevated midvein and secondary veins, the dimorphic flowers borne in a terminal, congested cyme or racemose-cyme, and the lack (or nearly so) of the style on the chasmogamous pistil. It is the only Mexican or Central American species with papillate to reticulate seeds and also the only species from this area lacking a separable membrane. Only three Mexican and Central American species of Helianthemum, H. concolor, H. Coulteri, and sometimes H. glomeratum, have leaves with prominent secondary veins on the lower surface. When Riley (Kew. Bull. 1923: 107.) described H. concolor, he indicated that it was closely related to H. Coulteri. However, the two species are separated geographically and they are very distinct morphologically. Helianthemum concolor can be readily distinguished from H. Coulteri by its leaves which appear bright green on both surfaces and whose lower surface is sparsely stellate-pubes- cent and hence the surface of the epidermis is readily visible. In contrast the densely hoary, stellate lower surface of H. Coulteri completely obscures the epidermis. In H. concolor the cleistogamous flowers are numerous in contrast to eight or fewer/cyme in H. Coulteri, and the inflorescence of H. concolor is densely covered with white but coarse stellate pubescence while that of H. Coulteri is covered with soft stellate pubescence intermixed with simple hairs. The seeds of H. concolor are smooth with a separable membrane while those of H. Coulteri are papillate to reticulate and lack a separable membrane. In the southern part of its range (Guatemala, Honduras and Nicaragua), H. Coulteri has been either confused with H. glomeratum or the specimens left unidentified. Helian- themum glomeratum may be readily distinguished from this species by its glomerate axillary and terminal cleistogamous flowers, by its chasmogamous flowers overtopping the cleis- 1965] Helianthemum — Wilbur 263 togamous ones which are usually solitary at the tip of the branches and branchlets, and by its smooth seeds which possess a thin, separable membrane when moistened. Watson (Proc. Am. Acad. 17: 323. 1882.) in his original description of this species indicated that Hemsley (Biol. Centr.-Amer. Bot. 1: 47. 1879.) had referred it to Chap- man's H. arenicola. However, the latter species is endemic to the Gulf Coast region of the United States. The lower surface of the leaf-blade cf the Gulf Coast species has ob- scure secondary veins and its ovary and capsule are stellate- pubescent in contrast to the prominently veined leaves and glabrous ovary of H. Coulteri. Helianthemum Coulteri varies in height, shape and size of leaves, size and number of flowers, size of fruit, and sculpture of seeds through its range. Additional collections are needed before we can evaluate the significance of these variations. The type of Halimium Berlandieri Briq. (G), which was most kindly loaned by the late Professor Charles Baehni, proves to be Helianthemum Coulteri and neither Halimium patens as questionably indicated by Standley (Contr. U.S. Nat. Herb. 23: 834. 1923.) nor even apparently a close relative of Halimium Pringlei with which Briquet compared it. REPRESENTATIVE SPECIMENS: MEXICO: HIDALGO: between Acaxochitlan & Puebla border on road to Huauchinango, Moore 2852 (GH, UC); District Zacualtipán Zacualtipán Moore 2359 (GH); between Pachuca & Real del Monte, Goodman 3408 (F, ISC). MEXICO: Sierra de las Cruces, Pringle 6672 (F, GH, ISC, MICH, MIN, MO, MSC, ND, UC). NUEVO LEON: Sierra Madre Oriental, Alanaca to Taray, about 15 mi. sw. of Galeana, Mueller & Mueller 980 a (GH, MICH, TEX) ; Cienaguillas to Puerto Santa Ana, about 15 mi. sw. of Galeana, Mueller & Mueller 902 (GH, MICH, TEX). PUEBLA: Honey Station, Pringle 8804 (F, GH, MIN, MO, MSC, PH, UC); about Honey Station, Pringle 10007 (F, GH, MIN, MO, PH, UC). SAN LUIS POTOSI: K. 50 road between San Luis Potosi & Rio V., Layman 4169 (PH); Alvarez, Sierra de Alvarez, Pennell 17832 (PH). VERACRUZ: Below Las Vigas, Perote, Balls 4792 (GH). GUATEMALA: CHIMALTENANGO: Barranco de La Sierra, southeast of Patzün, Standley 61531 (F). QUICHE: mountains nw. of Quiché, Grant 699 (F, GH, MICH). HONDURAS: Morazán: western 264 Rhodora [ Vol. 67 slopes of Cerro de Uyuca, along trail from Las Flores toward Tatum- bla, Standley 22712 (F, UC); Piedra Herrada, lower slopes of Cerro de Uyuca, Standley 11911 (F); open slopes near Haya Grande Drainage of the Rio Yeguare, at about Long. 87° W. & Lat. 14^N., Williams & Molina R. 13278 (F, GH); Mt. Uyuca, drainage of the Rio Yeguare, at about Long. 87^ W. & Lat. 14°N., Glassman 1930 (F, ILL, MIN, OKL). NICARAGUA: southwest of Jinotega, along road to Cantera & Los Pinos, Standley 10076 (F). 5. Helianthemum arenicola Chapm. Helianthemum arenicola Chapm., Fl. S. U.S. 35. 1860: Lectotype: Chapman (US #6889!). Isolectotypes: (GH, Mo #788712!). “West Florida,” Apalachicola. Helianthemum canadense (L.) Michx. var. obtusum Wood, Class- Book 246. 1861. Halimium arenicola (Chapm.) Gross., Pflanzenreich 14 (IV. 193): 49. 1908. Crocanthemum arenicola (Chapm.) Barnh. in Small, Man. SE. Fl. 879. 1933. Perennial herb 5-15(20) cm. tall with several stems arising from ascending to erect subterranean woody portion of an old stem. STEMS often partly buried in the loose sand, spreading to erect, covered with a depressed, canescent stellate pubescence. CAULINE LEAVES: petiole 0.5-2(2.4) mm. long; blade (4)8-17(28) mm. long, (1)2-4(10) mm. wide; oblanceolate to oblong to lanceolate, sometimes the lowermost spatulate; cinerous and densely stellate-pubescent, hoary and stellate- tomentose beneath; midvein elevated but the secondary veins obscure beneath; base of blade cuneate-obtuse, apex obtuse to acute. FLOWERS dimorphic (chasmogamous and cleistogamous), 2-10 in typically terminal or occasionally lateral umbellate clusters or rarely the lateral cluster reduced to a single flower. Bracts 1-3.5 mm. long, 0.3-0.6 mm. wide, linear. Pedicel and calyx stellate-canescent. CHASMOGAMOUS FLOWERS: pedicels (2.5)5-10(16) mm. long. OUTER SEPALS (free portion) 2-4 mm. long, about 0.5 mm. wide, linear; INNER SEPALS 4-8 mm. long, 3.5-4.8 mm. wide, ovate or ovate-elliptie, acute. COROLLA yellow, petals 8-9.5 mm. long, 7.5-9 mm. wide. STAMENS 17-29. PISTIL 1.2-1.5 mm. long; ovary 1-1.2 mm. long, 0.7-1.1 mm. in diameter, ovoid, stellate-pubescent on the upper half, style 0.1-0.3 mm. long; stigma c. 0.6 mm. wide, capitate. FRUITING CALYX 5.6-8(10) mm. long, 3.6-4(5.2) mm. in diameter, ovoid-ellipsoid. CAPSULE 3.6-5.4 mm. long, 2.4-8.6 mm. in diameter, ovoid-ellipsoid, greenish, stellate- pubescent near the top, glabrous beneath, 3-valved; each valve 1.8-2.8 mm. wide, elliptic, acute, slightly concave. SEEDS 14-17, dark brown, irregular, pebbled to somewhat papillate, without a separable 1965] Helianthemum — Wilbur 265 membrane; cohering together in a globular or ellipsoidal mass by interwoven funiculi. CLEISTOGAMOUS FLOWERS: pedicels 1-2 mm. long. OUTER SEPALS (free portion) 1-2 mm. long, c. 0.4 mm. wide, linear; INNER SEPALS 3.6-5.6 mm. long, 2.2-2.8 mm. wide, ovate, acute. STAMENS 4-6(8). PISTIL 1.1-1.4 mm. long; ovary ovoid, stellate-pubescent on the upper half. FRUITING CALYX 4-5.6 mm. long, 3-3.7 mm. in diameter, ovoid- ellipsoid. CAPSULE 3-4.6 mm. long, 2:4-3.4 mm. in diameter, ovoid, greenish, stellate-pubescent near the top, the lower part glabrous, 3-valved; each valve 2.2-2.7 mm. wide, ovate, acute, slightly concave. SEEDS (6)9-14, similar to those of the chasmogamous flowers. FLOWERING: March-April. HABITAT: open, sandy scrub barrens and sand-dunes. DISTRIBUTION: near or along the beach from western Florida to Mississippi. (Map 7.) The stellate-pubescent ovary and capsule and the typically terminal (or sometimes lateral) umbellate clusters composed of dimorphic flowers readily distinguish this species from all other North American species of Helianthemum. Helianthemum arenicola is most closely related to H. Nashii; the two species possess the following characteristics in common: (1) the ovary and capsule stellate-pubescent ; (2) the seeds dark brown, mostly coherent, pebbled to some- what papillate and lacking a separable membrane. In 1861 when Wood described this species as H. canadense var. obtusum, he was unaware of the previous description of Chapman’s H. arenicola, and indicated that “It may prove distinct." In 1871 he accepted Chapman’s epithet, recog- nizing its specific distinction and cited his variety as a synonym. During this study four collections* were examined that seemed possible hybrids between H. arenicola and H. corym- bosum. The flowers of these specimens had calyces bearing abundant simple hairs (a condition unknown in typical H. arenicola but characteristic of H. corymbosum). The chas- mogamous outer sepals of these collections were relatively broad approaching those of H. corymbosum instead of linear as they are in H. arenicola. The ovaries and capsules were more nearly glabrous instead of stellate-pubescent and their cleistogamous flowers were apparently more numerous than 266 Rhodora [Vol. 67 the few that characterize H. arenicola. One herbarium sheet [Biltmore Herbarium 5741 (NY)] has specimens of both the putative hybrid and H. corymbosum while Godfrey’s collection from Franklin County has both the possible hybrid and the typical H. arenicola represented among its numerous sheets. It must be admitted that evidence of hybridization or introgression is slight and perhaps this variation is normal within the species. Field observations can do much in resolving this question. The variation appears quite comparable to the phase of H. Nashii that has been called H. thyrsoideum. *FLORIDA: Franklin Co.: very abundant, evergreen scrub-oak sand pine barrens, 1.5 miles e. of St. Theresa, 30 April 1960, Godfrey 59485b (DUKE); Pine palmetto woods, % mi. s. of Panacea, 20 March 1955, Adams 4 (GA). Dry sands, near the Coast, Florida, Biltmore Herbarium 5741 (NY). Drifting sands, near the coast of West Florida, March-April, Biltmore Herbarium 4028 (GH). REPRESENTATIVE SPECIMENS: FLORIDA: Escambia Co., Pensacola, 24 Apr. 1898, Baker (NCU, NY); Franklin Co., near the beach, Alligator Point, Godfrey 53202 (DUKE, FSU, GA, IA, NCSC, NCU, NY, USF, VDB) ; near Appalachicola, Curtiss 226 (F, FLAS, GA, GH, KANU, MIN, MO, NY, OKL, PH, SMU, UARK); Gulf Co., near Port Saint Joe, Perkins & Hall 2440 (cU); Okaloosa Co., % mi. w. of Mary Esther, Godfrey 56668 (FSU, IA, UC, USF). ALABAMA: Baldwin Co.; Fort Morgan, Tracy 7787 (F, GH, ISC, MIN, MO, MSC, NY, OS, PENN, US); Mobile Co., Mobile, 20 March 1883, Mohr (GH). MISSISSIPPI: Harrison Co.; near Handsboro, Channell 1301 (vpB); Jackson Co., Horn Island, Godfrey & Channell 53713 (FSU, GH, NY). 6. Helianthemum Nashii Britt. Helianthemum Nashii Britt., Bull. Torrey Club 22: 147. 1895. [as H. Nashi] Lectotype: Nash 815 (NY!). Isolectotypes: (CU, F, GH, MIN, MO, MSC, UC, US!). Vicinity of Eustis, Lake Co., Florida. Halimium Nashii (Britt.) Gross., Pflanzenreich 14 (IV. 193): 49. 1903. Helianthemum thyrsoideum Barnh. in Small, Fl. SE. U.S. 797, 1335. 1903. Type: Barnhart 2740 (NY!). Sutherland, Florida. Crocanthemum thyrsoideum (Barnh.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 307. 1925. Crocanthemum Nashii (Britt.) Barnh. in Small, Man. SE. Fl. 879. 1933. Perennial herb (14)25-30(41) cm. tall with few to numerous decumbent to ascending or more commonly erect, depressed stellate- 1965] Helianthemum — Wilbur 267 tomentulose stems arising from a caudex or from a subterranean horizontal rootstock. CAULINE LEAVES: petioles (0.6)1-2(3) mm. long; blade (5)23-27(38) mm. long, (0.9)3.8-5.7(8.4) mm. wide; oblance- elliptic on the lower half of the stem and becoming elliptic-lanceolate near the apex; densely stellate-canescent above and hoary and stellate- tomentose beneath; midvein elevated and the secondary veins but very slightly to moderately elevated beneath; base of blade cuneate, apex obtuse to acute; margins entire, slightly revolute. INFLORESCENCE: an elongate, leafy thyrse. FLOWERS dimorphic (chasmogamous and cleistogamous) in each cymule. Bracts 1-3.2 mm. long, 0.2-0.6 mm. wide, linear-lanceolate. Pedicels and calyx stellate-canescent or stel- late-canescent intermixed with abundant hirsute trichomes about 0.7-1.5 mm. long. CHASMOGAMOUS FLOWERS: pedicels (2)3-8(10) mm. long. OUTER SEPALS (free portion) 1-2.5(3) mm. long, 0.2-0.4 mm. wide, linear; INNER SEPALS 3.5-5.0 mm. long, 2.7-3.7 mm. wide, obliquely ovate- elliptic. COROLLA yellow, petals 5-9.4 mm. long, 3-6.6 mm. wide, broadly cuneate. STAMENS 12-18. PISTIL 1-1.8 mm. long; ovary 0.8-1.5 mm. long, 0.8-1.3 mm. in diameter, ovoid, stellate-pubescent; style 0.2-0.3 mm. long; stigma 0.4-0.7 mm. wide, capitate, FRUITING PEDICELS 3.6-7.2(12) mm. long. FRUITING CALYX 4-5.5 mm. long, 3.8- 4.6 mm. in diameter, ovoid to nearly globose. CAPSULE 3-4.8 mm. long, 2.4-3.8 mm. in diameter, broadly ovoid, elliptic in cross-section, apex blunt, stellate-pubescent on the upper half, gradually becoming glabrous on the lower half, 2-valved; each valve 2.4-3.8 mm. wide, broadly ovate, obtuse, concave. SEEDS 6-10, dark brown, irregular, cohering together into a globular mass by the interwoven funiculi, pebbled to somewhat papillate and lacking a separable membrane. CLEISTOGAMOUS FLOWERS: pedicels 0.5-2.4 mm. long. OUTER SEPALS (free portion) 0.5-1.2 mm. long, c. 0.5 mm. wide, linear; INNER SEPALS 0.5-3.4 mm. long, 1-2.3 mm. wide, obliquely ovate-elliptic. STAMENS 5(7). OVARY ovcid, stellate-pubescent. FRUITING PEDICELS 1.5-4.5 mm. long. FRUITING CALYX 3-3.6 mm. long, 2.4-3.6 mm. in diameter, ovoid. CAPSULE 3-4 mm. long, 2.4-3.5 mm. in diameter, otherwise similar to the chasmogamous capsule. SEEDS 5-8, similar to those of the chasmogamous flowers. FLOWERING: March-June. HABITAT: sand dunes and dry sandy pine woods and scrub. DISTRIBUTION: peninsular Florida. (Map 7.) The elongate, leafy, thyrsoid inflorescence and the bi- carpellate, stellate-pubescent ovary and capsule make H. Nashii most distinctive. This species and H. arenicola are the only New World 268 Rhodora [Vol. 67 species of Helianthemum with stellate-pubescent ovaries and capsules. It appears that H. Nashii is the only species of Helianthemum with a typically 2-valved capsule (or in any event, it is certainly the only North American species). Apparently all other species of Cistaceae characteristically have a 3 or 5-10-valved capsule. Barnhart (in Small, Fl. SE. U.S. 797, 1335. 1903.) de- scribed H. thyrsoideum as a new species. In 1933 (in Small, Man. SE. Fl. 880.) he transferred it to the segregate genus Crocanthemum overlooking the fact that Janchen had made the transfer eight years earlier. The species was there described as follows: “6. C. thyrsoideum Barnhart. Plant similar to C. Nashii in habit, but the inflorescence less widely branched, and the sepals densely hirsute." We have seen perhaps a dozen specimens fitting these characteristics including the type of Barnhart’s species. The only difference noted between H. thyrsoideum and H. Nashii was the presence of the hirsute hairs on the calyx of the former species. The variant with a hirsute calyx is known from Orange, Pinellas, Polk and Seminole Counties where the form with short-pubescent calyces is also to be found. There is no distinct geographic or ecologic range of these forms and both have been collected within the same stand. Therefore Barnhart's species, H. thyrsoideum, is here treated as a synonym of H. Nashii. There is a similar variant of the closely related H. arenicola, but in that species there seems to be some slight evidence that the hirsute calyces might be due to introgression from H. corymbosum. In H. arenicola, that possibly was introgressed by H. corym- bosum, the ovary or capsules were less pubescent and the bracts seemed to vary somewhat in the direction of H. corymbosum. REPRESENTATIVE SPECIMENS: FLORIDA: Brevard Co., west of Eau Galle, Hood 4305 (FLAS); Collier Co., Marco Island, 15 April 1954, West, Arnold & Cooley (FLAS); Duval Co., Jacksonville and vicinity, 1875, Hogg (NY); Hernando Co. at Weekiwachee Springs, Cooley & Eaton 6602 (FSU, USF); Highlands Co., Avon Park, Small 12662 (USF); Lake Co., Vicinity of Eustis, Nash 815 (CU, F, GH, MIN, 1965] Helianthemum — Wilbur 269 MO, MSC, NY, US); Levy Co., Cedar Key, Godfrey 56603 (IA, FSU, UC, USF, VDB); Manatee Co., Bradentown, 10 May 1936, Cuthbert (FLAS); Martin Co., near Darkeytown, 31 Jan. 1917, Atwood (CU); Orange Co., Windermere, Blanton 6506 (US); Osceola Co., Kissimmee, Singel- tary 218 (DUKE, NCSC); Palm Beach Co., just n. of Delray, 18 March 1945, Fox (NCSC, WVA); Pinellas Co., Gulfport, Schallert 3995 (SMU); Polk Co., Dear Lake scrub, McFarlin 5177 (MICH); Putnam Co., south end of Reserve Welaka, 19 June 1941, Laessle (FLAS); Seminole Co., at Sanford, Godfrey, Eaton & Ray 7547 (USF). 7. Helianthemum Bicknellii Fern. Helianthemum canadense (L.) Michx. var. Walkerae Evans, Bot. Gaz. 15: 211. 1890. Lectotype: Walker (US!) Douglas Co., Colorado. Helianthemum majus in sense of Bickn., Bull. Torrey Club 21: 259. 1894 but not of the basionym Lechea major L., Sp. Pl. 90. 1753, whose type was found to be conspecific with Cistus canadensis L.; nor Helianthemum majus (L.) BSP., Prel. Cat. N.Y. 6, 1888, a nomenclatural transfer (covering all northeastern entities) based on page priority. Halimium majus in sense of Gross., Pflanzenreich 14 (IV. 193) : 50. 1903. Crocanthemum majus Britt. in Britt. & Brown, Ill. Fl. ed. 2. 2: 540. 1913. Helianthemum Bicknellii Fern., Rhodora 21: 36. 1919. Type: F'ernald, 7 August 1908 (GH!). Bangor, Maine. Heteromeris major in sense of Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 169. 1921. (No basionym cited.) Crocanthemum Bicknellii (Fern.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 307. 1925. Helianthemum Walkerae (Evans) Lyon, [as Walkedae and without basionym or even indication that a new combination was being made], Am. Midl. Nat. 12: 278. 1927. Perennial herb, (12)20-46(67) cm. tall with many clustered stems arising from a multicipital caudex. Stems ascending to erect, stellate-pubescent, simple or sparsely branched at first anthesis; branches moderately ascending to appressed, restricted to the upper third of the stem, later in the growing season becoming densely leafy and crowded with cleistogamous flowers. CAULINE LEAVES: petiole 1-3(4) mm. long; blade (8)18-32(40) mm. long, (2)4-7(10.5) mm. wide; oblance-elliptic to elliptic, green and stellate-pubescent above, hoary stellate-tomentose beneath; base of blade cuneate, apex suba- cute to acute; margin entire, the upper cauline leaves slightly revolute; midvein and secondary veins prominent beneath. FLOWERS dimorphic (chasmogamous and cleistogamous); maturing at different times during the growing season and upon different positions on the plant. At first anthesis with (2)6-10(18) chasmogamous flowers terminating 270 Rhodora [Vol. 67 the stem and the uppermost branches in a cymose cluster and these rarely surpassed by the lateral branches at maturity. Pedicel and calyx of the chasmogamous flowers densely covered with stellate pubescence and with apparently simple (although mostly unequally stellate), pilose hairs. Bracts 3-7 mm. long, 0.5-1 mm. wide, lanceolate. Later in the growing season the lateral, ascending to depressed, leafy branches becoming crowded with sessile to subsessile cleistogamous flowers in axillary and terminal glomerules, Pedicels and calyces of the cleistogamous flowers stellate-pubescent. CHASMOGAMOUS FLOWERS: pedicels (1.2)3-8(12) mm. long, mostly shorter than calyx. OUTER SEPALS (free portion) (2.4)3.5-4.5(8) mm. long, nearly as long as the inner ones, 0.4-1.0 mm. wide, linear, acute; INNER SEPALS (3.4)5-7(8) mm. long, 2.4-4.0 mm. wide, ovate-elliptic, acute. PETALS 8-12 mm. long, 5-10 mm. wide, obovate. STAMENS (18) 26- 38. PISTIL 1.5-2.6 mm. long; ovary 1.0-1.6 mm. long, 0.8-1.4 mm. in diameter, ovoid, glabrous; style 0.4-0.8 mm. long; stigma 0.6-0.8 mm. wide, capitate. FRUITING CALYX (4.5)5.2-7.2(9.6) mm. long, (3.4)4.0- 5.5(6.5) mm. in diameter, ovoid. CAPSULE (2.4)3.5-5.0(5.6) mm. long, (2.2)3.0-4.0(5.0) mm. in diameter, ovoid-triquetrous, sometimes abruptly short-pointed; 3-valved (but a few capsules seen with 4 or 5 valves) ; each valve (2.2)3.0-4.0(4.4) mm. wide, ovate-elliptic, acute, slightly concave. SEEDS 12-26, ovoid to inequilateral, brown-spotted and indistinctly reticulate with very slight relief. CLEISTOGAMOUS FLOWERS: sessile to subsessile. FRUITING PEDICELS 0.4-0.7(1.3) mm. long. FRUITING CALYX 1.7-2.5(3.0) mm. long, 1.6- 2.2(2.5) mm. in diameter, ovoid. OUTER SEPALS (free portion) (0.3)0.6- 1.2(1.8) mm. long, 0.2-0.3 mm. wide, linear; INNER SEPALS 1.7-2.5 (3.0) mm. long, 1.5-2.3 mm. wide, ovate-elliptic, acute. STAMENS 3(5). CAPSULE 1.5-2.2(2.8) mm. long, 1.2-2.0(2.4) mm. in diameter, ovoid- triquetrous, apiculate, glabrous, 3-valved (few capsules seen with 4- valves); each valve 1.2-2.0(2.4) mm. wide, ovate-elliptic, acute, slight- ly concave, mostly dehiscing for about two-thirds distance towards base and spreading at maturity. SEEDS 1-2(3), globose( when one seed per capsule), half globose (when two), or three-sided with the outer convex (when three per capsule), obscurely reticulate with low relief and reddish brown. FLOWERING: chasmogamous flowers, June-July; cleistog- amous flowers, July-September. HABITAT: sandy and rocky shores, sand dunes or barrens, open dry sandy fields or prairies, and open mountain tops. DISTRIBUTION: Maine to northern Georgia and westward into Minnesota, eastern Wyoming, and Colorado. (Map 6). H. Bicknellii has a wider distribution and has been more 1965] Helianthemum — Wilbur 271 extensively collected than any North American Helianthe- mum. This species is rather uniform throughout its range excepting perhaps some specimens from Colorado which show somewhat wider cauline leaves on the lower half of the stem. However, this species has a remarkably different appearance from first anthesis, when the chasmogamous flowers terminate the stem and whose fruits drop as soon as they mature, to the end of the growing season when the branches are crowded with the mature cleistogamous fruits. This extreme variation causes difficulty in identification. The species with which H. Bicknellii is most easily con- fused is H. propinquum and they were in fact judged con- specific by Blake (Rhodora 20: 50. 1918.). Fernald (Rhodora 21: 36-37. 1919.), after considerable field study was finally convinced of their specific differences. Helianthemum pro- pinquum shares with H. Bicknellii the following character- istics: (1) chasmogamous flowers in a terminal cymose cluster; (2) cleistogamous flowers sessile to subsessile and borne on the branches in axillary and terminal glomerules ; (3) cleistogamous capsules 1.5-2.0(2.4) mm. in diameter each with 1-2(3) reticulate seeds. . Despite the close similarity between the two, they may be readily distinguished as follows: H. Bicknellii H. propinquum 1. Chasmogamous outer sepals 1. Chasmogamous outer sepals fused with the inner sepals fused for up to two thirds near the base and its free of the length of inner sepals portion more than half the and its free portion half of length of the inner sepals. the length or less of the inner sepals. 2. Base of leaf-blade cuneate. 2. Base of leaf-blade attenuate. 3. Usually stems arising from 3. Stems arising from subterran- a multicipital caudex. ean rootstocks. 4. Fruiting cleistogamous outer 4. Fruiting cleistogamous outer sepals more than 3-times long- sepals about twice longer than er than wide. wide (knob-like and attached to the middle edge of the inner ones). 5. Cleistogamous capsule ovoid- 5. Cleistogamous capsule ovoid triquetrous. or nearly so. 272 Rhodora [Vol. 67 There has been much confusion concerning the nomencla- ture of this species and it therefore seems desirable to dis- cuss it briefly. Prior to 1894 the taxon now known as H. Bicknellu had been erroneously equated with H. canadense or treated as a named or unnamed variant. Gray (Man. Bot. N. U.S. 47-48. 1848.) treated plants belonging to this spe- cies under H. canadense as follows: “A variety is more hoary, and with a stronger tendency to multiply the minute clustered flowers.” Evans (Bot. Gaz. 15: 211. 1890.) de- scribed this taxon as H. canadense var. Walkerae basing his description upon specimens from Colorado. Bicknell (Bull. Torrey Club 21: 257-260. 1894.), studying the genus in the vicinity of New York, recognized two distinct species. The first was H. canadense and the second and the more common species of the two, he referred to “H. majus (L.)" stating: “As a matter of fact, however, the apetalous state of true H. Canadense Michx., as here defined, does not closely resem- ble a Lechea, while the corresponding state of the newly recognized species is, in a marked degree, suggestive of a plant of that genus. ... It would appear, therefore, not at all improbable that the Helianthemum described by Linnaeus as Lechea major was in reality the plant here in view, and that his Cistus Canadensis was after all a different plant. An examination of the type specimens in the Linnaean her- barium can alone settle the question, and may prove that the plant here called majus requires a different name; but for the purpose of more easily handling the plants in this note, I adopt provisionally the name H. majus (L.), for the newly recognized plant... .” Therefore from 1894 to 1918 this species was known as Helianthemum majus or Crocanthemum majus. Blake’s (Rhodora 20: 49+50. 1918.) examination of the Linnean specimens demonstrated that Lechea major L. was con- specific with Cistus canadensis as had already been suggested by Elliott, Torrey and Gray, and Hooker. Blake concluded that the plant to which Bicknell applied the name was con- specific with H. propinquum. Fernald (Rhodora 21: 36+ 37. 1919.) indicated that Helianthemum majus in the sense of 1965] Helianthemum — Wilbur 273 Bicknell (not H. majus (L.) BSP, a synonym of H. cana- dense maintained under the American Code due to page priority) was a “good” species amply distinct from H. pro- pinquum and he provided the new name Helianthemum Bicknellii. Specimens from northeastern Sonora reported by White (Lloydia 11: 287. 1948.) as this species (under the name H. majus) are H. Pringlei. REPRESENTATIVE SPECIMENS: CANADA: MANITOBA: Brokenhead, 12 mi. north of Beausejour, Scoggan 11550 (MIN). ON- TARIO: Essex Co., Windsor, Macoun 34070 (GH, MO, NY, US). UNITED STATES: MAINE: Penobscot Co., Orono, Fernald 336 (A, GH, MIN, MO, MSC, NY, PH, US, WIS). NEW HAMPSHIRE: Cheshire Co., Walpole, Fernald 88 (GH). VERMONT: Bennington Co., Pownal, 10 Oct. 1857, Ames (MICH). MASSACHUSETTS: Barnstable Co., Harwich, Fernald & Long 382 (DS, DUKE, F, GA, GH, IA, IND, ISC, KANU, MICH, MIN, MO, NO, NY, OKL, PENN, SMU, TENN, TEX, UARK, UC, UMO, US, WIS). RHODE ISLAND: Washington Co., Prudence Island, Narragansett Bay, Mearns 384 (US). CONNECTICUT: New London Co., behind dunes, Bluff Point, Groton, Travis 2022 (PENN). NEW YORK: Cayuga Co., sandy crest of Salmon Creek Ravine, Genoa, Wiegand 10304 (CU, GH). PENNSYL- VANIA: Lancaster Co., about the mouth of the Tucquan, 24 July 1901, Heller (cU, F, GH, IND, MO, US). NEW JERSEY: Passaic Co., Wanaque, Mackenzie 2710 (MO, NY). DELAWARE: New Castle Co., Serpentine e. of Mount Cuba, Pennell 1499 (PENN). MARYLAND: Cecil Co., drainage of Octoraro Creek, e. of Rocksprings, Long 28484 (GH, PH). VIRGINIA: Arlington Co., Tarrison, Steele 391 (F, NY). NORTH CAROLINA: Ashe Co., near top of Nigger Mountain, Fox & Godfrey 3347 (FSU, GA, GH, NCSC, US) ; Buncombe Co., summit of Cedar Cliff Mountain, Biltmore Herb. 5740b (F, GH, MO, NY, US). GEORGIA: Towns Co., on summit at Enotah Bald Mountain, Brasstown Bald, Pyron 521 (GA). OHIO: Wayne Co., Shreve, Duvel 804 (NCU, OS). KENTUCKY: Meade Co., with- out exact locality, 1842, Short (PH). TENNESSEE: Blount Co., edge of “bald”, Gregory Bald, Wilson 1941 (TENN); Knox Co., Knoxville, Moldenke 10705 (NY). MICHIGAN: Menominee Co., Menominee, Grassl 2886 (MICH, NY). INDIANA: Lagrange Co., 3 mi. e. of Mongo, Yuncker & Welch 10782 (COLO, GH, SMU). WISCONSIN: Greene Co., 5 mi. ese, of Monticello, Iltis & Greene 6720 (MIN, WIS). ILLINOIS: Lake Co., n. of Waukegan and e. of the glacial Glenwood Ridge, Gates 2752 (F, ILL, MICH). MINNESOTA: Hubbard Co., roadside to Park Rapids, Moyle 759 (CU, GH, IND, NY, US). IOWA: Guthrie Co., 12 mi. n. of Adair, Fay 5310 (KANU, UC, US). MISSOURI: Barry Co.: Eagle Rock, Bush 197 (GH, MO, NCU, NY, US). ARKANSAS: Hempstead Co., Washington City, collector unknown (MO). SOUTH DAKOTA: Custer Co., Black Hills, 274 Rhodora [Vol. 67 Rydberg 542 (NY, US). NEBRASKA: Brown Co., Long Pine, Clements 2940 (CU, ISC, NY, US). COLORADO: E] Paso Co., Black Forest, Living- ston 640 (DUKE). 8. Helianthemum propinquum Bickn. Helianthemum propinquum Bickn., in Britton Man. Fl. N. U.S. ed. 2. 1069. 1905. Type: Bicknell, August 1908 (Nv!). Valley Stream, Long Island, New York. Helianthemum georgianum sensu Britt. in Britt. & Brown, Ill. Fl. ed. 2. 2: 540. 1913. Crocanthemum propinquum (Bickn.) Bickn., Bull. Torrey Club 40: 615. 1918. Perennial herb, 8-27(35) cm. tall with few to many stems arising usually singly at intervals (or sometimes few together) from a hori- zontal, subterranean rootstock. STEMS slender, ascending to erect, stellate-tomentose, somewhat reddish when young, mostly simple at first anthesis; branches ascending, mostly restricted to the upper half of the stem, densely leafy late in the season and becoming congested with cleistogamous flowers. CAULINE LEAVES: petioles (1)2-4(5) mm. long; blade (5)12-28(37) mm. long, (1.5)3-6(10) mm. wide; oblanceo- late to narrowly oblance-elliptic; dark green and stellate-pubescent above, hoary stellate-tomentose beneath; midvein and secondary veins prominent beneath; base of blade attenuate, apex obtuse to acute; margin entire, sometimes the upper leaves slightly revolute. FLOWERS dimorphic (chasmogamous and cleistogamous) ; maturing at different times during the growing season and upon different positions on the plant. At first anthesis with 2-6 chasmogamous flowers terminating the stem in a cymose cluster and these surpassed by the lateral, as- cending, leafy branches densely crowded with sessile to subsessile cleistogamous flowers in axillary and terminal glomerules. Bracts sub- tending chasmogamous flowers 1.5-3.5 mm. long, 0.2-0.4 mm. wide, lanceolate. Pedicel and calyx covered with appressed, densely matted, elongate stellate-pubescence. CHASMOGAMOUS FLOWERS: pedicels (2)8-14(22) mm. long, mostly longer than calyx. OUTER SEPALS: fused with 1/2-2/3 of the edge of the inner sepals, free portion (0.7) 1.5-3.0(4.0) mm. long, 1/3-1/2 the length of the inner sepals, 0.4-0.9 mm. wide, linear; INNER SEPALS (4)5.0-7.5(8) mm. long, 2.3-4.5 mm. wide, ovate, acute. PETALS 7.5- 10.0(13.5) mm. long, 6-12 mm. wide, obovate. STAMENS 24-30, “anthers orange" (fide Fernald 17160). PISTIL 2.0-2.6 mm. long; ovary 1.5-1.8 mm. long, 1.0-1.4 mm. in diameter, ovoid, glabrous; style 0.5-0.8 mm. long; stigma 0.6-0.8 mm. wide, capitate. FRUITING CALYX (5) 6-8 (9.7) mm. long, 3.5-7.0 mm. in diameter, ovoid, CAPSULE 3.7-5.8 mm. long, 3-4 mm. in diameter, ovoid-triquetrous, glabrous, 3-valved (but a few capsules seen with 4 or even 5 valves); each valve (1.2)2.2-3.7 mm. wide, ovate-elliptic; capsules usually overtopped by the upper branches 1965] Helianthemum — Wilbur 275 and mostly falling promptly after maturity. SEEDS 12-15, ovoid to inequilateral, brown-spotted, reticulate or sometimes somewhat papil- late at the edges only. CLEISTOGAMOUS FLOWERS: sessile to subsessile. FRUITING PEDICELS up to 1.5 mm. long. FRUITING CALYX 2.0-2.5(2.7) mm. long, 1.5-2.2 (2.5) mm. in diameter, ovoid to subglobose. OUTER SEPALS (free portion) 0.2-0.5 mm. long, 0.2-0.3 mm. wide, rudimentary, knob-like, attached at the middle edge of the inner sepals; INNER SEPALS 2.0-2.5(2.7) mm. long, 1.5-2.2 mm. wide, ovate, acute, somewhat lustrous. STAMENS 3-4. CAPSULE 1.5-2.2 mm. long, 1.3-2.0 mm. in diameter, ovoid and usually rounded in cross-section (rarely weakly ovoid-triquetrous), glabrous, 3-valved; each valve 1.1-1.8 mm. wide, ovate-elliptic, subacute to acute, never spreading at maturity. SEEDS 1-2(3), globose (when one seed per capsule), half globose (when two), and three-sided with the outer side convex (in three-seeded capsule), brown-spotted and reticulate. FLOWERING: Chasmogamous flowers, late May-early July; cleistogamous flowers, July-September. HABITAT: fields, sandy places and dry, open woods. DISTRIBUTION: Eastern Massachusetts southward through eastern Pennsylvania to northern Virginia; also collected from the mountains of North Carolina, Tennessee and northern Georgia. (Map 5.) Previously, H. propinquum was believed to occur only from eastern Massachusetts southward to the District of Columbia. However, this species also occurs in the mountains of western North Carolina, eastern Tennessee and northern Georgia where it has usually been misidenti- fied as H. Bicknellii. The species with which H. propinquum is most likely to be confused is H. Bicknellii, as was done by Blake (Rhodora 20: 49+ 50. 1918.). The characteristics which these species have in common, as well as those which serve to distinguish them from each other, are indicated under the discussion of H. Bicknellii. Prior to its original description in 1905, H. propinquum was referred to as a stunted form of H. canadense. The clustered, chasmogamous flowers with calyces covered with appressed stellate pubescence, the 1-2(3)-seeded, cleistog- amous capsule, and the stem which arises from subterranean rootstock, readily distinguish this species from both H. canadense and also from H. dumosum. However, H. propin- 276 Rhodora [Vol. 67 quum resembles H. canadense in having somewhat lustrous cleistogamous fruiting calyces and in having outer sepals which are rudimentary or knob-like and attached to the middle edge of the inner sepals. Britton (in Britt. & Brown, Ill. Fl. ed. 2. 2: 540. 1913.) equated this species with H. georgianum which is morpho- logically quite unlike it and which has a very different range. REPRESENTATIVE SPECIMENS: MASSACHUSETTS: Barnstable Co. Harwich, Fernald 380 (DS, DUKE, F, FLAS, FSU, GA, GH, IA, IND, ISC, KANU, MICH, MIN, MO, MSC, NCSC, NCU, NO, NY, OKL, PENN, SMU, TENN, UARK, UC, UMO, US, WIS, WVA); Brewster, Fernald 381 (DS, DUKE, F, FLAS, GA, GH, IA, IND, ISC, KANU, MICH, MIN, MSC, NCSC, NCU, NO, NY, OKL, PENN, SMU, TENN, TEX, UARK, UC, UMO, US, WIS, WVA). CONNECTICUT: Hartford Co., Southington, Bissell 88 (GH). NEW YORK: Nassau Co., Valley Stream, August 1903, Bicknell (NY). PENNSYLVANIA: Bucks Co., Turkey Hill Long 21299 (PH). NEW JERSEY: Morris Co., Succasunna, Mackenzie 4104 (MO, US). DELAWARE: Sussex Co., e. of Ellendale, Pennell 12884 (MINN, US). MARYLAND: Cecil Co., 2 mi. w. of Elkton, Randolph & Randolph 128 (cv, GH). DISTRICT OF COLUMBIA: Pinehurst, 3 June 1919, Steele (F, GH, NY). VIRGINIA: Arlington Co., Arlington Forest, Balls 7804 (US). NORTH CAROLINA: Ashe Co., Bald, Nigger Mountain w. of West Jefferson, Radford 41133 (NCU); Avery Co., 0.6 mi. ne. of Avery-Mitchell Co. line on US 19E., Ahles & Duke 43433 (NCU); Buncombe Co., Biltmore, Biltmore Herbarium 1217 (CU, MIN, US) ; Henderson Co., Flat Rock, Schallert 8645 (DUKE); Stokes Co., 1.5 mi. n. of Belews Creek, Rad- ford 34511 (NCU); Transylvania Co., behind Pisgah Inn, Mt. Pisgah, Oosting 1779 (DUKE, FLAS, PH). TENNESSEE: Coffee Co., 10 mi. ne. of Tullahoma, Hardin 15761 (TENN); Cumberland Co., near Crossville, Rhoades 19 (cU); Monroe Co., near Mt. Vernon, Ford & Russell 2097 (SMU, TENN). GEORGIA: without locality “Northern Georgia, chiefly of the Mts. & upper Country, 1878" Vassey (PH). 9. Helianthemum Greenei Robins. Helianthemum occidentale Greene, Bull. Calif. Acad. Sci. 2: 144. 1886. Type: Greene, July & August 1886 (ND!), isotype (NY!) “on dry summit in the central part of the Island of Santa Cruz, Santa Barbara Co., California; not Nym., Consp. Fl. Eur. 72. 1878. Helianthemum Greenei Robins., Syn. Fl. N. Am. 1(1): 191. 1895. (nom. nov. for Greene's later homonym). Halimium occidentale (Greene) Gross., Pflanzenreich 14 (IV. 193): 35. 1903. Crocanthemum occidentale (Greene) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. 1965] Helianthemum — Wilbur 277 Suffruticose perennial, often somewhat spartioid, 14-30 cm. tall with few stems arising from a woody caudex. STEMS: ascending to erect, moderately branched, and varying from sparsely to densely finely stellate-pubescent; the pubescence below the inflorescence usually of dark, capitately glandular trichomes about 0.4-0.6 mm. long. LEAVES sessile or with petioles up to 3 mm. long; blade 7-30 mm. long, 0.8-4.0(5.0) mm. wide, oblanceolate-elliptic to linear-lanceolate near the top of the plant, green and stellate-pubescent on both surfaces; midvein prominent and secondary veins obscure beneath; base of blade attenuate to cuneate, apex acute; margin entire and sometimes slightly revolute. INFLORESCENCE: corymbose to short-paniculate. FLOWERS isomorphic (all chasmogamous). Inflorescence with sparse to dense stellate pubescence intermixed with coarse, red, glandular hairs mostly 0.4-0.6 mm. long. Bracts 3-10 mm. long, 0.5-1.5 mm. wide, lanceolate and nearly same shape as the outer sepals. CHASMOGAMOUS FLOWERS: pedicels 0.5-4.0(6.0) mm. long, usually shorter than calyx. CALYX with simple, white, villous trichomes about 0.8-1.5 mm. long intermixed with fewer, shorter (c. 0.2-0.4 mm.) dark reddish, capitately glandular trichomes. OUTER SEPALS (free portion) 2.5-4.0 mm. long, 0.5-1.0 mm. wide, lanceolate; INNER SEPALS 4.5-7.0 (8.0) mm. long, 3-4 mm. wide, ovate, acuminate, PETALS 5-8 mm. long, 3-4 mm. wide, obovate, yellow. STAMENS 20-25. PISTIL 1.8-2.2 mm. long; ovary 1.2-1.4 mm. long, 0.8-1.0 mm. in diameter, ovoid, glabrous; style 0.5-0.7 mm. long; stigma 0.4-0.6 mm. wide. FRUITING CALYX 4.5-8.0 mm. long, 3.5-4.0 mm. in diameter, ovoid. CAPSULE 4.3-6.0 mm. long, 3.0-3.5 mm in diameter, ovoid, glabrous, 3-valved, each 3.0-3.5 mm. wide, ovate-elliptic, acute. SEEDS c. 15, inequilateral and the thin membrane readily separating when seeds moistened. FLOWERING: April-May. HABITAT: dry rocky ridges. DISTRIBUTION: Known only from certain islands off the southern California coast. (Santa Catalina, Santa Cruz and Santa Rosa Islands). (Map 10.) This species is readily distinguished from the closely related H. scoparium by its inflorescence which is densely covered with both simple villous and red glandular hairs and by its lanceolate outer sepals. There is some evidence that limited hybridization has occurred between this species and at least one form of the baffingly variable H. scoparium. Certain plants (Abrams & Wiggins 155 [uc] and Hoffman 15 June 1930 [SBM], which in most other respects would have been quickly relegated to the last mentioned species, proved exceptional 278 Rhodora [Vol. 67 in possessing at least a moderate sprinkling of glandular trichomes very similar to those possessed by H. Greenei. At least three recognizable variants of H. scoparium are to be found within the confined range of H. Greenei. These are the so-called varieties scoparium, vulgare, and a compara- tively rare form, possessing extremely mealy-pubescent stems and foliage, which has never been formally named and which occurs apparently only on Santa Cruz Island and the adjacent mainland of Santa Barbara and Ventura Counties. A few of the insular specimens of this unnamed variant possess some glandular trichomes of the sort found in H. Greenei which apparently grows in close proximity. Ex- tensive field work within the Californian complex would doubtless prove most profitable. Grosser (Pflanzenreich 14 IV. 193: 33. 1903.) assigned this species, H. scoparium, and the endemic Chilean H. spartioides to the section Spartioides of the segregate genus Halimium. Those three species were distinguished by him from other New World species of Helianthemum by their spartioid habit and isomorphic (chasmogamous) flowers. The floral characteristic for such separation has lost some of its value inasmuch as H. nutans and H. patens also have only isomorphic (chasmogamous) flowers and yet they are non-spartioid. This interesting disjunction pattern has recently been the subject of a stimulating symposium (Quart. Rev. Biol. 38: 109-177. 1963.) REPRESENTATIVE SPECIMENS: CALIFORNIA: Los Angelos Co., Santa Catalina Island, ne. side of Black Jack Mt., Blakley 5433 (sBBG) ; Santa Barbara Co., Santa Cruz Island, July & August 1886 Greene (ND, NY), April 1888 Brandegee (GH, UC), Abrams & Wiggins 156 (F, UC), Clokey 5007 (GH, UC); Santa Barbara Co., Santa Rosa Island, Epling & Erickson 8 Aug. 1937 (LA). 10. Helianthemum scoparium Nutt. Helianthemum scoparium Nutt. in Torrey & Gray, Fl. N. Am. 1: 152. 1838. Type: Nuttall, not seen. “Dry hills around Monterey, California." Linum trisepalum Kellogg, Proc. Calif. Acad. Sci. 3: 42. 1863. Type: not seen. Bolander, “on the White Hills back of Oakland." Helianthemum Aldersonii Greene, Erythea 1: 259. 1893. Lectotype: 1965] Helianthemum — Wilbur 279 Alderson, June 1898. (JEPS!) "Mountains of the southern borders of San Diego Co., Calif.” Halimium scoparium (Nutt.) Gross., Pflanzenreich 14 (IV. 193) :35. 1908. Helianthemum mendocinensis Eastwood ex Gross., Pflanzenreich 14 (IV. 193) : 35. 1903. nom. illegit. Art. 34(4) ; mistakenly published in synonymy of Halimium occidentale (Greene) Gross., a synonym of H. Greenei Robins. Halimium Aldersonii (Greene) Standl., Contr. U.S. Nat. Herb. 23 (3): 832. 1923. Crocanthemum scoparium (Nutt.) Millsp., Field Mus. Publ. Bot. 5: 175. 1923. Helianthemum scoparium var. vulgare Jepson, Man. FI. Pl. Calif. 641. 1925. Lectotype: Jepson July 3, 1896, Coulterville [Mariposa Co., California]. (JEPS!) Crocanthemum Aldersonii (Greene) Janchen, Nat. Pflanzenfam. ed. 2. 21: 805. 1925. Helianthemum scoparium var. Aldersonii (Greene) Munz, Man. South. Calif. 316. 1935. Helianthemum suffrutescens Schreiber, Madrofio 5: 81. 1939. Type: Schreiber 2243 (UC!). Isotype: (GH!). Amador Co.: dry slope 5.5 miles west southwest of Bisbee Peak, elevation 500 feet. Suffruticose perennial, usually spartioid, (5)12-42(65) em. tall with many stems arising from a woody caudex. STEMS spreading to erect, usually green and sparsely stellate-pubescent to glabrate (rarely covered with a dense stellate tomentum). LEAVES sessile or on petioles up to 3 mm. long; blade (4)7-26(43) mm. long, (8)10-15(24) -times longer than wide, (0.5)0.8-2.0(6.0) mm. wide, sometimes elliptic but usually narrowly oblanceolate to linear, green and sparsely stellate- pubescent to glabrate on both surfaces (occasionally covered with dense stellate tomentum) ` midvein prominent and secondary veins obscure beneath; base of blade attenuate, apex obtuse to acute; margin entire, mostly revolute. FLOWER: isomorphic (chasmogamous), few to many, paniculate to racemose or sometimes even appearing solitary at the tip of the very short branch (of the current year's growth). Pedicels and calyces glabrous to moderately stellate-pubescent (or more rarely covered with a dense stellate tomentum). CHASMOGAMOUS FLOWERS: pedicels 2-12 mm. long, ranging from shorter to three-times longer than the length of the calyx. OUTER SEPALS (free portion) 0.5-4.5 mm. long, 0.1-0.8 the length of the inner sepals, 0.2-0.4 mm. wide, linear; INNER SEPALS 2.5-7.0 mm. long, 2.0-3.5 mm. wide, ovate, acuminate. PETALS 3-11 mm. long, 3-8 mm. wide, obovate, yellow. STAMENS 12-47. PISTIL 1.4-2.0 mm. long; ovary 0.8-1.2 mm. long, ovoid, glabrous; style 0.5-0.8 mm. long, sometimes nearly as long as the ovary; stigma 0.3-0.5 mm. wide. FRUITING 280 Rhodora [Vol. 67 CALYX 3-8 mm. long, 2-3 mm. in diameter, ovoid. CAPSULE 2.5-4.0 mm. long, 1.8-2.7 mm. in diameter, ovoid, glabrous, 3-valved, each 1.8-2.7 mm. wide, ovate, acute. SEEDS 4-10, inequilateral, when moistened the thin membrane separable and with white papillae and the embryo visible. FLOWERING: March-July. HABITAT: dry sandy or rocky soil of hills and ridges. DISTRIBUTION: along the Californian coast from Mendocino Co. southward into northwestern Baja California and along the Sierra Nevada mountains from El Dorado Co. southward into Fresno Co. ; also known from several off-shore islands (Santa Rosa, Santa Cruz and Santa Catalina). (Map 11.) Helianthemum scoparium is an unusually variable species, which may be deduced in part from the fact that since it was first described, two proposed species and several varie- ties have been segregated from it. In addition, several collections were designated as new entities in herbaria but apparently never published. Even this indicated diversity fails to express the observed variation. The variation is particularly pronounced in respect to the habit, pubescence, number of flowers per inflorescence, size of inflorescence, length of both inner and outer sepals and the petals, and in the size as well as shape of the leaves. All of these variable characters have been used as criteria in specific or varietal segregation. Considerable time has been spent by us in examining more than 750 herbarium speci- mens in an attempt to determine what importance should be assigned to these characters, as well as to feasibility of specific and/or varietal segregation. The plants collected along the immediate coast from Mendocino Co. southward into Santa Barbara Co. have a low, spreading to ascending habit, with but few compara- tively large flowers either solitary at the tip of the very short branchlets to few in leafy racemose cymes, Their outer sepals are mostly 3-5 mm. long. The type of the species was based upon such plants. Plants of the so-called var. vulgare are relatively taller than those mentioned above and are mostly ascending to erect, with more numerous but smaller flowers and with 1965] Helianthemum — Wilbur 281 outer sepals less than half as long as the inner ones. Such plants occur almost throughout the entire range of the species but are especially abundant along Coastal Ranges from Lake County south into northern Baja California. The third previously recognized variant is a typically tall plant with an open panicle and with relatively longer petals. Plants possessing these features range from western San Bernardino Co. south into northern Baja California. Greene (Erythea 1: 259. 1893.) described such plants from the "mountains of the southern border of San Diego Co." as H. Aldersonii. Robinson (in Gray’s Syn. Fl. N. Am. 1(1): 191. 1895.) treated Greene's H. Aldersonii as a synonym of H. scoparium stating “If Prof. Greene's species is repre- sented as appears from character by Dr. Palmer's No. 18 from the same region, it is with little doubt merely a south- ern and more leafy form of H. scoparium, at least such was Dr. Gray's view." The features employed by Munz to characterize var. Aldersonii do not appear distinctive when a large series of specimens is examined. It is certain that numerous speci- mens occur with the open inflorescence of Aldersonii but with smaller petals which fall within the range of vulgare. Specimens with characteristics designated by Munz for var. Aldersonii are found along the coast as well as in the inland area attributed to it. In many respects numerous plants from the lower slopes of the Central Sierra are intermediate between var. vulgare and var. Aldersonii in possessing the smaller petals of the former and the more open inflorescence of the latter. Schreiber (Madrono 5: 81. 1939.) described plants from the foothills of the Central Sierra calling the proposed taxon H. suffrutescens. Schreiber’s type was from Amador County but plants with these features were seen from four nearby counties. The suggested differentiating characters prove variable even among specimens designated by Schreiber as H. suffru- tescens. In addition, a comparison of the leaves of H. suffru- tescens and H. Aldersonii shows them to be often similar in 282 Rhodora [Vol. 67 size and shape although the leaves of Schreiber's plants were more pubescent. The greater degree of pubescence alone does not appear worthy of formal recognition. Schreiber herself admitted that there are no floral or seed differences between her species and H. scoparium var. vulgare. A key is presented below based largely upon that published by Schreiber supplemented by the more recent treatment by Munz summarizing the differences supposedly distinguishing these taxa. 1. Leaves narrowly linear, green, sparsely stellate-pubescent to glabrous, deciduous in summer. 2. Plants low and divaricate, matted, with ultimate twigs erect; flowers few; inflorescence usually very leafy; inner sepals 4-5 mm. long; petals 5-7 mm. long (coastal from Mendocino Co. to Santa Barbara Co.). .................... H. scoparium var. scoparium. 2. Plants tall and rush-like; flowers numerous; inflorescence sparse- ly leafy; inner sepals 2-6 mm. long; petals 4-12 mm. long. 3. Panicle usually narrow; petals about 4-6 mm. long; inner sepals 2-3.5 mm. long (from Lake Co. south into northern Baja California and on the lower slopes the Sierra from Placer Co. to Mariposa Co., also found on both Santa Catalina and Santa Cruz Islands.) ................... eee H. scoparium var. vulgare. 3. Panicle extremely open; petals 8-12 mm. long; inner sepals 5-6 mm. long (from southwestern San Bernardino Co. south into northern Baja California). .. H. scoparium var. Aldersonit. 1. Leaves linear-lanceolate or oblanceolate, densely stellate-pubescent; not deciduous in summer (foothills of the Sierra in Amador Co.). dsseeneusseucasecsssssonessusuavuucssesscseseaeuenussccsssesssesuvapsacsecesssssssusennesnse H. suffrutescens. At the present state of our knowledge we do not feel that the variation noted within this complex can best be treated by the recognition of formal infraspecific categories. The present system as illustrated by the above key is in our opinion unsatisfactory. For example, the erect var. vulgare apparently contains populations that have small flowers (as is called for by the key) and other populations whose large flowers closely approximate those of var. scoparium. Perhaps as many as one-fifth of the specimens examined possess a combination of characters which would make it difficult to refer them to any of the so-called varieties or species. Such plants occur nearly throughout the range of the species but 1965] Helianthemum — Wilbur 283 are most abundant and perplexing in the foothills of the central Sierras, southern California and the offshore islands. The present unsatisfactory treatment can best be resolved by extensive observation of populations in their natural habitats together with transplant studies. The progeny of a few plants collected in the wild and grown at the Rancho Santa Ana appear to be quite different from their seed parent. We have pointed out under H. Greenei the presence of an unpublished variant of this species which occurs on several of the Channel Islands and to a limited extent upon the adjacent mainland that is rendered conspicuous by its mealy pubescence. We can only conclude that the entire complex is most deserving of intensive field study and that the problem does not lend itself to solution by reference to the herbarium. REPRESENTATIVE SPECIMENS: UNITED STATES; CALI- FORNIA: Amador Co., Ione, Hoover 2415 (UC); Calaveras Co., 2.5 mi. nw. of Harmon Peak, Roseberry 209 (UC); Contra Costa Co., Mount Diablo, Ridge south of Uncle Sam Canyon (on Eagle Point Ridge), Bowerman 1598 (Uc); El Dorado Co., Sweetwater Creek, June 1907, Simpson (UC); Fresno Co.: Chaparral, at n. end of the sw. facing Big Sandy Bluffs, 2 June 1935, Quibell (coto); Lake Co.: hills west of Mirabel Mine, Mason 8324 (UC); Los Angeles Co.; dry ridges and hill sides, Mandeville Canyon, Santa Monica Mountains, Clokey & Templeton 4490 (F, GA, GH, ILL, MIN, MO, NO, PENN, UC, US, WIS) ; Santa Cataline Island, Nuttall 154 (uc); Marin Co., Mt. Tamalpais, Heller 5721 a (F, GH, MO, PH, US) ; Mariposa Co., Devil's Gulch, 4 July 1892, Congdon (MIN); Mendocino Co.: about Mendocino, Brown 785 (F, MIN, MO, UC); Monterey Co., from type locality along the railroad near seaside (beyond Del Monte), Heller 6751 (COLO, F, GH, IND, ISC, MIN, MO, UC, US, WIS); Napa Co., Sarco Creek 4 mi. above Napa, Ewan 8825 (NO, SMU); Orange Co., Santa Ana River Canyon, Munz & Johnston 5307 (UC); San Benito Co., 10 mi. below San Benito, Fos- berg 84992 (GH, KANU); San Bernardino Co., Etiwanda, Abrams 2664 (DS, GH, MO, PH, UC, US); San Diego Co., La Jolla, Clements & Clem- ents 40 (ARIZ, COLO, F, GH, ILL, PENN, PH, MIN, MO, UC); San Francisco Co. without exact locality, Michener & Bioletti (GH); San Luis Obispo Co., 1.5 mi. sw. of Los Berros, Lee 411 (uc); San Mateo Co., Pilarcitos Lake and Canyon, Davy 1157 (uc); Santa Barbara Co., Santa Cruz Island, Clokey 5006 (GH, ILL, MIN, PENN, UC, US) ; Carpin- teria, 26 March 1958, Pollard (ARIZ, TEX); Santa Rosa Island, June 1888, Brandegee (GH, UC); Santa Clara Co., foothills west of Los Gatos, Heller 7350 (F, GH, ILL, IND, ISC, MO, PH, UC, US, WIS); Santa 284 Rhodora [Vol. 67 Cruz Co. Pine Mt. Trail, Big Basin, Mason 2247 (UC); Sonoma Co., Vine Hill, w. of Santa Rosa, Heiser 1744 (UC); Ventura Co., near North Signal Street, 17 April 1945, Pollard (GH). MEXICO: BAJA CALIFORNIA: Burned over hills about 20 mi. e. of Ensenada, Wig- gins 11867 (DS, UC); Tecate, n. Baja California, Fosberg 8346 (DS, MO, PENN). 11. Helianthemum nutans T. S. Brandeg. Helianthemum nutans T. S. Brandeg., Proc. Calif. Acad. ser. 2. 2: 129. 1889. Type: Brandegee, 7 May 1889 (Uc!). Isotypes: (A, F, NY, US!). “very abundant in rocky soil about the plains of San Julian" Baja California. Halimium nutans (T. S. Brandeg.) Standl., Contr. U.S. Herb. 23(3) : 834. 1923. Crocanthemum nutans (T. S. Brandeg.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 207. 1925. Bushy shrub, 10-30 cm. tall with few to numerous stems arising from a thick woody root. STEMS woody, gray, ascending, much branched. BRANCHES ascending, woody, with numerous, bushily ar- ranged ultimate branchlets; those of the current growth light green and densely, short stellate-pubescent. LEAVES only on current season's growth, sessile; blade 5-8 mm. long, 0.8-1.5 mm. wide; spatulate- oblanceolate, broader above the middle; light green and very densely short stellate-pubescent on both surfaces and also ciliate with 1-3 simple hairs (1-1.5 mm. long) along both margins of the blade; mid- vein but slightly elevated and secondary veins obscure beneath; leaf- blade tapering to the base, apex obtuse to somewhat acute; margin entire, non-revolute. (The old branches covered with axillary fascicles of small leaves that persist long after the principal leaves have fallen). FLOWERS isomorphic (chasmogamous), few on each branchlet, borne in a sympodial inflorescence but appearing solitary, either alternating with or opposite the leaves. Pedicel and calyx stellate-puberulent, sometimes the calyx with few simple, elongate trichomes resembling those of the leaf margin. CHASMOGAMOUS FLOWERS: pedicels 7-15 mm. long, somewhat curved, and articulate at the base. OUTER SEPALS (free portion) 2-4 mm. long, 1/3-1/2 the length of the inner sepals, 0.2-0.3 mm. wide, linear; INNER SEPALS 5.5-7.0 mm. long, 2.8-3.2 mm. wide, ovate-lanceolate, twice longer than wide, acute, bright green. COROLLA bright yellow; petals 7-9 mm. long, 5-7 mm. wide, broadly cuneate. STAMENS 25-32. PISTIL 1.6-2.0 mm. long; ovary 1.2-1.4 mm. long, 0.8-1.0 mm. in diameter, ovoid, glabrous; style 0.4-0.6 mm. long; stigma 0.4-0.6 mm. wide, capitate. FRUITING CALYX 6-8 mm. long, 3.0-3.5 mm. in diameter, nar- rowly ovoid. CAPSULE 4.8-5.2 mm. long, 2.4-2.7 mm. in diameter, nar- rowly ovoid-triquetrous, about twice as long as wide, glabrous, 3-valved, each valve 2.4-2.7 mm. wide, narrowly ovate, acute, slightly 1965] Helianthemum — Wilbur 285 concave. SEEDS 20-26, inequilateral, the thin seed membrane with numerous white papillae especially on the edges of the seed; the membrane separable when moistened and then the embryo becoming somewhat visible. FLOWERING: February-April HABITAT: on dry, rocky soil of flats and gullies. DISTRIBUTION: southern Baja Cali- fornia, Mexico. (Map 16, large dots.) This little-collected species is endemic to Baja California. It can be easily distinguished from all other North American species of Helianthemum by its low, shrubby and much- branched habit, and by its spatulate leaves (0.5-1.5 mm. wide) whose margins bear on each side 1-3 simple hairs about 1.0-1.5 mm. long. Grosser (Pflanzenreich 14 (IV. 193): 52. 1903.) sup- posed, judging only from the original description, that this species might be closely related to H. scoparium. The unique features possessed by this species would seem to indicate a relatively isolated position within the genus. REPRESENTATIVE SPECIMENS: MEXICO: BAJA CALIFORNIA: Lower California, Plains of San Julian, 7 May 1889, Brandegee (A, F, NY, UC, US); 5 mi. n. of Mesquital Grande, 9 Feb. 1935, Haines & Stewart (A, ARIZ, DS, F, ILL, ND, RSA, TEX, UC); bushy gully in Creta- ceous Sediments, 4 mi. e. of Santa Catarina Landing, Dressler 610 (Mo) ; locally common on bare flat area near Agua de Higuera, Moran 10282 (DUKE, SD). 12. Helianthemum patens Hemsl. Helianthemum patens Hemsl., Diagn. Pl. Nov. Mex. et Centr. Amer. part 2: 20. 1879. Type: Parry & Palmer 30 (K). Isotypes: (F, GH, MO, NY, US!). “Mexico: in regione San Luis Potosi, alt. 6000-8000 ped." Halimium patens (Hemsl) Gross, Pflanzenreich 14 (IV. 193): 46. 1903. Perennial herb or somewhat suffruticose, (3) 10-17(30) cm. tall with one to few stems arising from a woody caudex. STEMS ascending to erect, glabrous or densely stellate-pubescent, branching from near the base. LEAVES (on current season's growth) sessile or on petioles up to 1.0 mm. long; blade 2-8(15) mm. long, 0.6-4.0(6.0) mm. wide, ovate- elliptic or lanceolate, glabrous or sparsely to densely stellate-pubescent on both sides, midvein prominent and secondary veins obscure beneath; base of blade cuneate, apex acute or pointed and lustrous; margin entire or nearly so, non-revolute. Glabrous leaf-blades generally 286 Rhodora [Vol. 67 lanceolate and 5-times or more longer than wide; the sparsely to densely stellate-pubescent leaf-blades generally ovate-elliptic and 5- times longer than wide or less, FLOWERS isomorphic (chasmogamous) and with but few on each branch, borne in sympodial inflorescences but appearing solitary, axillary, or subterminal and either alternating or opposite the leaves. CHASMOGAMOUS FLOWERS: pedicels 5-15 mm. long, (1)1.5-3 times longer than calyx, divergent to ascending, purple or greenish or even hoary in color, glabrous or stellate-pubescent, disarticulating near the base. CALYX either glabrous and lustrous and mostly purplish, or sparsely stellate-pubescent intermixed with simple villous hairs or densely and minutely stellate-pubescent. OUTER SEPALS (free portion) 1.0-2.4 mm. long, 4-% the length of the inner sepals, 0.2-0.4 mm. wide, linear; INNER SEPALS 2.6-4.0 mm. long, 1.6-2.5 mm. wide, ovate, acute. COROLLA pale yellow; petals 3-6 mm. long, 2.4-5.0 mm. wide, obovate. STAMENS 10-14(20). PISTIL 1.4-1.7 mm. long; ovary 0.8-1.2 mm. long, c. 0.8 mm. in diameter, ovoid, glabrous; style 0.2-0.5 mm. long; stigma 0.3-0.6 mm. wide, capitate. FRUITING CALYX 3-5 mm. long, 2.5-4.0 mm. in diameter, ovoid to subglobose. CAPSULE 3-4 mm. long, 2.4-3.2 mm. in diameter, ovoid to subglobose, 3-valved; each valve 2.4-3.2 mm. wide, ovate, acute, somewhat rounded at the back. SEEDS 10-13(20), inequilateral, light brown, with separable membrane when moistened and the embryo then visible. FLOWERING: May-August. HABITAT: dry open rocky, grassland hills ond old lava among oak scrub. DISTRIBUTION: Mexico: from Zacatecas and San Luis Potosí to south- eastern Puebla. (Map 16, small dots.) Helianthemum patens can be readily distinguished from all central Mexican species by its relatively few petaliferous flowers borne on divergent to ascendent pedicels 1-3 times longer than its calyces in a sympodial, axillary or subtermi- nal inflorescence. Helianthemum nutans of Baja California has the same kind of inflorescence but is easily distinguished from H. patens by its shrubby and much-branched habit, by its fasciculate leaves that persist long after the principal leaves have fallen, and by the few simple hairs (1.0-1.5 mm. long) on the margin of the leaf. Grosser (Pflanzenreich 14 (IV. 193) : 47. 1903.) treated this species as having dimorphic flowers, but no cleistog- amous flowers were seen by us on any of the specimens 1965] Helianthemum — Wilbur 287 examined. Helianthemum patens together with H. nutans, H. scoparium, and H. Greenei are the only North American species having only petaliferous flowers. Vegetatively this is a rather variable species. Specimens examined may be arranged into three pubescence types. Plants collected from near the center of distribution of the species are light green, glabrous and lustrous throughout and have lanceolate leaves. Plants collected from the mid- dle and southern portion of the species distribution are green, sparsely stellate-pubescent, somewhat lustrous and with ovate-elliptic leaves. The isotypes (and hence pre- sumably the type) and the majority of the specimens examined belong to this form. Plants collected from the central and northern range of the species distributed are green or sometimes hoary, densely stellate-pubescent throughout, with ovate-elliptic leaves, and more woody than the other two. Specimens of all three types have however been collected in the same locality. More collections are necessary before any attempt to evaluate these conspicuous variants would appear warranted. All North American species of Helianthemum are stellate- pubescent except for the few specimens of this species which were completely glabrous. REPRESENTATIVE SPECIMENS: MEXICO: JALIsco: sw. of Ojuelos on road to Aquascalientes, McVaugh 16799 (MICH), McVaugh 16849 (MICH), McVaugh 16850 (MICH); McVaugh 16851 (MICH). PUEBLA: vicinity of San Luis Tultitanapa, near Oaxaca, Purpus 3383 (F, GH, MO, UC); below Atzizintla (San Antonio), Balls 5315 (A, UC); Esperanza, Purpus 2486 (F, GH, MO, UC); Esperanza, Balls 5401 (GH, UC); SAN LUIS POTOSI: San Rafael, Schaffner 498 (F, ISC, MICH, MIN, PH, POM); chiefly in the region of San Luis Potosi, Parry & Palmer 28% (GH, PH), Parry & Palmer 30 (F, GH, MO, NY, US); Mts. w. of San Luis Potosí, Sierra Madre Oriental, Pennell 17630 (MICH, NY, PH, UC); Alvarez, Palmer 602 (F, GH); ZACATECAS: 25 mi. nw. of Fresnillo, Waterfall 15590 & 15591 (OKLA); Zacatecas, Aug. 1903, Purpus (UC). 13. Helianthemum corymbosum Michx. Helianthemum corymbosum Michx., Fl. Bor.-Am. 1: 307. 1803. Type: Michaux (P). Photograph of type (GH!). “In maritimis Caro- linae et Georgiae." Cistus corymbosus (Michx.) Poir., Encyc. Suppl. 2: 272. 1811. 288 Rhodora [Vol. 67 Heteromeris cymosa Spach, Ann. Sci. Nat. 2nd. ser. 6: 370. 1836. nom. nud. & illegit. since Helianthemum corymbosum Michx. is cited as a synonym; descr, Spach, Hist. Nat. Vég. Phanér. 6: 103. 1838, but Helianthemum corymbosum Michx. is again cited as a synonym. Halimium corymbosum (Michx.) Gross., Pflanzenreich 14 (IV. 193): 49. 1903. Crocanthemum corymbosum (Michx.) Britt. in Britton & Brown, Ill. Fl. ed. 2. 2: 541. 1913. Perennial herb or rarely suffruticose, (7)11-30(52) cm. tall with one to many stems arising from a caudex. STEMS erect or nearly so, covered with appressed stellate-pubescence but becoming glabrate. BRANCHES alternate, wiry, ascending, mostly restricted to the upper half of the stem. CAULINE LEAVES: petiole 1-5 mm. long; blades (9)18-35(47) mm. long, (1.5)4-10(13) mm. wide; obovate-elliptic, elliptic to elliptic-lanceolate near the apex, green and stellate- puberulent beneath, conspicuously discolorous; midvein and secondary veins prominent beneath; base of blade cuneate, apex obtuse or nearly So to acute; margin entire, upper leaves slightly revolute. INFLORES- CENCE: a compound dichasium terminating the stem and sometimes the branches. FLOWERS dimorphic; the long-pedicellate chasmogamous flowers overtopping the very shortly pedicellate, smaller, more nu- merous and congested cleistogamous flowers. Bracts 1.6-7 mm. long, 0.2-1.2 mm. wide, spatulate-linear. Pedicel and calyx covered with short stellate pubescence and also with longer (0.5-1.5 mm. long), simple, often very abundant, villous trichomes. CHASMOGAMOUS FLOWERS: pedicels 6-15(28) mm. long, 1-5 times longer than calyx. OUTER SEPALS (free portion) 2.4-4(5.2) mm. long, 0.7-1.2 mm. wide, spatulate-linear, apex obtuse or nearly so, often drying brownish; INNER SEPALS 3-6(8) mm. long, 2.5-3.6 mm. wide, ovate, acute to acuminate. PETALS 6-11 mm. long, 5-9.5 mm. wide, obovate. STAMENS 20-30. PrsTIL 1-1.4 mm. long, ovary 0.7-1.1 mm. long, 0.6-1 mm. wide, ovoid, glabrous; stvle 0.2-0.5 mm. long; stigma 0.5-1 mm. wide, capitate. FRUITING CALYX 4-6.5(7.8) mm. long, 1.5-5 mm. in diameter. CAPSULE 3.6-5.4 mm. long, 3-4 mm. in diameter, ovoid and somewhat angled, apiculate, glabrous, 3-valved; each valve 2.4-3.3 mm. wide, ovate, acute, slightly concave; funiculi and placentae re- maining attached to the receptacle after valves and seeds have fallen. SEEDS 15-30, ovoid to somewhat irregular, light to dark brown, smooth and with a thin, separable membrane. CLEISTOGAMOUS FLOWERS: numerous, subsessile, sepal tips (especially the outer ones) brown. STAMENS 4-6. FRUITING PEDICELS 1-2.5 mm. long, shorter than calyx. FRUITING CALYX 2.2-4.8 mm. long, 1.8-3.5 mm. in diameter, ovoid. OUTER SEPALS (free portion) 1.8-3 mm, long, 0.3-0.9 mm. wide, spatulate-linear, apex obtuse or nearly so and brown; INNER SEPALS 2.2-4.8 mm. long, 1.2-3 mm. wide, ovate, acute. CAPSULE 1.6-3.8 1965] Helianthemum — Wilbur 289 mm. long, 1.4-3 mm. in diameter, ovoid-triquetrous, apiculate, 3-valved; each valve 1.4-2.4 mm. wide, ovate-elliptic, acute, slightly concave; funiculi and placentae remaining attached to the receptacle after valves and seeds have fallen. SEEDS 4-8(10), similar to those in the chasmogamous fruit. FLOWERING: February-April. HABITAT: sand dunes and beaches, dry sandy pinelands, fields and openings in live-oak woodlands. DISTRIBUTION: along the Coastal Plain of North Carolina southward throughout peninsular Florida and west along the Gulf Coast into Mississippi. (Map 8.) REPRESENTATIVE SPECIMENS: NORTH CAROLINA: Carteret Co., Lennox Point, Beaufort, Godfrey 49843 (COLO, DS, DUKE, FSU, ILL, IA, IND, ISC, KANU, MIN, NCSC, NCU, NY, PENN, PH, SMU, TEX, UC, US, WIS, WVA); Dare Co., pineland at Frisco, Radford, Haesloop & Miller 7790 (NCU). SOUTH CAROLINA: Beaufort Co., Hunting Island State Park, Bell 2478 (FSU, NCU) ; Berkeley Co., 5 mi. w. of Pineville, Godfrey & Tryon 1134 (GH, NY, US) ; Charleston Co., Isle of Palms, Robinson 133 (GH); Georgetown Co., 3 mi. e. of Georgetown, Godfrey & Tryon 1085 (CU, DUKE, GH, MICH, MO, MIN, NY, PÉNN, TENN, PH, UC, US). GEORGIA: Charlton Co., at Camp Cornelia, Duncan 7428 (GA, NCSC, SMU); Chat- ham Co., Tybee Island, Harper 2173 (GH, MO, NY, US). FLORIDA: Broward Co., Hollywood, Moldenke 599 (DUKE, ILL, MO, NY, PENN); Citrus Co., s. of Inverness, Cooley, Wood & Wilson 5936 (NCU, US, USF); Clay Co., Hibernia, March 1869, Canby (F, GH, MO, MICH, NY, us); Dade Co., Humbugus Prairie, Small & Mosier 5602 (DUKE, FLAS, GH, NCU, NY, US); Dixie Co., 10 mi. n. of Suwannee, Godfrey 56476 (DUKE, FSU, IA, USF, UC, VDB); Duval Co., near Jacksonville, Curtiss 4601 (GH, ISC, MIN, MSC, NY, UC, US); Curtiss 227 (CU, F, FLAS, GH, IA, KANU, MIN, MO, NY, PH, SMU, UC, US); Franklin Co., Alligator Point, Godfrey 53201 (DUKE, FSU, GH, IA, NCSC, NCU, NY, USF, VDB) ; Lake Co., vicinity of Eustis, Nash 137 (GH, MIN, MSC, NY, PH, UC, US) ; Lee Co., w. of Fort Myers, Moldenke 926 (DUKE, ILL, MIN, MO, NY, PENN). MISSISSIPPI: Harrison Co., Cat Island, Ray 2728 (USF). 14. Helianthemum concolor (Riley) Ortega Halimium concolor Riley, Kew Bull. 1923: 107. 19 April 1923. Type: Gonzalez 842 (K) not seen, “San Ignaciei Cerro del Buen Retiro, 1660 M." Sinaloa. Halimium exaltatum Rose & Standl., Contr. U.S. Nat. Herb. 23 (3): 833. 18 July 1923; ibid., 23: 1675. 1926, Riley’s epithet cited as follows: “Page 833. Halimium exaltatum. A synonym is Hali- mium discolor [sic] Riley, Kew Bull. 1923: 107. 1923." Type: Pringle 10409 (Cell. Isotypes: (F, DUKE, GH, OKLA, MIN, MSC, sMU!). “dry hills above Uruapan, Michoacan, alt. 1,500 m.” Crocanthemum exaltatum (Rose & Standl.) Janchen, Nat. Pflanzen- fam. ed. 2. 21: 305. 1925. 290 Rhodora [Vol. 67 Helianthemum concolor (Riley) Ortega, Cat. Sist. Pl. Sinaloa 4. 1929. Helianthemum exaltatum (Rose & Standl.) Ortega, Cat. Sist. Pl. Sinaloa 4. 1929. Shrub up to 1 m. tall. STEMS erect, coarsely stellate-pubescent. CAULINE LEAVES: petiole 3-5 mm. long; blade 11-42 mm. long, 5-20 mm. wide; elliptic-obovate, green and sparsely stellate-pubescent on both sides, lower epidermis conspicuous and not masked by the pubescence; midvein and secondary veins conspicuous and mostly elevated beneath; base of blade attenuate, apex obtuse to acute, sometimes mucronate; margin entire, non-revolute. FLOWERS dimorphic (chasmogamous and cleistogamous) crowded at the end of the stem and branches. Chasmogamous flowers fewer and on pedicels sub- equaling to slightly longer than those of the cleistogamous flowers. Pedicel and calyx covered with short stellate-pubescence and with white and with much longer, coarse, stellate-pubescence, with the individual trichomes up to 2 mm. long. Bracts numerous, 2-6 mm. long, 0.2-0.5 mm. wide, linear to narrowly lanceolate. CHASMOGAMOUS FLOWERS: pedicels 3.5-9.5 mm. long. OUTER SEPALS (free portion) 2.5-5.0 mm. long, 0.2-0.3 mm. wide, very narrowly linear; INNER SEPALS 4.5-7.0 mm. long, 2.8-3.6 mm. wide, ovate, acumi- nate, sometimes with veins prominent at the back. COROLLA yellow, petals 5-8 mm. long, 5-6 mm. wide, obovate. STAMENS 20-30. PISTIL 1.6-2.0 mm. long; ovary 1.2-1.6 mm. long, 0.5-0.8 mm. in diameter, ovoid, glabrous; style 0.4-0.7 mm. long; stigma 0.3-0.4 mm. wide, capitate. Chasmogamous flowers with mature fruit not seen. CLEISTOGAMOUS FLOWERS: pedicels 2-3(up to 5 at fruiting) mm. long. Outer sepals (free portion) 2.0-3.6 mm. long, 0.1-0.3 mm. wide, narrowly linear; INNER SEPALS 3.5-5.0 mm. long, 1.6-2.0 mm. wide, ovate-lanceolate, acuminate. STAMENS 5-7. PISTIL 1.5-2.0 mm. long, ovary ovoid, glabrous. FRUITING CALYX 4.8-6.5 mm. long, 2.4-3.6 mm. in diameter, ovoid. CAPSULE 3.6-4.4 mm. long, about twice longer than wide, 1.7-2.5 mm. in diameter, ovoid-triquetrous, glabrous, 3-valved; each valve 1.7-2.4 mm. wide, ovate to ovate-lanceolate, acute. SEEDS 4-7, ovoid or depressed on both sides, with thin separable membrane and the embryo becoming visible when seed is moistened. FLOWERING: October-February. HABITAT: Dry hillsides. DISTRIBUTION: Mexico: southern Sinaloa, northwestern Jalisco, central and northwestern Michoacan, and south- western Mexico State. (Map 17, small dots). This is the only Mexican species with a leaf-blade so sparsely stellate-pubescent that the epidermis is readily visible beneath. SPECIMENS EXAMINED: MEXICO: JALISCO: San Sebastián, Segundo Arroyo, Mexia 1542 (UC); trail from Real Alto to San Sabestián, Mexia 1773 (F, UC). MEXICO: Dist. Temascaltepec, Pineda, 1965] Helianthemum — Wilbur 291 Hinton 3190 (ARIZ, GH, PH). MICHOACÁN: Uruapan, Pringle 13447 (GH, MICH); dry hills above Uruapan, Pringle 10409 (F, DUKE, GH, OKLA, MIN, MSC, SMU, US) ; Dist. Coalecoman, Coaleoman, Hinton 12960 (ARIZ, GH). SINALOA: 8.7 mi. e. of Portrerillos; ca. 36 mi. e. of Con- cordia, Breedlove 1679 (DUKE). 15. Helianthemum georgianum Chapm. Helianthemum georgianum Chapm., Fl. S. U.S. ed. 3. 36. 1897. “Bainbridge, Georgia, Mobile (Mohr).” Lectotype: Chapman (M0!). Halimium georgianum (Chapm.) Gross., Pflanzenreich 14 (IV. 193): 49. 1903. Crocanthemum stipulatum Janchen, Oster. Bot. Zeit. 71: 269. 1922. Type: Tracy 8060. (W, not seen.) Isotypes: (all seen F, GH, MIN, OS, TEX, UC). Weatherford, Texas. Crocanthemum georgianum (Chapm.) Barnh. in Small, Man. SE. U.S. Fl. 879. 1933. Perennial herb or very rarely suffruticose, (10)15-30(40) em. tall with few to many stems arising from a caudex, spreading by slender, shallowly placed, horizontal roots. STEMS ascending to erect, stellate- tomentose, divergently branched. BASAL LEAVES often in a rosette or forming a mat at the base of the stem, but frequently lacking; petiole 1-2 mm. long; blade 10-28 mm. long, 4.5-11.0 mm. wide; spatu- late-oblanceolate, stellate-tomentose on both surfaces; base of blade attenuate, apex obtuse to acute; margin entire, non-revolute. CAULINE LEAVES: estipulate (sometimes with smaller axillary fasciculate leaves at first resembling stipules) ; petiole 1-3 mm, long; blade (7)20-35 (44) mm. long, (1.2)4.5-8.5(12) mm. wide; oblanceolate and becoming nar- rowly elliptic near the top of the stem, green and stellate-pubescent above, hoary and stellate-tomentose beneath; midvein and secondary veins prominent beneath; base of blade attenuate to cuneate, apex obtuse to acute; margin entire, the upper leaves somewhat revolute. FLOWERS dimorphie (chasmogamous and cleistogamous), 2-7 in race- mose cyme at the end of the main stem and lateral branches. First chasmogamous flowers borne on longer pedicels. Bracts 1.2-5.0 mm. long, 0.3-1.0 mm. wide, lanceolate. Pedicel and calyx stellate-canescent. CHASMOGAMOUS FLOWERS: pedicels 5-12(15) mm. long, 1-3 times longer than calyx. OUTER SEPALS (free portion) 1.5-3.5 mm. long, 5-10 times longer than wide, 0.3-0.5 mm. wide, linear; INNER SEPALS 3.6-6.6 mm. long, 3.0-4.5 mm. wide, ovate, acute. PETALS 6-10(12) mm. long, 4-8(11) mm. wide, obovate. STAMENS (15)20-36. PISTIL 1.2-2.5 mm. long; ovary 0.8-1.7 mm. long, 0.8-1.2 mm. in diameter, ovoid, glabrous; style 0.3-0.7 mm. long; stigma 0.6-1.0 mm. wide, capitate. FRUITING PEDICELS 6-13(16) mm. long. FRUITING CALYX 5.0-7.2 mm. long, 4.2-5.4 mm. in diameter, ovoid. CAPSULE 3.8-5.7 mm. long, 3.2-4.5 mm. in diameter, ovoid, apiculate, glabrous, 3-valved; each valve 3.0-4.2 mm. wide, ovate, acute, slightly concave. SEEDS 20-35, ovoid-depressed to 292 Rhodora [Vol. 67 inequilateral, smooth and when moistened, the membrane separable and the embryo visible. CLEISTOGAMOUS FLOWERS: pedicels 0.6-3.0 mm. long. OUTER SEPALS (free portion) 1.4-2.2 mm. long, c. 0.8 mm. wide, linear; INNER SEPALS 3.0-4.2 mm. long, 2.6-3.8 mm. wide, ovate, acute. STAMENS (3)5(8), remaining attached to the base of the capsule. PISTIL 1.3-1.7 mm. long, ovary ovoid, glabrous. FRUITING PEDICELS 2-6 mm. long. FRUITING CALYX 4.0-5.4 mm. long, 3.2-4.8 mm. in diameter, ovoid. OUTER SEPALS (free portion) 1.9-3.0 mm. long, 5-10 times longer than wide, 0.3-0.5 mm. wide, linear; INNER SEPALS 4.0-5.4 mm. long, 3-4 mm. wide, ovate, acute. CAPSULE 3.0-4.2 mm. long, 2.4-3.3 mm. in diameter, ovoid, glabrous, with 3 valves; each valve 1.9-3.0 mm. wide, ovate, acute. SEEDS 12-20, similar to the seeds of the chasmogamous flowers. FLOWERING: April-June. HABITAT: Dry, open, sandy woods and fields, turkey oak sand-ridges, and sand-dunes along the beach. DISTRIBUTION: along the coast from North Carolina to northern Florida, westward along the Gulf Coast to Louisiana, and through most of eastern and central Texas; also known from southern Oklahoma and southern Arkansas. (Map2.) The 2-7 dimorphic flowers in racemose cymes at the end of the principal branches, and the frequent presence of basal leaves are enough to distinguish this species from any other Helianthemum in the United States or Canada. The inflo- rescence of H. georgianum is approached by that of the Mexican and Central American H. Pringlei. However, the leaves of H. georgianum are long-petiolate and with promi- nent secondary veins beneath in contrast to the sessile or short-petiolate leaves with obscure secondary veins of H. Pringlei. In addition basal leaves are often present in H. georgianum while they are always lacking in H. Pringlei. It was at first felt that an undescribed taxon rested un- detected under the binomial of this species. Specimens obtained from the sandy maritime forests from the Caro- linas and from the offshore islands in the Gulf of Mexico have a very different appearance from the majority of the specimens which resemble those collected in the eastern half of Texas. We were unable to find characters however to substantiate the different appearance that this extreme presents. They were noticeably different from the norm in 1965] Helianthemum — Wilbur 293 their generally low habit and but few (2(3)), mostly cleis- togamous flowers. Janchen (Oster. Bot. Zeit. 71: 269. 1922.) redescribed this species as Crocanthemum stipulatum, although all of the New World species of Helianthemum are estipulate. Janchen considered the small, fasciculate leaves found in the axil of the cauline leaves to be stipules and felt that this difference alone was of such importance that he had no hesitation to describe the new species on the basis of only this distinction. REPRESENTATIVE SPECIMENS: NORTH CAROLINA: Brunswick Co., Southport, Correll & Bloomquist 440 (PENN); Carteret Co., Lennox Point, e. of Beaufort, Fox & Godfrey 2694 (GH, NCSC, NY, PH); Dare Co.: maritime forest at Buxton, Radford, Haesloop & Miller 7580 (NCU). SOUTH CAROLINA: Horry Co.: Myrtle Beach, Wilbur 6974 (DUKE). GEORGIA: Decatur Co., Bainbridge, Chapman (IA, MO) ; Dougherty Co., Albany, 21 May 1930, Harper (ARIZ, F, GH, MICH, MO, NY, PH, US); McIntosh Co., in southeast section of Sapelo Island, Adams & Duncan 17887 (NCSC). FLORIDA: Gadsden Co., near River Junction, Curtiss 6392 (DS, F, GH, ILL, ISC, MIN, MO. NY, SMU, UC, US) ; Okaloosa Co., 0.5 mi. w. of Mary Esther, Godfrey 56671 (FSU, IA, MSC, NY, UC, USF); Wakulla Co., along the beach, Mashes Island, Godfrey 53203 (DUKE, FSU, GH, IA, NCSC, NCU, NY, TENN, USF, VDB) ; Walton Co., 5 mi. w. of Portland, Godfrey 56662 (FSU, IA, UC, VDB). ALABAMA: Mobile Co., Spring Hill, Mackenzie 4032 (GH, IND, MIN, NY). MISSIS- SIPPI: Harrison Co., Deer Island, Pollard 1184 (CU, F, GH, MO, NY, OS, US). LOUISIANA: Natchitoches Parish, Chopin, Palmer 7985 (NY, US). ARKANSAS: Ouachita Co., near Chidester, Moore 56-138 (NY). OKLAHOMA: Marshall Co., Lake Texoma, 20 June 1960, Penfound (OKL). TEXAS: Bastrop Co., West Point, Barkley et al 7039 (BUS, COLO, DUKE, FLAS, GH, ILL, ISC, MIN, PENN, TEX); Denton Co., near Roanoke, Lundell & Lundell 9539 (DS, GH, MICH, SMU); Llano Co., Enchanted Rock, Tharp et al 47273 (BUS, COLO, DUKE, FLAS, GH, ISC, MIN, TEX); Mason Co, 7 mi. n. of Mason, McVaugh 8318 (F, GH, MICH, SMU, TEX); Parker Co., Weatherford, Tracy 8060 (F, GH, MIN, MSC, OS, TEX, UC); Taylor Co., 1 mi. n. of Abilene, Tolstead 7499 (MO, NCSC, NY, SMU, TEX, UC); Travis Co., Austin, Tharp 44105 (COLO, DS, DUKE, FSU, IA, ILL, IND, ISC, MO, NCU, NO, NY, OKL, OKLA, PH, SMU, UARK, UC, VDB, WVA). 16. Helianthemum Pringlei S. Wats. Helianthemum Pringlei S. Wats., Proc. Am. Acad. 23: 268. 1888. Type: Pringle 1186, September 1887 (GH!). Isotypes: (F, MIN, MSC, NY, UC, US!). "On pine plains at the base of the Sierra Madre, Chihuahua." 294 Rhodora [Vol. 67 Halimium Pringlei (S. Wats.) Gross., Pflanzenreich 14 (IV. 193) : 46. 1903. Heteromeris Pringlei (S. Wats.) Ponzo, Nuovo Gior. Bot. Ital. n. s. 28: 169. 1921. (No basionym cited). Crocanthemum Pringlei (S. Wats.) Janchen, Nat. Pflanzenfam, ed. 2. 21: 305. 1925. Perennial herb or sometimes suffruticose, 10-42 cm. tall with few stems arising from an upright caudex (or rarely from a horizontal, subterranean rootstock). STEMS branched, erect to ascending, stellate- pubescent, but also sometimes intermixed with simple, villous hairs on the lower half. CAULINE LEAVES: mostly appressed to the stem, sometimes with but few and small fasciculate leaves in their axils; sessile or with petiole up to 1 mm. long; blade 4.5-25 mm. long, 2.0-5(7.0) mm. wide, elliptic or oblance-elliptic, rarely the lower- most obovate, grayish and stellate-tomentose on both surfaces, or occasionally intermixed with simple villous hairs (especially those on the lower half of the stem); midrib conspicuous and elevated beneath, secondary veins obscure; base of blade cuneate, the apex subacute to acute; margin entire, nonrevolute. FLOWERS: dimorphic (chasmogamous and cleistogamous), 2-5 in racemose cymes termi- nating the stem and the branches. Pedicel and calyx stellate-tomentose (the outer sepals very rarely villous). Bracts 2-7 mm. long, 0.5-1.5 mm. wide, linear-lanceolate or narrowly elliptic. CHASMOGAMOUS FLOWERS: pedicels 2-12 mm. long, shorter than to twice longer than calyx, articulate, sometimes purplish. OUTER SEPALS (free portion) 1.2-3.2 mm. long, 0.2-0.4 mm. wide, linear; INNER SEPALS 4-6 mm. long, 2-3 mm. wide, ovate, acute, sometimes purplish. PETALS 5-10 mm. long, 3.6-8.5 mm. wide, yellow or sometimes with purple spot at the middle of the upper edge. STAMENS 25-36. PISTIL 1.4-2.0 mm. long; ovary 1.0-1.4 mm. long, 0.8-1.0 mm. in diameter, ovoid, glabrous; style 0.4-0.6 mm. long; stigma 0.6-0.8 mm. wide, capitate. FRUITING PEDICELS 3-12 mm. long, shorter to twice longer than calyx, articulate. FRUITING CALYX 4.0-6.5 mm. long, 4-5 mm. in diameter, ovoid, sometimes with a distinct base. OUTER SEPALS (free portion) 2.2-3.2 mm. long, 0.2-0.4 mm. wide, linear; INNER SEPALS 4.0-6.5 mm. long, 2.8-3.2 mm. wide, ovate, acute, sometimes purplish. CAPSULE 3.5-6.0 mm. long, 3.0-4.5 mm, in diameter, ovoid-triquetrous, glabrous, 3-valved; each valve 3.0-4.5 mm. wide, ovate, acute. SEEDS 18-24, ovoid to laterally depressed or somewhat inequilateral, smooth and when moistened the membrane separable and the embryo visible. CLEISTOGAMOUS FLOWERS: fruiting pedicels 1.0-4.2 mm. long, shorter to nearly as long as calyx, articulate. FRUITING CALYX 3.2-4.5 mm. long, 2.4-3.0 mm. in diameter, ovoid, sometimes with a distinct base. OUTER SEPALS (free portion) 1.2-3.0 mm. long, 0.2-0.4 wide, linear; INNER SEPALS 3.2-4.5 mm. long, 2.2-2.8 mm. wide, ovate, acute, sometimes pur- plich. STAMENS 4-5. CAPSULE 2.4-4.5 mm. long, 2.4-3.2 mm. in diameter, 1965] Helianthemum — Wilbur 295. ovoid-triquetrous, glabrous, rarely exceeding the calyx in length, 3- valved; each valve 2.4-3.2 mm. wide, ovate, acute. SEEDS 4-8, similar to the seeds of the chasmogamous flowers. FLOWERING: June-October. HABITAT: Dry hillsides, pine- oak forest, moist meadows, and mountain summits. DISTRI- BUTION: eastern Sonora into Chihuahua and southward in scattered localities into Guanajuato; also occurring in Guatemala and Honduras. (Map 13.) A few specimens examined during this study varied con- spicuously from the typical plants which are herbaceous, erect and tall, and have numerous ascending branches. These few variants were dwarf plants possessing shorter internodes, smaller leaves, and whose crowded flowers are mostly borne on shorter pedicels than those of the typical plants. However, plants having intermediate characters be- tween the two extremes have been encountered and were even mounted on the same herbarium sheet. A few other specimens (some from Mexico and all those collected from Guatemala and Honduras) were suffruticose but most had received stem injury. REPRESENTATIVE SPECIMENS: MEXICO: CHIHUAHUA: Ma- jalea, 18-20 Aug. 1935, LeSueur (ARIZ, F, GH, MO, SMU, TEX, UC); El Cima, LeSueur 795 (F, TEX); pine plains, base of the Sierra Madre, Pringle 1186 (F, GH, MIN, MSC, ND, NY, RSA, UC, US); Mesa, west of Hop Valley, Sierra Madre Mts., 17 Sept. 1903, Jones (nS, UC) ; Loreto, Rio Mayo, Gentry 2578 (ARIZ, F, GH, MO, UC); Summit Sta. Clara Mts., Shreve 7949 (ARIZ, F). DURANGO: 20 mi. w. of Durango, Water- fall 12580 (OKLA); El Fuerte, on railroad west of Durango, Pennell 18247 (F); Otinapa, Palmer 442 (GH). GUANAJUATO: Dolores Hidalgo- Guanajuato Rd., Kenoyer 2138 (GH). SONORA: region of the Rio de Bavispe, White 3434 (GH, MICH, PH), White 4406 (ARIZ, GH, MICH), White 3513 (ARIZ, MICH), White 4262 (MICH). ZACATECAS: road to Huejuquilla el Alto, Jal, McVaugh 17745 (MIcH). GUATEMALA: QUICHE: between Quiché & San Pedro Jacopilas, Standley 62459 (F); HUEHUETENANGO: hills east of Aguacatan Standley 62555 (F); Mala- catán, Smith 3286 (GH). HONDURAS: EL PARAÍSO: moist meadow, south of Guinope, Standley 14863 (F) ; about 2 mi. nw. of Guinope, Wil- liams & Molina R. 11524 (F) ; Cumbre northwest of Guinope, Standley et al 2023 (F), Standley et al 2115 (F). MORAZÁN: Tanque,. Rodriquez 859 (F); Zamorano, Rodriquez 2166 (F). 17. Helianthemum chihuahuense S. Wats. Helianthemum chihuahuense S. Wats., Proc. Am. Acad. 23: 268. 1888. 296 Rhodora [Vol. 67 Type: Pringle 1187, October 1887. (GH!). Isotypes: (DS, F, MIN, NY, US!). “On pine plains at the base of the Sierra Madre, Chi- huahua." Halimium chihuahuense (S. Wats.) Gross., Pflanzenreich 14 (IV. 193) : 45. 1903. Heteromeris chihuahuensis (S. Wats.) Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 169. 1921. (No basionym cited.) Suffruticose perennial, 11-32(45) cm. tall with few to numerous stems arising from a woody caudex (or rarely from subterranean rootstock). STEMS sometimes decumbent, usually ascending to erect, covered with stellate pubescence intermixed with simple villous hairs (about 1.0-1.5 mm. long). CAULINE LEAVES sessile or with petioles up to 1 mm. long; blade 7-22 mm. long, 2.0-7.5 mm. wide, elliptic or sometimes elliptic-oblanceolate, densely covered on both surfaces with both short stellate pubescence and simple villous hairs 0.5-1.5 mm. long (the stellate tomentum decreases as the leaf ages and hence the lower leaves sometimes appear lustrous with only villous hairs on both sides), sometimes tinged with purplish color ; midvein conspicuous and elevated beneath, secondary veins obscure beneath ; (usually with small and numerous fasciculate leaves in the axil of the principal ones; those often persisting throughout the season). FLOWERS: dimorphic with the chasmogamous and cleistogamous usually borne upon dif- ferent branches. Chasmogamous flowers few, glomerate or sometimes in a racemose cyme terminating the main stem and sometimes the major branches. Cleistogamous flowers l-few, glomerate at the end of the branches (sometimes appearing in the axils of the leaves). Pedi- cel and calyx covered with minute stellate pubescence and intermixed with simple villous hairs 0.5-1.5 mm long. Bracts 1.5-5.4 mm. long, 0.2-0.6 mm. wide, linear. CHASMOGAMOUS FLOWERS: pedicels 4-5 ram. long. OUTER SEPALS (free portion) 1.6-3.0 mm. long, 0.3-0.4 mm. wide, linear; INNER SEPALS 5.5-7.0 mm. long, 2.4-3.0 mm. wide, ovate, acute to acuminate. PETALS 7-10 mm. long, 5-6 mm. wide, obovate. STAMENS 24-30. PISTIL c. 1.8 mm. long, ovary c. 1.2 mm. long, c. 0.9 mm. in diameter, ovoid, glab- rous; style 0.5-0.8 mm. long; stigma c. 0.6 mm. wide, capitate. FRUITING PEDICELS up to 10 mm. long, articulate. FRUITING CALYX 6-8 mm. long, 2.8-3.5 mm. in diameter, ovoid. OUTER SEPALS (free portion) c. 3 mm. long, c. 0.4 mm. wide, linear; INNER SEPALS 6-8 mm. long, 3-4 mm. wide, ovate, acute to acuminate. CAPSULE c. 6 mm. long, c. 4 mm. in diameter, ovoid-triquetrous, glabrous, 3-valved; each valve c. 4 mm. wide, ovate, acute. SEEDS: none seen. CLEISTOGAMOUS FLOWERS: fruiting pedicels 1.6-5.0 mm. long, articu- late. OUTER SEPALS (free portion) 1.2-3.6 mm. long, 0.2-0.4 mm. wide, linear; INNER SEPALS 3.6-5.6 mm. long, 2.4-3.2 mm. wide, ovate, acute to acuminate. STAMENS 5-6. CAPSULE 2-5 mm. long, 1.6-3.2 mm. in diameter, ovoid-triquetrous, glabrous, 3-valved; each valve 1.6-3.2 mm. 1965] Helianthemum — Wilbur 297 wide, ovate, acute. SEEDS (2)4-11, inequilateral, smooth, when seeds moistened the membrane separable and the embryo visible. FLOWERING: June-October. HABITAT: pine-oak forest and wooded hillside. DISTRIBUTION: southwestern Chihuahua and southern Durango, and Hidalgo; also known from Guatemala, Honduras and Costa Rica. (Map 14.) The simple, villous cauline hairs, the suffruticose habit, the obscure secondary veins on the lower surface of the cauline leaves, and the smal] fasciculate axillary leaves that persist on the plant long after the principal leaves have fallen are all features which in combination would readily distinguish this species from other American species of Helianthemum. Helianthemum Pringlei is the only species included in this study whose leaves, when villous, strongly suggest those of H. chihuahuense. But although the flowers of both H. Pringlei and H. chihuahuense are dimorphic, they differ in arrangement. The chasmogamous flowers of H. chihua- huense are borne at the end of the main stem and sometimes the major branches, while its cleistogamous flowers are soli- tary or glomerate at the end of the branches; in H. Pringlei the dimorphic flowers are borne together at the end of the main stem and branches. The habit of H. chihuahuense is suffruticose while that of H. Pringlei is usually herbaceous. The cauline leaves of H. chihuahuense have small fasciculate leaves in their axil and this is rare indeed in H. Pringlei. South of Mexico H. chihuahuense has been often mis- takenly identified as H. glomeratum, but the latter species lacks the simple villous hairs and its chasmogamous flowers differ in being usually solitary at the end of the branches and branchlets and decidedly overtop the glomerate, cleis- togamous ones. REPRESENTATIVE SPECIMENS: MEXICO: CHIHUAHUA: Vi- cinity of Areponapuchic, slopes of Barranca de Urique, Knoblock 1312 (MICH, MSC); Memelichi, Rio Mayo, Standley 2791 (F) ; base of Sierra Madre, Pringle 1187 (DS, F,GH, MIN, ND, NY, RSA, US). DURANGO: Coyo- tes, Sierra Madre Occidental, Pennell 18269 (PH) ; 6 mi. southwest of El Salto, Waterfall 15486 (OKLA, SMU). HIDALGO: Trinidad, Pringle 11880 (GH). GUATEMALA: CHIMALTENANGO: near San Martin, Jilo- tepeque, Standley 64392 (F); HUEHUETENANGO: Sierra de los 298 Rhodora [Vol. 67 Cuchumatanes above Chiantla, Standley 65609 (F); 10 km. e. of Hue- huetenango, Standley 82067, 82085, & 82117 (F) ; nw. of Malacatancito, at km. 8 of the highway from Huehuetenango, Standley 82220 (F, GH) ; along trail between San Juan Atitlán & San Sebastian, Sierra de Cuchumatanes, Steyermark 52043 (F). JALAPA: Between Guisiltepeque & Potrero Carrillo, Steyermark 33026 (F, GH) ; QUICHÉ: along road south of Chichicastenango, Standley 62364 (F). SOLOLÁ: Volcan San Pedro, north facing slopes towards Lago de Atitlán, Steyermark 47191 (F). TOTONICAPÁN: along road between San Francisco and El Alto Momostenango, Standley 83983 (F). HONDURAS: MORAZÁN : slopes of Cerro de Uyuca, region of El Valle Encantado, Standley et al 947 (F), Standley & Williams 104 (F); Hoya Grande, along road between El Zamorano & Suyapa, Standley & Williams 1432 (F); near Hoya Grande, drainage of the Rio Yeguare, Williams & Molina R. 13278 (GH), Williams & Molina R. 10993 (GH, MO, MICH); Zamorano, Rod- riguez 554 (F), Rodriguez 618 (F). COSTA RICA: Los Friales, Valerio 1371 (F). 18. Helianthemum glomeratum (Lag.) Lag. ex Dunal Cistus glomeratus Lag., Gen. et Sp. Nov. 16. 1816. Type: not seen. “prope Acapulco et Cimapan in Nova Hispania.” Trichasterophyllum hyssopifolium Link, Jahr. Gewachskunde 3: 69. 1820. nom. nud., cited in synonymy by Grosser. Helianthemum glomeratum (Lag.) Lag. ex Dun., Prodr. 1: 269. 1824. Helianthemum obcordatum Moc. & Sessé ex Dun., Prodr. 1: 284. 1824. Type: MA on loan to F! “In Mexico.” Heteromeris mexicana Spach, Ann. Sci. Nat. 2nd ser. 6: 370. 1836. Helianthemum glomeratum “Lag.” cited as a questionable synonym; descr. Spach, Hist. Nat. Vég. Phanér. 6: 104. 1838. Type: not seen. “Cette plante croit au Mexique; nous l'avons décrite d'aprés des échantillons de l'herbier de M.P.B. Webb, les uns trouvés par M. Andrieux à Toluca, les autres envoyés par Pavon sous le nom de Cistus mexicanus, sans indication précise de localité." Taeniostema micranthum Spach, Ann. Sci. Nat. 2nd ser. 6: 371. 1836. nom. nud.; descr. Spach, Hook. Comp. Bot. Mag. 2: 289. 1836 [1 May 1837]. Type: not seen. Helianthemum glomeratum cited as questionable synonym in Annales and as a synonym in the Histoire. *I have described it from dried specimens of plants cultivated in the Jardin des Plantes at Paris." Cistus Mexicanus Sessé & Moc., Pl. Nov. Hisp. 180. 1894. Type: MA?, not seen. “ad Oppidum S. Rosae, prope Guanaxuatum." Halimium glomeratum (Lag.) Gross, Pflanzenreich 14 (IV. 193): 47. 1903. Heteromeris glomerata (Lag.) Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 169. 1921. (No basionym cited.) 1965] Helianthemum — Wilbur 299 Crocanthemum glomeratum (Lag.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 805. 1925. Suffruticose perennial, 1.0-5.5 (8.0) dm. tall with few to many stems arising from a woody, branched caudex and occasionally spreading by a horizontal, woody stem up to 1.5 dm. long. STEMS ascending to erect, stellate-tomentose, becoming glabrate and with numerous branches and ultimate branchlets. LEAVES on the current season's growth sessile to subsessile or with petiole up to 2 mm. long (if so, secondary veins prominent beneath); blade 1.0-2.5(3.5) em. long, 2-6(14) mm. wide; oblanceolate or rarely elliptic to obovate, hoary and stellate-tomentose on both surfaces; midvein mostly prominent and secondary veins obscure or very slightly elevated or prominent beneath; base of blade attenuate, apex rarely obtuse, subacute to acute; margin entire, nonrevolute. FLOWERS: dimorphic (chasmoga- mous and cleistogamous) but rarely with only cleistogamous flowers and lacking the chasmogamous ones, or with chasmogamous and but very few cleistogamous ones, Chasmogamous flowers (mostly solitary) at the tip of the branches and branchlets, borne on filiform pedicel (1.5)2-3 times longer than calyx and overtopping the subsessile cleis- togamous flowers. Cleistogamous flowers sub-sessile in axillary and terminal glomerules. Bracts 1.6-5.0 mm. long, 0.2-0.8 mm. wide, linear to oblanceolate, few to very numerous. Pedicel and calyx stellate- tomentulose. CHASMOGAMOUS FLOWERS: pedicels (7)10-20 mm. long, (1.5)2-3 times longer than calyx. OUTER SEPALS (free portion) 0.6-4.0 mm. long, 1/5-2/3 the length of the inner sepals, 0.2-0.4 mm. wide, linear; INNER SEPALS 3-7 mm. long, 1.8-3.2 mm. wide, ovate or ovate-lanceolate, acute or acuminate, sometimes with somewhat elevated veins at the back. PETALS 4-9 mm. long, 3.5-6.0 mm. wide, obovate, “pale yellow with slight orange flush at base." [Ball 4449 GH!] STAMENS 20-34. PISTIL 1.2-2.4 mm. long; ovary 0.6-1.6 mm. long, 0.5-0.9 mm. in diameter, ovoid, glabrous; style 0.5-1.0 mm. long; stigma 0.3-0.7 mm. wide, capitate. FRUITING CALYX 4.5-7.5 mm. long, 2.6-4.5 mm. in diameter, ovoid. CAPSULE 3.5-4.5 mm. long, 2-3 mm. in diameter, ovoid-triquetrous, glabrous, 3-valved; each valve 2-3 mm. wide, ovate, acute, slightly concave. SEEDS (6)10-20, ovoid, somewhat depressed on both sides to inequilateral, slightly pebbled, with thin separable membrane and the embryo becoming: visible when seeds moistened. CLEISTOGAMOUS FLOWERS: subsessile. FRUITING PEDICELS 0.3-3.6 mm. long, shorter than calyx. FRUITING CALYX 2.4-5.0 mm. long, 2-3(4) mm. in diameter, ovoid, sometimes with yellowish base. OUTER SEPALS (free portion) 0.6-3.0 mm. long, 0.2-0.4 mm. wide, linear; INNER SEPALS 2.4-5.0 mm. long, 1.2-3.6 mm. wide, ovate or ovate-lanceolate, acute or acuminate. STAMENS 3-6. CAPSULE 1.6-3.6 mm. long, 1.2-2.4(3.0) mm. in diameter, ovoid-triquetrous, glabrous, 3-valved; the valves 1.2-2.4(3.0) mm. wide, ovate, acute. slightlv concave. SEEDS (1)2-7(11), similar to those of the chasmogamous flowers. 300 Rhodora [Vol. 67 FLOWERING: throughout the year. HABITAT: grassland with scattered pine or oak or in open forests or on dry rocky hills. DISTRIBUTION: southwestern Texas, southern Baja California southward into Guatemala, (Map 15.) Helianthemum glomeratum has a wide distribution and has been more extensively collected than any other species of the genus found in Mexico or Central America. This is the only species mainly restricted to Mexico and Guatemala which extends northward in the United States. It is found in the Chisos Mountains of Brewster Co., Texas. The suffruticose habit and the densely glomerate, nearly sessile, cleistogamous flowers, often overtopped by the chas- mogamous flowers possessing pedicels (1.5)2-3 times longer than its calyx, are characteristics which taken col- lectively readily distinguish this species from all American species of Helianthemum. This species exhibits a remarkable variation in foliage, flowers, fruits and seeds. The foliage differs in blade size, length of petiole and extent of elevation of the secondary veins on the lower surface of the blade. The flowers differ in both kind and quantity; some specimens have both chas- mogamous and cleistogamous ones, and still other plants have cleistogamous flowers only. The fruits, especially those derived from the cleistogamous flowers, differ greatly in size and number of seeds. This variation appears to be nearly continuous. Following his two segregate genera, Spach listed two new species, Heteromeris mexicana and Taeniostema micran- thum. For both of these species Helianthemum glomeratum was cited as a questionable synonym. Heteromeris mexicana was based upon specimens with both chasmogamous and cleistogamous flowers. Spach's (Hist. Nat. Vég. Phanér. 6: 104. 1838.) account of the latter species leaves no doubt that Helianthemum glomeratum was the plant being de- scribed. Taeniostema micranthum was based on specimens with cleistogamous flowers only. Spach (Hist. Nat. Vég. Phanér., 6: 104. 1838.), however, equated the latter species with Helianthemum glomeratum. 1965] Helianthemum — Wilbur 301 REPRESENTATIVE SPECIMENS: UNITED STATES: TEXAS: Brewster Co., top of Mt. Emory, Chisos Mts., Warnock 20976 (TEX) ; Flat Top Mountain, Chisos Mountains, Warnock 1116 (Us). MEXICO: AGUASCALIENTES: road to Calvillo, 18 mi. west of Aguascalientes, MecVaugh & Koelz 114 (MICH). Sur Baja California: La Laguna, Sierra Laguna, Gentry 4378 (ARIZ, DS, GH) CHIAPAS: Tstapa an son- nigen Hanger, Seler 2086 (F, GH). CHIHUAHUA: Santa Eulalia Mts., Pringle 300 (F, GH, MICH, PENN, RSA). DURANGO: Santiago Papas- quiaro, Palmer 56 (F, GH, UC). FEDERAL DISTRICT: lava fields near Eslaba, Pringle 11371 (F, GH, SMU). GUANAJUATO: Puerto Nieto near San Miguel Allende, Kenoyer 2049 (GH). GUERRERO: west of Chilpan- cingo, Richands et al 3353 (SMU). HIDALGO: Trinidad, Pringle 11881 (GH). JALISCO: hillside above Etzatlán, Pringle 11372 (F, GH, MSC, SMU); hills near Guadalajara, Pringle 2361 (ARIZ, F, GH, MSC, NY, PH, RSA, UC); San Sebastián, w. to Mascota, Mexia 1414 (DS, F, GH, MICH, MIN, UC). MEXICO: Dist. Temascaltepec, Hinton 3033 (F, UC), Hinton 8401 (F, GH), Hinton 8990 (DS, F, ILL, MICH, SMU, TEX), Hinton et al 2720 (UC), Hinton et al 8916 (ARIZ, F, GH, TEX). MEXICO-PUEBLA: Ixtaccihuatl, Purpus 1815 (F, UC). MICHOACÁN: grassy hillslopes, Zitacuaro-Las Canoas, Dist. Zitacuaro, Hinton et al 13548 (F, ILL, MICH, UC); Tancitaro, Dist. Uruapan, Hinton et al 15733 (ARIZ, GH, TEX). MORELOS: Cuernavaca, Kenoyer A467. NAYARIT: Tepic, Palmer 2017 (GH). NUEVO LEON: about 15 mi. sw. of Galeana, Mueller & Mueller 980 (F, GH, MICH, TEX). OAXACA: Sierra de San Felipe, Smith 809 (F). PUEBLA: San Manuel de la Sierra, Sierra Negra, Balls B4449 (GH, UC). QUERETARO: 15 mi. se. of San Juan del Rio, Waterfall 13978 (OKLA, SMU). SAN LUIS POTOSI: Alvarez, Palmer 56 (F, GH, UC) ; near San Luis Potosi, Palmer 35 (GH, UC). SINALOA: Sierra Monterey, Gentry 5891 (ARIZ, DS, GH, MICH). SONORA: Alamos, Palmer 342 (GH). TLAXCALA: Sta. Ana Chiantauapan, Arsene 1738 (MO). ZACATECAS: Zacatecas, Purpus 502 (POM, UC). GUATEMALA: CHIMALTENANGO: Johnston 1773 (F). HUEHUETENANGO: Sierra de Los Cuchumatanes, above Chiantla, Standley 65620 (F), 65604 (F); Rio Pucal, Standley 65826 (F, GH, MICH). JALAPA: hills between Guisiltepeque and Potrero Carrillo, Steyermark 33024 (F). QUEZALTENANGO: Cerro La Pedrera, south of Quezaltenango, Standley 66460 (F). QUICHE: without exact locality, Aguilar 1444 (F). SAN MARCOS: Slopes of Volcán Tajumulco, Steyermark 36917 (F). ZACAPA: Sierra de Las Minas, Steyermark 42754 (F, UC). 19. Helianthemum rosmarinifolium Pursh Helianthemum rosmarinifolium Pursh, Fl. Am. Sept. 2: 364. 1814. Type: “Enslen. July. v.s. in Herb. Enslen.", not seen; “In pine barrens, Georgia.” not Helianthemum rosmarinifolium Presl, Symbolae Botanicae 1: 32. Tab. 21. 1832, which was from Pales- tine. Heteromeris polifolia Spach, Hook. Comp. Bot. Mag. 2: 291. 1836 302 Rhodora [Vol. 67 [1 May 1837]. Type: “. .. in the Province of Texas by Mr. Drummond (third collection, No. 20!), as also by M. Berlandier.", not seen. Isolectotype: Drummond, 3rd collection No. 20 (GH, NY!) and Berlandier (US-1169410!) also seen. Anthelis (Horanthes) podanisia Raf., New Fl. N. Am. 3: 30. 1836 [1838]. Horanthes podanisia (Raf.) Raf., Sylva Tell. 133. 1838. Helianthemum polifolium (Spach) T. & G., Fl. N. Am. 1: 151. 1838; not Helianthemum polifolium DC., Lam. & DC. Fl. Fr. 4 (2): 823. 1815. Helianthemum capitatum Nutt. ex Engelm. & Gray, Bost. Journ. Nat. Hist. 5: 212. 1847. A substitute for the preoccupied Helianthemum polifolium (Spach) T. & G. Halimium rosmarinifolium (Pursh) Gross, Pflanzenreich 14 (IV. 193): 49. 1903. Halimium domingense Urb., Symb. Ant. 7: 286. 1912. Type: von Tuerckheim 3430, not seen. Isotype: von Tuerckheim 3430 (NY!). “in Sto. Domingo prope Constanza in Valle nuevo, 2200 m." Halimium stenophyllum Urb., Symb. Ant. 7: 524. 1913. Type: Fuertes 1919, not seen. Isotypes: Fuertes 1919 (NY, US!). *in Sto. Domingo prov. Azua ad Las Canitas 1400 m. alt." Crocanthemum domingense (Urb.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. Crocanthemum stenophyllum (Urb.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. Crocanthemum rosmarinifolium (Pursh) Janchen, Nat. Pflanzenfam. ed. 2. 21: 307. 1925. Perennial herb (13)20-40(51) cm. tall with several stems arising from a woody caudex. STEMS ascending to erect, stellate-tomentose, with numerous ascending branches. BASAL LEAVES sometimes present, if so, 10-22 mm. long, 3-5 mm. wide, obovate. CAULINE LEAVES: petiole 1-4 mm. long; blade (4.5) 14-38(48) mm. long, 5-14 times as long as wide, (0.7)2.3-5.2(7.8) mm. wide, oblanceolate to elliptic to linear- lanceolate, green and stellate-pubescent above, hoary and stellate- tomentose beneath; midvein elevated and secondary veins but slightly elevated beneath; base of blade attenuate, apex subacute to acute; margin entire, somewhat revolute. FLOWERS: dimorphic (both chas- mogamous and cleistogamous). Chasmogamous flowers solitary at the tip of the main branches borne on filiform pedicel (2-3 times longer than calyx) overtopping the subsessile, cleistogamous flowers. Cleistog- amous flowers subsessile in axillary and terminal glomerules. Bracts up to 2 mm. long and c. 0.3 mm. wide, linear. Pedicel and calyx stellate- tomentose. CHASMOGAMOUS FLOWERS: pedicels 10-22 mm. long, 2-4 times longer than calyx. OUTER SEPALS (free portion) 1.3-2.0 (2.5) mm. long, 0.2-0.3 mm. wide, linear; INNER SEPALS 2.5-4.3 mm. long, 1.3-2.0 mm. wide, 1965] Helianthemum — Wilbur 303 ovate, acute and slightly oblique. PETALS 4-5 (6.4) mm. long, 3.6-4.8 (5.4) mm. wide, yellow, obovate. STAMENS 15-24. PISTIL 0.8-1.5 mm. long; ovary 0.5-1.0 mm. long, 0.4-0.7 mm. in diameter, ovoid, glabrous; style 0.3-0.4 mm. long; stigma c. 0.8 mm. wide, capitate. FRUITING CALYX: 2.5-4.5 mm. long, 1.5-3.0 mm. in diameter, ovoid to nearly pyriform, with somewhat yellowish base. CAPSULE 2-3 mm. long, 1.4-1.8 mm. in diameter, ovoid, glabrous, 3-valved; each valve 1.0-1.4 mm. wide, ovate- elliptic, acute, separating to the middle at maturity. SEEDS 1-3(6), ovoid or somewhat appressed on both sides to inequilateral, smooth and when moistened with a thin separable membrane and then the embryo visible. CLEISTOGAMOUS FLOWERS: subsessile. FRUITING PEDICELS 0.5-3.0 mm. long. FRUITING CALYX 1.5-1.8 mm. long, 1.2-1.5 mm. in diameter, some- what obovoid with yellowish base. OUTER SEPALS (free portion) 0.5-1.0 mm. long, c. 0.2 mm. wide, linear; INNER SEPALS 1.5-1.8 mm. long, 1.0-1.5 mm. wide, ovate, acute. STAMENS 3-5. CAPSULE 1.3-1.7 mm. long, 1.0-1.3 mm. in diameter, somewhat obovoid, glabrous, 3-valved; each valve 0.8-1.0 mm. wide, somewhat obovate, subacute, separating to the middle at maturity. SEEDS 1(2), similar to those of the chasmoga- mous flowers. FLOWERING: May-July. HABITAT: open sandy pinelands, sandhills, and fields. DISTRIBUTION: North Carolina south- ward into northern Florida and westward into the eastern half of Texas; also found in the West Indian Island of Santo Domingo. (Map 3.) Helianthemum rosmarinifolium can be readily recognized by its solitary chasmogamous flowers borne at the top of the main branches on filiform pedicels 2-4 times longer than the calyx at anthesis, its cleistogamous capsules less than 1.5 mm. in diameter with the valves separating only to the middle, and the midcauline leaves 5-14 times as long as wide. In spite of differences in habit and foliage, this taxon ap- pears most closely related to H. glomeratum. Both of these taxa possess terminal, solitary, chasmogamous flowers on filiform pedicels 2-4 times longer than their calyces, over- topping the subsessile cleistogamous flowers which occur in axillary and terminal glomerules. The fruit and associated sepals derived from the cleistogamous flowers are quite vari- able in H. glomeratum and even one of its races approaches those of H. rosmarinifolium in shape and size of calyx, num- ber of seeds, and seed characteristics. 304 Rhodora [Vol. 67 Helianthemum argenteum is similar to H. glomeratum and H. rosmarinifolium in having solitary, terminal, chas- mogamous flowers on filiform pedicel (1) 1.5-2 times longer than the calyx. But it can be readily distinguished by its de- cumbent habit and its silvery pubescence. The axillary, ses- sile to subsessile, solitary (rarely two) cleistogamous flowers in the axil of the narrow (1.2 mm. wide or less) sessile leaves also serve to make it readily separable. Although neither the type of the species nor a photograph of it has been available in this study, the original description seems sufficiently diagnostic to make it certain that this is the plant that Pursh described as H. rosmarinifolium. This species has previously been considered endemic to the southeastern United States. However, Urban (Symb. Ant. 7: 286. 1912.) described a supposedly new species col- lected from the West Indies (Santo Domingo) naming it Halimium domingense. One year later, Urban (Symb. Ant. 7: 525. 1913.) proposed another species as new from the same general area, calling it Halimium stenophyllum. The isotypes of both of Urban's species, Halimium domin- gense (NY!) and Halimium stenophyllum (NY! US!), were studied, and in spite of the geographical isolation, no dif- ference could be found except that the Dominican specimens are not quite so erect as specimens collected in the United States. Therefore both of Urban's proposed species are here treated as synonyms of H. rosmarinifolium. This is the only species of Helianthemum known from the West Indies. The distribution of Sabatia calycina (Lam.) Heller is rather similar as both species range throughout the southern Coastal Plain and reappear in central Hispaniola. This Helianthemum is not known from peninsular Florida while the Sabatia ranges well into the peninsula and is known also from the Oriente Province of Cuba. REPRESENTATIVE SPECIMENS: NORTH CAROLINA: Richmond Co., roadside, Hoffman, Gupton 1610 (NCU). SOUTH CAROLINA: Berke- ley Co., Gravel Hill Lake, ene. of Bonneau, Ahles 30788 (DUKE, NCU); Edgefield Co. 3 mi. ene. of Trenton, Radford 26416 (DUKE, NCU). CEORGIA: Baker Co., 10 mi. sw of Newton, Thorne 4512 (GA, IA, CU); Candler Co., 10 mi. n. of Metter, 25 June 1950, Sargent (GH, KANU, 1965] Helianthemum — Wilbur 305 MIN, NCSC, OKL, WIS, WVA) ; Decatur Co., near Bainbridge, Curtiss 6477 (DS, F, GH, ILL, ISC, MIN, MO, NY, SMU, UC, US) ; McIntosh Co., along the Altamaha River near Darien, Correll 5470 (DUKE, GA, NCSC, TENN); Screven Co., Oliver, Curtiss 6838 (CU, GH, ILL, MIN, MO, NY). FLORIDA: Gadsden Co., without exact locality, Boomhour 627 (DUKE). MISSIS- SIPPI: Clarke Co., De Soto, Mohr (Ny, Us); Lowndes Co.: Columbus, Mohr (MO, NY). ARKANSAS: Drew Co., Wilmar, Demaree 24526 (SMU); Johnson Co., Piney, Palmer 8162 (MO, PH, US). OKLAHOMA: Marshall Co., Lake Texoma, Goodman 7044 (DUKE, OKL). TEXAS: Bas- trop Co., about 5 mi. s. of Bastrop, Lundell & Lundell 8975 (Ds, GH, MICH, NY, SMU); Brazos Co., Bryan, Palmer 7796 (MO, NY, US) ; Dallas Co., near Dallas, Curtiss 228 (CU, F, GH, KANU, MICH, MIN, NY, OKL); Tarrant Co., without exact locality, Ruth 496 (CU, F, GH, ILL, ISC, MICH, MSC, NY, PH, SMU, US, WIS); Waller Co., Hempstead, Hall 30 (F, MO, NY, US). SANTO DOMINGO: Prope Constanza in Valle nuevo, 2200 m., Tuerckheim 3430 (NY); prov. Azua ad Las Caffitas 1400 m. alt., Fuertes 1919 (NY, US). 20. Helianthemum argenteum Hemsl. Helianthemum argenteum Hemsl., Diagn. Pl. Nov. Mex. et Centr.- Amer. part 2: 20. 1879, Type: Parry & Palmer 29. (K) Isotypes: (GH, NY, US!) “Mexico: in regione San Luis Potosi, alt. 6000- 8000 ped." Halimium argenteum (Hemsl.) Gross., Pflanzenreich 14 (IV. 193) : 47. 1903. Heteromeris argentea (Hemsl.) Ponzo, Nuovo Gior. Bot. Ital. n.s. 28: 170. 1921. (No basionym cited.) Crocanthemum argenteum (Hemsl.) Janchen, Nat. Pflanzenfam. ed. 2. 21: 305. 1925. Suffruticose perennial, 5-15 cm. tall, arising from a thick woody root. STEMS short, erect, with numerous, decumbent to erect branches clothed with fine, silvery, appressed elongate tomentum, LEAVES (on the current season's growth) estipulate but with smaller leaves in their axil, sessile; blade 5-7 mm. long, 0.8-1.2 mm. wide; narrowly elliptic to linear, densely covered with fine, silvery, appressed elongate tomen- tum on both surfaces; midvein but slightly elevated and secondary veins obscure beneath; base of blade somewhat attenuate, apex acute to pointed and somewhat lustrous; margin entire, non-revolute. FLOWERS dimorphic (both chasmogamous and cleistogamous) : the chas- mogamous solitary at the end of the branches (mostly accompanied by two sessile cleistogamous flowers) borne on pedicel (1)1.5-2 times longer than calyx. Cleistogamous flowers solitary (sometimes two together), sessile in the leaf axil. Pedicel and calyx covered with fine, silvery, depressed elongate tomentum. CHASMOGAMOUS FLOWERS: Pedicels 5-7 mm. long. OUTER SEPALS (free portion) 1.5-2.0 mm. long, about half the length of the inner sepals, 306 Rhodora [Vol. 67 Maps 1-3. Map 1. H. carolinianum. Map 2. H. georgianum. Map 9. H. rosmarinifolium with inset of the island of Hispaniola. Maps 4-6. Map 4. H. canadense. Map 5. H. propinquum. Map 6. H. Bicknellii. 308 Rhodora [Vol. 67 Maps 7-11. Map 7. H. arenicola (open circles) and H. Nashii (solid dots). Map 8. H corymbosum. Map 9. H. dumosum. Map 10. H. Greenei. Map. 11. H. scoparium. 0.3-0.4 mm. wide, linear; INNER SEPALS 3-4 mm. long, 1.2-1.6 mm. wide, ovate-lanceolate, acuminate. COROLLA yellow, petals 3-4.5 mm. long, 3.0-3.5 mm. wide, obovate. STAMENS 10-15. PISTIL 1.2-1.6 mm. long; ovary 0.8-1.2 mm. long, 0.4-0.5 mm. in diameter, narrowly ovoid; glab- rous; style 0.1-0.2 mm. long; stigma about 0.2 mm. wide. FRUITING PEDICELS 7-10 mm. long. FRUITING CALYX narrowly ovoid. OUTER SEPALS (free portion) 1.5-2.5 mm. long, about half that of the inner, 0.3-0.4 mm. wide, linear; inner sepals 5.0-5.5 mm. long, 1.5-2.0 mm. wide, ovate-lanceolate, acuminate, about twice longer than wide. CAPSULE 3.6-4.0 mm. long, about twice as long as wide, 1.2-1.6 mm. in diameter, narrowly ovoid-triquetrous, glabrous, very thin-walled, 3-valved; each valve 1.2-1.6 mm. wide, ovate-lanceolate, acute, somewhat spreading at maturity. SEEDS 3, ovoid or nearly so, covered with white papillae borne on the thin separable membrane, the embryo visible when seed moistened. CLEISTOGAMOUS FLOWERS: Sessile to subsessile. FRUITING PEDICELS up to 0.8 mm. long. FRUITING CALYX 2.8-3.6 mm, long, 1.4-1.7 mm. in 1965] Helianthemum — Wilbur 309 Maps 12-15. Map. 12. H. Coulteri. Map 13. H. Pringlei. Map 14. H. chihuahuense. Map 15. H. glomeratum. (Open circles on all maps indicate that the exact locality is unknown.) 310 Rhodora [ Vol. 67 Maps 16-17. Map 16. H. nutans (large dots) and H. patens (small dots). Map 17. H. concolor (small dots) and H. argenteum (large dot). diameter, ovoid. OUTER SEPALS (free portion) 1.2-1.6 mm. long, about half the length of the inner sepals, 0.2-0.3 mm. wide, linear; INNER SEPALS 2.8-3.6 mm. long, about twice as long as wide, 1.0-1.4 mm. wide, ovate-lanceolate, acuminate. STAMENS 3. CAPSULE 1.2-1.4 mm. long, about twice longer than its diameter, 0.5-0.7 mm. in diameter, nar- rowly ovoid-triquetrous, very thin-walled, glabrous, 3-valved; each valve 0.5-0.7 mm. wide, ovate-lanceolate, acute. SEEDS 1-2, other charac- ters similar to those of the chasmogamous seeds, 1965] Helianthemum — Wilbur 311 FLOWERING: June. HABITAT: rocky hillsides and summits. DISTRIBUTION: known only from the State of San Luis Po- tosi, Mexico. (Map 17, large dot). Helianthemum argenteum is rare and has been little col- lected. Only three collections were seen by us. It is the only Mexican species of Helianthemum with the following com- bination of features: an elliptic-linear leaf blade, less than 1.5 mm. wide and 5-7 times longer than wide covered with depressed silvery pubescence and with solitary (or rarely two together) cleistogamous flowers which are sessile to sub- sessile in the leaf axil and whose chasmogamous flowers are solitary at the end of the branches. SPECIMENS EXAMINED: MEXICO: SAN LUIS POTOSI: chiefly in the region of San Luis Potosi, Parry & Palmer 29 (GH, NY, US); Cerro Tepetate, southwest of San Luis Potosi, Pennell 17656 (PH), S. Miquelito Mts., Schaffner 606 (PH). DOUBTFUL AND EXCLUDED NAMES Anthelis (Horanthes) arenaria Raf., New Fl. N. Am. 3: 31. 1836. [1838]; later Horanthes arenaria (Raf.) Raf., Sylva Tell. 133. 1838. The original description of Rafinesque’s species is quoted here in full: “551 Anthelis (Horanthes) arenaria Raf. many pro- cumbent stems nearly smooth. Leaves linear acute sometimes ciliate, pedicels axillary uniflore equal to leaves, 3 sepals mem- branaceous rufous smooth trinerve ovate acute, 2 linear subciliate — in sands of Pine barrens in South New Jersey and Florida, first discovered by Kin in 1801. Root large deep, many short diffuse stems, leaves small, flowers rather large.” There is not a single species of Helianthemum occurring in both “South New Jersey and Florida." However the following charac- teristics: “. . . leaves. . . sometimes ciliate . . . uniflore . . . in sands of Pine barrens in South New Jersey . . . flowers rather large” make it probable that Rafinesque was redescribing at least in part what was already known as Helianthemum canadense (L.) Michx. Cistus Rosmarinifolius Mog. E Sessé, Fl. Mex. 130. 1894. Type: ? MA, not seen. “in oppido S. Andrés de el Valle haud procul Temascal- tepec, ubi vulgo Romerillo audit.” Helianthemum astylum Moc. et Sessé ex Dunal, Prodr. 1: 284. 1824. Treated as a synonym of Halimium glomeratum (Lag.) Grosser (Pflanzenreich 14 (IV. 193) : 47. 1903.) And previously Hemsley (Biol. Cent. Am. 1: 47. 1879) considered it a probable synonym of “one of the states of H. glomeratum.” Helianthemum tripetalum Moc E Sessé ex Dun., Prodr. 1: 284. 1824. 312 Rhodora [Vol. 67 — Lechea tripetala (Moc. & Sessé ex Dun.) Britt. Bull. Torrey Club 21: 252. 1894. Horanthes tripetala attributed by Merrill (Ind. Raf. 169. 1949.) to Raf., New Fl. N. Am. 3: 30. 1836 [1838]. who under the genus Anthelis and subg. Horanthes merely indicates that *H. tripetala of Mexico is probably a Lechea!" The *H." doubtless stood for Helianthemum and referred to the publication of Dunal cited above. DEPARTMENT OF BOTANY, DUKE UNIVERSITY DURHAM, NORTH CAROLINA THE EXTENDED DISTRIBUTION OF ERAGROSTIS TRACYI FROM SANIBEL ISLAND, FLORIDA' A. S. Hitchcock in 1934 (1), based the description of Eragrostis tracyi on specimens of S. M. Tracy (no. 7168) collected on Sanibel Island, Florida, 1901. This endemic of narrow distribution was collected again from the type lo- cality by George R. Cooley (no. 2608) (1954) (2). Since its discovery by Tracy in 1900, a period of 64 years, the species has been known only from Sanibel Island (3). In 1964, the species was twice encountered in other locali- ties. The 20th of April, collection no. 27403 was made on Mound Key, Estero Bay, Lee Co., only about 15 mi. south- east of Sanibel. Two pinkish plants 5 cm. tall, with basal branches and spreading panicles were growing in dry, com- pacted soil of the wooded trailside on the top of the Mound. They were the only ones available; pressure of time pre- vented wider search. The 6th of October a population was found on Longboat Key, Sarasota Co., about 60 mi. north of Sanibel; collection no. 27566. It is established on a low terrace of the Gulf of Mexico on an extensive shell-beach with a remarkable asso- ciation of Sabal palmetto, Agave decipiens and Juniperus silicicola. Frequency and size of the shrubs in the under- growth give the appearance of a primary condition, or at ‘Contribution 19, Botanical Laboratories, University of South Florida. 1965] Eragrostis tracyi — Lakela 313 least absence of recent disturbance. Conspicuous elements are Yucca aloifolia, Harrisia Aboriginum, Opuntia Dillenii, Rapanea guianensis, Randia aculeata, and Forestiera poru- losa. In the strand vegetation Eragrostis tracyi was spotted by its panicles among Sporobolus virginicus, Sesuvium portu- lacastrum, Flaveria floridana and Amaranthus australis. This habitat presents a sharp contrast to that of the bare, disturbed soils of Sanibel Island. However, strand vegetation is subject to disturbance by various factors. The 10th of Feb- ruary the site was checked for signs of perennial growth for added information on the life duration of Eragrostis tracyi. The grasses were prostrate and matted. With scattered de- bris it appeared as if the area had been flooded. Many tufts of Eragrostis were dead, but in others tips of new green leaves were visible among the old dead ones. Two tufts were potted in the greenhouse. The growth of new leaves con- tinues and, obviously, Eragrostis tracyi perennates from un- derground parts. OLGA LAKELA, UNIVERSITY OF SOUTH FLORIDA, TAMPA LITERATURE CITED 1. HiTCHCOCK, A. S. 1934. Eragrostis Tracyi. Am. Jour. Bot. 21: 130. f.1. 2. COOLEY, GEORGE R. 1954. Vegetation of Sanibel Island. Rhodora 57: 269-289. 3. HITCHCOCK, A. S. rev. AGNES CHASE, 1950. Manual of the Grasses of the United States. pp. 162-163. Erratum for Rhodora Volume 67, Number 770 d p. 183 Fig. 1 is Plate 1311 Volume 67, No, 771 including pages 217-313, issued September 28, 1965. FARLOW REFERENCE LIBRARY Dote JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL ) STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER \ Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. Vol. 67 October-December, 1965 No. 772 CONTENTS: Nomenclatural Changes and Additions to Britton and Wilson's *Flora of Porto Rico and the Virgin Islands" Brother Alain LAO ier ................. e terrre erro eno eren rores 315 The Status of Carex incomperta Bickn. E E E er EE 9361 The Distribution Patterns of Northern and Southern Elements in the Flora of Newfoundland RK tegen, Een Eed 363 The Mountain Element in the Flora of the Peninsula ot Virginia. A, M, Harvill Jaeger 393 i (Continued on Inside Cover) The Nef England Botanical Club, Inc. Botanical Museum, Oxford St., Cambridge 38, Mass. CONTENTS: — continued Discovery of the Massachusetts Fern in Wisconsin Thomas G. Hartley |... esee 399 A Third Subspecies in the Artemisia tridentata Complex Alan A. Beetle and Alvin Young ........ ees 405 Geocaulon lividum in New England Stuart K. Harris «eese 407 Further Notes on Chamaecyparis thyoides in New Hampshire Henry I. Baldwin .......... 409 Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 67 October-December, 1965 No. 772 NOMENCLATURAL CHANGES AND ADDITIONS TO BRITTON AND WILSON’S “FLORA OF PORTO RICO AND THE VIRGIN ISLANDS" BROTHER ALAIN LIOGIER, (") Our knowledge of the flora of Puerto Rico and the Virgin Islands comes from its main and most recent source, the Flora published by N. L. Britton and P. Wilson between 1923 and 1930, as part of the Scientifie Survey of Porto Rieo and the Virgin Islands, sponsored by the New York Academy of Sciences. Already, as the last parts were being prepared, some plants not recorded in the previous parts had to be cited in an Appendix (1926) and a Supplement (1930). Many of the scientific names used by Britton and Wilson are now obsolete, either because the authors had a different concept of the generic lines, used a different Code of Nomenclature, or because new monographs having been published, some nomenclatural changes have proved neces- sary; also some species have been collected in the area for the first time, and have to be added to the flora; and a few species have been described as new from the Island of Puerto Rico. The species count in Britton and Wilson's Flora was 2464; our new count goes up to 2582, and the author is convinced that there are still some species to be collected in the area, that are not recorded so far. In this paper, I shall cite all the changes in nomenclature, 'Henorary Curator of West Indian Botany, The New York Botanical Garden. 315 316 Rhodora [Vol. 67 new records and new species, and also the reductions to synonymy made necessary since the publication of the Flora. Whenever possible, the bibliography will be given, for a family, a genus or a species. The pages refer to Brit- ton & Wilson’s Flora, as part of the Scientific Survey, Vols. Vand VI. The first name cited is the correct name as here interpreted, and the name in parenthesis and in italics is the name applied by Britton & Wilson (except that the par- enthetical authorities have been removed) in the Flora, or in a few occasions the name used in other Floras. For new rec- ords and new species, the locality is given, unless the plant is fairly common in the Island. For most cases, no attempt has been made by the author to make a critical study of each group; this kind of work would require several years of study and would amount to the preparation of a new Flora. Only occasionally has attention been given to the changes in nomenclature in the cultivated plants. I am grateful to the New York Botanical Garden for the facilities given to me both in the Herbarium and in the Li- brary, and also for financial support for some field work in Puerto Rico; my thanks also to the University of Puerto Rico, particularly the College of Agriculture and Mechanical Arts at Mayagiiez, for the facilities in collecting and drying the plants, extended to me through the kindness of Dr. José A. Ramos, BASIC BIBLIOGRAPHY LANJOUW, J., et al.—International Code of Botanical Nomenclature. 1961, REGNUM VEGETABILE. Vol. 36. 1964. LEON, HNO. & ALAIN, HNo.—Flora de Cuba. 5 vols. 1947-1962. STANDLEY, P. & STEYERMARK, J.A. et al.—Flora of Guatemala, 1946- 1962. Woopson, R.E., SHERY, R.W. et al—Flora of Panama, 1943-1962. LITTLE, E. & WADSWORTH, F.—Common trees of Puerto Rico and the Virgin Islands. 1964. WILLIAMS, R.O., et al.—Flora of Trinidad and Tobago. 1928-1955. PULLE. A.A. et al.—Flora of Surinam. 1965] Flora of Porto Rico — Liogier Ota VOLUME V OF THE SCIENTIFIC SURVEY TYPHACEAE p. 9 Typha domingensis (Pers.) Kunth (Typha angustifolia L., as to the Antillean plant). POTAMOGETONACEAE (Zannichelliaceae; includes Cymodoceaceae) p. 10 Ruppia maritima L. includes R. anomala Ostenfeld. as a syn- onym. p. 11 Potamogeton insulanus Hagstr. (Potamogeton epihydrus Raf., as to the Puerto-Rican plant). p. 11 Potamogeton nodosus Poir. (Potamogeton fluitans Roth) p.12 Syringodium filiforme Kurtz (Cymodocea manatorum Aschers.) p. 12 Halodule beaudettii (Den Hartog) Den Hartog. To be added: St. Thomas, St. Croix. See Blumea XII: 289-312. 1964. ALISMATACEAE (Alismaceae) Ref.: Rhodora 57: 133-156; 178-188; 202-212. 1955. p. 13-14 Echinodorus tenellus (Mart.) Buch. var. latifolius (Seu- bert) Fassett (Helianthium tenellum Britt.) p. 14 Echinodorus berteroi (Spreng.) Fassett var. berteroi (Echino- dorus cordifolius Griseb.) HYDROCHARITACEAE p. 17 Limnobium stoloniferum (G.F.W. Meyer) Griseb. (Hydromis- tria stolonifera G.F.W. Meyer) GRAMINEAE (Poaceae) Ref.: HITCHCOCK, A. E CHASE, A.—Manual of the Grasses of the West Indies. 1956. p. 23 Hackelochloa granularis (L.) Kuntze ( Rytilix granularis Skeels) p. 24 Andropogon brevifolius Sw. (Schizachyrium brevifolium Nees) p. 24 Andropogon gracilis Spreng. (Schizachyrium gracile Nash) p. 25 Andropogon hirtiflorus (Nees) Kunth (Schizachyrium dom- ingense Nash) p. 25 Andropogon semiberbis (Nees) Kunth (Schizachyrium semi- berbis Nees) p. 26 Andropogon annulatus Forsk. (Dichanthium annulatum Stapf.) p. 27 Andropogon saccharoides Sw. (Amphilophis saccharoides Nash) p. 27 Andropogon pertusus (L.) Willd. (Amphilophis ischaemum Nash) 3518 Rhodora [Vol. 67 To be added: Andropogon condylotrichus Hochst. Bayamon, Andro- p. p. p. p. 28 . 29 . 29 . 29 . 31 . 32 . 92 . 98 33 34 34 pogon virginicus L. Maricao. Vetiveria zizanioides (L.) Nash ex Small (Anatherum zizani- oides H. & C.) Sorghum halepense (L.) Pers. (Holcus halepensis L.) Sorghum saccharatum (L.) Moench. (Holcus sorghum L.) Sorghastrum setosum (Griseb.) Hitche. (Sorghastrum parvi- florum H. & C.) Tragus berteronianus Schult. (Nazia aliena Scribn.) Trichachne insularis (L.) Nees (Valota insularis Chase) Trichachne eggersii (Hack.) Henr. (Valota eggersii H. & C.) Digitaria adscendens (HBK.) Henr. (Syntherisma sanguinalis Dulac) Digitaria violascens Link (Syntherisma chinensis Hitche.) Digitaria ischaemum (Schreb.) Schreb. (Syntherisma ischae- mum Nash) Digitaria horizontalis Willd. (Syntherisma digitata Hitchc.) Digitaria argillacea (Hitchc. & Chase) Fern. (Syntherisma argillacea H. & C.) Digitaria panicea (Sw.) Urb. (Syntherisma filiformis Nash) To be added: Digitaria villosa (Walt.) Pers. Tortuguero, San Juan, p. 35 37 . 38 . 98 . 98 . 99 . 39 . 43 . 43 p. 48 Campo Alegre. Eriochloa polystachya HBK. (Eriochloa subglabra Hitchc.) Paspalum boscianum Flügge (Paspalum melanospermum Desv.) See Sci. Surv. VI: 524. Paspalum rupestre Trin. includes Paspalum leoninum Chase, as a synonym. Paspalum molle Poir. (Paspalum portoricense Nash) Paspalum saugetii Chase (Paspalum poiretii R. & S. as to the Puerto-Rican plant). Paspalum blodgetii Chapm. (Paspalum simpsoni Nash) Paspalur: laxum Lam. (Paspalum glabrum Poir.) Setaria utowanaea (Scribn.) Pilger (Panicum utowanaeum Scribn.) Panicum muticum Forsk (Panicum barbinode Trin.) Panicum angustifolium Ell. (Panicum fusiforme Hitchc.) To be added: Panicum strigosum Muhl. Santana (Vélez 984) Panicum capillare L. St. Croix. Lasiacis patentiflora Hitche. & Chase. To be added: St. Croix. Oplismenus setarius (Lam.) R. & S. To be added: Humacao. Echinochloa colona (L.) Link (Echinochloa colonum, incorrect spelling) Echinochloa crus-pavonis (HBK.) Schltr. (Echinochloa erus- galli Beauv.) 1965] Flora of Porto Rico — Liogier 319 Setaria barbata (Lam.) Kunth (Chaetochloa barbata Hitche. & Chase) Setaria geniculata (Lam.) Beauv. (Chaetochloa geniculata Millsp. & Chase) Setaria tenacissima Schrad. (Chaetochloa tenacissima Hitche. & Chase) Setaria magna Griseb. (Chaetochloa magna Scribn.) Setaria setosa (Sw.) Beauv. (Chaetochloa setosa Seribn.) Setaria rariflora Mikan (Chaetochloa rariflora Scribn.) Setaria vulpiseta (Lam.) R. & S. (Chaetochloa vulpiseta Hitche. & Chase) added: Setaria lutescens (Weigel) F. T. Hubb. Maricao and Guilarte. Setaria pradana (León) León. Cayo Muer- tos, Mona Island. Setaria leiophylla (Nees) Kunth. Ponce, Desecheo Island. Pennisetum purpureum Schum. To be added, as an escape, very abundant. Cenchrus myosuroides HBK. (Cenchropsis myosuroides Nash) Cenchrus incertus M. A. Curtis (Cenchrus carolinianus Walt.) Cenchrus brownii R. & S. (Cenchrus viridis Spreng.) Leersia hexandra Sw. (Homalocenchrus hexandrus Kuntze) Leersia monandra Sw. (Homalocenchrus monandrus Kuntze) Aristida chaseae Hitche. To be added: Boquerón. Sporobolus tenuissimus (Schrank) Kuntze (Sporobolus mura- lis Hitche. & Chase) Add to Puerto Rico: Near Adjuntas. Sporobolus poiretii (R. & S.) Hitche. (Sporobolus berteroanus Hitche. & Chase) Sporobolus pyramidatus (Lam.) Hitche. Kunth) Omit: Sporobolus domingensis (Trin.) Kunth. The Cabo Rojo specimen (Sintenis 549) is Sporobolus pyramidatus (Lam.) Hitchc. Cynodon dactylon (L.) Pers. (Capriola dactylon Kuntze) Chloris inflata Link (Chloris paraguayensis Steud.) Chloris petraea Sw. (Eustachys petraea Desv.) Omit: Chloris sagraeana A. Rich., not in St. Croix. Leptochloa panicea (Retz.) Ohwi (Leptochloa filiformis P. Beauv.) Leptochloa fascicularis (Lam.) A. Gray (Diplachne fasci- cularis Beauv.) Pappophorum pappiferum (Lam.) Kuntze (Pappophorum. alopecuroideum Vahl) Phragmites communis Trin. (Phragmites phragmites Karst.) Eragrostis glutinosa (Sw.) Trin. To be added: St. Croix. Eragrostis urbaniana Hitchc. To be added: Anegada. (Sporobolus argutus Ref.: p. 80 p. 81 p. p. p. 85 85 86 p. 87 p. p. p. 87 88 88 p. 88 p. To be 89 . 91 . 91 . 91 . 92 Rhodora [Vol. 67 Bambusa bambos (L.) Voss (Bambos vulgaris Schrad.) Melinis minutiflora Beauv. To be added, as an escape. Las Mesas, Mayagiiez. CYPERACEAE KÜKENTHAL, G.—Bot. Jahrb. 1949-1950. (Rhynchospora) SVENSON, H.—N. Amer. Flora 18, parts 8 and 9. 1947-1957. Koyama, T.—Journ. Fac. Sci. Univ. Tokyo, Sect. III, Bot. VIII. 1961. GONZALEZ Mas, ARTURO.— The Cyperaceae of Puerto Rico. 1964. Cyperus brevifolius (Rottb.) Hassk. (Kyllinga brevifolia Rottb.) Cyperus obtusatus (Presl) Mattf. & Kük. (Kyllinga pungens Link) Cyperus peruvianus (Lam.) F. N. Wiliams (Kyllinga peru- viana Lam.) Cyperus densicaespitosus Mattf. & Kük. (Kyllinga pumila Michx.) Omit: Kyllinga odorata Vahl, not in Puerto Rico. Cyperus polystachyos Rottb. (Cyperus odoratus L., sensu Britt.) Cyperus imbricatus Retz. (Cyperus radiatus Vahl) Cyperus flavus (Vahl) Nees (Cyperus cayennensis Britt.) Cyperus nanus Willd. (Cyperus granularis Britt.) Cyperus nanus var. subtenuis Kük. (Cyperus tenuis Sw.) Cyperus mutisii (HBK.) Griseb. (Cyperus incompletus Link) Cyperus planifolius L. C. Rich. (Cyperus brunneus Sw.) Cyperus unifolius Bóckl. (Cyperus filiformis Sw.: Cyperus calcicola Britt., No. 23, p. 86). Cyperus odoratus L. (Cyperus ferax L. C. Rich.) Cyperus flexuosus Vahl (Cyperus vahlii Steud.) added: Cyperus iria L. Quite abundant, in wet places. Cyperus difformis L. Mayagüez. Cyperus digitatus Roxb. Mayagüez Cyperus unioloides R. Br. Lares. Cyperus swartzii (Dietr.) Böckl. & Kük. Peñuelas. Cyperus eggersii Böckl. San Sebastián. Eleocharis flavescens (Poir.) Urb. Add synonym: Eleocharis flaccida Urb., No. 4. Eleocharis geniculata (L.) R. & S. (Eleocharis caribaea Blake; Eleocharis atropurpurea Kunth, as to the Puerto-Rican plant.) Eleocharis montana (HBK.) R. & S. (Eleocharis nodulosa Schultes) Elocharis sintenisii Böckl. (Eleocharis yunquensis Britt.) p. í Flora of Porto Rico — Liogier 321 Eleocharis minutiflora Bóckl. (Eleocharis microcarpa Torr.) Eleocharis elegans (HBK.) R. & S. (Eleocharis geniculata R. & S. in part) added: Elocharis fallax Weath. Guanica. Eleocharis rostellata Torr. Vega Baja. Eleocharis radicans (Poir. Kunth. Not collected by recent collectors. Based on a Ledru collection, before 1804. Fimbristylis vestita (Kunth) Hemsl. (Stenophyllus vestitus Britt.) Fimbristylis portoricensis (Britt.) Alain (Stenophyllus por- loricensis Britt.; Bulbostylis pauciflora Clarke). Fimbristylis curassavica (Britt.) Alain (Stenophyllus curas- savicus Britt.) To be added: Guanica. Fimbristylis cymosa R. Br. (Fimbristylis spathacea Roth.) Fimbristylis dichotoma (L.) Vahl (Fimbristylis diphylla Vahl) Fimbristylis monostachyos (L.) Hassk. (Abildgaardia mono- stachya Vahl) Scirpus lacustris L. ssp validus (Vahl) T. Koyama (Scirpus validus Vahl) Omit: (Fuirena squarrosa Michx., not in Puerto Rico). Scirpus fuirena T. Koyama (Fuirena umbellata Rottb.) Rhynchospora stellata Griseb. (Dichromena colorata Hitchc.) Rhynchospora nervosa (Vahl) Böckl. (Dichromena ciliata Vahl) Rhynchospora radicans (S. & C.) Pfeiff. (Dichromena radi- cans S. & C.) Rhynchospora nitens (Vahl) A. Gray (Psilocarya portori- censis Britt.; Psilocarya nitens Wood.) Rhynchospora polyphylla (Vahl) Vahl (Rhynchospora jamai- censis Britt.) Rhynchospora hispidula (Vahl) Böckl. (Rhynchospora tri- chodes Clarke in Urb.) Rhynchospora pusilla (Sw.) Griseb. (Rhynchospora berterü Clarke) Rhynchospora intermixta Wr. ex Sauv. (Rhynchospora bru- neri Britt.) Rhynchospora podosperma Wr. ex Sauv. (Rhynchospora filiformis Vahl) Rhynchospora brachychaeta Wr. ex Sauv. (Rhynchospora blauneri Britt.) Rhynchospora alba (L.) Vahl (Rhynchospora luquillensis Britt.) S p. p. p. p. 107 109 Rhodora [Vol. 67 Rhynchospora fascicularis (Michx.) Vahl (Rhynchospora distans Vahl) Rhynchospora globularis Chapm. (Rhynchospora cymosa Ell.) Rhynchospora rugosa (Vahl) Gale (Rhynchospora glauca Vahl) Rhynchospora marisculus Lindl. & Nees (Rhynchospora borin- quensis Britt.) Rhynchospora lindeniana Griseb. var. bahamensis (Britt.) Gale (Rhynchospora bahamensis Britt.) Rhynchospora odorata Wr. ex Griseb. (Rhynchospora mari- sculus Lindl. & Nees, sensu Britton & Wilson). added: Rhynchospora racemosa Wr. ex Sauv. Maricao to Rosario. Rhynchospora microcarpa Baldw. ex Gray. Vega Baja. Cladium jamaicense Crantz (Mariscus jamaicensis Britt.) Cyperus pedunculatus (R. Br.) Kern (Remirea maritima Aubl.; Remirea pedunculata R. Br.) Lagenocarpus guianensis Lindl. (Lagenocarpus portoricensis Britt.) Scleria setacea Poir. in Lam. Add to synonymy: Scleria stevensiana Britt., No. 2. Scleria eggersiana Böckl. (Scleria grisebachii Clarke) To be added: Scleria triglomerata Michx. Western Puerto Rico. Ref.: p. p. p. 113 . 113 Scleria ciliata Michx. Las Mesas, Mayagüez. PALMAE Gentes Herbarum. 1940-1963. Prestoea montana (R. Grah.) Nichols. (Euterpe globosa Gaertn.) Calyptronoma rivalis (O. F. Cook) Bailey (Calyptrogyne occidentalis Maza, as to the Puerto-Rican plant). Aiphanes acanthophylla (Mart.) Burrett (Bactris acantho- phylla Mart.) Acrocomia media O. F. Cook (Acrocomia aculeata Lodd in Mart., as to the Puerto-Rican plant.) Thrinax ponceana O. F. Cook. To be added: South coast, near Ponce. Coceothrinax alta Bece. To be added: St. Thomas, P. Rico. Coccothrinax eggersiana Bece. To be added: St. Thomas St. Croix. Coccothrinax discreta Bailey & Moore. To be added: Vieques. Coccothrinax sancti-thomae Becc. To be added: St. Thomas. ARACEAE Philodendron hederaceum (Jacq.) Schott (Philodendron 1965] p. 132 Ref.: p. p. ; 401 135 135 . 188 . 139 . 141 142 . 148 Flora of Porto Rico — Liogier 323 scandens C. Koch & H. S.. Sello; Philodendron oxycardium Schott, No. 6, p. 124). Philodendron nechodomi Britt. To be added: Luquillo Mts. Cf. Sci. Surv. VI: 325 and 569. Colocasia esculenta (L.) Schott (Caladium colocasia W. F. Wight) Caladium bicolor (Ait.) Vent. (Cyrtospadix bicolor Britt. & Wils.) Xanthosoma nigrium (Velloso) Stellfeld (Xanthosoma viola- ceum Schott) XYRIDACEAE Xyris carolinensis Walt. (Xyris jupicai L. C. Rich.) BROMELIACEAE SMITH, LyMAN B.—Bromeliaceae N. Am. Flora 19: 61-228. 1938. Aechmea lingulata (L.) Baker (Wittmachia lingulata Mez) Ananas comosus (L.) Merrill (Ananas ananas Cockerell) Catopsis floribunda (Brongn.) L. B. Sm. (Catopsis nutans Griseb.) Tillandsia usneoides L. (Dendropogon usneoides Raf.) Tillandsia flexuosa Sw. (Tillandsia aloifolia Hook.) Tillandsia valenzuelana A. Rich. in Sagra (Tillandsia sublaxa Baker) Vriesea macrostachya (Bello) Mez (Neovriesia macrostachya Britt.) Vriesea sintenisii (Baker) Smith & Pitt. (Thecophyllum sintenisii Mez) COMMMELINACEAE Ref.: Woopson, R. E.—Ann. Mo. Bot. Gard. 29: 145. 1942. p. p. p. p- p. p- 144 145 146 146 147 149 . 149 Phaeosphaerion persicariaefolius (DC.) C. B. Clarke (Athyro- carpus persicariaefolius Hemsl.) Commelina diffusa Burm. f. (Commelina longicaulis Jacq.) Aneilema geniculatum (Jacq.) Woods. (Tradescantia geni- culata Jacq.) Tripogandra elongata (G. F. W. Meyer) Woods. (Tradescan- tia elongata G. F. W. Meyer). Rhoeo spathacea (Sw.) Stearn (Rhoeo discolor Hance) PONTEDERIACEAE Eichhornia crassipes (Mart.) Solms (Piaropus crassipes Britt.) Eichhornia diversifolia (Vahl) Urb. (Piaropus diversifolius P. Wils.) 324 Rhodora [Vol, 67 LILIACEAE (includes Smilacaceae) p. 150 Sanservieria metalica Geróme and Labory. (Cordyline guine- ensis Britt.) p. 151 Aloe barbadensis Mill. (Aloe vulgaris Lam.) p. 154 Smilax lanceolata L. (Smilax domingensis Willd. Cf. Flora of Guatemala 3: 95. 1952) AMARYLLIDACEAE Ref.: HUME, H.—Herbertia VI: 121-135. 1939 (Zephyranthes) p. 156 Hypoxis wrightii (Baker) Brackett. To be added: Catano, Tortuguero. Dorado, Bayamón. Cf. Rhodora 25: 140. 1928; Sci. Surv. VI: 338. p. 158 Zephyranthes puertoricensis Traub. (Atamosco tubispatha Maza, as to the Puerto-Rican plant.) p. 159 Zephyranthes rosea Lindl. (Atamosco rosea Greene) p. 159 Zephyranthes grandiflora Herb. (Atamosco carinata P. Wils.) p. 160 Pancratium declinatum Jacq. (Hymenocallis declinata M. Roem.) DIOSCOREACEAE p. 1685 Dioscorea villosa L. (Dioscorea cayennensis Lam.; Dioscorea esculenta Burkill) p. 163 Dioscorea bulbifera L. To be added: Cf. Sei. Surv. VI: 531. HAEMODORACEAE p. 165 Trimezia martinicensis (Jaeq.) Herb. To be added: Baya- món, naturalized. Cf. Sci. Surv. VI: 531. IRIDACEAE p. 166 Eleutherine bulbosa (Mill.) Urb. (Galathea bulbosa Britt.) MUSACEAE ». 167 Heliconia caribaea Lam. (Bihai bihai Griggs, as to the Puerto Rican plant). ZINGIBERACEAE (Alpiniaceae) p. 171 Alpinia speciosa (Wendl) K. Schum. (Languas speciosa Small). p. 172 Renealmia antillarum (R. & S.) Gagnepain (Alpinia antil- larum R. & S.) p. 172 Renealmia aromatica (Aubl) Griseb. (Alpinia aromatica Aubl.) p. 173 Renealmia exaltata L. f. (Alpinia exaltata R. E S.) 73 Zingiber officinalis Rose. (Zingiber zingiber Karst.) — — 1965] Flora of Porto Rico — Liogier 325 BURMANNIACEAE Ref.: JONKER, F. P.—A Monograph of the Burmanniaceae. Utrecht, 1938. p. 179 Gymnosiphon germainii Urb. (Ptychomeria portoricensis Schltr.) p. 179 Gymnosiphon sphaerocarpus Urb. To be added: Maricao. p. 179 Apteria aphylla (Nutt.) Burm. ex Small (Aperia hymenan- thera Miq.) ORCHIDACEAE Ref.: ScHULTES, R. E.—Native Orchids of Trinidad and Tobago. 1960. DUNSTERVILLE, G. C. K.—Orquideas de Venezuela. 1960. p. 184 Pogonia macrophylla Lindl. (Psilochilus macrophyllus Ames). Cf. Brittonia 14: 443. 1962. p. 185 Vanilla dilloniana Correll (Vanilla eggersii Rolfe) p. 186 Spiranthes tortilis (Sw.) L. C. Rich. (Ibidium tortile House) p. 186 Spiranthes fawcettii Cogn. (Hapalorchis tenuis Schlecht.) p. 187 Spiranthes lucayana (Britt.) Cogn. (Mesadenus lucayanus Sehlecht.) p. 187 Spiranthes elata (Sw.) L. C. Rich. (Beadlea elata Small) p. 188 Spiranthes cranichoides (Griseb.) Cogn. (Beadlea crani- choides Small) p. 188 Spiranthes adnata (Sw.) Fawe. & Rendle (Pelexia adnata Spreng.) p. 189 Spiranthes speciosa (J. F. Gmelin) A. Rich. in Sagra (Stenorrhynchus speciosus A. Rich.) p. 189 Spiranthes orchioides (Sw.) A. Rich. (Stenorrhynchus lanceo- latus Griseb.) p. 189 Spiranthes amabilis Ames. To be added: Maricao State Forest. p. 189 Centrogenium setaceum (Lindl) Schlecht. (Eltroplectris acuminata Raf.) p. 189 Waullschlaegelia aphylla (Sw.) Rchb. f. To be added: Luquillo Mts. p. 190 Erythrodes plantaginea (L.) Fawe. E Rendle (Physurus plantagineus Lindl.) p. 190 Erythrodes hirtella (Sw.) Fawe. & Rendle (Physurus hir- tellus Lindl.) p. 191 Cranichis diphylla Sw. To be added: Maricao State Forest. p. 194 Polystachya extinctoria Rchb. f. ex Walp. (Polystachya minuta Britt.) Cf. Taxon IX: 150. 1960. p. 195 Polystachya cerea Lindl. (Polystachya minor Fawe. & Rendle) p. 195 Scaphyglottis modesta (Rehb. f.) Schlecht. (Tetragamestus modestus Rchb. f.) 326 p. 196 p. 196 p. 197 p. 197 p. 197 p. 197 p. 198 p. 198 p. 199 p. 199 p. 200 p. 200 p. 200 p. 200 p. 201 p. 201 p. 201 p. 202 p. 202 p. 202 To be p. 202 p. 203 p. 206 p. 206 p. 206 Rhodora [Vol. 67 Dilomilis montana (Sw.) Summerhayes (Octadesmia montana Benth.) Ref.: Taxon X: 253. 1961. Coelia triptera (Smith) G. Don (Hormidium tripterum Cogn.) Jacquiniella teretifolia (Sw.) Britt. & Wils. To be added: Cerro de Punta, Guilarte. Jacquiniella miserrima (Rchb. f.) Stehlé. To be added: Carite. Epidendrum fucatum Lindl. (Encyclia fucata Britt. & Millsp.) Epidendrum sintenisii Rchb. f. (Encyclia sintenisii Britt.) Epidendrum brittonianum A. D. Hawkes (Eneyclia papilion- acea, Schlechtr.) Epidendrum cochleatum L. (Anacheilium cochleatum Hof- fmg.) Epidendrum ottonis Rchb. f. (Nidema ottonis Britt, E Millsp.) Epidendrum ciliare L. (Auliza ciliaris Salisb.) Epidendrum difforme Jacq. (Amphiglottis difformis Britt.) Epidendrum moirianum A. D. Hawkes (Amphiglottis corym- bosa Britt.) Epidendrum necturnum Jacq. (Amphiglottis nocturna Britt.) Epidendrum anceps Jacq. (Amphiglottis anceps Britt.) Epidendrum secundum Jacq. (Amphiglottis secunda Britt.) Epidendrum lacerum Lindl. (Amphiglottis lacera Britt.) Epidendrum pallidiflorum Hook. (Amphiglottis pallidiflora Britt.) Epidendrum rigidum Jacq. (Spathiger rigidus Small) Epidendrum ramosum Jacq. (Spathiger ramosus Britt.) Epidendrum krugii Bello (Encyclia krugii Britt. & Wils. Sci. Surv. VI: 532) To be added: S.—W. of Puerto Rico; doubt- fully different from Epidendrum brittonianum A. D. Hawkes; to be studied further). added: Epidendrum vicentinum Lindl. Guilarte and Guavate. Epidendrum belvederense Fawe. & Rendle; locality unknown. Domingoa haematochila (Rchb. f.) Carabia (Domingoa hymenodes Schlecht.) To be added: Mona Island Cf. Sci. Surv. VI: 340-341. Brassavola cucullata (L.) R. Br. (Epidendrum cucullatum L.) Pleurothallis foliata Griseb. (Pleurothallis broadwayi Ames) To be added: Cerro de Punta. Cf. Bull, Torrey Bot. Club 90: 187. 1963. Pleurothallis appendiculata Cogn. To be added: Maricao. Pleurothallis longissima Lindl. To be added: Adjuntas. 1965] Flora of Porto Rico — Liogier 327 p. 208 Eulophia alta (L.) Fawe. & Rendle (Platypus altus Small) p. 209 Bulbophyllum pachyrrhachis (A. Rich.) Griseb. (Bolbophyl- laria pachyrrhachis Rchb. f.) p. 210 Maxillaria coccinea (Jacq.) L. O. Wms. (Ornithidium coc- cineum Salisb.) p. 210 Maxillaria conferta (Griseb.) Schweinf. (Ornithidium con- fertum Griseb.) p. 212 Oncidium lemonianum Lindl. (Oncidium intermedium Bert.) p. 214 Cochleanthes flabelliformis (Sw.) R. E. Schultes & Garay (Warszewiczella flabelliformis Cogn.) p. 216 Campylocentrum fasciola (Lindl.) Cogn. (Campylocentrum sullivani Fawc. & Rendle) PIPERACEAE Ref.: YUNCKER, T. G.—The Cuban Species of Peperomia. Rev. Soc. Cub. Bot. VI. 1949. The Piperaceae of Northern South America. 1950. p. 221 Piper hispidum Sw. (Piper scabrum Sw.) p. 221 Piper wydlerianum (Miq.) DC. (Piper citrifolium Lam.) p. 222 Piper glabrescens (Miq.) DC. (Piper treleaseanum Britt & Wils.) p. 227 Peperomia sintenisii C. DC. (Peperomia dendrophila Schl.) p. 228 Peperomia rhombea R. & P. (Peperomia myrtillus Miq.) p. 228 Peperomia megapoda Trel. To be added: Cayey to Guayamas. Cf. Sci. Surv. VI: 533. p. 228 Peperomia tenella A, Dietr. To be added: Cerro de Punta, Jayuya. p. 229 Piper peltatum L. (Pothomorphe peltata Miq.) p. 229 Piper umbellatum L. (Pothomorphe umbellata Miq., to be segregated from the preceding, same distribution). MYRICACEAE p. 232 Myrica holdridgeana Lundell. To be added: Sierra de Lu- quillo, EI Toro. Cf. Contr. Univ. Michigan Herb. 7: 5. 1942. ULMACEAE p. 284 Celtis iguanaea (Jacq.) Sarg. (Momisia iguanaea Rose & Standl.) MORACEAE p. 237 Ficus trigonata L. (Ficus crassinervia Desf.; F. stahlit Warb. in Urb.) p. 237 Ficus citrifolia P. Mill. (Ficus laevigata Vahl) p. 237 Ficus perforata L. (Ficus sintenisti Warb.) 328 Rhodora [Vol. 67 p. 239 Ficus retusa L. (Ficus nitida Thunb.) p. 241 Artocarpus heterophylla Lam. (Artocarpus integrifolia 1..) OLACACEAE p. 254 Ximenia americana L. To be added: Guanica. LORANTHACEAE p. 255 Dendropemon bicolor Krug & Urb. (Phthirusa bicolor Engler) p. 255 Dendropemon sintenisii Krug & Urb. (PLthirusa sintenisü Engler) p. 255 Dendropemon purpureum (L.) Krug & Urb. (Phthirusa pur- purea Engler) p. 256 Dendropemon caribaeum Krug & Urb. (Phthirusa caribaea Engler) p. 256 Eremolepis wrightii Griseb. (Ixidium wrightii Eichl.) p. 259 Phoradendron berterianum (DC.) Griseb. (Phoradendron dichotomum Krug & Urb.) p. 260 Dendrophthora flagelliformis (Lam.) Krug & Urb., include as a synonym: Dendrophthora wrightii Eichl. in Mart., No. 1, p. 259. Cf. Wentia 6: 1-133, 1961. ARISTOLOCHIACEAE p. 262 Aristolochia bilabiata L. (Aristolochia oblongata Jacq.) p. 263 Aristolochia grandiflora Vahl (Aristolochia galeata Mart. & Zuce.; Aristolochia ringens Vahl) p. 263 Aristolochia cordiflora Mutis ex HBK. (Aristolochia grandi- flora Sw.) To be added: Aristolochia littoralis Parodi (Aristolochia elegans Masters). St. Kitts, Aristolochia odoratissima L. St. Jan, St. Thomas. POLYGONACEAE tef.: HOWARD, R, A.— The species of Coccoloba from Puerto Rico and the Virgin Islands. Jour. Arn. Arb. 38: 211-242. 1957. p. 264 Polygonum punctatum Ell. (Persicaria punctata. Small) p. 265 Polygonum acuminatum HBK. (Persicaria acuminata Maza) p. 265 Polygonum densiflorum Meisn. (Persicaria portoricensis Small) p. 265 Polygonum segetum HBK. (Persicaria segetum Small) p. 266 Antigonon guatemalense Meisn. (Antigonon macrocarpum Britt. & Small) p. 266 Coccoloba (Coccolobis) p. 267 Coccoloba swartzii Meisn. f. urbaniana (Lindl) Howard (Coccolobis borinquensis Britt.) p. . 298 296 Flora of Porto Rico — Liogier 329 Coccoloba diversifolia Jacq., add synonym: Coccolobis lauri- folia Jacq., no. 8. Coccoloba microstachya Willd. (Coccolobis obtusifolia Jacq.) Coccoloba pubescens L. (Coccolobis grandifolia Jacq.) Coccoloba costata Wr. ex Sauv. (Coccolobis rupicola Urb.) Coccoloba swartzii var. portoricensis Meisn. is synonym to C. swartzii f. urbaniana (Lindl.) Howard CHENOPODIACEAE Suaeda fruticosa (L.) Forsk. (Dondia fruticosa Northrop) AMARANTHACEAE Amaranthus viridis L. add synonym: Amaranthus gracilis Desf., No. 4. Achyranthes aspera L. (Centrostachys aspera Standl.; C. indica Standl.) Alternanthera peploides (H. & B.) Urb. (Achyranthes pep- loides Britt.) Alternanthera paronychoides St. Hil. (Achyranthes poly- gonoides Lam.) Alternanthera sessilis (L.) R. Br. ex DC. (Achyranthes sessilis Steud.) Alternanthera portoricensis Kuntze (Achyranthes portori- censis Standl.) Alternanthera ficoidea (L.) R. Br. ex R. & S. (Achyranthes ficoidea Lam.) This species is probably a synonym to Alter- nanthera polygonoides (L.) R. Br. Alternanthera ficoidea var. bettzickiana (Mich.) Baker Achyranthes bettzickiana Standl.) Alternanthera ramosissima (Moq.) Chod. (Achyranthes ramosissima Standl.) Gomphrena decumbens Jacq. (Gomphrena dispersa Standl.) Iresine diffusa H. & B. ex Willd. (Iresine celosia L.) NYCTAGINACEAE Boerhaavia diffusa L. (Boerhaavia coccinea Mill.) The species of Torrubia are being studied and will probably be transferred to Guapira. Cf. Woodson, R. E. Ann. Mo. Bot. Gard. XLVIII: 61. 1961. PHYTOLACCACEAE Stegnosperma cubense A. Rich. To be added: Aibonito. AIZOACEAE Tetragonia tetragonioides (Pallas) O. Kuntze (Tetragonia expansa Murr.) 330 Rhodora [Vol. 67 CARYOPHYLLACEAE (Alsinaceae) p. 297 Stellaria antillana Urb. (Alsine antillana Britt. & Wils.) PORTULACACEAE Ref.: LEGRAND, DrEGO.—Las Especies americanas de Portulaca. Ann. Mus. Hist. Nat. Montevideo, 2 ser. VII (3): 1-147. 1962. p. 300 Portulaca rubricaulis HBK. (Portulaca phaeosperma Urb.) p. 3900 Portulaca teretifolia HBK. (Portulaca poliosperma Urb.) BASELLACEAE p. 301 Anredera leptostachys (Moq.) Steen. (Boussingaultia lepto- stachys Moq.) p. 301 Anredera baselloides (HBK.) Baill. (Boussingaultia basel- loides HBK.) p. 801 Basella alba L. (Basella rubra L.) NYMPHAEACEAE (includes Cabombaceae) p. 304 Nymphaea ampla (Salisb.) DC. (Castalia ampla Salisb.) p. 304 Nymphaea pulchella DC. (Castalia pulchella Britt.) p. 304 Nymphaea odorata Dryand (Castalia odorata Woodw. & Woodw.) p. 305 Nymphaea amazonum Mart. E Zucc. (Castalia amazonum Britt. & Wils.) p. 305 Nymphaea rudgeana G. F. W. Meyer (Castalia rudgeana Britt. & Wils.) ANNONACEAE Ref., FRIES, R. E.—Acta Horti Bergiani. 1931-1939. p. 311 Guatteria blainii (Griseb.) Urb. (Cananga blainii Britt.) p. 311 Guatteria caribaea Urb. (Cananga caribaea Britt.) LAURACEAE (includes Cassythaceae) Ref.: Kopp, LUCILLE.—Revision of Persea. Unpublished. KOSTERMANS, A. J. G. H.—Reinwardtia 6: 17-24. 1961. p. 316 Beilschmiedia pendula (Sw.) Benth. & Hook. (Hufelandia pendula Nees) p. 317 Licaria salicifolia (Sw.) Kosterm. (Acrodiclidium salicifolium Griseb.) p. 8317 Licaria triandra (Sw.) Kosterm. (Misanteca triandra Mez) p. 817 Licaria brittoniana Allen & Gregory. To be added: Maricao & Susua. Brittonia VII: 267-269. 1951. p. 318 Persea americana Mill. (Persea persea Cockerell) p. p. p. . 324 . 327 . 328 . 328 329 329 Flora of Porto Rico — Liogier 331 Persea urbaniana Mez (Persea portoricensis Britt. & Wils.) To be added: Luquillo & Cerro de Punta. Cf. Sci. Surv. VI: 346. Cinnamomum montanum (Sw.) Berchthold & Presl (Phoebe montana Griseb.) Cinnamomum elongatum (Vahl) Kosterm. (Phoebe elongata Nees) Cinnamomum zeylanicum Blume (correct author citation) BRASSICACEAE Brassica rapa L. (Brassica campestris L.) Brassica kaber (DC.) L. C. Wheeler (Sinapis arvensis L.) Rorippa portoricensis (Spreng.) Stehlé (Radicula portori- censis Britt.) Rorippa palustris (L.) Bess. (Radicula palustris Moench.) Rorippa nasturtium-aquaticum (L.) Hayek. (Sisymbrium nasturtium-aquaticum L.) To be added: Coronopus didymus (L.) Smith. Villalba. p. . 332 . 332 337 . 942 . 844 . 344 . 945 . 347 . 347 . 348 . 350 . 350 350 CAPPARACEAE (Capparidaceae) Cleome rutidosperma DC. (Cleome ciliata Schum. & Thonn.) Cleome viscosa L. (Cleome icosandra L.) MORINGACEAE Moringa oleifera Lam. (Moringa moringa Millsp.) ROSACEAE (includes Amygdalaceae) Rubus probus Bailey, found in the central mountains of Puerto Rico; probably a good species, not a hybrid. Prunus myrtifolia (L.) Urb. (Laurocerasus myrtifolia Britt.) Prunus occidentalis Sw. (Laurocerasus occidentalis Roem.) Chrysobalanus icaco L., add to synonymy: Chrysobalanus pellocarpus G. F. W. Meyer. LEGUMINOSAE-MIMOSOIDEAE Inga vera Willd. (Inga inga Britt.) Inga fagifolia (L.) Willd. (Inga laurina Willd.) Pithecellobium unguis-cati (L.) Benth. (correct author cita- tion) Calliandra caracasana (Jacq.) Benth. (Anneslia portori- censis Britt.) Calliandra haematostoma (Bertero) Benth. (Anneslia haema- tostoma Britt.) Calliandra purpurea (L.) Benth. (Anneslia purpurea Britt.) p. 373 p. 373 p. 374 p. 374 Rhodora [Vol. 67 Acacia farnesiana (L.) Willd. (Vachellia farnesiana Wight & Arn.) Leucaena leucocephala (Lam.) DeWit (Leucaena glauca Benth.) Desmanthus virgatus (L.) Willd. (Acuan virgatum Medic.; Acuan depressum Kuntze; Acuan insulare Britt. & Rose; cf. Sci. Surv. VI: 539). Schrankia portoricensis Urb. (Morongia portoricensis Britt.) Mimosa pigra L. To be added: Mayagüez. Prosopis limensis Benth. 'To be added: Vieques, St. Croix. Anadenanthera peregrina (L.) Speg. (Piptadenia peregrina Benth.) Albizia (correct spelling) CAESALPINIOIDEAE Cassia diphylla L. (Chamaecrista diphylla Greene) Cassia glandulosa var. swartzii (Wickstr.) | Macbride (Chamaecrista swartzii Britt.) Cassia mirabilis (Pollard) Urb. (Chamaecrista mirabilis Pollard) Cassia portoricensis Urb. (Chamaecrista portoricensis Cook & Collins) Cassia granulata (Urb.) Macbride (Chamaecrista granulata Britt.) Cassia grammica Spreng. (Chamaecrista grammica Pollard) Cassia diffusa DC. (Chamaecrista chamaecrista Britt.) Cassia aeschynomene DC. (Chamaecrista aeschynomene Greene) Cassia javanica L, (Cassia nodosa Hamilt.) Cassia antillana (Britt. & Rose) Alain (Chamaecrista antil- lana Britt. & Rose) Cassia fruticosa Mill. (Chamaefistula bacillaris G. Don) Cassia stahlii Urb. (Adipera stahlii Britt. & Rose) Cassia bicapsularis L. (Adipera bicapsularis Britt, & Rose) Cassia laevigata Willd. (Adipera laevigata Britt. & Rose) Cassia obtusifolia L. (Emelista tora Britt. & Rose) Cassia hirsuta L. (Ditremexa hirsuta Britt. & Rose). Col- lected again near Bayamón. Cassia occidentalis L. (Ditremexa occidentalis Britt. & Rose; Ditremexra sophera Britt. & Rose) Cassia polyphylla Jaeq. (Peiranisia polyphylla Britt. & Rose) Cassia obovata Collad. (Senma obovata Batka) Cassia emarginata L. (Jsandrina emarginata Britt. & Rose) Cassia alata L. (Herpetica alata Raf.) To be added: Cassia patellaria DC. (Chamaecrista patellaria Greene) 1965] p. T . 916 . 378 . 378 . 379 . 379 . 379 . 380 . 380 . 380 . 981 . 386 . 990 Ook z091 . 391 . 392 . 394 . 994 . 395 . 395 . 395 . 395 396 397 e 399 Flora of Porto Rico — Liogier 333 See Sci. Surv. VI: 540. Abundant, Western and South- ern Puerto Rico. Cassia exunguis Urb. (Chamaecrista exunguis Britt.) See Sci. Surv. VI: 541. Between Aibonito and Coamo. Cassia rotundifolia Pers. Cerro Gordo. Caesalpinia pulcherrima (L.) Sw. (Poinciana pulcherrima L.) Caesalpinia coriaria (Jacq.) Willd. (Libidibia coriaria Schl.) Caesalpinia crista L. (Guilandina crista Small) Caesalpinia bonduc (L.) Roxb. (Guilandina bonduc L.) Caesalpinia divergens Urb. (Guilandina divergens Britt.) Caesalpinia melanosperma (Eggers) Urb. (Guilandina melan- ospermum Eggers) Caesalpinia portoricensis (Britt. & Wils.) Alain (Guilandina portoricensis Britt. & Wils.) Caesalpinia culebrae (Britt. & Wils.) Alain (Guilandina culebrae Britt. & Wils.) Caesalpinia decapetala (Roth.) Alst in Trimen. (Biancaea sepiaria Todaro) Peltophorum inerme (Roxb.) Naves (Peltophorum ferrugi- neum Benth.) PAPILIONOIDEAE Crotalaria saltiana Andr. (Crotalaria striata DC.) Dalea domingensis DC. (Parosela domingensis Millsp.) Tephrosia cinerea (L.) Pers. (Cracca cinerea Morong) Tephrosia senna HBK. (Cracca cathartica Britt. & Millsp.) Tephrosia candida DC. To be added: Ponce, Carolina, prob- ably an escape. Gliricidia sepium (aen) Kunth ex Walp. (correct author citation) Corynella paucifolia DC. (Correct spelling, not C. pauciflora) Cracea caribaea (Jacq.) Benth. (Benthamantha caribaea Kuntze) Sesbania sevicea (Willd.) DC. (Sesban sericea Rydb.) Sesbania sesban (L.) Fawc. & Rendle (Sesban sesban Britt.) Sesbania emerus (Aubl.) Urb. (Sesban emerus Rydb.) Sesbania bispinosa (Jacq.) Steud. ex Fawe. & Rendle (Ses- bania spinosa Spreng.) To be added: Guayanilla. See Sci. Surv. VI: 352. Sesbania grandiflora (L.) Pers. (Agati grandiflora Desv.) Aeschynomene gracilis Vogel (Aeschynomene portoricensis Urb.) See Contr. U. S. Nat. Herb. 32. 1955. Zornia reticulata Sm. (Zornia diphylla Pers., as to the Puerto- Rican plant). . 404 Rhodora (Vol, 67 Zornia gemella (Willd.) Vogel. To be added: Guanajibo, Mayagüez. See Webbia XVI. 1961. Desmodium barbatum (L.) Benth. & Oerst. (Meibomia barbata Kuntze) Ref: Contr. Gray Herb. CXXIX. 1940; CXXXV. 1941. Desmodium adscendens (Sw.) DC. (Meibomia adscendens Kuntze) Desmodium canum (J. F. Gmel.) Schinz & Thell. (Meibomia supina Britt.) Desmodium affine Schl. (Meibomia affinis Kuntze) Desmodium axillare (Sw.) DC. (Meibomia axillaris Kuntze) Desmodium axillare var. acutifolium (Kuntze) Urb. (Meibo- mia wmbrosa Britt.) Desmodium axillare var. stoloniferum (Rich. ex Poir.) Schub. (Meibomia sintenisii Britt.) Desmodium wydlerianum Urb. (Meibomia wydleriana Britt.) Desmodium scorpiurus (Sw.) Desv. (Meibomia scorpiurus Kuntze) Desmodium glabrum (Mill) DC. (Meibomia mollis Kuntze) Desmodium tortuosum (Sio.) DC. (Meibomia purpurea Vail. ex Small) Desmodium procumbens (Mill) Hitche. (Meibomia procum- bens Britt.) Desmodium triflorum (L.) DC. (Sagotia triflora Duch. & Walp.) To be added: Desmodium cubense Griseb. (Meibomia eubensis Schindl.) . 405 . 406 . 406 . 408 . 409 . 410 . 411 . 412 . 412 . 413 . 413 Coamo. See Sci. Surv. VI: 352. Desmodium intortum (Mill. Fawe. & Rendle. Maricao. Dalbergia monetaria L. f. (Securidaca volubilis L.) Dalbergia ecastaphyllum (L.) Taub. (Ecastophyllum ecas- tophyllum Britt.) Machaerium lunatum (L. f.) Ducke (Drepanocarpus lunatus G. F. W. Meyer) Lonchocarpus pentaphyllus (Poir.) DC. (Lonchocarpus lati- folius HBK.) See Sargentia VIII: 154. 1949. Piscidia piscipula (L.) Sarg. (lIchthyomethia — piscipula Hitche.) Abrus precatorius L. (Abrus abrus W. F. Wight) Clitoria rubiginosa Juss. (Martiusia rubiginosa Britt.) Clitoria laurifolia Poir. in Lam. (Martiusia laurifolia Britt.) Centrosema plumieri (Turp.) Benth. (Bradburya plumieri Kuntze) Centrosema pubescens Benth. (Bradburya pubescens Kuntze) Centrosema virginianum Benth. (Bradburya virginiana Kuntze) 1965] Flora of Porto Rico — Liogier 335 p. 414 Cajanus cajan (L.) Millsp. (Cajan cajan Millsp.) p. 415 Rhynchosia phaseoloides (Sio.) DC. (Dolicholus pyramidalis Britt. & Rose) p. 415 Rhynchosia reticulata (Sw.) DC. (Dolicholus reticulatus Millsp.) p. 415 Rhynchosia minima (L.) DC. (Dolicholus minimus Medic.) p. 416 Calopogonium mucunoides Desv. (Calopogonium orthocarpum Urb.) p. 418 Canavalia (correct spelling, instead of Canavali). See Brit- tonia 16: 106-181. 1964. p. 419 Canavalia nitida (Cav.) Piper (Canavali rusiosperma Urb.) p. 419 Canavalia ensiformis (Jacq.) DC. (correct author citation) To be added: Canavalia gladiata (Jacq.) DC. Virgin Islands. Canavalia brasiliensis Mart. ex Benth. Virgin Islands. p. 420 Phaseolus schottii Benth. (Phaseolus trichocarpus Wr. ex Sauv., with doubt). p. 421 Macroptilium lathyroides (L.) Urb. (Phaseolus lathyroides L.) p. 421 Phaseolus coccineus L. (Phaseolus multiflorus Willd.) p. 422 Vigna luteola (Jacq.) Benth. (Vigna repens Kuntze) p. 423 Phaseolus peduncularis HBK. (Vigna peduncularis Fawe. & Rendle) p. 424 Pachyrrhizus erosus (L.) Urb. (Cacara erosa Kuntze) p. 424 Pueraria phaseoloides Benth. To be added: abundant, es- caped from cultivation. p. 426 Mucuna pruriens (L.) DC. (Stizolobium pruritum Piper) p. 427 Erythrina eggersii Kruk. & Mold. (Erythrina horrida Eggers) OXALIDACEAE p. 431 Oxalis intermedia A. Rich. (Jonoxalis intermedia Small) p. 431 Oxalis corymbosa Zuce. (Ionoxalis martiana Small) p. 431 Oxalis corniculata L. (Xanthoxalis corniculata Small) To be added: Oxalis barrelieri L. Eastern Puerto Rico. ERYTHROXYLACEAE p. 433 Erythroxylum rotundifolium Lunan (Erythroxylum brevipes DC.) p. 434 Erythroxylum rufum Cav. To be added: Maricao. See Sci. i Surv. VI: 354. MALPIGHIACEAE Ref.: NIEDENZU, F.— Pflanzenreich IV'". 1928 p. 438 Tetrapteris citrifolia (Sw.) Pers. (Tetrapteris inaequalis Cav.) p. 438 Banisteria lucida Rich. (Banisteriopsis lucida Small) p. 438 Tetrapteris buxifolia Cav. (Adenoporces buxifolia Small). 336 Rhodora [Vol. 67 To be added: St. Thomas. Cf. Sci. Surv. VI. 545. p. 439 p. 439 p. 440 p. 440 p. 441 p. 442 p. 444 p. 444 p. 446 p. 447 p. 448 p. 461 tef.: p. 469 p. 470 p. 470 p. 471 p. 471 Heteropteris purpurea (L.) Kunth (Banisteria purpurea L.) Heteropteris wydleriana A. Juss. (Banisteria wydleriana C. B. Robins.) Heteropteris laurifolia (L.) Juss. (Banisteria laurifolia L.) Stigmaphyllon diversifolium A. Juss. (Stigmaphyllon ledi- folium Small) Stigmaphyllon periplocifolium (Desf.) Juss. (Stigmaphyllon lingulatum Small) Galphimia gracilis Bartl. (Thryallis glauca Kuntze, as to the Puerto-Rican plant) Malpighia shaferi Britt. & Wils. Add locality: Maunabo, Puerto Rico. Cf. Sci. Surv. VI: 354. Malpighia angustifolia L. (Malpighia linearis Jacq.) Byrsonima lucida (Sw.) DC. (Byrsonima cuneata P. Wils.; Byrsonima horneana Britt. & Small, No. 2). Byrsonima spicata (Cav.) DC. (Byrsonima ophiticola Small, No. 3). Byrsonima wadsworthii Little. To be added: Luquillo Mts., and Cerro de Punta, Jayuya. Cf. Phytologia 4: 417. 1953 RUTACEAE Murraya exotica L. (Chalcas exotica Millsp.) Citrus limon (L.) Burm. f. (Citrus limonum Risso) Citrus maxima (J. Burm.) Merrill (Citrus grandis Osbeck) SIMAROUBACEAE (includes Surianaceae, p. 451) Castela erecta Turp. (Castelaria nicholsoni Small) Picrasma antillana (Eggers) Urb. (Aeschrion antillana Small) Picrasma excelsa (Sw.) Planch. (Aeschrion excelsa Kuntze) BURSERACEAE Bursera simaruba (L.) Sarg. (Elaphrium simaruba Rose) POLYGALACEAE BLAKE, S. F.—N. Amer. Flora 25: 305-379. 1924. Polygala cowellii (Britt.) Blake (Phlebotaenia cowellii Britt.) Polygala portoricensis (Britt.) Blake (Badiera portoricensis Britt.) Polygala penaea L. (Badiera penaea DC.) Securidaca virgata Sw. (Elsota virgata Kuntze) Securidaca diversifolia (L.) S.F. Blake (Elseta diversifolia Blake) 1965] Ref.: p. 473 Flora of Porto Rico — Liogier 337 EUPHORBIACEAE WEBSTER, GRADY.—Journ. Arnold Arb. 1956-1959. (Phyllanthus) According to herbarium notes, Andrachne cuneifolia Britton must be placed into Securinega. Webster has not made the combination yet. Phyllanthus juglandifolius Willd. (Asterandra grandifolia Britt.) Flueggea virosa (Willd.) Baill. (Conami portoricensis Britt.) Phyllanthus acidus (L.) Skeels (Cicca disticha L.) Phyllanthus epiphyllanthus L. (Xylophylla epiphyllanthus Britt.) Phyllanthus lathyroides HBK. has not been collected in Puerto Rico. The specimens so named belong to Phyllanthus niruri L. Phyllanthus stipulatus (Raf.) Webster (Phyllanthus diffusus Klotzsch) Phyllanthus pentaphyllus Wr. ex Griseb. ssp. polycladus (Urb.) Webster (Phyllanthus polycladus Urb.) Phyllanthus pentaphyllus ssp. pentaphyllus. To be added: Susua State Forest. To be added: Phyllanthus urinaria L. Abundant in the whole island of Puerto Rico. p. 483 p. 484 p. 484 p. 485 p. 485 p. 485 p. 486 p. 487 p. 488 p. 493 p. 493 p. 496 p. 496 Croton trinitatis Millsp. To be added: St. Thomas. See Sci. Surv. VI: 547. Jatropha curcas L. (Curcas curcas Britt. & Millsp.) Jatropha hernandiaefolia Vent. (Curcas hernandiaefolius Britt.) Jatropha gossypifolia L. (Adenoropium gossypifolium Pohl) Jatropha multifida L. (Adenoropium multifidum Pohl) Jatropha integerrima Jacq. (Adenoropium hastatum Britt. & Wils.) Argythamnia fasciculata (Vahl) Muell. Arg. (Ditaxis fasci- culata Vahl) Bernardia dichotoma ( Willd.) Muell. Arg. (Adelia bernardia L.) Adelia ricinella L. (Ricinella ricinella Britt.) Manihot utilissima Pohl (Manihot manihot Cockerell) Sebastiania corniculata (Vahl) Pax. To be added: Hato Tejas, Cerro Gordo. Pedilanthus tithymaloides var. angustifolius (Poit.) Dressler (Pedilanthus angustifolius Poit.) Pedilanthus tithymaloides var. padifolius (L.) Dressler (Pedilanthus padifolius Poit.) 338 Rhodora [Vol. 67 p. 497 Pedilanthus tithymaloides var. parasiticus (L.) Dressler (Pedilanthus latifolius Millsp.) p. 499 Euphorbia oerstediana (Kl. & Garcke) Boiss. in DC. (Dichy- lium oerstedianum Britt.) p. 500 Euphorbia petiolaris Sims (Aklema petiolare Millsp.) The genus Chamaesyce has not been revised. Will report later. BUXACEAE p. 507 Buxus laevigata (Sw.) Spreng. (Tricera citrifolia Willd.) p. 508 Buxus vahlii Baill. (Tricera vahlii Britt.) p. 508 Buxus portoricensis Alain. To be added: Maricao, Moca. ANACARDIACEAE p. 510 Metopium linnaei Engl. (Metopium toxiferum Krug & Urb.) AQUIFOLIACEAE (Ilicaceae) p. 515 Ilex sideroxyloides var. occidentalis (Macfad.) Loes. To be added: Luquillo. To be added: Ilex cookii Britt. & Wils. Cerro de Punta, Jayuya. Cf. Sci. Surv. VI: 357. CELASTRACEAE p. 5317 Torralbasia cuneifolia (C. Wr.) Krug & Urb. To be added: Cerro de Punta and Luquillo Mts. Cf. Sci. Surv. VI: 358. p. 517 Crossopetalum rhacoma Crantz (Rhacoma crossopetalum L.) p. 519 Cassine xylocarpa Vent. (Elaeodendron xylocarpum DC.) HIPPOCRATACEAE Ref.: SMITH, A. C.—Brittonia 3: 341-555. 1940. p. 520 Pristimera caribaea (Urb.) A. C. Sm. (Hippocratea caribaea Urb.). Collected several times, mainly in the western part of Puerto Rico; also in Hispaniola. SAPINDACEAE (includes Dodonaeaceae) Ref.: RADLKOFER, L.— Sapindaceae, in Pflanzenreich 14. 165. 1931- 1934. p. 521 Dodonaea viscosa Jacq., add synonym: Dodonaea ehrenbergii Schl.) No. 1. p. 525 Cardiospermum halicacabum var. microcarpum (HBK.) Blume (Cardiospermum microcarpum HBK.) p. 525 Thouinia striata Radlk. (Thyana striata Britt.) p. 526 Thouinia portoricensis Radlk. (Thyana portoricensis Britt.) p. 528 Matayba apetala (Macf.) Radlk.. (Matayba oppositifolia Britt.) p. 529 Melicoccus bijugatus Jacq. (Melicocca bijuga L.) 1965] Flora of Porto Rico — Liogier 339 RHAMNACEAE Ref.: JOHNSTON, MARSHALL C.—Wrightia 3: 91-96. 1963; Amer. JOURN. Bot. 50: 1021. 1963: 51: 1113-1118. 1964. p. 534 Ziziphus rignonii Delp. (Sarcomphalus domingensis Krug & Urb.) p. 534 Ziziphus reticulata (Vahl) DC. (Sarcomphalus reticulatus Urb.) p. 534 Ziziphus taylori (Britt.) M. C. Johnst. (Sarcomphalus tay- lori Britt.) To be added: Mona Island. See Sci. Surv. VI: 358. p. 535 Ziziphus mauritiana Lam. (Ziziphus jujuba Lam.) Add to Puerto Rico: Guanica. p. 536 Colubrina arborescens (Mill) Sarg. (Colubrina colubrina Millsp.) p. 537 Colubrina urbanii M. C. Johnst. (Hybosperma spinosum Urb.) VITACEAE p. 541 Cissus tuberculata Jacq. To be added: Limestone Mts., N. Puerto Rico. ELAEOCARPACEAE p. 542 Sloanea amygdalina Griseb. To be added: Maricao. TILIACEAE Ref.: Ko Ko Lay.—Ann. Mo. Bot. Gard. 37: 315-395. 1950. p. 544 Triumfetta bartramia L. (Add synonym: Triumfetta excisa Urb Now) p. 545 Triumfetta bogotensis DC. (Triumfetta hispida A. Rich.) To be added: St. Croix. See Sci. Surv. VI: 550. MALVACEAE p. 547 Abutilon americanum (L.) Sweet (Abutilon abutiloides Garcke) p. 548 Wissadula contracta (Link) Fries, is cited from Puerto Rico in Flora of Trinidad and Tobago. I have seen no specimen of this species from the area of the Flora. p. 550 Malvastrum americanum (L.) Torrey (Malvastrum spicatum A. Gray) p. 551 Sida salviaevolia Presl (Sida erecta Macf.) > Sida acuta Burm. f. (Sida carpinifolia L. f.) p. 552 Sida alba L. To be added: Cabo Rojo. See Sci. Surv. VI: 550. p. 560 Pavonia fruticosa (Mill) Fawe. & Rendle (Thyphalea fruti- cosa Britt.) uo) or c to 340 Rhodora [Vol. 67 p. 560 Pavonia scabra (Vogel) Stehlé & Quentin (Malache scabra B. Vogel) p. 561 Pavonia paniculata Cav. (Lebretonia paniculata Britt.) p. 564 Hibiscus tiliaceus L. (Parit tiliaceaum St. Hil.) p. 567 Gossypium arboreum L. var. nadam (Watt.) Prokh. (Gos- sypium barbadense L.) p. 567 Malvaviscus arboreus Cav. (Malvaviscus malvaviscus Britt. & Wils.) To be added: St. Themas. BOMBACACEAE Ref.: Bull. Jard. Bot. Bruxelles XXXIII: 1-315. 1963. p. 570 Pachira insignis (Sw.) Sw. (Pachira aquatica Aubl.) STERCULIACEAE p. 571 Melochia tomentosa L. (Moluchia tomentosa Britt.) p. 572 Melochia pyramidata L. (Moluchia pyramidata Britt.) p. 572 Melochia villosa (Mill) Fawc. & Rendle (Riedlea hirsuta DC.) 3. Waltheria indica L. (Waltheria americana L.) p. 974 Ayenia insularis Cristobal (Ayenia pusilla L., as to the plant). See Opera Lilloana IV: 1-230, 1960. p. 575 Guazuma ulmifolia Lam. (Guazuma guazuma Cockerell) OCHNACEAE Ref.: DWYER, J. D.—Lloydia 7: 121-145. 1944. p. 578 Ouratea striata (v. Tiegh.) Urb. To be added: Guavate. p. 578 Ouratea ilicifolia (DC.) Baill. Virgin Islands. THEACEAE Ref.: KOBUSKI, CLARENCE E.—Journ. Arnold Arb. XXII: 395-416. 1941; XXIV: 60-76. 1945; XXX: 166-186. 1949. p. 580 Ternstroemia peduncularis DC. (Taonabo peduncularis Britt.) p. 580 Ternstroemia luquillensis Krug & Urb. (Taonabo luquillensis Britt.) p. 581 Ternstroemia heptasepala Krug & Urb. (Taonabo heptasepala Britt.) p. 581 Ternstroemia subsessilis (Britt.) Urb. (Taonabo subsessilis Britt.) p. 581 Ternstroemia stahlii Krug & Urb. (Taonabo stahlii Britt.; Taonabo pachyphylla Britt.) p. 582 Cleyera albopunctata (Griseb.) Krug & Urb. (Eroteum. albo- punctatum Britt.) p. 582 Laplacea portoricensis (Krug & Urb.) Dwyer (Haemocharis portoricensis Krug & Urb.) GUTTIFERAE (Clusiaceae, includes Hypericaceae) 1965] p. p: p. jp 584 584 588 597 597 Flora of Porto Rico — Liogier 341 Calophyllum calaba L. (Calophyllum antillanum Britt.) See Howarp, R. A.—Journ. Arnold Arb. XLIII: 397-398. 1962. Rheedia portoricensis Urb. [Rheedia acuminata (Spreng.) Tr. & PL, not Rheedia acuminata (R. & P.) Tr. & PL] Clusia grisebachiana (Pl. & Triana) Alain (Clusia krugiana Urb.) Hypericum hypericoides (L.) Crantz (Ascyrum hypericoides L.) COCHLOSPERMACEAE Cochlospermum vitifolium (Willd.) Spreng. (Maximiliania vitifolia Krug & Urb.) FLACOURTIACEAE Homalium racemosum Jacq. (Homalium pleiandrum Blake; Homalium leiogynum Blake) Xylosma schwaneckeanum (Krug & Urb.) Urb. (Myroxylon schwaneckeanum Krug & Urb.) Xylosma paehyphyllum (Krug & Urb.) Urb. (Myroxylon pachyphyllum Krug & Urb.) Xylosma buxifolium A. Gray (Myroxylon buxifolium Krug & Urb.) Xylosma schaefferioides A. Gray. To be added: Maricao. Laetia procera (Poepp. & Endl.) Eichl. (Casearia bicolor Urb.) Lunania buchii Urb. To be added: Maricao. VIOLACEAE Hybanthus linearifolius (Vahl) Urb. (lonidium linearifolium Britt.) Hybanthus portoricensis Urb. (lonidium portoricense Krug & Urb.) PASSIFLORACEAE Ref.: Kittie, E. P.—Field Mus. Nat. Hist. Bot. Ser. XIX. 1938. p. p. p. p. 602 602 602 604 . 608 612 617 Passiflora murucuja L. To be added: Quebradillas. See Sci. Surv. VI: 363. Passiflora anadenia Urb. To be added: Guanica. Passiflora suberosa L. (Passiflora pallida L.) Passiflora edulis Sims. To be added: Bayamón. CACTACEAE Pereskia aculeata (L.) Mill. (Pereskia pereskia Karst.) Pilosocereus royeni (L.) Byles & Rowley (Cephalocereus royeni Britt. & Rose) Melocactus intortus (Mill.) Urb. (Cactus intortus Mill.) 342 Rhodora [Vol. 67 p. 617 Mammillaria nivosa Link (Neomammillaria nivosa Britt. & Rose) SCIENTIFIC SURVEY OF PUERTO RICO AND THE VIRGIN ISLANDS. VOLUME VI. MELASTOMATACEAE Ref.: WILLIAMS, Lours O.—Fieldiana, Bot. 29 10: 545-586. 1963. p. 2 Acisanthera quadrata Juss. (Acisanthera acisanthera Britt.) p. 5 Miconia serrulata (DC.) Naud. (Tamonea macrophylla Krasser) p. 6 Miconia mirabilis (Aubl.) L. O. Wms. (Tamonea guianensis Aubl.) p. 7 Miconia microcarpa DC. (Miconia affinis DC.) To be added: Graffenrieda ottoschulzii (Urb. & Ekm.) Urb. & Ekm. (Miconia ottoschulzii Urb. & Ekm): Isabón. Cf. Sci. Surv. VI: 555. p. 15 Henriettea macfadyenii (Triana) Alain (Henriettella mac- fadyenii Triana) p. 15 Henriettea membranifolia (Cogn.) Alain (Henriettella mem- branifolia Cogn.) p. 16 Henriettea fascicularis (Sw.) G. Maza (Henriettella fascicu- laris Wr. ex Sauv.) p. 16 Henriettea triflora (Vahl) Alain. To be added: Guavate. LYTHRACEAE p. 20 Cuphea parsonsia (L.) R. Br. (Parsonsia parsonsia Britt.) p. 21 Cuphea micrantha HBK. (Parsonsia micrantha Jennings) p. 21 Lagerstroemia speciosa (J. A. Murr.) Pers. (correct author citation) COMBRETACEAE (Terminaliaceae, p. 22) MYRTACEAE Ref.: McVauGH, RoGERS.—Fieldiana, Bot. 29 (8): 395-532. 1968. p. 26 Psidium amplexicaule Pers. Add locality: Utuado, Puerto Rico. p. 26 Psidium cattleyanum Sabine, Add Locality: Utuado, Puerto Rico. An escape. p. 27 Pimenta racemosa (Mill) J. W. Moore (Amomis caryo- phyllata Krug & Urb.) p. 28 Pimenta racemosa var. grisea (Kiaersk.) Fosberg (Amomis grisea Britt.) p. 28 Aulomyrcia citrifolia (Aubl.) Amsh. (Myrcia citrifolia Urb.; Eugenia saviaefolia Alain). 1965] Flora of Porto Rico — Liogier 343 28 Aulomyrcia leptoclada (DC.) Berg (Myrcia leptoclada DC.) 29 Myrcia fallax (Rich.) DC. (Myrcia berberis DC.) 30 Siphoneugenia densiflora Berg (Plinia dussii Urb.) 31 Calyptranthes pallens (Poir.) Griseb. To be added: Maricao, Guavate. To be added: Calyptranthes triflorum Alain. Maricao. Bull. Torrey Bot. Club 90: 189. 1963. Calyptranthes peduncularis Alain. Maricao. Bull. Torrey Club 90: 189. 1963. Calyptranthes luquillensis Alain. Luquillo Mts. Bull. Torrey Club 90: 189. 1963. Calyptranthes dumetorum Alain. Susua. 34 Eugenia domingensis Berg. (Eugenia aeruginea DC.) p. 85 Eugenia maleolens Poir. (Eugenia buxifolia Willd.) To be added: Eugenia farjardensis Krug & Urb. Fapardo, Dorado. Symb. Ant. 9: 109. 1923. Eugenia margarettae Alain. Doña Juana. Bull. Tor- rey Bot. Club 90: 190. 1923. Eugenia haematocarpa Alain. Luquillo Mts. Bull. Torrey Club 90: 190. 1923. Eugenia glabrata (Sw.) DC. Maricao. p. 39 Myrciaria floribunda (West) Berg (Eugenia floribunda West) p. 41 Jambosa jambos (L.) Millsp. (Jambos jambos Millsp.) p. 42 Myrcianthes fragrans (Sw.) McVaugh (Anamomis fragrans Griseb.) p. 42 Pseudanamomis umbellulifera (Britt.) Kassel (Anamomis umbellulifera Britt.) The presence of this plant in Puerto Rico has been confirmed. p. 42 Pimenta dioica (L.) Merrill (Pimenta pimenta Cockerell). Escaped: Bayamón. RHIZOPHORACEAE p. 44 Cassipourea guianensis Aubl. (Cassipourea alba Griseb.) ONAGRACEAE Ref.: Hara, HiRosur.—4Journ. Jap. Bot. 28: 289-294. RAvEN, PETER H.—Kew Bull. 15: 476. 1961. p. 45 Ludwigia palustris (L.) Elliot (Jsnardia palustris L.) p. 45 Ludwigia peploides (HBK.) Raven (Jussiaea repens L.) p. 45 Ludwigia leptocarpa (Nutt) H. Hara (Jussiaea leptocarpa Nutt.) p. 46 Ludwigia erecta (L.) H. Hara (Jussiaea erecta L.) p. 46 Ludwigia peruviana (L.) H. Hara (Jussiaea peruviana L.) p. 46 Ludwigia octovalvis (Jacq.) Raven (Jussiaea angustifolia Lam.) MTM M > 344 Rhodora [Vol. 67 ARALIACEAE p. 48 Didymopanax gleasonii Britt. & Wils. Sci. Surv. VI: 365, 574. To be added: Cerro de Punta, Maricao. UMBELLIFERAE (Ammiaceac) p. 51 Hydrocotyle verticillata var. triradiata (A, Rich.) Fern. To be added. p. 51 Hydrocotyle bonariensis Lam. To be added. p. 51 Centella erecta (L. f.) Fern. (Centella asiatica Urb.) p. 52 Apium leptophyllum (Pers.) F. Muell. (Cyclospermum lepto- phyllum Sprague) p. 52 Petroselinum crispum (Mill.) Mansfeld (Apium petroselinum L.) p. 52 Foeniculum vulgare Mill. (Foeniculum foeniculum Karst.) p. 59 Anthriscus cerefolium (L.) Hoffm. (Cerefolium cerefolium Britt.) ERICACEAE (includes Vacciniaceae) p. 54 Lyonia rubiginosa (Pers.) G. Don (Xolisma rubiginosa Small) p. 54 Lyonia stahlii Urb. (Xolisma stahlii Small) p. 55 Hornemannia racemosa Vahl (Thibaudia krugii Urb. & Hoerold) p. 56 Gonocalyx portoricensis (Urb.) A. C. Smith (Ceratostoma portoricensis Hoerold) MYRSINACEAE p. 57 Ardisia glauciflora Urb. (Jcacorea glauciflora Britt.) p. 57 Ardisia obovata Desv. (Icacorea guadalupensis Britt. & Wils.) p. 58 Ardisia luquillensis (Britt.) Alain (leacorea luquillensis Britt.) p. 59 Wallenia yunquensis (Urb.) Mez (Petesioides yunquense Britt.) p. 99 Wallenia pendula (Urb.) Mez (Petesioides pendulum Britt.) THEOPHRASTACEAE p. 62 Jacquinia stenophylla Urb. To be added: Guanica. SAPOTACEAE Ref.: CRONQUIST, ARTHUR.—Bull. Torrey Club 72: 191-204. 1945; 72: 550-562. 1945; 73: 465-471. 1946; Lloydia 9: 241-292. 1946; Journ. Arnold Arb. XXVI: 485-471, 1945. p. 65 Manilkara zapota (L.) v. Royen (Sapota achras Mill.) p. 65 Mieropholis curvata Urb. is synonym to Micropholis chryso- phyiloides Pierre. 1965] p. 66 Ds Ol p. 67 O S PAOI p. 68 p. 70 Dare ek pDJ(2 DAT DA hon 9 p. T4 DATTA DONS p. 78 nou p. 79 De ag p79 Flora of Porto Rico — Liogier 345 Mastichodendron foetidissimum (Jacq.) Cron. (Sideroxylon foetidissimum Jacq.; Sideroxylon portoricense Urb., No. 2). Pouteria multiflora (A. DC.) Baehni (Lucuma multiflora A. DC.) Pouteria mammosa (L.) Cron. (Achras zapota L.) Pouteria dictyoneura (Griseb.) Radlk. ssp. fuertesii (Urb.) Cron.(Paralabatia portoricensis Britt. & Wils.) To be added: Limestone hills, N. Puerto Rico. See Sci. Surv. VI: 556. Pouteria hotteana (Urb. & Ekm.) Baehni. To be added: Maricao. Chrysophyllum bicolor Poir. (Chrysophyllum eggersti Pierre, No. 4. p. 69) Dipholis cubensis (Griseb.) Pierre (Dipholis sintenisii Pierre) Bumelia obovata var. krugii (Pierre) Cron. (Bumelia krugii Pierre) Manilkara balata (Aubl.) Dubard (Manilkara nitida Dubard; Manilkara bidentata Cher.) Manilkara pleeana (Pierre) Cron. (Manilkara duplicata Dubard) Manilkara jaimiqui (Wr. ex Griseb.) Dubard. To be added: Lajas to Parguera. See Sci. Surv. VI: 366. (Manilkara emarginata Britt. & Wils.) EBENACEAE Diospyros sintenisii (Krug & Urb.) Standl. (Maba sintenisii Krug & Urb.) Diospyros digyna Jacq. (Diospyros ebenaster Retz.). See: Howard, Richard A.—Journ. Arnold Arb. XLII: 430. 1961; XLIII: 101. 1962. Diospyros philippensis (Desrousseaux) Guercke (Diospyros discolor Willd.) OLEACEAE Haenianthus salicifolius var. obovatus (Krug & Urb.) Knobl. (Haenianthus obovatus Krug & Urb.) Linociera domingensis (Lam.) Knobl. (Mayepea domingen- sis Krug & Urb.) Linociera caribaea (Jacq.) Knobl. (Mayepea caribaea Kuntze) Linociera axilliflora Griseb. (Mayepea axilliflora Krug & Urb.) Linociera ligustrina Sw. To be added: El Jobo, San Antonio. Linociera holdridgii Camp & Monachino, To be added: Guanica. Lloydia 2: 223. 1939. 346 p. 85 Ref.: p. 88 p. 89 p. 90 p. 91 p. 91 p. 92 p. 94 p. 94 p. 94 p. 96 p. 97 p. 97 p. 97 p. 97 p. 97 p. 98 p. 99 p. 99 p. 100 p. 100 Rhodora [Vol. 67 Menyanthaceae in GENTIANACEAE. APOCYNACEAE Woopson, ROBERT E.—N. Amer. Flora 29 (2). 1938. Plumeria obtusa L. (Plumeria portoricensis Urb.; Plumeria krugii Urb.) Tabernaemontana divaricata (L.) R. Br. (Tabernaemontana coronaria Willd.) Rauvolfia nitida Jacq. (Rawvolfia tetraphylla L.) Rauvolfia viridis R. & S. (Rauvolfia lamarckii A. DC.) Thevetia peruviana (Pers.) K. Schum. (Cerbera thevetia L.) Prestonia agglutinata (Jacq.) Woods. (Echites agglutinata Jacq.) Forsteronia portoricensis Woods. To be added: common in the mountains. Forsteronia corymbosa G. F. W. Meyer is represented by a doubtful specimen. Carissa macrocarpa (Ecklon) A. DC. (Arduina grandiflora E. Meyer) ASCLEPIADACEAE Cynanchum ephedroides (Griseb.) Alain (Metastelma ephe- droides Schltr.) Cynanchum monense (Britt.) Alain (Metastelma monense Britt.) Cynanchum lineare (Bello) Alain (Metastelma lineare Bello) Cynanchum anegadense (Britt.) Alain (Metastelma anega- dense Britt.) Cynanchum parviflorum Sw. (Metastelma parviflorum R. Br.) Cynanchum cheesmanii Woods. (Metastelma decipiens Schltr.) Cynanchum grisebachianum (Schltr.) Alain, comb. nov. (Met- astelma grisebachianam Schltr. in Urb. Symb. Ant. 5: 469. 1908; Metastelma decaisneanum Schltr. in Urb. Symb. Ant. 1: 250. 1899, not Cynanchum decaisneanum R. Holm, 1953; Cynanchum decaisneanum Alain, Mem. Soc. Cub. Hist. Nat. 22: 119. 1955.) Matelea maritima (Jacq.) Woods. (Ibatia maritima Dene.) Sarcostemma clausum (Jacq.) R. & S. (Funastrum clausum Schltr.) Gonolobus stephanotrichus Griseb. (Vincetoxicum stephano- trichum Britt.) Matelea variifolia (Schltr.) Woods. (Vincetoxicum varii- folium Britt.) 1965] Flora of Porto Rico — Liogier 347 p. 100 Matelea sintenisii (Schltr.) Woods. (Vincetoxicum sintenisit Britt.) CONVOLVULACEAE (includes Cuscutaceae) Ref.: OosTsTROM, S. J. van.—A Monograph of the genus Evolvulus. Meded. Mus. Herb. Univ. Utrecht 14: 1-267. 1934. O’DONELL, C. A.—Revisión de las especies americanas de Merremia. Lilloa VI: 467-554. 1941. p. 104 Evolvulus alsinoides L. var. linifolius (L.) Baker (Evolvulus linifolius L.) St. Thomas, St. Croix. p. 105 Evolvulus tenuis Mart. var. longifolius (Choisy) v. Ooststr. (Evolvulus bocasanus Britt.) p. 106 Jaequemontia obcordata (Millsp.) House (Jacquemontia sub- salina Britt.) p. 107 Jacquemontia tamnifolia (L.) Griseb. (Thyella tamnifolia Raf.) p. 107 Jacquemontia canescens (HBK.) Benth. To be added: Guanica. . 107 Ipomoea alba L. (Calonyction aculeatum House) . 108 Ipomoea tuba (Schltr.) G. Don (Calonyction tuba Colla) 108 Ipomoea quamoclit L. (Quamoclit quamoclit Britt.) 108 Ipomoea hederifolia L. (Quamoclit coccinea Moench) Ipomoea solanifolia L. (Exogonium solanifolium Britt.) 109 Ipomoea repanda Jacq. (Exogonium repandum Choisy) 110 Ipomoea steudelii Millsp. (Hxogonium arenarium Choisy) 111 Merremia umbellata (L.) Hall. f. (Ipomoea polyanthes R. & S.) p. 111 Merremia dissecta (Jacq.) Hall. f. (Ipomoea dissecta Pursh) p. 111 Merremia aegyptia (L.) Urb. (Ipomoea aegyptia L.) p. 112 Ipomoea setifera Poir. (Ipomoea rubra Millsp.) Add syno- nym: Ipomoea palustris Urb. Cf. Sci Surv. VI: 368 p. 112 Ipomoea acuminata (Vahl) R. & S. (Ipomoea cathartica Poir. in Lam.) p. 113 Ipomoea purpurea A. W. Roth; correct author citation. p. 113 Ipomoea pes-caprae var. brasiliensis (L.) v. Oostr. To be added: St. Thomas. p. 114 Merremia quinquefolia (L.) Hall. f. (Ipomoea quinquefolia L.) p. 114 Ipomoea spirillus House (Jpomoea heptaphylla Vast) p. 115 Merremia tridentata (L.) Hall. f. ssp. angustifolia (Jacq.) v. Oostr. (Ipomoea angustifolia Jacq.) p. 116 Ipomoea batatas (L.) L.; correct author citation. p. 117 Ipomoea violacea L. (Ipomoea tricolor Cav.) Cf. Blumea III: 541. 1940. p. 117 Ipomoea ochroleuca Spanoghe (Ipomoea kentrocarpa Hochst.) GkKKEKKEKKER m © e 348 . 118 . 119 . 121 . 121 p. 121 . 122 Rhodora [Vol. 67 To be added: San German. Merremia tuberosa (L.) Rendle (Operculina tuberosa Meissn. in Mart.) Stictocardia tiliaefolia (Desr.) Hall f. (Rivea campanulata House) Cuscuta obtusiflora HBK. var. glandulosa Engelm. (Cuscuta glandulosa Small) Cuscuta campestris Yuncker (Cuscuta pentagonia Engelm.) Cuscuta globulosa Benth. To be added: quite abundant, in Puerto Rico and the Virgin Islands. HYDROPHYLLACEAE Nama jamaicensis L. (Marilaunidium jamaicense Kuntze) BORAGINACEAE (includes Ehretiaceae) Cordia alliodora (R. & P.) Oken (Cerdana alliodora R. & P.) Cordia alba (Jacq.) R. & S. (Calyptracordia alba Britt.) Cordia gerascanthus L. To be added: Yauco Cordia sebestena L. (Sebesten sebestena Britt. & Small) Cordia rickseckeri Millsp. (Sebesten rickseckeri Britt.) Cordia collococea L. (Cordia glabra L.) Cordia stenophylla Alain (Varronia angustifolia West) Cordia polycephala (Lam.) Johnst. (Varronia corymbosa Desv., pro parte) Cordia globosa (Jacq.) HBK. (Varronia globosa Jacq.) Cordia bahamensis Urb. (Varronia bahamensis Millsp.) Cordia lima (Desv.) R. & S. (Varronia lima Desv.) Cordia rupicola Urb. (Varronia rupicola Britt.) Cordia bellonis Urb. (Varronia bellonis Britt.) Bourreria succulenta var. revoluta (HBK.) O. E. Schult. (Bourreria revoluta HBK.) Tournefortia gnaphalodes (L.) R. Br. (Mallotonia gnapha- lodes Britt.) Tournefortia maculata Jacq. (Tournefortia peruviana Poir.; Tournefortia laurifolia Vent.) Tournefortia volubilis L. Add synonym: Tournefortia miero- phylla Bert. ex Spreng. No. 7. Heliotropium indicum L. (Tiaridium indicum Lehm.) Heliotropium angiospermum Murray (Schobera angiosperma Britt.) Heliotropium procumbens Mill. (Heliotropium inundatum Sw.) Heliotropium microphyllum Sw. (Heliotropium crispiflorum Urb.) 1965] Flora of Porto Rico — Liogier 349 p. 137 Heliotropium arborescens L. (Heliotropium peruvianum L.) p. 137 Heliotropium amplexicaule Vahl (Cochranea anchusaef olia Gürcke; Heliophytum amplexicaule Britt. & Wils.) To be added: Cynoglossum furcatum Wall. in Roxb. Maricao. VERBENACEAE Ref.: MoLpENKE, HAROLD, N.— The known geographic distribution of the Verbenaceae, etc, New York, 1949. Suppl. 1-13. p. 138 Verbena tenuisecta Briq. To be added, cited by Moldenke, perhaps only cultivated. p. 138 Verbena bonariensis L. To be added: Guavate. p. 139 Ghinia boxiana Mold. (Ghinia spinosa Britt. & Wils., as to the Puerto Rican plant) p. 139 Lantana camara var. aculeata (L.) Mold. (Lantana aculeata L.) p. 139 Lantana camara var. mista (L.) L. H. Bailey. To be added: Bayamón, Mayagüez, St. Thomas. p. 1389 Lantana camara var. mutabilis (Hook.) L. H. Bailey. To be added: Trujillo. p. 139 Lantana camara var. sanguinea L. H. Bailey. To be added, cited by Moldenke. p. 189 Lantana camara f. parvifolia Mold. To be added: Bayamón. p. 139 Lantana strigosa (Griseb.) Urb. To be added: S. Puerto Rico. p. 189 Lantana fucata var. antillana Mold. To be added: San Juan. p. 139 Lantana reticulata Pers. To be added: S. Puerto Rico. p. 139 Lantana arida Britt. To be added: abundant in P. Rico; St. Croix. p. 139 Lantana insularis Mold. To be added: St. Croix. p. 139 Lantana involucrata var. odorata (L.) Mold. To be added: P. Rico, St. Croix, St. Thomas. p. 139 Lantana montevidensis (Spreng.) Briq. To be added: P. Rico; St. Thomas. p. 141 Lippia micromera Schau var. helleri (Britt.) Mold. (Lippia helleri Britt.) p. 142 Phyla nodiflora (L.) Greene (Lippia nodiflora Michx.) p. 142 Phyla nodiflora var. reptans (HBK.) Mold. (Lippia reptans HBK.) p. 142 Phyla stoechadifolia (L.) Small (Lippia stoechadifolia HBK.) p. 142 Phyla scaberrima ( A. L. Juss.) Mold. To be added, cited by Moldenke p. 142 Phyla strigulosa f. parvifolia (Mold.) Mold. To be added: S. P. Rico, St. Croix. p. 143 Bouchea prismatica var. longirostra Grenz. To be added: Ponce. p. p. Rhodora (Vol 67 Bouchea prismatica var. brevirostra Grenz. To be added: St. Thomas. Stachytarpheta cayennensis (L. C. Rich.) Vahl (Valeria- noides cayennense Kuntze) Stachytarpheta cayennensis var. albiflora Mold. To be added: Pueblo Viejo. Stachytarpheta X hybrida Mold. To be added: S. P. Rico, Culebrita. Stachytarpheta urticaefolia (Salisb.) Sims. To be added: Mayagiiez. Stachytarpheta jamaicensis (L.) Vahl (Valeranoides jam- aicense Kuntze) Stachytarpheta strigosa Vahl (Valerianoides strigosum Britt.) Citharexylum fruticosum var. subvillosum Mold. To be added: St. Thomas, Culebra. Citharexylum fruticosum var. villosum (Jacq.) O. E. Schulz. To be added: Puerto Rico, Virgin Gorda, Tortola. Citharexylum pentandrum Vent. To be added: in Herb. Ventenat. Citharexylum X perkinsii Mold. To be added: Cerro de Punta. Citharexylum tristachyum Turez. To be added: St. Thomas. Aegiphila elata Sw. To be added, cited by Moldenke. Vitex parviflora A. L. Juss. To be added: Mayagüez to Maricao. Vitex negundo var. intermedia (P’ei) Mold. To be added: St. Croix. Clerodendrum aculeatum (L.) Schlecht. (Volkameria aculeata L.) Clerodendrum bungei Steud. To be added: Vega Baja, probably an escape. Clerodendrum indicum (L.) Kuntze (Siphonanthus indicus L.) Clerodendrum inerme (L.) Gaertn. To be added: St. Croix. Avicennia germinans (L.) L. (Avicennia nitida Jacq.) Congea tomentosa Roxb. To be added: Trujillo, Cultivated. Petrea kohautiana Presl. To be added: Mayagiiez, St. Thomas. Cultivated. Tektona grandis L. (Tectona grandis incorrect spelling). LABIATAE (Lamiaceae) Salvia misella Kunth. To be added: Wydler (1826). Salvia micrantha Vahl. To be added: St. Croix, Mona Island. Hyptis spicigera Lam. (Hyptis americana Urb.) Hyptis escobilla Urb. To be added: Bayamón. 1965] p. PPP 159 . 163 166 . 167 169 170 171 171 171 171 172 172 . 173 LIS ETT . 174 SE bs: . 176 176 . 181 191 Flora of Porto Rico — Liogier 351 Hyptis scoparia Poit. To be added: Guilarte. SOLANACEAE Lyeium tweedianum Griseb. var. chrysocarpum (Urb. & Ekm.) L. C. Hitche. (Lycium americanum Jacq.) Solanum caribaeum Dunal. To be added: Comerio. Solanum erianthum D. Don (Solanum verbascifolium L.) Solanum campechiense L. (Solanum guanicense Urb.) Solanum ficifolium Ort. (Solanum torvum Sw.) Solanum elaegnifolium Cav. To be added: Ensenada. Solanum antillarum O. E. Schulz. To be added: Guilarte. Solanum ciliatum Lam. To be added: Maricao. Solanum pterocaulon Dunal. To be added. Lycopersicon lycopersicum (L.) Karst. (Lycopersicon lycoper- sicon Karst.) Saracha antillana Krug & Urb. To be added: Cerro de Punta. Solandra grandiflora Sw. (Swartzia grandiflora Gmel.) Datura suaveolens H. & B. (Brugmansia suaveolens Bercht. & Presl) Datura candida (Pers.) Safford (Brugmansia arborea Steud.) Datura inoxia Mill. (Datura metel L. of Britt. & Wils.) (aura metel L. (Datura fastuosa L. of Britton & Wilson) Cestrum salicifolium Dunal. To be added: Bayamón. Cf. Candollea 6: 360. 1935. Cestrum daphnoides Griseb. To be added: St. Thomas. SCROPHULARIACEAE Mecardonia dianthera (Sw.) Pennell (Mecardonia procum- bens Small), as to the plants from Hispaniola southward in the Lesser Antilles. Bacopa repens (Sw.) Wettst. (Macuillamia repens Pennell) Bacopa monnieri (L.) Pennell (Bramia monnieri Drake) Bacopa stricta (Schrad.) Robins. (Caconapea stricta Britt.) Bacopa innominata (Maza) Alain (Herpestis rotundifolia Gaertn. f.) Lindernia diffusa (L.) Wettst. (Vandellia diffusa L.) Lindernia dubia (L.) Wettst. (Ilysanthes dubia Barnhart) Lindernia crustacea (L.) F. Muell. To be added: common in waste grounds. Micranthemum umbrosum (Walt.) Blake (Globifera umbrosa J. F. Gmel.) LENTIBULARIACEAE Utricularia subulata L. (Setiscapella subulata Barnh.) p. . 199) 202 p. 224 Rhodora [Vol. 67 Utricularia pusilla Vahl (Setiscapella pusilla Barnh.) Utricularia juncea Vahl (Stomoisia juncea Barnh.) BIGNONIACEAE Doxantha unguis-cati (L.) Rehder (Batocydia unguis Mart.) Tabebuia pallida Miers (Tabebuia heterophylla Britt., No. 4). Schlegelia brachyantha Griseb. (Sehlegelia portoricensis Britt.) Tynnanthus caryophylleus (Bello) Alain, comb. nov. (Big- nonia caryophyllea Bello, Anal. Soc. Esp. Hist. Nat. 10: 293. 1881; Tynnanthus myrianthus Bur. & K. Schum in Mart. Fl. Bras. 8 (2): 197. 1896). Catalpa longissima (Jacq.) Dum.-Cours. To be added: St. Thomas. PEDALIACEAE Sesamum indicum L. (Sesamum orientale L.) GESNERIACEAE Alloplectus ambiguus Urb. (Crantzia ambigua Britt.) Cf. Contr. U. S. Herb, 29: 16. 1944. Gesneria sintenisii Urb. (Duchartrea sintenisii Britt.) Gesneria albiflora (Dene.) Kuntze (Pentarhaphia albiflora Dene.) ACANTHACEAE Blechum pyramidatum (Lam.) Urb. (Blechum blechum Millsp.) Ruellia tweediana Griseb. To be added: common in western Puerto Rico. Teliostachya alopecuroidea (Vahl) Nees (Lepidagathis alope- curoidea R. Br.) Stenandrium tuberosum (L.) Urb. (Gerardia tuberosa L.; Gerardia portoricensis Britt. & Wils., No. 2.) Dicliptera assurgens (L.) Juss. (Diapedium assurgens Kuntze) Dicliptera krugii Urb. (Diapedium krugii Britt.) Justicia pectoralis Jacq. (Stethoma pectoralis Raf.) Justicia verticillaris (Nees) Urb. (Stethoma verticillaris Britt.) Justicia comata (L.) Lam. (Stethoma comata Britt.) added: Asvstasia gangetica (L.) T. Anders. Escaped and common. RUBIACEAE Oldenlandia corymbosa L. To be added to Puerto Rico: San Flora of Porto Rico — Liogier 353 Juan, Bayamón. Cf. Sci. Surv. VI: 563. Oldenlandia lancifolia (K. Schum.) DC. (Oldenlandia herba- cea DC.) Oldenlandia callitrichoides Griseb. To be added: Bayamón. Lucya tetrandra (L.) K. Schum. (Clavenna tetrandra Standl.) Phialanthus myrtilloides Griseb. To be added: Maricao, Susua. Phialanthus grandifolius Alain. To be added: Maricao. Exostema ellipticum Griseb. To be added: Maricao, Villalba. Cf. Sci. Surv. VI: 371. Schradera vahlii Steyerm. (Urceolaria exotica Gmelin, pro parte). Cf. Mem. N. Y. Bot. Gard. 10: 277. 1963. Gonzalagunia spicata (Lam.) Maza (Duggena hirsuta Britt.) Coccocypselum herbaceum Aubl. (T'ontanea herbacea Standl.) Randia aculeata L. (Randia mitis L.) Hamelia patens Jacq. (Hamelia erecta Jacq.) Guettarda valenzuelana A. Rich. (Guettarda laevis Urb.) Antirhea obtusifolia Urb. (Stenostomum obtusifolium Britt. & Wils.) Antirhea coriacea (Vahl) Urb. (Stenostomum coriaceum Griseb.) Antirhea lucida (Sw.) Benth. & Hook. (Stenostomum lucidum Gaertn. f.) Antirhea portoricensis (Britt. & Wils.) Standl. (Stenostomum portoricense Britt. & Wils. Sci. Surv. VI: 564. 1930). To be added: Limestone hills, N. Puerto Rico. Antirhea sintenisii Urb. (Stenostomum sintenisii Britt. & Wils.) Antirhea acutata (DC.) Urb. (Stenostomum acutatum DC.) Terebraria resinosa (Vahl) Sprague (Laugeria resinosa Vahl) Chione seminervis Urb. & Ekm. To be added: Cerro de Punta, Jayuya. Seolosanthus multiflorus (Sw.) Krug & Urb. (Scolosanthus grandifolius Krug & Urb.) Seolosanthus versicolor Vahl (Scosanthus versicolor, in- correct spelling) Scolosanthus densiflorus Urb. To be added: Mariaco. Psychotria microdon (DC.) Urb. (Psychotria pinularis Sessé & Moc.) Palicourea crocea (Sw.) R. & S. Add synonym: (Palicourea brevithyrsa Britt. & Standl.) Lasianthus lanceolatus (Griseb.) Urb. (Lasianthus moralesit Wr. ex Sauv.) 354 Rhodora (Vol, 67 p. p. p. p. p. p. . 251 Geophila repens (L.) I. M. Johnst. (Geophila herbacea Schum.) 253 Richardia brasiliensis Gomez. To be added: Cerro Gordo. 254 Diodia serrulata (P. Beauv.) G. Tayl. (Diodia maritima Thonn.) 254 Hemidiodia ocymifolia (Willd. ex R. & S.) K. Schum. Cor- rect spelling and author citation. 255 Borreria brachysepala Urb. (Borreria laevis var. sintenisii Urb.). To be added . 256 Spermacoce confusa Rendle (Spermacoce tenuior L., sensu Britt. & Wils.) 256 Spermacoce tenuior L. (Spermacoce riparia C. E S.) 258 Mitracarpus villosus (Sw.) DC. (Mitracarpus hirtus DC.) CUCURBITACEAE 262 Lagenaria siceraria (Molina) Standl. (Cucurbita lagenaria L.) . 264 Citrullus lanatus (Thunb.) Mansfeld (Citrullus citrullus Karst.) . 265 Cucurbita moschata (Duch.) Duchesne & Poir. (Pepo moschata Britt.) CAMPANULACEAE (includes Lobeliaceae as a sub-family) 272 Lobelia rotundifolia Juss. ex A. DC. To be added: Cerro de Punta. . 272 Lobelia assurgens var. portoricensis (A. DC.) Urb. (Tupa robusta A. DC.) 272 Lobelia portoricensis (Vatke) Urb. (Tupa portoricensis Vatke) . 273 Laurentia longiflora (L.) Peterm. (Isotoma longiflora. Presl.) COMPOSITAE (includes: Cichorieae, Ambrosiaceae, CarduaceaeQ . 276 Lactuca intybacea Jacq. (Brachyramphus intybaceus DC.) . 277 Xanthium strumarium L. (Xanthium chinense Mill.) . 279 Ambrosia confertiflora DC. To be added: Lajas. . 284 Piptocoma antillana Urb. (Piptocoma rufescens Cass., as to the Puerto-Rican plant) . 284 Elephantopus scaber L. (Elephantopus mollis HBK.) . 285 Pseudo-elephantopus spicatus (Aubl.) Gleas. Correct author citation. . 287 Eupatorium macrophyllum L. (Hebeclinium macrophyllum DC.) . 287 Eupatorium odoratum L. (Osmia odorata Sch.-Bip.) . 288 Eupatorium corymbosum Aubl. (Osmia corymbosa Britt. & Wils.) . 909 . 911 . 312 . 913 . 313 . 314 . 316 . 317 . 319 . 320 Flora of Porto Rico — Liogier 355 Eupatorium geraniifolium Urb. (Osmia geraniifolia Britt. & Wils.) Eupatorium sinuatum Lam. (Osmia sinuata Britt. & Wils.) Eupatorium ivaefolium L. (Osmia ivaefolia Sch.-Bip.) Eupatorium borinquense (Britt.) Robins. (Osmia borinquen- sis Britt.) Eupatorium portoricense Urb. (Critonia portoricensis Britt. & Wils.) added: Eupatorium iresinoides Kunth. Villalba. Eupatorium oteroi Monachino. Mona Island. Phyto- logia 2: 406. 1948. Mikania micrantha var. congesta (DC.) Robins. (Mikania con- gesta DC.) Conyza canadensis (L.) Cron. var. pusilla (Nutt.) Cron. (Leptilon pusillum Britt.) Conyza bonariensis (L.) Cron. (Leptilon bonariense Small) Conyza floribunda HBK. (Leptilon linifolium Small) Conyza apurensis HBK. (Leptilon chinense Britt.) Pluchea carolinensis (Jacq.) D. Don (Pluchea odorata Cass.) Gnaphalium americanum Mill. (Gnaphalium spicatum Lam.) Lagascea mollis Cav. (Nocca mollis Jacq.) Zinnia peruviana (L.) L. (Crassina elegans Kuntze; Cras- sina multiflora L.) Helichrysum bracteatum (Vent.) Willd. To be added: Toro Negro. Cf, p. 323. Melanthera nivea (L.) Small. Add to synonymy: Melanthera calcicola Britt.; Melanthera canescens O. E. Schulz; Melan- thera montana O. E. Schulz; Melanthera confusa Britt. Verbesina alata L. (Tepion alatum Britt.) Verbesina encelioides (Cav.) Benth. & Hook. (Ximenesia encelioides Cav.) Bidens cynapiifolia var. tenuis O. E. Schulz. To be added. Bidens cynapiifolia var. portoricensis (Spreng.) O. E. Schulz. To be added. Bidens reptans var. urbanii (Greenm.) O. E. Schulz (Bidens urbanii Greenm.) Flaveria trinervia (Spreng.) C. Mohr To be added: Guanica. Porophyllum ruderale (Jacq.) Cass. (Porophyllum porophyl- lum Kuntze) Pectis floribunda A. Rich. To be added: Puéblo Viejo. Cf. Sci. Surv. VI: 365. Erechtites valerianaefolia (Wolf) DC. To be added: Guavate, Maricao. 356 p. 324 . 326 Rhodora [Vol. 67 Chrysanthemum indicum L. (Chrysanthemum morifolium Ram.) TAXACEAE Podocarpus coriaceus L. C. Rich. (Nageia coriacea Kuntze) OPHIOGLOSSACEAE Ophioglossum palmatum L. (Cheiroglossa palmata Presl) SCHIZAEACEAE Schizaea pennula Sw. (Actinostachys pennula Hook.) Schizaea poeppigiana Sturm. in Mart. (Lophidium poeppigi- anum Underw.) e GLEICHENIACEAE Gleichena flexuosa (Schad.) Mett. (Dicranopteris flexuosa Underw.) Gleichenia pectinata (Willd.) Presl (Dicranopteris pectinata Underw.) Gleichenia bifida (Willd.) Spreng. (Dicranopteris bifida Maxon) CYATHEACEAE Cyathea horrida (L.) J. E. Sm. (Hemitelia horrida R. Br.) Cyathea wilsoni (Hook. in Hook. & Baker) Domin (Hemitelia wilsoni Hook.) Cyathea escuquensis (Karst.) Domin (Hemitelia escuquensis Karst.) Cyathea borinquena (Maxon) Domin (Alsophila borinquena Maxon) Cyathea aquilina (Christ) Domin (Alsophila aquilina Christ) Lophosoria quadripinnata (Gmel.) C. Chr. (Alsophila quad- ripinnata C. Chr.) POLYPODIACEAE Elaphoglossum pumilum (Mett.) C. Chr. (Elaphoglossum piloselloides Moore) Elaphoglossum rigidum (Aubl.) Urb. (Elaphoglossum flac- cidum Moore) Elaphoglossum martinicense (Desv.) Moore (Elaphoglossum underwoodianum Maxon) Add: Elaphoglossum longifolium (Jacq.) J. Sm. (Elaphoglos- sum rigidum Urb., sensu Maxon in Flora of Puerto Rico). Elaphoglossum maxonii Underw. & Maxon (Elaphoglossum pteropus C. Chr., as to the Puerto-Rican plant). Flora of Porto Rico — Liogier 357 Elaphoglossum serpens Maxon. To be added: Cerro de Punta, Jayuya. Elaphoglossum crinitum (L.) Christ (Hymenodium crinitum Fée) Peltapteris peltata (Sw.) Morton (Rhipidopteris peltata Schott) Vittaria graminifolia Kaulf. (Vittaria filifolia Fée) Polytaenium cajenense (Desv.) Benedict (Antrophytum cayennense Kaulf.). Determination confirmed; the plant be- longs to the Flora of Puerto Rico. Grammitis hessii (Maxon) Alain (Polypodium hessii Maxon) Grammitis serrulata (Sw.) Sw. (Polypodium duale Maxon) Grammitis myosuroides (Sw.) Sw. (Polypodium myosuroides Sw.) Grammitis trifurcata (L.) Copel. (Polypodium trifurcatum L.) Grammitis taenifolia (Jenm.) Proctor (Polypodium taeni- folium Jenm.) Grammitis asplenifolia (L.) Proctor (Polypodium aspleni- folium L.) Grammitis jubaeformis (Kaulf.) Proctor (Polypodium jubae- forme Kaulf.) Grammitis mollissima (Fée) Proctor (Polypodium mollissi- mum Fée). Grammitis taxifolia (L.) Proctor (Polypodium taxifolium L.) To be added: Grammitis suspensa (L.) Proctor. Luquillo Mts. Grammitis hartii (Jenm.) Proctor. Cerro de Punta, Jayuya. Polypodium phyllitidis var. latum (Moore) Proctor (Poly- podium latum Sodiro) Cheilanthes trichomanoides (L.) Mett. (Notholaena tricho- manoides R. Br.) Cheilanthes paupercula (Kuntze) Mett. (Adiantopsis pauper- cula Fée) Lonchitis hirsuta L. (Anisosorus hirsutus Underw. & Maxon) Pteridium aquilinum var. caudatum (L.) Urb. (Pteridium caudatum Maxon) Pteridium aquilinum var. arachnoideum (Kaulf.) Brade (Pteridium arachnoideum Maxon) Pityrogramme trifoliata (L.) Tryon (Trismeria trifoliata Diels) Diplazium cristatum (Desv.) Alston (Diplazium arboreum Presl) Blechnum polypodioides (Sw.) Kuhn (Struthiopteris poly- podioides Trev.) p. p. p. p. Rhodora [Vol. 67 Blechnum divergens (Kunze) Mett. (Struthiopteris exaltata Broadh.) Blechnum underwoodianum (Broadh.) C. Chr. (Struthiop- teris underwoodiana Broadh.) Blechnum lineatum (Sw.) C. Chr. (Struthiopteris lineata Broadh.) Lomariopsis amydrophlebia (Slosson ex Maxon) Holttum (Stenochlaena amydrophlebia Slosson ex Maxon) Lomariopsis kunzeana (Underw.) Holttum (Stenochlaena kunzeana Underw.) Lomariopsis sorbifolia (L.) Fée (Stenochlaena sorbifolia J. Smith) Bolbitis nicotianaefolia (Sw.) Alst. (Leptochilus nicotianae- folius C. Chr.) Bolbitis pergamentacea (Maxon) Ching (Leptochilus per- gamentaceus Maxon) Bolbitis cladorrhizans (Spreng.) Ching (Leptochilus clador- rhizans Maxon) Bolbitis aliena (Sw.) Alst. (Leptochilus alienus C. Chr.) Lomogramma guianensis (Aubl.) Ching (Leptochilus guian- ensis C. Chr.) Polystichum echinatum (Gmel.) C. Chr. (Polystichum tri- angulum Fée, as to the Puerto-Rican plant). Rumohra adiantiformis (Forst.) Ching (Polystichum adianta- forme J. Smith) Thelypteris opposita (Vahl) Ching (Dryopteris opposita Urb.) Thelypteris sancta (L.) Proctor (Dryopteris sancta Kuntze) Thelypteris sancta var. portoricensis (Kuhn) Morton. To be added. Thelypteris piedrensis (C. Chr.) Morton (Dryopteris pied- rensis C. Chr.) Thelypteris diplazioides (Moritz ex Mett.) Ching (Dryop- teris linkiana Maxon) Thelypteris germaniana (Fée) Proctor (Dryopteris ger- maniana C. Chr.) Thelypteris balbisii (Spreng.) Ching (Dryopteris sprengelii Kuntze) Thelypteris decussata (L.) Proctor (Dryopteris decussata Urb.) Thelypteris deltoidea (Sw.) Proctor (Dryopteris deltoidea Kuntze) Thelypteris gongylodes (Schkuhr) Small (Dryopteris gongy- lodes Kuntze) 1965] P Flora of Porto Rico — Liogier 359 Thelypteris dentata (Forsk.) E. St. John (Dryopteris dentata C. Chr.) Thelypteris patens (Sw.) Small (Dryopteris patens Kuntze) Thelypteris normalis (C. Chr.) Small (Dryopteris normalis C. Chr.) Thelypteris serra (Sw.) R. St. John (Dryopteris serra Kuntze) Thelypteris oligophylla (Maxon) Proctor (Dryopteris oligo- phylla Maxon) Thelypteris poiteana (Bory) Proctor (Dryopteris poiteana Urb.) Thelypteris tetragona (Sw.) Small (Dryopteris subtetragona Maxon) Thelypteris megalodus (Schkuhr) Proctor (Dryopteris meg- alodus Urb.) Thelypteris guadalupensis (Wikstr.) Proctor (Dryopteris domingensis Maxon) Thelypteris sclerophylla (Kuntze) Morton (Dryopteris sclero- phylla C. Chr.) Thelypteris reptans (J. F. Gmel.) Morton (Dryopteris reptans C. Chr.) Thelypteris brittonae (Slosson) Alain (Dryopteris brittonae Slosson) Thelypteris nephrodioides (Klotzsch) Proctor (Dryopteris guadalupensis Kuntze) Thelypteris hastata (Fée) Proctor (Dryopteris hastata Urb.) Thelypteris leptocladia (Fée) Proctor (Dryopteris leptocladia Maxon) Thelypteris angustifolia (Willd.) Proctor (Dryopteris angus- tifolia Urb.) Thelypteris serrata (Cav.) Alst. (Dryopteris serrata C. Chr.) Thelypteris reticulata (L.) Proctor (Dryopteris reticulata Urb.) Ctenitis hirta (Sw.) Copel. (Dryopteris hirta Kuntze) Ctenitis nemorosa (Willd.) Copel. (Dryopteris nemorosa Urb.) Ctenitis subincisa (Willd.) Copel. (Dryopteris subincisa Urb.) Polystichopsis chaerophylloides (Poir.) Morton (Dryopteris chaerophylloides C. Chr.) Ctenitis effusa (Sw.) Copel. (Dryopteris effusa Urb.) Tectaria incisa Cav. (Tectaria martinicensis Copel.) Orthiopteris domingensis (Spreng.) Copel. (Saccoloma dom- ingense Prantl.) 360 Rhodora [Vol. 67 p. 490 Orthiopteris inaequale (Kunze) Copel. (Saccoloma inaequale Mett.) p. 492 Dennstaedtia obtusifolia (Willd.) Moore (Dennstaedtia ordi- nata Moore) p. 492 Dennstaedtia bipinnata (Cav.) Maxon (Dennstaedtia adian- toides Moore) p. 498 Dennstaedtia cicutaria (Sw.) Moore (—Dennstaedtia rubi- ginosa Moore) HYMENOPHYLLACEAE Ref.: J. G. WESSELS BoER.—Acta Bot. Neerl. 11: 277-330. 1962. Morton, C. V.—Contr. U. S. Nat. Herb. 29 (3). 1947. p. 496 Trichomanes punctatum ssp. sphenoides (Kunze) W. Boer (Trichomanes sphenoides Kunze) To be added: Trichomanes robustum Fourn. Cerro de Punta, Jayuya. p. 497 Trichomanes angustifrons (Fée) W. Boer in Kramer (cited by Maxon as Trichomanes pusillum Sw.). Cidra, Puerto Rico. p. 497 Trichomanes pusillum Sw. To be added: Las Cruces; Andubo. Puerto Rico. To be added: Trichomanes ovale (Fourn.) W. Boer. Trichomanes kapplerianum Sturm. p. 506 Hymenophyllum hirsutum (L.) Sw. (Hymenophyllum cilia- tum. Sw.) To be added: Hymenophyllum axillare Sw, Toro Negro. Hymenophyllum elegantulum var. petiolulatum Morton Cerro de Punta. Hymenophyllum undulatum (Sw.) Sw. Cerro de Punta, Jayuya. Hymenophyllum lanatum Fée. Fide Maxon. Hymenophyllum sieberi (Presl) v.d. Bosch El Yunque, Luquillo Mts. LYCOPODIACEAE p. 516 Lycopodium sintenisii (Herter) Maxon (Urostachys sintenisii Herter) Determination and record confirmed for Puerto Rico. SELAGINELLACEAE p. 518 Selaginella armata Baker (Selaginella plagiochila Baker) p. 518 Selaginella plumosa (L.) C. Presl (Selaginella stolonifera Spring) p. 519 Selaginella substipitata Spring (Selaginella portoricensis A. Braun) p. 520 Selaginella tenella (Beauv.) Spring (Selaginella albonitens Spring) 1965] Carex incomperta — Hermann 361 p. 520 Selaginella subcaulescens Bak. (Selaginella sintenisii Hieron.) To be added: Selaginella flabellata (L.) Spring. Selaginella stipitata Spring. Toro Negro. THE NEW YORK BOTANICAL GARDEN BRONX, NEW YORK THE STATUS OF CAREX INCOMPERTA BICKN. F. J. HERMANN In 1908 Eugene P. Bicknell, equally well-known as a student of the New England flora and of its avifauna, de- scribed a sedge from Nantucket Island, Massachusetts, as Carex incomperta. It had been generally passing as C. steri- lis Willd., and it is still a problematical plant, showing affinities, as pointed out by Mackenzie (North American Flora 18(2) :107.1931), with such diverse allies as C. cepha- lantha, C. angustior, C. howei, C. muricata, C. interior, C. mohrii, C. atlantica, C. ruthii, C. ormantha, C. wiegandii and C. phyllomanica. By Mackenzie and most current au- thors it has been rightly treated as most nearly related to C. atlantica Bailey (1893). From this it has been held to differ chiefly in characteristics of the pistillate scales. These are detailed in the following key which is a composite of those of Mackenzie (op. cit.), Fernald (Gray's New Manual of Botany, 8th ed., 1952) and Gleason (New Britton and Brown Illustrated Flora, 1952). Pistillate scales sharply keeled, acutish to short-cuspidate, two- thirds to fully as long as the perigynium-body, the raised, prom- inent midrib extending to the tip; culms slender, sharply tri- angular, 1.5-2.5 mm. thick at the base; leaf-blades 1.5-2.5 mm. BTA Co REE M LI ON aN E ERE TIE C. incomperta. Pistillate scales flat or very obscurely keeled, obtuse, half to two- thirds as long as the perigynium-body, the midrib not raised, obsolete or nearly so at the tip; culms stiff, obtusely triangular below, 2-3.5 mm. thick at the base; leaf-blades 1.5-4 mm. wide .. Dp NR UU MEHRHEIT ER rere C. atlantica. 362 Rhodora [Vol. 67 Both plants occur in swamps and bogs, particularly in peaty soils, and have a broad range in eastern North Amer- ica, from Nova Scotia to Florida westward to Texas, but C. atlantica is found chiefly on the coastal plain, whereas C. incomperta extends inland to Michigan and Indiana. In their extreme forms these two taxa are fairly distinct, but it is not often that a specimen will be found to be con- sistent in all of its purported diagnostic characteristics, and throughout most of their over-lapping geographic ranges (particularly in southern New England, I am in- formed by Mr. Richard J. Eaton) the proportion of inter- mediates is bewildering. Since correlation of the distinctions is so frequently impossible, it would appear that varietal status would better represent the relationship of Carex incomperta to C. atlantica, as well as facilitating identifica- tion of the plants, so it is here proposed as Carex atlantica Bailey var. incomperta (Bickn.) F. J. Herm., comb. nov. (C. incomperta Bickn., Bull. Torrey Bot. Club 35: 494. 1908). FOREST SERVICE HERBARIUM RANGE AND WILDLIFE HABITAT RESEARCH U. 8. DEPARTMENT OF AGRICULTURE, WASHINGTON, D. C. THE DISTRIBUTION PATTERNS OF NORTHERN AND SOUTHERN ELEMENTS IN THE FLORA OF NEWFOUNDLAND' by A. W. H. DAMMAN INTRODUCTION The flora of the Island of Newfoundland has attracted much attention, mainly as a result of the explorations of Fernald (1911, 1918a, 1918b, 1924, 1925) in the beginning of this century. The vivid descriptions of his field trips, and especially his explanation of the origin of the New- foundland flora, have focused the attention of plant geog- raphers on the island and the general region of the Gulf of St. Lawrence. His theories have had a great impact on plant geographical thinking in America as well as Northern Europe. The most conspicuous features of the flora of Newfound- land Rouleau, 1956; Arsène, 1927) are the occurrence of many species with arctic affinities, the absence of many temperate-boreal species present on Cape Breton Island, and the common occurrence of the so-called Coastal Plain spe- cies, very often in close association with the arctic species. In addition, the flora contains a number of species which have their main distribution area in western North Amer- ica. This Cordilleran element is mainly restricted to the mountains and unstable soils of western Newfoundland. Most of the data on species distribution in Newfoundland, used in this paper, were obtained by personal observation and collection. However, I am grateful to Dr. E. Rouleau of the University of Montreal for showing me many of his un- published distribution maps of Newfoundland vascular plants. These maps were particularly helpful in checking ‘Department of Forestry, Canada, Forest Research Branch Con- tribution No... 363 364 Rhodora [Vol. 67 my observations on the absence of a species from certain parts of the island. Some published accounts of botanical expeditions, in particular those of Fernald (1926, 1933), provided useful additional information on the distribution of a few rare speces. Thanks are also due to Dr. T. Ahti, University of Hel- sinki, Dr. W. S. Benninghoff, University of Michigan, Dr. T. C. Brayshaw and Dr. J. S. Rowe, Department of Forestry of Canada, and Dr. W. S. Maass, National Research Coun- cil, for helpful suggestions and valuable criticism on a draft of the manuscript, and to Dr. M. E. Hale, Smithsonian In- stitution for identification of some of the lichens. In this paper an attempt will be made to analyse the southern and northern elements in the flora of Newfound- land. Before doing this, it will be necessary to describe briefly the geography of the island. GEOGRAPHIC SETTING The Island of Newfoundland is located off the east coast of the American continent between 46°30’ and 51°30’ N. Lat. and between 59°30’ and 52°45’ W. Long. It covers an area of a little over 42,000 square miles. It is separated in the southwest from Cape Breton Island by the Cabot Strait, and in the northwest from Labrador by the Strait of Belle Isle, 70 and 11 miles wide at their narrowest points re- spectively (Figure 1). The island consists essentially of a tilted plateau of mod- erate to slight relief. It rises gradually to the west, where elevations of over 2,500 feet can be found. Calcareous soils and limestone outcrops and cliffs pre- vail along the west coast from St. George’s Bay to the north- ernmost tip of the Great Northern Peninsula. Acidic bedrock, and glacial till derived from it, cover most of the remainder of the island. Serpentine mountains occur along the west coast, between St. George’s Bay and Bonne Bay, and also in the northern part of the Great Northern Penin- sula. 1965] Newfoundland Flora — Damman 365 50° 2 GULF OF N ST. LAWRENCE ^ Lu La © PRINCE EDWARD [v] CAPE BRETON j ISLANO 45? e \ gd N L A T A 65° 60° peo ted Figure 1. Location of the Island of Newfoundland in relation to the American continent. In spite of its location, the island does not have a hyper- oceanic climate. Southwestern and western winds prevail throughout the year. Consequently, the climate is greatly influenced by the American continent. This is clearly em- phasized by the occurrence near the coast of many species considered continental or sub-continental in Europe, e.g., Chamaedaphne calyculata, Rhododendron lapponicum, and 366 Rhodora [Vol. 67 many others. Therefore, the climate of the coastal areas should be considered sub-oceanic at most. The cold Labrador Current flows along the shores of New- foundland; this current brings ice floes along the shores in winter and spring, and icebergs during spring and summer. The coastal ice fields greatly retard the arrival of spring (Figure 2). The vegetative period starts from 30 to 50 days later in Newfoundland than in Montreal (Hare, 1952). May 30 Figure 2. Start of vegetative season based on threshold temperature of 43°F. (After Hare 1952) 1965] Newfoundland Flora — Damman 361 Very low summer temperatures prevail along the coast owing to the frequent occurrence of fog and the cooling effect of the sea ; this phenomenon is particularly pronounced in the southern and southeastern parts of the island (Figure SPB In winter, the sea has a moderating effect on the tempera- tures; the winters of Newfoundland are considerably milder than those of the adjacent continent. This is es- Figure 3. Mean annual maximum temperature in °F. The isotherms have been drawn on the basis of sea level temperatures, and thus show the temperature irrespective of elevation. 368 Rhodora [Vol. 67 pecially true for the areas along the ice-free south coast and the Avalon Peninsula which is not usually surrounded by ice until March (Figures 4 and 5). The precipitation is well distributed throughout the year in all parts of the island. It ranges from over 55 inches in the southern and southwestern parts to less than 35 inches in the northern areas. The driest area, with a precipitation of 30 inches, is found in the northwestern tip of the Great Northern Peninsula. f f Figure 4. Mean? monthly-minimum-temperature for January in °F. (After Hare 1952) $n 1965] Newfoundland Flora — Damman 369 Figure 5. Mean monthly minimum temperature for January in ^F. (After Hare 1952) The island is situated within the Boreal forest region (Rowe, 1959). The central and western parts are heavily forested with treeless barrens restricted to the higher eleva- tions. Most of the southern and eastern parts are covered by dwarf shrub heath and bogs (Ahti, 1959). 370 Rhodora [Vol. 67 PLANTS WITH SOUTHERN AFFINITIES This group includes the so-called Coastal Plain element, a group of species restricted to oligotrophic bogs and dry sandy soils, and occurring southward along the Atlantic coast to the pine barrens of New Jersey. Although it is the most talked-about element, it is by no means the only south- ern element in the flora of Newfoundland; temperate-boreal species of both Canadian and Alleghanian range also occur. The southern element is defined here to include those species occurring in Newfoundland but having their main range south and west of the island, and whose continuous range on the Canadian mainland does not reach as far north as southern Labrador. Thus defined, the southern element includes those species which, under the present climatic conditions, can reach Newfoundland only across the Cabot Strait; a point of interest in Fernald’s theories on the origin of the Newfoundland flora, and in the dis- cussion in the final part of this paper. A few species be- longing to the southern element, as defined above, but occurring rather commonly up to the northern tip of the Great Northern Peninsula have been left out of considera- tion in the following discussion. What are the ecological factors controlling the distribu- tion of southern species in Newfoundland? Both climatic and edaphic factors are important in this respect; they will be briefly discussed below before describing the types of distribution to be found on the island. CONTROLLING FACTORS 1. SUMMER TEMPERATURES AND LENGTH OF VEGETATIVE SEASON. Most of the southern plants have the main part of their range in areas with considerably warmer summers and a longer vegetative season than that found in Newfound- land. Therefore, this is the most obvious factor controlling their distribution. Species thus controlled will either show a clear northern boundary in Newfoundland, or they are restricted to some of the warmer valleys and sites. 1965] Newfoundland Flora — Damman 5ygl It is worth mentioning that the areas with the longest and generally warmest vegetative season do not coincide with those with the highest annual maximum temperatures. The former occur in the valleys of the southern part of the west coast, whereas the latter are found in the northern central parts of the island. The annual maximum tempera- tures do not appear to be a relevant factor in determining the distribution of southern plants; at least, no species were found whose distribution was closely related to it. 2. WINTER TEMPERATURES. Most of the southern species reaching Newfoundland have a range extending far into areas with a much more continental climate. Therefore, they are adapted to winters much colder than those of New- foundland, and it is obvious that low winter temperatures will not prevent any of these species from occurring on the island. On the other hand, many species sensitive to low winter temperatures will be able to grow in Newfoundland because of the relatively mild winters of the coastal areas. These species concentrate especially on the southeastern penin- sulas and along the south coast. 3. SPRING TEMPERATURES. One of the typical features of a continental climate is the rapid change from winter to sum- mer, and the absence of a long period with alternating cold and warm spells so frequently encountered in northern oceanic climates. The occurrence of a cold spell following a mild period can be very detrimental to species which start their growing season as soon as the first mild days occur. This factor probably limits the distribution of a large num- ber of continental species. The phenomenon of a spring with alternating cold and mild spells is most common in the typically oceanic parts of the Island, e.g., the south coast and the eastern penin- sulas, but it is also of great importance in central New- foundland and along the northern shore. The west coast is the area most favorable for species sensitive to this factor. 372 Rhodora [Vol. 67 4. SOIL CONDITIONS. Edaphic factors play an important role in determining the distribution of plants. The distribu- tion of calcareous soils causes the greatest regional differ- ences in the flora of Newfoundland, and many southern plants are restricted to western Newfoundland for this rea- son only. Nutrient-rich soils are also more abundant in the western parts, but, although this limits the frequency of occurrence, it will rarely eliminate a species completely from the in- fertile eastern parts. Dry, sandy soils also are warm soils, and a few southern species with low nutrient and moisture requirements can be found exclusively on these sites. A few other southern spe- cies occur only on unstable talus or terrace slopes and other sites with limited competition. 5. BIOTIC FACTORS. Some species are so dependent on the habitat condition created by the plant community in which they grow that the absence of one or more of their dominant associates can be the factor limiting their distribution. This factor is indirectly controlled by the climate. An extreme example of this kind of dependence is given by a number of liverworts, such as Mylia anomala and Odontoschisma sphagni, which occur almost exclusively in clumps of Sphag- num fuscum and S. rubellum. In analogy with this, it is quite possible that the occurrence of many species of north- ern hardwood forests, such as Erythronium americanum, Medeola virginiana, Uvularia "SessiliRora, and Sanguinaria canadensis, are dependent on the occurrence of this type of forest. Sugar maple-yellow birch forests occur locally on rich soils in Cape Breton Island but they are lacking on Newfoundland, and with them many species characteristic of this forest type. TYPES OF DISTRIBUTION It will be obvious, that the distribution of the southern species is rarely limited by any single factor but as a rule x SESSILI FOLIA 1965] Newfoundland Flora — Damman 373 the result of the simultaneous action of several of them. The following distribution types can be recognized: A.COMMON OR FREQUENT THROUGHOUT BUT ABSENT ON THE GREAT NORTHERN PENINSULA AND AT HIGHER ELEVATIONS. This group is clearly limited by the length and warmth of the vegetative season. A large number of species belong to this group only a few of them are mentioned below. Osmunda regalis Aster umbellatus Trillium cernuum Sambucus pubens Dulichium arundinaceum Alnus rugosa Carex intumescens Prunus virginiana Carex debilis Vaccinium macrocarpa. ` Carex gracillima Senecio aureus Carex crinita Solidago rugosa Carex castanea Acer rubrum Corylus cornuta Pogonia ophioglossoides Nemopanthus mucronatus? Arethusa bulbosa Viburnum cassinoides? Rhododendron canadense? Diervilla lonicera Cypripedium acaule Nymphaea odorata? B. RESTRICTED TO THE WEST COAST SOUTH OF DEER LAKE, THE DEEPER INLETS OF THE SOUTH COAST, NOTABLY BAY D'ESPOIR, AND SHELTERED PARTS OF THE SOUTH- ERN AVALON PENINSULA. This group is also clearly limited by the length and warmth of the vegetative season, but its species demand somewhat higher summer temperatures than those of group A. Betula lutea Agrimonia striata Scutellaria lateriflora C. RARE OR LOCALLY OCCURRING SPECIES WHOSE DISTRIBU- TION IS CLEARLY LIMITED BY LENGTH AND WARMTH OF THE VEGETATIVE SEASON. *Species extremely rare on the Great Northern Peninsula, 374 Rhodora [Vol. 67 a. Restricted to Warm, Favorable Locations on the West Coast. (Extreme examples of groups A and B.) Claytonia caroliniana Geranium robertianum Fraxinus nigra (always as a small tree or shrub) Cornus alternifolia b. Limited to a Few Dry, Sandy Areas in Central New- foundland. Most likely these are relic species for it is difficult to understand how they could other- wise have reached their present locations, This applies particularly to Pinus resinosa. Carex houghtonii Pinus resinosa Gaultheria procumbens c. Halophytes Restricted to Southwestern Newfound- land. The water of the Gulf of St. Lawrence is considerably warmer than that of the other coasts. It could well be that this rather than the air tem- perature determines the distribution of these species. Scirpus validus Spartina pectinata Scirpus maritimus? d. Very Scattered, but Clearly Limited by the Length of the Vegetative Season. Typha latifolia — This species is restricted to a relatively warm belt extending from Port-aux-Basques along the west coast to Bonne Bay, and then to the northern part of Cen- tral Newfoundland; it occurs also in the central parts of the Avalon Peninsula. D. RESTRICTED TO WESTERN NEWFOUNDLAND, SOUTH OF BONNE BAY. "New addition to the Newfoundland flora discovered by the author in a salt marsh near Boswarlos, Port au Port Bay, during the field season of 1961. 1965] Newfoundland Flora — Damman 375 This type of distribution appears to be caused by two sets of conditions: a. The Occurrence of Calcareous or Highly Basic Soils. Cypripedium reginae Dryopteris robertiana Scirpus acutus Salix glaucophylloides b. Rather Continental Course of Spring Temperatures. Epigaea repens is the most conspicuous example of this. The species occurs very commonly in western Newfound- land south of Bonne Bay, and even reaches rather high eleva- tions at the east side of the Lewis Hills and in the mountains east of Grand Lake. It is very rare in the forested interior around Red Indian Lake and Victoria Lake. At high eleva- tions the species is restricted to sites well covered with snow in winter and early spring. Consequently, it starts its vegetative period very late, and at a time that severe cold spells are unlikely to occur. An example of a similar protection against early sprouting is seen in Smilacina racemosa*, It occurs in an Alnus crispa thicket on the southwest slope of the Lewis Hills at about 1000 feet elevation. It is not a particularly warm site, but it is usually covered with snow until the middle of June. Moreover, the conditions on the forest floor show an in- teresting resemblance to those of a hardwood forest. E. CONCENTRATED IN SOUTHEASTERN NEWFOUNDLAND, THE SOUTH COAST, AND THE BOGS OF THE CARBONI- FERIOUS SANDSTONE AREA OF ST. GEORGE'S BAY, BUT ALSO PRESENT IN BOGS ON THE WEST COAST MOUNTAINS AND OF THE INTERIOR BARRENS. This group includes the Coastal Plain species. They are moderately oceanic species, apparently sensitive to low win- ter temperatures, adapted to cool summers, and possibly author in 1961. In 1963, Dr. E. Rouleau (oral communication) found this species in an Alnus rugosa swamp along Mollichienic Brook, Western Newfoundland. 316 Rhodora [Vol. 67 pond species of this group occur, rather sparingly, at the higher elevations. In these areas, as in the coastal areas, snow drifting is severe in winter time, and thick snow layers cover the valleys and depressions. The occurrence of these species at higher altitudes could well be dependent on this protective snow cover. The species with a very conspicuous concentration in the oceanic parts of the island, i.e., on the Avalon Peninsula, the Carboniferous sandstone area of St. George's Bay, and to a lesser degree along the south coast, are listed under b. a. Schizaea pusilla Xyris montana Myriophyllum tenellum Bartonia paniculata Nymphoides lacunosa Aronia floribunda Utricularia geminiscapa Juncus militaris Sphagnum molle” Juncus pelocarpus Sphagnum strictum’ Scirpus subterminalis Potamogeton oakesianus In contrast to the other species of this group, both Sphag- num molle and Sphagnum strictum have never been found on the West Coast north of Port-aux-Basques. b. Gaylussacia dumosa Habenaria blephariglottis Calopogon pulchellus Four species with a much less oceanic distribution farther south have a very similar range in Newfoundland as the species of group Eb, viz. Gaylussacia baccata, Carex fol- liculata, Habenaria orbiculata, and Dryopteris novebora- censis. Species which also have an oceanic distribution in Amer- ica, Lycopodium inundatum and Sphagnum pylaesii, go much farther north along the Atlantic coast. The latter, especially, is often peculiarly abundant in bog pools on very cold peaks. "Until recently, Sphagnum molle was known from only two localities north of New Jersey, both on Newfoundland. Dr. W. S. G. Maass of the Atlantic Regional Laboratory, National Research Council, dis- covered the species on a number of localities in recent years, most of them on the Avalon and Burin Peninsulas. 1965] Newfoundland Flora — Damman 377 F. RESTRICTED TO AVALON AND BURIN PENINSULAS. Apparently an extreme form of group Eb. Sphagnum macrophyllum® This type of distribution is also found in many species which do not reach their northern limit in Newfoundland. Most conspicuous among them are some European species such as Nardus stricta, Pedicularis sylvatica, Juncus bulbo- sus, and Sieglingia decumbens. For some of them south- eastern Newfoundland is their only foothold in North America. G. SPECIES KNOWN FROM ONLY ONE OR TWO LOCALITIES. Their occurrence is either due to accident or to historical factors. Obviously, they are not very well adapted to the climatic conditions and the competition relationships exist- ing on the island. Aster acuminatus Smilacina racemosa Mitchella repens Carex lacustris’ West Coast Dryopteris marginalis’ Scirpus maritimus Dennstaedtia punctilobula ` | Oxalis montana PLANTS WITH ARCTIC OR SUB-ARCTIC AFFINITIES The flora of Newfoundland contains a very considerable arctic element. Many of these species reach their southern limit at sea level in Newfoundland, and occur farther south ‘In 1962 Dr. W. S. G. Maass collected Sphagnum macrophyllum, thus far not known north of Nova Scotia, from five localities on the island, four of them on the Avalon Peninsula. 'Collected by Dr. R. Tuomikoski along the Harry's River in 1949. sOccurs on a forested but unstable talus slope east of Barachois Pond. This station was discovered by the author in 1964. 378 Rhodora [Vol. 67 only in alpine habitats, whereas others occur nowhere south of the island. Their distribution in Newfoundland can largely be ex- plained by a combination of climatic and edaphic factors, but lack of competition is an important factor for some spe- cies. In the following the factors thought to be controlling their distribution will be briefly discussed, followed by the description of a few typical distribution types. CONTROLLING FACTORS 1. COMPETITION. It seems strange that more optimal physiological conditions, such as higher temperatures, should eliminate species in more southern latitudes. There- fore, the restriction of arctic species to northern latitudes is often believed to be a result of competition, The northern species, being species with a large temperature amplitude but unable to compete successfully with the more vigorous southern species, are expelled to areas with less severe competition. 2. HIGH SUMMER TEMPERATURES. Dahl (1951) lists a num- ber of species which grow very poorly in botanical gardens at lower altitudes. He made the interesting observation that these species generally grow well in the autumn, and develop fine buds in spring. However, during the summer they be- come yellowish or brownish along the margins of the leaves, and after a few years they die. Dahl considers high summer temperatures detrimental to these species, and found a very good correlation between the isotherms of the annual maxi- mum temperature and the southern limit of these species. This intolerance for high temperatures also explains the occurrence of many arctic-alpine species in closed plant communities along the outer coast line far south of their in- land range. 3. LIGHT REQUIREMENTS. Arctic species are plants of open, unforested areas, and usually they are species with high light requirements, unable to survive under shrub or forest canopy. 1965] Newfoundland Flora — Damman 379 4. SOIL CONDITIONS. This affects the arctic element to the same extent as all other plant species. The distribution of serpentine and limestone has a very pronounced effect on the occurrence of a large group of species. 5. PHOTOPERIODISM. Little information is available on the effect of day length on the distribution of arctic species. However, its effect seems to be negligible with respect to our problem. This factor can at most prevent the occurrence of arctic species on Newfoundland, and it will certainly never cause the occurrence of an extremely long day plant at more southern latitudes. 99 6. ARIDITY OF CERTAIN COASTAL HABITATS. Fernald (1955, p. 123-7), commenting on the occurrence of the arctic ele- ment at sea level in northwestern Newfoundland, remarks that its presence is a result of the aridity and the calcareous nature of the soil rather than the coldness of the climate. The calcareous soils are undoubtedly an important factor govern- ing the distribution of certain arctic plants, but it is hard to believe that the aridity of these sites is more important than the low summer temperatures. The coastal zone is remarkably cold, and the distribution of most species ap- pears to be closely related to it. Those species whose restric- tion to the northwestern section of the Great Northern Peninsula does not appear to be controlled by low summer temperature only are also species of wet sites, such as Carex microglochin, C. capitata, and C. chordorrhiza. 7. CONTINENTALITY OF CLIMATE. The species mentioned above have a preference for continental climates. They may require low winter temperatures, or a rapid change from winter to summer. The warmer summers of continental climates can hardly play a role here. If this were the case, then these species would undoubtedly occur also at low ele- vations farther south along the west coast. TYPES OF DISTRIBUTION A. ON EXPOSED HEADLANDS ALONG THE COAST, EXPOSED PARTS OF THE BARRENS OF EASTERN NEWFOUNDLAND, 380 Rhodora [Vol. 67 AND AT HIGH ELEVATIONS ELESW HERE, BECOMING PRO- GRESSIVELY MORE COMMON TOWARD THE NORTHERN PART OF THE LONG RANGE MOUNTAINS. This type of distribution is shown by species which can find suitable soil conditions in any part of the province, i.e., they are species of acidic soils and those indifferent in this respect. They seem to be limited chiefly by high summer temperatures, but thrive well in both oceanie and more con- tinental parts of the island. Loiseleuria procumbens Diapensia lapponica Empetrum eamesii Silene acaulis’ Lycopodium selago Lycodium annotinum var. pungens Carex bigelovii Betula borealis Juncus trifidus Vaccinium uliginosum Arctostaphylos alpina Cetraria nivalis Sphagnum lindbergii | Alectoria nigricans Cetraria cucullata Alectoria nitidula Alectoria ochroleuca Lobaria scrobiculata (as a ground inhabiting species) B. RESTRICTED TO THE HIGHLANDS OF ST. JOHN, AND SOMETIMES A FEW OTHER HIGH AREAS IN THE WESTERN PART OF THE ISLAND. This group consists of species of acid soils. Their occur- rence is also chiefly controlled by summer temperatures, they require definitely lower temperatures than the species of Group A. Salix herbacea Cassiope hypnoides Sibbaldia procumbens Phyllodoce caerulea Deschampsia atropurpurea Hierochloe alpina C. ALONG THE WEST COAST FROM ST. GEORGE’S BAY TO THE NORTHERNMOST TIP OF THE GREAT NORTHERN PENIN- SULA. These species occur at higher elevations in the south but they come down to sea level in the northern areas. They are all basiphilic species, and their distribution is deter- "Distinctly more common in limestone and serpentine areas. 1965] Newfoundland Flora — Damman 381 mined by the occurrence of limestone or serpentine. Some are restricted to calcareous habitats ( Ca), others occur on both serpentine and calcareous soils (Ca + Mg). Oxytropis foliolosa (Ca) Hedysarum alpinum (Ca) Dryas integrifolia (Ca) Carex rupestris (Ca) Carex terrae-novae (Ca) Saxifraga oppositifolia (Ca) Salix vestita (Ca) Cerastium beeringianum (Ca + Mg) Saxifraga aizoon (Ca) Arenaria dawsonensis (Ca) Arenaria humifusa (Ca + Mg) Armeria labradorica (Ca + Mg) Potentilla nivea (Ca) Kobresia simpliciuscula (Ca) Arenaria rubella (Ca + Mg) Castilleja septentrionalis (Ca + Mg) Salix cordifolia (Ca + Mg) Polygonum viviparum (Ca) Cetraria tilesii (Ca) Solidago multiradiata (Ca + Mg) Rhododendron lapponicum (Ca + Mg) D. RESTRICTED TO THE NORTHERN PART OF THE GREAT NORTHERN PENINSULA. High summer temperatures appear to be the main factor limiting these species to this part of the island. Another factor, here vaguely described as continentality, is also im- portant, at least for the first two species. Carex chordorrhiza Pyrola grandiflora (Ca) Carex capitata Primula egaliksensis (Ca) Salix reticulata (Ca) Habenaria straminea (Ca) Bartsia alpina (Ca) Carex bicolor (Ca) Carex microglochin (Ca) Saxifraga cespitosa (Ca) E. RESTRICTED TO SERPENTINE AREAS. The serpentine areas have a very sparse vegetation cover, and competition is of little importance on these sites. They are bleak, exposed habitats, locally with severe frost churn- ing in fall and winter but the average summer temperatures 382 Rhodora [Vol. 67 in the low lying areas (300-400ft.) are a few degrees higher than those prevailing in the coastal parts of southeastern Newfoundland, Evidently, these species do not require lower summer temperatures than the species of group A. They occur on these sites because of the lack of competition, the basic soils, and their ability to tolerate frost churning. Lychnis alpina Adiantum pedatum var. aleuticum A large number of basiphilic species occur both on the serpentine and limestone soils (see under C). It should be realized that there are several other species limited to the serpentine soils which do not belong to the floral elements discussed in this paper. F. SUB-ARCTIC HABITATS, BUT ALSO ON UNSTABLE SOILS, CLIFFS, AND RIVER BEDS AT LOWER ELEVATIONS. Competition appears to be the main factor controlling the distribution of these species, although some of them require basic soils. Trisetum spicatum Carex atrofusca Saxifraga aizoides Anemone parviflora Luzula spicata Carex scirpoidea Epilobium latifolium (known only from river gravels in Newfoundland.) G. RATHER FREQUENT THROUGHOUT, ALTHOUGH DISTINCTLY MORE COMMON AT HIGHER ELEVATIONS. A very large number of species belongs to this category. The following are a few examples. Thalictrum alpinum Scirpus hudsonianus Rubus chamaemorus Carex capillaris Selaginella selaginoides Rubus arcticus Rubus acaulis Tofieldia pusilla A number of species show distribution patterns inter- mediate between the groups discussed above. Intermediate ranges between groups A and B are most common ; to this category belong: Betula minor, Betula glandulosa, Salix uva- ursi, and Carex rariflora. All of them show a very marked 1965] Newfoundland Flora — Damman 383 increase in abundance to the north, and they are rare or absent east of the Long Range Mountains. GLACIAL AND POST-GLACIAL HISTORY AS A FACTOR IN EXPLAINING THE DISTRIBUTION PATTERNS Most of the distribution patterns of northern and southern elements on the Island of Newfoundland can be explained adequately by the existing climatic and edaphic conditions. The ecological factors controlling this species distribution were analyzed in the previous section; in the following, the role of the historical factor will be discussed in more detail. The importance of historical factors is often overem- phasized in explaining the distribution of species (Bócher, 1943, 1951). Undoubtedly, the Wisconsin glaciation and the post-glacial climatic changes have had a pronounced influence on the composition of the present flora of New- foundland, as they have had on the floras of all glaciated regions. There are probably a few species on the island today whose presence is the result of the post-glacial climatic optimum ; there are others which may have taken advantage of the low sea level during late glacial times, and there are certainly many species which are absent due to the glacial epoch. However, it is well to remember that there is a large speculative element in all theories based on history. Often these theories are very attractive, but equally often it is not necessary to resort to far-fetched hypotheses to explain the present distribution of species, provided one pays proper attention to their ecology. It is almost impossible to write about the origin of the Newfoundland flora, without mentioning the admirable pio- neer work of Fernald. An extensive literature has built it- self around his controversial theories, and the last word has certainly not been said about it. I will restrict myself to a very brief description of Fernald's ideas as far as they con- cern the subject of the paper, followed by my own views. A detailed discussion of this subject can be found in Marie- Victorin (1938), Raup (1941), Deevey (1949), Dahl 384 Rhodora [Vol. 67 (1946), Dansereau (1950), Wynne-Edwards (1937-1939), Rousseau (1949). A discussion on the distribution and the origin of the so-called *Cordilleran" element in the flora of Newfoundland has been omitted here on purpose since the problem is of an entirely different nature. A point of interest in Fernald's theories on the origin of the Newfoundland flora (1911, 1918a, 1918b, 1925, 1926, 1933) is his explanation of the presence of the Coastal Plain element. In his early writings, Fernald (1918) believed that the Coastal Plain element had migrated northward toward the end of the glacial period over a land bridge, approxi- mately following the continental shelf. This land bridge was very sandy, and only species with low nutrient require- ments could use it. Thus it was well suited for the species of acid soils which make up the Coastal Plain element but not for other southern species. Later Fernald (1933, p. 95-107) revised this theory, mainly since his nunatak theory (Fernald, 1925), as an ex- planation of the occurrence of the Cordilleran element in eastern Canada, had received so much approval of both geol- ogists and botanists, whereas his post-glacial land bridge theory had few supporters. He postulated that the Coastal Plain species had reached Newfoundland during the Ter- tiary period over the then emerged continental shelf. Sub- sequently, they survived the glacial period on nunataks, unglaciated refugia occurring as islands in the continental ice sheet. At present it is obvious that all of Newfoundland, includ- ing its highest peaks, has been covered with ice at some time or another during the glacial epoch (MacClintock and Twenhofel, 1940). The other nunatak areas in northeastern North America are equally unjustified (Flint et al., 1942; Odell, 1938; Abbe, 1936; Deevey, 1949). In addition to this, many so-called nunatak indicators have been found in the "gap" of the Canadian Shield during recent years (Rous- seau, 1948, 1949; Scoggan, 1950; Raymond, 1950). AI- though the existence of genuine nunataks on Newfoundland is extremely unlikely, it is quite well possible that certain 1965] Newfoundland Flora — Damman 385 floral elements survived the glaciation in coastal habitats, especially along the Atlantic shore. However, it is not necessary to resort to such speculations to explain the distribution of these species. The Coastal Plain element does not appear to be a relic flora, which is restricted to the bleak eastern and southern parts of the island owing to the occurrence of acidic soils (Fernald, 1911, 1918). In my opinion, it occurs on Newfoundland because it is made up of oceanic and sub-oceanic species which are unable to tolerate the severe winters of most of the adjacent continent. Severe snow drifting in the barren areas may even permit these species to occur farther inland than would be otherwise possible. A few species with an unusual gap in their distribution, e.g. Sphagnum molle and S. macrophyl- lum, could well have covered more of the island during the post-glacial climatic optimum. A second point of interest in Fernald’s theories is his ex- planation of the absence of many species of nutrient-rich soils occurring commonly on the adjacent Canadian main- land. He attributed their absence to the inability of these species to cross the Cabot Strait between Cape Breton Island and Newfoundland. He reasoned further, that owing to their higher demands on soil fertility these species could not make use of the land bridge as the Coastal Plain spe- cies did. "Thus", he states, “typical Canadian plants, unless their range extends to the north side of the Strait of Belle Isle, are absent from the island" (Fernald, 1911). Clearly this point of view cannot be defended any longer. A large group of species, notably those with high nutrient requirement in groups A, B, C, and D do not occur on the Northern Peninsula and consequently they must have reached Newfoundland from the south (see also p. 32). Thus the Cabot Strait cannot be considered a very effective barrier. "This Strait separates Newfoundland from southern Labrador; it is only eleven miles at its narrowest point and thus it can be much easier bridged by diaspores than the seventy mile wide Cabot Strait. 386 Rhodora [Vol. 67 Those species present on the adjacent mainland (Roland, 1942; Dore and Roland, 1941) but absent on the Island of Newfoundland are all species of tolerant hardwood forests, e.g., Acer saccharum, Fagus grandifolia, Epifagus virgini- ana, Quercus borealis, Ulmus americana and Sanguinaria canadensis. Even in the climatically favorable southwestern part of Newfoundland the vegetative season starts almost two weeks later than in Cape Breton Island. Therefore, it seems more reasonable to attribute the absence of these species to the shortness of the vegetative season rather than to their inability to cross the Cabot Strait. On Newfoundland the drier alder thickets are the habitats most closely resembling the tolerant hardwood forests. It is interesting to note that the few tolerant hardwood spe- cies which do reach Newfoundland all occur in alder thickets, e.g., Claytonia caroliniana and Smilacina racemosa. There is one group of species with southern affinities whose distribution is rather enigmatic. It includes species as Pinus resinosa, Carex houghtonii and Gaultheria pro- cumbens (Group Cb) but also Chimaphila umbellata. All of them occur on a number of widely separated or isolated stations where they can be very common. It is difficult to understand the present distribution of these species without assuming that they once covered a larger part of the island. This could have been the case during a warm post-glacial period. Evidence for such a climatic optimum has been col- lected for other parts of northeastern America (Deevey, 1949). Livingstone and Livingstone (1958) and Schofield (1960) have shown that a distinctly warmer period has occurred on Cape Breton Island. During this period. Quer- cus and several other hardwoods were much more common than today, and even Carya pollen has been found in these “Recent addition to the flora of Newfoundland. Collected by R. S. van Nostrand at New Bay Lake (central Newfoundland), and by the author near Flat Bay Brook (West Coast), both in 1958. There is an unverified record of its occurrence near Hall’s Bay. 1965] Newfoundland Flora — Damman 387 deposits. This period was followed by a slightly cooler and more humid period in which Tsuga was a prominent com- ponent of the forest, and from then on the climate gradually deteriorated. Although no evidence for a climatic optimum is available for Newfoundland, it seems reasonable to assume that this warmer climate also affected the island, thus creating con- ditions for a more widespread occurrence of species such as Pinus resinosa. In the absence of pollen analytical studies or fossil records from Newfoundland we can only speculate on this subject. It can only be said with confidence that the present climatic conditions are unfavorable for these species for otherwise they would have spread from their present locations. The latter applies also to the southern species which are known from a single locality only (Group G.) Oxalis montana, a species restricted to LaPoile Bay, an isolated bay on the south coast may be an exception in this respect. It has to cross a large area of bleak barren land to reach the forests of the west coast, and it is conceivable, though unlikely, that this is the reason for its absence from the rest of the island. The occurrence of a post-glacial climatic optimum on Newfoundland may throw some doubt on the previous state- ment that some of southern species in groups A, B, C, and D must have reached the island across the Cabot Strait. Could not their northern limit have occurred in Labrador during this warm period, so that they could have crossed the Strait of Belle Isle and spread into Newfoundland from the north? Wenner's (1947) pollen analytical studies in Labrador and northern-most Newfoundland seem to indicate that this is very unlikely. He found no evidence of important climatic changes after the area became covered with a forest vegeta- tion. Presumably, the hypsothermal period occurred before the climate in southern Labrador had ameliorated sufficient- ly to permit the growth of a forest vegetation. The widespread occurrence of a great number of arctic and sub-arctic species occurring on the 4sland can be satis- factorily explained by the climatic conditions, as has been 388 Rhodora [Vol. 67 done in the description of the distribution types. Their common occurrence at sea level in coastal areas is due to the very low summer temperatures prevailing in these re- gions. Temperature measurements with sucrose ampoules (Pallmann et al, 1940) carried out by the author in 1961, have shown that the average temperatures during the veg- etative season in coastal barrens are equal to those on the highest peaks of the Table Mountain at Bonne Bay, the Lewis Hills and the Gaff Topsails. A peculiar mingling of arctic and southern species can be seen in the coastal habitats, as already remarked by Fernald (1911). However, in the author’s opinion this is a result of the present day climatic conditions rather than the presence of a relic Coastal Plain element in an essen- tially northern vegetation. The arctic species occur there because of the low summer temperatures, whereas the Coastal Plain species are present because of the oceanic cli- mate, and in spite of the low summer temperatures to which they are adapted. SUMMARY The flora of Newfoundland contains many species with northern and southern affinities. The ecological factors con- troling the distribution of these floral elements are dis- cussed in some detail followed by a description of the distribution patterns within Newfoundland. Most of the species distributions can be satisfactorily explained by the present climatic and edaphic conditions. The Coastal Plain element, should not be considered as a relic flora restricted to southern Newfoundland because of soil conditions but rather as a flora consisting of oceanic species unable to tolerate extreme winter temperatures. The absence of some southern species, notably those of northern hardwood forests should be attributed to unfavor- able climatic conditions and the absence of the hardwood forest floor habitat rather than to isolation. The number of species which did reach Newfoundland across the Cabot 1965] Newfoundland Flora — Damman 389 Strait is so great that it can hardly be considered an effective barrier for most species. The peculiar distribution of Pinus resinosa, Chimaphila umbellata, Gaultheria procumbens and Carex houghtonii can only be explained by assuming that these species were widespread during the post-glacial climatic optimum. The subsequent deterioration in climate restricted these species to a few isolated areas with relatively warm soils. SOMMAIRE La flore de Terre-Neuve comprend un grand nombre de plantes du sud et de plantes arctiques et arctiques-alpines. Les facteurs écologiques que régissent la répartition des éléments de la flore de l'ile font l'objet d'une étude détaillée que l'auteur fait suivre d'une description du mode naturel de répartition des diverses plantes dans lle de Terre-Neuve. Pour la plupart des plantes, la répartition des espéces peut étre attribuée ans crainte d'erreur aux conditions locales actuelles du climate et du sol. La flore de la plaine cótiére ne doit pas étre regardée comme une flore reliquale restreinte au sud-est de Terre-Neuve par suite des conditions particu- lières du sol, mais plutôt comme une flore composée d'espéces océaniques qui ne peuvent pas survivre aux tem- pératures hivernales extrémes. L’absence de certaines espéces du sud, l'absence notam- ment de certaines espèces des foréts de feuillus du nord, est attribuable aux conditions climatiques défavorables et à absence du milieu caractéristique des foréts de feuillus, plutót qu'a lisolement. Le nombre d'espèces qui se sont établies dans l'ile de Terre-Neuve aprés avoir traversé le détroit de Cabot est tellement éléve que l'on croit difficile- ment que le détroit puisse constituer un sérieux obstacle à la propagation de la plupart des espéces. La répartition bizarre des especes Pinus resinosa, Chima- phila umbellata, Gaultheria procumbens et Carex houghtonii ne peut étre attribuée que à la répartition de ces especes dans toute l'ile durant une période post-glaciaire, alors que 390 Rhodora [Vol. 67 les conditions climatiques étaient les plus favorables à leur reproduction. Le climat beaucoup plus rude qui a suivi cette période a confiné la survivance de ces espéces dans quelques poches isolées oü le sol est rélativement moins froid. RESEARCH OFFICER, FOREST RESEARCH BRANCH, DEPARTMENT OF FORESTRY, CANADA. ST. JOHNS, NEWFOUNDLAND LITERATURE CITED ABBE, E. C. 1936. Botanical results of the Grenfell-Forbes Northern Labrador Expedition, 1931. Rhodora 38: 102-164. AnTI, T. 1959. Studies on caribou lichen stands of Newfoundland. Ann. Botan. Soc. Zoo. Botan. Fennicae *Vanamo" 30: (4): 1-44 ARSÉNE, L. 1927. Contribution to the flora of the islands of St. Pierre et Miquelon. Rhodora 29: 117-133, 144-158, 173-191, 204-220. BocHER, T. W. 1943. Nordische Verbreitungstypen. Svensk Bot. Tidskr. 37: (4): 352-370. . 1951. Distributions of plants in the cireum-polar area in relation to ecological and historical factors. J. Ecology 39: (2): 376-395. Dant, E. 1946. On different types of unglaciated areas during the ice ages. New Phytologist 45: 225-242. - 1951. On the relations between summer temperature and the distribution of alpine vascular plants in the lowlands of Fen- noscandia. Oikos 3: (1): 22-52. DANSEREAU, P. 1950. Flora and vegetation on the Gaspé Peninsula. Wild Flower, April 1950: 26-39. DEEVEY, E. S. 1949, Biogeography of the Pleistocene. Bull. Geol. Soc. Am. 60: 1315-1416. DoRE, W. G. and A. E. ROLAND. 1941. The grasses of Nova Scotia. Nova Scotian Inst. Sci. Proc. 20: 177-288. FERNALD, M. L. 1911. A botantical expedition to Newfoundland and Southern Labrador. Rhodora 13: 109-162. 1918a. The geographic affinities of the vascular floras of New England, the Maritime Provinces and Newfound- land. Am. J. Botany 5: 219-236. ————. 1918b. The contrast in the floras of Eastern and Western Newfoundland. Am. J. Botany 5: 237-247. — ——— — ————. 1924. Isolation and endemism in Northeastern Amer- ica and their relation to the age and area hypothesis. Am. J. Botany 11: 558-572. 1965] Newfoundland Flora — Damman 391 . 1925. Persistence of plants in unglaciated regions of boreal America. Mem. Amer. Acad. Sci. N. S. 15: 237-342. . 1926. Two summers of botanizing in Newfoundland. Rhodora 28: 49-63, 74-87, 89-111, 115-129, 145-155, 161-178, 181-204, 210-225, 234-241. . 1933. Recent discoveries in the Newfoundland flora. Rhodora 35: 1-16, 47-63, 80-107, 120-140, 161-185, 208-2283, 231- 247, 265-283, 298-315, 327-346, 364-384, 395-403. FLINT, R. F., M. DEMOREST, and A. L. WASHBURN. 1942. Glaciation of Shickshock Mountains, Gaspé Peninsula. Bull. Geol. Soc. Am. 53: 1211-1230. Hare, F. K. 1952. The climate of the Island of Newfoundland. Can. Dep. Mines Tech. Surv., Geogr. Bull. 2: 36-88. LIVINGSTONE, D. A. and B. G. R. LIVINGSTONE. 1958. Late glacial and post-glacial vegetation from Gillis Lake in Richmond County, Cape Breton Island, Nova Scotia. Am. J. Sci. 256: 341-359. MacCuintock, P. and W. H. TWENHOFEL. 1940. Wisconsin glaciation in Newfoundland. Bull. Geol. Soc. Am. 51: 1729-1756. MARIE-VICTORIN, FRERE. 1938. Phytogeographical problems of East- ern Canada. Am. Midland Naturalist 19: (3): 489-558. ODELL, N. E. 1938. The great ice age and its effects. In Forbes, A. Northernmost Labrador mapped from the air. Am. Geol. Soc. Spec. Publ. 22: 204-215. PALLMAN, H., E. EICHENBERGER, und A. HASLER. 1940, Eine neue Methode der Temperaturmessung bei ókologischen und boden- kundlichen Untersuchungen. Ber. Schweiz. Botan, Ges. 50: 337- 362. Raup, H. M. 1941. Botanical problems of Boreal America. Botan. Rev. 7: 147-248. RAYMOND, M. 1950. Esquisse phytogeographique du Québec. Mem. Jard. Bot. Montreal 5: 1-147. RoLAND, A. E. 1942. The flora of Nova Scotia. Nova Scotian Inst. Sci. Proc. 21: 95-642. RovLEAU, E. 1956. A check-list of the vascular plants of the Province of Newfoundland. Contr. Inst. Bot. Univ. Montreal 69: 41-106. RovssEAU, J. 1948. The vegetation and life zones of George River, Eastern Ungava, and the welfare of the natives. Arctic 1: (2): 93-106. RECEN . 1949. The value of botany as indicator of unglaciated areas. Pacific Science Congr. New Zealand. Feb. 1949. RowE, J. S. 1959. Forest regions of Canada. Can. Dep. Northern Affairs Nat. Resources, Forestry Branch Bull. 123: 71 p. SCHOFIELD, W. B. and H. ROBINSON. 1960. Late-glacial and post- glacial plant macrofossils from Gillis Lake, Richmond County, Nova Scotia, Canada. Am. J. Sci. 258: 518-522. 392 Rhodora [Vol. 67 SCOGGAN, H. J. 1950. The flora of Bic and the Gaspé Peninsula, Québec. Nat. Museum Canada, Bull. 115: 399 p. WENNER, CARL-GOSTA. 1947. Pollen diagrams from Labrador. Geogra- fiska Annaler 29: (3-4): 137-373. WYNNE-Epwarps, V. C. 1937. Isolated arctic-alpine floras in eastern North America. Roy. Soc. Can. Trans. 31, Sec. V, ser. 3: 1-26. 1939. Some factors in the isolation of rare alpine plants. Roy. Soc. Can. Trans. 33, Sec, V, ser. 3: 35-42. GREAT NORTHERN Highlands PENINSULA of St.John GULF OF ST. LAWRENCE Port au Port Bay/ en's BONAVISTA PENINSULA PORT AUX BASQUES [4 ° Poite 8g 55° 60° Map of Newfoundland showing geographical features men- Figure 6. tioned in text. THE MOUNTAIN ELEMENT IN THE FLORA OF THE PENINSULA OF VIRGINIA* A. M. HARVILL, JR. The Peninsula of Virginia is a narrow neck of land about 60 miles long lying southeast of Richmond. It is bounded by Hampton Roads and the Chesapeake Bay on the east, by the York River estuary on the northeast, and by the James River on the southwest. Although some of the earliest floris- tic work in this country was done on The Peninsula of Vir- ginia, little was accomplished after the colonial period until Professor and Mrs. E. J. Grimes made extensive collections in 1921-22. Professor M. L. Fernald and his colleagues did some collecting in the area some years later, but it was largely by-passed by them for the richer possibilities to the south of the James River. The writer was fortunate to have the opportunity to study the distribution of plants on The Peninsula while at the College of William and Mary in Wil- liamsburg in the summer of 1964. A very intriguing phytogeographic element in the flora of the region is composed of species which occur on the coastal plain and are rare or unknown on the piedmont of Virginia, but reoccur in the mountains of the Virginia-Carolina area. This element has a long history in the literature and is discussed in some detail by Pennell (1935), Braun (1937), Fernald (1937), and by other botanists. Recent botanical activity in Virginia and the Carolinas en- able us to gain a clearer picture of the phytogeographic rela- tions and history of the coastal plain - mountain element and some eighteen species with this distribution pattern are discussed in this paper. These species are placed in 4 groups, (1) wide-ranging, (2) chiefly of northern distribution, (3) chiefly of southern distribution, or (4) endemic to the Vir- ginia-Carolina area. Within these groups the species are *The writer is indebted to Professor A. B. Massey for information on some species in the Virginia Polytechnic Institute herbarium, and to Dr. C. F. Lane of Longwood College for reading the manuscript. 393 394 Rhodora [Vol. 67 arranged according to their distribution in the Carolinas (Radford, Ahles and Bell, 1964) because the distribution of the flora of the Carolinas appears to offer some possibilities for an explanation of some of the disjunct distributional pat- terns in the Virginia flora. DISJUNCT SPECIES OF THE PENINSULA OF VIRGINIA I. Wide-ranging species north and south. Species of the piedmont and mountains in the Carolinas: Actaea pachypoda Ell; Agrimonia pubescens Wallr.; Campanula americana L.; Dirca palustris L.; and Sanicula marilandica L. Species chiefly of the coastal plain and mountains in the Caro- linas: Habenaria ciliaris (L.) R. Br. II. Species generally of northern distribution. Species of the piedmont and mountains in the Carolinas: Jsotria medeoloides (Pursh) Raf. and Lathyrus venosus Muhl. Species of the coastal plain, piedmont and mountains in the Carolinas: Comptonia peregrina (L.) Coulter. Species chiefly in the mountains in the Carolinas: Aralia race- mosa L. III. Species generally of southern distribution. Species of the piedmont and mountains in the Carolinas: Mono- tropsis odorata Ell. Species of the coastal plain, piedmont and mountains in the Caro- linas: Callicarpa americana L.; Galax aphylla L.; Helenium brevi- folium (Nutt.) Gray (including H. curtissii Gray) and Magnolia tripetala L. Species chiefly in the mountains in the Carolinas: Parnassia asarifolia Vent. and Stewartia ovata (Cav.) Weath. IV. Endemic to the Virginia-Carolina area: Chelone cuthbertti Small. Apparently bicentric, the known occurrence of the Chelone is on the coastal plain of Virginia and in the mountains of North Caro- lina. Other species of The Peninsula belong to the coastal plain- mountain element but their distribution is as yet not so well documented. Also, numerous other species found south of the James River on the coastal plain have disjunct colonies in the mountains but are not included in this study. Five of the eighteen species mapped in this paper are known from the mountains of the Carolinas, but not from those of Virginia; they are Helenium brevifolium, Isotria medeoloides, Monotropsis odorata, Stewartia ovata and Chelone cuthbertit. 1965] Peninsula of Virginia — Harvill 395 Figs. 1-4. DISTRIBUTION IN VIRGINIA AND THE CAROLINAS OF SPECIES WHICH ARE DISJUNCT ON THE PENINSULA OF VIRGINIA. The Peninsula is solid black. Shaded areas represent the occurrence of all species mapped in the figure. The coastal plain is east of the dotted line; the mountains are west of the dashed line; between them is the pied- mont. Fig. 1. e — Magnolia tripetala; X = Galax aphylla; and Comptonia peregrina. Fig. 2. e — Sanicula marilandica; X — Cam- panula americana; and Actaea pachypoda. Fig. 3. e — Dirca palus- tris; and X = Agrimonia pubescens. Fig. 4. € = Lathyrus venosus; and X = Habenaria ciliaris. The species Galax aphylla occurs very locally in the south- ern piedmont of Virginia with stations in Campbell and Pittsylvania counties. The type of distribution shown by Galax is characteristic of several species occurring on The Peninsula; i.e., widespread on the coastal plain and in the mountains but with very local colonies on the piedmont of 396 Rhodora [Vol. 67 Figs. 5-9. Explanation under Figs. 1-4. Fig. 5. X — Aralia race- mosa; and Parnassia asarifolia. Fig. 6. X — Chelone cuthbertii; and Stewartia ovata. Fig. T. 9 = Isotria medeoloides; and X = Helenium brevifolium. Fig. 8. € — Callicarpa americana. Fig. 9. € — Mono- tropsis odorata. Virginia. Baldwin (1941) shows that the coastal plain popu- lations in Virginia are tetraploid and those of the mountains are diploid. It would be interesting to know the chromosome numbers of the very local piedmont populations. The chro- mosome complements of isolated populations of the other species with disjunct distribution would also be of consider- able interest. Significant genetical differences can be ex- pected in plants which are isolated in small colonies. 1965] Peninsula of Virginia — Harvill 397 Mapping the eighteen species discussed in this paper sug- gested some conclusions on the reasons for the disjunct distribution of the species and may provide some clues to the history of floristic movements in the area which is now Virginia. The coastal plain of Virginia is relatively rich in species, the mountains somewhat poorer, and the piedmont is the poorest. Moreover, the piedmont of Virginia is poorer floristically than the piedmont areas to the south of it. Many species generally distributed in the Carolinas become more restricted in distribution in Virgnia. Of the three physio- graphic provinces dealt with here, this northward restriction or limitation in distribution is most marked in the piedmont. The piedmont of Virginia appears as an island which is poor in numbers of species within the richly diversified floras surrounding it. Many of the Virginia species with a disjunct distribution have persisted in bogs and in soils influenced by underlying Miocene shells. It would seem from the available data that the soil conditions, topography, and climatic conditions of the coastal plain and the mountains of Virginia provided refugia where many plants persisted during the climatic stresses of the Pleistocene, such refugia being rare or absent in the piedmont. Isolation could be expected to decrease the number of biotypes in the colonies (Hultén, 1937) and result in populations with little variability and a lack of aggres- siveness. Such populations apparently have narrow edaphic tolerances and are specialized for the soil conditions in which they have persisted and cannot compete successfully with other species in areas of differing edaphic values. REFERENCES BALDWIN, J. T., JR. 1941. Galax: The genus and its chromosomes. Jour. Hered. 32: 249-254. BRAUN, E. L. 1937. Some relationships of the flora of the Cumberland Plateau and the Cumberland Mountains in Kentucky. Rhodora 39: 193-208. DEEVEY, E. S., JR. 1949. Biogeography of the Pleistocene. Bull. Geol. Soc. Am. 60: 1315-1416. 398 Rhodora [Vol. 67 ERLANSON, EILEEN W. 1924. Flora of the Peninsula of Virginia. Papers Mich. Acad. Sci., Arts and Letters 4: 115-182. FERNALD, M. L. 1937. Local plants of the inner Coastal Plain of southeastern Virginia. Part III, Phytogeographic considerations. Rhodora 39: 465-491. HARVILL, A. M., JR. 1965. Plants from The Peninsula of Virginia. Jour. Va. Acad. Sci. 15 (in press). HULTEN, Eric. 1937. Outline of the History of Arctic and Boreal Biota During the Quaternary Period, Stockholm, 168 pp. MassEY, A. B. 1961. Virginia Flora. Blacksburg, Va., 258 pp. PENNELL, F. W. 1935. The Scrophulariaceae of Eastern Temperate North America. Philadelphia, 650 pp. RADFORD, A. E., H. E. AHLES and C. R. BELL. 1964. Guide to the Vascular Flora of the Carolinas. Chapel Hill, 383 pp. LONGWOOD COLLEGE FARMVILLE, VIRGINIA DISCOVERY OF THE MASSACHUSETTS FERN IN WISCONSIN THOMAS G. HARTLEY An interesting discovery made in a recent survey of the flora of the “Driftless Area"* was the Massachusetts fern, Thelypteris simulata (Davenp.) Nieuwl. An abundance of this species was found growing in low sandy woods in the old bed of Glacial Lake Wisconsin in Jackson County, Wis- consin. Collections were made in 1958 and 1960 at two localities about eight miles apart: near Rudd's Hills, Sec- tion 35, Millston Twp. (Hartley 4029), and north of Wild- cat Mound, Section 22, Millston Twp. (Hartley 4819 and 9600). Duplicates of these collections have been distributed to the following herbaria: State University of Iowa, Uni- versity of Wisconsin, Gray Herbarium, United States Na- tional Herbarium, Rancho Santa Ana Botanic Garden, University of Minnesota, University of Michigan, Univer- sity of Illinois, Wisconsin State College at Eau Claire, and Wisconsin State College at LaCrosse. My collections appear to be the first authentic records of this species west of West Virginia. There are reports of it from the dune country of northwestern Indiana by Pepoon (1927) and Peattie (1930), but Fassett (1933) has pointed out that the Umbach collection upon which these were based was misidentified marsh fern, Thelypteris palustris. The accompanying map shows the distribution of Thelyp- teris simulata as represented by the Wisconsin collections and collections at the Gray Herbarium, New England Bo- tanical Club Herbarium, United States National Herbarium and the Wiegand Herbarium of Cornell University. The “Driftless Area” of the Upper Midwest is completely surrounded by glaciated territory and apparently was not covered by continental glaciation during the Pleistocene epoch. Within the belts covered by continental ice sheets of *A dissertation for the degree of Ph. D. at the State University of Iowa, Iowa City, was based on this survey. 599 400 Rhodora [Vol. 67 Massachusetts fern, Thelypteris simulata (Davenp.) Nieuwl., in Jackson County, Wisconsin. (Hartley 9600.) Plate 1317 northeastern North America and northwestern Europe there is no similar region of substantial size which was left bare of glacial ice. As delimited in this survey, the “Drift- less Area” covers approximately 15,000 square miles of southwestern Wisconsin and adjacent Minnesota, Iowa and Illinois. The inclusion of northeastern Iowa and southeast- ern Minnesota requires figurative usage of the name since older ice sheets, generally believed to have been Kansan and Nebraskan, are thought to have covered these areas. The old bed of Glacial Lake Wisconsin covers approxi- mately 1,825 square miles in the northeastern section of the 1965] Massachusetts Fern — Hartley 401 0 > b oe ls | x OMS Distribution of the Massachusetts fern, Thelypetris simulata (Davenp.) Nieuwl. The broken line surrounding the Wisconsin sta- tions indicates the limits of the “Driftless Area." 402 Rhodora [Vol. 67 “Driftless Area." The lake is believed to have been formed when glacial ice of the Cary substage of the Wisconsin stage of the Pleistocene dammed up the Wisconsin River in the region of the Baraboo Range. Radio-carbon dates from material in various Cary deposits in the North Central States (Flint and Deevey, 1951 and Flint and Rubin, 1955) indicate that this occurred about 15 thousand years ago. Pollen studies from bogs in the region (Hansen, 1937) show that the lake had drained and was supporting vegeta- tion before the advance of the Valders ice into northeastern Wisconsin, an event that is dated at about 11 thousand years. Botanically this old lake bed is extremely interest- ing. There are vast areas of black spruce bogs with sur- rounding meadows of ericads and open sphagnum; cold, soft-water streams and flowages; dry, sandy jack pine- scrub oak barrens; and open sandy plains, prairies and blowouts. Woodlands of the type occupied by Thelypteris simulata are frequent in eastern Jackson County. The dominant trees of these woodlands are Acer rubrum, Pinus strobus, Quercus alba and Betula papyrifera, with an under- story of Nemopanthus mucronata, Ilex verticillata, Hama- melis virginiana and Pyrus americana. The woodland floor is often sphagnous and otherwise characterized by such species as Lycopodium clavatum, Lycopodium complanatum, Dryopteris cristata, Osmunda cinnamomea, Osmunda re- galis, Carex brunnescens, Carex debilis, Carex emmonsii, Carex folliculata, Carex intumescens, Carex trisperma, Cin- na latifolia, Clintonia borealis, Cypripedium acaule, Panax trifolius, Cornus canadensis, Trientalis borealis, Coptis groenlandica, Rubus hispidus and Viola incognita. The pres- ence of white oak is interesting and perhaps indicates a less highly acid environment than is found in most poorly- drained areas of this region. Morphologically, Thelypteris simulata is more or less intermediate between Thelypteris palustris and the New York fern, Thelypteris noveboracensis. W. H. Wagner, Jr. (1963), nas suggested that it may have originated as an allopolyploid, pointing out that T. palustris has n=35 and € 1965] Massachusetts Fern — Hartley 403 T. noveboracensis has n—2' and that in material of T. simulata from Maryland he found that »—64. Dr. Wagner reasons that the two extra chromosomes (n should = 62 in the allopolyploid) may indicate a considerable lapse of time since the original hybridization during which two aneuploid changes might have taken place. He also studied the Wis- consin material (Hartley 9600) and noted that the chromo- some number “.... could not be determined exactly, but the number is very close to or the same" as that of the Mary- land material. This information provides us with the attractive possi- bilitv of independent origin as explanation for the disjunct distribution of this species. Though only one of the parents, Thelypteris palustris, is known to occur in Wisconsin, there is the possibility that they were both there at some time during or since the Pleistocene. The extant range of T. noveboracensis extends to counties bordering the eastern and southern shores of Lake Michigan in Michigan, Indiana and Illinois and these areas are similar, floristically, to the old bed of Glacial Lake Wisconsin. Another possible explanation for this distribution is that the Wisconsin population is a relict of a more extensive late or post-Pleistocene distribution. Considerable fluc- tuation in size of the Great Lakes occurred during late Wis- consin times and this apparently provided, at times, more or less continuous habitats along which many species could migrate inland. When subsequent changes destroyed inter- vening habitats, disjunct patterns of distribution resulted. This explanation is particularly applicable to a number of Atlantic Coastal Plain species of open sandy habitats in the old bed of Glacial Lake Wisconsin and may apply to Thelyp- teris simulata as well. Long distance dispersal must, of course, be considered as a third possibility, though it seems less likely since the pre- vailine winds are from the west rather than the east and woodlands of the type occupied by this species are more or less “closed” with regard to competition. 404 Rhodora [Vol. 67 The curators of the Gray Herbarium, New England Bo- tanical Club Herbarium, United States National Herbarium and the Wiegand Herbarium of Cornell University are thanked for the loan of herbarium material, information and permission to examine specimens. Thanks are also ex- tended to Dr. Rolla M. Tryon, Jr., for his assistance in the identification of the Wisconsin material of this species and to the National Science Foundation for financial aid. ARNOLD ARBORETUM, HARVARD UNIVERSITY LITERATURE CITED FaAssETT, N. C. 1933. Notes from the Herbarium of the University of Wisconsin IX, Rhodora 35: 199-203. FLiNT, R. F. and E. S, DEEVEY 1951. Radiocarbon dating of late- Pleistocene events. Amer. Jour. Sci. 249: 257-300. FLINT, R. F. and M. RUBIN 1955. Radioactive carbon dates of pre- Mankato events in eastern and central N. America. Science 121: 649-658. HANSEN, H. P. 1937, Pollen analysis of two Wisconsin bogs of different ages. Ecology 18: 136-148. PEATTIE, D. C. 1930. Flora of the Indiana Dunes. Field Museum of Natural History, Chicago. 432 pp. PEPOON, H. S. 1927. An annotated flora of the Chicago Area. Chicago. 554 pp. WAGNER, W. H. JR. 1963. A biosystematic survey of United States ferns — preliminary abstract. Amer. Fern Jour. 53: 1-17. A THIRD SUBSPECIES IN THE ARTEMISIA TRIDENTATA COMPLEX* ALAN A. BEETLE AND ALVIN YOUNG A recent monograph of the Section Tridentatae of Artemi- sia recognized two subspecies of Artemisia tridentata (Beetle, 1960). Need for the naming of a third subspecies in the A. tridentata complex is now recognized. This sub- species is intermediate in ecology, morphology, and distribu- tion between A. tridentata subsp. vaseyana and A. tri- dentata subsp. tridentata. In some field situations all may grow together. Credit must be given to the techniques of thin-layer chromotography for final separation of these closely related types. Young (1965) found this undescribed subspecies to have the “widest distribution of any occurring in Sublette County, Wyoming.” He says it is “a dwarf shrub with small leaves, often suggestive of A. nova, with flower and fruit char- acters similar to representatives of A. tridentata subsp. tridentata. The chromatographic data show that it is more closely related to A. tridentata subsp. tridentata than to A. nova. The most distinguishable feature of the chromato- gram is the presence of a heavy absorption cone extending from R:-.12 to R:-.30. In the field this newly recognized subspecies is found associated with A. tridentata subsp. tridentata but always occupying the poor sites (e.g. shallow soils of hilltops and flats). A. tridentata subsp. (wyomingensis) subsp. nov. Subspecies inter A. tridentata subsp. tridentata et A. tridentata subsp. vaseyana intermedia; arbuscula, usque ad 1 m. longa; numquam propagata ut A. tridentata subsp. vaseyana; folia angustata cuneata, tridentata, 1-2 em. longa; panicula angusta; floreres 5 - 8. “Type: Wyoming, Sublette County, North of Pinedale, and % mile North of Daniel Junction, growing on Pleistocene gravels, and forming a sagebrush-thickspike-wheatgrass community, collected July 20, 1964 by Alvin L. Young No. 105, and deposited in the Rocky Mountain Herbarium. *Published with approval of the Director, Wyoming Agricultural Experiment Station, as Journal Paper No. 263. 405 406 Rhodora [Vol. 67 This collection was chosen as the type because it was it- self identified through thin-layer chromatograhpy and be- cause it is representative of the subspecies. The particular subspecific epithet for it is given because the major portion of the distribution of the plant is within the state of Wyoming and because the research leading to its recognition was carried out there. Recognition of this third subspecies in the A. tridentata complex will help to sharpen the management picture since the sites occupied by the three subspecies are somewhat dif- ferent, and the increase in grass following their control by chemicals is, therefore, different. ARTEMISIA TRIDENTATA subspecies tridentata subspecies wyomingen- basin big sagebrush subspecies vaseyana sis mountain big sagebrush Wyoming big sagebrush the commonest big sage- the common big sage- the common big sage brush at 5,000 ft. brush in Montana and brush at 7,000 feet and or below Wyoming and n. Colo- above rado at 5,000 to 7,000 feet no tendency to layer no tendency to layer strong tendency to layer dry, deep-soil sites heads few-flowered heads paniculate leaves narrowly lanceolate dry, shallow-soil sites heads intermediate heads intermediate leaves narrowly cuneate RANGE MANAGEMENT SECTION UNIVERSITY OF WYOMING LARAMIE REFEREN CES deep-soil snowfall areas of mountnain slopes heads many-flowered heads spikate leaves broadly cuneate BEETLE, A. A. 1960. A study of sagebrush. Wyo. Agr. Exp. Sta. Bul. 368: 1-85. YOUNG, ALVIN. 1965, A chemical study of the taxonomy of Section Tridentatae of genus Artemisia. Wyo. Range Mgt. Issue 198: 2 - 9. GEOCAULON LIVIDUM IN NEW ENGLAND In July 1964 the Gray Herbarium asked me if I could assist the late Dr. Hans Stauffer of Zurich, Switzerland in finding Geocaulon lividum (Richards.) Fern, in the White Mountains. Dr. Stauffer came to my camp in Dummer, New Hampshire, and we spent two days searching for it on Imp Mt. and Mt. Clinton with no success. Geocaulon has not been collected frequently in New Eng- land; the herbarium of the New England Botanical Club contains fifteen collections, the Gray Herbarium five, three of which are duplicates of sheets in the Botanical Club, and the herbarium of the Boston Society of Natural History has seven, all duplicates. The most recent collection was made in 1941 but most of the collections were made prior to 1910. These collections come from twelve localities, Lubec and Roque Bluffs in coastal Washington County, Maine, Katah- din in Piscataquis County, Bald Mt. in Somerset County, Mts. Abraham and Saddleback in Franklin County and Goose Eye in Oxford County; Mt. Ingalls, Imp Mt., Mt. Clinton and the Basin Rim in Cóos County, New Hampshire; and Mt. Mansfield in Lamoille County, Vermont. My acquaintance with the species was limited to finding it once on Katahdin in 1928. On 27 August 1964 I searched for the plant on the Basin Rim and found it growing on the slopes of both Mt. Meader and Mt. Royce and a week later I visited Goose Eye and Mt. Carlo in the Maine portion of the Mahoosuc Trail and found five stations. In June 1965 I revisited Mt. Clinton and finally found a few sterile stems about a quarter of a mile south of the new Mizpah Hut. A month later I again found the species on the Bondcliff Trail in Lincoln, Grafton County, New Hampshire. Apparently this is the first record for Grafton County. I was unable to find Geocaulon in what appeared to be favorable sites on Mt. Lafayette and Mt. Kinsman in Grafton County. The follow- ing collections are on deposit in the herbarium of the New England Botanical Club: MAINE, Oxford County, north slope of Mt. Cario, Riley, Harris 26803, 1 September 1964; NEW 407 408 Rhodora [Vol. 67 HAMPSHIRE, Coos County, Webster Cliff Trail Lu mile south of Mizpah Hut, Bean Grant, Harris 27387, 22 June 1965; Grafton County, Bondcliff Trail near trail to Guyot Shelter, Lincoln, Harris 27687, 20 July 1965. Geocaulon lividum is not an easy species to find. The leaves, borne on low herbaceous stems, resemble very closely the leaves of blueberries and the two grow frequently to- gether. However my initial discovery of the plant on the Basin Rim came when I noticed some foliage in the under- growth with the characteristic purplish color that the leaves of the closely related Comandra umbellata assume in the late summer. The flowers are inconspicuous and drop early in the season, on the Basin Rim on 11 June 1965 they were past their prime. The berry-like fruit are showy but apparently scarce in the White Mountains. I could find only about a dozen fruit on over 500 stems I examined on the Basin Rim in August 1964 and the same ratio seemed to hold on Goose Eye and Mt. Carlo. On 31 August 1965 on the Basin Rim I could not find a single fruit on more than 1000 stems al- though in June some of the stems bore as many as twenty flowers or immature fruit. Gray’s Manual describes the fruit as being scarlet; all I have seen have been yellow- orange. Pease in the Flora of Northern New Hampshire states the habitat as, “mossy bogs on the upper parts of the secondary mountains, local (somewhat similar in distribution to Rubus Chamaemorus)." I found one station on Mt. Carlo in a mossy bog but all the other stations were in damp but not boggy, rather open low spruce and fir forest. On Mt. Meader some of the plants are growing in thin mossy wooded soil on ledges. Rubus Chamaemorus is abundant in the Mahoosues and on Mt. Clinton but I never found Geocaulon growing with it. The stations varied in elevation from about 2500 feet on the Basin Rim to 4100 feet on Mt. Guyot. On Mt. Royce the stand of Geocaulon, which is the largest one I have found, is at an altitude only a few feet higher than a grove of mature fruiting Quercus rubra and Epigaea repens, var. glabrifolia is growing with the Geocaulon. Many of à Thea» 1965] Chamaecyparis thyojtes — Baldwin 409 the black spruces forming the forest cover of this station are heavily infected with Arceuthobium pusillum. A thorough search for the plant in favorable habitats may reveal that Geocaulon lividum is a much more common plant in northern New England than present collections indicate. STUART K. HARRIS BOSTON UNIVERSITY FURTHER NOTES ON CHAMAECYPARIS THYOIDES IN NEW HAMPSHIRE Since the publication of my note (Rhodora 63:281-285), some additional stations in central New Hampshire have been visited, and it may be of interest to describe these briefly. These will be numbered consecutively as additions to the list previously reported. 6a. Robb Reservoir, Stoddard. Elevation 1,275. A half mile from the previously recorded station for Chamaecyparis thyoides in Stoddard thirteen additional clumps of these trees were found in April 1963. They were mingled with a thin stand of red maple on the northeast side of an open bog thru which the outlet of Rye Pond flows into Robb Reser- voir. The cedars appeared to be sprouts from old stumps and are possibly survivors of a former larger stand that may have been killed by flooding. No reproduction or young trees were observed. 7. Ring Brook, Sutton. Elevation 950 feet. This station for Chamaecyparis thyoides covers an area of about 22 acres in a swamp forming the headwaters of Ring Brook. It begins about one quarter mile west of the intersection of Baker Hill Road with Chalk Pond road leading from Sutton to Lake Sunapee. The stream was dammed by beavers a few years ago resulting in the death or severe injury of red spruce and white pine but leaving red maple and the cedar little affected. On the West side of the swamp the red maple 410 Rhodora [Vol. 67 has suppressed the cedars which are consequently very small. Many of the larger trees in this station display cankers due to infection with Gymnosporangium sp. The ground is either bare or covered with a growth of sphagnum which indicates the severity of the recent flooding. 8. Moose Brook, Hancock. Elevation 825 feet. This small stand of cedars is located along the first half mile of Moose Brook as it leaves Norway Pond and flows thru a rather wide flood plain. This plain has been the site of beaver activity in the past and is now occupied by a dam a few hundred yards upstream from the bridge on Longview Road. A seven-stem clump of cedars 4” to 6” in diameter along with a smaller double stemmed tree occur 150 ft. southeast of the dam while on the opposite bank 200 ft. downstream there are three 5” cedars. These trees have been over-topped by a high forest of white pine, red maple and hemlocks and it is doubtful if they can long sur- vive this competition. Farther downstream are two groups opposite each other only 100 yards from the highway. How- ever, they have been girdled one to six inches wide near the base possibly by beavers. It is reported that there was formerly an extensive cedar swamp on this stream and if so these few trees are doubtless the remnants. The associated vegetation near the beaver dam consists almost wholly of white pine, gray birch, and red maple with some alder and one pitch pine. The ground cover of Cas- sandra, Kalmia angustifoliia and other ericaceous shrubs is interesting as it suggests a former wet swamp soil that has been gradually built up by the repeated flooding of beaver dams until upland species have been able to invade the site. 9. Shedd Brook, Hillsboro. Elevation 970 feet. Scattered trees of Chamaecyparis thyoides occur along Shedd Brook beginning about 300’ east of the Hillsboro-Windsor town line. The brook here meanders thru a marshy plain which shows evidence of intermittent flooding probably by beavers. Much of the sparse stock of red maples is dead as well as some of the cedars. There are five or six large cedars 8 to ee 1965] Chamaecyparis thyoites — Baldwin 411 12 inches d.b.h. and up to fifty ranging from two to five inches in diameter. Some of these latter appear to be sprouts from old stumps. The trees are scattered along both sides of the brook for half a mile with rarely more than three or four in a clump. The finding of Chamaecyparis thyoides in these localities emphasizes the point that one should not be dogmatic in stating that the tree does not occur in any wetland in this region without a thorough search. Specimens from the above stations are deposited in the Herbaria of the University of New Hampshire and the New England Botanical Club. HENRY I. BALDWIN HILLSBORO, NEW HAMPSHIRE Volume 67, No. 772 including pages 815-437, was issued December 31, 1965. EARLOW REFERENCE LIBRARY APR 1319060 ` 1965] Index to Volume 67 413 INDEX TO VOLUME 67 New scientific names and combinations are printed in bold face type Abies balsamea 92 Abildgaardia monostachya 321 Abrus abrus 334; precatorius 334 Abutilon abutiloides 339; ameri- canum 339 Acacia farnesiana 332 Acer rubrum 373, 402; saccharum 386 Achras zapota 345 Achyranthes aspera 329; bettzic- kiana 329; ficoidea 329; pep- loides 329; polygonoides 329; portoricensis 329; ranosissima 329; sessilis 328 Acisanthera quadrata 342 Acrocomia aculeata 322; media 322 Acrodiclidium salicifolium 830 Actaea pachypoda 394, 395 fig. Actinostachys pennula 356 Acuan depressum 332; insulare 332; virgatum 332 Adelia bernardia 337; ricinella 337 Adenoporces buxifolia 335 Adenoropium gossypifolium 337; hastatum 337; multifidum 387 Adiantopsis paupercula 357 Adiantum pedatum, var. aleuti- cum 382 Adipera bicapsularis 332; laevi- gata 332; stahlii 832 Aechmea lingulata 323 Aegiphila elata 350 Aeschrion antillana 336; excelsa 336 Aeschynomene gracilis 333; por- toricensis 333 Agati grandiflora 333 Agave decipiens 312 Agrimonia pubescens fig.; striata 373 acisanthera 3842; 394, 395 Agropyron repens 89 Aiphanes acanthophylla 322 Aklema petiolare 338 Alectoria nigricans 380; nitidula 380; ochroleuca 380 Algae of Forty Ponds Under Con- struction at Ithaca, New York, Soil 242 Alloplectus ambiguus 352 Alnus crispa 375; rugosa 373, 375 note Aloe barbadensis 324; 324 Alpinia antillarum 324; aromatica 824; exaltata 324; speciosa 324 Alsine antillana 330 Alsophila aquilina 356; borin- quena 356; quadripinnata 356 Alternanthera ficoidea 329; var. bettzickiana 329; peploides 329; paronychoides 329; portoricen- sis 329; ramosissima 329; ses- silis 329 Amaranthus arenicola 181; aus- tralis 313; gracilis 329; viridis 329 Ambrosia confertiflora 354 Ammophila breviligulata 89 Amomis caryophyllata 342; grisea 342 Ampelopsis arborea 181 Amphiglottis anceps 326; corym- bosa 326; difformis 326; lacera 326; nocturna 326; pallidiflora 326; secunda 326 Amphilophis ischaemum 317; sac- charoides 817 Anabaena affinis 252; cylindrica 252, 253; flos-aquae 252, 253; oscillaroides 252, 3253; sub- cylindrica 252; torulosa 252 Anabaenopsis elenkinii 252, 253 vulgaris 414 Anacharis Alsinastrum 12, 21, 26, 166; callitrichoides 166; cana- densis 12, 25, 31, 166, 169; var. Planchonii 12, 23; chilensis 166; densa 157, 166; Hillariana 166; linearis 29, 166; Matthew- sii 166; Naias 166; Nutallii 6, 166, 169; occidentalis 6, 167; Planchonii 12, 167; pomeranica 167; potamogeton 167; tripteris 167 Anacheilium cochleatum 326 Anadenanthera peregrina 332 Anamomis fragrans 343; umbel- lulifera 343 Ananas ananas 328; comosus 323 Anatherum zizanioides 318 Andrachne cuneifolia 337 - Andropogon annulatus 317; brevi- folius 317; condylotrichus 318; gracilis 317; hirtiflorus 317; pertusus 317; saccharoides 317; semiberbis 317; virginicus 317 Aneilema geniculatum 328 Anemone parviflora 92, 93, 382 Angiospermous Leaves, Determi- nation of the Developmental Pattern of 49 Anisosorus hirsutus 357 Anneslia haemostoma 331; por- toricensis 331; purpurea 331 Anredera baselloides 330; lep- tostachys 330 Anthelis arenaria 311; podanisia 302 Antigonon guatamalense 328; macrocarpum 328 Antirhea acutata 353; coriacea 353; lucida 353; obtusifolia 353; portoricensis 853; sintenisii 353 Anthriscus cerefolium 344 Antrophytum cayennense 357 Apalanthe granatensis 167; guy- annensis 167; Schweinitzii 2, 167 Aperia hymenanthera 325 Rhodora [Vol. 67 Aphanocapsa elachista 252 Apios americana 26; tuberosa 26 Apium leptophyllum 344; pet- roselinum 344 Apteria aphylla 325 Aralia racemosa 394, 396 fig. Arceuthobium pusillum 409 Arctic Species in Southeastern New Brunswick, New Records of 92 Arctostaphylos alpina 380 Ardisia glauciflora 344; luquillen- sis 344; obovata 844 Arduina grandiflora 346 Arenaria dawsonensis 381; humi- fusa 381; rubella 381 Arethusa balbosa 373 Argythamnia fasciculata 337 Aristida chaseae 319 Aristolochia bilabiata 328; cordi- flora 328; elegans 328; galeata 828; grandiflora 328; littoralis 328; oblongata 328; odoratis- sima 328 Armeria labradorica 381 Aronia floribunda 376 Artemisia nova 405; tridentata 405, 406; Complex, a Third Subspecies in the 405; subsp. tridentata 405, 406; subsp. vaseyana 405, 406; subsp. wyomingensis 405, 406 Artocarpus heterophylla 328; in- tegrifolia 328 Ascolepis brasiliensis 99 Ascyrum hypericoides 341 Aster acuminatus 377; divarica- tus 231; hookerianus 235; um- bellatus 373 Asterandra grandifolia 337 Asystasia gangetica 852 Atamosco carinata 324; rosea 324; tubispatha 324 Athyrocarpus 323 Auliza ciliaris 326 persicariaefolius 1965] Aulomyrcia citrifolia 342; lepto- clada 342 Avicennia germinans 850; nitida 350 Ayenia insularis 340; pusilla 340 Babingtonia pestifera 167 Bacopa innominata 351; mon- nieri 351; repens 351; stricta 351 Bactris acanthophylla 322 Badiera penaea 836; portoricensis 336 Baldwin, Henry I., Further Notes on Chamaecyparis thyoides in New Hampshire 409 Bambos vulgaris 320 Bambusa bambos 320 Banisteria laurifolia 336; lucida 335; purpurea 836; wydleriana 336 Banisteriopsis lucida 335 Bartsia alpina 381 Bartonia paniculata 376 Basella alba alba 330; rubra 830 Batocydia unguis 352 ` Beadlea cranichoides 325; 325 Beaman, John H. and D. C. D. DeJong, A New Species of Lag- enophora (Compositae) from Guatemala 36 Becquerelia glomerulata 125 Beetle, Alan A. and Alvin Young, A ‘Third Subspecies in the Artemisia tridentata Complex 405 Beilschmiedia pendula 330 Benitoa occidentalis 218, 224 Benthamantha caribaea 383 Bernardia dichotoma 337 Betula borealis 380; glandulosa 382; lutea 92, 373; minor 382; papyrifera 402 Bianeaea sepiaria 333 Bidens cernua 61, 62; Notes on elata Index to Volume 67 415 Varieties of Bidens connata and a Hybrid with 59; connata 60, 61, 62; var. connata 61; var. gracilipes 59, 62; var. pinnata 59, 62; var. saint-vincentii 60, 61; var. submutica 61; X B. cernua 62; cynapiifolia, var. portoricensis 355; var. tenuis 355; reptans, var. urbanii 355; urbanii 355 Bignonia caryophyllea 352 Bihai bihai 324 Blechnum divergens 358; lineatum 358; polypodioides 357; under- woodianum 358 Blechum blechum 352; pyramida- tum 852 Boerhaavia coccinea 329; diffusa 329 Bolbitis aliena 358; cladorrhizans 358; nicotianaefolia 358; perga- mentacea 358 Bolivia, Studies in the Flora of, III — Cyperaceae, Part 1 97 Borreria brachysepala 354; levis, var. sintenisii 354 Bouchea prismatica, var. brevi- rostra 850; var. longirostra 349 Bourreria revoluta 348; succul- enta, var. revoluta 348 Boussingaultia baselloides leptostachys 330 Brachyramphus intybaceus 354 Bradburya plumieri 384; pube- scens 334; virginiana 334 Bramia monnieri 351 Brassavola cucullata 326 Brassica campestris 331; kaber 331; rapa 331 Browne, Elizabeth M. and E. T. Browne, Jr., Flowering Plants New to or Rare in Kentucky 180 Brugmansia arborea 351; suaveo- lens 351 Bulbophyllaria pachyrrachis 327 330; 416 Bulbophyllum pachyrrachis 327 Bulbostylis boliviana 114; capil- laris 111, 113; var. tenuifolia 113; Funckii 110; hirtella 111, 113; junciformis 111, 114; jun- coides 111, 112; var. ampliceps 111, 112; papillosa 111, 113; paradoxa 111; pauciflora 821; sphaerocephala 111, 112; var. macrocephala 112; sphaerolepis 112; tenuifolia 113 Bumelia krugii 345; obovata, var. krugii 345 Burch, Derek, Two Species of Chamaesyce (Euphorbiaceae) new to the United States 185 Bursera simaruba 336 Buxus laevigata 338; portoricen- sis 338; vahlii 338 Byrsonima cuneata 336; horneana 336; lucida 336; ophiticola 386; spicata 336; wadsworthii 336 Cacalia atriplicifolia 55, 56; and C. muhlenbergii, Natural and Artificial Hybrids of 55; muhl- enbergii 55, 56; suaveolens 55, 56; tuberosa 55, 56, 57 fig. Cacara erosa 335 Caconapea stricta 351 Cactus intortus 341 Caesalpinia bondue 383; coriaria 333; crista 333; culebrae 383; divergens 383; melanosperma 333; portoricensis 333; pulcher- rima 383; decapetala 333 Cajan cajan 334 Cajanus cajan 334 Caladium bicolor 323; 328 Calliandra caracasana 331; hae- matostoma 331; purpurea 331 Calliearpa americana 394, 396 fig. Calonyetion aculeatum 347; tuba 347 Calophyllum colocasia antillarum 340; Rhodora [Vol. 67 calaba 840 Calopogon pulchellus 376 Calopogonium mucunoides 335; orthocarpum 335 Calothrix stagnalis 252 Calyptrocarya fragifera 125; glo- merulata 125 Calyptracordia alba 348 Calyptranthes dumetorum 343; luquillensis 343; pallens 343; peduncularis 343; triflorum 348 Calyptrogyne occidentalis 322 Calyptronoma rivalis 322 Campanula americana 394, 395 fig.; rotundifolia 92 Campylocentrum fasciola 327; sul- livani 327 Cananga caribaea 380; blainii 330 Canavalia brasiliensis 335; ensi- formis 335; gladiata 335; nitida 335; rusiosperma 335 Capriola dactylon 319 Cardiospermum halicacabum, var. microcarpum 338; microcarpum 338 Carex angustior 361; annectens 198, 199; atlantica 361, 362; var. incomperta 362; atrofusca 382; bicolor 381; bigelovii 380; boliviensis 134, 137; Bonplandii 134, 135; brunnescens 402; capillaris 382; capitata 379, 381; castanea 373; cephalantha 361; chordorrhiza 379, 381; cladostachya, var maxima 134, 135; crinata 373; debilis 373, 402; decidua, var. Brehmeri 134, 136; emmonsii 402; fe- cunda 134, 137; fissa 198, 199; ((SMultiflorae), An Eastern Variety of 198; var. aristata 198, 199; var. fissa 199; folli- culata 376, 402; Goodenowii, var. stolonifera 136; gracillima 373; houghtonii 374, 386, 389; howei 361; incomperta 361, 1965] 362; Bickn., The Status of 361; interior 361; intumescens 373, 402; Jamesonii, var. subfulva 134, 137; lacustris 377; Mando- niana 135; microglochin 379, 381; mohrii 361; muricata 361; nebularum 134; ormantha 361; phalaroides 184, 138; var. moesta 138; phyllomanica 361; pichinchensis 134, 136; var. obtusisquamea 137; pinetorum 134, 135; rariflora 382; rupe- stris 381; ruthii 361; scirpoidea 382; sterilis 361; terrae-novae 381; trisperma 402; wiegandii 361 Carissa macrocarpa 346 Carum Carvi 196 Casearia bicolor 341 Cassandra 410 Cassia aeschynomene 832; alata 332; antillana 332; bicapsularis 331; emarginata 331; exunguis 833; diffusa 332; diphylla 332; fruticosa 332; glandulosa, var. Swartzii 332; grammica 332; granulata 332; hirsuta 832; javanica 332; laevigata 332; mirabilis 332; nodosa 832; obovata 332; obtusifolia 332; occidentalis 332; patellaria 832; polyphylla 332; portoricensis 332; rotundifolia 333; stahlii 332 Cassine xylocarpa 338 Cassiope hypnoides 380 Cassipourea alba 843; guianensis 343 Castalia amazonum 330; ampla 330; jamesoniana 330; odorata 830; pulchella 330; rudgeana 330 Castela erecta 336 Castelaria nicholsoni 336 Castilleja septentrionalis 381 Catalpa longissima 352 Index to Volume 67 417 Catopsis floribunda 328; nutans 323 Celtis iguanaea 327 Cenchropsis myosuroides 319 Cenchrus brownii 319; carolin- ianus 319; incertus 319; myosu- roides 319; viridis 319 Centaurea Jacea 197; nigra, var. radiata 197 Centella asiatica 344; erecta 344 Centrogenium setaceum 325 Centrosema plumieri 334; pubes- cens 334; virginianum 334 Centrostachys aspera 329; indica 329 Cephalocereus royeni 341 Cerastium beeringianum 381 Ceratophyllum demersum 196 Ceratostoma portoricensis 344 Cerbera thevetia 346 Cerdana alliodora 348 Cerefolium cerefolium 344 Cestrum daphnoides 851; salici- folium 351 Cetraria cucullata 380; nivalis 380; tilesii 381 Chaetochloa barbata 319; geni- culata 319; magna 319; rariflora 319; setosa 319; tenacissima 319; vulpiseta 319 Chalcas exotica 836 Chamaecrista aeschynomene 332; antillana 332; chamaecrista 332; diphylla 332; exunguis 833; grammica 332; granulata 332; mirabilis 832; patellaria 332; portoricensis 332; swartzii 332 Chamaecyparis thyoides 409, 410, 411; in New Hampshire, Fur- ther Notes on 409 Chamaedaphne calyculata 365 Chamaefistula bacillaris 332 Chamaesaracha villosa 194; New to Texas, 194 Chamaesyce adenoptera 185; (Eu- phorbiaceae), New to the 418 United States, Two Species of 185; maculata 185; mendezii 185, 186; prostata 185; thymi- folia 185, 186 Characium obtusum 251 Cheilanthes paupercula trachomanoides 357 Cheiroglossa palmata 9856 Chelone cuthbertii 394, 396 fig. Chenopodium pumilio 181 Chimaphila umbellata 386, 389 Chinia boxiana 349; spinosa 349 Chione seminervis 353 Chlamydomonas aggregata 248, 252; snowii 252; typica 248, 252 Chlorella ellipsoidea 251; vulgaris 251 Chloris inflata 319; paraguayen- sis 319; petraea 319; sagraena 319 Chlorococcum humicolum 249 Chlorosarcinopsis eremi 248, 251 Chroococcus minor 252 Chrysanthemum indicum 856; morifolium 356 Chrysobalanus icaco 331; pello- carpus 331 Chrysophyllum bicolor 345; eg- gersii 345 Chrysopsis berlandieri 87 fig.; camporum 87 fig.; (Compositae- Astereae), A Second Character Distinguishing Heterotheca s. str. from 86; divaricata 231; fulerata 87 fig.; graminifolia 88; stenophylla 87 fig.; villosa 87 fig. Cicca disticha 387 Cinna latifolia 402 Cinnamomum elongatum 331; montanum 331; zeylanicum 331 Cissus tuberculata 339 Cistus canadensis 64, 255, 258, 269, 272; carolinianus 211; corymbosus 287; glomeratus 298; Mexicanus 298; rosmarini- 357; Rhodora Coccolobis [Vol. 67 folius 311; Virginianus 260 Citharexylum fruticosum, var. subvillosum 350; var. villosum 350; pentandrum 350; tris- tachyum 350; X perkinsii 350 Citrullus citrullus 354; lanatus 354 Citrus grandis 336; limon 336; limonum 336; maxima 836 Cladium jamaicense 114; 322 Clarkia davyi 230; prostrata 230 Clavenna tetrandra 353 Claytonia caroliniana 374, 386 Cleome ciliata 331; icosandra 381; rutidosperma 331; viscosa 331 Clerodendrum aculeatum 350; bungei 850; indicum 350; in- erme 350 Cleyera albopunctata 340 Clintonia borealis 402 Clitoria laurifolia 334; rubiginosa 334 Clusia grisebachiana 341; giana 341 Coccocypselum herbaceum 853 Coccoloba costata 329; diversifolia 329; microstachya 329; pube- scens 329; swartzii 328; var. portoricensis 329; b. urbaniana 329 kru- borinquensis 328; grandifolia 329; laurifolia 329; obtusifolia 329; rupicola 329 Coccothrinax alta 322; discreta 322; eggersiana 322; sancti- thomae 322 Cochleanthes flabelliformis 327 Cochlospermum vitifolium 341 Cochranea auchusaefolia 349 Coelastrum microsporum 251 Coelia triptera 326 Coleman, James R., Natural and Artificial Hybrids of Cacalia atriplicifolia and C. muhlen- bergii 55 Colocasia esculenta 323 1965] Colubrina arborescens 339; col- ubrina 339; urbanii 339 Comandra umbellata 408 Commelina diffusa 323; longi- caulis 328 Comptonia peregrina 394, 395 fig. Conami portoricensis 337 Congea tomentosa 350 Conyza apurensis 355; bonarien- sis 355; canadensis, var. pusilla 355; floribunda 355 Coptis groenlandica 402 Cordia alba 847; alliodora 348; bahamensis 348; bellonis 348; collococca 348; dentata 348; gerascanthus 348; glabra 348; globosa 348; lima 348; poly- cephala 348; rickseckeri 348; rupicola 348; sebestena 348; stenophyllum 348 Cordyline guinensis 324 Coreopteris Tripteris 167 Cornus alternifolia 374; densis 402 Coronopus didymus 331 Corylus cornuta 373 Corynella pauciflora 338 Costa Rica, A New Species of Senecio from 239 Cracca caribaea 333; cathartica 333; cinerea 833 Cranichus diphylla 325 Crantzia ambigua 352 Crassina elegans 355; multiflora 355 Crepis capillaris 197 Critonia portoricensis 355 Crocanthemum Aldersonii 279; arenicola 264; argenteum 305; Berlandieri 260; Bicknellii 269; brasiliense 65; canadense 255; carolinianum 65, 201, 212; corymbosum 288; domingense 302; dumosum 214; exaltatum 289; georgianum 291; glomera- tum 299; majus 269, 272; Nashii cana- Index to Volume 67 419 266, 268; nutans 284; occiden- tale 276; Pringlei 294; propin- quum 274; rosmarinifolium 302; scoparium 279; stenophyl- lum 302; stipulatum 70, 291, 293; thyrsoideum 266, 268 Crossing Diagrams, Interpreta- tion of 146 Crossopetalum rhacoma 388 Crosswhite, Frank S. and Shoichi Kawano, Chromosome Numbers in Penstemon (Scrophularia- ceae) I: New Mitotic Counts 187 Croptilon divaricatum 217, 218, 281; var divaricatum 218; var. hirtellum 218, 229; var. hook- erianum 218, 235; hookeria- num 235 Crotalaria saltiana 333; striata 333 Croton trinitatis 387 Ctenitis effusa 359; hirta 359; nemorosa 359; subincisa 359 Cuphea micranthea 342; par- sonsia 342 Cureas curcas 387; hernandiae- folius 337 Cucurbita lagenaria 354; moscata 354 Cuscuta campestris 348; gland- ulosa 348; globosa 348; obtusi- flora, var. glandulosa 348; pent- agona 348 Cyathea aquilina 356; borinquena 356; escuquensis 356; horrida 356; wilsoni 356 Cyclospermum leptophyllum 344 Cylindrospermum catenatum 252; marchicum 252 Cymodocea manatorum 817 Cynanchum anegadense 346; cheesmanii 346; decaisneanum 346; ephedroides 346; grise- bachianum 346; lineare 346; monense 345; parviflorum 346 420 Cynodon dactylon 819 Cynoglossum furcatum 349 Cyperaceae, Part I, Studies in the Flora of Bolivia III 97 Cyperus brevifolius 320; brunneus 320; calcicola 820; cayennensis 320; densicaespitosus 320; dif- formis 320; digitatus 320; eg- gersii 320; ferax 320; flavus 320; flexuosus 320; filiformis 320; granularis 320; imbricatus 320; incompletus 320; iria 320; mutisii 320; nanus 820; var. subtenuis 320; obtusata 320; odoratus 320; pedunculatus 320; peruvianus 320; planifolius 320; polystachyos 820; radiatus 320; swartzii 320; tenuis 320; uni- folius 320; unioloides 320; vahlii 320 Cypripedium acaule 373, reginae 375 Cyrtospadix bicolor 323 Cystopteris bulbifera 92 402; Dactylococcopsis smithii 252 Dalbergia ecastophyllum monetaria 334 Dalea domingensis 333 Damman, A. W. H., The Distri- bution Patterns of Northern and Southern Elements in the Flora of Newfoundland 363 Daoud, H. S. and Robert L. Wil- bur, A Revision of the North American Species of Helianthe- mum (Cistaceae) 63, 201, 255 Daphne Mezereum 196 Datura candida 351; fastuosa 351; inoxia 851; metel 351; suaveo- lens 351 Davis Edw. L., Determination of the Developmental Pattern of Angiospermous Leaves 49 DeJong, D. C. D. and John H. Beaman, A New Species of 334; Rhodora [Vol. 67 Lagenophora (Compositae) from Guatemala 36 Dendropemon bicolor 328; cari- baeum 328; purpureum 328; sintenisii 328 Dendrophthora flagelliformis 328; wrightii 328 Dendropogon usneoides 328 Dennstaedtia adiantoides 360; bipinnata 360; cicutaria 360; obtusifolia 360; ordinata 360; punctilobula 377; rubiginosa 360 Deschampsia atropurpurea 380 Desmanthus virgatus 332 Desmodium adscendens 834; affine 334; axillare 334, var. acuti- folium 334; var. stoloniferum 334; barbatum 834; canum 334; cubense 334; glabrum 334; intortum 834; procumbens 334; tortuosum 334; scorpiurus 334; triflorum 334; wydlerianum 334 Diapedium assurgens 352; krugii 352 Diapensia lapponica 380 Dichanthium annulatum 317 Dichromena ciliata 123, 124, 321; colorata 321; Kuntzei 116; Mandonii 118; radicans 321 Dichylium oerstedianum 338 Diclipptera assurgens 352; bra- chiata 181; krugii 352 Dicranopteris bifida 356; flexuosa 356; pectinata 356 Didymopanax gleasonii 344 Diervilla lonicera 373 Digitaria adscendens 318; argil- lacea 318; horizontalis 818; ischaemum 318; panicea 318; villosa 318; violascens 318 Dilomilis montana 326 Diodia maritima 354; 354 Dioscorea bulbifera 824; cayan- serrulata 1965] nensis 324; esculenta 324; vil- losa 324 Diospyros digyna 345; discolor 345; ebenaster 345; philippen- sis 345; sintenisii 345 Dipholis cubensis 345; sintenisii 345 Diplachne fascicularis 319 Diplacrum longifolium 126 Diplandra potamogeton 167 Diplazium arboreum 357; crista- tum 357 Diplopappus divaricatus 231 Dirca palustris 394, 395 fig. Dissecting Equipment and Mate- rials for the Study of Minute Plant Structures 95 Ditaxis fasciculata 337 Ditremexa hirsuta 332; occident- alis 332; sophera 332 Dodonaea ehrenbergii 338; viscosa 338 Dolicholus minimus 335; pyra- midalis 335; reticulatus 835 Domingoa haematochila 326; hymenodes 326 Dondia fruticosa 329 Doxantha unguis-cati 352 Drepanocarpus lunatus 334 Dryas integrifolia 92, 93, 381 Dryopteris angustifolia 359; brit- tonae 359; chaerophylloides 359; cristata 402; decussata 358; deltoidea 358; dentata 359; domingensis 359; effusa 359; germaniana 358; gongy- lodes 358; guadalupensis 359; hastata 359; hirta 359; lepto- cladia 359; linkiana 358; mar- ginalis 377; megalodus 359; nemorosa 359; normalis 359; noveboracensis 376; oligophylla 359; opposita 358; patens 359; piedrensis 358; poiteana 359; reptans 359; reticulata 359; robertiana 375; sancta 358; Index to Volume 67 421 sclerophylla 359; serra 359; serrata 359; sprengelii 358; subincisa 359; subtetragona 359 Duchartrea sintenisii 352 Duggena hirsuta 352 Dulichium arundinaceum 192, 373 Ecastophyllum ecastophyllum 834 Echinochloa colona 318; colonum 318; crus-galli 318; crus-pav- onis 318 Echinodorus berteroi, var. ber- teroi, 317; cordifolius 317; ten- ellus, var. latifolius 317 Echites agglutinata 346 Egeria densa 2, 26, 28, 156, 157, 161, 163, 166, 168, 170; Naias 166, 168, 169 Eichhornia crassipes 323; diversi- folia 323 Elaeodendron xylocarpum 338 Elaphoglossum crinitum 357; flac- cidum 856; longifolium 356; martinicense 356; maxonii 356; philoselloides 356; pteropus 356; pumilum 356; rigidum 356; serpens 357; underwood- ianum 356 Elaphrium sinaruba 386 Eleocharis albibracteata 105, 107; var. nubigena 107, 108; atro- purpurea 320; boliviana 107; Brehmeriana 107; capitata 106; caribaea 320; costulata 105; crinalis 105, 107; Dombeyana 105, 107; elegans 104, 108, 321; exigua 105; fallax 321; filicul- mis 105, 109; fistulosa 104, 105; flaccida 320; flavescens 320; geniculata 104, 106, 108, 320, 321; interstincta 104, 105; maculosa 105, 106; microcarpa 321; minima 105, 108; minuti- flora 321; montana 320; nodu- losa 104, 108, 820; nubigena 107; obtusa, var. jejuna 195; 422 radicans 104, 105, 321; rostel- ocreata, var. flaccida 106; lata 321; Sellowiana 105, 106; sintenisii 320; sulcata 109; yunquensis 320 Elephantopus mollis 354; scaber 354 Eleutherine bulbosa 324 Elodea bifoliata 165, 167; Bran- degeae 3, 164, 165, 167; calli- trichoides 161, 164, 166, 167; canadensis 2, 5, 12, 13 pl., 14 fig., 15, 21, 22, 23, 24, 25, 26, 27, 28, 29, 31, 33 fig., 156, 159, 161, 162, 163, 165, 166, 167, 168, 169, 170, 171; var. angustifolia 6, 168; var. B angustifolia 167; var. gigantea 156, 168; var. latifolia 12, 166, 168; sub. var. repens 12, 166; var. Planchonii 12, 166; capensis 168; chilensis 168; chinensis 168; columbiana 11, 165, 168; crispa 168; densa 26, 156, 168; var. crispa 168; var. longifolia 157, 168; Ernstae 161, 165, 166, 168; gigantea 28, 156, 157, 168; granatensis 8, 159, 164, 165, 166, 167, 168, 169, 170, 171; guyanensis 21, 168; var. dicranoides 168; f. longi- folia 168; ioensis 12, 168; iowen- sis 12, 168; Kochii 168; latifolia 12, 21, 169; linearis 2, 23, 29, 30 fig., 31, 33 fig., 165, 166,169, 170; longivaginata 2, 30 fig., 32 fig., 34, 164, 165, 166, 169; Matthewsii 161, 165, 166, 169; minor 6, 169; Monograph of the Genus: Part 4 and Summary 1, 155; Naias 166, 168, 169; neva- densis 165, 169; Nutallii 2, 5, 6, 11, 12, 14 fig., 21, 26, 29, 31, 32 fig., 34, 155, 161, 164, 165, 166, 167, 168, 170, 171, 195; oblongifolia 12, 15, 169; occiden- talis, 6, 7, 169; orinocensis 169; Rhodora [Vol. 67 paraguayensis 169; peruvien- sis 165, 169; Planchonii 12, 21, 22, 23, 24, 169; Potamogeton 164, 165, 166, 167, 168, 169, 170; Richardii 164, 166, 169; Schweinitzii 2, 3, 4, 5, 6, 21, 25, 30 fig., 32 fig., 164, 166, 167, 170; titicacana 164, 165, 170; verticillata 170 Elsota diversifolia 386; virgata 336 Eltroplectris acuminata 325 Emelista tora 332 Emilia coccinea 355; sagittata 355 Empetrum eamesii 380 Encyclia fucata 826; krugii 326; papilonacea 326; sintenisii 326 Epidendrum anceps 326; belveder- ense 326; brittonianum 326; ciliare 326; cochleatum 326; conopseum 48; and Eulophia ecristata in Mississippi 48; cucullatum 326; difforme 826; fucatum 326; krugii 326; lacerum 326; moirianum 326; nocturnum 326; ottonis 326; pallidiflorum 326; ramosum 326; rigidum 326; secundum 326; sintenisii 326; vicentinum 326 Epifagus virginiana 386 Epigaea repens 375; var. glabri- folia 408 Epilobium latifolium 382 Equisetum fluviatile 195 Eragrostis glutinosa 319; multi- caulis 195; pectinacea 195; tracyi 312, 313; from Sanibel Island, Florida, The Extended Distribution of 312; urbaniana 319 Erechtites valerianaefolia 355 Eremolepis wrightii 328 Erigeron hyssopifolius 92 Eriochloa gracilis 181; poly- stachya 318; subglabra 318 1965] Eroteum albopunctatum 340 Erythrina eggersii 335; horrida 335 Erythrodes hirtella 325; planta- ginea 325 Erythronium americanum 372 Erythroxylum brevipes 335; rotundifolium 335; rufum 335 Eugenia aerunginea 3943; buxi- folia 343; domingensis 343; far- jardensis 343; floribunda 343; glabrata 843; haematocarpa 343; maleolens 343; margaret- tae 343; saviaefolia 342 Eulophia alta 327; ecristata 48 Euphorbia adenoptera 185 Eupatorium borinquense 354; corymbosum 354; geraniifolium 355; iresinoides 355; ivaefolium 355; macrophyllum 354; odora- tum 354; oteroi 355; portori- censis 855; sinuatum 355 Euphorbia adenoptera 185; oer- stediana 338; petiolaris 338 Eustachys petraea 319 Euterpe globosa 322 Evolvulus alsinoides, var. lini- folius 347; bocasanus 347; lini- folius 847; tenuis, var. longi- folius 347 Exogonium arenarium 347; re- pandum 347; solanifolium 347 Exostema ellipticum 353 Fagus grandifolia 386 Fern in Wisconsin, Discovery of the Massachusetts 399 Ficus citrifolia 327; crassinervia, f. stahlii 327; laevigata 927; nitida 328; perforata 327; retusa 328; sintenisii 327; tri- gonata 327 Fimbristylis aestivalis 109, 110; annua 109; curassavica 321; cymosa 321; dichotoma 821; diphylla 321; monostachya 109, Index to Volume 67 423 110, 321; portoricensis 321; spathecea 321; vestita 321 Flaveria floridana 313; trinervia 855 Flora of Newfoundland, The Dis- tribution Patterns of Northern and Southern Elements in the 363 Flora of Oklahoma, Additions to the Vascular 191 Flora of the Peninsula of Vir- ginia, The Mountain Element in the 393 Florida, The Extended Distribu- tion of Eragrostis tracyi from Sanibel Island 312 Flueggea virosa 337 Foeniculum foeniculum 344; vul- gare 344 Forsteronia corymbosa 346; por- toricensis 346 Forestiera porulosa 313 Foster, Robert C., Studies in the Flora of Bolivia III — Cy- peraceae, Part I 97 Fraxinus nigra 374 Fuirena squamosa 321; umbellata 1083, 821 Funastrum clausum 346 Galathea bulbosa 324 Galax aphylla 394, 395 fig. Galium triflorum 92 Galphimia gracilis 336 Gaultheria procumbens 374, 386, 389 Gaylussacia baccata 376; dumosa 376 Geocaulon lividum 407, 408, 409; in New England 407 Geranium robertianum 374 Gillett, George W., Genotypic Variations in the Phacelia hir- suta Complex 42; Genotypic Variation in the Phacelia stricti- flora Complex 139 424 Geophila herbacea 354; 354 Gerardia portoricensis 852; tu- berosa 352 Gesneria albiflora 352; sintenisii 352 Gleichenia bifida 356; 356; pectinata 356 Gliricidia sepium 338 Globifera umbrosa 351 Gloeocystis ampla 252; gigas 252; vesiculosa 251 Gnaphalium americanum spicatum 355 Gomphrena decumbens 329; dis- persa 329 Gonium pectorale 252 Gonocalyx portoricensis 344 Gonolobus stephanotrichus 846 Gonzalaguinia spicata 353 Gossypium arboreum, var. nadam 340; barbadense 340 Graffenrieda ottoschulzii 342 Grammitis asplenifolia 357; hartii 357; hessii 357; jubaeformis 357; mollissima 357; myosu- roides 357; serrulata 357; sus- pensa 357; taenifolia 357; taxi- folia 357; trifurcata 357 Gratiola aurea 197 Guatemala, A New Species of Lag- enophora (Compositae) from 36 Guatteria blainii 330; caribae 330 Guazuma guazuma 340; ulmifolia 340 Guettanda laevis 353; valenzue- lana 353 Guilandina bonduc 333; crista 333; culebrae 333; divergens 333; melanospermum 333; por- toricensis 333 Gymnosiphon germanii sphaerocarpus 325 Gymnosporangium sp. 410 repens flexuosa 355; 325; Habenaria blephariglottis 376; Rhodora [Vol. 67 ciliaris 394, 395 fig.; conopsea 35; orbiculata 376; straminea 381 Hackelochloa granularis 317 Haemocharis portoricensis 340 Haenianthus obovatus 345; salici- folius, var. obovatus 845 Halimium Aldersonii 279; areni- cola 264; argenteum 305; Ber- landieri 260, 263; canadense 255; carolinianum 212; chihua- huense 296; concolor 289; cory- mbosum 288; Coulteri 260; dis- color 289; domingense 302, 304; exaltatum 289; georgianum 291; glomeratum 298, 311; majus 269; Nashii 266; nutans 284; occidentale 67, 276, 279; patens 263, 285; Pringlei 263, 294; rosmarinifolium 302; scoparium 67, 279; spartioides 67; steno- phyllum 302, 304 Halodule beaudettii 317 Hamamelis virginiana 402 Hamelia erecta 353; patens 353 Hapalorchis tenuis 325 Haplanthe Schweinitzii 170 Haplopappus, Section Isopappus (Compositae), Taxonomy of 217 Haplopappus bartlettii 218; di- varicatus 217, 218 219 220 fig., 221, 222, 223 fig., 224, 225, 226, 227, 228, 229, 281, 232, 235, 237; X graniticus 222; X torreyi 222; X validus 222; var. hookerianus 235; hirtellus 230; hookerianus 235; occidentalis 218, 224; rigidifolius 217, 218 fig., 220 fig., 221, 222, 223, 224, 225, 226, 227, 228, 229, 230, 231, 234, 235, 237; X divari- catus 227; validus 225, 226, 227, 228; subsp. graniticus 217, 220 fig., 221, 222, 223, 224, 225, 226, 229, 233, 236, 237; subsp. torreyi 217, 220 fig., 222, 223, 1965] Index to Volume 67 425 225, 226, 227, 229, 232, 233, 237; X graniticus 222; X H. rigidi- folius 223; X subsp. validus 217, 220 fig., 221, 222, 223, 224, 225, 226, 229, 283, 235, 237; X grani- ticus 222; X torreyi 226 Harms, Vernon L., A Second Char- acter Distinguishing Hetero- theca s. str. from Chrysopsis (Compositae: Astereae) 86 Harris, Stuart K., Geocaulon lividum in New England 407; Notes on the Flora of Coos County, New Hampshire 195 Harrisia Aboriginum 313 Hartley, Thomas G., Discovery of the Massachusetts Fern in Wis- consin 399 Harvill, A. M., Jr., The Mountain Element in the Flora of the Peninsula of Virginia 393 Hebeclinium macrophyllum 354 Hedysarum alpinum 381 Helenium brevifolium 3894, 396 fig.; curtissii 394 Helianthemum Aldersonii 278, 281; arenicola 67, 71, 74, 75, 76, 77 fig., 79, 203, 260, 263, 264, 265, 266, 267, 268, 308 fig., argenteum 70, 71, 74, 76, 77 fig., 81, 203, 304, 305, 310 fig., 311; astylum 65, 311; Bicknellii 66, 69, 71, 78, 75, 76, 77 fig., 80, 207, 210, 211, 255, 257, 258, 269, 270, 271, 272, 273, 275, 307, fig.; brasiliense 63, 65, 213; canadense 65, 66, 69, 70, 71, 73, 76, 77 fig., 78, 79, 201, 206, 209, 211, 215, 216, 255, 257, 258, 272, 278, 275, 276, 307 fig., 311; var. obtusum 264, 265; var. sabulonum 255, 259; var. Walk- erae 269, 272; capitatum 66; carolinianum 65, 67, 69, 70, 71, 72, 74, 76, 76, 77 fig., 78, 79, 201, 204, 211, 212, 218, 306 fig.; chihuahuense 69, 70, 71, 77 fig., 81, 204, 295, 297, 809 fig.; (Cistaceae), A Revision of the North American Species of 63, 201, 255; concolor, 71, 77 fig., 81, 204, 262, 289, 290, 310 fig.; corymbosum 64, 65, 66, 71, 72, 73, 77 fig., 80, 81, 204, 265, 266, 268, 287, 308 fig.; Coulteri 69, 70, 71, 78, 74, 75, 76, 77 fig., 79, 204, 207, 208, 210, 260, 262, 263, 809 fig.; dumosum 69, 70, 71, 76, 77 fig., 78, 79, 206, 209, 211, 214, 215, 216, 257, 258, 275, 308 fig.; exaltatum 290; georgianum 70, 71, 77 fig., 81, 206, 208, 210, 274, 276, 291, 292, 306 fig.; glomeratum 64, 65, 66, 71, 72, 74, 77 fig., 81, 201, 206, 208, 210, 262, 297, 298, 300, 303, 304, 309 fig.; Greenei 67, 69, 70, 71, 72, 77 fig.; 80, 203, 276, 278, 279, 283, 287, 308 fig.; hirsutissimum 64; majus 66, 255, 269, 272, 273; mendocinen- sis 279; Nashii 67, 71, 73, 74, 15, 16, 77 fig., 79, 203, 265, 266, 267, 268, 308 fig.; nutans 69, 70, 71, 72, 76, 77 fig., 80, 203, 278, 284, 286, 287, 810 fig.; obeordatum 65, 298; occiden- tale 276; patens 69, 70, 72, 73, 74,77 fig., 80, 203, 205, 278, 285, 286, 287, 310 fig.; polifolium 302; polygalaefolium 65; Prin- glei 69 70, 71, 74, 75, 77 fig., 81, 204, 205, 273, 292, 293, 297, 309 fig.; propinquum 69, 71, 73, 15, 76, 77 fig., 80, 207, 210, 211, 257, 258, 271, 273, 274, 275, 307 Do: ramuliflorum 65, 255; rosmarinifolium 64, 65, 66, 70, 71, 75, 77 fig., 81, 205, 207, 208, 301, 308, 304, 306 fig.; scopa- rium 64, 67, 70, 72, 78, 77 fig., 80, 203, 205, 277, 278, 280, 281, 426 285, 287; var. Aldersonii 279, 281, 282; var. scoparium 282; var. vulgare 279, 280, 281, 282; spartioides 64, 278; suffrute- scens 68, 279, 281, 282; tenellum 317; thyrsoideum 266, 268; tri- petalum 65, 511; Walkerae 269 Helichrysum bracteatum 355 Heliconia caribaea 324 Heliophytum amplexicaule 349 Heliotropium amplexicaule 349; angiospermum 348; arbor- escens 349; crispiflorum 348; indicum 348; inundatum 848; microphyllum 348; peruvianum 349; procumbens 348 Hemicarpha micrantha 99 Hemidiodia ocymifolia 354 Hemitelia escuquensis 356; hor- rida 366; wilsoni 356 Henriettea fascicularis 342; mac- fadyenii 342; membranifolia 342; triflora 342 Henriettella fascicularis 342; macfadyenii 342; membranifolia 342 Hermann F. J., An Eastern Vari- ety of Carex fissa ( Multiflorae) 198; The Status of Carex in- comperta Bickn 361 Herpestris rotundifolia 351 Herpetica alata 832 Heteromeris argentea 305; cana- densis 201, 255; carolinianum 212; chihuahuense 296; cymosa 288; glomerata 298; major 255, 269; mexicana 298, 300; Mi- chauxii 255; polifolia 301; Pringlei 294; ramuliflorum 255 Heteropteris laurifolia 336; pur- purea 336; wydleriana 336 Heterotheca bartlettii 218; lati- folia 87 fig., 181; psammophila 87 fig.; s. str. from Chrysopsis ( Compositae-Astereae), A Sec- ond Character Distinguishing Rhodora [Vol. 67 86; subaxillaris 87 fig. Hibiscus tiliaceus 340 Hierochloe alpina 380 Hieracium aurantiacum 197 Hippocratea caribaea 388 Hippurus vulgaris 196 Hodgdon, A. R., Mountain Flow- ers of New England (Review) 84 Holcus halepensis 318; sorghum 318 Homalium leiogynum 341; pleian- drum 341; racemosum 341 Homalocenchrus hexandra 319; monandrus 319 Horanthes arenaria 311; poda- nisia 302; tripetala 312 Hormidium flaccidum 246 pl., 251; tripterum 826 Hornemannia racemosa 344 Hufelandia pendula 330 Humulus lupulus 50, 51 pl, 52 Hybanthus linearifolius 341; por- toricensis 341 Hybosperma spirosum 339 Hydora lithuanica 171 Hydrilla verticillata 167, 170, 171; var. crispa 168 Hydrocotyle bonariensis ` 344; verticillata, var. triradiata 844 Hydromistria stolonifera 317 Hymenocallis declinata 324 Hymenodium crinitum 357 Hymenophyllum ` axillare 360; ciliatum 360; elegantulum, var. petiolatum 360; hirsutum 360; lanatum 360; sieberi 360; un- dulatum 360 Hypericum hypericoides 341 Hypoxis wrightii 324 Hyptis americana 350; escobilla 350; scoparia 351; spicigera 350 Ibatia maritima 346 Ibidium tortile 325 Icacorea glauciflora 344; guada- 1965] lupensis 344; luquillensis 344 Icthyomethia piscipula 334 Ilex cookii 338; sideroxyloides, var. occidentalis 338; verti- cillata 402 Ilysanthes dubia 351 Inga fagifolia 330; inga 331; laurina 331; vera 381 Inula divaricata 231 Ionidium linearifolium 341; por- toricensis 341 Ionoxalis intermedia 335; tiana 3385 Ipomoea acuminata 347; aegyptia 347; alba 347; amnicola 200; angustifolia 347; batatas 347; cathartica 347; (Convolvula- ceae), A South American Waif in Missouri 200; dissecta 347; hederifolia 847; heptaphylla 200, 347; kentrocarpa 347; och- roleuca 347; palustris 347; pes- caprae, var. brasiliensis 347; polyanthes 347; pulchella 200, 347; purpurea 847; quamoclit 347; quinquefolia 347; repanda 347; rubra 347; setifera 347; solanifolia 347; spinillus 347; stendelii 347; tricolor 347; tuba 347; violacea 347; Wrightii 200 Iresine celosia 329; diffusa 329 iris psuedacorus 196 Isandrina emarginata 332 Isnardia palustris 83, 348 Isopappus divaricatus 231; var. hookerianus 235; var. hirtellus 229; hookerianus 235; validus 233 Isotoma longiflora 354 Isotria medeoloides 394, 396 fig. Ixidium wrightii 328 mar- Jacquinia stenophylla 344 Jacquiniella miserrima 326; ter- etifolia 326 Jacquemontia canescens 347; ob- Index to Volume 67 427 cordata 847; subsalina 347; tamnifolia 347 Jambos jambos 343 Jambosa jambos 348 Jatropha curcas 337; dioica, var. graminea 194; gossypifolia 387; hernandiaefolia 337; in- tegerrima 337; multifida 337 Juncus bulbosus 377; militaris 376; pelocarpus 376; trifidus 380 Juniperus silicicola 312 Jussiaea angustifolia 343; erecta 343; leptocarpa 181, 343; per- uviana 343; repens 347 Justicia comata 852; pectoralis 352; verticilaris 352 Kalmia angustifolia 410 Kawano, Shoichi and Frank S. Crosswhite, Chromosome Num- bers in Penstemom (Schrophu- lariaceae) I: New Mitotic Counts 187 Kentucky, Flowering Plants New to or Rare in 180 King, R. M., A New Species of Senecio from Costa Rica 239 Kingsbury, John M. and Grace D. Lanciani, Soil Algae of Forty Ponds Under Construction at Ithaca, New York 242 Kobresia simpliciuscula 381 Kyllinga brevifolia 320; odorata 320; peruviana 320; pumila 320; pungens 320 Lactuca intybacea 354 Laetia procera 841 Lagarosiphon major 168 Lagascea mollis 355 Lagenaria siceraria 354 Lagenocarpus guianensis portoricensis 322 Lagenophora andina 40, 41; (Com- positae) from Guatemala, A 822; 428 New Species of 36; cuchu- matanica 36, 38, 39 pl., 40, 41; mauiensis 40; panamensis 40 Lagerstroemia speciosa 342 Lakela, Olga, The Extended Dis- tribution of Eragrostis tracyi from Sanibel Island, Florida 312 Lanciani, Grace D. and John M. Kingsbury, Soil Algae of Forty Ponds Under Construction at Ithaca, New York 242 Languas speciosa 324 Lantana aculeata 349; arida 349; camara, var. aculeata 349; var. mista 349; var. mutabilis 349; f. parvifolia 349; var. sanguinea 349; fucata, var. antillana 349; insularis 349; involucrata, var. odorata 349; montevidensis 349; reticulata 849; strigosus 349 Laplacea portoricensis 340 Larix laricina 195 Lasiacis patentiflora 318 Lasianthus lanceolatus 353; mor- alesii 353 Lathyrus japonicus, var. glaber 89; venosus 394, 395 fig. Laugeria resinosa 853 Laurentia longiflora 354 Laurocerasus myrtifolia 331; oc- cidentalis 331 Lebretonia paniculata 340 Lechea major 64, 255, 258, 269, 272; tripetala 312 Leersia hexandra 319; monandra 819 Lemna minor 195 Lepidagathis alopecuroidea 352 Leptilon bonariense 355; chinense 355; linifolium 355; pusillum 355 Leptochilus alienus 358; clado- rhizans 358; guianensis 358; nicotianaefolius 358; perga- Rhodora [Vol. 67 mentaceus 358 Leptochloa fascicularis 319; fili- formis 319; panicea 319 Leucaena glauca 332; leucoce- phala 332 Libidibia coriaria 333 Licaria brittoniana 330; salici- folia 330; triandra 330 Limnobium stoloniferum 317 Limonium Nashii 89 Lindernia crustacea 351; diffusa 350; dubia 351 Linociera axilliflora 345; caribaea 345; domimgensis 345; hold- ridgii 345; ligustrina 345 Linum catharticum 196; imbri- catum 192; trisepalum 278 Liogier, Brother Alain, Nomen- clatural Changes and Additions to Britton and Wilson’s “Flora of Porto Rico and the Virgin Islands” 315 Lipocarpha Sellowiana 98 Lippia helleri 349; micromera, var. helleri 349; nodiflora 349; reptans 349; stoechadifolia 349 Lobaria scrobiculata 380 Lobelia assurgens, var. portori- censis 354; portoricensis 354; rotundifolia 354 Loiseleuria procumbens 380 Lomariopsis amydrophlebia 358; kunzeana 358; sorbifolia 358 Lomograma guianensis 358 Lonchitis hirsuta 357 Lonchocarpus latifolius 334; pen- taphyllus 834 Lophidium poeppigianum 356 Lophosoria quadripinnata 356 Lotus corniculatus 196 Luchia Berteroniana 170 Lucuma multiflora 345 Lucya tetrandra 353 Ludwigia erecta 343; leptocarpa 343; natans 83; octovalvis 343; (Onagraceae), An Earlier 1965] Name for 83; palustris 83, 343; peploides 343; peruviana 348; repens 83, 84 Lunania buchii 341 Luzula spicata 382 Lychnis alpina 382 Lycium americanum 351; tweedi- anum, var. chrysocarpum 351 Lycopodium annotinum, var. pun- gens 380; clavatum 402; var. megastachyon, f. furcatum 195; complanatum 402; inundatum 376; selago 380 Lycopersicon esculentum 50 Lycopersium esculentum lycopersicum 351 Lycopodium sintenisii 860 Lyngbya limnetica 252; marten- siana 252; versicolor 252 Lyonia rubiginosa 344; stahlii 344 196; Maba sintenisii 345 Machaerium lunatum 334 Macmillamia repens 351 Macroptilium lathyroides 335 Magnolia tripetala 394, 395 fig. Malache scabra 340 Mallotonia gnaphalodes 348 Malpighia angustifolia 336; lin- earis 336; shaferi 336 Malvastrum americanum Spicatum 339 Malvaviscus arboreus 340; mal- viscus 340 Mammillaria nivosa 342 Manihot manihot 337; ultissima 337 Manilkara balata 345; bidentata 345; duplicata 345; emarginata 345; jaimiqui 345; nitida 345; pleeana 345; zapota 344 Marilaunidium jamaicense 848 Mariscus jamaicensis 322 Martiusia laurifolia 334; ginosa 334 Massachusetts Fern in Wisconsin, 339; rubi- Index to Volume 67 429 Discovery of the 399 Mastichodendron ` foetidissimum 345 Matayba apetala 339; oppositi- folia 838 Matelea maritima 346; sintenisii 347; variifolia 346 Matricaria inodora 230 Mayepea axilliflora 345; caribaea 345; domingensis 345 Maxillaria coccinea 327; conferta 327 Maximiliania vitifolia 341 Mecardonia dianthera 351; pro- cumbens 351 Medeola virginiana 372 Meibomia adscendens 334; affinis 334; axillaris 334; barbata 334; cubensis 334; mollis 334; pro- cumbens 334; purpurea 384; scorpiurus 334; sintenisii 334; supina 384; umbrosa 334; wyd- leriana 334 Melinis minutiflora 320 Melocactus intortus 341 Melicocca bijugata 338 Melicoccus bijugatus 388 Melochia pyramidata 340; tomen- tosa 340; villosa 340 Melanthera calcicola 355; cane- scens 355; confusa 355; mon- tana 355; nivea 855 Merremia aegyptia 347; dissecta 347; quinquefolia 347; triden- tata, ssp angustifolia 347; tuberosa 348; umbellata 347 Mesadenus lucayanus 325 Metastelma anegadense 846; de- caisneanum 346; decipiens 346; ephedroides 346; grisebachia- num 346; lineare 346; monense 345; parviflorum 346 Metopium linnaei 338; toxiferum 338 Miconia affinis 342; microcarpa 842; mirabilis 342; ottoschulzii 430 342; serrulata 342 Micranthemum umbrosum 851 Micropholis chrysophylloides 344; curvatus 344 Microspora stagnorum 251 Mikania congesta 355; micrantha, var. congesta 355 Mimosa pigra 382 Misantheca triandra 330 Mitchella repens 377 Mitella nuda 92 Mitracarpus hirtus 854; villosus 354 Moluchia pyramidata 340; tomen- tosa 340 Momisia iguanaea 327 Moneses uniflora 196 Monotropsis odorata 394, 396 fig. Moringa moringa 331; oleifera 331 Morongia portoricensis 832 Mucuna pruriens 335 Mulcahy, David L., Interpretation of Crossing Diagrams 146 Murraya exotica 336 Mylia anomala 372 Myrcia berberis 343; citrifolia 342; fallax 343; leptoclada 343 Myrcianthes fragrans 343 Myrciaria floribunda 343 Myrica holdridgeana 327; pensyl- vanica 377 Myriophyllum tenellum 196, 376; verticillatum, var. pectinatum 196 Myroxylon buxifolium 341; pachy- phyllum 341; schwaneckeanum 341 Nageia coriacea 356 Nama jamaicensis 348 Nardus stricta 377 Nazia aliena 318 Nemochloa millegrana 124 Nemopanthus mucronata 373, 402 Neomammillaria nivosa 342 Neovriesia macrostachya 323 Rhodora [Vol. 67 New Brunswick, New Records of Arctic Species in Southeastern 92 New England, Mountain Flowers of (Review) 84 Newfoundland, The Distribution of Northern and Southern Ele- ments in the Flora of 363 New Hampshire, Notes on the Flora of Coös County 195 New York, Soil Algae of Forty Ponds Under Construction at Ithaca 242 Nidema ottonis 326 Nocco mollis 855 Nodularia harveyana 252; spumi- gena 252 Nostoc commune 252, 253; linckia 252; muscorum 252; paludosum 252; sphaericum 252; verruco- sum 252 Notholaena trichomanoides 357 Nymphaea ampla 330; amazonum 330; jamesoniana 330; odorata 330, 373; pulchella 330; rudge- ana 320 Nymphoides lacunosa 376 Octadesmia montana 326 Odontoschisma sphagni 372 Oklahoma, Additions to the Vas- cular Flora of 191 Oldenlandia callitrichoides 353; corymbosa 352; herbacea 353; lancifolia 853 Oncidium intermedium 327; lem- onianum 327 Oocystis crassa 251; parva 251 Operculina tuberosa 348 Ophioglossum palmatum 356 Oplismenus setarius 318 Opuntia Dillenii 313 Ornithidium coccineum 327; con- fertum 327 Orthiopteris domingensis 359; in- equale 358 Oscillatoria acutissima 252; agar- 1966] dhii 252, 258; amoena 252; am- phibia 252; angusta 252, 253; angustissima 252; limnetica 252, 253; rubescens 252; splendida 252, 253; subbrevis 252; tenuis 252; terebriformis 252 Osmia borinquense 355; corym- bosa 354; geraniifolia 355; ivae- folia 355; odorata 354; sinuata 355 Osmunda cinnamomea 402; rega- lis 373, 402 Ouratea ilicifolia 340; striata 340 Oxalis barrelieri 335; corniculata 146, 147 fig., 148 fig., 149 fig., 150, 151, 335; corymbosa 355; dillenii 146, 147 fig., 148 fig., 149 fig.; grandis 146, 147 fig., 148 fig. 149 fig.; intermedia 335; montana 377, 387; priceae 146, 147 fig., 148 fig., 149 fig., 150, 151; stricta 146, 147 fig., 148 fig., 149 fig., 151; suksdorfii 146, 147 fig., 148 fig., 149 fig. Oxytropis foliolosa 381 Pachira aquatica 340; insignis 340 Pachyrrhizus erosus 335 Palicourea brevithyrsa 353; crocea 853 Palmella mucosa 251 Panax trifolius 402 Pancratium declinatum 324 Panicum angustifolium 818; bar- binode 318; capillare 318; fusi- forme 318; muticum 318; stri- gosum 318; utowanaeum 318 Pappophorum alopecuroideum 319; pappiferum 319 Paralabatia portoricensis 345 Pariti tiliaceus 340 Parnassia asarifolia 394, 396 fig. Parosela domingensis 333 Parsonsia micrantha 342; sonia 342 Paspalum blodgettii 318; boscia- par- Index to Volume 68 431 num 318; glabrum 318; laxum 318; leoninum 318; melanosper- mum 317; molle 317; poirettii 317; portoricensis 318; rupestre 318; saugetii 318; simpsoni 318 Passiflora anadenia 841; edulis 341; murucuja 341; pallida 341; suberosa 341 Pavonia fruticosa 339; paniculata 840; scabra 340 Pectis floribunda 355 Pedicularis sylvatica 377 Pedilanthus angustifolius 337; latifolius 338; padifolius 337; tithymaloides, var. angustifolius 337; var. padifolius 337; var. parasiticus 337 Peiranisia polyphylla 332 Pelexia adnata 325 Peltapteris peltata 356 Peltophorum ferrugineum 323; inerme 3833 Pennisetum purpureum 319 Penstemon arkansanus 187, 188 fig.; calycosus 188 fig. 189; digitalis 188 fig., 189; gracilis, ssp gracilis 189; ssp wisconsin- ensis 188 fig. 189; grandiflorus 189; harbourii 188 fig., 189; lax- iflorus 188 fig. 189; multicaulis 187; pachyphyllus 188 fig.; ssp pachyphyllus 189; pallidus 188 fig., 189; (Scrophulariaceae) I: New Mitotic Counts, Chromo- some Numbers in 187; teucri- oides 188 fig. 190; tubaeflorus 188 fig. 190; wherryi 187 Pentarhaphia albiflora 352 Peperomia dendrophila 32: megapoda 327; myrtillus 327; rhombea 327; sintenisii 327; tenella 327 Pepo moschata 354 Pereskia aculeata 341; pereskia 341 Persea americana 330; persea 432 330; portoricensis 831; ur- baniana 331 Persicaria acuminata 327; por- toricensis 328; punctata 328; segetum 828 Petesioides pendulum 344; yun- quense 344 Petrea kohautiana 350 Petroselinum crispum 344 Phacelia gilioides 42, 43, 44 fig., 45, 46 fig., 47; hirsuta 42, 43, 44 fig., 45, 46 fig., 47; Com- plex, Genotypic Variation in the 42; patuliflora 139, 140 fig., 141, 142 fig., 148, 144; var. patuli- flora 139; sericea 47; stricti- flora 139, 140 fig, 141, 142 fig., 143, 144; Complex, Geno- typic Variation in the 139; var. lundelliana 139 Phaeosphaerion persicariaefolius 323 Phaseolus coccineus 385; lathy- roides 335; multiflorus 335; peduncularis 335; schotti 835; trichocarpus 335 Phialanthus grandifolius 358; myrtilloides 353 Philodendron hederaceum 322; nechodomi 323; oxycardium 323; scandens 323 Philotria angustifolia 170; cana- densis 12, 28, 170; densa 157, 170; granatensis 170; guyan- nensis 170; iowensis 12, 170; linearis 29, 31, 170; minor 6, 169, 170; Nuttallii 6, 11, 12, 170; occidentalis 6, 170; orinocensis 170; Planchonii 12, 170 Phlebotaenia corvellii 336 Phleum pratense, f. viviparum 195 Phoradendron berterianum 328; dichotomum 328 Phoebe elongata 331; montana 331 Phormidium minnesotense 252, 253 Rhodora [Vol. 68 Phragmites communis 319; phrag- mites 319 Phthirusa bicolor 328; caribaea 328; purpurea 328; sintenisii 828 Phyla nodiflora 349; var. reptans 349; scaberrima 349; stoechadi- folia 349; strigulosa. f. parvi- folia 349 Phyllanthus acidus 337; diffusus 337; epiphyllanthus 337; ju- glandifolius 337; lathyroides 337; niruri 337; pentaphyllum, ssp. pentaphyllum 337; ssp. polycladus 337; | polycladus 337; stipulatus 337; urinaria 837 Phyllodoce caerulea 380 Physalis hederaefolia, var. heder- aefolia 194 Physurus hirtellus 325; planta- gineus 325 Piaropus crassipes 323; diversi- folius 323 Picea mariana 92; rubens 92 Pierasma antillana 336; excelsa 336 Pike, R. B., Common Trees of Puerto Rico and the Virgin Islands (Review) 93 Pilosocereus royeni 341 Pimenta dioica 343; pimenta 343; racemosa 342; var. grisea 342 Pinus resinosa 374, 386, 387, 389; rudis 37; strobus 402 Piper citrifolium 327; glabrescens 327; hispidum 327; peltatum 327; scabrum 327; treleaseanum 327; umbellatum 327; wydleria- num 327 Piptadenia peregrina 332 Pipptocoma antillana 354; rufes- cens 354 Piscidia piscipula 334 Pithecellobium unguis-cati 331 Pityrogramme trifoliata 357 1966] Platypus atus 327 Pleurostachys minuscula 124; puberula, var. Buchtienii 124 Pleurothallis appendiculata 326; broadwayi 326; foliata 326; longissima 326; Plinia dusii 348 Pluchea carolinensis 355; odorata 355 Plumeria krugii 346; obtusa 345; portoricensis 346 Podocarpus coriaceus 356 Pogonia macrophylla 325; ophio- glossoides 373 Pohl, Richard A., Dissecting Equipment and Materials for the Study of Minute Plant Structures 95 Poinciana pulcherrima 333 Polygala cowellii 336; penaea 336; portoricensis 336 Polygonum acuminatum 328; den- siflorum 328; punctatum 328; segetum 328; viviparum 381 Polypodium asplenifolium 357; duale 357; hessii 357; jubae- forme 357; latum 357; mollis- simum 357; myosuroides 357; phyllitidis, var. latum 9857; taenifolium 357; taxifolium 357; trifurcatum 357 Polystachya cerea 325; extinctoria 325; minor 325; minuta 325 Polystichopsis chaerophylloides 359 Polystichum ` adiantiforme 358; echinatum 358; triangulum 358 Polytaenium cajanense 357 Porophyllum porophyllum 355; ruderale 355 Porto Rico and the Virgin Is- lands", Nomenclatural Changes and Additions to Britton and Wilson's “Flora of 315 Portulaca phaeosperma 330; polio- sperma 330; rubricaulis 330; Index to Volume 68 433 teretifolia 330 Potamogeton | epihydrus 317; fluitans 817; insulanus 317; natans 195; nodosus 317; oake- sianus 376; oblongifolium 84; rotundifolium 84 Potamogeton natans 195; ob- longifolium 84; rotundifolium 84 Potentilla fruticosa 92; nivea 381 Pothomorphe peltata 327; umbel- lata 327 Pouteria dictyoneura, ssp fuer- tesii 345; hotteana 345; mam- mosa 345; multiflora 345 Prestoea montana 322 Prestonia agglutinata 846 Primula egaliksensis 381 Pristimera caribaea 338 Prosopis limensis 332 Protoderma viride 251 [Protosiphon botryoides 251 Prunus myrtifolia 331; occiden- talis 331; virginiana 373 Pseudanamomis umbellulifera 843 Pseudoelephantopus spicatus 354 Psidium amplexicaule 342; cat- tleyanum 342 Psilocarya nitens 322; portoricen- sis 322 Psilochilus macrophyllus 325 Psoralea esculenta 26 Psychotria microdon 353; pinu- laris 353 Ptychomeria portoricensis 325 Pteridium aquilinum, var. arach- noideum 357; var. caudatum 357; arachnoideum 357; cauda- tum 357 Pueraria phaseoloides 335 Puerto Rico and the Virgin Islands, Common Trees of (Re- view) 93 Pyrethrum bonplandianum 182 Pyrola grandiflora 381 Pyrus americana 402 434 Quamoclit coccinea 347; quamo- clit 847 Quercus alba 402; borealis 386; rubra 408 Radicula palustris 331; portori- censis 331; portoricensis 331 Randia aculeata 313, 353; mitis 353 Ranunculis acris, var. latisectus 196 Rapanea guianensis 318 Rauwolfia lamarckii 346; nitida 346; tetraphylla 346; viridis 346 Raven, Peter H., An Earlier Name for Ludwigia natans (Onagraceae) 83 Remirea maritima 822; culata 322 Renealmia antillarum 324; matica 324; exaltata 324 Rhacoma crossopetalum 338 Rhamnus volubilis 84 Rheedia acuminata 341; icensis 341 Rhipidopteris peltata 357 Rhododendron canadense lapponicum 365, 381 Rhoeo discolor 323; spathacea 323 Rhynchosia minima 335; phaseo- loides 335; reticulata 335 Rhynchospora alba 321; andina 116, 119; aristata 115, 117; var. latilaminata 117, 118; bahamen- sis 322; berterii 321; blauneri 321; boliviensis 116, 120; borin- quensis 322; brachychaeta 321; bruneri 321; capillacea 192; ciliata 124; confinis 115, 116, 123; corymbosa 115, 118; cy- mosa 322; cyperoides 115; 119; distans 322; emaciata 116, 122; exaltata 116, 119; var. cephalo- phora 119; var. ovalis 119; fascicularis 322; filiformis 321; glauca 115, 121, 322; globosa pedun- aro- portor- 3738; Rhodora [Vol. 68 115, 120; var. Loefgrenii 120; globularis 322; hirta 115, 120; hispidula 321; intermixa 821; jamaicensis 321; lindeniana, var. bahamensis 322; luquillensis 321 luzuliformi 116, 122; macro- chaeta 115, 116; var. colum- biensis, f. condensata 117; var. Ruiziana 117; var. quinque- spicata 117; Marisculus 115, 121, 322, microcarpa 322; mille- grana 115, 124; nervosa 115, 123, 321; var. Jelskiana 124; nitens 821; odorata 322; podo- sperma 321; polyphylla 115, 118, 321; var. laxa 118; var. longispieulosa 118; puberula 115, 124; pusilla 321; racemosa 322; radicans, 115, 321; robusta 116, 121; rugosa 322; Schei- deana 115, 117; setacea 116, 123; stellata 321; tenerrima 123; tenuis 116, 122; var. ema- ciata 122; umbraticola, var. Kuntzei 115, 116; trichodes 321; velutina, var. Sellowiana 116, 122 Richardia brasiliensis 354 Ricinella ricinella 337 Riedlea hirsuta 340 Rivea companulata 848 Roberts, Patricia R., New Rec- ords of Arctic Species in South- eastern New Brunswick 92 Rorippa nasturtium-aquaticum 331; palustris 331; portoricen- sis 331 Rubus acaulis 382; arcticus 382; chamaemorus 382, 408; hispi- dus 402; pensilvanicus 196; probus 331 Ruellia twediana 852 Rumohra adiantiformis 358 Ruppia anomala 317; maritima 317 Rytilix granularis 317 1966] Sabatia calycina 304 Saccoloma domingense 359; in- equale 360 Sagotia triflora 334 St. John, Harold, Monograph of the Genus Elodea: Part 4 and Summary 1, 155 Salix cordifolia 381; glaucophyl- loides 375; herbacea 380; myr- tillifolia 92, 93; reticulata 381; uva-ursi 382; vestita 381 Salvia micrantha 850; nusella 350 Sambucus pubens 373 Sanguinaria canadensis 372, 386 Sanicula marilandica 394, 395 fig. Sansevieria metalica 324 Sapota achras 344 Saracha antillana 351 Sarcomphalus domingensis 339; reticularis 389; taylori 339 Sarcostemma clausum 346 Sargent, F. H., Eulophia ecristata and Epidendrum conopseum in Mississippi 48 Saxifraga aizoides 382; aizoon 381; cespitosa 381; oppositifolia 381 Scaphyglottis modesta 325 Scenedesmus hystrix 252; obli- quus 252; quadricauda 252, 253 Schizachyrium brevifolium 317; domingense 317; gracile 317; semiberbis 317 Schizaea pennula 356; pigiana 356; pusilla 376 Schizothrix calcicola 252 Schlegelia brachyantha 352; por- toricensis 352 Schobera angiosperma 348 Schradera vahlii 353 Schrankia portoricensis 332 Scirpus acaulis 100, 102; acutus 375; americanus 100; var. poly- phyllus 102; ssp. polyphyllus 102; asper 100, 102; ataca- mensis 100, 101, 102; boli- poep- Index to Volume 68 435 viensis 100, 101; californicus 100, 103; cernuus 100, 101; deserticola 100, 103; fuirena 321; hudsonianus 382; inunda- tus 100, 101; lacustris, ssp. validus 321; macrantha 99; maritimus 374, 377; rigidus 100, 102; riparius 103; subter- minalis 376; validus 321, 374 Scleria amphigaea 125, 126; arun- dinacea 126, 130, 182; bracteata 126, 131; f. simplicior 131; cas- tanea 125, 127; ciliata 322; composita 126, 128; cyperi- noides 126, 129; eggersiana 322; grisebachii 322; hirtella 125, 126; lagoénsis 126, 181; latifolia 126, 130, 132; lepto- stachya 125, 126, 127; lith- osperma, 125, 127; macrophylla 125, 128; melaleuca 126, 132; microcarpa 125, 128; mitis 125, 129; obtusa 126, 129; panicoides 126, 180; paludosa 128; pleo- stachya 127; pterota 126, 132; reflexa 133; secans 126, 133; setacea 126, 131, 321; steven- siana 322; tenacissima 126, 131; triglomerata 322; vaginata 126, 132 Scolosanthus densiflorus 853; grandiflorus 353; multiflorus 353; versicolor 353 Scosanthus versicolor 353 Scudday, James F., Chamaesara- cha villosa New to Texas 194 Scutellaria lateriflora 373 Sebastiania corniculata 337 Sebesten rickseckeri 347; sebes- tena 348 Securidaca diversifolia 336; vir- gata 336; volubilis 334 Selaginella albonitens 860; ar- mata 360; flabellata 361; plagi- ochila 360; plumosa 860; por- toricensis 360; selaginoides 436 382; sintenisii 361; stipitata 361; stolonifera 360; subcau- lescens 361; substipitata 360; tenella 360 Senecio aureus 873; costaricensis 239, 240 pl, 241; from Costa Rica, A New Species of 239; townsendii 241 Senna obovata 832 Serpicula canadensis 12, 171; oc- cidentalis 6, 167, 169, 170, 171; verticillata, f. 8, var. angusti- folia 6, 171 Sesamum indicum 352; orientale 352 Sesban emerus 333; sericea 333; sesban 833 Sesbania bispinosa 333; emerus 333; grandiflora 333; sesban 833; sericea 333; spinosa 333 Sesuvium portulacastrum 313 Setaria barbata 319; Faberi 195; geniculata 319; leiophylla 319; lutescens 319; magna 319; pradana 319; rariflora 319; setosa 319; tenacissima 819; utowanaea 318; viridis 195; vulpiseta 319 Setiscapella pusilla 352; subulata 351 Shepherdia canadensis 92 Sherff, Earl Edward, Note on Varieties of Bidens connata and a Hybrid with B. cernua 59 Shinners, Lloyd H., Ipomoea amnicola (Convolvulaceae), A South American Waif in Mis- souri 200 Sibbaldia procumbens 380 Sida acuta 339; alba 339; carpini- folia 339; erecta 339; salviae- volia 389 Sideroxylon foetidissimum 345; portoricensis 345 Sieglingia decumbens 377 Silene acaulis 380; pumilio 196 Rhodora [Vol. 68 Sinapis arvensis 331 Siphonanthus indicus 350 Siphoneugenia densiflora 343 Sisymbrium nasturtium-aquati- cum 331 Sloanea amygdalina 339 Smilacina racemosa 375, 377, 386 Smilax domingensis 324; lanceo- lata 324 Smith, Edwin B., Taxonomy of Haplopappus, Section Isopap- pus (Compositae) 217 Solandra grandiflora 351 Solanum antillarum 351; campe- chiense 351; caribaeum 351; cilatum 851; elaegnifolium 351; erianthum 351; ficifolium 351; guanicense 351; pterocaulon 351; torvum 351; verbasci- folium 351 Solbrig, Otto T., The Typification of Xanthocephalum (Compos- itae) 182 Solidago multiradiata 92, 93, 381; rugosa 373; sempervirens 89 Sorghastrum parviflorum 318; setosum 318 Sorghum halpense 818; sacchara- tum 318 Spartina pectinata 374 Spathiger ramosus 326; rigidus 326 Spermacoce confusa 354; riparia 354; tenuior 354 Sphaerellopsis fluviatilis 252 Sphagnum fuscum 372; lindbergii 380; macrophyllum 377, 385; molle 376, 385; pylaesii 376; rubellum 372; strictum 376 Spiranthes adnata 325; anabilis 325; cranichoides 325; elata 325; fawcettii 325; lucayana 325; orchioides 325; speciosa 325; tortilis 325 Spirulina major 252, 253 Sporobolus argutus 819; bertero- 1966] anus 319; cryptandrus 180; domingensis 319; muralis 319; poiretii 319; pyramidatus 319; tenuissimus 319; virginicus 313 Stachytarpheta cayennensis 850; var. albiflora 350; jamaicensis 350; strigosa 350; urticaefolia 350; X hybrida 350 Stegnosperma cubense 329 Stellaria antillana 330 Stenandrium tuberosum 852 Stenochlaera amydrophlebia 358; kunzeana 358; sorbifolia 358 Stenophyllus curvassavicus 321; portoricensis 321; vestitus 321 Stenorrhyncus lanceolatus 325; speciosa 325 Stenostomum acutatum 853; cori- aceum 353; lucidum 353; obtusi- folium 353; portoricense 353; sintenisii 853 Stethoma comata 352; pectoralis 352; verticillaris 352 Stewartia ovata 394, 396 fig. Stichococcus bacillaris 251 Stictocardia tiliaefolia 348 Stigmaphyllon diversifolium 336; ledifolium 336; lingulatum 856; periplocifolium 336 Stizolobium pruritum 335 Stomoisia juncea 352 Struthiopteris exaltata 358; lin- eata 858; polypodioides 357; underwoodiana 358 Suaeda fruticosa 329 Succisa pratensis 230 Swartzia grandiflora 351 Synechococcus aeruginosa 252 Synechocystis aquatilis 252, 253; crassa 252; sallensis 252 Syntherisma argillacea 318; chin- ensis 318; digitata 318; fili- formis 318; ischaemum 818; sanguinalis 318 Syringodium filiforme 317 Tabebuia heterophylla 352; pal- Index to Volume 68 437 lida 352 Tabernaemontana coronaria 346; divaricata 346 Taeniostema micranthum 65, 201, 298, 300 Tamonea guianensis 342; rophylla 342 Taonabo heptasepala 340; luquil- lensis 340; pachyphylla 340; peduncularis 340; stahlii 340; subsessilis 340 Taraxacum officinale 89 Taylor, R. John and Constance, Additions to the Vascular Flora of Oklahoma 191 Tectaria incisa 359; martinicen- sis 859 Tectona grandis 350 Teliostachya alopecuroidea 852 Tephrosia candida 333; cinerea 333; senna 333 Tepion alatum 355 Terebraria resinosa 353 Ternstroemia heptasepala 840; luquillensis 340; peduncularis 340; stahlii 340; subsessilis 340 Tetraclea Coulteri 194 Tetragamestus modestus 325 Tetragonia expansa 329; tetra- gonioides 329 Tetrapteris buxifolia 385; citri- folia 335; inaequalis 335 Thalictrum alpinum 382 Thecophyllum sintenisii 323 Thelypteris angustifolia — 359; balbisii 358; brittonae 359; decussata 358; deltoidea 358; dentata 359; diplazioides 358; germaniana 358; gongylodes 358; guadalupensis 359; has- tata 359; leptocladia 359; megalodus 359; nephrodioides 359; normalis 359; novebora- mac- censis 402, 403; oligophylla 359; opposita 358; palustris 399, 402, 403; patens 359; piedrensis 358; poiteana 359; 438 reptans 359; reticulata 359; sancta 358; var. portoricensis 358; sclerophylla 359; serra 359; serrata 359; simulata 399; 400PI., 401 fig., 402, 403; tetra- gona 359 Thevetia peruviana 346 Thibaudia krugii 344 Thouinia portoricensis 338; stri- ata 338 Thrinax ponceana 322 Thryallis glauca 836 Thyana portoricensis 338; striata 338 Thyella tamnifolia 347 Thyphalea fruticosa 339 Tiarella cordifolia 50 Tiaridium indicum 348 Tiffney, W. N. and W. N. Tiffney, Jr. Tulostoma brumale Pers., A New Record for New Eng- land 89 Tigridia purpurea 192 Tillandsia aloifolia 823; flexuosa 323; sublaxa 323; usneoides 323; valenzuelana 323 Tofieldia pusilla 382 Tontanea herbacea 853 Torralbasia cuneifolia 338 Tournefortia gnaphalodes 348; laurifolia 348; maculata 348; microphylla 348; peruviana 348; volubilis 348 Tradescantia elongata 823; geni- culata 323 Tragus beteronianus 818 Tribulus terrestris 181 Tricera citrifolia 338; vahlii 338 Trichachne eggersii 318; insularis 318 Trichasterophyllum hyssopifolium 298 Trichomanes angustifrons 360; kapplerianum 360; ovale 360; punctatum, ssp. sphenoides 360; pusillum 360; robustum 360; Rhodora [Vol. 68 sphenoides 360 Trientalis borealis 402 Trillium cernuum 373 Trimezia martinicensis 324 Tripogandra elongata 823 Trisetum spicatum 382 Trismeria trifoliata 357 Triumfetta bartramia 339; bo- gotensis 339; excisa 339; his- pida 389 Trochisia reticularis 251 Tsuga canadensis 92 Tulostoma brumale 89, 90; A New Record for New England 89; imbricatum 90; mammosum 90; pedunculatum 90; simulans 90; squamosum 90 Tupa portoricensis 354; robusta 354 Tynnanthus caryophylleus myrianthus 352 Typha angustifolia 317; domin- gensis 317; latifolia 374 352; Udora brasiliensis 171; canaden- sis 7, 12, 171; granatensis 171; guyannensis 171; lithuanica 171; occidentalis 21, 171; orino- censis 171; pomeranica 171; surinamensis 171; verticillata 171; var. minor 170, 171 Ulmus americana 386 Ulothrix belkae 247 pl., 248, 251; subtillissima 251 Uncinia hamata 133; 133 Urceolaria exotica 358 Urostachys sintenisii 360 Utricularia geminiscapa 376; in- flata, var. minor 197; juncea 352; pusilla 352; subulata 351 Uvularia sessilifolia 372 phleoides Vaccinium macrocarpon 373; uli- ginosum 380 Vachellia farnesiana 332 Valerianoides cayennense 350; Missouri Botanical 3 00 1753 1966] jamaicense 850; strigosum 350 Vallisneria americana 196 Valota eggersii 318; insularis 318 Vandellia diffusa 351 Vanilla dilloniana 325; eggersii 325 Varronia angustifolia 348; ba- hamensis 348; bellonis 348; corymbosa 848; globosa 348; lima 348; rupicola 348 Verbascum Blattaria 197 Verbena bonariensis 349; tenui- secta 349 Verbesina alata 355; 355 Vetiveria zizanioides 318 Viburnum cassinoides 373 Vigna luteola 335; peduncularis 335; repens 335 encelioides Vincetoxicum sintenisii 347; stephanotrichum 346; varii- folium 346 Viola incognita 402 Virgin Islands, Common Trees of Puerto Rico and (Review) 93; Nomenclatural Changes and Additions to Britton and Wil- son’s “Flora of Porto Rico and the 315 Virginia, The Mountain Element in the Flora of the Peninsula of 393 Vitex negundo, var. 350; parviflora 350 Vittaria filifolia 357; graminifolia 357 Volkameria aculeata 350 intermedia Vriesea macrostachya 323; sin- tenisii 823 Wilbur, Robert L. and H. S. Daoud, A Revision of the North American Species of Helian- themum (Cistaceae) 63, 201, 255 Wallenia pendula 344; yunquen- sis 344 Index to Volume 68 mim Waltheria americana 340; indica 340 Warszewiczella flabelliformis 327 Wisconsin, Discovery of the Mas- sachusetts Fern in 399 Wissadula contracta 339 Wittmachia lingulata 328 Wullschlaegelia aphylla 325 Xanthium chinense 354; stru- marium 354 Xanthocephalum bonplandianum 184; centauroides 182, 183 pl., 184; (Compositae), The Typi- fication of 182; humile 182 Xanthocoma humile 182, 184 Xanthosoma nigrium 323; viola- ceum 323 Xanthoxalis corniculata 335 Ximenia americana 328; ence- lioides 355 Xolisma rubiginosa 344; stahlii 344 Xylophylla epiphyllanthus 337 Xylosma buxifolium 341; pachy- phyllum 341; schaefferioides 341; schwanecheanum 341 Xyris carolinensis 323; judicai 323; montana 376 Young, Alvin and Alan A. Beetle, A Third Subspecies in the Artemisia tridentata Complex 405 Yucca aloifolia 313 Zephyranthes grandiflora 324; puertoricensis 324; rosea 324 Zingiber officinalis 324; zingiber 324 Zinnia peruviana 355 Zizania aquatica, var. anguisti- folia 195; interior 11 Ziziphus jujuba 339; mauritiana 339; reticulata 339; rignonii 339; taylori 339 Zornia diphylla 333; gemella 334; reticulata 333