Hodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ROLLA MILTON TRYON ) RADCLIFFE BARNES PIKE STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY ALFRED LINN BOGLE f Associate Editors RICHARD EDWIN WEAVER S Vol. 76 March, 1974 No. 805 CONTENTS: The Phytogeography and Systematics of Xanthisma tex- anum D. C. (Asteraceae); Proper Usage of Intraspecific Categories John C. Semple EMEND 1 A White-Flowered Form of Utricularia purpurea C. Barre Hellquist eet 19 On the Scientific Name of the Longleaf Pine BEE Ey Ward T R in 20 The Occurrence of Bald Cypress (Taxodium distichum (L.) Richard) in Suffolk County, Long Island, New York Edward J. Hehre and Peter M. Bolid Leeds: 25 The Nef England Botanical Club, Ine. Botanical Museum. Oxford St.. Cambridge, Mass. 02188 RHODORA. — A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. 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Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire 03824 Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to RHODORA, Botanical Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive the next number of RHODORA, changes of address must be received prior to the first day of March, June, September or December. Second Class Postage Paid at Boston, Mass. MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 76 March, 1974 No. 805 THE PHYTOGEOGRAPHY AND SYSTEMATICS OF XANTHISMA TEXANUM DC. (ASTERACEAE): PROPER USAGE OF INFRASPECIFIC CATEGORIES? JOHN C. SEMPLE? The monotypic genus Xanthisma DC. (Asteraceae) in- cludes sufficient variation in its single species X. texanum DC. to illustrate the three levels of taxonomically recog- nized infraspecific variation: subspecifie, varietal and for- mal. Proper usage of the three taxonomic ranks requires consideration of morphological and distributiona] data ac- cording to a consensus of international opinion. Although much has been published on the B-chromosome of the spe- cies (see Semple, 1972a), no modern systematic treatment of X. texanum has been published. Examination of some five hundred herbarium collections and experimental data indicate that a revision of Gray's 1852 treatment, which was based on only a few specimens, is necessary. A detailed diseussion of the distribution of the species as a whole and each of the infraspecific taxa is warranted for two reasons: first, to document the allopatric condition of the two subspecies, and second, to show that the distri- "The work was part of an NDEA Title IV Fellowship sponsored doctoral dissertation submitted to Washington University, St. Louis, Mo. Aug. 1972: Dr. Walter H. Lewis advisor. 2Current Address: Dept. of Biology, University of Missouri-St. Louis, St. Louis, Mo. 63121. 2 Rhodora [Vol. 76 bution of the species is not predictable from the general- ized vegetation distributions given by Gould (1962). The Edwards Plateau and its eroded portions are basic con- siderations in both cases. A taxonomic treatment consider- ing all aspects of the problem is given at the end of the discussion of distribution and morphological variation. All populations of a species ought to be interfertile to some degree. The validity of treating all the morphological forms of Xanthisma as a single species was tested by hy- bridization experiments involving greenhouse plants grown from fruiting material collected at wild populations. No barriers to fertilization were found and normal chromosome homologue pairing in meiotic prophase was observed in F, generation hybrids, which also set viable seed. Cross pol- lination was accomplished following the procedure of Smith & Parker (1971) and was facilitated by self-incompatibil- ity. Selfing could be induced, but yielded progeny of low fitness. Clearly, Gray's combining all taxa of Xanthisma into a single species is acceptable. DISTRIBUTION OF THE SPECIES Xanthisma texanum grows in a portion of the central southwest of the United States: Texas, Oklahoma and east- ern New Mexico. The known distribution of the species is illustrated in Figure 1 and is based on herbarium collections from GH, KSU, MO, NY, OKLA, SMU, TEX, TTC, US, WIS, and Arizona State University-Tempe.’ Collections of subsp. drummondii (T. & G.) Semple and subsp. texanum are rep- resented by dots and stars respectively; var. texanum and var. orientalis Semple of subsp. texanum are represented by encircled and unenclosed solid stars; putative hybrids between the two subspecies are represented by open stars. Treatment of Gray's (1852) varieties as subspecies, a term he did not use (Fernald, 1940), is based upon involucral bract morphology and the nearly allopatric distribution of the two taxa, while the distribution and bract morphology "The assistance of Frederick Utech and Timothy Love at the last two herbaria, respectively, is gratefully acknowledged. 1974] Xanthisma — Semple 3 Distribution of Xanthisma texanum DC ye subspeci(ic hybrids subsp drummondii @® subsp texanum var texanum * subsp texanum var. orientalis atc Figure 1. The distribution of Xanthisma texanum DC. Locations of collections have been determined from herbarium collections from eleven herbaria cited in the text. A key to the symbols used is given. Collections are plotted on a county outline map of Texas and Okla- homa. 4 Rhodora [Vol. 76 variation within subsp. texanum is properly treated at the varietal level. Variation in environmental factors accounts for the species distribution. Climatic factors determine the extent of the species’ range in the east, northwest and south. Too much rain in the east and too little in the north and west are the ap- parent limiting factors, while increasingly tropical condi- tions become limiting to the south (Visher, 1954). Local variation in geological factors accounts for the distribution within the range and secondarily along the margins of the range. The western margin of the eastern deciduous forest (Bray, 1906; Tharp, 1926; Braun, 1950; and Gould, 1962) forms the eastern margin of the range of X. texanum, which is a member of the ecotone flora sensu latu between the forests and grasslands. Reasons for the present loca- tion of the forest-prairie ecotone are not appropriate for discussion in this paper. A detailed treatment of geological influences on the spe- cies’ range is given by Semple (1972b) and summarized for each taxon under each appropriate heading. The Rio Grande Valley supports a noticeably distinct vegetation community (Foscue, 1932). Xanthisma texanum is excluded from the community, which apparently forms a barrier to southward expansion of the range of X. texanum, since no collections have yet been made in Mexico. Either the Rio Grand Valley community is an impassible barrier or no inhabitable locations exist in Mexico for X. texanum. DIAGNOSTIC MORPHOLOGY AND DISTRIBUTION OF THE TWO SUBSPECIES A subspecies is characterized by all members exhibiting a particular morphology distinct from other individuals in the species and by the allopatric distribution of these mem- bers from the rest of the species (Du Reiz, 1930; Clausen, 1941; van Steenis, 1957; Valentine & Lóve, 1958; and Ehrendorfer, 1968). Those morphologically intermediate individuals between the subspecies and the rest of the spe- 1974] Xanthisma — Semple 5 $ € Figure 2. Geographie distribution of involucral bract variation of Xanthisma texanum DC. Bracts from plants from 68 populations are illustrated in outline on a map of Texas and Oklahoma. Bracts 1-48 are from individuals of subsp. drummondii (T. & G.) Semple; bracts 52-68 are from individuals of subsp. texanwm; and bracts 49-51 are from putative hybrids between the two subspecies. 6 Rhodora [Vol. 76 cies only occur in populations geographically located be- tween the subspecies and the remainder of the species. Within X. texanum all members of subsp. drummondit (T. & G.) Semple are distinguished by involucral bracts that are lanceolate and taper to an acute apex. Subspecies texanum includes other members of the species and is char- acterized by involucral bracts that are depressed-ovate in shape and shorter than those of subsp. drummondii. Diagrammatic representations (Figure 2) of involucral bracts typical of those occurring in the middle of the series forming the involucre are superimposed on an outline map of Texas and Oklahoma and were taken from plants occur- ring in wild populations at the locations indicated. Bracts 1-48 show the similarity that occurs within subsp. drum- mondii (see Figure 3.C for a detailed illustration of a typi- cal involucral bract). Bracts 52-68 illustrate the range of variation in involucral bract shape found in subsp. texanum (Figures 3.A and 3.B), Bracts 49-51 illustrate involucral bracts from putative hybrids between the two subspecies and are the only bracts that are intermediate in shape. The allopatric distribution of the two subspecies is due to the presence of the Edwards Plateau and the Blacklands Prairies between the two taxa. Geologically and topo- graphically the Edwards Plateau is broken up into five regions (Fenneman, 1931; Oetking, 1959; and Raisz, 1957), although Gould (1962) included all the varied vegetation types within a single vegetation region. With a few excep- tions, in part due to man's influences, subsp. drummondii does not grow in areas overlying uneroded portions of the Edwards Plateau, since the upper strata limit the amount of water available to the shallow rooted plants. Depending upon the extent of erosion and the nature of the underlying bedrock, subsp. drwmmondii grows in varying degrees of commonness in the areas of the eastern portion of the plateau (colloquial sense) in the following manner: com- mon at the lower elevations of the Central Mining District, common throughout the Lampasas Cut Plains and rare on the more northeastern Comanche Plateau. Both subspecies 1974] Xanthisma — Semple 7 are absent in the southeastern portion of the Edwards Plateau (the Texas Hill Country). Thus differences in ge- ology account for the different numbers of collections in this well botanized area and also explain the means by which the two subspecies are isolated north and west of San Antonio, Texas. The presence of the Blacklands Prairies isolates the two subspecies north and east of San Antonio. The prairies, which run from San Antonio to Dallas and further north- eastward, are characterized by dark, high clay content soils (Sellards et al., 1966) and are dry and rock hard in the summer. Typically, Xanthisma occurs on sandy, light brown or reddish soils. Field work in the central Texas region showed that X. texanum is very rare throughout the entire length of the prairies, except for a few individu- als surviving on disturbed or introduced soils along high- ways and river banks. Clearly, Gray's varieties are exem- plary of what are today treated as subspecies. A narrow zone of isolated subsp. drummondii populations occurs in Caldwell, Gonzales and Wilson Counties, but the small populations only thrive in favorably moist years. Interestingly, Heterotheca pilosa (Nutt.) Shinners and Bradburia hirtella T. & G. are common in these counties and further east, the former being the most prevalent yel- low flowered member of the Astereae. Similarity in super- ficial appearance of these two species with X. texanum undoubtedly explains confusion about the distribution of X. texanum in this critical region between the two sub- Species. DIAGNOSTIC MORPHOLOGY AND DISTRIBUTION OF THE TWO VARIETIES OF SUBSP. TEXANUM A variety is characterized by all members of a population exhibiting a particular morphology distinct from other in- dividuals in the species. The distribution of these popula- tions is sympatric with populations whose members are not within the same variety, and also many populations of 8 Rhodora [Vol. 76 morphological intermediates exist (Du Reiz, 1930; Clausen, 1941; van Steenis, 1957; Valentine & Lóve, 1958; and Ehrendorfer, 1968). By this definition varieties can be recognized within a species, which does not have subspecies. Furthermore, variation that is mot known to be confined to an isolated group of populations can not be given sub- specific status. Van Steenis (loc. cit.) described varietal level variation as being continuous with other varieties, although the continuum would have pronounced modes. The variation between subspecies can be described as dis- continuous, except for the few hybrids. Figure 1 illustrates the sympatric condition of the two varieties of subsp. texanum: var. texanum with obtuse involucral bract apices (Figure 3.A) and var. orientale Semple with cuspidate involucral bract apices (Figure 3.B). FORMAL LEVEL VARIATION IN VAR. ORIENTALIS Formal level morphological variation is generally re- garded as some phenotype possessed by only some members of a population, whereas the subspecific and varietal level phenotypes are possessed by all members of a population. Those individuals in populations of subsp. texanum var. orientale which exhibit red pigmentation along the lateral margins of the involucral bracts forming the middle and inner series of the involucre have been designated as forma rubrum Semple (Figure 3.B). Such individuals are rare, but do occur in populations throughout the range of var. orientalis. GENETIC BASIS OF DIAGNOSTIC CHARACTERISTICS Experiments were conducted to show that all diagnostic characters have a genetic basis. Size but not general shape of the involucral bracts was found to be variable. Lack of sufficient light, available water, amount of soil or a high degree of crowding all resulted in smaller bracts being formed than those bracts formed by the individual when sufficient quantities of soil, light and water were provided. 1974] Xanthisma — Semple 9 The maximum size was determined by heredity and not the environment. Therefore size should be considered second- arily to shape in aetermining the proper identity of a par- ticular collection. The red pigment of f. rubrum was found to be passed from one generation to the next in var. orientale, but was never seen in var. texanum or in subsp. drummondii. Al- though the pigment is not always present in detectable quantities, its importance warrants some taxonomic status, since Xanthisma has always been previously described as monochromous. However, in some experimental plants heads were occasionally produced which manifested the red pigment in the distal portion of the ray straps on the adaxial surface and on the lobes of the disc florets, but only on the last day or two of disc anthesis. Most individuals produced enough pigment to give the rays an orange hue, but one plant produced rays that were deep rust red. Such variation in intensity of color does not occur on bract mar- £ins. No documentation of bicolor rays in wild plants exists. HABIT AS AN EXAMPLE OF VARIATION NOT WARRANTING TAXONOMIC STATUS Despite the great range of variation in habit occurring in the species, no one form was found to warrant taxonomic recognition. Within a single population long-stemmed and short-stemmed individuals were found in nature. The dis- tribution patterns of the short-stemmed plants (Figure 4.C) and the stemless plants (Figure 4.D) suggest that these forms are ecotypes, since they have a genetic basis. The extremes of short-stemmedness and stemlessness only occur in populations of var. orientale and western popula- tions of subsp. drummondii, respectively. The existence of à full range of intermediates between the extremes and the long-stemmed forms (Figures 4.B, 4.F and 4.G) in most populations makes taxonomic recognition unjustified. [Vol. 76 Rhodora 10 $a! npuowwnjp "esans Sijejuaio “ea WAVEIA) "fSqns whuera) “eA = whueray ULL 1974] Xanthisma — Semple 11 A COMMENT ON CHEMICAL STUDIES The taxonomic treatment that follows is based on mor- phology and distribution. Two dimensional paper chroma- tograms were run on leaf and flower extracts of specimens of each taxon. In general, subspecies texanum had fewer compounds than subspecies drummondii, and var. orientale has fewer compounds than var. texanum. In var. orientale individuals of forma rubrum had one compound, an uniden- tified anthocyanin (Semple, 1972b), not found in other individuals of the species. The full significance of the chemical studies can only be known upon complete identifi- cation of each compound. The preliminary results show that chemical data support the taxonomy. The chemical studies will have meaning in work on determining the evo- lution of the taxa described. TAXONOMIC TREATMENT The most recently published treatment of the genus Xan- thisma is in Correll & Johnston (1970). The following detailed description of the genus is more complete and ac- Figure 3. Involucral Bracts of Infraspecific Taxa. Each bract shown. is typical of those in the middle of the series forming the involucre: (A) subsp. texanum var. texanum; (B) subsp. texanum var. orientale Semple (dashed lines indicate the extent of red pigment on the margins of bracts of f. rubrum Semple); (C) subsp. drum- mondii (T. & G.) Semple. Figure 4. Habit Variation of Xanthisma texanum DC. The base- line under each silhouette equals one decimeter. Each form shown il- lustrates the habits of experimental plants and represents naturally occurring forms. Plants were grown under similar conditions from fruiting material collected at wild populations. All plants were from different populations, except (B) and (C). See the text for a full discussion. Figure 5. Heterocarpic Fruit of Xanthisma texanum DC. Ray and disc florets produce different kinds of fruit: (A), (B) and (C) are radial, tangential and diagrammatic cross-sectional views of a ray fruit; (D) and (E) are radial and diagrammatic cross-sectional views of a central disc fruit. Peripheral disc fruit are larger and angular ovate in cross-section. 12 Rhodora [Vol. 76 curate in light of the findings of my investigations. The previously unreported heterocarpic fruit condition is also described (Figure 5). A complete list of specimens exam- ined in the study is given in Semple (1972b) and only rep- resentative specimens are cited here. Xanthisma DC., Prodr. 5:94-95. 1836. Centauridium T. & G., Fl. N. Amer. 2:246. 1842. Taprooted annuals, rarely biennial and flowering the second season; branches ascending, few to many. Leaves alternate, the upper sessile, the lower terminally serrate; basal leaves petioled, lobed or pinnatifid to bipinnatifid. Heads solitary; involuere turbinate to hemispheric or cam- panulate; involucral bracts broad in 3-4 series, imbricate, stiff, the margins pale; ray florets pistillate, fertile, yellow; dise florets perfect, fertile, yellow. Fruit heterocarpic with pappus bristly and margins finely serrate; both fruit kinds with fine ascending white hairs, straw-colored when mature and darkening with age. Receptacle slightly convex, with subulate chaff reticulately dispersed between the fruits. The genus is monotypic and is endemic to the central southwestern areas of the United States. Xanthisma texanum DC., Prod, 5:95. 1836. Lectotype: Texas; Bexar County, in woods near Medina, May 1828, Berlandier 2039d (G!). Isolectotypes; G! (4), GH! (1), NY! (1), and us! (2). Herbs 2-7 (rarely more) dm tall, branching near the middle to ground level from a main stem 0.1-3 dm long, or rarely acaulescent. Upper leaves 1-2 cm long, linear, entire; basal leaves 5-8 cm long, ovate to obovate in outline, inter- grading above. Heads solitary, not crowded; involucre 5-10 mm high, involucral bracts 1.5-4 mm broad, rarely recurved, margins pale (the lateral margins very rarely red), the central portion green, reticulate venation pro- nounced, the inner-most bracts chartaceous, the outer-most linear grading into the peduncle bracts; ray florets 10-30, the straps 8-15 mm long, yellow adaxially, pale yellow 1974] Xanthisma — Semple 13 abaxially (rarely red-tinted abaxially) ; disc florets many 40-150, slightly ampliate to tapering, the lobes short. Ray floret cypselas curved obpyramidal, triangulate in cross- section, the pappus grading from short linear to long basally broad scales; disc floret cypselas obovate radially and narrowly-so tangentially, the central fruits rhombic in ercss-section, the outer obtrullate, the pappus in two whorls, the inner members ca. 6 mm long, linear, 8-15, the outer members 3 mm long or less many. The species is characterized by the following combination of characters: heterocarpic fruit with a pappus of bristly seales only, fruit pubescence of long white ascending hairs, receptacle slightly convex with a persistent reticulate net- work of subulate scales. The lectotype of the species was chosen from five speci- mens seen on microfilm only (1DC micro edition of Candolle Prodromi Herbarium). Since all five specimens are similar in quality, the specimen with the complete label giving col- lection number, date and location, and floral color (fourth from the left on the microfilm) is designated lectotype. Torrey (1859) first noted the typographic error in the Pro- dromus, which cited the type collection as Berlandier 2639 rather than Berlandier 2039. Key to Infraspecific Taxa I. Involucral bracts shorter than wide above the widest part, 1.5-3.5 mm wide and 1-2 mm long above the widest We eS, ur. spe Feats d yr g e raa subsp. texanum (1) II. Involucral bracts rounded with little or no flare 1 mm from the apex, occasionally recurved. i gu ck na ease esse es ewer’ var. texanum (2) II. Involueral bracts obtusely pointed, the deltoid por- tion 2-3 mm wide and 1.5-2 mm long above the widest part. ............-...-- var. orientale (3) III. Lateral margins of bracts whitish. .......... JI NRORURUIOPNBA TO ROS .... f. orientale: (38) III. Lateral margins of bracts red. f. rubrum (3b) 14 Rhodora [Vol. 76 I Involucral bracts longer than wide above the widest part; deltoid portion 2.5-5 mm wide and 3.5-8 mm long in bracts of the middle series. .. ssp. drummondii (4) 1. Xanthisma texanum spp. texanum Xanthisma texanum var. berlandieri Gray, Smiths. Contrib. Knowl. 3(5) : 98. 1852. Xanthisma berlandieri (Gray) Small, Fl. SE. U.S. p. 1184. 1903. This subspecies includes the type collection of X. texanum and is distinguished by small involucral bracts with rounded or obtusely pointed apices. Two varieties are recognized within the subspecies. 2. Xanthisma texanum ssp, texanum var. texanum The type variety of the species is distinguished by its small apically rounded involucral bracts 1.5-2 mm wide, appressed or rarely recurved; generally 30-35 bracts are visible without removing the outer series. The variety is endemic to the Nueces Plains of southern Texas from San Antonio to the south and west. East of San Antonio this variety forms hybrids with subspecies drummondit. SOME REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Atascosa Co., Charlotte, 4 May 1919, Schulz 39 (Us). Bexar Co., San Antonio, 4 June 1948, Burr 21 (NY, TEX), in woods near the Medina River, July 1829, Berlandier 629 (GH, MO, NY, US). Dimmit Co., Big Wells, 21 April 1945, Shinners 7,400 (NY, OKLA, SMU, TEX), in front of cemetery, 6 May 1964, Turner 4,996 (TEX). Frio Co., Dilley, 6 mi. W, 8 July 1970 Lewis 7,634 (Mo). La Salle Co., Artesia Wells, 3 mi. W., 7 April 1963, Dickey 170 (SMU, TEX). Medina Co., Farm Rd. 1343, 4.4 mi S of U.S. Hwy. 90, 29 June 1972, Semple & Shea 720 (MO) ; Devine, 3 mi. SW, 1 May 1954, Tharp & Turner 3,439 (OKLA, TEX). Zavala Co., Batesville, 6 mi. E, 6 May 1964, Turner 5,002 (TEX). 9. Xanthisma texanum ssp. texanum var. orientale Semple, var. nov. Involucri bracteae parvae manifeste depresse ovatae, apice obtuse, cuspidatae, interdum marginibus lateralibus rubris. 1974] Xanthisma — Semple 15 HOLOTYPE: Texas: Hidalgo Co.: 13.5 mi. N. of Edinburg on U. S. Hwy. 183, 8 July 1970, W. Lewis 1642 (Mo). Iso- types: four, to be distributed. The involucral bracts are small and pronouncedly shield- shaped, occasionally with red lateral margins. The apices are obtusely cuspidate. This variety is distinguished by the 2-3.5 mm wide in- volucral bracts having obtuse, cuspidate apices. The short stem habit is also frequently encountered. Hybridization with var. texanum is common, but is unknown with ssp. drummondii. The name is taken from its distribution in the eastern half of the range of ssp. texanum. Two forms are recognized. 3a. Xanthisma texanum var. orientale Semple f. orientale The type form is the most common form of the variety. SOME REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Aransas Co., Aransas Refuge, 29 Sept. 1944, Cory 45, 895 (Ny, TEX). Bee Co., Beeville, 21 June 1935, Drushel 9,941 (NY). Brooks Co., Hebbron- ville, 15 mi. E, 25 Nov. 1962, Dohnke T (sMU). Calhoun Co., Port Lavaca, 14 June 1953, Johnston 53.280.157 (OKLA, TEX). Goliad Co., Goliad, 15.9 mi. S, 9 July 1970, Lewis 7,655 (Mo). Hidalgo Co., Rio Grande Valley, 8 Aug. 1942, Walker 70 (KSU, TEX). Jim Hogg Co., Hebbronville, 8 mi. E, 8 July 1970, Lewis 7,638 (MO). Jim Wells Co., Premont, 12 mi. N, 24 Nov. 1954, Johnston 542,144 (TEX). Kenedy Co., King Ranch, Norias Div. 24 Nov. 1953 Johnston 53.280.154 (OKLA, TEX.) Kleberg Co., Kingville, 1940, Sinclair s.n. (TEX). Refugio Co., 5 mi. S of county line on U. S. Hwy. 77A, 12 June 1971, Semple 581 (Mo). San Patricio Co., Aransas Pass, 24 May 1922, Schulz 868 (TEX, US). Near San Patricio, May 1834, Berlandier 2578 (GH, MO, NY). Webb Co., Mirando City, 31 Aug. 1936, Baird s.n. (NY). Willacy Co. Yturria Station, 6 Aug. 1924, Runyon 664 (TEX, US). 3b. Xanthisma texanum var. orientale f. rubrum Semple, forma nov. Involucri bracteae cum marginibus later- alibus rubris. HOLOTYPE: Texas: San Patricio Co., 0.5 mi. E. of Ingleside on Texas Hwy. 361, 13 June 1971, Semple 602 (Mo). Isotype: one to be distributed. This form is distinguished from the type form by having bracts with red lateral margins. 16 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Aransas Co., Aransas Refuge, 29 Sept. 1944, Cory 45,895 (Mo). Goliad Co., General Zaragoza State Park, 1 mi. S, 12 June 1971, Semple 579 (mo). Kenedy Co., Raymondville, 16 mi. N, 2 Dec. 1945, Cory 51,493 (us). Willacy Co., Port Mansfield, 0.5 mi. W of beach by garbage dump, 9 July 1970, Lewis 7,644 (MO). 4. Xanthisma texanum DC. ssp. drummondii (T. & G.) Semple, stat. nov. Centauridium drummondii T. & G., Fl. N. Amer. 2: 462. 1842. SYNTYPES: Texas, north of San Antonio, Riddell s.n. (NY) ; Texas, exact locality unknown, Nov. 1835, Drummond 227 (GH!, NY!). LECTOTYPE: Drummond 227 (GH). Isolectotypes: GH, NY. Xanthisma texanum var. drummondii (T. & G.) Gray, Smiths. Contrib. Knowl. 3(5) : 98. 1852. Xanthisma drummondii (T. & G.) Hooker f., Curtis’ Bot. Mag. 33: t. 6275. 1877. Subspecies drummondii is distinguished by its involucral bracts, which are wider than those of ssp. texanwm and are distinctly lanceolate. Populations occur in much of Texas north of San Antonio, and in Oklahoma and eastern New Mexico. Drummond 227 (GH) was selected lectotype because it is the largest and most complete specimen. The Riddell col- lection cited by Torrey & Gray (1842) is fragmentary and a putative hybrid between the two subspecies. SOME REPRESENTATIVE SPECIMENS EXAMINED. NEW MEXICO: Roose- velt Co., Elida, 5 mi. NE, 28 Sept. 1965, Anderson 3,004 (KSU). Lingo, 1.2 mi. N, Semple & Shea 705 (MO). OKLAHOMA: Beckham Co., Carter, 5 mi. S, 5 July 1965, Thomas 79 (OKLA). Caddo Co., Cement, 27 June 1936, Demaree 18, 133 (NY). Comanche Co., Cache, June 1927, Ortenburger s.n. (U.S.) Creek Co., Drumright, 10 June 1935, Fry 8 (OKLA). Custer Co., Clinton, 28 June 1966, Ganz s.n. (ARIZ. st. U.). Grady Co., Verdun, 25 June 1963, Pearce 885 (OKLA, SMU). Greer Co., Granite, 17 June 1939, Grawmanns 85 (OKLA). Harmon Co., Vinson, Shinners 31,644 (sMU). Jackson Co., Duke, 1.7 mi W. 15 July 1970, Flyr s.n. (MO). Kay Co.; 26 July 1898, White s.n. (us). Kiowa Co. Mt. Park, 23 June 1913, Stevens 1,293 (OKLA). 1974] Xanthisma — Semple 17 Lincoln, 23 Aug. 1895, Blankenship 17,334 (wis). Logan Co., Guthrie, 12 July 1916, Keyser 6,036 (Ny). Osage Co., 8 mi. S of Kans. state line on Okla. Hwy. 99, 24 June 1972, Semple & Shea 667 (MO). Oklahoma Co., Oklahoma City, 27 May 1937, Waterfall 582 (OKLA, NY). Pawnee Co., Pawnee, 8 mi. E, 8 June 1934, Brodell s.n. (TEX). Payne Co., Mulhall, 1.1 mi. N, 25 June 1972, Semple & Shea 674 (MO). Stephens Co., Duncan, 8 mi. N, 27 May 1960, Waterfall 15,911 (OKLA). Washita Co., 2.2 mi. E of county line on Okla. Hwy. 152, 20 Aug. 1970, Semple & Love 264 (Mo). TEXAS: Andrews Co., Andrews, 4 mi. W, 28 May 1956, Turner 3,957 (TEX). Archer Co., Mankins, 2 mi. NE, 8 June 1956, Shinners 23,758 (SMU). Baylor Co., Seymour, 4 Aug. 1955, Shinners 20,776 (SMU). Blanco Co., Silverton, Farm Rd. 145, 26 June 1972, Semple & Shea 699 (M0). Burleson Co., Caldwell, 14 June 1971, Semple 625 (Mo). Burnet, 8 mi. W, 19 May 1955 Turner & Johnston 2,480 (TEX). Callahan Co., Baird, Aug. 1882, Letterman 26 (Mo, US). Childress Co., Childress, Biology Class C. High School 12 (TEX). Coke Co., Robert Lee, 3 mi. S, 4 Oct. 1968. Gary 192 (TrC). Comanche Co. Comyn, Theney School s.n. (TEX), Dallas Co., Dallas, near Comanche Peak, Aug. 1877, Reverchon s.n. (vs). Dewitt Co., western part of Co., 20 July 1941, Reidel s.n. (TEX). Erath Co., 1922, Gaugh 29 (vs). Fisher Co., Rotan, 10 Sept. 1933, Brooker s.n. (TEX). Gaines Co., Lamesa, 15.1 mi. W, 14 Sept. 1946, Whitehouse 16,788 (NY, SMU). Gillespie Co., Cherry Spring, 18??, Jermy 639 (Mo). Gonzales Co., Cost, 11.5 mi. SW, 29 June 1972, Semple & Shea 730 (Mo). Hall Co., Turkey, 5 mi. S, 12 June 1950, Tilton T.550.604 (TTC). Hood Co., 6 Aug. 1877, Reverchon 1368 (KsU. NY, US). Howard Co. Big Spring, 22 mi. S, 27 April 1963. Dallas 64 (OKLA). Kerr Co.. 6 June 1929, Whitehouse 7.056 (TEX). Limestone Co.. Kosse, 3.5 mi. N, 14 June 1971, Semple 626 (MO). Lubbock Co., Lubbock, Demaree 7,715 (MO, TEX, TTC. US). McCullogh Co., 1 mi. S of Colorado R., 28 June 1972, Semple & Shea 716 (MO). Navarro Co., Corsicana, 21 June 1872, Hall 311 (GH, MO, NY, Us). Robertson Co., Calvert, 8 May 1904, Tyler s.n. (US). Tar- rant Co. Lake Fort Worth, 16 July 1923, Ruth 742 (GH, KSU, US, wis). Travis Co. Austin, 9 June 1908, Biltmore s.n. (US). Van Zandt Co., Edom, 3 July 1939, Ball 7 (sMv). Washington Co., 20 May 1938, Brackett s.n. (TEX). Young Co. Belknap, 6 May 1858, Hayes 397 (NY). LITERATURE CITED Braun, E. L. 1950. Deciduous Forests of Eastern North America. Blakiston Co. Philadelphia. Bray, W. L. 1906. Distribution and Adaptation of the Vegetation of Texas. Bull. of U. of Tex. No. 82, Sci. Ser. No. 10. 18 Rhodora [Vol. 76 CANDOLLE, A. P. DE. 1836. Prodromus 5: 94-95. CLAUSEN, R. T. 1941. On the terms “subspecies” and “variety.” Rhodora 43: 157-167. CORRELL, D. S. & M. C. JoHNSTON. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation Press. Austin. Du RErTZ, G. E. 1930. The fundamental units of biological tax- onomy. Svensk. Bot. Tidskr. 24: 333-428. EHRENDORFER, F. 1968. Geographical and ecological aspects of infraspecific differentiation. In V. H. Heywood, ed. Modern Methods of Plant Taxonomy. Academic Press. New York. FENNEMAN, N. M. 1931. Physiography of the Western United States. MeGraw-Hill Book Co. New York. FERNALD, M. L. 1940. Some spermatophytes of eastern North America. Rhodora 42: 239-278. FoscuE, E. J. 1932. The natural vegetation of the lower Rio Grande Valley, Texas. Field & Lab. 1: 25-30. GOULD, F. W. 1962. Texas Plants— a checklist and ecological summary. Texas A. & M. — Tex. Agr. Ex. Sta. College Station, Texas. GRAY, A. 1852. Plantae Wrightiana Texano — Neo-Mexicanae. Part I. Smiths. Contrib. Knowl. 3: 98. Hooker, J. D. 1877. Curtis’ Bot. Mag. 33: t.6275. OETKING, P. F. 1959. Geological Highway Map of Texas. Dallas Geological Soc. Dallas. Rarsz, E. 1957. Land forms of the United States. Jn Oetking, P.F. 1959. Geological Highway Map of Texas. Dallas Geological Soc. Dallas. SELLARD, E. H., W. S. ADKINS & R. B. PLUMMER. 1966. Geology of Texas. Vol. I. Stratigraphy. U. of Tex. Bull. No. 3232. SEMPLE, J. C. 1972a. Behavior of B-chromosomes in Xanthisma texanum DC.: a non-random phenomenon. Science 175: 666. 1972b. Cytology, Flavonoid Chemistry and System- aties of the Texas Sleepy Daisy Xanthisma texanum DC. (As- teraceae). Ph.D. Dissertation. Washington University. St. Louis. SMALL, J. K. 1903. Flora of the Southeastern United States of America. Pub. by Small. New York. p. 1184. SMITH, E. B. & H. M. Parker. 1971. A biosystematic study of Coreopsis tinctoria and C. cardaminefolia. Brittonia 23: 161-170. STEENIS, C. G. G. J. VAN. 1957. Specific and infraspecific delimina- tion. Flora Malesiana Ser. 1, 5: clxvii-eexxxix. THARP, B. C. 1926. Structure of the Texas Vegetation East of the 98th Meridian. U. of Tex. Bull. No. 2606. Torrey, J. 1859. In Emory, W. H., ed. Report on the United States and Mexican Boundary Survey. Nicholson Press. Washington. Vol. II. pp. 80-81. 1974] Xanthisma — Semple 19 . & A. GRAY. 1842. A Flora of North America. Vol. II. facsimile ed., 1969. Hafner Pub. Co. New York. p. 246. VALENTINE, D. H. & A. LóvE. 1958. Taxonomic and biosystematic categories. Brittonia 10: 153-166. VISHER, S. S. 1954. Climatic Atlas of the United States. Harvard U. Press. Cambridge. MISSOURI BOTANICAL GARDEN ST. LOUIS, MO. 63110 A WHITE-FLOWERED FORM OF UTRICULARIA PURPUREA FROM NEW HAMPSHIRE. A white- flowered form of Utricularia purpurea Walter was found while conducting research in a small pond in New Hamp- shire. This form was extremely abundant forming exten- sive mats on the surface of this shallow pond. None of the normal purple-flowered plants were observed. Plants commonly associated with the white form were Utricularia vulgaris L., Utricularia intermedia Hayne and Najas flexilis Rostk. & Schmidt. The pond had a pH of 6.5 and methyl orange alkalinity reading of 8.0 mg/1. The following form is described. Utricularia purpurea Walter forma alba Hellquist, forma nova. differt forma purpurea quantum flores albos non purpureos habet. TYPE: NEW HAMPSHIRE: CARROLL COUNTY: small unnamed pond southeast of Dorr Pond on N.H. Route 153 at the settlement of Woodman, Town of Wakefield (U.S.G.S. quadrangle Newfieid, Me.-N.H.), C. B. Hellquist 8935 (NHA). The holotype has been placed at the University of New Hampshire Herbarium (NHA). Isotypes are in the her- barium of the New England Botanical Club (NEBC) and Boston State College Herbarium. C. BARRE HELLQUIST BIOLOGY DEPARTMENT BOSTON STATE COLLEGE BOSTON, MASSACHUSETTS 02115 ON THE SCIENTIFIC NAME OF THE LONGLEAF PINE' DANIEL B. WARD The Longleaf Pine of the southeastern United States has been spared the numerous changes in scientific name that have cast confusion over many of our other tree species. From pioneer days the dominant practice, both among botanists and foresters, has been to apply the name Pinus palustris Mill. to the Longleaf. Prior to the 1940's the only author of significance to do otherwise was J. K. Small (1933) who interpreted P. palustris Mill. to be the northern variety of slash pine, P. elliottii Engelm. Small’s Manual of the Southeastern Flora, however, achieved only regional impact, and his use of P. australis Michx. f. for the Long- leaf did not gain a wide following. A much stronger impetus for associating the Longleaf with Pinus australis was generated in 1948 by M. L. Fer- nald who advocated this name in two detailed and strongly worded articles (Fernald & Schubert, 1948; Fernald, 1948). Fernald's position was reinforced by the publica- tion of his monumental reworking of Gray's Manual of Botany (1950), and was further entrenched when the inde- pendent H. A. Gleason adopted the same name in his New Britton and Brown Illustrated Flora (1952) as did A. Cronquist in their companion field guide, the Manual of Vascular Plants of Northeastern United States and Adja- cent Canada (Gleason & Cronquist, 1963). These three publications totally dominate the field of regional floras in the Northeast, and their consistent use of Pinus australis inevitably suggests that this name is supported by general professional agreement and by nomenclatural legitimacy. Yet in publications other than these or their derivatives, the Longleaf remains known by the name Pinus palustris "This paper is Florida Agricultural Experiment Station Journal Series No. 4775. 20 1974] Longleaf Pine — Ward 21 Mill. This is the name found in state or district floras such as the Manual of the Vascular Flora. of the Carolinas (Radford et al., 1968), the Manual of the Vascular Plants of Texas (Correll & Johnston, 1970), and A Flora of Tropical Florida (Long & Lakela, 1971), as well as in the more basic modern treatments of the genus by Mirov (1967) and by Little & Critchfield (1969). The U. S. Forest Service, following the guidance of its dendrologist, E. L. Little, has remained steadfast with P. palustris, and the state forest services have done likewise. The rationale behind Fernald's heterodox displacement of Pinus palustris was his view that Philip Miller (1768), superintendent of the Chelsea Physic Garden and British author of a series of widely used and authoritative horti- cultural dictionaries, did not have the Longleaf when he described an American tree under this name. The original description contained phrases that do not apply to the stately and versatile Longleaf: “I have been informed that they grow to a height of twenty-five or thirty feet. I have not heard the wood was of any use but for fuel." However, Miller reported these statements as hearsay, not as observed fact, while his description also contained such passages as ‘“‘Pine-tree with the longest leaves growing by threes out of each sheath" and “Their leaves are a foot or more in length, growing in tufts at the end of the branches, so have a singular appearance." These phrases were given without qualification and form a brief but unmistakable description of the Longleaf, Fernald further relied on the habitat of Pinus palustris reported by Miller, “swamps in many parts of North America," as excluding the Longleaf. He proposed instead that Miller may have had the Loblolly Pine, P. taeda L. Actually however, although the Longleaf usually occurs on dry well-drained soils, it not uncommonly is found associated with the Slash Pine in the extensive level undrained swamps known as flatwoods and throughout its range probably occurs as frequently in wet soils as Fernald's proposed substitute, the Loblolly. Of couse, Miller's use of an epithet 22 Rhodora [Vol. 76 meaning “swampy”, even if it were totally unsuitable, would not be justification for discarding his P. palustris; Article 62 of the International Code of Botanical Nomen- clature (Stafleu, 1972) provides that: “a legitimate name or epithet must not be rejected merely because it is in- appropriate . . ." No information is available from specimens preserved by Miller; Dr. Schubert examined the Miller collections of the British Museum and was unable to find a specimen that could be considered a type. Fernald, by correspondence, did locate a specimen he presumed to be of P. taeda that Miller may have seen and that someone, possibly Miller, had noted as “palustris.” It seems improbable that Miller would have so misidentified P. taeda since this species was well known to him and was treated in some detail in the several editions of his “Dictionary.” Fernald did demonstrate at some length that the name Pinus palustris was not clearly understood and was even misused by Miller's contemporaries and other early workers, but this again does not provide justification for discarding the name. There seems to be essentially no sound rationale for interpreting this name to mean anything other than, in the conventional usage, the Longleaf Pine. In the event, moreover, that Miller's brief but clear diagnosis should be considered ambiguous, no alternative name is available. Pinus lutea Walter, whimsically sug- gested by Fernald, is probably P. taeda. Pinus australis Michx. f. is unmistakably the Longleaf, and was selected as the correct name for this species by Small and by Fernald. Little (1948), however, accurately pointed out prior to the publication of Fernald's first paper on the subject, that Article 63 (then Article 60) of the Inter- national Code makes this name illegitimate since it was superfluous when published. Not only did Michaux (1810) cite P. palustris as a synonym, but he specifically explained that he was replacing this name with P. australis, using an epithet (“southern”) that he thought more appropriate. It is precisely such actions as this that Article 63 is de- 1974] Longleaf Pine — Ward 23 signed to prevent, and Fernald's protestations that Michaux was “definitely defining a new species" are contradicted by Michaux' own words, in translation: *I have thought like- wise that the specific name ‘australis’ was preferable to that of ‘palustris’, under which this species has been de- scribed by botanists; for this last gives an absolutely false idea of the nature of the soil where this tree grows.” The only other specific name apparently ever given the Longleaf was Pinus longifolia Salisb., but here again the epithet used was superfluous, for Salisbury (1796) was un- abashedly providing a substitute name for P. palustris and thereby formed an illegitimate and unusable combination. The person who would apply a scientific name to the Longleaf Pine, therefore, has the choice, should he wish to follow the Code, of either accepting Miller’s brief descrip- tion of Pinus palustris as adequate, or of discarding it as confused and coining and publishing a name of his own creation. Neither P. australis Michx. f. nor P. longifolia Salisb. may be legitimately used for this tree, and prudence and practicality, as well as historical precedent, indicate strongly the advisability of retaining the Longleaf Pine under the name Pinus palustris Mill. LITERATURE CITED CORRELL, D. S. & M. C. JOHNSTON. 1970. Manual of the Vascular Plants of Texas. Texas Research Found. 1881 p. FERNALD, M. L. 1948. The confused bases of the name Pinus palustris. Rhodora 50: 241-249. . 1950. Gray's Manual of Botany, 8th ed. Ameri- can Book Co. 1632 p. , & B. G. SCHUBERT. 1948. Studies of American types in British herbaria. Rhodora 50: 181-190. GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora. Lancaster Press. Vol. 1. ,& A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Van Nos- trand Co. 810 p. LITTLE, E. L. 1948. Notes on nomenclature of trees. Phytologia 2: 457-458. 24 Rhodora [Vol. 76 , & W. B. CRITCHFIELD. 1969. Subdivisions of the genus Pinus (pines). U. S. Dept. of Agric. For. Ser. misc. publ. 1144. 51 p. Lonc, R. W., & O. LAKELA. 1971. A Flora of Tropical Florida. Univ. of Miami Press. 962 p. MicHAUX, F. A. 1810. Histoire des Arbres Forestiers de l'Amerique Septentrionale. 1: 64. MiLLER, P. 1768. The Gardeners Dictionary, 8th ed. London. Mirov, N. T. 1967. The Genus Pinus. Ronald Press. 602 p. RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the Vascular Flora of the Carolinas. Univ. N. Car. Press. 1183 p. SALISBURY, R. A. 1796. Prodromus Stirpium in Horto ad Chapel Allerton Vigentium. London. SMALL, J. K. 1933. Manual of the Southeastern Flora. The author, New York. 1554 p. STAFLEU, F. A. 1972. International Code of Botanical Nomencla- ture. Utrecht. 426 p. DEPARTMENT OF BOTANY AGRICULTURAL EXPERIMENT STATION UNIVERSITY OF FLORIDA GAINESVILLE, FLORIDA THE OCCURRENCE OF BALD CYPRESS (TAXO- DIUM DISTICHUM (L) RICHARD) IN SUFFOLK COUNTY, LONG ISLAND, NEW YORK. In October, 1972, Mr. Richard Fackovec, a student in an aquatic botany course at Southampton College, brought to our attention a specimen of Taxodium distichum that he col- lected along the shore of Long Pond in Sag Harbor, New York. We returned to investigate the region in November and found eight bald cypress seedlings, ranging in height from one to 214 meters and growing close to shore in 0.5-0.7 meters of water. The four smallest trees (approxi- mately one meter tall) are growing close together at the southern end of the pond at the edge of a large marsh. The other cypress trees are located at the southern end of the pond also, but along the western shore. The two largest trees (215 and 114 meters tall) are at the edge of a large stand of Typha latifolia. Most of the remaining shoreline of Long Pond consists of very dense vegetation composed primarily of shrubs and small trees (e.g. Myrica gale, Decodon verticillata, Cassandra calyculata, Vaccinium corymbosum, Nyssa sylvatica, Acer rubrum). It is within this shrub zone that we found two additional bald cypress seedlings. One had become encompassed so completely by encroaching shoreline vegetation that we removed it; sub- sequently, it has been replanted along the shore of a pond in the Morton Wildlife Refuge, a Federal Wildlife Preserve in Noyac, New York. We have communicated with Dr. Richard Stalter, plant taxonomist at St. Johns University in Jamaica, New York, regarding the northern distribution of Taxodium distichum. As far as we have been able to determine, this is the first record of a natural population of bald cypress in New York State. The closest naturally-occurring populations of Taxodium are in Delaware, where there are a few large colonies (Stalter, personal communication). There was, at one time, a colony in New Jersey, but it has been un- known in that state for several years (Stalter, personal communication). 25 26 Rhodora [Vol. 76 Accompanied by Dr. Stalter, we returned to Long Pond in December in an unsuccessful effort to locate mature bald cypress trees. We share the opinion with Dr. Statler that this colony of Taxodium distichum represents a natu- ralized population. The presence of a few bald cypress seedlings in relatively inaccessible portions of the shore- line strongly suggests that they were not planted. The apparent absence of large bald cypress in the area leaves unanswered the question as to how the seedlings became established; hopefully, further investigation will afford us an answer. The voucher specimen collected by Mr. Fackovec is de- posited in the Southampton College Herbarium. EDWARD J. HEHRE PETER M. BOLLARD DEPARTMENT OF BIOLOGY SOUTHAMPTON COLLEGE SOUTHAMPTON, NEW YORK 11968 STUDIES ON THE BRYOPHYTES OF SOUTHERN MANITOBA III. COLLECTIONS FROM GRAND BEACH PROVINCIAL PARK PAUL W. STRINGER AND MURIEL H. L. STRINGER Grand Beach Provincial Park lies approximately 57 miles north of Winnipeg. The entire park, which covers 59,000 acres, is in the Manitoba Lowlands Section of the Boreal Forest Region (Rowe, 1959). Three miles of the area fronts on Lake Winnipeg. 'The underlying bedrock is palaeozoic limestone, and the overlying beds are modified till, or lacustrine clays deposited in glacial Lake Agassiz. Most soils have a high lime content. On the flat, poorly drained areas, black spruce (Picea mariana (Mill. BSP.) and tamarack (Larix laricina (Du Roi) K. Kich) forest prevail, while on the better drained alluvial strips bordering the rivers and creeks, white spruce (Picea glauca (Moench) Voss), aspen (Populus tremuloides Michx.) and balsam poplar (Populus balsamifera L.) pre- dominate. On the low, well-drained ridges, jack pine (Pinus banksiana Lamb.) is dominant, with aspen, paper birch (Betula papyrifera Marsh.) and some bur oak (Quercus macrocarpa Michx.) present. No bryophytes have previously been reported for the park. In the present study, several sites were visited, mostly in the vicinity of 96°35’W and 50°33’N. The sites were chosen as representative of the major vegetation types in the area, and the following seven, visited in July, 1972, are reported as representative of the area studied. Several of the vascular collections could be determined to genus only, because of lack of flowering or fruiting material. 1. Sand dunes and dune slacks behind the East Beach. Dominant shrubs were Salix spp., especially in the slacks, with Prunus pumila L. conspicuous on the drier areas. Other prominent vascular plants on the dry areas included Carex spp. Rhus radicans L. var. rydbergii (Small) 27 28 Rhodora [Vol. 76 Rehder, and Artemisia spp. Total plant cover was rather thin (about 20%) except in the slacks, and most bryophyte cover was on wet litter in the slacks. Small turf mosses such as Bryum spp. were, however, common in dry habi- tats. 2. Aspen-dominated forest just s. of the parking lot at the East Beach. Most mosses were from tree bases. 3. Mixed mature aspen — jack pine forest on w. side of the road n. of main park entrance. 4. Black spruce bog with some tamarack and white cedar (Thuja occidentalis L.). The understory was dominated by Ledum groenlandicum Oeder and the terrain consisted of pools and Carex hummocks. Just s. of Highway 12, 14 mile from its junction with Highway 59. 5. Open jack pine with some birch, on s. side of Highway 12, 2 miles from its junction with Highway 59. 6. Tamarack — black spruce bog on e. side of Highway 59, 2 miles n. of its junction with Highway 12. Drier than site 4, and with a tall shrub layer of Alnus sp., as well as a short shrub layer dominated by Ledum groenlandicum. Deadfall was abundant. 7. Mature, open jack pine forest on w. side of Highway 59 4 miles n. of its junction with Highway 12. There was an open short shrub and herb layer dominated by Vaccinium angustifolium Ait., Chimaphila umbellata (L.) Bart. and Amelanchier alnifolia Nutt., with Linnaea borealis L. and Vicia americana Muhl. prominent. The forest floor was dominated by Plewrozium schreberi and Dicranum spp., especially D. polysetum. The many large humus-covered boulders present all had an extensive, species-rich bryo- phyte cover. Numbers in the species list refer to the sites described above. Taxonomic and field work were shared equally by both authors. Voucher specimens have been deposited in the authors’ own herbarium and at the University of Winnipeg. Nomenclature for Sphagnobrya and Eubrya follows Crum, Steere, and Anderson (1965) with modification according to Crum (1971). Hepatic nomenclature follows Schuster 1974] Bryophytes — Stringer & Stringer 29 (1969), supplemented by Schuster (1953) with abbrevia- tions of authorities amended to conform with the list of Sayre, Bonner and Culberson (1964). Vascular plant no- menclature follows Scoggan (1957). Hepaticae Calypogeia muelleriana (Schiffn.) K. Miill. (6). Three collections, two on humus mixed with well-rotted wood, and one on a rotten stump with Lophocolea heterophylla, Cephalozia media and Hypnum pallescens. Only once pre- viously recorded for the province (Longton, 1972). Cephalozia media Lindb. (6). One collection, on a rotten stump. | Cephaloziella rubella (Nees) Douin (4). On humus over rotten wood, with Amblystegium juratzkanum, Chiloscy- phus pallescens and Thuidium recognitum. Chiloscyphus pallescens (Ehrh.) Dum. (4). Only once pre- viously recorded for the province, in Spruce Woods Pro- vincial Park (Stringer & Stringer, a, in preparation). Chiloscyphus polyanthus (L.) Corda (3, 4). On bark at the base of an aspen tree (3) with Hypnum pallescens, and on humus with Distichium inclinatum. Recorded previ- ously only in Spruce Woods Provincial Park (Stringer & Stringer, in preparation). Lophocolea heterophylla (Schrad.) Dum. (3, 6, 7,). On humus or rotten wood, often with Pohlia nutans or Hyp- num pallescens. Previously recorded only in the Spruce Woods area by Bird (1969) and by Stringer & Stringer (in preparation). Lophocolea minor Nees (6). On thick humus (dead Aula- comnium palustre), with Pohlia nutans and Eurhynchium pulchellum. Not previously recorded for the province. Ptilidium ciliare (L.) Nees (5, 6, 7,). On loose humus or litter with Dicranum spp. and Pleurozium schreberi. One collection (7) on loose conifer needles over rock, with Hedwigia ciliata. Ptilidium pulcherrimum (Web.) Hampe (7). One collec- tion, on fallen birch bark. 30 Rhodora [Vol. 76 Radula complanata (L.) Dum. (7) One collection, on packed humus at the base of a jack pine tree. Riccardia pinguis (L.) S. Gray (4). On thick, moist humus mixed with silt, with Myurella julacea, Amblystegium juratzkanum and Platydictya jungermannoides. Previ- ously recorded only in the Spruce Woods area by Bird (1969) and Stringer & Stringer (in preparation). Sphagnobrya Sphagnum capillaceum (Weiss) Schrank (6). In deep hum- mocks, with Pleurozium schreberi. Eubrya Amblystegium juratzkanum Schimp. (1, 2, 4, 5, 6, 7). On rotten wood or humus, especially at tree bases, where it was often found with Pylaisiella polyantha and Brachy- thecium salebrosum. Amblystegium varium (Hedw.) Lindb. (1, 2, 6). On thick humus and rotten wood, with Brachythecium spp. and Mnium cuspidatum. Aulacomnium palustre (Hedw.) Schwaegr. (4, 5, 6). On thick humus, in deep pure sods, or with Pleurozium schreberi. One collection (5) on rotting birch bark. Brachythecium campestre (C. Müll.) B. S. G. (5, 6, 7). On humus, usually over well-drained sandy soil. Previously recorded in Manitoba only in the Winnipeg and Spruce Woods areas by Stringer & Stringer (1973) and Stringer & Stringer (in preparation). Brachythecium rutabulum (Hedw.) B. S. G. (1, 2, 5, 6, 7). On humus or rotten wood in moist habitats. Previously recorded only by Mueller-Dombois (1964), Longton (1972), and by Stringer & Stringer (in preparation), in the Spruce Woods area. Brachythecium salebrosum (Web. and Mohr) B. S. G. (1, 2, 3, 5, 6, 7). Common in many habitats, especially on tree bases. Bryoerythrophyllum recurvirostrum (Hedw.) Chen (4). On thick humus, with Amblystegium juratzkanum, Myurella julacea and Platydictya jungermannoides. 1974] Bryophytes — Stringer & Stringer 31 Bryum angustirete Kindb. ex Mac. (1). On humus over sand, alone, and with Encalypta procera. Bryum argenteum Hedw. (1, 3). On dry humus and sand. Often with Funaria hygrometrica. Bryum creberrimum Tayl. (3, 5). On humus mixed with sand. Often with Ceratodon purpureus. Bryum pallescens Schleich. ex Schwaegr. (I, 2,3). Oh humus in moist habitats. Bryum pseudotriquetrum (Hedw.) Gaertn. Meyer and Scherb. (2, 4). On thick, moist humus with Campylium stellatum and Brachythecium rutabulum. Callicladium haldanianum (Grev.) Crum (5, 6, 7). On humus and moist rotten wood. Previously reported only in the Winnipeg and Spruce Woods areas by Bird (1969) and by Stringer & Stringer (a, in preparation). Calliergon giganteum (Schimp.) Kindb. (4). On water- logged litter, in large masses mixed with Campylium stellatum. Campylium chrysophyllum (Brid.) J. Lange (4). On well- rotted wood and humus. Campylium hispidulum (Brid.) Mitt. (4). One collection, on humus. with Amblystegium juratzkanum. Campylium stellatum (Hedw.) C. Jens. (4, 6). On wet humus or litter, often with Mnium affine or Thuidium delicatulum var. radicans. Ceratodon purpureus (Hedw.) Brid. (1, 3, 4, 5, 7). On humus over dry sand, with Bryum spp., Funaria hygro- metrica, or Polytrichum juniperinum. Dicranum bonjeanii De Not. ex Lisa (7). On thick humus over rotten wood, with D. polysetum and Ptilidium cili- are. Dicranum drummondii C. Müll. (6, 7). In thick, deep, pure clumps on humus, or as small plants on rotten wood with such typical species of decayed wood as Pohlia nutans and Tetraphis pellucida. Dicranum fuscescens Turn. (5, 7). On humus or rotten wood, usually in pure tufts. 32 Rhodora [Vol. 76 Dicranun polysetum Sw. (4, 5, 6, 7). Common, especially in jack pine forests of the area. Forms deep pure tufts. Often found also with Pleurozium schreberi. Distichium inclinatum (Hedw.) B. S. G. (4). Common in this collecting area and most often found with Myurella julacea, Amblystegium juratzkanum, and Platydictya jungermannoides. Several of the collections had mature capsules, and the large warty spores (354-404) distin- guish this species from the closely related Distichium capillaceum (Grout, 1936). Drepanocladus aduncus (Hedw.) Warnst. (1). On wet lit- ter under willows. Drepanocladus aduncus (Hedw.) Warnst. var. polycarpus . (Bland. ex Voit) Roth (1, 2). More commonly found . than the preceding, this variety occurred on moist humus or litter, often with Leptodictyum riparium. Drepanocladus revolvens (Sw.) Warnst. (4). Several col- lections made on wet litter or humus, in pure mats or with Pleurozium schreberi and Thuidium recognitum. Drepanocladus uncinatus (Hedw.) Warnst. (4, 5, 6, 7). On rotten wood or humus, in drier habitats than other mem- bers of the genus. With Brachythecium spp. and Bryum Spp. Encalypta procera Bruch (1). One collection, on thin hu- mus over dry sand. Eurhynchium pulchellum (Hedw.) Jenn. (4, 6, 7). On thick humus or litter in both wet and dry habitats. Fissidens adianthoides Hedw. (4). One small collection, with Campylium stellatum, on loose, moist humus. Not previously recorded for the province. Funaria hygrometrica Hedw. (1, 4, 5, 7). Frequent on sandy soil, often mixed with ashes of old fires. Com- monly with Bryum spp., Ceratodon purpureus, and Lepto- bryum pyriforme. Haplocladium microphyllum (Hedw.) Broth. (2, 3, 6, 7). Almost all collections on rotten wood but one collection (7) on humus. 1974] Bryophytes — Stringer & Stringer 33 Hedwigia ciliata (Hedw.) P. Beauv. (7). Several collec« tions, all on humus over rock or loose conifer needles over rock. Hylocomium splendens (Hedw.) B. S. G. (4, 6, 7). On litter and loose humus, often with Pleurozium schreberi. Hypnum lindbergii Mitt. (4, 5, 6). Usually on wet, well- rotted wood or wet humus. Hypnum pallescens (Hedw.) P. Beauv. (3, 5, 6, 7). Com- mon on live and fallen bark of both conifers and hárd- woods. Often associated with Pylaisiella polyantha. Hypnum pratense Koch ex Spruce (1). One collection, on wet rotten bark with Drepanocladus aduncus var. poly- carpus. Isopterygium turfaceum (Lindb.) Lindb. (7). One collec- tion, on thick humus over granite rock, with Pohlia nu- tans and Brachythecium rutabulum. Leptobryum pyriforme (Hedw.) Wils. (1, 2, 4, 7). Com- mon on a wide variety of substrates and with a wide range of associated species. Leptodictyum riparium (Hedw.) Warnst. (1, 2). On wet humus or litter, often with Drepanocladus aduncus or Amblystegium varium. Leptodictyum trichopodium (Schultz) Warnst. var. kochii (B. S. G.) Broth. (1, 2). On deadfall or humus in moist habitats. Leskea polycarpa Hedw. (1, 2,5). On bark or humus, found most often with Amblystegium varium or Leptodictyum trichopodium var. kochii. Mnium affine Bland. ex Funck (4, 6). On wet humus or disintegrated rotten wood. Mnium cuspidatum Hedw. (1, 2, 3, 5, 6). On humus, rotten wood or litter in drier habitats than M. affine, often alone, or with Brachythecium salebrosum. Myurella julacea (Schwaegr.) B. S. G. (4). Common in wet habitats in this collecting area and often found with Campylium stellatum and Aulacomnium palustre. Orthotrichum obtusifolium Brid. (3). One collection, on poplar bark, with Hypnum pallescens. 34 Rhodora [Vol. 76 Orthotrichum speciosum Nees ex Sturm (3, 7). On live poplar bark, often with Pylaisiella polyantha, but per- sisting on deadfall even in advanced stages of decay where it often occurs with Haplocladium microphyllum. Platydictya jungermannioides (Brid.) Crum (4). Fairly common in this collecting area, with Myuwrella and Dis- tichium. Recorded only once previously for the province, as Amblystegiella sprucei (Bruch) Loeske, by Crum and Schofield (1959). Pleurozium schreberi (Brid.) Mitt. (3, 4, 5, 6, 7). Com- mon, and often forming large pure colonies in both wet and dry coniferous forests. Pohlia nutans (Hedw.) Lindb. (5, 6, 7). Usually on well- rotted wood but sometimes (7) on humus. Polytrichum juniperinum Hedw. (5, 6, 7). Usually on hu- mus over sand in drier habitats. Polytrichum piliferum Hedw. (7). One collection on humus over sand. Ptilium crista-castrensis (Hedw.) De Not. (5, 7). Usually a forest floor species of dry coniferous forests in the area, occurring on loose humus or needle litter over sand. Pylaisiella polyantha (Hedw.) Grout (2, 3, 5, 6, 7). Com- mon with Orthotrichum speciosum, especially on bark of live hardwoods, but also on humus (6, 7) or at the base of jack pine (5). Tetraphis pellucida Hedw. (6). On well-rotted wood, with Pohlia nutans and Drepanocladus uncinatus. Thuidium delicatulum (Hedw.) B. S. G. var. radicans Crum, Steere & Anderson (4). On loose wet humus and litter with Hypnum lindbergii, Campylium stellatum and Tomenthypnum nitens. Thuidium recognitum (Hedw.) Lindb. (4, 6). On loose hu- mus and needle litter in wet or moist habitats. Tomenthypnum nitens (Hedw.) Loeske (4). On loose, wet humus and litter. Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. (7). On thin humus over sand. Stringer & Stringer 35 1974] Bryophytes DISCUSSION A search of the literature reveals that little work has been done on the bryophytes of Manitoba. Bird (1966), who gives a comprehensive list of the recordings for the province, lists only 289 taxa of bryophytes for the whole of Manitoba. This list has been somewhat augmented by Bird (1969) and Longton (1972), but published work on the bryophyte flora of the province is still greatly lacking. This situation is especially unfortunate as Manitoba is particu- larly interesting from a phytogeographic point of view, ly- ing as it does at a “crossroads of floral elements, from both north and south, east and west" (Lóve, 1959). Approximately half of the species recorded in the present study are pan-North American in distribution, according to the designations given by Bird (1969) and Bird and Won Shic Hong (1969). However, the Boreal element is repre- sented by species such as Dicranum fuscescens, D. polyse- twm, Tetraphis pellucida, Myurella julacea, Thuidium deli- catulum var. radicans, Thuidium recognitum, Pleurozium schreberi, Ptilium crista-castrensis, and Pylaisiella poly- antha. The Arctic Boreal element is represented by Di- cranum | bonjeanüi, Bryoerythrophyllum | recurvirostrum, Bryum pallescens, Orthotrichwm speciosum, Calliergon gi- ganteum, Campylium stellatum, Eurhynchium pulchellum, Tomenthypnum nitens, Hylocomium splendens, Polytrichum juniperinum, and Polytrichum piliferum. A few Eastern Boreal species are also present, i.e. Dicranum drummondii, Leptodictyum trichopodium var. kochii and Callicladium haldanianum. The hepatics are largely pan-North American species. Cephaloziella rubella, however, is Boreal and Ptilidium ciliare is Arctic Boreal (Bird & Won Shic Hong, 1969). Calypogeia . muelleriana and Cephalozia media have been previously recorded only from the north-east part of the province, from Churchill and York Factory respectively (Longton, 1972). The only previous record of Cephaloziella rubella was from Spruce Woods Provincial Park ( Stringer 36 Rhodora [Vol. 76 & Stringer, in preparation). Lophocolea minor, recorded from Grand Beach as new to the province, has since been found at several stations in Birds Hill and Whiteshell Pro- vineial Parks, where Radula complanata is also quite fre- quent. Riccardia pinguis has only two previous records, both from Spruce Woods Provincial Park, although subse- quent work by the authors shows that it is common on the calcareous clays of the southern part of the province. All of these liverworts are probably much more widely dis- tributed throughout the province than the scant records indicate. Several moss species which, according to the literature, should have a wide distribution have few records in Mani- toba. This is especially true of Brachythecium spp., per- haps because of the apparent scarcity of fertile material and the consequent difficulty in making a positive identifi- cation. Brachythecium rutabulum, common in the Grand Beach aréa studied, was reported as new to the province by Longton (1972), although Mueller-Dombois (1964) de- scribed it as one of the mosses forming the typically con- tinuous carpet in one of the jack pine — black spruce forest types in southeastern Manitoba. The only other records of Brachythecium campestre are also from the southern part of the province (Stringer & Stringer, in preparation, 1973) although this species appears to have a pan-North Ameri- can distribution. Callicladium haldanianum, reported for the first time in the province by Bird (1969) as Heterophyllium haldani- anum (Grev.) Kindb., reaches its western limit in Mani- toba. The Grand Beach record is further north than either of the Bird records. Fissidens adianthoides, a new record for Manitoba, ap- pears to be rare in western Canada (Bird 1966), although Grout (1936) gives its distribution as from Cape Breton Island to Vancouver Island and south to California and Florida. It is intended that the information presented in this study should, in conjunction with information presented in 1974] Bryophytes — Stringer & Stringer 37 further papers in this series, assist in describing the bryo- phyte flora of Manitoba and its relationships with the floras of adjacent regions. ACKNOWLEDGEMENTS The field work was supported in part by Grant No. 140- 118 from the University of Winnipeg. The authors also wish to express their gratitude to Dr. Dale H. Vitt, Univer- sity of Alberta, for his prompt assistance in identification of some of the Brachythecium material. LITERATURE CITED Birp, C. D. 1966. A catalogue of the bryophytes reported from Alberta, Saskatchewan and Manitoba. Processed publication, University of Calgary, Calgary, Alberta. . 1969. Bryophytes of the aspen parkland of west- central Canada. Can. Jour. Bot. 47: 187-212. . & Won Suic Hone. 1969. Hepaticae of southwestern Alberta, Can. Jour. Bot. 47: 1727-1746. Crum, H. 1971. Nomenclatural changes in the Musci. The Bryol- ogist 74: 165-174. . & W. B. SCHOFIELD. 1959. The mosses of Gillam, Mani- toba. Bull. No. 160, Nat. Mus. Can., Ottawa, .„ W. C. STEERE & L. E. ANDERSON. 1965. A list of the mosses of North America. The Bryologist 68: 377-434. Grout, A. J. 1936. Moss flora of North America. Vol. I, Part I. Newfane, Vermont. Published by the author. LoNGTON, R. E. 1972. Nineteen bryophytes new to Manitoba. The Bryologist 75: 348-350. LóvE, D. 1959. The postglacial development of the flora of Mani- toba: a discussion. Can. Jour. Bot. 37: 547-585. MUELLER-DoMBors, D. 1964. The forest habitat types in south- eastern Manitoba and their application to forest management. Can. Jour. Bot. 42: 1417-1444. RowE, J. S. 1959. The forest regions of Canada. Bull. 125, Dept. Northern Affairs and Nat. Resources, Forestry Branch. Ottawa. SAYRE, G., C. E. B. BONNER, & W. L. CULBERSON. 1964. The authori- ties for the epithets of mosses, hepatics, and liehens. The Bry- ologist 67: 113-135. ScHUSTER, R. M. 1969. The Hepaticae and Anthocerotae of North America east of the hundredth meridian. Vol. II. 1062 pp. Colum- bia University Press, New York. 38 Rhodora [Vol. 76 1953. Boreal Hepaticae, a manual of the liver- worts of Minnesota and adjacent regions. Amer. Midland. Nat. 49(2) : i-v, 257-684, SCOGGAN, H. J. 1957. Flora of Manitoba. Bull. 140, Biol. Ser. No. 47, Nat. Mus. Can., Ottawa. STRINGER, M. H. L. & P. W. STRINGER. Studies on the bryophytes of southern Manitoba. I. Collections from Spruce Woods Pro- vincial Park, In preparation. & . 1973. Studies on the bryo- phytes of southern Manitoba. II. Collections from the Winnepeg area. Can. Field Nat. 87: 141-144. DEPARTMENT OF BIOLOGY UNIVERSITY OF WINNIPEG 515 PORTAGE AVENUE WINNIPEG 2, MANITOBA, CANADA SOME NEW OR NOTEWORTHY VASCULAR PLANT RECORDS FROM NORTHWESTERN SASKATCHEWAN VERNON L. HARMS AND JOHN H. HUDSON During the course of recent botanical studies in north- western Saskatchewan, several new or otherwise interest- ing plant records were collected. Most of these were col- lected from along the Green Lake-La Loche Road during a study supported by the Institute for Northern Studies, Uni- versity of Saskatchewan, or from the Lake Athabasca area. The purpose of this paper is to comment on these records in order to make this information generally available to taxonomists and phytogeographers. The collection numbers are those of the senior author unless otherwise indicated. All specimens cited have been deposited in the Fraser Her- barium, University of Saskatchewan (SASK). SPARGANIUM CHLOROCARPUM Rydb. 7 mi. s.-s.e. of La Loche, Mile 60 Buffalo Narrows-La Loche Road, Harms 17635; McAneely Creek, 2 mi. s.w. of Turnor Lake village, Harms 18390; Bear Creek, Mile 41 Buffalo Narrows-La Loche Road, Harms 19661. This species has never been listed for Saskatchewan by Fraser and Russell (1937), Fraser and Russel] (revised by Russell, Ledingham and Coupland (1954)), or Breitung (1957). The specimens of the species were for the most part confused with Sparganium multipedunculatum. While a taxonomic problem exists between the two taxa, and even- ually both of them, together with S. angustifolium, may be merged as varieties under an enlarged S. simplex, there is no question that many of our specimens best fit S. chloro- carpum as it is now recognized. Other specimens in the Fraser Herbarium now identified as S. chlorocarpum in- clude the following: Melfort, J. B. Millar 64-163, 64-12; Saskatoon, J. H. Hudson 2488, 2548, J. B. Millar 65-217, W. P. Fraser 1930; 39 40 Rhodora [Vol. 76 Pike Lake, 20 mi. s. of Saskatoon; W. P. Fraser 1 Sept. 1937, R. C. Russell, et al., 4 August 1937; St. Denis, J. B. Millar 62-72; W. of Whitefox, W. P. Fraser & G. F. Leding- ham 28 July 1936; Swift Current, J. L. Bolton 8 July 1936; Dahlton, W. P. Fraser and. G. F. Ledingham 20 July 1937; Egg Creek, s. of Cumberland House, G. H. Townsend 55; Waskesiu, W. P. Fraser 24 July 1938; Lake Waskesiu Road, W. P. Fraser July 1932; Mile 23 Waskesiu Road to Prince Albert, W. P. Fraser 24 August 1934; Meadow Lake Pro- vincial Park, Waterhen River, J. H. Hudson 2427; Sylvan Lake, Candle Lake Road, J. K. Jeglum 18 July 1964; 1.7 mi. n. of forks at Candle Lake, J. K. Jeglum 9 July 1964. CALAMAGROSTIS LAPPONICA (Wahl.) Hartm. 3 mi. s. of La Loche, Mile 64 Buffalo Narrows-La Loche Road, Harms 17532, 17565; 2 mi. s. of La Loche, Mile 65 Buffalo Nar- rows-La Loche Road, Harms 17547, 17543; Island in Wil- liam River, south shore Lake Athabasca, R. Hermesh 485; Thompson Bay, south shore of Lake Athabasca, R. Hermesh 582. This northern species has never been previously reported for Saskatchewan. There seems to be intergradation be- tween Calamagrostis lapponica and C. neglecta (Ehrh.) Gaertn. as indicated by an intermediate collection from Taylor Lake, Mile 19 Buffalo Narrows-LaLoche Road (18160), and between C. lapponica and both C. neglecta and C. inexpansa A. Gray as indicated by intermediate specimens from the same area (18142, 18147). The whole C. lapponica, C. neglecta, and C. inexpansa complex appears in need of careful taxonomic study. CAREX ADUSTA Boott. Turnor Lake, Harms 18364-B, 18361-A. In the Fraser Herbarium, there are three specimens from the Waskesiu area or the road to Montreal Lake, Prince Albert National Park (W. P. Fraser July 8, 1939, June 25, 1940, June 22, 1941). This species has been collected by J. H. Hudson at Flin Flon, Manitoba, near the Saskatche- wan border. Breitung (1957) reports it also from Meadow 1974] Saskatchewan Plants — Harms & Hudson 41 Lake but a substantiating voucher is missing from the Fraser Herbarium. The above record represents a con- siderable northwestward range extension in Saskatchewan. CAREX RUFINA Drej. Beach at Thomson Bay, south shore Lake Athabasca, R. Hermesh 421. This represents the first report for this species in Sas- katchewan. Characteristic material of this species comes from Greenland and Iceland. Scoggan (1957, p. 179) re- lates that A. E. Porsild collected typical material from southern Keewatin, Northwest Territories, while Baldwin found abberrant material at Nueltin Lake, northern Mani- toba, about 125 miles south of Porsild's station. Porsild has annotated Baldwin’s sheet as follows: “Differs from typical C. rufina by paler scales having more prominent midveins, paler and more prominently nerved perigynia, and by terminal spikelets being not so strictly gynaecan- drous (in some even androgynous). Perigynia only rarely with a few teeth in the upper part." The present Lake Athabasca material agrees well with the characters men- tioned in these statements. This collection represents about a 350 mile westward range extension for the species. HABENARIA ORBICULATA (Pursh) Torr. Lac La Plonge, 5 mi. e. of Beauval, Harms 17959-B. In the Fraser Herbarium there are specimens of this species from Big Sandy Lake, Hansen Lake Road (G. W. Argus 4237), from La Ronge (J. S. Maini 611), and from Pinkney Lake, 35 mi. n.e. of Candle Lake (J. M. A. Swan 65-45). Breitung (1957) also reports it from Torch River and Amisk Lake, the latter based on a collection by J. H. Hudson. The present record is the first one from the west- ern part of the province and represents a considerable west- ward extension of the main species range. It is disjunctly present in British Columbia. SPIRANTHES GRACILIS (Bigel) Beck. (S. lacera Raf.). Little Amyot Lake area, Mile 69 Green Lake-Buffalo Nar- rows Road, Harms 17736; 2 mi. e. of Beauval, Harms 42 Rhodora [Vol. 76 17776; Taylor Lake, Mile 19 Buffalo Narrows-La Loche Road, Harms 18139-B. There are specimens of this species in the Fraser Her- barium from Lake Waskesiu, Prince Albert National Park (Anonymous July 1932) and from 20 miles south of Mea- dow Lake (A. J. Breitung 8164). The above records am- plify the known range of the species and extend it to north- western Saskatchewan. SPERGULA ARVENSIS L. 2 mi. w. of Beauval, Harms 17846. Breitung (1957) does not report this plant for Saskatche- wan, and Scoggan (1957) does not report it for Manitoba. Boivin (1968) listed this species as doubtful for Saskatche- wan. Hudson (1972) reported it as new to Saskatchewan on the basis of a collection from Gronlid (B. Zuk 1971, DAO). Boivin, in personal correspondence to J. H. Hudson, indicated the presence of an earlier collection from Sylvania (K. Drake Juiy 26, 1966, DAS). Mrs. H. D. Bobier recently sent a collection of the species from Rapid View (west of Meadow Lake) to J. H. Hudson. A map by J. F. Alex indi- cates two infestations of this weed near Outlook, Saskatche- wan but these are not substantiated by voucher specimens. Thus the above record represents only the fourth report for Saskatchewan and the only one from this far northwest in the province. CALTHA NATANS Pallas. 7 mi. s.-s.e. of La Loche, Mile 60 Buffalo Narrows-La Loche Road, Harms 17633. There are no Saskatchewan specimens of this species in the Fraser Herbarium, Breitung (1957) reported it from Windrum Lake (56°02’N, 104°W) and Amisk Lake, the latter based on a collection by J. H. Hudson. The present collection presumably represents the third report for Sas- katchewan. RANUNCULUS HYPERBOREUS Rottb. Open beach, William's Point, south shore Lake Athabasca, R. Hermesh 415. This represents the first record of this northern butter- cup species in Saskatchewan. 1974] Saskatchewan Plants — Harms & Hudson 43 MYRIOPHYLLUM ALTERNIFLORUM DC. Little Amyot Lake, Mile 70 Green Lake-Buffalo Narrows Road. Breitung (1957) omits this species from his list of Sas- katchewan plants. Boivin (1968) reports only 3 collections from the prairie provinces: Cochrane River, Reindeer Lake, and Lake Axis. The present collection would there- fore represent only the second Saskatchewan record for the species. EUPHRASIA ARCTICA Lane var. DOLOSA Boivin (E. subarctica Raup, E. disjuncta Fern. & Wieg. var. dolosa Boivin). 2 mi. s. of La Loche, Mile 65 Buffalo Narrows-La Loche Road, Harms 17550. In the Fraser Herbarium, the only Saskatchewan speci- men of this species is from Stony Rapids, north of Fond-du- lac River (J. S. Maini & M. Swan Aug. 3, 1961). Breitung (1957) cites a Raup report from Lake Athabasca. The above record represents about a 200 mile southward range extension for the species in Saskatchewan. LOBELIA DORTMANNA L. Little Amyot Lake, Mile 70 Green Lake-La Loche Road, Harms 16972. In the Fraser Herbarium we have specimens of this spe- cies from Little Gull Lake, south shore Lake Athabasca (G. W. Argus 553-63) and Carswell Lake, 40 mi. south of Lake Athabasca (G. W. Argus 628-62). Breitung (1957) reports the species also from Windrum Lake, north of the Churchill River, 56°02’N, 104°W. Therefore, the above rec- ord represents the third general locality for the species in Saskatchewan. ADOXA MOSCHATELLINA L. 17 mi. n. of Meadow Lake, J. H. Hudson 2729 (SASK). Boivin (1972, p. 5) reports this species in Saskatchewan from Pasquia Hills and Candle Lake; the latter specimen, which he collected, was the first record for Saskatchewan. The present material is evidently the third record of the species for the province and represents more than a 100 mile westward range extension in the province. 44 Rhodora [Vol. 76 ANAPHALIS MARGARITACEA (L.) B. & H. 3 mi. s. of Buffalo Narrows, Mile 124 Green Lake-Buffalo Narrows Road, Harms 18244. There are various specimens of this species in the Fraser Herbarium from the Cypress Hills area of Saskachewan. Breitung (1957) reports it also from Cut Knife but a sub- stantiating voucher has not been found. Therefore the above record represents at least a 200 mile, if not a 300 mile northward range extension. BIDENS BECKII Torr. Little Amyot Lake, Mile 70 Green Lake-Buffalo Narrows Road, Harms 16978. There are two specimens of this species in the Fraser Herbarium from the Cumberland House area on the Sas- katchewan River (A. E. Etter Aug. 8, 1934; D. Dabbs 126- 66) in east-central Saskatchewan. The above report there- fore represents the second known Saskatchewan locality for the species and is a considerable westward range extension in the province. LITERATURE CITED BorviN, B. 1968. Flora of the Prairie Provinces, Part II. Reprinted from Phytologia 16-18. 1972. Flora of the Prairie Provinces, Part III. Re- printed from Phytologia 22-23. BREITUNG, A. J. 1957. Annotated Catalogue of the Vascular Flora of Saskatchewan. The American Midland Naturalist 58: 1-72. FRASER, W. P. & R. C. RUSSELL. 1937. An Annotated List of the Plants of Saskatchewan. University of Saskatchewan, Saska- toon. HUDSON, J. H. 1972. 1971 Botanical Records for Saskatchewan. The Blue Jay 30: 122-124. RUSSELL, R. C., G. F. LEDINGHAM, & R. T. COUPLAND. 1953. Re- vision of Fraser, W. P. and R. C. Russell, An Annotated List of the Plants of Saskatchewan. University of Saskatchewan, Saskatoon. SCOGGAN, H. J. 1957. Flora of Manitoba. National Museum of Canada Bulletin No. 140, Ottawa. FRASER HERBARIUM UNIVERSITY OF SASKATCHEWAN SASKATOON, SASKATCHEWAN, CANADA PARASITIC WITCHWEED: STRIGA ASIATICA VERSUS S. LUTEA (SCROPHULARIACEAE) F. N. HEPPER When plants of a witchweed were discovered in 1956 parasitizing Zea mays L. in North and South Carolina, specimens were sent to Kew for identification. I determined them as Striga asiatica (L.) O. Kuntze and communicated the name to Dr. S. F. Blake and other authorities in the United States, who used it until a paper by Dr. C. J. Saldanha (1963) proposed the rejection of that name in favour of S. lutea Lour. (Smith 1966). The purpose of this paper is to reinstate S. asiatica on the basis of typification of the Linnean basionym Buchnera asiatica, rather than on the opinions of other taxonomists. Linnaeus’ description of Buchnera asiatica in ‘Species Plantarum' (1753) is unusually full for that work and entirely devoid of any references to literature. This indi- cates that he had before him a specimen, or specimens, from which he drew up his description. Examination of his herbarium at the Linnean Society, London, revealed six sheets bearing the name ‘Buchnera asiatica! in Linnaeus’ hand and now numbered according to Savage's Catalogue as 790/10, 11, 12, 13, 14, & 15. Sheets 13, 14 and 15 can be discounted, since they are annotated as having been collected by Thunberg (13) some- time after 1772, and by Koenig (14 & 15) in 1777 according to the date on the sheet. They are referable to Striga bilabiata (Thunb.) O. Kuntze (syn. S. thunbergii Benth.) and S. euphrasioides Benth., respectively. Sheet 12 bears a single simple plant with pale flowers that would be known in India as Striga lutea. The only annotation on the sheet is ‘Indica,’ indicating its country of origin. The specimen does not fit Linnaeus' description, and it can be ruled out as the type. Five specimens are glued to sheet 10, including a well- branched plant with dark-coloured flowers that fits the 45 16 Rhodora [Vol. 76 original description admirably. Two other specimens match it except for their simple habit. The remaining two are only the inflorescences of Striga densiflora Benth. The specimen on sheet 11 is very similar to the large branched one on sheet 10, but there is no annotation apart from the name. On sheet 10, however, Linnaeus has added the locality “ins. Johan.," and although the ‘Species Plantarum’ cites the distribution as “Habitat in Zeylona, China” this reference to “ins. Johan.” provides a clear indication of the provenance and history of what must be regarded as the type material of Buchnera asiatica. Where, then, is “ins. Johan.” and who collected the specimen? Several voyages to China were undertaken before 1753. including one by Peter Osbeck who wrote an account of it in Swedish (1757) which was translated into German (1765) and from the German edition into English, and published in 1771 under the title ‘A voyage to China and the East Indies, together with a voyage to Suratte by Olof Toreen in a series of letters to Doctor Linnaeus.’ Toreen (or Torén), in fact, continued his voyage as far as China and it is his letters, and not Osbeck’s text, which are of particular interest since he tells (p.166) how, on 16 August 1750, he landed at the island of “St. Joanna” in the Comoros near Madagascar. It is evident that Linnaeus worked out the collections made by Toreen and Osbeck, attributing them al] to China in the appendix to Osbeck and Toreen’s travels entitled ‘Flora Sinensis.’ There, on p. 356, is listed Buchnera asiatica with the reference to ‘Species Plantarum.’ It is interesting to note that the same island was visited by Koenig in 1768 where he gathered the plant Linnaeus (1771) described as Baccharis arborea (which Brenan (1968) identified as Vernonia grandis (DC.) Humbert var. comorensis Humbert), yet Linnaeus gave the occurrence as the “East Indies.” Striga asiatica (L.) O. Kuntze, Rev. Gen. Pl, 2: 466. 1891. Type. Comoro Islands, St. Joanna, Toreen LINN. (Sav- age Cat. No. 790/10 partly). 1974] Striga — Hepper AT Buchnera asiatica L., Sp. Pl. 630. 1753. Striga lutea Lour., Fl. Cochinch. 22. 1740. Buchnera hirsuta Benth., Scroph. Ind. 41. 1835. Striga hirsuta, (Benth.) Benth. in Hook., Comp. Bot. Mag. 1: 363. 1836. Bentham's work was completed long before an interna- tional code of botanical nomenclature had been agreed, and certainly his concepts of typification were not in accord with modern attitudes. Hence, he coined the new names to replace Buchnera asiatica which seemed to him, as to Saldanha, to apply to a number of different species. The more so since he considered S. lutea Lour. to be a distinct species. Saldanha considered Striga asiatica and its basionym as nomena rejicienda under Art. 69 since “any attempt to decide a lectotype is bound to be arbitrary and open to question." On the contrary, this species is one of the few Linnean species that may be typified and localized with certainty. I am grateful to my colleague Mr. R. D. Meikle for his helpful advice and assistance. LITERATURE CITED BRENAN, J. P. M. 1968. The identity of Baccharis arborea L. Kew Bull. 21: 427. LiNNAEUS, C. 1753. Species Plantarum, 630. 1771. Mantissa Plantarum, 284. OsBECK, P. 1771. A voyage to China and the East Indies. (English ed.). SALDANHA, C. J. 1963. The genus Striga Lour. in Western India. Bull. Bot. Surv. India 5 (1) : 67-70. SMITH, C. EARLE. 1966. Identity of witchweed in the southeastern United States. Rhodora 68 (774) : 167. THE HERBARIUM ROYAL BOTANIC GARDENS KEW, ENGLAND CACALIA RUGELIA: A NEW COMBINATION FOR A NORTH AMERICAN SENECIONOID* T. M. BARKLEY AND ARTHUR CRONQUIST The problematic taxon long-called Senecio rugelia (Shuttl. ex Chapman) A. Gray (Compositae) is a distinctive species occurring in coniferous associations along the higher ridges of the Great Smokies astride the North Carolina-Tennessee border. It is a tall, coarse herb with several conspicuous discoid heads and dull, dirty-white corollas. The species was introduced to science as Rugelia nudicaulis Shuttl. ex Chapman in 1860, but in 1883 Asa Gray transferred it to Senecio, where it has remained. Many botanists have recognized the incongruity of in- cluding this entity in Senecio, but the morphological toler- ances of Senecio have been usually treated as very wide, and so the matter has not been pursued. However, Green- man (1901) and Alexander (1937) noted the possible relationship to Cacalia. The species referable to Cacalia have had their systematic woes, being variously regarded as constituting several gen- era, or as members of the single genus Cacalia, or as a part of Senecio. Compounding these difficulties has been a dis- agreement concerning the typification of the name Cacalia. A review of these matters is beyond the scope of this paper, but we are treating Cacalia in the expanded sense, and we accept Cacalia hastata L. as the generic type (Vuilleumier & Wood, 1969). In general, Cacalia differs from Senecio by the two floral features of discoid heads and white to creamy corollas. Both are features which occur independently as evolution- ary tendencies in other alliances within Senecio. In Cacalia, however, the two occur together, presumably defining a natural assemblage. *Contribution no. 1193. Division of Biology, Kansas Agr. Expt. Station, Manhattan. Financial support from the National Science Foundation is gratefully acknowledged. 48 1974] Caealia — Barkley and Cronquist 49 Cacalia is here regarded as a derivative of Senecio section Palmatinervii of Mexico, a group of coarse herbs strongly resembling Cacalia in aspect. Nearly all species of the Palmatinervii for which chromosome counts are reported have n=30 (Gibson, 1969). Cacalia has n—30, and a re- duction series of n=28, 27, 26, 25, and 20 (Ornduff et al., 1963, 1967). The species here offered as Cacalia rugelia has »—28, a number unreported for Senecio (Fedorov, 1969). In a recent study of Mexican cacalioids, Pippen (1968) recognizes four genera, but he offers a table summarizing the distinctions between these ‘“‘Cacalioid genera” and Sene- cio (table 2, p. 371). Incorporated into the table are the cacalioid features of discoid heads and white or creamy florets. The table also shows that the *Cacalioid genera" characteristically have deeply lobed disk corollas, a feature which we find in Cacalia rugelia as well. Furthermore, in Pippen's key the present species falls into the assemblage treated as the Mexican segregate-genus Odontotrichum, and it is morphologically compatible with that group. The floristic relationships between the Southern Appalachian region and some of the mountains of Mexico have been noted by other authors (e.g. Sharp, 1946). In the light of the foregoing considerations, the following transfer is proposed: Cacalia rugelia (Shuttl. ex Chapman) Barkley & Cronquist, comb. nov. Type: “Smoky Mountains, Tennessee. Rugel, Buckley." Paratype: Buckley, NY! Senecio rugelia A. Gray, Proc. Am. Acad. 19: 54. 1885. Rugelia nudicaulis Shuttleworth ex Chapman, FI. S. U. S. 246. 1860. non Cacalia nudicaulis (Less.) O. Kuntze, 1891. non Senecio nudicaulis Buck.-Ham. ex D. Don, 1825. LITERATURE CITED ALEXANDER, E. J. 1937. Senecio rugelia. Addisonia 20: 29-30, pl. 655. 50 Rhodora [Vol. 76 FEDOROV, AN. A., ed. 1969. Chromosome numbers of flowering plants. Soviet Sciences Press. Leningrad. GIBSON, E. S. 1969. A revision of the section Palmatinervii of the genus Senecio (Compositae) and its allies. Unpub. doctoral dis- sertation, Kansas State Univ. Manhattan. pp. 1-178 + VI. GREENMAN, J. M. 1901. Monogr. nord-und centralam. Arten der Gattung Senecio. Leipzig. (Reprinted in Englers Bot. Jahrb. 32: 1-33. 1902.) ORNDUFF, R. P. H. Raven, D. W. KvHos & A. R. KINCHEBERG. 1963. Chromosome numbers in Compositae III. Senecioneae. Am. Jour. Bot. 50: 131-139. OnNDUFF, R., T. Mosquin, D. W. KvHos & P. H. RAVEN. 1967. Chromosome numbers in Compositae VI. Senecioneae II. Am. Jour. Bot. 54: 205-213. PIPPEN, R. W. 1968. Mexican “Cacalioid” genera allied to Senecio (Compositae). Contr. U. S. Natl. Herb. 34: 365-447. SHARP, A. J. 1946. Informe preliminar sobre alqunos estudios fitogeograficos efectuados en Mexico y Guatemala. Rev. Soc. Mexicana Hist. Nat. 7: 35-39. figs. 1-5. VUILLEUMIER, B. S. & C. E. Woop. 1969. Lectotypification of Caca- lia L. (Compositae-Senecioneae). Jour. Arnold Arb. 50: 268-275. HERBARIUM, DIVISION OF BIOLOGY, KANSAS STATE UNIVERSITY, MANHATTAN, KANSAS THE NEW YORK BOTANICAL GARDEN, BRONX, NEW YORK TYPIFICATION AND APPLICATION OF THE NAMES SCIRPUS AMERICANUS PERS., S. OLNEYI GRAY, AND S. PUNGENS VAHL ALFRED E. SCHUYLER The names Scirpus americanus Pers. (Syn. Pl. 1: 68. 1805) and Scirpus pungens Vahl (Enum. Pl. 2: 255. 1805) are usually applied to the same species by Cyperologists (Beetle, 1947; Koyama, 1963). The former name for plants commonly known as Three-square in North America pres- ently receives widespread usage because of priority. Data from type specimens and original descriptions now make it apparent that these names should be applied to different species. The type of S. americanus (Hab. in Carolina in- feriore, Herb. Michx., P) is conspecific with plants usually treated (Beetle, 1947; Koyama, 1963) as S. olneyi Gray (Bost. Journ. Nat. Hist. 5: 238. 1845). The type of S. pungens (Erhart misit, Herb. Vahl, C) is conspecific with plants usually treated (op. cit.) as S. americanus. In his original description of Scirpus americanus, Per- soon cites Michaux's description of S. triqueter L. (Fl. Bor.- Am. 1: 30. 1803.) and gives the habitat as "in Carolina inferiore." The specimens on the sheet thus named and labeled in Michaux's herbarium at P have strongly winged culms, short involucral bracts, shallowly cleft scales, small achenes, and broad bristles characteristic of plants resem- bling the type of S. olneyi (Rhode Island, Providence, Olney, GH). Thus the correct name for plants usually treated as S. olneyi is S. americanus. In the original description of Scirpus pungens, Martin Vahl gave the habitat as, *in Europa, in insula Borboniae. Commerson." The specimen from Vahl's herbarium desig- nated as the type probably was collected in Europe and also contains the word “pungens Vahl" presumably in the hand- writing of Jens Vahl, son of Martin Vahl. No specimen labeled, *in insula Borboniae," or collected by Commerson has been located. The triangular culms, long involucral 51 52 Rhodora [Vol. 76 bracts, deeply cleft scales, and narrow bristles are char- acteristics of plants usually but incorrectly treated as S. americanus. Scirpus pungens is the earliest available name for these plants. The literature cited by Vahl is also in accordance with his description and the designated type specimen. The cited illustrations of Morrison (Pl. Hist. Univ. Oxon. s. 8, t. 10, f. 20. 1699.) and Plukenet (Algm. Bot. t. 40, f. 1. 1696.) closely resemble the type specimen of Scirpus pungens. Roth's descriptions of S. mucronatus L. (Tent. Fl. Germ. 1: 28. 1788, and 2: 60. 1789.) and S. triqueter L. (Neue Beytr. 1: 91. 1802.) cited by Vahl best apply to plants resembling the type specimen of S. pungens. Apparently Roth was confused about the identity of S. mucronatus and S. triqueter, and incorrectly applied the names to plants of S. pungens. ACKNOWLEDGMENTS I am grateful to H. Heine (P), J. Raynal (P), and A. Skovsted (C) for courtesies enabling me to examine critical specimens and data at their respective institutions, and to John Braxton (PH) and Helena Greenwood (PH) for their assistance. Abbreviations of institutions are those given by Lanjouw and Stafleu (1964). LITERATURE CITED BEETLE, A. A. 1947. Scirpus. N. Am. Fl. 18: 481-504. KovAMA, T. 1963. The genus Scirpus Linn., critical species of the section Pterolepis. Canad. Jour. Bot. 41: 1107-1131. LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum. Part 1. The herbaria of the world, International Association for Plant Taxonomy, Utrecht. 251 pp. DEPARTMENT OF BOTANY ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA PHILADELPHIA, PENNSYLVANIA 19103 THE CORRECT AUTHORITY FOR CARDAMINE CLEMATITIS (CRUCIFERAE) T. R. DUDLEY! The most recent entry in the literature of Cardamine clematitis appears on page 508 of the Manual of the Vas- cular Flora of the Carolinas by Radford, Ahles and Bell (1968). They cite “Shuttlew.” as the authority for this taxon. This authority refers to Robert James Shuttle- worth, a famed naturalist and collector who was the spon- sor of Ferdinand Rugel, a prolific plant collector in eastern North America from 1840 and for many years thereafter until his death in 1879. The various references in the literature to Cardamine clematitis have consistently overlooked the fact that Shut- tleworth cannot be cited as the sole publishing authority for the specific epithet, clematitis. In addition to Radford, Ahles and Bell (1968), several other earlier North Ameri- can references also provided descriptions and occasionally illustrations of C. clematitis, and all accepted Shuttleworth as the sole publishing authority. These include: Chapman, Flora of the Southeastern United States, ed. 2, Supplement p. 605. 1887; ibid., ed. 3, p. 25. 1897; Britton & Brown, Illust. Flora of the Northern United States and Canada, ed. 1. 2: 130. fig. 1730. 1897; ibid., ed. 2. 2: 185. fig. 2088. 1913; Small, Flora of the Southeastern United. States, ed. 1, p. 482. 1903; ibid., ed. 2, p. 568. 1933; Fernald in Gray's Manual of Botany, p. 721. 1950; Gleason and Cronquist in Britton and Brown, Illust. Flora of the Northeastern United States and Adjacent Canada 2: 230, fig. p. 231. 1952; Gleason & Cronquist, Manual of Vascular Plants of North- eastern United States and Adjacent Canada, p. 340. 1963; and Radford, Ahles and Bell, Guide to the Vascular Flora of the Carolinas, p. 173. 1964. ‘Research Botanist, Herbarium, U.S. National Arboretum, Agricul- tural Research Service, Northeastern Region, U.S. Department of Agriculture, Washington, D.C. 20002. 53 54 Rhodora [Vol. 76 In the course of evaluating the Ferdinand Rugel collec- tions in the Isaac C. Martindale Herbarium at the U.S. National Arboretum, F. G. Meyer called my attention to an unidentified specimen of Cardamine collected by Rugel and accompanied by a signed holograph label that reads “Top of Smoky Mts. on the first Branch Nord Carolina Side. May 1866." Although this Rugel specimen clearly repre- sents C. clematitis, it is not a part of the type. The first published reference to Cardamine clematitis is on page 53 of Sereno Watson's Bibliographic Index to North American Botany (Smithsonian Miscellaneous Col- lections No, 258, 1878) that reads as follows: “C. clematitis Shuttl. in Herb. Gray, ined." It is evident that Watson took the name from Shuttleworth’s printed herbarium exsiccata label which reads “Cardamine clematitis Shuttle. n. sp.” and that name was used only as a nomen nudum. Watson’s reference does not incorporate a validating description. On this basis alone, clematitis cannot be used as a valid specific epithet. It is listed, however, in Index Kewensis 1: 421. 1895 as: “Clematitis Shuttle. ex S. Watson" ! Later, in a paper by Asa Gray entitled “Some New North American Genera, Species, &c." (Proc, Am. Acad. Arts Sci. n.s., 7:45. 1880, a valid Latin description is pro- vided. Gray precisely identified the plant as “Cardamine clematitis Shuttleworth in coll. distrib. Rugel," and explains that the original collection was from “wet ground along streamlets in the higher Iron or Smoky Mountains of North Carolina and Tennessee, collected in 1844 by Rugel .. .” When dealing with nomenclatural problems of this na- ture, we are guided by Article 46 and its accompanying recommendations (particularly Recommendation 46C) in the International Code of Botanical Nomenclature, p. 46. 1972. 'The correct authority citation for Cardamine clema- titis is C. clematitis Shuttleworth ex A. Gray, although it may be shortened, particularly in floristic treatments, to cite only the publishing author. Under no circumstances can Shuttleworth be assigned solitary authorship. The cor- rect citation of the name is: 1974] Cardamine clematitis — Dudley 55 Cardamine clematitis Shuttleworth ex A. Gray, Proc. Amer. Acad. Arts Sci, new series, 7: 45. 1880 — non Shuttl. ex Watson, Bibliog. Index North American Bot., p. 53, 1878. The original collection and designated type of Cardamine clematitis is Rugel No. 19 collected in 1844 in the Smoky Mts. of Tennessee. The holotype is deposited at GH, and cited duplicates (isotypes) are to be found at G-Herb. Bois- sier and W. TYPE: Tennessee, Smoky Mountains [‘‘in locis humidis et ad regionis super. montium Smoky Ms., Tennessee, Mai 1884.”] May 1844, Ferdinand Rugel No. 19 (holotype, GH; isotypes BM, G, W). The most complete set of Rugel’s material, purchased from Shuttleworth, is at the British Museum; however, reference to the existence of such a specimen has not previously ap- peared in the literature. N. K. B. Robson of the British Museum (Natural History) assures me that a duplicate is retained at BM. The one Rugel collection of Cardamine clematitis not previously reported in the literature is a specimen collected, probably a unicate, after Rugel’s professional contacts with Shuttleworth had ceased. The specimen was maintained in Rugel’s personal herbarium until that herbarium was pur- chased in 1881 from Rugel’s heirs by Isaac C. Martindale: North Carolina: “Top of Smoky Mts. on first Branch, Nord Carolina Side", Ferdinand Rugel, May 1866 (NA — from herbarium of Isaac C. Martindale). The astute monographer of Cardamine and indefatigable expert on the Cruciferae, D. E. Schulz, in Monographie der Gattung Cardamine” (Bot. Jahrb. 32: 440. 1903.) recog- nized Shuttleworth as responsible for the epithet clematitis, but also recorded that Asa Gray provided a description of the species in Proc. Am. Acad. Arts Sci. n.s. 7: 45. 1880. Schulz examined the original 1844 material of C. clematitis collected by Rugel (GH, G & w — but not BM), which was annotated and distributed by Shuttleworth; he also cited numerous additional collections made by J. K. Small, N. L. 56 Rhodora [Vol. 76 & E. G. Britton & A. M. Vail, S. B. Buckley, J. K. Small & A. A. Heller, W. M. Canby, M. E. Hyams and J. W. Chicker- Ing. SYNONYMS OF CARDAMINE CLEMATITIS A point should also be discussed that concerns the synon- omy of Cardamine clematitis Shuttleworth ex A. Gray as presented by the Radford, Ahles and Bell publications (1964 & 1968). These works refer “C. flagellaris” (= C. flagellifera Schulz, Bot. Jahrb. 32: 405. 1903.) into synon- omy under C. clematitis. Although the type specimen of C. flagellifera (Biltmore Herbarium 7756) was originally annotated as C. clematitis, it is not conspecific with C. clematitis. Not only does C. flagellifera stand morphologi- cally and ecologically distinct from C. clematitis, but it was also assigned by Schulz within Cardamine to Sect. Macro- phyllum, whereas C. clematitis was referred by Schulz to the typical section. Small (1903) described C. hugeri that Radford, Ahles & Bell (1964 and 1968) sank under C. clematitis. However, if referrable into synonomy at all, C. hugeri, a low altitude plant, more logically pertains to C. flagellifera. In Watson’s reference (loc. cit.) to Cardamine clematitis, he cites as a synonym a “Nasturtium officinale” that ap- peared in the Supplement to Torrey and Gray, A Flora of North America 1: 666. 1843. In addition to validating, describing and typifying C. clematitis, Gray (loc. cit.) explains Watson’s confusion in incorrectly citing this “Nasturtium officinale” as a synonym of C. clematitis. The original Rugel material, annotated and distributed from Switzerland by Shuttleworth, was a mixed collection. A part representing C. clematitis “was mixed up with a Flor- ida species intermediate between Cardamine and Nastur- tium, first received from Leavenworth without fruit, and referred in the supplement to the first volume of Torrey and Gray's Flora to N. officinale.” This element of “Nasturtium officinale”, according to Gray, was later received from S. B. 1974] Digitaria sanguinalis — Ebinger 57 Buckley; again later received from Shuttleworth’s dis- tributed collections of Rugel as Cardamine curvisiliqua Shuttleworth; and yet again received from Shuttleworth as Nasturtium stylosum Shuttleworth! The current fate of Leavenworth’s “Nasturtium officinale’ is not within the scope of this paper; however, both Chapman (1887) and Small (1903) equate Leavenworth’s “Nasturtium officinale” with Cardamine curvisiliqua Shuttleworth. U.S. NATIONAL ARBORETUM WASHINGTON, D.C. 20002 A NEW FORM OF DIGITARIA SANGUINALIS. As a result of extensive field work in east-central Illinois an unusual form of the common crabgrass was found. This form differs from typical Digitaria sanguinalis (L.) Scop. in that long, spreading, papillose-based hairs are found scattered along both sides of the flattened rachis. These colorless hairs are 3-6 mm long (rarely 1 cm long) and on the specimens examined usually 1 to 5 hairs are found on each cm of rachis length. As a result of this difference the following form is described. Digitaria sanguinalis (L.) Scop. forma illinoensis Ebinger forma nova. A forma sanguinalis differt pilis base papillosis in rhachidi. TYPE: ILLINOIS: DOUGLAS CO.: 4 miles east of Hindsboro, Sargent Twp. (NE!4, Sect. 3, RIOE, T14N), in open field at edge of road, J. E. Ebinger 6845 (EIU). This area was revisited on 9 October 1969 and a second collection (J. E. Ebinger 9282) was made at that time (EIU, ISM). Long, papillose-based hairs are known in a few species of Digitaria. The presence of these hairs is an important diagnostic characteristic used by Hitchcock (1935), Hen- rard (1950), and many others to separate the tropical Digi- taria horizontalis Willd. from other members of the genus. The hairs in this species are similar in all respects to those 58 Rhodora [Vol. 76 found in D. sanguinalis f. illinoensis. The two taxa are easily separated however, since in D. horizontalis the nearly glabrous spikelets are usually less than 2.4 mm long, ex- tremely narrow and with an acuminate apex while D. san- guinalis has pubescent spikelets that are longer (2.5-3.5 mm long) and broader and with an acute apex. The only other closely related taxon in which these hairs have been re- ported is D. adscendens (HBK) Henrard var. rhachiseta Henrard. The differences between this taxon and D. san- guinalis have been studied by Ebinger (1962, 1965). LITERATURE CITED EBINGER, J. E. 1962. Validity of the grass species Digitaria adscendens. Brittonia 14: 248-253. . 1965. Digitaria sanguinalis in South America. Trans. Ill. St. Acad. Sei. 58: 255-258. HENRARD, J. T. 1950. Monograph of the genus Digitaria. xii + 999 pp. Leyden, Universitare Pers Leiden. Hitcucock, A. S. 1935. Manual of the grasses of the United States. U. S. Dept. of Agric. Misc. Publ. 200. 1040 pp. (Ed. 2, revised by Agnes Chase, 1950). JOHN E. EBINGER EASTERN ILLINOIS UNIVERSITY CHARLESTON, ILLINOIS THE GROWTH OF SOME NEW ENGLAND PERENNIAL SEAWEEDS NORMAN B. REYNOLDS Growth studies on Fucus vesiculosus L., Ascophyllum nodosum (L.) LeJolis, Rhodymenia palmata (L.) Greville, Laminaria digitata (L.) Lamouroux and Laminaria sac- charina (L.) Lamouroux were initiated at Dover Point, Dover, New Hampshire and Jaffrey Point (Fort Stark), Newcastle Island, Portsmouth, New Hampshire in July 1968. Dover Point is an estuarine tidal rapid area whereas Jaffrey Point is a semi-exposed open coastallocation. Both areas have been studied extensively and much is known of their flora and ecology (Mathieson, Hehre & Reynolds, in press; Mathieson, Reynolds & Hehre, in press; Reynolds, 1971). Approximately 20 plants of each species were tagged at both locations with plastic forester's tape and small white tags. The monthly growth of F. vesiculosus, A. nodosum and R. palmata was recorded by measuring their total length, while in L. digitata and L. saccharina growth was recorded by determining the monthly increase in distance between the intercalary meristem and paired holes (Sun- dene, 1964). The holes were made with a cork borer 1 inch above the meristem. New holes were made every 2 months, because the terminal portions of the lamina sloughed off continuously. Specimens of L. saccharina, L. digitata and R. palmata were transplanted from the high to the low current areas at Dover Point, and their growth and longevity was recorded. During the 16 months of the ex- periment, nearly all the initial plants were lost due to human interference, grazing of snails, wave action or degeneration. Consequently, it was necessary to continu- ously tag and measure new plants. The growth of Rhodymenia palmata was sporadic in the areas of low and high currents at Dover Point. Frag- mentation occurred extensively during the late winter, The 59 60 Rhodora [Vol. 76 area of low currents was not suitable for the sustained growth of R. palmata, for when plants were transplanted there they died in a few months. All specimens of R. palmata grew at least one inch during June 1968. The best growth of Fucus vesiculosus and Ascophyllum nodosum occurred during the summer while the poorest growth occurred during cold weather. The growth of F. vesiculosus and A. nodosum at Jaffrey Point was consistently lower than at Dover Point. The maximum growth of F. vesiculosus was 2 inches per month, and it occurred from May 15 to August 15, 1968, in the area of low cur- rents at Dover Point. Figures 1 and 2 show the growth of Laminaria digitata and L. saccharina. Because of the extreme variability in growth rates between individual plants, these rates are described only in relative terms as follows: (1) good growth — over 3 inches/month (2) average growth — 2-3 inches/month (3) poor growth — up to 2 inches/month. The growth of Laminaria digitata in the low current areas at Dover Point was quite variable. When trans- plants were made from the high to the low current areas, the growth was initially high, but the plants eventually degenerated. The probable cause is siltation and the physi- ological decrease of oxygen and nutrients due to the de- creased currents. Good growth of L. digitata occurred in the spring and summer at Jaffrey Point and in the area of high currents at Dover Point. Poor growth was particu- larly evident in the winter at the same locations. In general, L. digitata had a higher rate of growth and it was sustained for a longer time in the high current areas at Dover Point than at Jaffrey Point. An interesting phenomenon was observed when L. digi- tata was transplanted from the high to the low current areas at Dover Point. No dissection of the newly formed parts of the blades took place, and in a few months it was difficult to distinguish the plants from L. saccharina, ex- cept for the identification tags. The mechanical force of 1974] Seaweeds — Reynolds H è 5 3 B & 3 z £ 3 ii ii RELATIVE GROWTH OF LAMINARIA DIGITATA 61 1- 1- | ee s d ; v 2 o 3 E $ 3 $ $ i d oi i $ d H1MO39 Figure 1 RELATIVE GROWTH OF Laminaria digitata LEGEND: Poor Growth 0-2 inches per month Average Growth 2-3 inches per month Good Growth 38-54% inches per month poor 62 RELATIVE GROWTH OF LAMINARIA SACCHARINA Rhodora high current area low current area = à b [j] » o O Jaffrey tt Dover Pt. [Vol. T T T "CT | EN . , ¢ , i1 : i i | i o: OB HLMOYS Figure 2 RELATIVE GROWTH OF Laminaria saccharina LEGEND: Poor Growth 0-2 inches per month Average Growth 2-3 inches per month Good Growth 3-5 % inches per month 6 1974] Seaweed — Reynolds 63 the current was not high enough to cause the final separa- tion of the frond segments. The growth rate of L. saccharina was good for most of the year in the low current areas at Dover Point. A slight decrease in growth occurred during the winter. Good growth of L. saccharina occurred during the summer and early fall at Jaffrey Point and in the high current areas at Dover Point. Poor growth was evident during the late fall and early winter. Laminaria saccharina exhibited good growth at all areas during the summer. Laminaria saccharina dominated the kelp beds at Dover Point during the winter, but L. digitata was dominant the rest of the year. It can be concluded that L. saccharina is better adapted to the estuarine en- vironment than is L. digitata. The penetration of L. sac- charina into the estuary is dependent on the presence of currents to compensate for the reduced salinities. LITERATURE CITED MATHIESON, A. C., E. HEHRE & N. B. REYNOLDS. Investigations of New England marine algae I. A floristic and descriptive eco- logical study of the marine algae of Jaffrey Point, New Hamp- shire. Nova Hedwigia, In press. MATHIESON, A. C., N. B. REYNOLDS & E. HEHRE. Investigations of New England marine algae. II. Distribution of benthonic marine algae in the Great Bay Estuary System. Nova Hedwigia, In press. REYNOLDS, N. B. 1971. The ecology of a New Hampshire estuarine tidal rapid. Ph.D. Thesis, University of New Hampshire Li- brary, 101 pp. SUNDENE, O. 1964. The ecology of Laminaria digitata in Norway in view of transplant experiments. Nytt Mag. Bot. 11: 83-107. DEPARTMENT OF BIOLOGICAL SCIENCES STATE UNIVERSITY OF NEW YORK COLLEGE AT CORTLAND CORTLAND, NEW YORK 13045 CHROMOSOME NUMBERS IN KOSTELETZKYA PRESL (MALVACEAE) ORLAND J. BLANCHARD, JR. Heretofore, knowledge of the cytology of Kosteletzkya Presl has consisted of a single chromosome count from K. hastata Presl (Skovsted, 1941). This note corrects the number erroneously reported by Skovsted and adds counts from two African species, K. adoensis (Hochst. ex A. Rich.) Mast., and K. buettneri Gürke, and four American species, K. coulteri A. Gray, K. pentasperma (Bertero ex DC.) Griseb. K. paniculata Benth., and K. virginica (L.) Presl ex A. Gray (Table 1). Counts were made from pollen mother cells of buds col- lected in the wild or taken from plants grown from seeds collected in the wild. Methods of fixation, preparation, and documentation are those reported by Bates and Blanchard (1970). Voucher specimens have been deposited in the L. H. Bailey Hortorium (BH). The species counted represent a rather wide range of morphological diversity in the genus as well as a con- siderable geographical range. Kosteletzkya paniculata be- longs to the specialized sect. Orthopetalum Benth. of western Mexico, which is notable in its convolute, tubular corolla and its exserted staminal column. The remaining species, both African and American, belong to the wide- spread sect. Kosteletzkya, but within that section repre- sent a diversity of forms. The fact that counts made from all these species are the same, n = 19, suggests that the number in the genus may be constant, At variance with this conclusion is Skovsted's report of 2n = 34 in K. has- tata, a species closely related to K. pentasperma. How- ever, an examination of the illustration by Skovsted (1941, fig. 99) shows 38 chromosomes rather than the reported 34. It may therefore be assumed that the gametic chromo- some number of K. hastata is also » — 19. 64 1974] Kosteletzkya — Blanchard 65 The error that entered Skovsted's paper is of further interest because he used the erroneous count to support Hochreutiner's suggestion (1900) that Kosteletzkya could be related to Hibiscus sect. Pterocarpus Garcke ex Hochreutiner, a section known cytologically only from H. vitifolius L. with a chromosome number of n = 17 (Skovsted, 1935, 1941). There are ample reasons on morphological grounds for relating Kosteletzkya and Hibiscus sect. Pterocarpus, e.g., they share depressed, pentagonal, 5-crested capsules, the valves of which fall away completely from the floral axis at maturity, as well as similarities in calyx, involucre, and vegetative char- acters. Now, however, any proposal to unite the two taxa must take into account the difference in chromosome numbers. Acknowledgments Field collections were aided by funds from National Science Foundation Grant GB-8759 to David M. Bates. I also wish to thank Paul A. Fryxell for providing trans- portation in Mexico. LITERATURE CITED Bates, D. M., & O. J. BLANCHARD, JR. 1970. Chromosome numbers in the Malvales. II. New or otherwise noteworthy counts rele- vant to classification in the Malvaceae, tribe Malveae. Amer. J. Bot. 57: 927-934. HOCHREUTINER, B. P. G. 1900. Revision du genre Hibiscus, An- nuaire Conserv. Jard. Bot. Genéve 4: 23-191. SKOVSTED, A. 1935. Chromosome numbers in the Malvaceae. I. J. Genet. 31: 263-296. 1941. Chromosome numbers in the Malvaceae. II. C. R. Lab. Carlsb. Physiol. 23: 195-242. 66 Rhodora [Vol. 76 Table 1. Chromosome counts in Kosteletzkya Species n-number Locality and Collector K. adoensis 19 Cultivated, BH-69:555. Seed source — Angola: Huambo Distr., Instituto de Investigação Agronómica. de Angola K. buettneri 19 Cultivated, BH-72:100. Seed source — Zambia: “C Pro- vince," Robinson 6706 K. coulteri 19 Mexico: Sinaloa, Fryxell, Bates & Blanchard 1544 K. pentasperma 19 Mexico: Jalisco, Fryxell, Bates & Blanchard 1610 19 Mexico: Michoacan, Fryzell, Bates & Blanchard 1650 19 Mexico: Nayarit, Fryxell, Bates & Blanchard 1563 K. paniculata 19 Cultivated, BH-71:119. Seed source — Mexico: Jalisco, Fryxell, Bates & Blanchard 1590 K. virginica 19 USA: Florida, Collier Co., Blanchard & Blanchard 303 19 USA: New Jersey, Ocean Co., Blanchard & Blanchard 284 19 Cultivated, BH-69:430. Seed source — USA: Texas, Jeffer- son Co., Knight 88 L. H. BAILEY HORTORIUM CORNELL UNIVERSITY ITHACA, NEW YORK 14850 NAME CHANGES FOR THE SEED PLANTS IN THE BAHAMA FLORA WILLIAM T. GILLIS The author has been working in collaboration with Richard A. Howard of the Arnold Arboretum and George R. Proctor of the Institute of Jamaica towards a revision of Britton and Millspaugh’s “Bahama Flora," published first in 1920 and reprinted without changes in 1962. A recent paper by the three of us (Gillis, Howard, and Proctor, 1973) has reported additions to the flora of the Bahamas based on personal collections, new collections available to us, and reports from the literature. The present paper offers updated annotations on the correct scientific names to be applied to the species listed by Britton and Millspaugh whose work was completed under the provisions of the American Code of Botanical Nomenclature. We encourage others to use this list and its supporting bibliography, and to report additional changes or additions to me. We would particularly value contribu- tions which specialists in non-vascular plants could make to the listings in Britton and Millspaugh for mosses and liverworts, algae, fungi, slime molds, and lichens. Our efforts will consider only the flowering plants and pteri- dophytes. We therefore encourage workers in these other fields of botany to update knowledge of their disciplines in the Bahamas too. The value of lists of nomenclatural corrections has been evident to us in the work of Alain (1965) on the flora of Puerto Rico and the Virgin Islands, Additional nomen- clatural changes have been obtained from monographs or the recent floras of Jamaica (Adams, 1971) and Barbados (Gooding et al., 1965). In all of these, however, the rea- sons for the changes of names are not always explained, requiring the curious or careful worker to reinvestigate each and every problem. I believe the explanations which aecompany the changes reported here will be of value to 67 68 Rhodora [Vol. 76 other workers in subtropical New World floras and par- ticularly in the West Indies. Reasons given for changes are usually: use of an earlier epithet, avoidance of a tautonym, replacement of a later homonym, correct application of a name, etc. When name changes have been made by others than myself, I have cited the publication upon which this information was based. Conversely, when I have had reason not to accept a particular treatment, then reasons have been stated. The listing is by no means complete. Certain groups within the Boraginaceae, Rubiaceae, Loranthaceae, and Asclepiadaceae need intensive study and many more obser- vations in the field, especially in such genera as Helio- tropium, Borreria, Dendropemon, Phthirusa, and Cynan- chum. A new look must be given to the genus Agave for the West Indies, inasmuch as the most recent treatment is now 60 years old (Trelease, 1913). Other groups in- cluding the ferns are being studied by others at the present time and we await publication of the conclusions of these investigators. The Turks and Caicos Islands are politically a separate Crown Colony from the newly independent Commonwealth of the Bahama Islands. Britton and Millspaugh included all within the Bahama Flora and such is the geographical sense employed here in that it includes the Turks and Caicos Islands as well. For ease of reference this paper will follow the order of species presented in Britton and Millspaugh’s Bahama Flora, which will be Monocotyledons first, followed by the Dicotyledons, and then the Gymnosperms. Again for brevity, Britton and Millspaugh’s flora is referred to by the expression B&M in the text of this paper. The figure in the left margin refers to the page in B&M on which the taxon in question is discussed. The name following the arrow (—) is the name considered to be correct. These names are not necessarily nomenclatural or taxonomic equivalents, hence the reason for not employing an equals 1974] Bahama Flora — Gillis 69 sign (—). In some instances, B&M misidentified the plant or used a binomial incorrectly. Where monographs or significant studies support the use of names as they were employed in B&M, such references are cited under the generic name, e.g., Guaiacum (Porter, 1972). In a few instances where the name changes have involved closely allied genera, or where the existing keys in B&M are wholly inadequate, particularly for use in the field, I have provided new keys. They are designed to employ characters of the floral, fruiting, and vegetative portions of the plant so that they can be of optimum use in working with specimens, whether living or in the herbarium, and whether reproductive or vegetative. ACKNOWLEDGEMENTS I should like to acknowledge the help of my associates Dr. Howard and Mr. Proctor in drawing up this list, of Dr. Carroll E. Wood, Jr. and Mr. George N. Avery, who read the manuscript and made valuable suggestions, of my col- league Dr. Kenneth R. Robertson who gave advice, and of the many workers who have studied genera that have representatives in the Bahamas. Special appreciation is due to the librarians at the Arnold Arboretum and Gray Herbarium of Harvard University and for the use of the bibliographic files of the Generic Flora of the Southeastern United States project, made available by Dr. Wood. I should also like to acknowledge with thanks the continuing co- operation of the personnel of the Ministry of Agriculture and Fisheries, Commonwealth of the Bahama Islands. This study was made while the author was a Research Fellow at the Arnold Arboretum and I express my grati- tude for this opportunity. A generous gift from an anony- mous donor interested in the work on the Bahama flora. made this study possible. 70 PAGE IN B&M 4 Rhodora [Vol. 76 TYPHACEAE Typha angustifolia > Typha domingensis (Pers.) Kunth, These are two different valid species accord- ing to Hotchkiss and Dozier (1949) with only Typha domingensis occurring in the Bahamas. POTAMOGETONACEAE (ZANNICHELLIACEAE) Potamogeton heterophyllus > Potamogeton gramineus L. var. gramineus. On the basis of both herbarium specimens and abundant fresh material from a pond on San Salvador Island (Gillis 8864), I have con- sidered all the Potamogeton material on the Islands to be the same (see Ogden, 1948). The question of good characters, reliable under all conditions, for separating Ruppia maritima from R. cirrhosa, is still open. Various treatments have been consulted (Fernald and Wiegand, 1914; Hag- strom, 1916; McCann, 1945; Setchell, 1946; Reese, 1962a and b; Gamerro, 1968; and Schwanitz, 1967) for specimens collected in the Bahamas. For the moment, it appears that both species seem to be present. Therefore, R. cirrhosa is an addition to the flora, rather than merely a different name for the species in B&M. Collectors are urged to look for R. cirrhosa inasmuch as flowering and fruiting material are as yet unknown in the Bahamas. An adaptation of Gamerro's key follows: Peduncle of inflorescence more than 10 cm. (8-60 em.), coiled until anthesis; anther sacs reniform, 1.7 mm. long; fruits rounded at apex; leaves 1 mm. broad; leaf epidermal cells 16-194; 2n = 40 .... TEN Ruppia cirrhosa (Petagna) Grande. Peduncle of inflorescence less than 5 cm., not coiled in spiral until anthesis; anther sacs elliptic, 0.6-0.7 mm. long; fruits irregularly pointed; leaves 0.5 1974] 6 13 14 Bahama Flora — Gillis 71 mm. broad; epidermal cells of leaves 12.5-16.0,5 Oe ee et Mert a tad Ruppia maritima L. ZANNICHELLIACEAE (CYMODOCEACEAE) Cymodocea manatorum —> Syringodium filiforme Kütz. This change of name follows Dandy and Tandy (1939) and den Hartog (1970). Halodule wrightii Aschers. is indeed in the Bahamas, but so is H. beaudettei (den Hartog) den Hartog (1964). These two species can be separated as fol- lows: Leaf tips bicuspidate; leaf blades 1/3-4/5 mm. wide; leaves with linear teeth, concave on the inner side ... Halodule wrightii Aschers. Leaf tips tricuspidate; leaf blades 2/3-114 mm. wide; leaves with linear lateral teeth ... Halodule beaudettei (den Hartog) den Hartog. HYDROCHARITACEAE (ELODEACEAE) Halophila — consult Hartog (1959). ALISMATACEAE Echinodorus cordifolius — Echinodorus berteroi (Spreng.) Fass. (See Fassett, 1955). GRAMINEAE There is some question among agrostologists whether Schizachyrium should be segregated from Andropogon as it was in B&M. Chase (1951) joins the two genera. For the moment, however, I shall follow treatments by Gould in maintaining them as separate genera (see Gould, 1967, 1968, and 1969). Nazia aliena > Tragus berteronianus Schult. Tragus is conserved over Nazia. The species epithet changed from aliena to berteronianus because the type of Lappago alienus Spreng., the basionym, is referred to Pseudechinolaena. polystachya (H.B.K.) Stapf, and not to this Bahamian plant (Chase, 1951). 72 15 16 16 16 18 18 18 19 20 Rhodora [Vol. 76 Syntherisma filiformis > Digitaria panicea (Sw.) Urb. The species on which the epithet filiformis is based is a different species from the one of concern here. The earliest name for this species is Milium panicea Sw. which was transferred to Digitaria by Urban. The combination was made twice in the same year (1920), by Urban in February and by Fernald in June, The Urban combination obviously has priority. Digitaria is an earlier generic name than Syntherisma. Syntherisma digitata — Digitaria horizontalis Willd. Milium digitatum Sw. (1788) is the earliest basionym, but is is preoccupied in Digitaria by D. digitata Buse (1854). The next earliest name available is D. kori- zontalis Willd. (1809). Syntherisma sanguinalis > Digitaria sanguinalis (L.) Scop. Valota insularis > Trichachne insularis (L.) Nees. Valota is inadequately published (Chase, 1951). Paspalum poiretii R. et S. ^ Paspalum sagetii Chase. Paspalum caespitosum — Paspalum poiretii R. et S. The work of Chase (1929) sorts out the names of these Paspalum species by typifying them. It is thus apparent that, although the name P. poiretii is still used in the Bahama flora, it must be applied to a different species from the one to which it was as- signed by B&M. Paspalum portoricense ^ Paspalum molle Poir. in Lam. The change is required because of an earlier epithet (Chase, 1951). Paspalum simpsonii > Paspalum blodgettii Chapm. Paspalum blodgetti is an earlier name for P. simp- sonii Nash, and not a synonym of P. caespitosum Chase, 1951). Paspalum glabrum — Paspalum laxum Lam. The change is required because of an earlier epithet (Chase, 1951). Paspalum distichum L. is attributed by B&M to the 1974] 22 22 22 24 24 27 29 Bahama Flora — Gillis T8 Pugil. Pl. Jam., one of the dissertations of Linnaeus (1759). It actually appeared earlier in the same year in the Systema Naturae, ed 10, and should be so attributed. Panicum barbinode > Panicum purpurascens Raddi. The change is required because of an earlier epithet (Hitchcock, 1936). Panicum distantiflorum — Setaria distantiflorum (A. Rich. in Sagra) Pilger. Panicum chapmani — Setaria chapmanii (Vasey) Pilger. Rominger (1962) has shown that these two species properly belong in Setaria. Panicum dichotomiflorum — Panicum _ bartowense Scribn. et Merr. These names were treated as synony- mous by B&M. Hitchcock (1935) and Chase (1951) felt that they are separate species and that the Baha- mian plant is P. bartowense, Fernald (1934) treated the taxon as P. dichotomiflorum var. bartowense Schribn. et Merr.) Fern. Panicum coerulescens > Panicum coerulescens Hack. ex Hitehe. This change is simply a corrected spelling. Oplismenus hirtellus (L.) R. & S. 1817 — Oplismenus hirtellus (L.) Beauv., Ess. Agrost. 54, 168. 1812. An author change is needed for this earlier publication of the binomial. Chaetochloa geniculata — Setaria geniculata (Lam.) Beauv. Chaetochloa setosa — Setaria setosa (Sw.) Beauv. Chaetochloa macrosperma — Setaria macrosperma (Scribn. et Merr.) K. Schum. Setaria is the older and preferred name over Chaetochloa. Setaria Beauv. is conserved over Setaria Michx. Cenchropsis mysuroides — Cenchrus mysuroides H.B.K. Cenchropsis is not now considered distinct from Cenchrus (Chase, 1936 and Delisle, 1963). Cenchrus carolinianus ^ Cenchrus incertus M. A. Curtis. The use of Cenchrus carolinianus in B&M was a misapplication of the name. That name is 74 29 29 30 31 31 32 38 33 Rhodora [Vol. 76 now considered a synonym of C. longispinus, but is correctly used for a plant not found in the Bahamas. The plant in question is C. incertus which is wide- spread in the Bahamas (DeLisle, 1963). Cenchrus microcephalus > Cenchrus incertus M. A. Curtis. DeLisle (ibid.) felt that there was so little difference between the *endemic" race of Cenchrus with small fruits and the more widespread species that he treated them as conspecific. The small fruited form should be considered an insular form. Cenchrus viridis —^ Cenchrus brownii R. et S. This name change is required because of an earlier epithet (DeLisle, ibid.). Stenotaphrum — Consult Sauer, 1972. Aristida gyrans > Aristida vilfifolia Henr. After publication of B&M, Aristida vilfifolia was segregated as a distinct species from A. gyrans which was the name in B&M (Henrard, 1926-33; 1929-33). It is the segregate which indeed is the one in the Bahama flora. Aristida scabra > Aristida ternipes Cav. This change is required because of an earlier epithet. Sporobolus indicus — Sporobolus jacquemontii Kunth. Sporobolus berteroanus — Sporobolus indicus (L.) R. Br. In the Sporobolus indicus complex, there are two Caribbean species which are often confused: S. jac- quemontii, which has an obtuse upper glume and is a fully tropical species, and S. indicus, which has an acute upper glume and is a more subtropical species. According to the description in B&M, these taxa have been confused (Clayton, 1964 and personal communi- cation). Sporobolus atrovirens — Sporobolus bahamensis Hack. Sporobolus atrovirens is a different plant from the one in the Bahamas; it is a Mexican plant, is a perennial, and has smaller spikelets and glumes than S. baha- mensis. The latter is a Bahamian endemic and an annual. 1974] 33 34 36 39 39 41 Bahama Flora — Gillis 75 Sporobolus argutus > Sporobolus pyramidatus (Lam.) Hitehe. This name change is required because of an earlier basionym. The citation for Domingan drop- seed-grass should be Sporobolus domingensis (Trin.) Kunth, Rév. Gram. 1, Suppl.; xvii. 1830, a citation three years earlier than that cited in B&M. Capriola dactylon > Cynodon dactylon (L.) Pers. Cynodon is conserved over Capriola, Chloris polydactyla > Chloris dandyana Adams. Adams (1971) published a new name for this plant without giving the background reasons for the illegiti- macy of the original name. The background is as follows: There is a Jamaican plant with the legiti- mate name Andropogon barbatus L. Some time after describing this Jamaican grass, Linnaeus applied the same binomial to an East Indian grass. The epithet barbatus (referring to the East Indian species) was transferred to Chloris by Swartz, an act which pre- vents the transfer of the Jamaican species epithet to Chloris. In the meantime, Linnaeus proposed a super- fluous name for the first Andropogon barbatus, i.e., A. polydactylos. Swartz’s transfer of this super- fluous name to Chloris is illegitimate; hence Adams’s publication of a new name for the species in the West Indies. (I should like to acknowledge personal com- munication from C. D. Adams and J. E. Dandy for the historical background on this matter.) Phragmites phragmites ^5 Phragmites australis (Cav.) Trin. ex Steud. Clayton (1968) pointed out that the correct name for the pan-tropical reed is the com- bination used above. Diplachne fascicularis > Leptochloa fascicularis (Lam.) A. Gray. Current views of agrostologists suggest that Diplachne should be contained within Leptochloa. Eragrostis cilaris (L.) Link 1827 Eragrostis cili- aris (L.) R.Br. in Tuckey, 1818. Chase (1951) re- corded the earlier publication of this combination. 76 42 44 45 46 AT 49 Rhodora [Vol. 76 - Uniola virgata > Leptochloópsis virgata — (Poir.) Yates. Yates (1966) has subdivided Uniola into Uniola, Chasmanthium, and Leptochlodpsis on the basis of a number of anatomical, cytological, morpho- logical, and embryological characters. Because this species of spike-grass is considered close to Lepto- chloa, Yates erected Leptochlodpsis to contain it and another species from Ecuador. CYPERACEAE Cyperus paniculatus > Cyperus polydactylos Rottb. These names originate from the same date and are considered to be synonymous. Corcoran (1941) chose C. polydactylos to be the inclusive name. The Bahama populations might be designated as C. polydactylos var. texensis (Torr.) Fern. (McLaughlin, 1944). Cyperus cuspidatus ^ Cyperus aristatus Rottb. True C. cuspidatus does not occur in the Bahamas, but in Cuba. (See Horvat, 1941; McLaughlin, 1944). Cyperus pseudovegetus > Cyperus distinctus Steud. Cyperus pseudovegetus was a name misapplied to the plant of the Bahamas (O’Neill, 1939). Cyperus brunneus — Cyperus planifolius L. C. Rich. This name change is required by an earlier epithet (McGivney, 1938; McLaughlin, 1944). Cyperus ferax — Cyperus odoratus L. var. odoratus. O'Neill (1940) demonstrated that these two names apply to the same species, and that C. odoratus is the earlier name. He further thought that the typical variety was the one found in the Bahamas. According to the treatment by McLaughlin (1944), the populations of Cyperus filiformis Sw. in the Bahamas are of the typical variety (var. filiformis). Stenophyllus wilsonii > Bulbostylis floccosa (Griseb.) Clark in Urb. Stenophyllus is merged with Bulbo- stylis in the treatment by Kral (1971). Bulbostylis floccosa is the earliest name. It is not endemic to the 1974] 50 51 52 53 54 54 55 55 56 Bahama Flora — Gillis T4 Bahamas as stated in B&M, but also is found in Cuba and Hispaniola. Fimbristylis diphylla — Fimbristylis dichotoma (L.) Vahl. This change is required because of an earlier basionym. According to B&M, the record of Fimbristylis hirta in the Bahamas is very tenuous. Neither Kral nor I have seen any material from the Bahamas. If in- deed it does occur there, its more appropriate name should be F. squarrosa Vahl (see Kral, 1971). Abildgaardia monostachya > Abildgaardia ovata (Burm. f.) Kral. This change is required because of an earlier epithet (Kral, ibid.). There are cyperologists who wish to merge Dichro- mena with Rhynchospora. On the basis of the striking white bracts and insect pollination characteristic of Dichromena, I am, for the moment at least, retaining it as distinct. Rhynchospora cyperoides > R. cyperoides (Sw.) Mart. var. cyperoides. Rhynchospora tracyi —^ Rhynchospora cyperoides var. triceps (Vahl) Bock. This treatment follows that of Kükenthal (1949). Rhynchospora perplexa > Rhynchospora microcarpa Baldw. ex Gray. Gale (1944)maintained that true R. perplexa was not in the Bahamas and that all the material which Britton and Millspaugh called by this name was misdetermined. Rhynchospora bahamensis Britt. ~ Rhynchospora lin- deniana var. bahamensis (Britt.) Gale. Gale's treat- ment (ibid.) of the West Indian Rhynchospora species reduced Britton’s species to a variety of the more widespread R. lindeniana. Rhynchospora marisculus > Rhynchospora odorata C. Wright ex Griseb. Gale (ibid.), in examining the same material which Britton and Millspaugh did, determined that the use of R. marisculus in B&M was a misapplication of the name. 78 56 60 61 67 67 Rhodora [Vol. 76 Mariscus jamaicensis > Cladium jamaicense Crantz. Most treatments now segregate Cladium from Mariscus. Scleria. Consult Jackson, 1949. PALMAE (ARECACEAE) Thrinax parviflora > Thrinax floridana Sarg. Thri- nax parviflora is an endemic species of Jamaica. Furthurmore, the treatment of Thrinax by Read in Adams (1971) will be superseded now that additional type material has been examined. The correct name for the small thatch of South Florida and the northern West Indies with leaf segments broadest at the point of fusion, with scattered, fimbriate, centrally trans- lucent seales, and glabrous axes should be T. floridana (Read, personal communication). Thrinax parviflora > Thrinax floridana Sarg. Thri- (Jacq.) L. H. Bailey. True C. argentea may exist in the southern Bahamas, but this has yet to be demon- strated. It is now treated as being indigenous to Hispaniola. (See Moore, 1963). Paurotis wrightii > Acoelorrhaphe wrightii (Griseb. ex H. Wendl.) H. Wendl. ex Becc. The preferred name for the Paurotis palm, Spanish-top, or Ever- glades palm is now Acoelorrhaphe (Moore, 1963). Pseudophoenix vinifera > Pseudophoenix sargentii subsp. saonae (Cook) Read. True P. vinifera is in- digenous to Hispaniola and is not known from the Bahamas. (Read, 1968). COMMELINACEAE Commelina longicaulis ^ Commelina diffusa Burm. f. Commelina elegans —^ Commelina virginica L. Name changes are made here in accord with the treatment by Brashier (1969). The following key may be more easily used than that in B&M: Plant annual, trailing; spathes not united at the base, margins ciliate; internodes glabrous; sta- 1974] 68 69 71 78 78 79 83 83 84 84 84 Bahama Flora — Gillis 79 minodes Qoi m Commelina diffusa Burm. f. Plant perennial, tufted and erect; spathes united at the base, margins membranous; internodes puberu- lent; staminodes 3 ...... Commelina virginica L. Rhoeo discolor ^ Rhoeo spathacea (Sw.) Stearn. This name change follows Stearn (1957) who discovered that the earlier epithet of Swartz applies to this plant. AGAVACEAE (LILIACEAE) Cordyline guineensis 5 Sansevieria hyacinthoides (L.) Druce. This change follows Wijnands (1973). SMILACACEAE Smilax. Consult Coker, 1944. AMARYLLIDACEAE Atamosco rosea — Zephranthes rosea Lindl. Atamosco cardinalis > Zephyranthes cardinalis C. Wright. Zephyranthes is conserved over Atamosco. Hymenocallis caymanensis ^ Hymenocallis latifolia (Mill.) Roemer. This change follows the treatment by Sealy (1954). ORCHIDACEAE Carteria corallicola — Basiphyllaea corallicola (Small) Ames. Carteria is a later homonym for a genus of algae. Basiphyllaea was erected as a genus to correct this situation. Vanilla eggersii — Vanilla dilloniana Correll. Vanilla articulata — Vanilla barbellata Reichb. f. This treatment of Vanilla in the Bahamas follows Cor- rell (1950). Pelexia adnata — Spiranthes adnata (Sw.) Benth. Although Pelexia is a conserved name, it is also a generic synonym of Spiranthes (Garay and Sweet, 1972). Pelexia setacea > Eltroplectris calcarata (Sw.) Garay 80 85 85 86 87 87 89 89 90 91 Rhodora [Vol. 76 et Sweet. Luer (1972) changed Pelexia setacea to Centrogenium setaceum but Garay and Sweet (1972) maintained that this combination is based on an ille- gitimate name and therefore should be Eltroplectris calcarata (Sw.) Garay et Sweet. Ibidium tortile > Spiranthes tortilis (Sw.) L. C. Rich. Jbidiwm is considered to be a generic synonym of Spiranthes. (Garay & Sweet, 1972). Ibidium lucayanum — Spiranthes polyantha Reichb. f. These names are considered synonymous by Luer (1972). Stenorrhynchus lanceolatus — Spiranthes lanceolata (Aubl) Léon. Stenorrhynchus is now treated as a generic synonym of Spiranthes. (See Dunsterville and Garay, 1965). Ponthieva brittonae > Ponthieva racemosa var. brit- tonae (Ames) Luer. Two varieties of Ponthieva racemosa occur in the Bahamas. One is the typical variety (var. racemosa) as treated by Luer (1972) and reported by him as new to the flora. Var. brit- tonae is the reduction made by Luer of the name in B&M. Physurus querceticola —^ Erythodes querceticola (Lindl.) Ames. Physurus is considered to be a gene- ric synonym of Erythodes (Garay and Sweet, 1972). Polystachya minuta — Polystachya flavescens (Lindl.) J. J. Smith. Although the epithet minuta (from Epidendrum minutum Aubl.) is the oldest name available, it is preoccupied in Polystachya, and hence, P. flavescens must be used. Spathiger | rigidus > Epidendrum rigidum Jacq. Spathiger is considered a generic synonym of Epi- dendrum. Auliza nocturna > Epidendrum nocturnum Jacq. Auliza is considered a generic synonymn of Epiden- drum (Dunsterville and Garay, 1965; Luer, 1972). There is much discussion among orchid taxonomists whether Encyclia should be segregated from Epiden- 1974] 91 92 93 93 95 95 103 104 Bahama Flora — Gillis 81 drum or not. For the moment, I accept the treat- ment of Dressler (1961; 1966) and Beckner (1970) in maintaining Encyclia at the generic level as in B&M. Encyclia rufa > Encyclia bahamensis (Griseb.) Britt. et Millsp. Specimens labeled E. rufa from the Ba- hamas are now all best considered E. bahamensis. Encyclia rufa is, as B&M suspected, still not known from the archipelago. Encyclia diurna — Encyclia hodgeana (A. D. Hawkes) Beckner. This name change is required because of more complete understanding of the taxa involved. The species previously called E. diurna or E. gracilis is, in fact, a hybrid between E. hodgeana and E. tampensis. (See Beckner, 1970). Epicladium boothianum — Encyclia boothiana var. erythronioides (Small) Luer. Epicladiwm is a generic synonym of Encyclia. Anacheilium cochleatum — Encyclia cochleata var. triandra (Ames) Dressler. Anacheiliwm is a generic synonym of Encyclia. Laeliopsis domingensis ^ Broughtonia lindenii (Lindl.) Dressler. This change is made in accord with the treatment by Dressler (1961). Limodorum simpsoni — Calopogon tuberosus (L.) Britten, Stearns, and Poggenberg. This treatment follows Luer, 1972. PICRODENDRACEAE Picrodendron macrocarpum — Picrodendron baccatum (L.) Krug et Urb. in Engl. I presently have Picro- dendron under study. It appears that this genus should be placed within the Euphorbiaceae. There is likely only one species, which would be the typical one, P. baccatum. ULMACEAE Trema lamarckiana — Trema lamarckianum (R. et S.) Blume. The gender of Trema should be neuter. 82 Rhodora [Vol. 76 MORACEAE 105 Ficus jacquinifolia > Ficus perforata L. Fieus brevifolia — Ficus citrifolia Mill. Recent treat- ments of Ficus (DeWolf, 1969 and Condit, 1969) have equated F. perforata with F. jacquinifolia A. Rich. in Sagra. Moreover, they have also equated F. brevi- folia and F. citrifolia. B&M’s F. brevifolia Nutt. (lectotype at BM) must become F. citrifolia Mill., the earlier name (lectotype at BM, chosen by Dandy). I have further studied F. perforata which DeWolf and Condit have given as the correct name for F. jac- quinifolia (which itself should be written jacquinii- folia). The Linnaean name must be typified by an illustration in Plumier associated with the disserta- tion on the plants of Surinam (1775). Although there is a specimen of F. perforata at LINN (Savage Cata- log No. 1240: 9), the only reference in the Disserta- tion (and the later Amoenitates) in this context is that to Plumier. The specimen at LINN appears to be of the tree of Central America and northern South America, F. pertusa L. f. I should like to acknowledge the consultation and discussion with Dr. Gordon De- Wolf which proved very valuable in application of names in Ficus. A key to the Bahama material fol- lows: 1. Syconia sessile .......... Ficus aurea Nutt. 1. Sycania stalked 2. Syconia 7-10 mm. in diameter; petioles less than 1 cm.; blades obovate; lateral veins flush with lower surface; leaf base cuneate Lee eee eee eee ee eee Ficus perforata L. 2. Syconia 8-18 mm. in diameter; petioles longer than 1 cm.; blades ovate; lateral veins + prominent, somewhat elevated on lower surface; leaf blade base rounded ......... Lee eee ee eee eee eee Ficus citrifolia Mill. 1974] 108 113 114 114 115 115 117 TIT DT 121 121 121 Bahama Flora — Gillis 83 LORANTHACEAE Generic study of Dendropemon, Phthirusa, and Phora- dendron is needed before good species concepts and correct nomenclature emerge. ARISTOLOCHIACEAE Aristolochia passifloraefolia — Aristolochia passiflori- folia A. Rich. in Sagra. With the exception of the orthograhic change (above) made in accord with rules adopted at the Seattle Congress, the names for Aristolochia species in the Bahamas remain as they were in B&M (Pfeifer, 1966; 1970). POLYGONACEAE Persicaria hydropiperoides > Polygonum hydropipe- roides Michx. Persicaria punctata — Polygonum punctatum Ell. Persicaria portoricensis Polygonum densiflorum Meissn. Persicaria is treated by Graham and Wood (1955) as a section of Polygonum. Fagopyrum fagopyrum —> Fagopyrum esculentum Moench. This change is necessary in order to avoid a tautonym. Coccolobis laurifolia — Coccoloba diversifolia Jacq. Coccolobis bahamensis — Coccoloba tenuifolia L. Coccolobis diversifolia —^ Coccoloba swartzii Meisn. Coccoloba is conserved over Coccolobis. All Coccoloba species therefore should change spelling to conform. The 3 species mentioned above are changed further in accord with Howard (1957), From Howard's treatment also comes the change of citation for Coccoloba uvifera (L.) L., Syst. Nat., ed. 10, p. 1007. 1759. CHENOPODIACEAE Dondia linearis > Suaeda linearis (Ell.) Moq. Dondia fruticosa —> Suaeda fruticosa (L.) Forsk. Dondia insularis ^ Suaeda insularis (Britt.) Urb. et Ekm. Dondia is considered to be a taxonomic synonym 84 124 126 126 126 127 128 128 130 131 131 132 132 Rhodora [Vol. 76 of Suaeda, Suaeda Forsk. ex Scopoli remains in the list of conserved names even though the reason for its inclusion is no longer valid. AMARANTHACEAE Amaranthus gracilis > Amaranthus viridis L. Reed (1968) and Merrill (1936) considered these species to be taxonomic synonyms. Amaranthus viridis L. is the earlier name. Centrostachys indica — Achyranthes aspera var. in- dica L. Centrostachys seems better treated as a generic synonym Achyranthes. Achyranthes maritima — Alternanthera maritima St. Hil. Achyranthes polygonoides — Alternanthera polygo- noides (L.) R. Br. Achyranthes repens —> Alternanthera pungens H.B.K. This treatment follows Reed, 1968. The epithet pungens is used for the last species rather than the earlier epithet repens because Achyranthes repens is a nomen confusum according to Reed. Philoxerus remains the same except for citing an earlier date of publication: Philoxerus vermicularis (L.) R. Br. Prodr. Fl. Nov. Holl. 416. 1810. Iresine celosia — Iresine diffusa Humb. et Bonpl. ex Willd. (Sp. Pl., ed. 4, 4: 765. 1806). Iresine celosia is an illegitimate name. NYCTAGINACEAE Boerhaavea coccinea Mill. > Boerhavia coccinea Mill. Boerhaavea erecta L.— Boerhavia erecta L. Boer- havia is the original spelling. Torrubia obtusata > Guapira obtusata (Jacq.) Little. Torrubia cokeri > Guapira obtusata (Jacq.) Little. Torrubia longifolia > Guapira longifolia (Heimerl) Little. Torrubia bracei > Guapira bracei (Britt.) Little. If one considers that blollies are similar to Pisonia 1974] 138 139 143 144 144 146 147 Bahama Flora — Gillis 85 species, then Torrubia merges with Pisonia. If one wishes to recognize both as distinct genera, then one must take up Guapira, an earlier name than Torrubia. The Committee for Spermatophytes rejected a pro- posal to conserve Torrubia, hence a paper by Little (1968) which made the appropriate transfers to Guapira, It is currently felt that Torrubia cokeri is synonymous with Guapira obtusata; it has been so treated here. This is an exceedingly variable species with different forms having been named in the past. PORTULACACEAE Portulaca phaeosperma —> Portulaca rubricaulis H.B.K. Legrand (1952) chose the earlier name for this species. NYMPHAEACEAE Castalia pulchella > Nymphaea pulchella DC. Wood (1959) has treated Castalia as a subgenus of Nym- phaea. LAURACEAE Ocotea coriacea > Nectandra coriacea (Sw.) Griseb. Recent treatments have merged Ocotea and Nectandra (Wood, 1958; Allen, 1966). Persea pubescens Persea palustris (Raf.) Sarg. Kopp (1966) pointed out that palustris is an earlier epithet than pubescens for this species. CASSYTHACEAE Cassytha americana —> Cassytha filiformis L. This change is necessitated because filiformis is the earlier epithet. CRUCIFERAE (BRASSICACEAE) Hitchcock (1945) recognized several varieties of Lepidium virginicum. If this treatment is followed, the Bahama one is var. virginicum. Sinapis arvensis Brassica kaber (DC.) L. C. Wheeler. Wheeler (1938) thought that Sinapis should 86 148 148 151 152 153 154 Rhodora [Vol. 76 properly be considered a part of Brassica. Sinapis arvensis of B&M cannot be transferred to Brassica because there is already a B. arvensis. Brassica kaber appears to be an older combination than B. will- denovii Boiss., as attributed to the Bahamas in Adams, 1972. Radicula brevipes —> Rorippa portoricensis var. pu- mila (O. E. Schulz) Stuckey. The name used in B&M and its synonyms are misapplied to the Bahamian taxon, according to Stuckey (1972). Stuckey inad- vertently maintained a neuter ending for the varietal epithet in his treatment; this is corrected above. Canara didyma — Coronopus didymus J. E. Smith. Coronopus (1757) is an older name than Canara (1792). Coronopus Zinn. (1757) is conserved over Coronopus Miller (1754), a genus of Plantaginaceae. MORINGACEAE Moringa moringa — Moringa oleifera Lam. In avoid- ing the tautonym, Moringa moringa, Ernst (1963) has picked up Moringa oleifera for the horseradish- tree. CRASSULACEAE Bryophyllum pinnatum — Kalanchoë pinnata (Lam.) Pers. Treatments of Kalanchoë and Bryophyllum (Baldwin, 1938; Hamet, 1907) have recognized Kalanchoë as the inclusive genus. ROSACEAE (AMYGDALACEAE) Laurocerasus myrtifolia > Prunus myrtifolia (L.) Urb. The cherry-laurel is best considered in the larger genus Prunus especially in view of other recent floras which have so placed it. CHRYSOBALANACEAE (AMYGDALACEAE) Chrysobalanus pellocarpus ^ Chrysobalanus icaco L. Prance (1972) thought that there were insufficient means to separate C. pellocarpus from C. icaco and has treated them as taxonomic synonyms. 1974] 155 158 159 159 Bahama Flora — Gillis 87 LEGUMINOSAE (MIMOSACEAE) Pithecellobium is conserved over Pithecolobium, and hence all species change to this orthography. The ram’s horn has been a source of confusion for a number of years. Pithecellobium keyense Britt. ex Coker in Shattuck was proposed to replace P. guada- lupense because the latter name was not applicable. Britton did not state why he considered the name inapplicable. In creating the new name, Britton (actually Coker) had no description and hence, created a nomen nudum. This situation was rectified in 1928 by Britton and Rose. Although the type of Mimosa guadalupensis Pers., the basionym, cannot be found at present in the Jussieu Herbarium at P, there appears to be no valid reason why this name cannot be taken up, pending proof that this plant does not represent the concept of the species known as “ram’s horn" in the West Indies and South Florida. Isely (1972) referred to specimens at the Delessert Herbarium and at Leiden which should be “critical,” but inasmuch as the basionym is linked by Persoon with “Hab ad Guadalupam (Herb. Juss.)," the lecto- type material should be sought in the Jussieu Her- barium. Why Britton and Rose indicated that this species was “not Inga guadalupensis Desv." (which is based on the same basionym) is unknown. The only Pithecellobium which occurs on Guadaloupe is this one. I am therefore retaining the name in B&M, but adding the parenthetical reference to the author of the basionym which was omitted in B&M: Pithe- cellobium guadalupense (Pers.) Chapm. The Lysiloma species remain as in B&M (Gillis and Stearn, in press), except for correcting gender to neuter. Anneslia haematostoma — Calliandra haematomma (Bert.) Benth. Anneslia formosa > Calliandra formosa (Kunth) Benth. Calliandra is conserved over Anneslia. The 88 160 161 162 163 163 164 164 166 Rhodora [Vol. 76 specific epithet haematostoma in B&M appears to be a misprint, judging from the basionym they cite. Despite efforts of Britton and Rose (1928) to frac- tionate Acacia into Bahamia and Lucaya, it seems best to maintain the three species of Acacia in B&M under the same names as in the flora: Acacia acuifera, A. choriophylla, and A. macracantha. Vachellia farnesiana — Acacia farnesiana (L.) Willd. Vachellia is best considered a generic synonym of Acacia. Leucaena glauca — Leucaena latisiliqua (L.) Gillis et Stearn. Based on typification of an earlier name than either the epithet glauca or leucocephala (deWit, 1961). Gillis and Stearn (in press) have shown that latisiliqua should be taken up. Acuan virgatum — Desmanthus virgatus (L.) Willd. var. virgatus. Acuan depressum —> Desmanthus virgatus var. de- pressus (Willd.) Turner. Desmanthus is conserved over Acuan. The two Bahama plants probably are best considered distinct only at the infraspecific level; therefore the treatment of Isely (1970) is fol- lowed, Neptunia — consult Windler (1966). Prosopis — consult Johnston (1962). LEGUMINOSAE (CAESALPINIACEAE) Cassia. tora, —^ Cassia obtusifolia L. B&M suggest that Cassia tora and C. obtusifolia are synonyms and chose the former as the name to be used in the Bahama flora. Typification shows that such is not the case. DeWit (1955) selected a specimen in the Linnaean Herbarium as the type of C. tora L., but Brenan (1958) objected to this choice and selected a speci- men in the Hermann Herbarium (at BM) as the type. Based on the position of the petiolar gland, length of flowering and fruiting pedicels, anther shape, and presence of a 1.2-2 mm. areole on both 1974] 168 169 169 169 170 170 172 172 172 Bahama Flora — Gillis 89 sides of the seed, Brenan distinguished these two as distinct species. There is probably no Cassia tora in the New World according to Brenan. Chamaecrista chamaecrista — Cassia nictitans L. De- spite the efforts of Britton and Rose to fractionate Cassia into segregate genera Chamaecrista, Ditre- mexa, Peiranisia, ete., I maintain Cassia as an in- clusive genus. This necessitates different names for the six species placed in Chamaecrista in B&M. Cassia chamaecrista L. is based on a composite collection according to Pennell (1917). Therefore the correct name for the annual plant that is distributed through- out Eastern United States is generally accepted as C. fasciculata Michx. “Chamaecrista chamaecrista" of B&M should be rather the perennial species, Cassia nictitans L. I believe it is necessary to retypify all the names used in this complex in order to apply these names properly. Chamaecrista riparia > Cassia caymanensis C. D. Adams. Adams (1970) has shown that Cassia riparia is an illegitimate name, and therefore this plant must have a new epithet in Cassia. Chamaecrista lucayana — Cassia lucayana Britt. Chamaecrista caribaea — Cassia caribaea Northrop. Chamaecrista inaguensis > Cassia inaguensis Britt. Chamaecrista lineata > Cassia lineata Sw. These species revert to their original names in Cassia. Guilandina crista > Caesalpinia bonduc (L.) R. Br. Gray nickerbean has been shown by Dandy and Exell (1938) to be Caesalpinia (or Guilandina) bonduc. The epithet crista as used by Linnaeus refers to an Old World species. Guilandina bonduc — Caesalpinia divergens Urb. Guilandina ovalifolia — Caesalpinia ovalifolia Urb. Current thought among those working with this group of scrambling shrubs of the coastal regions of the West Indies is to treat them as a section of Caesalpinia, rather than a separate genus Guilandina. 90 173 177 178 181 181 Rhodora [Vol. 76 For some long period of time, G. bonduc was used to refer to the yellow-seeded nickerbean. As we have seen, proper typification by Dandy and Exell has shown that the name bonduc is misapplied when used thus. Among the synonyms given by B&M for this species, most apply to still other species. Caesal- pinia divergens seems to be the earliest name available for this species. The yellow-seeded nickerbeans in the Bahamas (with the exception of a new species to be described by Gillis and Proctor from Inagua) can be separated as follows: 1. Seeds spherical, yellow; leaflets elliptic to sub- orbicular, obtuse or emarginate ............. Lee ee eee Caesalpinia ovalifolia (Urb.) Britt. 1. Seeds oblong, bronze-brown; leaflets ovate, acute or acuminate ............. nee I accept B&M’s treatment of 1920 instead of the later treatment of Britton and Rose (1928) which recog- nized the segregate genus Vicarago. I see no reason to separate this taxon from Caesalpinia. LEGUMINOSAE (FABACEAE) Ateleia cubensis > Ateleia gummifera (Bert. ex DC.) Dietr. The epithet gummifera is an earlier epithet for this species (Rudd, 1968). Yakovlev (1967 a and b) has recognized a number of subspecific units in Sophora tomentosa L. For the moment, these seem difficult to accept as other than insular forms, and therefore I consider the taxon to be the single, variable S. tomentosa throughout the archipelago as in B&M. Cracca cinerea > Tephrosia cinerea (L.) Pers. Cracca cathartica — Tephrosia senna H.B.K. Although Cracca Benth. in Benth. et Oersted is conserved over Cracca L., Tephrosia Pers. is conserved over Cracca L., Needhamia Scopoli, and Reineria Moench. The 1974] 182 182 182 183 184 184 184 185 Bahama Flora — Gillis 91 correct generic name for the plants in the Bahamas is Tephrosia. Because the key to species in B&M is somewhat inadequate, a revised one follows: 1. Leaflets 9-15; racemes opposite the leaves; pod 2.5-4 mm. broad .. Tephrosia cinerea (L.) Pers. 1. Leaflets 5-9; racemes terminal and opposite the leaves; pod 4-5 mm. broad ................. Tephrosia senna H.B.K. e «.4* 9» 9 6,8 he 6 6 8 à 5 € 8 9.0, 9 Sesban sericea — Sesbania sericea (Willd.) DC. Sesban occidentalis —^ Sesbania emerus (Aubl.) Urb. Agati grandiflora — Sesbania grandiflora (L.) Pers. Sesbania is conserved over Sesban and Agati. The second of the species mentioned above was known only from fruit at the time of the writing of B&M. From collections made recently from the same popu- lation, I have determined it to be S. emerus. Stylosanthes — consult Mohlenbrock (1957). Meibomia supina —^ Desmodium canum (Gmel.) Schinz et Thell. Meibomia tortuosa —^ Desmodium tortuosum ( Mill.) DC. Meibomia mollis ^À Desmodium glabrum (Mill.) DC. Desmodium is conserved over Meibomia. Supina cannot be used as an epithet for the first species above because it is based on a name which is a later homonym (Hedysarum supinum Sw. non H. supinum Chaix ex Villars). Incanum has often been taken up as a specific epithet for this plant also (based on Hedysarum incanum Sw.), but the basionym is a later homonym for H. incanum Thunb. Desmodium canum is based on the earliest available legitimate epithet. Desmodium glabrum is based on an earlier basionym than Meibomia mollis of B&M. Alysicarpus nummularifolius > Alysicarpus vaginalis (L.) DC. Alysicarpus vaginalis has been used for the false moneywort rather than a combination based on Hedysarum nummularifolium L. Schindler (1926) 92 186 186 187 188 188 191 192 Rhodora [Vol. 76 interprets the Linnaean material as being Indigofera echinata, in part, and Alysicarpus monilifer, in part, hence the choice of Hedysarum vaginale L. as the basionym rather than the earlier one, Ecastophyllum ecastophyllum > Dalbergia ecastophyl- lum (L.) Taub. Dalbergia’s having been conserved over Ecastophyllum eliminates the problem of the tautonym in B&M. Ichthyomethia | piscipula — Piscidia piscipula (L.) Sarg. Piscidia has been conserved over I[chthyome- thia. (See Rudd, 1969). Abrus abrus — Abrus precatorius L. The later epithet of Linnaeus is used to avoid the tautonym. Bradburya floridana — Centrosema floridanum (Britt.) Lakela. Bradburya virginiana — Centrosema virginianum (L.) Benth. Bradburia has been conserved for a genus of Compositae over Bradburya of the Leguminosae. See also Lakela (1963). Canavali lineata — Canavalia rosea (Sw.) DC. Canavali bahamensis > Canavalia nitida (Cav.) Piper. Canavalia is conserved over Canavali. Sauer (1964) revised species of Canavalia, but apparently over- looked Johnston’s note (1949b) on publication of names referring to C. maritima. The basionym of this oft-used binomial has been treated as Dolichos maritima Aubl., but Thouars, in making the combina- tion Canavalia maritima, did not in fact cite the Aublet name; hence it is not a new combination, but a new name dating from 1813. Aublet’s name cannot then be transferred to Canavalia because it is pre- émpted by the Thouars combination, albeit for the same species. Alas, another name proposed in the in- terval is the oldest which must be used, Canavalia rosea (Sw.) DC. Species 2 of B&M (Canavali gladi- ata) is probably not in the Bahamas. All specimens bearing this determination by either Britton or Mills- 1974] 192 193 193 193 194 195 195 197 197 Bahama Flora — Gillis 93 paugh have been annotated by Sauer as C. maritima (=C. rosea). Cajan cajan —> Cajanus cajan (L.) Millsp. Cajanus is conserved over Cajan. The tautonym of B&M can thus be avoided. Dolicholus reticulatus ^ Rhynchosia reticulata (Sw.) DC. Dolicholus swartzii > Rhynchosia swartzii (Vail) Urb. Dolicholus minima > Rhynchosia minima (L.) DC. Rhynchosia is conserved over Dolicholus. Dr. John Grear indicates (personal communication) that Rhynchosia caribaea based on Glycine caribaea Jacq. is not in the flora, thus agreeing with B&M. The latter is ironically not native to the New World, but is restricted to South Africa. Phaseolus lathyroides ^ Macroptilium lathyroides (L.) Urb. On the basis of having five calyx teeth rather than four, Macroptilium is segregated from the large genus Phaseolus. Dolichos lablab > Lablab purpureus L. Many workers (including Adams, 1972) use the generic segregate Lablab in preference to Dolichos. If one follows this trend, then the change above is indicated. Otherwise, Dolichos lablab may continue to be used. Dolichos insularis ^ Oxyrhynchus volubilis Brande- gee. This change follows studies by Rudd (1967). Vigna repens > Vigna luteola (Jacq.) Benth. in Mart. Although the epithet repens is the oldest name, it is invalidated in Vigna by V. repens Baker, published in 1876 (Merrill, 1910). OXALIDACEAE Inoxalis intermedia > Oxalis intermedia A. Rich. Xanthoxalis corniculata > Oxalis corniculata L. Mod- ern treatment of Oxalidaceae would merge /noxalis and Xanthoxalis into Oxalis (Eiten, 1963). 94 200 200 200 201 201 202 205 Rhodora [Vol. 76 LINACEAE Cathartolinum curtissii ^À Linum medium var. texa- num (Planch.) Fern. Cathartolinum corallicola —^ Linum bahamense var. corallicola (Small) Rogers. Cathartolinum bahamense > Linum bahamense North- rop var. bahamense. Cathartolinum bracei —^ Linum bahamense var. bracei (Small) Rogers. Cathartolinum lignosum — Linum bahamense North- rop. var. bahamense. Rogers (1963; 1968) has restored Linum as the correct name for the yellow-flowered species in the West Indies. I have followed his 1963 treatment chiefly. ZYGOPHYLLACEAE Guaiacum — consult Porter (1972). MALPIGHIACEAE Byrsonima cuneata — Byrsonima lucida (Mill.) DC. It is generally agreed that the plum-berry or locust- berry should be Byrsonima lucida, based on Malpighia lucida, an earlier epithet than cuneata. But upon whose M. lucida? Most authors attribute the name to Swartz, but his M. lucida is a later homonym for M. lucida Mill. Both names can be typified by material at BM. DeCandolle (1824) specifically indicated the Swartz material as basionym, but it is illegitimate as a later homonym. Kunth (1923) indicated that he thought that M. lucida should be a Byrsonima but did not make the transfer in accord with the Code. Rather than presume to make a new combination at this date myself, I interpret this situation in the light of his- tory. Swartz undoubtedly knew of Miller’s name and probably even examined herbarium material from the Chelsea Physic Garden upon which the lectotype of M. lucida Mill. (chosen by Gillis) has been selected. That he (Swartz) did not specifically indicate that 1974] 211 219 220 Bahama Flora — Gillis 95 he was taking up Miller's name is not important, it seems to me. There is ample precedent with Jacquin and Linnaeus, for instance, in assuming that the later author knew of the earlier work but did not, under existing convention, always cite the author of the earlier binomial, or place of publication. With this assumption, I shall follow Robertson (1971), in at- tributing the combination to DeCandolle with a lecto- type of Miller's name at BM. SIMAROUBACEAE Cronquist's treatment of Alvaradoa (1944) would make the Bahamian populations Alvaradoa amorpho- ides subp. psilophylla (Urb.) Cronq. BURSERACEAE Elaphrium inaguense > Bursera inaguensis Britt. Elaphrium simaruba — Bursera simaruba (L.) Sarg. Bursera is conserved over Elaphrium. POLYGALACEAE Badiera oblongata > Polygala oblongata — (Britt.) Blake. It seems desirable not to segregate the woody species of Polygala into the genus Badiera. EUPHORBIACEAE Cicca disticha — Phyllanthus acidus (L.) Skeels. This name change follows Webster (1956-58). Margaritaria bahamensis > Margaritaria tetracocca (Baill) Webster. Webster (1956-58: J. Arnold Arbor. 38: 66) found that an earlier epithet applied to this species. Xylophylla epiphyllanthus —^ Phyllanthus epiphyllan- thus L. This treatment follows Webster (1967). Phyllanthus pruinosus > Phyllanthus caroliniensis subsp. saxicola (Small) Webster. Phyllanthus pentaphyllus > Phyllanthus pentaphyllus C. Wright subsp. pentaphyllus. This treatment fol- lows Webster (1970). 96 223 225 225 225 228 229 229 230 232 234 235 Rhodora [Vol. 76 Croton flocculosus Geisl. > Croton flavens var. bal- samiferus (Jacq.) Muell.-Arg. It appears to me that Croton flocculosus is best treated as a variety of the widespread Croton flavens. It matches well the lecto- type of Croton balsamiferus Jacq. (lectotype from Martinique at BM), the basionym. Curcas curcas > Jatropha curcas L. Placement of the physie-nut in Jatropha seems to be the best place- ment taxonomically, and also avoids a tautonym. Adenoropiwm gossypifolium — Jatropha gossypifolia L. Adenoropium is best treated as a subdivision of Jatropha. Argythamnia — consult Ingram (1967). The place of publication of Acalypha alopecuroidea Jacq. should be Collect. 3: 196. 1790. Acalypha ostryaefolia > Acalypha ostryifolia Ridd. Pera bumeliaefolia > Pera bumeliifolia Griseb. These orthographic changes are required by the Seattle Code. Manihot manihot > Manihot esculenta Crantz. This name is the earliest available to replace the tautonym used in B&M. Gymnanthes lucida — Ateramnus lucidus (Sw.) Rothm. Rothmaler (1944) and Dandy (1967) have resur- rected Ateramnus P. Browne to replace the later Gymnanthes Sw. Although I am of the opinion that Gymnanthes should be conserved over Ateramnus, conservation was already rejected once by the Special Committee for Pteridophyta and Phanerogamae (Taxon 3: 241. 1954). 'There appears to be no choice but to adopt the new combination in Ateramnus. Adenorima gymnonota > Euphorbia gymnonota Urb. Adenorima is treated as a section of Huphorbia by Webster (1967) and that treatment is followed here. Arthrothammus | cassythoides > Euphorbia cassytho- ides Boiss. Arthrothamnus differs too little from other forms of Euphorbia to be segregated from it. 1974] 235 236 238 238 238 238 243 244 247 248 248 248 249 Bahama Flora — Gillis 97 Tithymalus | trichotomus > Euphorbia trichotoma H.B.K. Tithymalus is treated by Webster (1967) as a section of Euphorbia. Aklema petiolaris > Euphorbia petiolaris Sims. Aklema differs too little from other forms of Eu- phorbia to be segregated from it. Chamaesyce wilsonit > Chamaesyce lecheoides var. wilsonii (Millsp.) Burch. Chamaesyce | lecheoides > Chamaesyce — lecheoides Millsp. var. lecheoides. Chamaesyce insulae-salis ^ Chamaesyce centunculo- ides (HBK.) Millsp. Chamaesyce exumensis —^ Chamaesyce lecheoides var. exumensis (Millsp.) Burch. These name changes fol- low the treatment by Burch (1966). BUXACEAE Tricera bahamensis ^ Buxus bahamensis Baker in Hook. Tricera is best treated by the earlier name Buxus (Howard, 1962). ANACARDIACEAE Poison-ivy in the Bahamas is the typical subspecies: Toxicodendron radicans (L.) Kuntze subsp. radicans (see Gillis, 1971). CELASTRACEAE Rhacoma coriacea, ^ Crossopetalum coriaceum North- rop. Rhacoma — crossopetalum — Crossopetalum rhacoma (Sw.) Hitchc. Rhacoma aquifolia + Crossopetalum aquifolium (Griseb.) Hitchc. Rhacoma ilicifolia > Crossopetalum ilicifolium (Poir.) Kuntze. Crossopetalum was revived by Brizicky (1964) as the earliest name for this genus. Although several recent rorkers have accepted Cas- sine as a widespread genus that would include Elaeodendron (Adams, 1972 and Ding Hou, 1963), 98 252 253 253 257 257 258 258 263 264 266 Rhodora [Vol. 76 I have chosen to follow Robson (1965) to recognize the segregate genus Elaeodendron for the Bahamas, thus leaving Cassine sensu stricto for a small African genus. SAPINDACEAE Thyana discolor > Thouinia discolor Griseb. Thou- inia Poit. is an earlier name than Thyana Hamilt. and is conserved over Thouinia Thunb. ex L.f. Allophylus cominia — Allophylus cobbe (L.) Raeusch. Leenhouts (1967) makes a strong case for recognizing only one worldwide species in this genus. He has found no morphological gaps which can be used to distinguish separate populations in any geographical sense. Having chosen to recognize only one species, Leenhouts chose A. cobbe as the all-encompassing name from two Linnaean names of the same date. Melicocca bijuga — Melicoccus bijugatus (L.) Jacq. The correct spelling of this name was worked out by Brizicky (1963). RHAMNACEAE Sarcomphalus taylori — Ziziphus taylori (Britt.) M. C. Johnston. Johnston’s treatment (1964) merges Sarcomphalus with Ziziphus. Colubrina reclinata > Colubrina elliptica (Sw.) Briz. et Stern. Colubrina cubensis > Colubrina cubensis var. flori- dana M. C. Johnston. Colubrina colubrina > Colubrina arborescens (Mill.) Sarg. These changes follow the monographic treat- ment by Johnston (1971). TILIACEAE Triumfetta — consult Ko Ko Lay, 1950. MALVACEAE Phymosia — consult Fryxell (1971). Gayoides crispum — Herissantia crispa (L.) Brizicky. 1974] 267 268 268 268 271 271 273 274 Bahama Flora — Gillis 99 The earliest generic name available for this segre- gate of Abutilon is Herissantia (See Brizicky, 1968). Sida ciliaris — Sida ciliaris L. var. ciliaris. Sida carpinifolia —^ Sida acuta subsp. carpinifolia (L.f.) Borss. Sida, spinosa, — Sida spinosa var. angustifolia (Lam.) Griseb. Sida hederaefolia > Sida javanensis Cav. emend. Borss. Recent treatments of Sida (Kearney, 1954b; Clement, 1957; Borssum Waalkes, 1966) do not all agree on treatment of the species in the Bahamas. I have followed Clement and Borssum Waalkes here. Malache scabra — Pavonia spicata Cav. var, spicata. Malache bahamensis > Pavonia bahamensis Hitchc. Pavonia is conserved over Malache. Kearney’s (1954a) treatment is followed here except for one point. He overlooked the fact that Pavonia scabra (B. Vogel) Juble et Quentin is a later homonym of P. scabra Presl. It must be replaced by P. spicata var. spicata. Malvaviscus sagraeanus > Malvaviscus arboreus var. mexicanus Schlecht. Schery (1942) has found an older name than the one given in B&M. Hibiscus bahamensis — Hibiscus brittonianus Kear- ney. Because H. bahamensis Britt. is a later homo- nym, Kearney (1954c) published a new name. Although a number of recent publications place okra in the genus Hibiscus as H. esculentus, Bates (1965) and Borssum Waalkes (1966) retained it in Abel- moschus, as in B&M, on the basis of the spathelike calyces which are basically adnate to the staminal tube and corolla, and which are circumscissilely de- ciduous. Pariti tiliaceus — Hibiscus tiliaceus L. subsp. elatus (Hochr.) Borss. Borssum Waalkes (1966) is followed in his treatment of this taxon. Gossypium barbadense — Gossypium arborescens var. nadam (Watt.) Prockh. 100 Rhodora [Vol. 76 214 Gossypium punctatum > Gossypium hirsutum var. 276 punctatum (Schum.) Hutch. Hutchinson (1943) is followed for the treatment of indigenous cotton STERCULIACEAE The use of names in Melochia has been thoroughly considered by Goldberg (1967) who considered Mo- luchia and Melochia synonymous. The only changes from B&M therefore are orthographic, Because the keys in B&M are inadequate, a new key to the Bahama species is given here: 1. Flowers in sessile glomerules, many per cluster; sinus between calyx teeth acute; fruit a 5-parted coccus less than 3.5 mm. in diameter ........ SNMP Melochia nodiflora Sw. . Flowers pedicelled, 3-10 per cluster; sinus be- tween calyx teeth rounded to truncate; fruit a 5-angled capsule, greater than 5 mm. in diameter .......... ee RII 2. 2. Capsule with rounded angles and unbranched hairs; inflorescences mostly in axillary cymes or appearing terminal; foliage leaves glabrous or with scattered, simple hairs; leaves only slightly lighter in color on lower surface than on upper ........... enne IM Melochia pyramidata L. Capsule sharply angled, with dense stellate hairs; inflorescences mostly opposite the leaves, never terminal; foliage leaves densely stellate; leaves tending toward being dis- colorous, much lighter on the lower surface than upper due to dense pubescence ...... 2.3232 *2»*«***3** Melochia tomentosa L. 278 Waltheria americana —^ Waltheria indica L. An ex- amination of Linnaean material and resultant lecto- typification has resolved the question of a name for 1974] 278 280 284 284 Bahama Flora — Gillis 101 the common Waltheria in accord with Brizicky (1966) and the example used in the International Code (Art. 57). The lectotype (chosen by Gillis) of Waltheria indica L. is in the Hermann Herbarium at BM. The lectotype of Waltheria americana L. is at LINN (Savage Catalog No. 852: 1). They represent the same taxon; both names originate from the same date (1753). Robert Brown (in Tuckey, Narr. Exp. River Zaire 484. 1818) appears to be the first to adopt the name W. indica L. for the combined species, and therefore should be followed. The descriptions and separation of the two species of Waltheria in B&M are ambiguous. The foliage of W. bahamensis is decidedly bronze in the field; her- barium specimens do not show this character well, but it is useful as a field character. The following key should assist in making determinations: Foliage green; branches of stellate hairs 2-4 mm. Ing 550500 Lot Waltheria indica L. Foliage bronze; pubescence in small tufts, branches of stellate hairs less than 1 MIN ...... ers Waltheria bahamensis Britt. Ayenia pusilla > Ayenia insulicola Cristobal. Except for an orthographic change necessitated by the Seattle Code, the Ayenia species should be named according to Cristobal (1960). HYPERICACEAE Ascyrum linifolium > Hypericum hypericoides (L.) Crantz. The nomenclature of Ascyrum has under- gone much discussion (Adams, 1957; Adams and Robson, 1961). The treatment of Adams and Robson (ibid.) has been followed here, FLACOURTIACEAE Myroxylon ilicifolium > Xylosma ilicifolia Northrop. Myroxylon bahamense —^ Xylosma bahamensis (Britt.) Standl. Xylosma G. Forst. has been conserved over 290 300 302 303 305 Rhodora [Vol. 76 Myroxylon J. R. et G. Forst. as a genus of Flacourtia- ceae (Myroxylon L. has been conserved in the Legu- minosae). The question of whether these two species are, in fact, distinct still needs to be explored further. PASSIFLORACEAE Passiflora pallida > Passiflora suberosa L. The two names above are synonyms of the same date; they were united by Killip (1938). See also Brizicky, 1961. Passiflora species in the Bahamas will be evaluated in a separate paper. LOASACEAE Mentzelia — consult Darlington (1934). LYTHRACEAE Parsonsia parsonsia — Cuphea parsonsia (L.) R. Br. Parsonsia is conserved for a genus of Apocynaceae; it therefore cannot be used for a genus of Lythraceae. COMBRETACEAE (TERMINALIACEAE) Conocarpus erecta > Conocarpus erectus L. The generic name Conocarpus should be treated as mascu- line in accordance with the International Code (Recommendation 75A). In Stearn (1966) the sub- stantive termination -carpus is consistently treated as masculine. Stearn (personal communication) sug- gests that Linnaeus may have treated Conocarpus as feminine by analogy with many classical second- declension names of trees. MYRTACEAE Eugenia buxifolia > Eugenia foetida Pers. The Myrtaceae need to be reworked in the light of Mc- Vaugh’s generic guidelines and definitions (1956; 1968). The change indicated above is made in accord with his analysis of types (McVaugh, 1973). Pimenta pimenta — Pimenta dioica Merr. This change is made to avoid a tautonym. 1974] 306 306 306 310 313 315 316 316 Bahama Flora — Gillis 103 Anamomis longipes > Psidium longipes (Berg) Mc- Vaugh var. longipes. Anamomis bahamensis — Psidium longipes var. orbi- culare (Berg) McVaugh. Anamomis lucayana — Myrcianthes fragrans (Sw.) McVaugh. The first two of these changes is in accord with the interpretation of these species by MeVaugh (1973). Psidium longipes var. orbiculare is further elaborated in the same paper by McVaugh by de- scribing unnamed races which can be distinguished. Myrcianthes fragrans is defined in an earlier paper by MeVaugh (1963). ONAGRACEAE Jussiaea suffruticosa > Ludwigia octovalvis subsp. sessiliflora (Micheli) Raven. This name change fol- lows the treatments of Ludwigia by Raven (1962; 1968). UMBELLIFERAE (AMMIACEAE) Foeniculum foeniculum — Foeniculum vulgare Gaertn. This change is made to avoid a tautonym. MYRSINACEAE Icacorea paniculata — Ardisia escallonioides Cham. et Schl. Icacorea guadalupensis > Ardisia obovata Desv. Ardisia is conserved over Icacorea. The Bahama plants need further study and interpretation, but they appear to be named best as stated above. Rapanea guianensis > Myrsine floridana A. DC. Rapanea guianensis Aubl. may be typified by a speci- men of Aublet's in P (lectotype of Gillis). It repre- sents a plant with lustrous, large leaves from northern South America. The northern West Indian and Flo- ridian species has dull, smaller leaves. Futhermore, it represents the group of species with a globose stigma, a character which has been used to separate Myrsine from Rapanea. Stearn (1969) made the 104 318 320 321 322 322 323 323 323 Rhodora [Vol. 76 combination Myrsine punctata (Lam.) Stearn for this species, but the name is a later homonym for a species of the Pacifie region which had been published several years earlier. The earliest available name for this plant is Myrsine floridana A. DC. If one prefers this species in Rapanea, then its name should be R. punctata (Lam.) Lundell. PRIMULACEAE Samolus floribundus — Samolus parviflorus Raf. This change is made in aecordance with Channell and Wood (1959) ; it is based on an earlier epithet. SAPOTACEAE Chrysophyllum — consult Cronquist (1945). Sideroxylon foetidissimum — Mastichodendron foeti- dissimum (Jacq.) Cronq. This change is made in ac- cord with the treatment by Cronquist (1946a). Lucuma serpentaria > Pouteria domingensis (Gaertn.) Cronq. var. domingensis. This change follows the treatment by Cronquist (19462). Dipholis salicifolia > Bumelia salicifolia (L.) Sw. Although Cronquist (1945) retained Dipholis as a segregate genus, Stearn (1968) merged it with Bumelia. It is the latter treatment which is followed here. Bumelia angustifolia > Bumelia celastrina H.B.K. Bumelia loranthifolia > Bumelia americana (Mill.) Stearn subsp. americana. Bumelia bahamensis > Bumelia americana (Mill.) Stearn subsp. americana. The Bumelia complex in the West Indies has been treated by Stearn (1968), a treatment followed here. The following key should help clarify the species in the Bahamas: 1. Ovary glabrous; endosperm copious; leaves longer than 7 cm.; terminal shoots never be- coming thorns .. Bumelia salicifolia (L.) Sw. 1. Ovary pubescent; endosperm little or none; 1974] Bahama Flora — Gillis 105 leaves shorter than 6 em. ; terminal shoots often thorny a C ok oie CE ELE 2. 2. Leaves narrow, linear to spathulate, 2-10 mm. broad (juveniles may be larger) ; fruit oblong; plant inhabiting shores, mangal, and SUNG areas. va ck ee ts re 2. Leaves obovate to oblanceolate, 1-4 mm. broad; fruits globose; inhabiting uplands .. A cR Bumelia americana, (Mill.) Stearn subsp. americana. 324 Mimusops emarginata — Manilkara bahamensis (Baker) Lam. et Meeuse. Probably no species in the Bahamas is easier to identify and yet harder to name than the wild dilly! In addition to the long list of synonyms given in B&M, it has been called Achras emarginata (L.) Little, Mimusops jamaiqui (Wright) Dubard, M. jamaiqui subsp. emarginata, (L.) Cronq., and Manilkara parvifolia (Nutt.) Dubard, to name a few. For the moment, I have selected Manilkara bahamensis (Baker) Lam. et Meeuse. Thus, it re- sides in the same genus preferred by Moore and Stearn (1967) for its close relative, the sapodilla, and also preferred by Cronquist (1945). 324 Sapota achras — Manilkara zapota (L.) P. van Royen. By the same token as with the wild dilly (above), the cultivated sapodilla has had a variety of names, many of which are permutations of each other. Moore and Stearn (ibid.) analyzed the problem thoroughly and leave a choice of names depending upon the breadth of one's generic concepts. Along with them, I choose the one indicated above. EBENACEAE 325 Maba crassinervis > Diospyros crassinervis (Krug et Urb.) Standl. Maba is treated as being a synonym of Diospyros. The name given above in Diospyros is temporary. A new combination will soon be pub- lished by Mr. Frank White (Oxford). 106 328 331 331 332 333 334 334 334 335 335 Rhodora [Vol. 76 OLEACEAE The Mayepea bumelioides of B&M will be placed in Chionanthus by Stearn at a later date, at which time a new combination will be published. GENTIANACEAE Based on Shinners’s typification (1957), the name of the marsh gentian remains the same, but the author of the binomial and place and date of publication change from what is given in B&M: Eustoma exalta- tum (L.) Salisb., Parad. Lond. t. 34. 1806. Sabbatia campanulata — Sabatia stellaris Pursh. Sabbatia simulata — Sabatia stellaris Pursh, Wilbur (1955) and Perry (1971) have studied the taxonomy of Sabatia in detail. As a consequence, one species only is considered to be in the Bahamas. The correct spelling of the generic name should also be noted. MEN YANTHACEAE Nymphoides aureum — Nymphoides grayana (Griseb.) Kuntze. This name change is made in accord with the treatment by Ornduff (1969). It should be noted that this so-called endemic Nymphoides (according to B&M) is also found in Cuba. APOCYNACEAE Plumiera obtusa — Plumeria obtusa L. var. obtusa. Plumiera inaguensis > Plumeria obtusa L. var. ob- tusa. Plumiera bahamensis —^ Plumeria obtusa L. var. ob- tusa. Plumiera rubra — Plumeria rubra L. Plumiera sericifolia, + Plumeria obtusa var. sericifolia (C. Wright) Woodson, Woodson (1938) reworked the species of Plumeria, taking up the correct spelling of the generic name. He considered several of the "species" of B&M to be insular races of P. obtusa var. obtusa. 1974] 335 336 336 337 337 538 Bahama Flora — Gillis 107 Woodson (1936) recognized Neobracea bahamensis as endemic, but also noted that there are three other species in Cuba, i.e, it is not a monotypic genus as stated in B&M. Stearn (1964) confirmed the name of the periwinkle as Catharanthus roseus (L.) G. Don, Echites echites > Echites umbellata Jacq. var. um- bellata. Rhabdadenia paludosa > Rhabdadenia biflora (Jacq.) Muell.-Arg. Rhabdadenia sagraei —^ Angadenia berteri (A. DC.) Miers. Urechites lutea — Urechites lutea var. serica Long. Woodson (1936) redefined the generic limits of a number of American Apocynaceous plants. His treat- ment is followed here. Long (1970a), in studying populations in South Florida, treated Urechites such that the Bahamian populations are recognized as a distinct pubescent population, U. lutea var. sericea. Because the Apocynaceous vines are not adequately keyed in B&M, and because the names of all of them have been changed, the following key has been prepared for use in separating them: KEY TO BAHAMIAN APOCYNACEOUS VINES 1. Corolla white or ivory, usually more than 5 cm. long; leaves remaining flat or folding along the midrib when pressed ; follicles of pair divergent, not tending to curve back like pincers, 4-10 mm. ihiek sou ily doe rer EDITI 2. 2. Follicles of a pair widely divergent (greater than 180?) ; tube of corolla salverform with rotate lobes; cylindric part of corolla 10-15 times the length of the sepals; calyx lobes free nearly to base, linear to linear-lanceolate . Echites umbellata Jacq. var. umbellata. 2. Follicles of a pair divergent less than 90°; tube of corolla funnelform; cylindric part of 108 359 389 Rhodora [Vol. 76 corola 2-3 times the length of the sepals; calyx lobes free scarcely more than 2/3 the length, ovate, to 2.5 mm. broad .......... .. Rhabdadenia biflora (Jacq.) Muell.-Arg. 1. Corolla yellow, usually less than 5 cm. long; leaves tending to curl under at the edges when pressed ; follicles of a pair diverging at an angle of less than 60°, curving back to touch near their tips, resembling a pair of pincers, 1.5-3 mm. thick .............eleele eh 3. 3. Vine scarcely 1 m. long; corolla 2-4 cm. long; leaves oblong, usually less than 1 cm. broad; pedicels glabrous; calyx lobes free less than half their length, 1-1.5 mm., acumi- nate; cylindric part of corolla visible most of its length (i.e, not hidden by sepals); petiole 2 mm. long ..................... TEE Angadenia berteri (A.DC.) Miers 3. Vine up to 3.5 mm. long; corolla 4-5 cm. long; leaves obovate or elliptic to 3 cm. broad; pedicels densely pubescent; calyx lobes free nearly to base, 8-12 mm. long, linear; cylindric portion of corolla hidden by sepals; petiole 1 em. long ............. ZEE Urechites lutea var. sericea Long Rauwolfia ` tetraphylla > Rauvolfia nitida Jacq. Rao (1956) thought that Rauvolfia tetraphylla is indige- nous to the Greater Antilles, Central America, and South America as far as Peru and Venezuela, The Bahamian species is R. nitida, which is not a synonym of R. tetraphylla as suggested in B&M. Vallesia glabra — Vallesia antillana Woodson. Wood- son (1937, 1938b) found that the true Vallesia glabra is a plant of Mexico and the Pacific Coast of South America. The species of southern peninsular Florida and the West Indies that has corollas twice the length of those of V. glabra needed a name. The name V. glabra of B&M is thus misapplied. 1974] 342 342 343 343 343 343 344 347 B47 347 Bahama Flora — Gillis 109 ASCLEPIADACEAE Metastelma | northropiae > Cynanchum northropiae (Schltr.) Alain, Metastelma hamatum > Cynanchum caribaeum Alain. Metastelma inaguense > Cynanchum inaguense (Vail) Howard et Dunbar. Metastelma linearifolium > Cynanchum savannarum Alain. Metastelma eggersii > Cynanchum eggersii (Schltr.) Alain. Metastelma palustre > Cynanchum angustifolium Pers. Metastelma species are considered synonymous with Cynanchum. (See Woodson, 1941; Alain, 1955). Much work needs to be done on the Bahamian species, but almost certainly several will be united. I shall refrain from making transfers to Cynanchum of several names which have not already been trans- ferred because I feel certain at this point that these names are synonyms of other names in the flora. (See also Howard and Dunbar, 1964). Merrill and Hu (1949) discovered that Muhlenberg had found Cynanchum angustifolium to be the eariest legitimate name for the last species listed above. They indicated the extensive synonyms for this species, a few of which had never ben included in the standard indices. Philibertella clausa > Sarcostemma clausum (Jacq.) R. & S. Sarcostemma is an earlier name than Phili- bertella (Woodson, 1941). CONVOLVULACEAE Evolvulus glaber —^ Evolvulus convolvuloides ( Willd.) Stearn. Evolvulus alsinoides > Evolvulus alsinoides var. gri- sebachianus Meissn. in Mart. Evolvulus linifolius > Evolvulus alsinoides var. lini- folius (L.) Baker. Stearn (1972) has modified the nomenclature of several species of Evolvulus in the flora in the course of typification of their names. The 110 255 2356 357 Rhodora [Vol. 76 last taxon listed above is now believed not to occur in the flora. Jacquemontia jamaicensis ^ Jacquemontia havanensis (Jacq.) Urb. Dr. Kenneth Robertson (personal com- munication) has pointed out that this name change is required because of an earlier basionym. Calonyction aculeatum — Ipomoea alba L. Calonyction tuba — Ipomoea macrantha R. et S. Both Ooststroom (1940) and Gunn (1972) have worked with Calonyction. Gunn’s nomenclature is followed here. Quamoclit quamoclit ^ Ipomoea quamoclit L. Quamoclit coccinea > Ipomoea hederifolia L. Quamo- clit, like Calonyction, is treated as a section of Ipo- moea. (see Ooststroom, 1953). Ipomoea cathartica — Ipomoea acuminata (Vahl) R. et S. Ipomoea pes-caprae > Ipomoea pes-caprae subsp. bra- Siliensis (L.) Ooststr. This treatment of Ipomoea follows that of Ooststroom (1940) and St. John (1970). The citation for Ipomoea stolonifera in B&M is incorrect; there is an earlier publication date of the combination: Ipomoea stolonifera (Cyrill.) Gmel., Syst. Veg. I: 345. 1796. Ipomoea dissecta + Merremia dissecta (Jacq.) Hall. f. Merremia is segregated from Ipomoea on the basis of its smooth pollen and its corolla without a red or purple eye (O'Donell, 1941; Ooststroom with Hoog- land, 1953; Verdcourt, 1963). Dichondra. Consult Tharp and Johnston (1961). CUSCUTACEAE Cuscuta pentagona — Cuscuta campestris Yuncker. This species needed a new name because of confusion in the old one (Yuncker, 1932). HYDROPHYLLACEAE Marilaunidium jamaicense > Nama jamaicensis L. 1974] 357 358 358 359 359 360 361 362 363 365 366 Bahama Flora — Gillis 111 Nama is an earlier name than Marilaunidium. Fur- thermore, Nama jamaicensis L. is the conserved type species. BORAGINACEAE (EHRETIACEAE) Sebesten sebestena — Cordia sebestena L. Varronia globosa > Cordia globosa var. humilis (Jaeq.) Johnston. Varronia bahamensis > Cordia bahamensis Urb. Varronia brittonii > Cordia brittonii (Millsp.) Mac- bride. Varronia lucayana — Cordia lucayana (Millsp.) Mac- bride. Sebesten and Varronia should be treated as synonyms of Cordia (Macbride, 1916; Johnston, 1949). Rochefortia bahamensis — Rochefortia spinosa (Jacq.) Urb. According to Lefor (1968), the Rochefortia in the Bahamas is not an endemic species, but the same as that found in Cuba. There is debate over the generic selection for bay lavender. For the moment, I maintain Mallotonia, as in several other recent floras (Gooding, Loveless, and Proctor, 1965; Adams, 1972). Heliotropium parviflorum — Heliotropium angiosper- mum Murray. The change of name is required because H. angiospermum is an earlier name. Heliotropium inundatum — Heliotropium procumbens Mill. Heliotropium procumbens is an earier name for the same species. Both names can be typified by specimens at BM; the lectotypes have been selected by me. VERBENACEAE Valerianoides fruticosa > Stachytarpheta fruticosa (Millsp.) B. L. Robinson. Valerianoides jamaicensis —> Stachytarpheta jamai- censis (L.) Vahl. Stachytarpheta is conserved over Valerianoides. 112 366 367 368 368 369 369 371 Rhodora [Vol. 76 Bouchea prismatica > Bouchea prismatica var. longi- rostra Grenzen. This treatment follows the revision of Bouchea by Grenzenbach (1926). Moldenke (1971) believes that the typical variety is also present in the Bahamas. Priva — consult Kobuski (1926). Lippia stoechadifolia > Phyla stoechadifolia — (L.) Small. Lippia nodiflora — Phyla nodiflora (L.) Greene var. nodiflora. Lippia reptans — Phyla nodiflora var. reptans (H.B.K.) Moldenke. These changes are made in accord with the treatment of Phyla by Moldenke (1965). Lippia geminata of B&M is probably not found within the archipelago. The appropriate treatment of this bi- nomial would be Lantana microcephala A. Rich. in any event. According to treatment by Long (1970a) our variety of Lantana ovatifolia would be the typical variety (var. ovatifolia). Using Moldenke's treatments (1958a, b, and c) of Citharexylum, I have prepared the following key to distinguish the Bahamian species: Pedicels less than 1.2 mm. long during anthesis; pyremes 2-loculate, fruit not shiny; leaves + pilose, closely reticulate-veined, the reticulation prominent above and below ... Citharexylum fruticosum L. Pedicels 2 mm. long or more during anthesis; pyremes 1-loculate, fruit shiny; leaves glabrous, widely reticulate-veined, reticulation obscure above Citharexylum caudatum L. Moldenke (1958a; 1971) has considered 5 infra- specific taxa of C. fruticosum to exist in the Bahamas: var. fruticosum, var. smallii Moldenke, var. subvil- losum Moldenke, var. villosum (Jaeq.) O. E. Schultz, and forma bahamense Moldenke, Having made field 1974] 373 373 374 375 376 377 378 378 Bahama Flora — Gillis 113 studies on the characters upon which these taxa are based, I consider these to be insular forms or popula- tions without names. For those who wish to name these variations, an appropriate key is found in Moldenke (1958a). Callicarpa — consult Moldenke (1936). Petitia — consult Moldenke (1937). Volkameria aculeata —^ Clerodendrum aculeatum (L.) Schlecht. var. aculeatum. Volkameria is treated by Moldenke (1971) as a section of Clerodendrum. The place of citation for C. aculeatum is incorrect in B&M. The author of the combination is Schlechten- dahl, and the citation is Linnaea 6: 750. 1831. Clerodendrum fragrans > Clerodendrum philippinum Schauer. Howard and Powell (1968) found that C. fragrans was an illegitimate name. Moldenke (1971) considers the Bahamian population to be var. pleni- florum Schauer. Avicennia nitida — Avicennia germinans (L.) L. Stearn (1958) found this earlier epithet which ap- plies to the species of black mangrove in the West Indies, but failed to note that its earliest combination was by Linnaeus himself. LABIATAE (LAMIACEAE) Melosmon cubense — Teucrium cubense Jacq. Epling (1925) united Melosmon and Teucrium. Leonurus — consult Epling (1925). The place of publication for Leonotis nepetifolia (L.) R.Br. is the Prodromus florae Novae Hollandiae, p. 504. 1810, and not as given in B&M (Epling, 1925). In Epling’s treatment (1938-39), Salvia serotina L. and S. micrantha Vahl (indicated as synonyms in B&M) were separated on rather minor characters. Through the courtesy of Dr. Mildred Mathias and Dr. Raymond Harley, I have seen an unpublished manuscript of Epling and Carlos Jativa in which 114 385 388 Rhodora [Vol. 76 these species were reunited. It seems best to continue to treat them as synonymous. (Permission of Carlos Jativa to refer to this unpublished material is ac- knowledged with appreciation.) On the basis of rather minor characters of the calyx, Shinners (1962) reworked the Micromeria brownei complex, recognizing the Bahamian populations as M. bahamensis Shinners. Because my own collections from Inagua more closely resemble the taxon which Shinners cites from Hispaniola, I believe more work needs to be done before the matter is resolved. Chiefly because of the minor nature of the characters which are used in defining Shinners's species, I continue to recognize a broadly treated M. brownei (Sw.) Benth. SOLANACEAE Physalis angulata > Physalis angulata L. var. angu- lata. Physalis turbinata — Physalis cordata Mill. Physalis pubescens > Physalis barbadensis Jacq. These changes are in accord with the treatment by Waterfall (1967). Solanum verbascifoltum — Solanum erianthum D. Don. Roe (1968) found that the old name for this plant did not apply to the species in our flora. Lycium spathulifolium > Lycium tweedianum var. chrysocarpum (Urb. et Ekm.) Hitche. In the mono- graph of Lycium (Hitchcock, 1932), the Bahamian plant required a name change. Although it is possible that the species of the Florida Keys (L. carolinianum Walt.) is in the Bahamas, it has yet to be found. SCROPHULARIACEAE Maurandya antirrhinaeflora > Maurandya antirrhini- flora H. et B. ex Willd. This orthographic change is required by the Seattle Code. I can now report this species to be on the far side of the archipelago from New Providence as recorded in B&M: Gillis 11780 from Grand Turk. 1974] 389 390 392 393 394 395 399 401 Bahama Flora — Gillis 115 Bramia monnieri > Bacopa monnieri (L.) Pennell. Bacopa has been conserved over Brami and Moniera. It is an earlier name than Bramia. Although Pennell (1935) segregated Erinus from Mecardonia, I believe that Mecardonia is best treated as an inclusive genus, thus maintaining Mecardonia procumbens (Mill.) Small as in B&M. Afzelia cassioides > Seymeria cassioides (Walt.) Blake. Seymeria is conserved over Afzelia. Buchnera elongata —> Buchnera floridana Gandoger. Philcox (1965) thinks that B. elongata is a mis- applied name for the Florida and Bahama popula- tions. LENTIBULARIACEAE Setiscapella subulata — Utricularia subulata L. Stomoisia cornuta — Utricularia cornuta Michx. Tay- lor (1967) includes Setiscapella and Stomoisia within Utricularia. BIGNONIACEAE Macrocatalpa punctata — Catalpa punctata Griseb. var. punctata. This change is made in accord with the treatment of Catalpa by Paclt (1952). PEDALIACEAE Sesamum orientale —^ Sesamum indicum L. These names are synonymous binomials of the same publica- tion date. Sesamum indicum L. has been used more commonly than the other name. Hill (1939) pointed out that DeCandolle (Pl. Rar. Jard. Genève 18, t.5 — 1889) was the first to unite the two names under Sesamum indicum. ACANTHACEAE Blechum brownei should be retained despite the at- tempt of some to change the name of this species to Blechum pyramidatum. The Linnaean binomial is Ruellia bltechum, Lamarck renamed the species Bar- 116 402 402 403 404 407 408 409 410 Rhodora [Vol. 76 laria pyramidata, but this name is illegitimate, being superfluous when published. Jussieu later published a new name in Blechum; this is accepted because the tautonym which would be created by transferring the Linnaean epithet is illegitimate (Bremekamp, 1938; Long, 1970b). Gerardia droseroides > Stenandrium droseroides Nees in DC. Gerardia bracteosa — Stenandrium bracteosum (Britt. et Millsp. Britt. ex Leonard). Stenandrium is con- served over Gerardia. Anthacanthus spinosus — Oplonia spinosa (Jacq.) Raf. Stearn (1971) has studied the generic complex of which this species is a part and has recognized our plant to be an Oplonia. Diapedium assurgens Dicliptera assurgens (L.) Juss. Dicliptera is conserved over Diapedium (see also Long, 1970b). I have chosen the lectotype to be at LINN (No. 28:23 according to the Savage Catalog, 1945). RUBIACEAE The Rubiaceae are in need of in-depth treatment as a whole. Names accepted here may be subject to re- vision as more work is done. Oldenlandia callitrichoides > Hedyotis callitrichoides (Griseb.) Lewis. Merrill and Metcalf (1942) and Lewis (1961) have united Oldenlandia and Houstonia with Hedyotis. Lewis’s treatment is followed here. Rachicallis americana — Rhachicallis americana (Jacq.) Hitche. The original spelling of the generic name has been restored. Despite the fact that it has been written without the first “h” ever since the original publieation, there seems no reason not to accept the original orthography considering the Greek origin. Nickerson and Tripp (1973) have reported this species to have either male or perfect flowers, never female. Randia mitis — Randia aculeata L. These two names, 1974] 411 412 413 413 414 415 Bahama Flora — Gillis 117 synonyms of the same date of publication, have been united under R. aculeata. Catesbaea, parviflora — Catesbaea parviflora var. sep- tentrionalis Krug et Urb. ex Urb. The Bahamian populations fit this variety better than do the typical species form. Hamelia erecta > Hamelia patens Jacq. These names are synonyms of the same date of publication. Thomas Elias (personal communication) has informed me that L’Héritier was the first to unite them, and should be followed. Guettarda taylori —^ Guettarda nashii Britt. et Millsp. Guettarda inaguensis > Guettarda nashii Britt. et Millsp. There does not seem to be sufficient difference to maintain three distinct species of Guettarda sup- posedly endemic to the vicinity of Matthew Town, Inagua. Having studied both the type specimens (at NY) and material in the field, I have concluded that Britton and Millspaugh named specimens, not species. Guettarda inaguensis is represented by a type that appears to have come from an aberrant form with long lengths of bare branches and leaves only near the tips. Guettarda taylori is represented by a type that has no flowers left and for which no fruits were ever known. Because of the inadequancy of these two specimens, it seems best to select the name for the united species by the type possessing the most ade- quate material. Inasmuch as these three species names originate from the same data, I am choosing G. nashii as the name to be used when all three are united. The populations of Guettarda in the vicinity of Matthew Town, although variable, appear to me to be best treated as a single species. Stenostomum lucidum — Antirhea lucida (Sw.) Hook. f. in Benth. et Hook. f. Stenostomum myrtifolium — Antirhea myrtifolia (Griseb.) Urb. 118 Rhodora [Vol. 76 415 Stenostomum densiflorum — Terebraria resinosa (Vahl) Sprague. Stenostomum as used by B&M is better divided into Antirhea and Terebraria. Antirhea is an older name than Stenostomum and hence must be the name used for those species traditionally placed in Stenostomum. The tortuous history of the name of Terebraria resinosa is recounted thoroughly by Sprague (1932). Sprague recognized two West In- dian species of Terebraria, differing only in shape of the areoles of the leaf venation. In his treatment, the Bahamian population would be T. densiflora. I consider these differences to represent only insular forms, not worthy of nomenclatural separation. A key to the species in the Bahamas of the old Stenos- tomum follows: 1. Ovary 2 (-4) loculate; flowers without bracts; anthers half-exserted; stipules deciduous; foli- age without resinous secretions .... Antirhea lucida (Sw.) Hook. f. in Benth. et Hook. f. 1. Ovary 4-6 loculate; flowers subtended by minute bracts; anthers included; stipules persistent; foliage resinous-viscid ................0-. 2. 2. Leaves 3 cm. long or less, elliptic to obovate- oblong; diffusely-branched shrub with leaves crowded near ends of branches; midvein very pronounced, often white; 2-4 flowers per branch of inflorescence; stipules ovate, not forming a collar around stem; inflorescences paired on either side of stem, but not branched ................ eee 2. Leaves 4-9 cm. long, oblong-lanceolate ; coarsely-branched shrub or small tree with leaves widely scattered; midvein only slightly more pronounced than lateral veins; more than 6 flowers per branch of inflorescence; stipules forming a collar 2-4 mm. broad 1974] 419 420 421 Bahama Flora — Gillis 119 around stem; inflorescence bifid with flowers only on inside (adaxial surface) of branches Terebraria resinosa (Vahl) Sprague. Psychotria undata > Psychotria nervosa Sw. Jacquin described P. undata in the Plantarum rariorum horti caesarei schoenbrunnensis. According to Stafleu (1967), the title page date of 1798 is probably in error, and actual date of publication may have been as late as 1803. In any event, Swartz in his Prodro- mus (1788) published the name Psychotria nervosa for the same species, clearly the earliest date of pub- lication. Psychotria nervosa Benth, (1841) and P. nervosa D. Don (1825) are later homonyms. Ernodea cokeri > Ernodea taylori Britton. Ernodea nashii o Ernodea millspaughii Britton. It is difficult to make hard-and-fast decisions on the Ernodea problems in the Bahamas at this juncture. Whether E. littoralis and E. angusta are truly distinct is questionable. Long (1970a) has treated E. angusta as a variety of E. littoralis which may be the best way to handle these perplexing taxa. Until intensive study is made of these two species with comparisons to the other Ernodea taxa in the Bahamas, I shall continue to recognize these two as species. Distinc- tions among the other four species do not seem to hold up when examined under field conditions. Al- though more study will be carried out on Ernodea in the Bahamas, under a tentative arrangement which seems reasonable at this stage, E. cokeri is considered to be a synonym of E. taylori, and E. nashii is merged with E. millspaughii. The latter name was chosen from the two originating at the same date of publi- cation because it is represented by the type specimen possessing both flowers and fruits. The type of E. taylori has no flowers at all; the type of E. nashii must have had flowers at the time Britton described the species, but they are missing from the type 120 422 425 426 Rhodora [Vol. 76 specimens now (types are at NY). The differences as observed in the type specimens of these micro- species seem inadequate for maintenance in this ex- ceedingly variable genus (also see Britton, 1908, for descriptions of “races”? within this complex). A key to identify the Bahamian species of Ernodea within my species concept at the moment follows: 1. Calyx lobes nearly as long as the fruit, or longer .............. RR RRRRRRI 2. 2. Corolla white to very pale pink .......... SENE Ernodea littoralis Sw. 2. Corolla red ...... Ernodea angusta. Small. 1. Calyx lobes less than half as long as the fruits 2.0... 0 eee 3. 3. Leaves 6-8 mm. wide ................... MEIN Ernodea millspaughii Britton. 3. Leaves 1-9 mm. wide ............... less ZEE Ernodea taylori Britton. Borreria saxicola > Borreria brittonii Standl. Un- like Ernodea, the eight species of Borreria in the Bahamas — or, at least most of them — seem to be good biological species with a high degree of ende- mism in the southern islands, especially Inagua and the Caicos group. Whereas actual treatment of this genus is deferred for the moment, one name change needs to be cited. Following Standley (1931), the name B. saxicola Britt. is replaced because it is a later homonym. Spermacoce — consult Bacigalupo (1972). CUCURBITACEAE Anguria pedata > Psiguria pedata (Jacq.) Howard. In his treatment of modern names for plants dis- cussed in Jacquin’s Selectarum, Howard (1978) found that the name Anguria Jacq. was a later homonym for Anguria Mil. The next available generic name is Psiguria; Howard made the neces- sary new combinations for this species, 1974] 428 429 431 432 432 436 440 441 443 Bahama Flora — Gillis 121 LOBELIACEAE Lobelia — consult MeVaugh (1943). GOODENIACEAE Scaevola plumierii > Scaevola plumieri (L.) Vahl. This orthographic change is made to conform to the present International Code. COMPOSITAE (AMBROSIACEAE AND CARDUACEAE) Xanthium chinense — Xanthium strumarium L. Xanthium strumarium is an earlier name than X. chinense. Ambrosia paniculata —^ Ambrosia artemisiifolia L. I have selected a lectotype of Ambrosia paniculata from material in the Michaux Herbarium at Paris. Furthermore, I have examined the lectotype of A. artemisiifolia which was selected by Payne (1970) at LINN, and concur with Payne that these plants are conspecific. The Linnaean name is obviously the older one and should be used. Iva — consult Jackson (1960). Ageratum latifolium —> Ageratum conyzoides subsp. latifolium (Cav.) M. F. Johnson. This name change has been made in accord with the treatment by John- son (1971). The typical subspecies has also been reported from the Bahamas, thus adding a taxon to the flora. Chrysopsis graminifolia > Heterotheca graminifolia (Michx.) Shinners. Shinners (1951), Wagenknecht (1960), and Harms (1964) have argued for the merger of Heterotheca and Chrysopsis. Aster bracei — Aster tenuifolius var. aphyllus Long. Long (1970a) has treated the Bahamian populations in connection with those of South Florida which he was studying. This taxon seems best treated as a variety of the more widespread A. tenuifolius. Leptilon linifolium — Conyza floribunda H.B.K. 124 Rhodora [Vol. 76 CUPRESSACEAE (PINACEAE) 462 Juniperus lucayana — Juniperus bermudiana L. More work needs to be done on typification and on popula- tion studies in Juniperus to place the Bahamian plant properly. For the moment, it is treated in accord with Moore, 1966. CYCADACEAE 463 A thorough monograph of Zamia is needed. In a manuscript by Chamberlain, left unpublished at his death, Z. lucayana has been referred to the Cuban Z. guttierezii Sauv. So little material of this plant, as well as of Z. angustifolia and Z. tenuis, is avail- able at this time, that it is not appropriate to make a nomenclatural judgment now. The recent redis- covery of the Long Island population of “Z. lucayana" by S. R. Hill (in press) and its subsequent cultiva- tion will help to make material available for further study. LITERATURE CITED ACUNA, J. and J. T. Rorc. 1962, Psychotriae Cubenses Novae (Rubiaceae). Brittonia 14: 224-229. ADAMS, C. D. 1970. Miscellaneous additions and revisions to the flowering plants of Jamaica. Phytologia 20: 309-14. 1971. Miscellaneous additions and revisions to the flowering plants of Jamaica. III. Phytologia 21: 405-10. 1972. Flowering plants of Jamaica. University of the West Indies, Mona, Jamaica. 848 pp. ADAMS, WILLIAM P. 1957. A revision of the genus Ascyrum (Hy- pericaceae). Rhodora 59: 73-95. ADAMS, WILLIAM P. and NoRMAN K. B. RoBSON. 1961. A re-evalua- tion of the generic status of Ascyrum and Crookea (Gutttiferae). Rhodora 63: 10-16. ALAIN, HERMANO. 1962. Flora de Cuba, Vol. 5 — Rubiales - Valeri- anales - Cucurbitales - Campanulales - Asterales. Editorial Uni- versitaria, Universidad de Puerto Rico, Rio Piedras. ALLEN, CAROLINE K. 1966. Notes on Lauraceae of tropical America. I. The generic status of Nectandra, Ocotea, and Pleurothyrium. Phytologia 13: 221-27. 1974] Bahama Flora — Gillis 125 AMES, OAKES. 1910. A new Ponthieva from the Bahamas. Torreya 10: 90-91. BACIGALUPO, NELINDA M. 1972. Observaciones sobre algunas es- pecies de los géneros Spermacoce L. y Spermacoceodes O.K. (Rubiaceae). Darwiniana 17: 341-57. BAEHNI, C. 1965. Mémoires sur les Sapotacées. III. Inventaire des genres. Boissiera 11: 1-262. BALDWIN, J. T., JR. 1938. Kalanchoé: the genus and its chromo- somes. Amer. J. Bot. 25: 572-79. Bates, DAvID M. 1965. Notes on the cultivated Malvaceae. I. Hi- biscus. Baileya 13: 56-130. BEAUVOIS, A. M. F. J. PaLisoT DE. 1812. Essai d'une nouvelle Agrostrographie. Paris. BECKNER, JOHN. 1970. Notes on Florida and West Indian orchids. Phytologia 20: 217. BELCHER, R. O. 1956. A revision of the genus Erechtites (Compos- itae) with inquiries into Senecio and Arrhenechthites. Ann. Mis- souri Bot. Gard. 43: 1-85. BoGLE, A. LINN. 1970. The genera of Molluginaceae and Aizoaceae in the southeastern United States, J. Arnold Arbor. 51: 431-62. BoRSSUM, WAALKES, J. VAN. 1966. Malesian Malvaceae revised. Blumea 14: 1-213. BRASHIER, CLYDE K. 1966. A revision of Commelina (Plum.) L. in the U.S.A. Bull. Torrey Bot. Club 93: 1-19. BREMEKAMP, C. E. B. 1938. Notes on the Acanthaceae of Surinam. Rec. Trav. Bot. Neerl. 35: 130-170. + 4 pl. BRENAN, J. P. M. 1958. New and noteworthy Cassias from Tropical Africa. Kew Bull. 13: 231-252. BRITTON, N. L. 1908. The genus Ernodea Swartz: a study of species and races. Bull. Torrey Bot. Club 35: 203-208. 1911. The botanical name of the wild sapodilla. Torreya 11: 128-29. 1918. Flora of Bermuda. Charles Scribner's Sons. New York. and CHARLES FREDERICK MILLSPAUGH. 1920. The Bahama flora. Published by the authors. Reprinted 1962, Hafner Publishing Co., New York. and JOSEPH NELSON RosE. 1928. Rosales - Mimo- saceae. North American Flora. Vol. 23, Pt. 1. 1923-30. Scientific survey of Puerto Rico and the Virgin Islands. Botany of Puerto Rico and the Virgin Islands. Descriptive flora. Part 1 — New York Acad. 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Crossability values within the slash-Caribbean Pinus species complex. Euphytica 19: 184-89. SPRAGUE, T. A. 1932. Laugeria “Vahl” = Terebraria Kuntze. Kew Bull. 1926: 349. STAFLEU, FRANS A. 1967. Taxonomic Literature. Regnum Vegeta- bile 52. 556 pp. Utrecht. STAFLEU, FRANS et al. 1972. International Code of Botanical No- menclature. Reg. Veg. 82: 1-426. STEARN, WILLIAM T. 1957. The boat-lily (Rhoeo spathacea). Bai- leya 5: 195-98. 1958. A key to West Indian mangroves. Kew Bull. 1958: 33-37. 1964. Catharanthus roseus, the correct name of the Madagascar periwinkle. Lloydia 27: 196-200. 1966. Botanical Latin. Thomas Nelson and Sons, Ltd. London. 566 pp. 1968. Jamaican and other species of Bumelia (Sapotaceae). J. Arnold Arbor. 49: 280-89. 1969. A synopsis of Jamaican Myrsinaceae. Bull. Brit. Mus. Nat. Hist. Bot. 4: 143-78 -- 8 pl. 1971. A survey of the tropical genera Oplonia and Psilanthele (Acanthaceae). Bull. Brit. Mus. Nat. Hist. Bot. 4: 259-323. 1974] Bahama Flora — Gillis 137 1972. Typification of Evolvulus nummularius, E. convolvuloides, and E. alsinoides (Convolvulaceae). Taxon 21: 647-50. STUCKEY, RONALD L. 1972. Taxonomy and distribution of the genus Rorippa (Cruciferae) in North America. Sida 4: 279-430. TAYLOR, PETER. 1967. Lentibulariaceae. Mem. N.Y. Bot. Gard. 17: 201-28. THARP, B. C. and MARSHALL C. JOHNSTON. 1961. Recharacteriza- tion of Dichondra (Convolvulaceae) and a revision of the North American species. Brittonia 13: 346-60. TRELEASE, W. 1918. Agave in the West Indies. Mem. Natl. Acad. Sci. 11: 1-55. 1916. The genus Phoradendron: a monographie re- vision. Univ. Illinois Press. Urbana. VERDCOURT, B. 1963. Convolvulaceae. in: Flora of Tropical East Africa, C. E. Hubbard and E. Milne-Redhead. eds.; Crown Agents for Overseas Governments and Administrations. WAGENKNECHT, BURDETT L. 1960. Revision of Heterotheca, section Heterotheca (Compositae). Rhodora 62: 61-76; 99-107. WATERFALL, U. T. 1967. Physalis in Mexico, Central America, and the West Indies. Rhodora 69: 82-102; 203-40; 319-29. WEBSTER, GRADY L. 1956-58. A monographie study of the West Indian species of Phyllanthus. J. Arnold Arbor. 37: 91-122; 217- 68; 340-59; pls. 1-14; 38: 51-80; 170-98; 295-373; pls. 15-23; 39: 49-100; 111-212; pls. 24-32. 1967. The genera of Euphorbiaceae in the southeastern United States. J. Arnold Arbor. 48: 303-61; 363-430. 1970. A revision of Phyllanthus (Euphorbi- aceae) in the continental United States. Brittonia 22: 44-76. WHEELER, Louis C. 1938. The names of three species of Brassica. Rhodora 40: 306-309. WIJNANDS, D. O. 1973. Typification and nomenclature of two spe- cies of Sansevieria (Agavaceae). Taxon 22: 109-114. WILBUR, ROBERT L. 1955. A revision of the North American genus Sabatia (Gentianaceae). Rhodora 57: 1-33; 43-71; 78-104. WILSON, KENNETH A. 1960. The genera of Convolvulaceae in the southeastern United States. J. Arnold Arbor. 41: 298-317. WINDLER, D. R. 1966. A revision of the genus Neptunia (Legum- inosae). Aust. J. Bot. 14: 379-420. Woop, CARROLL E., JR. 1958. The genera of the woody Ranales in the southeastern United States. J. Arnold Arbor. 39: 296-346. 1959. The genera of the Nymphaeaceae and Ceratophyllaceae in the southeastern United States. J. Arnold Arbor. 40: 94-112. 138 Rhodora [Vol. 76 and R. B. CHANNELL. 1960. The genera of the Ebenales in the southeastern United States. J. Arnold Arbor. 41: 1-35. Woopson, ROBERT E., JR. 1936. Studies in the Apocynaceae, IV. the American genera of Echitoideae. Ann. Missouri Bot. Gard. 23: 169-438 + 7 pl. 1937. New or otherwise noteworthy Apo- cynaceae of tropical America, V. Ann. Missouri Bot. Gard. 24: 11-16. 1938a. Studies in the Apocynaceae. VII. An evaluation of the genera Plumeria L. and Himatanthus Willd. Ann. Missouri Bot. Gard. 25: 189-224. 1938b. (Asclepiadales) Apocynaceae. North Amer. Flora 29: 103-192. YAKOVLEV, G. P. 1967a. Systematical and geographical studies of the genus Sophora L. and allied genera. Prob. Pharmacog. 21: 42-62. (In Russian). 1967b. Some characteristics of structure of seeds of Sophora L. and allied genera in connection with its systematics and phylogeny. Repr. Proc. Leningrad Chem.-Pharm. Inst. 21: 90-98. (In Russian). YATES, HARRIS O. 1966. Revision of grasses traditionally referred to Uniola I. Uniola and Leptochloópsis. Southwestern Nat. 11: 372-394. YUNCKER, TRUMAN GEORGE. 1932. The genus Cuscuta. Mem. Tor- rey Bot. Club 18: 113-331. ARNOLD ARBORETUM, HARVARD UNIVERSITY CAMBRIDGE, MASS. 02138 AND DEPARTMENT OF BIOLOGY UNIVERSITY OF MIAMI CORAL GABLES, FL. 33124 STONE'S PLANTS OF SOUTHERN NEW JERSEY: A REPRINT' To those who had always considered Witmer Stone as an ornithologist — and one of the country's greatest — the appearance in 1911 of his “Plants of Southern New Jersey" came as a distinct surprise. Had they known more about the man they would have realized that he was one of the last of the all-around naturalists, interested in mammals, insects and plants as well as birds. The recent facsimile reprint of Stone's book, by the Quarterman Publications, Inc., of Boston, makes available once more a work which for more than sixty years has been the standard reference for everyone who is interested in the flora of the Pine Barrens, as well as the other plant provinces of southern New Jersey. Originally published as Part II of the Annual Report of the New Jersey State Museum, this important volume has for many years been out of print and has, indeed, be- come something of a collector's item. Although one can not be other than grateful that this flora is once again in circulation, it is to be regretted that the Foreword was not written by a professional botanist, preferably one who was intimately acquainted with Witmer Stone. Since this reprint is a facsimile, every word, every punc- tuation mark and every misprint appears exactly as it did in the original. There are numerous discrepancies between its nomenclature and that in use today. It would have been helpful to the reader if Elizabeth M. Woodford, who wrote the Foreword, had explained the reasons for this. Partly because he was a zoologist and partly because of his friendship with Dr. N. L. Britton of the New York Botanical Garden, Stone's treatment of plant names follows the now-abandoned American Code of Botanical] Nomen- 'Stone, Witmer. The Plants of Southern New Jersey. Reprint. 1973. Quarterman Publications, Inc., Boston, Massachusetts. 828 pp. Illust. $25.00. 139 140 Rhodora [Vol. 76 clature. This code, like that of the zoologists, permits the use of tautonyms or duplicate binomials. Hence we find in Stone such combinations as Blephariglottis blephari- glottis, Hepatica hepatica, Linaria linaria and others. The American Code never embraced the principle of the conservation of generic names and Stone therefore con- sistently employed what he considered to be the earliest generic epithets. As a consequence we have scores of generic names which are unfamiliar to most present-day taxonomists. A few examples will suffice: Spathyema for Symplocarpus, Juncoides for Luzula, Abama for Narthe- cium, Vagnera for Smilacina, Gyrostachys for Spiranthes, Hicoria for Carya, Ammodenia for Arenaria (peploides), Meibomia for Desmodium, etc. Fortunately most of the presently accepted names are cited in synonymy (although not always with the correct authority), but it seems a pity that a republication of this work should not have suggested some revision and up- dating of its nomenclature. It might also have corrected such misprints as “Sandy Hood” for “Sandy Hook" (page 432) and “dandelon” for “dandelion” (page 817). Mrs. Woodford states that Stone’s collection of 12,000 specimens went to the herbarium of the Academy of Natu- ral Sciences of Philadelphia. Such is not the case. Realizing that many of his duplicates were already in the Academy and that he had frequently been in the field with members of the Philadelphia Botanical Club whose specimens had likewise been deposited there, I suggested to Dr. Stone that he leave his herbarium to the University of Pennsylvania. He readily agreed, and in the fall of 1942 the University received his collection of more than 14,000 sheets, rich not only in plants from southern New Jersey, but also in ma- terial from northern Pennsylvania. It was my pleasant task to supervise the incorporation of this collection into the herbarium of the University. Witmer Stone once told me that his manuscript for the New Jersey Flora had been prepared in a little more than four months. Doubtless he had made some preliminary 1974] Book Review — Fogg 141 efforts to put together his observations accumulated over many years of field work, but even so this was a prodigious accomplishment. He added that the Director of the New Jersey Museum had approached him in desperation to give them something for their Annual Report and urged him to prepare a paper on the Pine Barrens of New Jersey. Stone's reply was that this would be like treating one vegetational province in a vacuum and that he preferred to deal with the entire flora of the lower three-fifths of the state, at the same time emphasizing the geographic origins of the various elements represented therein. This he did superbly well, in a manner which certainly would have elicited the approbation of Asa Gray. Mrs. Woodford gives a picture of Witmer Stone as a warm, friendly human being with a wide breadth of in- terests and a well developed sense of humor. With this i can heartily agree. It was my good fortune for many years to spend my summers at Cape May, where Dr. Stone also had a cottage, and in many walks with him I came to appreciate his encyclopedic knowledge and great capacity for friendship. JOHN M. FOGG, JR. ARBORETUM OF THE BARNES FOUNDATION MERION, PA. 19066 THE ASIATIC SPECIES OF DESMODIUM: A REVIEW A new series of monographie studies, Ginkgoana,' was inaugurated early in 1973 with the publication of Number 1, “The Asiatic species of Desmodium and its allied genera (Leguminosae)," by Hiroyoshi Ohashi of the Department of Botany, University of Tokyo. The first study in the series is truly a book, printed in a clear and pleasing type face, and very well edited. The text is written in English. This study is a portion of the thesis presented by Mr. Ohashi for the degree of Doctor of Science. It is an inclusive treatment with clearly stated conclusions. Since the genus Desmodium (in the broad sense) is one in which I have long had a particular interest and since I know the Asiatic species less well than the American and African, I am pleased to see this detailed piece of work. Of special importance are the pollen studies and the meticu- lous investigations of floral parts, seeds, and seedlings. Dr. Ohashi confirms the conclusions of Ambrose? for Des- modium (i.e. the species of Desmodium ser. Americana have hypogaeous germination, while those of other species ob- served have an epigaeous type of germination), and pre- sents new information for additional species. The chapter considering observations on morphological characters demonstrates well the care and scope of Dr. Ohashi's observations on habit, leaves, inflorescences, bracts, bracteoles, androecia or stamens, pollen grains, fruits [*^pods"], seeds, seedlings, and chromosome numbers. He has illustrated with clarity the basic structures and prob- able evolutionary progression in the development of the 'Ginkgoana, Contributions to the Flora of Asia and the Pacific Region. No. 1, The Asiatic Species of Desmodium and its Allied Genera (Leguminosae), by Hiroyoshi Ohashi, 318pp., 86 figures, 76 plates [18.1 X 25.6 em.]. February 15, 1973. Academia Scientific Book Ine., Tokyo, Japan. "The Michigan Botanist 6: 97-99. 1967. 142 1974] Book Review — Schubert 143 inflorescence for the group (Desmodium sens. lat.) and also of the bracts and bracteoles. In both cases the tendencies indicated could be compared profitably with parallel or similar progressions in the New World and African groups of related genera. The resulting comparisons, particularly with New World genera and species, should be of much interest because of the numerous groups of closely related species which radiate from the centers of diversity in Mex- ico and Brazil. The taxonomie treatment has well constructed keys, de- tailed descriptions, ample citations of specimens, exceed- ingly good illustrations of floral parts and of fruits, and distribution maps. In general, I think Dr. Ohashi has rather over-classified the group he has studied so carefully, follow- ing Schindler and Hutchinson in large part. However, he has documented his data and stated his reasons so carefully that his occasionally debatable conclusions do not detract from the usefulness of the work. The treatment of Des- modium subgenus Sagotia is perhaps the one which troubles me most. Dr. Ohashi says “The subgenus Sagotia is one of the most polymorphic and widespread groups of the genus Desmodium. It is characterized by having mostly the ter- minal racemes composed of 2(-3)-flowered fascicles which are subtended usually only by the primary bracts (i.e. lack- ing the secondary bracts), ebracteolate flowers, diadelphous androecia and rather small-sized articles of the fruits.” He includes, however, species with secondary bracts, among them Desmodium velutinum, D. gangeticum, and D. dichoto- mum; some with bracteoles, such as D. caudatum and D. benthamii; some with flowers borne singly; and some with loment articles of considerable size. It seems to me that the only reason the genus Sagotia (sensu Ohashi) is so polymorphic is that Dr. Ohashi has assigned to it very unlike elements which do not have a close natural relationship. His arrangement may seem feasible when studying only Asian species, but on a world wide basis I think it would become wholly unworkable and its defects would be much magnified. In some infrageneric 144 Rhodora [Vol. 76 groups in Asia as for most of the genus Desmodiwm in Africa there are many unrelated, distinctive species in con- trast to the situation in North and South America where there are many small to large groups of closely related species. This makes for a very different aspect of a pre- sumed phylogeny, of proposed evolutionary series, and for an understanding of interspecific relationships in Asia and Africa on the one hand and in America on the other, even though there are some species which bridge the gap by natural or artificial means. The only serious omissions in this careful study are in the areas of ecology and phytogeography, discussion of the former being almost totally lacking and of the latter only minimal. This may indicate a lack of field experience which can be remedied later by an active field program. Field work might also give Dr. Ohashi a broader view of the taxa with which he is working and a somewhat different ap- proach to their interrelationships. In concluding this review I would stress the meticulous care of the observations Dr. Ohashi has made and the great gap he has filled in our knowledge of one large complex in the family Leguminosae. BERNICE G. SCHUBERT ARNOLD ARBORETUM HARVARD UNIVERSITY 22 DIVINITY AVENUE CAMBRIDGE, MASS. 02138 A MAGNIFICENT SWISS FLORA MANUAL' A REVIEW One of the most important tasks of botanists specializing in taxonomy and geobotany is the compilation of flora manuals for their colleagues in general and for the much more numerous amateur botanists in particular. In this way, the author himself gets a supplementary survey over the plants of the region in which he works and gathers experience that enables him to solve problems of unusual importance. At the same time he gets a rare opportunity to stimulate interest in his subject and to gather new prose- lytes among the younger generation to carry on in his foot- steps. Only a few botanists have the encyclopedic knowl- edge and ability to write floras, and still fewer want to take the risk of being unfairly criticized for this least gratifying of scientific occupations, because in no other field is it easier to be criticized for opinions and judgements. This is the perennial trouble for compilers of floras and also the main cause of the scarcity of good manuals. If the author also ventures to make deviations from the accepted delimita- tions, descriptions and names of the plants included, he runs the additional risk of condemnation by his more con- ventional colleagues, because scientists are no less con- servative than are other citizens. Ordinary manuals are always valuable, but unconventional floras ought to be doubly welcome, because nothing else can stimulate future research to a higher degree. Although Switzerland is one of the smaller European countries, it has fostered astonishingly many of the botani- cal giants. Several of its botanists certainly have been con- ventional and followed the trend of their times, whereas others have dared to have new ideas and have looked at the 'Hans Ernst Hess, Elias Landolt, Rosemarie Hirzel: Flora der Schweiz und angrenzender Gebiete. Three volumes, 29 X 23 cm., 2690 pages, with 9 colored plates and about 3450 drawings. Birk- háuser Verlag, Basel & Stuttgart, 1967, 1970, 1972. Price 450 Schweizer francs. 145 146 Rhodora [Vol. 76 plants of their country from new points of view. These latter we have to thank for the unusually large number of presently well-known species that were first described from Switzerland. Although many of the great Swiss botanists of the past are now always mentioned with reverence, there were times when their contemporaries called them names and were critical about their scientific abilities. Such is the fate of those who are born too early or try to be prophets in their own land, but time is their confederate. It would be difficult to mention all the many flora man- uals, which have been published in Switzerland in the past, and the list of those which have reached fame far outside the country because of scientific quality is far from being short. One such flora was Albrecht von Haller’s great *Historia stirpium indigenerorum Helvetiae inchoata", which was published in 1768 in three folio volumes. That work has stood in a class by itself for two centuries as the most monumental botanical manual ever written in the field of descriptive floristies. It still remains important for its clear descriptions and other information, but the conserva- tism in nomenclature and the reluctance to accept what would be the future in that field has detracted from the nomenclatural importance of the work though it always will be a text of literary superiority rarely achieved in scientific books. Although many good Swiss floras have been published since Haller's work, two centuries had to pass before a text of similar magnitude and quality again was compiled for this flora of more than 3500 species. This is the new “Flora der Schweiz," by H. E. Hess, E. Landolt and R. Hirzel, the three large volumes of which were published in 1967, 1970 and 1972. It is a work of extraordinary quality and quan- tity, based on fifteen years of intensive work by the two first authors and on equally intensive work of drawing the excellent pictures of more than 3450 of the species by the third author. The virtues of this manual are too many to be even sim- ply listed, and readers with different inclinations will al- 1974] Book Review — Lóve 147 ways find some matters of special interest. Considerable information on the methods and philosophy adopted in the compilation of this great work is given in an introductory chapter, which then is followed by almost thirty pages on the history and origin of the flora and its distribution, and more than forty pages on its ecology. Specialists are apt to find omissions in these chapters, and some may perhaps join issue with the authors as to some of the explanations given. That is to be expected in any work of this magnitude and concentration. But this reviewer, who believes he has gathered considerable knowledge about this flora through visits and reading numerous good books and reports during the past three decades, finds these chapters to be highly educational and of great interest from more points of view than he cares to mention. Numerous maps in these intro- ductory chapters ought to make it easy for those unfamiliar with the geography of the land to follow the detailed ac- count. The manual itself is arranged according to the Engler system as are most modern European floras. A general key to the higher groups leads to keys to the families of the major groups where these commence, and keys to genera and species are then given in appropriate places in the text. Families and genera are described in detail, followed under the latter by information about the size of the genus and its distribution, and about cytological information avail- able. After the generic key there are detailed reports of possible hybridization observed, frequently with explana- tions of the morphological, geographical, and cytological characteristics of the plants. Most species are represented by a good drawing, fre- quently with additional details. Their Latin names are followed by synonyms, if any, and German names, and a very detailed description is given of the morphological characteristics. For every species there is also information on chromosome number and other cytological peculiarities, and the authors have done an admirable work in looking up the original papers in order to be able to exactly inform the 148 Rhodora [Vol. 76 reader about the geographical origin of the material. This is the only place where such information has been compiled, and it is admirably corect, although some few mistakes have not been avoided. Then there are reports on the ecological occurrence of each species in Switzerland, and on the gen- eral distribution of the taxon elsewhere in the world. Fre- quently, the authors then add taxonomical and geographical remarks on the variation of the species within and outside their country, especially when they feel that it is to be regarded as a critical taxon in need of more close investiga- tion, with references to the most recent literature in every special case. It is easy to find points of disagreement in a work of this magnitude, although this reviewer is of the opinion that the authors have sueceeded unusually well in avoiding harsh judgements. They have made an effort to accept a modern species concept although sometimes this may not have suc- ceeded well enough, and the descriptions of the taxa are clear and moderately concise. It is a pity that they have found it impossible to mention subspecific categories, but since Switzerland abounds in such variations which fre- quently may be of doubtful value, it is understandable that the authors avoided spending perhaps twice as much time in evaluating these. The only serious remark to be made seems to be that the authors adopted, for present-day European botanists, an unusually conservative attitude towards the nomenclature, so that sometimes their eccentricity may look disturbing. Nevertheless, this is understandable, and the explanation of this given in the last volume is not unrea- sonable although I believe that a more liberal attitude would have made the work better and also more useful for a longer period of time. After the description of the last species in the third vol- ume, there are more than 90 pages of corrections and addi- tions, half a page with an epilogue, where some criticism of plant sociology and nomenclature by reviewers of the first two volumes are met, and four pages on which the termi- nology used is explained. There are good indexes of Latin 1974] Book Review — Lóve 149 and German names, and these are preceded by sixty pages of a very important bibliography. The three volumes are printed in the unusual size 29 X 23 em, with considerable empty space on every page that is partially used for the large drawings. The heavy volumes are strongly and tastefully bound in green linen, and they are certainly reasonably priced for their size and quality. Their weight would make it difficult to carry them into the field, but the authors are compiling a smaller manual with the keys only and new drawings of selected species that will be a handy companion on excursions in the Alps and elsewhere. The “Flora der Schweiz” is an unusually thorough and scholarly work which will long be the standard reference for this part of Europe. It is also an invaluable source of information for botanists from other parts of the continent, and also for specialists in other fields, especially that of cytotaxonomy. The text and pictures are so detailed and exact that the book can be highly recommended to those foreign botanists and institutions which need good and reliable information on European plants which may have been introduced overseas, because no other flora comple- ments the comprehensive Flora Europaea more excellently for southern and central European plant species. ÁSKELL LÓVE 473 HARVARD LANE BOULDER, COLORADO 80303 INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA Manuscripts must be double-spaced or preferably triple- spaced (not on corrasable bond), and a list of legends for figures and maps provided on a separate page. Footnotes should be used sparingly, as they are usually not necessary. Do not indicate the style of type through the use of capitals or underscoring, partieularly in the citations of specimens, except that the names of species and genera may be under- lined to indicate italies in discussions. Specimen citations should be selected critically especially for common species of broad distribution. Systematic revisions and similar papers should be prepared in the format of “The System- aties and Ecology of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- larly with reference to the indentation of keys and syno- nyms. Papers of a floristic nature should follow, as far as possible, the format of ‘Contribution to the Fungus Flora of Northeastern North America. V.," H. E. Bigelow & M. E. Barr, Rhodora 71: 177-203. 1969. For bibliographic cita- tions, a recommended list of standard journal abbreviations is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300. 1958. Volume 76, No. 805, including pages 1-150, was issued March 30, 1974. 150 CONTENTS: — continued Studies on the Bryophytes of Southern Manitoba III. Collections from Grand Beach Provincial Park Paul W. Stringer and Muriel H. L. Stringer .................... 27 Some New or Noteworthy Vascular Plant Records from Northwestern Saskatchewan Vernon L. Harms and John H. Hudson ...............5.4....« 39 Parasitic Witchweed: Striga asiatica Versus S. lutea (Scrophulariaceae) F. N. Hepper eee eene ennt ent tna th tta anne tnntn seen tine th atas tna ta nan 45 Cacalia rugelia: A New Combination for a North American Senecionoid T. M. Barkley and Arthur Cronquist ............. eee 48 Typification and Application of the Names Scirpus ameri- canus Pers. S. olneyi Gray, and S. pungens Vahl. Alfred E. Schuyler ............ eee eee eee eee eene te nent tn nennen nnn 51 The Correct Authority for Cardamine clematitis (Cruciferae) y JU C209 207,177 NERS EE EATE AA IE E E T 53 A New Form of Digitaria sanguinalis John E. Ebinger .rsccssosssssssessssssssecccserscsssnesensseterevssesacccssssssescescnns 57 The Growth of some New England Perennial Seaweeds Narman EEO ER aai iori d diassec itaq E a 59 Chromosome Numbers in Kosteletzkya Presl. (Malvaceae) Orland J. Blanchard, Jv. sscsssccersssescersrsssccccoveccessonconecssenseseceees 64 Name Changes for Seed Plants in the Bahama Flora William T. Gillis eaea aa eaan A 67 Book Review JORN EOI: IT: aeien oicseeri tis reei E 139 Book Review Bornio G uSOhWDOT 5 un udis aszsessensoes iic iaa neem: 142 Book Review Fer) ASE GS LEPC TYE) S EO E E ERE E TEM NE Dodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ROLLA MILTON TRYON ) RADCLIFFE BARNES PIKE STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY ALFRED LINN BOGLE RICHARD EDWIN WEAVER œ Associate Editors L Vol. 76 June, 1974 No. 806 CONTENTS: A Systematic Treatment of the Native Unifoliolate Crotalarias of North America (Leguminosae) Donald R. Wiéndler ....cccccrccccscescsssssercosscessconessscssecceccssscessccosasences 151 The Flora of Mount Monadnock, New Hampshire Er o e E A E T E E E T 205 Taxonomy of Perityle Section Perityle (Compositae-Peritylinae) NH MB AR eia A E A E ER A E A E 229 Carex exilis, Geocaulon lividum and Other Plants of Interest in Pittsburg, New Hampshire BEEN TONTO n A n a N eiii 307 (Continued on Inside Back Cover) The Nem England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02138 RHODORA. — A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price $10.00 per year, net, postpaid, in funds payable at par in the United States currency at Boston. Some back volumes, and single copies are available. For information and prices write RHODORA at address given below. Scientific papers and notes, relating directly or indirectly to the plants of North America, will be considered by the editorial com- mittee for publication. 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MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS QTRbooora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 76 June, 1974 No. 806 A SYSTEMATIC TREATMENT OF THE NATIVE UNIFOLIOLATE CROTALARIAS OF NORTH AMERICA (LEGUMINOSAE)"* DONALD R. WINDLER Crotalaria is a large genus of tropical, subtropical, and temperate plants comprised of about 550 species, over 400 of which occur in Africa. In 1939 Senn reported 31 species for North America with the majority growing in Mexico and the West Indies; only 21 of these species were con- sidered to be native. Eight of the 21 species, including the Linnean C. sagittalis, form a group that is so distinctive that Rafinesque (1837) separated it into the subgenus Iocaulon. The Iocaulon? Crotalarias are annual or perennial plants characterized by simple leaves; decurrent stipules, when present; terminal or leaf-opposed inflorescences ; and glabrous, inflated fruits that become black at maturity. Senn (1939) indicated that this is the most difficult group of North American Crotalarias in which to delimit species and suggested the need for further work. 'A portion of a dissertation submitted as partial fulfillment of the requirements for the Doctor of Philosophy Degree in the Department of Botany, University of North Carolina at Chapel Hill. "Contribution number 7 from Towson State College Herbarium. Cost of Publication of illustrations assisted by the Towson State College Faculty Research Fund. "The name Iocaulon will be used as an adjective throughout this paper to refer to the unifoliolate Crotalarias related to C. sagittalis. 151 152 Rhodora [Vol. 76 The history of the classification of Crotalaria has been reviewed by Baker (1914), Senn (1939), and Polhill (1968). While Baker treated the African species of Crota- laria primarily, he also reviewed the history of the classi- fication of the genus and discussed its separation from the closely related genus Lotononis. Senn reviewed rather briefly the nomenclatural history of Crotalaria in North America, and discussed the placement of the North Amer- ican species into sections described by Baker. Senn's treat- ment includes a key to the species, nomenclature and synonomy, Latin descriptions for newly described species, lists of specimens examined, and a few comments on each of the taxa. Most species lack descriptions and all lack illustrations, which makes it difficult to identify a plant if the key only narrows the choice to a group of species, Polhill (1968), while dealing primarily with the African species of Crotalaria, presented a discussion of the history of generic subdivisions, discussed typification of the genus, and proposed a new arrangement of species into sections based on combinations of flower characters. The characters used were: (1) type and position of standard appendages, (2) twisting of the keel, (3) lobing of the calyx, (4) calyx- keel length ratio, (5) pubescence of the style, (6) style shape, and (7) receptacle size. Polhill placed C. sagittalis and its relatives in the section Calycinae Wight & Arnott. All the characters that Polhill used for this placement were found to be consistently present in the Iocaulon Crotalarias studied. COLLECTIONS STUDIED In addition to the personal collections made in the United States and Mexico, specimens were borrowed from twenty- one herbaria. The names and locations of the herbaria from which specimens were borrowed appear below, ac- companied by the abbreviation of each as listed in Index Herbariorum, except in one case, marked by an asterisk, where no listing could be found for the herbarium and an arbitrary abbreviation is used. 1974] Crotalarias — Windler 153 A— Arnold Arboretum, Cambridge, Massachusetts. ALU — University of Alabama, University, Alabama. AUA — Auburn University, Auburn, Alabama. GA — University of Georgia, Athens, Georgia. GH — Gray Herbarium, Cam-. bridge, Massachusetts. LAF — University of Southwestern Louisiana, Lafayette, Louisiana. MICH — University of Michigan, Ann Arbor, Michigan. *MISS — University of Mississippi, University, Mississippi. NCU — University of North Carolina, Chapel Hill, North Carolina. NY — The New York Botanical Garden, New York, New York. SIU — Southern Illinois University, Carbondale, Illinois. TEX — University of Texas, Austin, Texas. UC — University of California, Berkeley, California. us — United States Na- tional Museum, Washington, D.C. usF— University of South Florida, Tampa, Florida. wis — University of Wis- consin, Madison, Wisconsin, BM — British Museum of Natural History, London, Eng- land. G-— Conservatoire et Jardin Botanique, Geneva, Switzerland. K-— Royal Botanic Gardens, Kew, Great Britain. MEXU — Universidad Nacional de México, Mexico City, Mexico. P — Museum Nacional d'Histoire Naturelle, Paris, France. Crotalaria plants from more than 60 locations in the United States and Mexico were grown from seed at the North Carolina Botanical Garden, Chapel Hill, during the summer of 1968. Data from the garden plantings were used in evaluating the status of taxa. Another paper de- tailing the results of the field and garden studies is published elsewhere (Windler, 1973). SYSTEMATIC TREATMENT CROTALARIA (Dillenius ex Linnaeus, Hort. Cliff. p. 218, 1731, pro parte) L., Sp. Pl. p. 714 pro maxima parte. Type species: Crotalaria lotifolia L. (see Polhill, 1968 pp. 174-5). GENERAL DESCRIPTION OF THE NATIVE IOCAULON CROTALARIAS Erect, ascending, spreading, decumbent, or procumbent 154 Rhodora [Vol. 76 terrestrial herbs or shrubs. Root a taproot with slender branches, nodules frequently present. Stems single in the erect species to many in the decumbent and procumbent species, the epidermis green, frequently becoming purple, with an indument varying from nearly glabrous to serice- ous, of spreading or appressed, simple trichomes, Stipules absent or varying from a pair of subulate lobes projecting from near the leaf base to inverted-sagittate, decurrent, foliaceous appendages which wing the stem for a portion of the subtending internode, a full internode, or for several of the subtending internodes, the lobes spreading, incurved toward the stem or parallel to the stem. Leaves alternate, unifoliolate, undivided, entire, ciliate, membranaceous to fleshy, broadly ovate to lanceolate or linear, 0.9-10.2 cm. long, 0.3-4.6 cm. wide, the adaxial surface dark green, gla- brous, or with spreading or appressed, simple trichomes, the abaxial surface usually lighter green than the adaxial and with spreading or appressed simple trichomes; petioles 0.5-3 mm. long. Inflorescences loose racemes, peduncles borne terminally, or opposite leaves and 0.0-3 cm. below the node. Bracts lanceolate, elliptic, subulate, or linear, to 14 mm. long, one at the base of each flower, frequently persistent after flower or fruit abscission. Flowers papilionaceous. Calyces bilabi- ate, deeply cleft, the upper two lobes free and broader than the lower three, the lower usually remaining coherent, the calyx bearing two lateral bracteoles, one on either side. Standards (vexillum, banner) orbicular, obovate, or ellip- tic-oblong, the apex rounded or retuse, 4 mm. shorter to 6 mm. longer than the upper calyx lobes, yellow, the adaxial surface frequently becoming red-tinged, lined or unlined, nearly glabrous with a few trichomes along the main vein on the adaxial surface, with lamelliform appendages at the base of the abaxial surface of the blade. Wings (alae) ob- long, attached by a stalk continuous with the lower margin (cleaver-shaped), the blades usually puckered between some of the veins, yellow. Keel (carina) petals twisted at the tip, white or greenish-white, lanate-ciliate. Stamens 10, 1974] Crotalarias — Windler 155 monadelphous, the staminal tube split on the upper side, anthers dimorphie, with basifixed long anthers alternating with medifixed short anthers, the short anthers positioned above the elongate at anthesis. Styles geniculate at the base, bearing a spiral line of trichomes transversing the length and culminating in a tuft of trichomes on the stigma. Fruit an inflated, sub-cylindric, short-stipitate legume, glabrous inside and out, becoming black at maturity. Seeds 7-46 per pod, oblique-cordiform, smooth. The systematic treatment presented here covers twelve native North American unifoliolate species of Crotalaria section Calycinae. The descriptions are based on materials collected during the study and on numerous herbarium specimens. Measurements of stipule width are taken at the base of the sinus between the two lobes and include the width of both at that point. Length measurements of the calyx and standard are made from the base of the torus to the tip of the respective structure in the undissected flower. Illustrations were prepared by Mrs. Jerry Meyer. Since data on cross-fertility between the numerous populations sampled is not available, taxa have been dis- tinguished primarily on patterns of morphological discon- tinuity. It is assumed that these discontinuities also reflect breeding discontinuities, especially in areas where species are sympatric. In mountainous areas, however, many pop- ulations are well isolated, frequently leading to morpholog- ical discontinuity between populations. In these instances greater emphasis has been placed on similarities in inter- preting relationships. Diagnostic characters that have been useful in the group are plant habit; general size; position of inflorescence; presence, absence, size, and shape of the stipules; length and position of trichomes; and flower size. Most of the diagnostic characters are present on the plants during the entire growing season. The synonomy includes only those names that were pos- sible to check either against types or clear descriptions. A 156 Rhodora [Vol. 76 special effort is made to point out all places where the present treatment differs from that of Senn's. l A set of duplicate specimens from the author’s field col- lections has been deposited in the herbarium of the Univer- sity of North Carolina at Chapel Hill (NCU). KEY TO THE SPECIES OF IOCAULON CROTALARIAS WHICH OCCUR IN NORTH AMERICA A. Inflorescence terminal, or terminal and also leaf- opposed. ...........leeee RR heh B. B. Plants with decurrent stipules. (West Indies, Cen- tral and South America). ........... 1. C. pilosa. B. Plants without decurrent stipules. ............ C. C. Shrubs and subshrubs with diffuse habit. .. D. D. Peduncles 9 cm. or more long; calyx 15-20 mm. long. (Mexico, Central and South America). .......... esses 2. C, nitens. D. Peduncles 6 cm. or less long; calyx 10-11 mm. long. (Northwestern Mexico). ..... 3. C. brevipedunculata. C. Herb with decumbent or ascending habit. (Western Mexico). ........ 4. C. polyphylla. A. Inflorescence not terminal, always leaf-opposed. ..... eee e hehe ehe E. E. Plants erect or strongly ascending. ........... F. F. Stem pubescence spreading; adaxial leaf sur- faces regularly pubescent. ............... G. G. Stipules with lobes recurved. (West Indies and South America). .... 5. C. stipularia. G. Stipules with lobes spreading, (Through- out). co.cc eee eee ee eee 6. C. sagittalis. F. Stem pubescence appressed, adaxial leaf sur- faces glabrous or with appressed trichomes. H. Upper leaf surface glabrous. ......... I. I. Plants perennial; stipules usually con- spicuous; peduncles 7.5-23 cm. long. 1974] Crotalarias — Windler 157 (Southern United States). .......... NEP EL o M NC 7. C. purshii. I. Plants annual; stipules absent or incon- spicuous; peduncles 2.5-8 cm. long. (Mexico and Central America). ...... "T c IEEE 8. C. quercetorum. H. Upper leaf surface bearing trichomes. .. J. J. Peduncles 0.5-2 cm. long, usually as thick as the stem; stipules present; trichomes dense, obscuring the stem. HMETICO)S S.V VI 9. C. mexicana. J. Peduncles 2.5 cm. long or longer, thinner than the stem; stipules absent; tri- chomes not obscuring the stem (except occasionally near the top). (Southern United States). .. 10. C. rotundifolia. E. Plants decumbent, procumbent or spreading. .. K. K. Mature plants decumbent or procumbent from a thick taproot, stems usually less than 6 dm. long; trichomes on stem 0.3-3.2 mm. long. (Mexico and Southeastern United States). .. eer er ere o o 10. C. rotundifolia. K. Mature plants diffuse or spreading, stems usu- ally more than 6 dm. long; trichomes on stem usually less than 1 mm. long. (Mexico). .. L. L. Calyx 13-20 mm. long; stipules 0.5-2.7 cm. broad. (Mexico). .. 11. C. bupleurifolia. L. Calyx 7.5-12 mm. long; stipules narrow or absent. (Mexico). .. 12. C. nayaritensis. 1. Crotalaria pilosa Miller, Garden Dict. No. 2. 1768, non C. pilosa Roxb. ex Mart. Denkschr. Konigl. Akad. Wiss. München 6: 156, 1920, non C. pilosa Thunb. Prod. Pl. Capensium 125. Crotalaria pterocaula Desv. J. Bot. (Desvaux) 3: 76. 1814. Type: not located. Crotalaria genistella HBK. Nov. Gen. et. Sp. Pl. 6: 398. 1824. Type: Humboldt 2050 (P, mc Microfiche observed). 158 Rhodora [Vol. 76 ac a Figure 1. Crotalaria pilosa. A. Habit and leaf variation B. Stem 159 Crotalarias — Windler 1974] Su2QWw '*7) 'suo2vM ‘DUNAS '? pue soyd +4 ‘syion16 ‘IBA ‘IVA SUazU 7140]0304/) jo uonnqujsuiq ‘T dey .. 06 001 On JEBEUE Pad | | Blueindijs'9 o | | eSo|Id') e | | Ssijioe48 uea suajiu *) X | su3]iu ') 9 j | |o hí a ee or we $ o | — 1 ia | ‘ NE. ® go | f MSN l a) f ? X | Qo. Tr, | q | 4. | $ | ` | 1 | re Mw | | | | | | | | | $ | ie I| 1 Ñ \ Ü e | | x | | | \ ? | n \ A | | ct pe b MED oL 08 06 ~ 001 E oll 160 Rhodora [Vol. 76 Crotalaria pilosa Miller var. skutchii Senn, Rhodora 41: 331. 1939. Type: Skutch 3071 (GH). Annual erect herb with a taproot to 0.4 cm. thick. Stem single, 2.8-9.5 dm. long, 1.3-4 mm. thick, the longest inter- node 2.8-8.2 cm. long; vestiture of appressed or loosely ap- pressed trichomes 0.5-1.5 mm. long. Stipules present, de- current more than the length of the internode, 0.3-0.8 cm. wide, the width constant, the lobes 2.1-9.7 mm. long, usually not spreading. Leaves elliptic, lanceolate, narrowly lanceo- late, or linear, 3.3-8.6 cm. long, 3.5-25 mm. wide, base rounded to cuneate, the apex acute; vestiture of both sur- faces of appressed trichomes 0.4-2 mm. long; petioles 1-1.3 mm. long. Inflorescences terminal or leaf-opposed and ter- minal, peduncles 4.5-12 cm. long. Bracts sessile, linear, or subulate, 3.1-4 mm. long, 0.3-0.5 mm. wide; pedicels 3-4.8 mm. long. Calyces 9-11.5 mm. long, the tube 2.5-3.1 mm. long, with appressed, loosely appressed, or somewhat spreading trichomes 0.5-1 mm. long; bracteoles elliptic, narrowly elliptic, or subulate, 2.5-4 mm. long, 0.4-0.5 mm. wide. Corollas yellow, the standards 9-12.8 mm. long, equaling or to 1.7 mm. longer than the upper calyx lobes; elongate anthers 1.5-2 mm. long, short anthers 0.3-0.5 mm. long ; style 5.5-5.8 mm. long. Fruits 2.5-3.6 cm. long, breadth not known; seeds tan, brown, or red-brown, 15-23 per pod, 2.9.5 mm. long. Chromosome number: n—16. Flowering time: July-March. Habitat: sandy and gravelly soils in fields and open pine woods, elevation sea level to 2,400 feet. Range: Costa Rica, Cuba, Jamaica, Panama, (Also widely distributed in South America.) Figure 1, Map 1. Type: “La Vera Cruz in New Spain" (Mexico, Cuba or Jamaica?), Houstoun (BM; photo., NY!). In spite of the fact that Miller, in his original descrip- tion, cited Crotalaria pilosa as “growing naturally at La Vera Cruz in New Spain", no specimens have been observed from Veracruz, Mexico. The citation appears to be in error. The probable true type locality was either in Cuba or Jamaica. Senn, in his treatment of C. pilosa, recognized the species as being composed of three varieties. Senn's 1974] Crotalarias — Windler 161 variety skutchii is here treated as being a part of variety pilosa; his variety robusta is transferred to C. bupleurifolia and treated as a variety of that species. No field collec- tions were made of C. pilosa during this study and no seed was obtained for garden plantings. Crotalaria pilosa is characterized by its erect habit, ter- minal or leaf-opposed and terminal inflorescences, and stipules that wing the stem for more than one node below the node of origin (Figure 1). Occasional specimens have spreading lobes at the summit of the stipules. Such speci- mens observed are Britton, Britton, & Shafer 734 (NY) and Bro. Leon & Fr. M. Rosca 8837 (NY), both from Cuba. Crotalaria pilosa is alone among the species covered in this treatment in possessing both terminal inflorescences and stipules. REPRESENTATIVE SPECIMENS: COSTA RICA. Cina Grande, entre San Ramon y Otevas, Brenes 11332 (NY). CUBA. Isle of Pines: pinelands, Britton, Britton, and Wilson 14615 (NY, US); PINAR DEL RIO: near Rio de las Vueltas, N of S. Diego de los Banos, Bro. Leon 4908 (GH). DOMINICAN REPUBLIC. TRUJILLO: between Baya- guana and Guerra, Howard and Howard 9924 (cH). JAMAICA. Upper Clarendon, James Hill Savannah, Harris 12844 (GH, NY, US). PANAMA. Ancon Hill, Canal Zone, open hillside, Killip 12075 (us). TRINIDAD. Without date or specific locality, de Suber 369 (P). 2. Crotalaria nitens HBK. Nov. Gen. et Sp. 6: 399. 1824. Crotalaria bracteata Schlecht. et Cham. Linnaea 5: 575. 1830, non Roxburgh, Hort. Bengal 54. 1814. (Type: Schiede 597, Hacienda de la Laguna, (B; photo., F!). Crotalaria schiedeana Steudel, Nomen. Botanicus. 1: 445. 1840. Type: same as for C. bracteata for which this was a nomen nov. Shrub with unknown root type. Stems 5-15 dm. long, 2-4 mm. thick, the longest internode 1-5.5 em. long; vestiture of appressed or loosely appressed trichomes, 0.5-1 mm. long. Stipules absent. Leaves elliptic, lanceolate, oblong, or oblanceolate, 2.5-7.5 (10) em. long, 3.5-18.5 mm. wide, base rounded to cuneate, the apex rounded, acuminate, or acute; vestiture of both surfaces of appresed or loosely appressed 162 Rhodora [Vol. 76 trichomes, 0.3-1 mm. long; petioles 1.3-3 mm. long. Inflor- escenses terminal, peduncles 6-51 cm. long. Bracts petiolate, lanceolate, the petioles 0.3-1 mm. long, the blades 4-14 mm. long, 0.6-3 mm. wide; pedicels 4.1-7 mm. long. Calyces 15-18 (20) mm. long, the tubes 3.3-5 mm. long, bearing appressed or loosely appressed trichomes 0.3-1 mm. long; bracteoles lanceolate, elliptic-lanceolate or linear, 4.5-9 mm. long, 0.5-1.4 mm. wide. Corollas yellow, the standards 13.5-19 (22) mm. long, equaling to 2.5 mm. longer than the upper calyx lobes; elongate anthers 2.2-3.6 mm. long, short anthers 0.6-1.2 mm. long; styles 8.5-11.5 mm. long. Fruits 2.5-3.2 em. long, breadth not known; seeds grey-brown, brown, or red-brown, 26-40 per pod, 1.8-2.8 mm. long. Chromosome number: not known. Flowering time: Sep- tember-December. Habitat: bare soil, dry open mountain- sides and pinelands, elevation 4,000-7,200 feet. Range: Guatemala; Mexico: Chiapas, Jalisco, Oaxaca, Veracruz (Also widely distributed in South America). Type: Mariquita (Colombia), Humboldt (P, nc Micro- fiche observed). A photograph of the type of Crotalaria nitens was ob- served in the Ipc microfiche photographs of the Humboldt, Bonpland, and Kunth herbarium (P). It bears only the name C. nitida and the location Mariquita which is also mentioned in the location sites for C. nitens. It compares favorably with the photograph examined by Senn (1939) of the “co-type” in the Willdenow Herbarium. Senn’s appli- cation of the name is followed in this treatment. Crotalaria nitens is characterized by its shrubby habit, lack of stipules, terminal inflorescences, and large flowers. Of the North American species, it most resembles C. brevi- pedunculata but differs from it in having a larger flower size and longer peduncles. Two varieties are recognized: Peduncles stout, 1-2 mm. thick; bracts 7.5-14 mm. long, 1.2-3 mm. wide. .......... 2a. C. nitens var. nitens. Peduncles slender, 0.5-0.6 mm. thick, bracts 4-5 mm. long, 0.6-0.8 mm. wide. ........ 2b. C. nitens var. gracilis. 1974] Crotalarias — Windler 163 mv, WIN (hi i | y Wt N N =— a Z Figure 2. Crotalaria nitens var. nitens. A. Habit and leaf variation B. Stem Crotalaria nitens var. gracilis. C. Habit D. Stem 164 Rhodora [Vol. 76 2a. Crotalaria nitens HBK var. nitens Figure 2, Map 1. REPRESENTATIVE SPECIMENS: BRITISH HONDURAS. EL CAYO: near Hydram, Rio Frio Augustine, Mt. Pine Ridge, Hunt 214 (us). GUATEMALA. VERAPAZ: Coban, 1,350 m., Türkheim 1948 (MICH, NY, US). HONDURAS. Comayagua, near Siguatepeque, 1,080-1,400 m., Standley 56370 (Us). MEXICO. CHIAPAS: Muncipio of Tenejapa, 6,000 ft, Breedlove 10976 (US); OAXACA: Entre Puerto Eligio a Comaltepec, Calderon 394 (MICH, US); VERACRUZ: Zacuapan, Purpus 8007 (NY, US). 2b. Crotalaria nitens HBK var. gracilis Windler, Phytologia 21: 263, 1971. Figure 2, Map 1. Type: Mexico: Jalisco; Sierra de Halo, logging road 7 miles south southwest of Tecalitlan and extending south- east toward San Isidaro, 13-16 miles from highway. Mc- Vaugh & Koelz 1188 (MICH). 3. Crotalaria brevipedunculata Windler, Phytologia 21: 257. 1971. Shrub or suffrutescent herb with root type unknown. Stems more than 3 dm. long, 3.5 mm. thick, the longest internode 1.2 em. long; vestiture of dense, short spreading trichomes 0.6-0.7 mm. long. Stipules absent. Leaves elliptic or elliptic-oblong, 2.1-4.4 em. long, 5-13.5 mm. wide, base euneate to widely acute, the apex acuminate; vestiture of loosely appressed trichomes 0.5-0.9 mm. long; petioles 1.6- 2.1 mm. long. Inflorescenses terminal and also leaf-opposed, peduncles 1.2-6.1 cm. long. Bracts sessile, linear or nar- rowly lanceolate, 4-4.4 mm. long, 0.3-0.6 mm, wide; pedicels 3.8-4.2 mm. long. Calyces 10.5-11 mm. long, the tubes 2.5 mm. long, bearing spreading trichomes 0.5-0.7 mm. long; bracteoles linear, 3.5 mm. long, 0.2-0.3 mm. wide. Corollas yellow, the standards 10.5-11 mm. long, equaling the upper calyx lobes; elongate anthers 1.6-2 mm. long, short anthers 0.3-0.4 mm. long; style 6.4 mm. long. Fruits and seeds not known. Chromosome number: not known. Flowering date: December 20. Habitat: shady canyon slope with oaks and palms, elevation 3,500 feet. Range: Mexico: Durango, Sinaloa. Figure 3, Map 2. = Type: Sierra Tres Picos, Durango, infrequent, scattered. Gentry 5311 (GH). 1974] Crotalarias — Windler 165 Figure 3. Crotalaria brevipedunculata. A. Habit B. Stem 166 Rhodora [Vol. 76 X C.brevipedunculata | € C.mexicana M LL—— — 6 C. polyphylla e C.quercetorum ——. | | Map 2. Distribution of Crotalaria brevipedunculata, C. mexicana, C. polyphylla, and C. quercetorum. Crotalaria brevipedunculata is characterized by its lack of stipules, its short terminal inflorescences and small flowers. It most nearly resembles C. nitens, but differs from it in its smaller flower size and shorter peduncles. In addition to the holotype only one other collection of this species has been observed: Mexico. SINALOA: Puerto a Tamiapa, Gentry 5815 (MICH, NY). 4. Crotalaria polyphylla Riley, Kew Bulletin 43: 333. 1923. Perennial decumbent or ascending herb with a taproot to 1.4 em. thick. Stems 4-10, 1.2-3 dm. long, 1-2.5 mm. thick, the longest internode 1-3.2 cm, long; vestiture of short appressed or long spreading trichomes, the appressed 0.3-0.6 mm. long, the spreading 0.8-1.2 mm. long. Stipules 1974] Crotalarias — Windler 167 Figure 4. Crotalaria polyphylla. A. Habit and leaf variation B. Stem 168 Rhodora [Vol. 76 absent. Leaves elliptic, oval, or elliptic-oblong, 1.6-4.2 em. long, 5-183 mm. wide, base rounded to cuneate, the apex obtuse, rounded, or acuminate; vestiture of both surfaces of loosely appressed trichomes 0.5-1.1 mm. long; petioles 1-2.5 mm. long. Inflorescences terminal or terminal and also leaf-opposed, peduncles 4-16 em. long. Bracts sessile, nar- rowly elliptic to narrowly lanceolate, 2.5-9.5 mm. long, 0.3-1.4 mm. wide; pedicels 4-6 mm. long, Calyces 9-13.5 mm. long, the tubes 2-3 mm. long, bearing appressed tri- chomes 0.3-1 mm. long; bracteoles linear, narrowly lanceo- late or narrowly elliptic, 3-7 mm. long, 0.4-1 mm. wide. Corollas yellow, the standards 10-14 mm. long, 0.5-1.5 mm. longer than the upper calyx lobes; elongate anthers 1.7-2.3 mm. long, short anthers 0.5-0.7 mm. long; styles 6-7.3 mm. long. Fruits 1.4-2.4 em. long, 0.5-0.9 em. broad; seeds green, grey-green, green-brown, brown, or tan, 7-18 per pod, 1.8-2.5 mm. long. Chromosome number n—16. Flowering time: June-October. Habitat: slopes in open pine woods, elevation 6,500-9,500 feet, Range: Mexico: Chihuahua, Durango, Jalisco, Sinaloa. Figure 4, Map 2. Type: Mexico: Sierra Madre, Seeman 2187 (K!). Material here treated as Crotalaria polyphylla was in- cluded by Senn (1939) in C, maritima var. maritima; however, its terminal inflorescences and complete lack of stipules clearly separate it from C. rotundifolia (for treat- ment of Senn's C. maritima, see C. rotundifolia var. rotundi- folia). The epithet polyphylla was used by Senn as the varietal name for Mexican plants he considered to be C. purshii (C. purshii var. polyphylla). Personal examination of the type has led to the recognition of C. polyphylla at the specific level. (For treatment of plants previously placed in C. purshii var. polyphylla by Senn see C. querce- torum.) Crotalaria polyphylla is characterized by its decumbent or ascending habit, lack of stipules, and terminal inflorescences. REPRESENTATIVE SPECIMENS: MEXICO. CHIHUAHUA: Memelichi, Rio Mayo, pine slopes, Standley 2701 (GH, MEXU, US). DURANGO: 1974] Crotalarias — Windler 169 32.7 mi. W of Durango in allano slope below pine forest, 8,500 ft., Johnston 2694 (MEXU, MICH). JALISCO: Sierra de la Campana, along road to Mascota, 1,900-2,000 m., McVaugh 13777 (MEXU, MICH, US). SINALOA: Ocurahui, Sierra Surotato, Gentry 6227 (MICH, NY). 5. Crotalaria stipularia Desvaux, J. Bot. (Desvaux) 3: 76. 1814. Crotalaria espadilla HBK. Nov. Gen. et Sp. Plantarum 6: 399. 1824. Type: Humboldt 1025, (P; observed IDC Microfiche). Crotalaria stipularia var. serpyllifolia DC., Prodromus 2: 124. 1825. Type: (G; observed DC Microfiche in Pro- dromus Herbarium). Crotalaria stipularis, orthographic error by Grisebach, Flora Brit. West Ind. Isl. 178. 1864. Crotalaria stipularis var. oblongata Grisebach, Flora Brit. West Ind. Isl. 178. 1864. Crotalaria stipularis var. sericea Grisebach, Flora Brit. West Ind. Isl. 178. 1864. Crotalaria stipularis var. espadilla Kuntze, Revisio Generum Plantarum 1: 175, 1891. Crotalaria stipularia forma eliptica Chodat & Hassler, 4: 835. 1904. Crotalaria stipularia var. grandifolia Senn, Rhodora 41: 333. 1939. Type: Hahn 239 (GH). Annual, erect herb with a taproot to 0.6 cm. thick. Stem solitary, 1-6.3 dm. long, 1.2-5 mm. thick, the longest inter- node 3-8 em. long; vestiture of loose appressed and/or spreading trichomes, 0.5-3 mm. long. Stipules present, decurrent the length of the internode, 1.1-3.2 cm. wide at the top, tapering to the subtending node, the stipule lobes ineurved 1.2-1.6 em. long. Leaves oval, oblong, elliptie, or lanceolate, 2.2-8.2 cm. long, 7.5-25 (-37) mm. wide, base rounded to cuneate, the apex retuse, rounded, acuminate, or acute; vestiture of both surfaces of appressed or loosely appressed trichomes 0.5-2.1 mm. long; petioles 0.5-1.5 mm. long. Inflorescences leaf-opposed, peduncles 3.1-10 cm. long. Bracts petiolate, elliptic, lanceolate to linear, the petioles 170 Rhodora [Vol. 76 Figure 5. Crotalaria stipularia. A. Habit and leaf variation B. Stem 1974] Crotalarias — Windler 171 1-1.6 mm. long, the blades 3-5 mm, long, 0.3-1.2 mm. wide; pedicels 2-4 mm. long. Calyces 8-10 mm, long, the tubes 2.2.3 mm. long, bearing appressed or loosely appressed trichomes 0.7-1.2 mm. long; bracteoles elliptic, lanceolate, or linear, 3-4.5 mm. long, 0.4-0.7 mm. wide, Corollas yel- low, the standards 6-10 mm. long, equaling to 1.5 mm. shorter than the upper calyx lobes; elongate anthers 1.1-1.3 mm. long, short anthers 0.3-0.5 mm. long; styles 4.5-5.2 mm. long. Fruits 2.5-3.3 cm. long, width not known; seeds tan, bronze, red-brown, or dark brown, 17-41 per pod, 2-3 mm. long. Chromosome number: n—16. Flowering time: February-December. Habitat: sandy soils, savannahs, clearings, fields and roadsides, elevations to 2,300 feet. Range: West Indies; Dominican Republic, Dominica, Guadeloupe, Haiti, Martinique, Puerto Rico, Trinidad. (Also widely distributed in South America.) Figure 5, Map 1. Type: Not located. Crotalaria stipularia, as treated by Senn, was divided into three varieties. Varieties "typica" and “orandifolia” are not different except in overall plant size. Both varia- tions are found in a single location mass collection from Puerto Rico sent to me by Dr. R. A. Howard of the Arnold Arboretum. This size variation is probably typical of plants of this species at most locations. Senn’s application of De Candolle’s C. stipularia var. serpyllifolia is to a plant in which all of the original leaves were lost during drought (or some other catastrophe) and new branches with small leaves and shorter internodes had developed before the plant was collected. The wide decurrent stipules were not lost when the leaves were lost, which accounts for the variety having large stipules. The IDC Microfiche photo- graph of De Candolle’s specimen in the Prodromus Her- barium shows a plant with smaller leaves, but not quite so small as those in Senn’s cited specimens. No field col- lections were made of C. stipularia during this study, but seeds were obtained for garden studies (Irwin 1050 and Stimson 4264). 172 Rhodora [Vol. 76 Crotalaria stipularia is characterized by its erect habit, leaf-opposed inflorescences, small flowers, and large stipules with incurved lobes. It most nearly resembles C. sagittalis, but the two taxa may be separated by stipule characters. REPRESENTATIVE SPECIMENS: DOMINICA. Grand Savannah, Lloyd 855 (NY). DOMINICAN REPUBLIC. Prov. de la Vega: near Gara- bawa at Rio Yaquir, Fuertes 1669 (A, NY, P). GUADELOUPE. Montebello, alt. 50 m., Questel 432 (P, us). HAITI. Dept. du Nord: east of Marmelade, grassy summit of mountain, Leonard 7222 (us). MARTINIQUE. Fort Vaillant, Hahn 239 (GH; Photo., Us). PUERTO RICO. Along railroad N of Mayaguez, Heller 4574 (MICH, NY, US). ST. VINCENT. near Barrovollic (Leeward), Smith & Smith 1053 (NY). TRINIDAD. south of Dabadie, Bulton & Hazen 702 (GH). 6. Crotalaria sagittalis L., Sp. Pl. p, 714. 1753. Crotalaria fruticosa Miller, Gard. Dict. (pages not num- bered) 1768. Type: Jamaica, Houstoun, (BM, photo., NCU). Anonymos sagittalis (L.) Walt., Flora Carol. 181. 1788. Type: same as for C. sagittalis L. Crotalaria parviflora Roth, Catalecta Bot. 1: 83. 1797. Type: not located. Crotalaria sagittalis var. B oblonga Michx. Flora Boreali- Am. 2: 55. 1803. Type locality: "Virginia et Carolina". Crotalaria platycarpa Link, Enum. Pl. Hort. Reg. Bot. Berol. Alt. part 2, 227. 1822. Type: destroyed in Berlin. Crotalaria pringlei A. Gray, Proc. Am. Acad. 17: 200. 1881-2. Type: Arizona, Pringle, summer 1881 (GH). Crotalaria sagittalis var. fruticosa (Miller) Fawcett and Rendle, Fl. Jamaica 4: 10. 1920. Crotalaria sagittalis var. blumeriana Senn, Rhodora 41: 339. 1939. Type: Blumer 1772 (GH). Crotalaria tuerckheimii Senn, Rhodora 41: 339. 1939. Type: von Tuerckheim II 1282 (GH). Crotalaria belizensis Lundell, Amer. Midl, Naturalist 29: 473. 1943. Type: Gentle 4144 at (NY). Annual or rarely perennial, erect herb with a taproot to 0.6 cm. thick. Stems 1 (-7), 0.25-4.1 dm. long, 1-2.5 mm. thick, the longest internode 0.4-4.2 em. long; vestiture of long spreading trichomes, 1.5-3 mm. long. Stipules present, 1974] Crotalarias — Windler his Figure 6. Crotalaria sagittalis. A. Habit and leaf variation B. Stem Rhodora [ Vol. Map 3. FER Oa DON A Tet "ese e! NA ir C. sagittalis - YT) = on - > J 252525» u o 1 ka d o i 3 “ee *] (e ^ — } 297... W " —7» Distribution of Crotalaria sagittalis. 1974] Crotalarias — Windler 175 decurrent from 1% to the full length of the internode, 0.35- 1.7 cm. wide at the top, tapering to the subtending node, the stipule lobes spreading 0.4-1.8 cm. long. Leaves elliptic, oblong, lanceolate, or linear-lanceolate, 1.1-10 cm. long, 2.5-15 (22.5) mm. wide, base rounded to cuneate, the apex obtuse, mucronate, acuminate, or acute; vestiture of both surfaces of spreading or loosely appressed trichomes 1-2.9 mm. long; petioles 1.2-2.5 mm. long. Inflorescences leaf- opposed, peduncles 2-8 (-12) cm. long. Bracts petiolate, lanceolate to elliptie lanceolate, the petioles 0.5-1.5 mm. long, the blades 4-8.1 mm. long, 0.5-1.3 mm. wide; pedicels 3.5-5.5 mm. long. Calyces 7-15 mm. long, the tubes 2.3-3.9 mm. long, bearing loosely appressed trichomes 0.8-2 mm. long; bracteoles elliptic lanceolate, lanceolate, or linear, 2.5-7 mm. long, 0.7-1.2 mm. wide. Corollas yellow, the standards 6-12.5 mm. long, equaling to 4 mm. shorter than the upper calyx lobes; elongate anthers 0.9-2 mm. long, short anthers 0.3-0.5 mm. long; styles 4-5.3 mm, long. Fruits 1-2.7 (3.6) cm. long, 0.5-1.1 cm. broad; seeds tan, tan-green, olive, green-brown, or coffee-brown, 7-32 per pod, 1.4-3 mm. long. Chromosome number: n=16. Flowering time: May-October. Habitat: road and railroad rights-of- way, disturbed ground, sand deposits, fields, prairies, lake margins, and dry oak and pine forests, elevation sea level — 8,500 feet. Range: Eastern and Central United States through Mexico and Central America to Panama and on the Greater Antilles. Figure 6, Map 3. Lectotype: “2 Kalm sagittalis" (LINN) (see Taxon 21: 545.) Type locality: “Brasilia, Virginia" as cited in Species Plantarum. Plants included here under Crotalaria sagittalis include material cited by Senn as variety blumeriana and variety fruticosa, as well as Senn’s variety "typica". Early leaves of plants of C. sagittalis tend to be broader than the later leaves, and all branches tend to have narrower leaves than the main stem. In addition, the further south a plant occurs, the longer the plant grows, and the more early leaves are 176 Rhodora [Vol. 76 lost before the end of the growing season. There is also a tendency for southern plants to have narrower early leaves than northern plants. Garden studies have shown Crotalaria sagittalis var. blumeriana to be a dry habitat ecophene of the typical species. Although a great deal of variation exists in C. sagittalis, especially in the Mexican representatives, little in the way of a clear pattern that would allow a useful segregation of the plants (and popu- lations) into varieties has been detected. Further study of this wide-spread but poorly understood species, to gain a more complete picture of the relationships between plant variation, geographic, and ecological factors, may prove helpful, especially in Mexico, Central and South America. REPRESENTATIVE SPECIMENS: BRITISH HONDURAS. Honey Camp, Lundell 665 (vus). COSTA RICA. Paturage a San Jose de Costa Rica, Tonduz 443 (MicH, P). CUBA. Pinar del Rio: Herradura, Britton, Britton, Earle, & Gager 6404 (NY). GUATEMALA. Alta Verpaz: Coban, alt. 4,300 ft., Tuerckheim 239 (GH, Us). HAITI. Massif de la Selle Nouvelle Touraine, alt. 1,400 m., Ekman 1501 (Ny, US). HONDURAS, Morazan, hills around Zamorano Valley, alt. 800-900 m., Swallen 11252, (us). JAMAICA. Castleton, 490 ft., Harris 11855 (GH, NY, P, US). NICARAGUA. Comarca ElCabo: LaTronquera, alt. 60 m., Molina 14881 (Ny). PANAMA. Panama: Sabana de Dormisolo, near Chepo, Pittier 4686 (Us). EL SALVA- DOR. Ahuachapan: Ahuachapan, alt. 800-1,000 m., Standley 19747 (GH, NY, US). MEXICO. Baja California: S side of La Carrerita, Sierra El Taste, 1,500-1,600 m., Carter & Chisaka 8504 (MEXU). Chiapas: Tenejapa, 7,800 ft., Breedlove 19718 (us). Chihuahua: Madera, Pennell 19228 (vs). Colima: Colima, Palmer 130 (vs). Durango: without locality, Garcia 942 (vs). Guanajuato: NE of Guanajuato, Ripley & Barneby 13351 (NY). Jalisco: near Chapala, Rose & Painter 7636 (NY). Mexico: Puerto del Salitre, 1,300 m., Hinton 1780 (GH, MEXU, NY). Michoacan: vicinity of Coalcoman, 1,000 m., Hinton 12862 (Ny, US). Morelos: Cuernavaca, Rose & Painter 6850 (GH, Ny, US), Nayarit: Cerro de la Cruz, E of Tepic, Mexia 669 (GH). Oaxaca: near city of Oaxaca, Rose & Hough 4623 (NY, US). Puebla: 72 km. SE of Mexico City, Weaver 995 (US). San Luis Potosi: 7-8 mi. NE of Xilitla, King 4360 (us). Sinaloa: near Mazatlan, Windler & Windler 2885 (NcU). Sonora: NE cf El Tigre, 6,000 ft, White 4370 (GH, MICH). Tamaulipas: vicinity of Gomez Farias, ca. 350 m., Palmer 341 (us). Veracruz: SE of Cate- maco, meadow, Dressler & Jones 114 (MEXU, MICH, NY, US), UNITED 1974] Crotalarias — Windler 177 STATES. Alabama: MOBILE C0.: Dauphin Island, Jones 585 (ALA). Arkansas: SALINE CO.: ridges, Benton, Demaree 37333 (stu). Con- necticut: MIDDLESEX CO.: Middlefield, Kofoid (GH). Georgia: JACK- SON CO.: 11 mi. N of Athens, Cronquist 4571 (GH). Illinois: JACKSON co.: roadside S of Elkville, Bailey & Swayne 462 (stu, NCU). Indiana: LAKE CO.: East Gary, Hull 18 July 1936 (GH). Iowa: JONES CO.: railroad right-of way, Sect. 7, Oxford Twp., Isely 5913 (GA). Kansas: RILEY Co.: ravines, Norton 80 (GH, P). Kentucky: GRAVES CO.: rail- road embankment, Fancy Farm, O'Dell & Windler 936 (stv). Louis- iana: CALCASIEU PARISH: 114 miles NNW of Starks, Sect. 26, Las- seigne 1579 (LAF). Maryland: Montgomery?, Wildwood, Dowell 1626 (GH). Massachusetts: MIDDLESEX CO.: shore of Winter Pond, Bartlett 224 (GA, GH, TEX, WIS). Michigan: KALAMAZOO CO.: 1 mi. E of Schoolcraft, Hanes 1796 (GH). Minnesota: CHISAGO CO.: Lindstrom, Anderson 6458 (GH, WIS). Mississippi: LAYAYETTE CO.: near Tula Cemetery, 18 mi. SE of Oxford, Pullen 64294 (GA, MISS, NCU). Missouri: BARTON CO.: prairies, Milford, elev. 750 ft., Demaree 39377 (GA, NCU, USF). New Jersey: BERGEN CO.: Hackensack, route 4, sandy field, Rush & Svenson 6769 (GH, wis). New York: SARATOGA co.: N of Crescent, Burnham 8 Sept. 1906 (GH). North Carolina: CALDWELL CO.: 1 mi. SE of Hudson, Randolph & Randolph 1096 (GH, NCU). Oklahoma: MUSKOGEE CO.: 2 mi. S of Braggs, Edmond 164 (TEx). Pennsylvania: DELAWARE CO.: Wayne, dry gravel bank, Bartram 1162 (GH). Rhode Island: Cumberland (?) without collec- tion name & number, Aug. 1840 (GH). South Carolina: GREENVILLE CO.: 215 mi. SW of Berry's Pond, Freeman 56577A (NCU). Tennessee: KNOX CO.: sandy soil near Knoxville, Ruth 661 (wis). Texas: JASPER co.: 3 mi. N of Evadale, Turner & Tharp 3079 (TEX). Vermont: WINDHAM CO.: Vernon, Blanchard 1901 (GH). Virginia: CULPEPER co.: 0.3 mi. N of Lakota on Co. Rt. 621, Ahles & James 61692 (NCU). West Virginia: HARDY co.: Lost River Bridge, wasteground, Core 18 Aug. 1931 (GH). 7. Crotalaria purshii DC., Prodromus 2: 124. 1825. Crotalaria sagittalis var. B L. Sp. Pl. 714. 1753. Crotalaria sagittalis var. a linearis Michx., Fl. Boreali- Am. 2: 55. 1808. Type locality: “Virginia et Carolina". Crotalaria laevigata Pursh, Fl. Am. Sept. 469, non Lam., Encyc., 2: 198. 1786. Type: Pursh specimen not located ; Syntype was plate in Plukenet, Phytographia-Amagestum Bot. t. 277, fig. 2. 1694. Specimen from which this figure was drawn is in Sloane collection at BM. Perennial, erect herb with a taproot to 0.7 cm. thick. 178 Rhodora [Vol. 76 Figure 7. Crotalaria purshii. A. Habit and leaf variation B. Stem Crotalarias — Windler C. purshii X Map 4. Distribution of Crotalaria purshii. HAA A as ae Pe HO UI i Aro - C : VETE CHI Lad Re 179 180 Rhodora [Vol. 76 Stems 1-4 (8), 1.5-5 dm. long, 1-1.5 mm. thick, the longest internode 3.3-8.5 cm. long; vestiture of short appressed or loosely appressed trichomes 0.4-0.8 mm. long. Stipules present, decurrent one-half to the full length of the inter- node, 0.3-0.9 cm. wide at the top tapering to the base, the stipule lobes spreading 0.5-1.7 cm. long. Leaves linear, oblong, linear-lanceolate or elliptic, 4.9-10 cm. long, 1.7-12 mm. wide, base rounded to cuneate, the apex obtuse or acute; the lower surface bearing appressed trichomes 0.3- 0.7 mm. long, the upper surface glabrous; petioles 0.5-2 mm. long. Inflorescences leaf-opposed, peduncles 7.5-23 cm. long. Bracts petiolate, elliptic-lanceolate to lanceolate, the petioles 0.5-1.6 mm. long, the blades 4.5-7 (8.5) mm. long, 0.6-1.8 (2) mm. wide; pedicels 4.5-6 mm. long. Calyces 9-12.65 mm. long, the tubes 2.5-3.5 mm. long, bearing ap- pressed trichomes 0.2-0.5 mm. long; bracteoles lanceolate or ellipite-lanceolate, 3.5-5 mm. long, 0.8-1.2 mm. wide. Corollas yellow, the standards 8-13.5 mm. long, 2 mm. shorter to 0.5 mm. longer than the upper calyx lobes; elongate anthers 1.2-1.6 mm. long, short anthers 0.3-0.6 mm. long; styles 2.7-4.6 mm. long. Fruits 1.4-2.8 cm. long, 0.6-1 cm. broad; seeds tan, olive, green-brown, or dark brown, 11-37 per pod, 1.8-2.9 mm. long. Chromosome num- ber: n=16. Flowering time: March-October. Habitat: savannahs, sandhills, and open pine woods, coastal plain. Range: United States: Alabama, Florida, Georgia, Louis- iana, Mississippi, North Carolina, South Carolina, Virginia. Figure 7, Map 4. Type: Based on Crotalaria laevigata Pursh. Type locality: Pine woods of Virginia and Carolina. Crotalaria purshii, named by De Candolle, was based on material from Virginia and Carolina. He cites C. laevigata of Pursh as being the same, but proposes the name C. purshii since C. laevigata had been previously used by Lamarck. Crotalaria sagittalis var. B L. is based on Plukenet's figure (Plukenet, 1694, t. 277, Fig. 2) which is purshii. Senn (1939) recognized a variety, polyphylla, under 1974] Crotalarias — Windler 181 Crotalaria purshii. Most specimens of this variety are here treated as C. quercetorum. For further information con- cerning the epithet polyphylla see the discussion under C. polyphylla. Crotalaria purshii is characterized by its erect habit; sparse, short, appressed pubescence; spreading stipule lobes; frequently narrow leaves, the upper surfaces of which are glabrous; and long peduncles. Crotalaria purshii is similar to C. sagittalis in habit and general appearance in parts of its range, but it may be distinguished from C. sagittalis by its glabrous upper leaf surfaces, appressed pubescence, and long peduncles. The extremely variable C. rotundifolia var. rotundifolia may occasionally bear a resemblance to C. purshii, but these plants lack glabrous upper leaf surfaces. REPRESENTATIVE SPECIMENS: UNITED STATES. Alabama: BALD- WIN co.: sandy field, Jones 496 (aua). Florida: CALHOUN CO.: Cypress Park, Dead Lakes, Godfrey & Clewell 62916 (LAF). Georgia: WAYNE CO.: open pinelands, 5 mi. S. of Jessup, Duncan 6941 (GA). Louisiana: ST. TAMMANY PARISH: 7% mi. S of town of Pearl River, Sect. 13, Lasseigne 1480 (LAF). Mississippi: PEARL RIVER CO.: 1 mile N of Picayune along Hwy. 12, low pine lands, Bollwinkel & Wun- derlin 9 (stu). North Carolina: CRAVEN CO.: pine savannah, 0.6 mi. SW of Havelock, Radford 34504 (NcU). Virginia: ISLE OF WIGHT CO.: dry sandy pine and oak woods ca. 1 mile SE of Zuni, Fernald & Long 6233 (GH). 8. Crotalaria quercetorum Brandegee, U. Cal. Pub. Bot. 10: 407. 1924. Crotalaria purshii var. polyphylla (Riley) Senn, Rhodora 41: 346. 1939. (see note below) Annual, erect, or ascending herb with a slender taproot to 0.25 em. thick. Stem solitary, 1.1-5.3 dm. long, 1-3.1 mm. thick, the longest internode 1.3-5.2 cm. long; vestiture of short appressed trichomes, 0.3-0.4 mm. long. Stiples usually absent or very narrow, decurrent 1/5 the length of the internode, 0.3-0.4 cm. wide at the top narrowing to the base, the stipule lobes spreading 0.1-0.25 cm. long. Leaves elliptic, narrowly oblong, narrowly elliptic, narrowly lanceo- 182 Rhodora [Vol. 76 Figure 8. Crotalaria quercetorum. A. Habit and leaf variation B. Stem 1974] Crotalarias — Windler 183 late, or linear, 3-6.8 cm. long, 2-5.5 (-8) mm. wide, base rounded to cuneate, the apex acute or acuminate; vestiture of appressed trichomes 0.2-0.5 mm. long, the upper surface usually glabrous; petioles 1.5-3 mm. long. Inflorescences leaf-opposed, peduncles 2.5-8 cm. long. Bracts sessile, nar- rowly elliptic, or linear, 2.5-3.5 mm. long, 0.2-0.3 mm. wide; pedicels 2.5-4.2 mm. long. Calyces 8.2-11 mm. long, the tubes 1.5-2 mm. long, bearing appressed trichomes 0.2-0.8 mm. long; bracteoles linear or narrowly elliptic, 1.5-3.5 mm. long, 0.2-0.3 mm. wide. Corollas yellow, the standards 7.5-10.5 mm. long, equaling to 2 mm. shorter than the upper calyx lobes; elongate anthers 0.8-1.4 mm. long, short anthers 0.2-0.3 mm. long; styles 3.5-3.6 mm. long. Fruits 1-2.4 cm. long, 0.4-0.7 cm. broad; seeds olive, green- brown, or brown, 24-30 per pod, 1.4-2 mm. long. Chromo- some number: n—16. Flowering time: August-March. Habitat: ridges and slopes in oak, oak-pine, and pine woods, elevation 2,600-7,000 feet. Range: Mexico: Chiapas, Jalisco, Michoacan, Nayarit, Oaxaca, Sinaloa. Figure 8. Map 2. Type: Mexico: Chiapas: Hacienda Monserrate, Purpus 9144 (uc!). Crotalaria quercetorum was included under C. purshii by Senn. Unfortunately, Senn chose to use the epithet polyphylla for what he recognized as a variety of C. purshit. Examination of the type of C. polyphylla Riley leads to the conclusion that the name is erroneously applied to the ma- terial here treated as C. quercetorum. The holotype of C. quercetorum is a mixed sheet with one specimen present which may be interpreted as intermediate between C. quercetorum and C. sagittalis. The isotype (US) examined is also composed of several elements, but all are referable to C. quercetorum. Crotalaria quercetorum is characterized by its short appressed pubescence, lack or near lack or stipules, usually narrow foliage, short peduncles and small flowers. Crota- laria quercetorum is similar to C. purshii but may be dis- tinguished by its much shorter peduncles and its usually 184 Rhodora [Vol. 76 smaller size. Because a number of specimens with inter- mediate morphology have been collected, hybridization between C. quercetorum and C. sagittalis is suspected in parts of their range where they are sympatric. Windler & Windler 2994 (NCU) is a collection in which some speci- mens seem to reflect this putative hybridization. REPRESENTATIVE. SPECIMENS: GUATEMALA. Exact locality not given, Bernoulli 282 (G). MEXICO. cnurAPAS: Hacienda Monser- rate, Purpus 9144 (uc, Us); Jalisco: 15-20 mi. SE of Autlan-Sierra Manantlan ca. 1,700 ft., McVaugh 13973 (MICH); Michoacan: 32 mi. W of Jiquilpan along road to Manzanillo, 7,000 ft, Fearing 1195 (TEX); Nayarit: 1 mi. from town of Santa Maria del Oro on way to Laguna Santa Maria, 3,700 ft., Windler & Windler 2917 (NCU); Oaxaca: 5-6 Km NE of Putla, 350 m., McVaugh 22248 (MICH); Sinaloa: Puerto a Tamiapa, 4,500 ft., Gentry 5815 (MEXU). 9. Crotalaria mexicana Windler, Phytologia 21: 259. 1971. Crotalaria sagittalis var. fruticosa (Miller) Fawcett and Rendle, 4: 10, 1920, pro parte, sensu Senn, non sensu typus. Annual erect herb with a slender taproot to 0.3 cm. thick. Stems solitary, 1.2-2.3 dm. long, 1.6-2.4 mm. thick, the longest internode 1-1.5 em. long; vestiture of dense, ap- pressed trichomes, 1.2-2.5 mm. long. Stipules absent. Leaves narrowly elliptic, linear-lanceolate or linear, 2.2-4.6 cm. long, 4-8 mm. wide, base rounded to cuneate, the apex rounded, acute or acuminate; vestiture of both surfaces of dense, loosly appressed trichomes 1.1-2.1 mm. long; petioles 0.5-0.6 mm. long. Inflorescences leaf-opposed, peduncles 0.8-2 cm. long. Bracts sessile, lanceolate, 3.3-3.6 mm. long, 0.7-0.8 mm. wide; pedicels 2.5-4 mm. long. Calyces 10.5-11.5 mm. long, the tubes 2-2.5 mm. long, bearing loosely ap- pressed and spreading trichomes 0.8-2 mm. long ; bracteoles linear-lanceolate, 4-4.5 mm. long, 0.5-0.6 mm. wide. Corollas yellow, the standards 10-10.5 mm. long, equaling to 0.5 mm. shorter than the upper calyx lobes; elongate anthers 1.5-1.7 mm. long, short anthers 0.5-0.6 mm. long; styles 4.8-5.3 mm. long. Fruits 2.1-2.5 em. long, 0.8-1 cm. wide; seeds brown, number per pod not known, 1.8-2 mm. long. Chromosome number: not known. Flowering time: Sep- 1974] Crotalarias — Windler 185 — EREE eee, = ee er te € JL WEEE [rJ Figure 9. Crotalaria mexicana. A. Habit and leaf variation B. Stem C. Leaf pubescence 186 Rhodora [Vol. 76 tember-October. Habitat: dry slopes of mountains, eleva- tion ca. 6,000 ft. Range: Mexico: Jalisco. Figure 9, Map 2. Type: Mexico: Jalisco, mountainside above Etzatlan, Pringle 8855 (holotype, GH; isotypes, TEX, US). Crotalaria mexicana is a new species, the representatives of which were referred by Senn (1939) to C. sagittalis var. fruiticosa (here treated as C. sagittalis var. sagittalis). Crotalaria mexicana is most similar to C. sagittalis and C. quercetorum. It differs from C. sagittalis in its lack of stipules and absence of spreading pubescence, and from C. quercetorum in its short thick peduncles and in its dense pubescence. Crotalaria mexicana is characterized by its erect habit, dense, appressed pubescence, lack of stipules, and extremely short, leaf-opposed peduncles. REPRESENTATIVE SPECIMENS: MEXICO. JALISCO: near Etzatlan, Pringle 8855 (—type), Pringle 11807 (GH, US), Rose & Painter 7571 (US); near Guadalajara, Rose & Painter 7469 (US). 10. Crotalaria rotundifolia (Walt.) Gmelin in Linne . Systema Naturae 2: 1095. 1792. Anonymos rotundifolia Walter, Flora Caroliniana, 181. 1788. Type: BM, photo., NCU! Crotalaria sagittalis var. ovalis Michx. Flora Boreali-Am. 2: 55. 1803. Type locality: “Virginia et Carolina". Crotalaria rotundifolia Poiret, Encycl. Meth. 2: 402. 1812, non Gmelin, 1792. (see Ward, 1962). Crotalaria ovalis (Michx.) Pursh, 469. 1814, nomen illegit. Crotalaria hookeriana A. DC. in A.P. & A. DC., Mem. Soc. Phys. Genéve 9: 97. 1841. Type: not observed. Crotalaria leptoclona Schauer, Linnaea 20: 737. 1847. Type: Mexico, Aschenborn 172, not located. Crotalaria maritima Chapman, Flora South. United States, Suppl. 614. 1883. Type: not located. Crotalaria linaria Small, Flora SE United States 679. 1933. Type: Pine Key, Fla., Blodgett (NY!). 1974] Crotalarias — Windler 187 Crotalaria maritima var. linaria (Small) Senn, Rhodora 41: 347. 1939. Crotalaria rotundifolia var. linaria (Small) Fernald & Schubert, Rhodora 50: 208. 1948. Perennial procumbent, decumbent, or ascending herb with a taproot to 1.2 cm. thick. Stems 2-20, 0.3-6.5 dm. long, 0.5-3.1 mm. thick, the longest internode 0.6-4.8 cm. long; vestiture of appressed, loosely appressed, or spreading trichomes, appressed 0.3-0.6 mm. long, spreading 0.8-3.2 mm. long. Stipules present, with only the stipule lobes present or the stipules decurrent from 1, to the full length of the internode, 0.13-2.3 cm. wide at the top tapering to the base, the stipule lobes spreading 0.15-1.7 cm. long. Leaves round, oval, oblong, ovate, lanceolate, elliptic, or linear, 0.9-5.5 em. long, 1.0-33.1 mm. wide, base rounded to widely cuneate, the apex rounded, obtuse, mucronate, acu- minate, or acute; vestiture of both surfaces of erect, ap- pressed, or loosely appressed trichomes 0.3-2.8 mm. long; petioles 0.5-3 mm. long. Inflorescences leaf-opposed, pe- duncles 1.3-33.9 cm. long. Bracts with or without petioles, elliptic, lanceolate, or linear, the petioles to 1.5 mm. long, the blade 2-7.1 mm. long, 0.3-2 mm. wide; pedicel 2-5 mm. long. Calyces 7-14.9 mm. long; the tubes 1.5-4.5 mm. long, bearing appressed, loosely appressed or spreading trichomes 0.2-1.5 mm. long; bracteoles broadly to narrowly elliptic, lanceolate, or linear, 1.8-6.5 mm. long, 0.3-1.3 mm. wide. Corollas yellow, the standards 6-15 mm. long, 2.5 mm. shorter to 1.9 mm, longer than the upper calyx lobes; elongate anthers 0.9-2.4 mm. long, short anthers 0.3-0.7 mm. long; styles 4-6.9 mm. long. Fruits 1.2-2.8 cm. long, 0.5-1.1 cm. broad; seeds tan, bronze, green-tan, olive, green- brown, or red-brown, 12-46 per pod, 1.5-2.7 mm. long. Chromosome number: n—16. Flowering time: United States, January-October; Mexico, July-March. Habitat: road and railroad rights-of-way, rocky slopes, sandridges, sandhills, open pine, and pine-hardwood forests, elevation sea level — 8,250 feet. Range: Costa Rica; Guatemala; Mexico: Chihuahua, Guanajuato, Hidalgo, Jalisco, Mexico, 188 Rhodora [Vol. 76 Michoacan, Morelos, Nayarit, Oaxaca, Puebla, San Luis Potosi, Sonora, Veracruz; El Salvador; United States: Alabama, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Carolina, Virginia. Type: In Walter Herbarium at BM, photo observed. Crotalaria rotundifolia was first recognized as a distinct species by Walter (1788) who also recognized it as being congeneric with C. sagittalis L. Walter apparently thought that C. rotundifolia and C. sagittalis represented a new genus distinct from Crotalaria, but, instead of describing a new genus, he placed them in a miscellaneous category under the genus name Anonymos. A few years later, Gme- lin placed Walter's Anonymos species under other genera (Sprague, 1939). Anonymos rotundifolia was placed with C. sagittalis L. under Crotalaria. In 1812, Poiret again published the combination, and he, rather than Gmelin, is frequently attributed the authorship of the combination C. rotundifolia. In 1814, Pursh also placed Walter’s Anonymos rotundifolia into Crotalaria. He, however, pro- posed a new species name, C. ovalis, and cited Walter’s name as a synonym, making C. ovalis a superfluous epithet when published and therefore illegitimate. Senn (1939), after examining a photograph of Miller's type of Crotalaria angulata, thought that it was synony- mous with C. rotundifolia, and the name has since been used for the American plants. In 1897, Britten and Baker examined the specimen and concluded it did not differ from C. biflora L. A comparison of the photograph of Miller’s specimen with an IDC microfiche of a C. biflora specimen in the Linnaean Herbarium leads me to agree with Britten and Baker. In addition, Miller indicated that the flowers were produced singly on the sides of the branches and that the leaves were sessile. Neither of these characters agrees with the type from the Miller Herbarium. It may also be possible that Miller made a mistake in indi- cating the location of the place of origin of the seed. It now seems better to recognize the epithet of Walter and to reject the Miller name for our American plants. 1974] Crotalarias — Windler 189 Fernald and Schubert (1948), after studying photo- graphs of Walter's Herbarium (taken by Schubert) at the British Museum of Natural History, suggested that the name Crotalaria rotundifolia actually refers to the plants with short appressed pubescence previously referred to as C. maritima. A more recent photograph of the specimen figured by Fernald and Schubert has been supplied by Dr. Norman K. B. Robson of the British Museum. Dr. Robson (personal communication) indicates that the plant has “appressed-sericeous” pubescence and would “run down easily in Senn's key to C. maritima Chap.". In light of this information, the name C. rotundifolia is recognized as ap- plying to the material previously recognized as C. maritima, as Fernald and Schubert suggested. Crotalaria rotundifolia may be distinguished by its pro- cumbent, decumbent, or occasionally ascending habit; leaf- opposed inflorescences; and the presence of stipules. It is an extremely variable species, especially in the Florida peninsula. Two varieties may be distinguished in the following way: Plants with appressed stem pubescence. .............. RONDE n 10a. C. rotundifolia var. rotundifolia. Plants with spreading stem pubescence, .............. 10b. C. rotundifolia var. vulgaris. 10a. Crotalaria rotundifolia var. rotundifolia Figure 10, Map 5. While plants of some populations in the higher Florida Keys and along the Florida Gulf coast have only narrow leaves, plants at most locations in the state are extremely variable. Crotalaria linaria Small was based on narrow leaved plants and was maintained by Senn as a variety of C. maritima. Since a narrow leaved plant usually cannot be differentiated as being from either the entirely narrow leaved populations or the heterogeneous populations on the peninsula, no purpose is served by maintaining the variety or by reducing it to the status of forma. 190 Rhodora Figure 10. Crotalaria rotundifolia var. rotundifolia. A., B. Habit and leaf variation C. Stem Crotalaria rotundifolia var. vulgaris D. Habit and leaf variation E. Stem 1974] Crotalarias — Windler 191 C. rotundifolia Map 5. Distribution of Crotalaria rotundifolia, var. rotundifolia. [Vol. 76 Rhodora 192 C. rotundifolia var vulgaris Map 6. Distribution of Crotalaria rotundifolia var. vulgaris. 1974] Crotalarias — Windler 193 REPRESENTATIVE SPECIMENS: PUERTO RICO. Vicinity of Dorado, white sand, Britton, Britton, & Brown 6653 (US). UNITED STATES. Florida: LEVY CO.: evergreen scrub oak sandridge, Cedar Key, Godfrey 56571 (GA, GH, USF). MONROE CO.: open dry sandy swale behind low coastal shell ridge, Middle Cape, Cape Sable, Everglades National Park, Ward 3939 (GA, LAF, NCU). Georgia: MCINTOSH CO.: sandhill thicket 1.3 mi. N of Ft. Barrington, Cox Road on ridge, Boseman 458 (NCU). Mississippi: JACKSON C0.: Ocean Springs and vicinity of Gulf Coast Research Laboratory, sandy soil under pines, west end of Deer Island, Channell 414 (usF). South Carolina: GEORGETOWN CO.: roadsides, 5.6 mi. S of Maryville, Radford 28670 (NCU). 10b. Crotalaria rotundifolia var. vulgaris Windler, Phyto- logia 21: 264. 1971. Crotalaria angulata Miller, 1768, sensu Senn, 1939. (See discussion above. Figure 10, Map 6. Type: South Carolina: Hampton County, about three miles northwest of Yemassee on South Carolina Highway 68. Sandhill. 23 July 1967, D. R. & B, K. Windler 2769 (NCU). Crotalaria rotundifolia var. vulgaris is distinguished by its spreading pubescence. Over most of its range var. vulgaris is also characterized by round to oval leaves, but in northern Florida and southern Georgia it intergrades with the usually narrower leaved var. rotundifolia. REPRESENTATIVE SPECIMENS: COSTA RICA. Cartago, Elev. 4,500 ft., Cooper 165 (us). GUATEMALA. Chimaltenango: alt. 2,100 m., Standley 61485 (NY). PANAMA. Panama: roadside, Rio Tapia, Standley 28214 (us). EL SALVADOR. vicinity of San Salvador, alt. 650-850 m., Standley 22444 (NY). MEXICO. Baja California: San Antonio, Cape region, Purpus 475 (US). Guanajuato: 20 mi. NW of Irapuato, Barkley, Rowel, & Parson 752 (TEx). Hidalgo: Dist. Zimapan, on road from Hwy. at km. 242 to Encarnacion, Moore & Wood 4319 (MEX). Jalisco: 5 mi. W of Guadalajara on Hwy. 15, beneath cliff face in road cut, brown sandy soil, Irwin 1257 (TEX). Mexico: Lecheria, Pringle 13411 (GH, US). Michoacan: growing in oak, 2 mi. above Tancitaro, elev. 7,000 ft, Leavenworth 517 (GH). Morelos: km. 56-57, road from D.F. to Cuernavaca, roadside em- bankment, Moore 125 (GH). Nayarit: 13 mi. SE of Tepic, in red clay, Irwin 1293 (TEX). Oaxaca: Valley of Oaxaca, alt. 5,500-7,500 194 Rhodora [Vol. 76 ft., Nelson 1481 (GH). Puebla: vicinity of Puebla, Arsene 1414 (Us). San Luis Potosi: 15 mi. E of Ciudad del Marz on Rt. 80, ca. 1,000 m., Mickel 544 (MICH). Sinaloa: dry hill, vicinity of Mazatlan, Rose, Standley, & Russell 13823 (US). Sonora: 3 mi. NE of Matape, Wiggins & Rollins 400 (GH). Veracruz: Misantla, Purpus 5907 (GH, NY, US). UNITED STATES. Alabama: CLARKE CO.: near Choctaw Lake, elev. 170 ft., Demaree 50694A (LAF, NCU). Florida: HERNANDO CO.: Chinsegut Hill Bird Sanctury, open area along margins of pine woods, Windler 1136 (NCU). Georgia: BEN HILL CO.: open pine-oak on sandy upland, 7.1 mi. N of Fitzgerald, Duncan & Hardin 14301 (GA). Louisiana: WASHINGTON PARISH: mixed woods, edge, along Pushepetappa Creek, 2 mi. WNW of Varnado, Thieret 25910 (LAF). Mississippi: LAMAR C0.: 12 mi. WSW of Hattiesburg along Black Creek, low area, Doughty 162 (MISS). North Carolina: BLADEN CO.: wet ditch on 701 to White Lake, Radford & Radford 2065 (NcU). South Carolina: BERKELEY CO.: common on sandy bank ea. 15 mi. S of Moncks Corner on 52, Wilbur 3547 (GA). Virginia: NANSEMOND CO.: dry white sand of pineland SW of Marsh Hill School S of South Quay, Fernald & Long 11050 (GH). 11. Crotalaria bupleurifolia Schlecht. & Cham., Linnaea 5: 575. 1830. Crotalaria heldiana A. DC. in A. & A.P. DC. Mem. Soc. Phys. Geneve 9: 97. 1841. Type: Grown from seed of un- known source in the garden at Carlsruhe, (G!). Annual or perennial spreading suffrutescent herb with a taproot to 2 cm. thick. Stems few-many, 5-27 dm. long, 2.3.5 mm. thick, the longest internode 3.5-10 cm. long; vestiture of spreading or loosely appressed trichomes 0.2- 1.3 mm. long. Stipules present, decurrent for 14, to the full length of the internode, 0.5-2.7 cm. wide at the top tapering slightly to or past the subtending node, the stipule lobes absent or spreading 0.5-3 cm. long. Leaves oval, ellip- tie, ovate, broadly lanceolate, or lanceolate, 3.2-10.2 cm. long, 7-46 mm. wide, base obtuse to cuneate, the apex obtuse, mucronate, acuminate, or acute; vestiture of both sides of spreading or loosely appressed trichomes 0.2-1.6 mm. long or nearly glabrous; petioles 2-3 mm. long. Inflor- escences leaf-opposed, peduncles 2-16 cm. long. Bracts sessile, linear, subulate, or narrowly lanceolate, 4.5-7.5 mm. long, 0.3-0.6 mm. wide; pedicels 4-5.5 mm. long. Calyces 1974] Crotalarias — Windler 195 13.5-19 mm. long, the tubes 2.5-5 mm. long, bearing loosely appressed or spreading trichomes 0.1-1.1 mm. long; bracteoles linear, subulate, or narrowly lanceolate, 3-6 mm. long, 0.2-0.7 mm. wide. Corollas yellow, the standards 16-25 mm. long, equaling to 6 mm. longer than the upper calyx lobes; elongate anthers 2-2.8 mm. long, short anthers 0.5-0.7 mm. long; styles 7-11 mm, long. Fruits 3-4.1 cm. long, breadth not known; seeds tan, green-brown, or brown, number per pod not known, 2.2-2.5 mm. long. Chromosome number: »—16. Flowering time: August-March. Habitat: North-facing slopes and bluffs and cloud forests with firs and broad leaf trees, elevation 2,500-7,500 feet. Range: Mexico: Chiapas, Guerrero, Jalisco, Mexico, Michoacan, Sinaloa, Veracruz. Type: Hacienda de la Laguna? Schiede et Deppe s.n. (G!). Crotalaria bupleurifolia was described by Schlechtendal and Chamisso in an article describing the collections of Schiede and Deppe in Mexico. A specimen collected by Schiede and Deppe at G is marked “Typus” and may well represent the type even though the location on the label reals *Mexique. Cuerta grande de Chiconquiaco" rather than “Hacienda de la Laguna" cited with the original description of C. bupleurifolia. The number 596 cited with Schiede's name in Senn (1939) refers to the species number which accompanied Schlechtendal et Chamisso’s description. Crotalaria bupleurifolia is characterized by its generally large size, unusual stipules, and large flowers and habit. It most closely resembles C. nayaritensis but may be dis- tinguished from it by the large flower size, larger overall size, and usually wider stipules. Two varieties may be recognized: Stipules present only at the base of peduncles, decurrent for only a single mnternodes ys a es sete ee ee es LAM 11a. C. bupleurifolia var. bupleurifolia. Stipules present at the base of most leaves, frequently decurrent for more than one internode. ............ 11b. C. bupleurifolia var. robusta. 196 Rhodora [Vol. 76 Figure 11. Crotalaria bupleurifolia var. bupleurifolia. A. Habit and leaf variation B. Stem Crotalarias — Windler 197 \ ? ` 2 | ^ | x K éinioleh b diina E: ho i x H ) Lor mi M irs os Pe o 3 . r = z a3 g i ] SA e © z PEERS m t VA e > LOJ g hee v Ec: . 'e o S t. = [d E — 3 3s $ i / 9 9 a a : @ 3 3 uU A A 2m 9. — E / rr c3 i 5 e GO te fi JS TU m——— 2 | e © C.nayaritensis o 10 Map. 7. Distribution of Crotalaria bupleurifolia var. bupleurifolia, C. bupleurifolia var. robusta, and C. nayaritensis. 198 Rhodora [Vol. 76 lla. Crotalaria bupleurifolia var. bupleurifolia Figure 11. Map 7. REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: NE of Bochil, Miranda 5746 (MEXU). Guerrero: San Antonio Buenos Aires, Hinton 11708 (GH, TEX, US). Jalisco: near Santa Monica, 1,950-2,050 m, McVaugh 14086 (MEXU, MICH, US). Mexico: Rincon del Carmen, Dist. Tamascaltepec, 1,340 m, Hinton 1730 (MEXU, NY, US). Michoacan: NW of Aguililla, 15 Km S of Aserradero Dos Aguas, 1,650-1,700 m, McVaugh 22767 (MICH). San Luis Potosi: Las Canoas, Pringle 8 (MEXU). Sinaloa: Batel, Concordia, Dehesa 1619 (US). Tamaulipas: 10 Km NW of El Progresso, 1,450 m, Stanford, Retherford, & North- craft 1009 (GH). Veracruz: Barranca de Tenampa, Zacuapan, Pur- pus 3663 (GH, NY, US). llb. Crotalaria bupleurifolia var. robusta (Senn) Windler, Phytologia 21: 265. 1971. Crotalaria pilosa var. robusta Senn, 1939, 41: 331. Figure 12, Map 7. Type: Temascaltepec, Cumbre de Tejupilco, Hinton 2686 (Us!). REPRESENTATIVE SPECIMENS: MEXICO. Jalisco: 10 Km al N de La Cuesta, sobre el camino a Talpa, 1,100 m, Rzedowski 15134 (MEXU). Mexico: Plaza de Gallos, 1,200 m. Hinton 4595 (GH, NY). Sinaloa: Km 1206 on Mexico Hwy. 40, ca. 30 mi, E of Mazatlan- Guadalajara Junction, 2,700 ft., Windler & Windler 2869 (NCU). 12. Crotalaria nayaritensis Windler, Phytologia 21: 261. 1971. Annual or perennial spreading herb with a taproot to 1.5 em. thick. Stems 1-many, 6-12 dm. long, 1.5-2.5 mm. thick, the longest internode 3.8-10 cm. long; vestiture of short appressed trichomes, 0.2-0.7 mm. long. Stipules present, decurrent the length of the internode, 0.15-1.1 em. wide at the top tapering to or past the subtending node, the stipule lobes parallel with the stem or spreading, 0.1- 1.3 cm. long. Leaves oval, elliptic, ovate, narrowly ovate, oblong or lanceolate, 3.5-7.8 cm. long, 7-26 mm. wide, base obtuse to cuneate, the apex obtuse, mucronate, acuminate or acute; vestiture of both surfaces of appressed or loosely appressed trichomes 0.3-0.8 mm. long; petioles 1.2-2.5 mm. long. Inflorescences leaf-opposed, peduncles 3.2-16 cm. long. 1974] Crotalarias — Windler 199 Figure 12. Crotalaria bupleurifolia var. robusta. A. Habit and leaf variation B. Stem 200 Rhodora [Vol. 76 Braets sessile, linear or elliptic lanceolate, 3-4.5 mm. long, 0.2-0.5 mm. wide; pedicels 2.8-3.8 mm. long. Calyces 7.5- 12 mm. long, the tubes 2-3.2 mm. long, bearing appressed trichomes 0.1-0.5 mm. long; bracteoles linear or narrowly lanceolate, 1.5-3 mm. long, 0.2-0.3 mm. wide, Corollas yel- low, the standards 7-12.5 mm. long, 2 mm. shorter to 1 mm. longer than the upper calyx lobes; elongate anthers 1.3-2.1 mm. long, short anthers 0.4-0.5 mm. long; styles 5-6 mm. long. Fruits 1.3-2.3 em. long, 0.5-0.8 em. broad; seeds tan, grey-tan, green-brown, brown or red-brown, 30-35 per pod, 1.6-2.4 mm. long. Chromosome number: n=16. Flowering time: August-February. Habitat: steep moist slopes and pine woods, elevation 2,500-6,600 feet. Range: Mexico: Jalisco, Nayarit. Figure 13, Map 7. Type: Mexico: Nayarit, North of Compostella (near Km. 24), about 7 miles southwest of Tepic, along road between Tepic and Compostella. Road-cut through moun- tain on moist steep slope, Windler & Windler 2902 (NCU). Crotalaria nayaritensis is a new species named for the Mexican state from which the holotype was collected. It is characterized by its spreading or diffuse habit, leaf-opposed inflorescences, small flower size, and appressed pubescence. Of the Mexican species it most nearly resembles C. bupleu- rifolia, but differs from it in having a smaller flower, appressed pubescence and usually narrower stipules. REPRESENTATIVE SPECIMENS: MEXICO. Jalisco: 13 mi. SW of Autlan, 1,000 m, MceVaugh 19886 (MICH); Llano Verde, municipio de Tecalitlan, 1,600 m, Rzedowski 17417 (MiCH); 3 mi. S of Maza- mitla, 2,100-2,200 m, MeVaugh 12997 (micH, US); San Sebastian, W to Mascota, 1,425 m, Mexia 1408 (vs); Tepic, Palmer 1869 (NY, US). Nayarit: 10 mi. SE of Ahuacatlan, 1,100-1,200 m, Fedema. 287 (MICH) ; N of Compostella, 3,000 ft., Windler & Windler 2902 (NCU); Mina Esperanza Rosa Morada, Ortega 6682 (US); 2 mi. N of Tepic, 3,000 ft., Windler & Windler 2897 (NCU). PUTATIVE HYBRIDS BETWEEN CROTALARIA SPECIES During the course of the study a number of putative hybrids were observed, The following presentation sets 1974] Crotalarias — Windler 201 Figure 18. Crotalaria nayaritensis. A. Habit and leaf variation B. Stem 202 Rhodora [Vol. 76 forth the probable parents and lists of representative speci- mens which appear to be the results of crosses between the parents. Crotalaria sagittalis X C. stipularia ST. KITTS. Pastures near Sandy Point, Britton & Cowell 128 (NY). Crotalaria sagittalis X C. rotundifolia var. vulgaris UNITED STATES. Alabama: Mobile, Mohr s.n. 1873 (SIU). Florida: BAY CO., E of Panama City, Windler & Windler 2723 (NCU); N of Betts, Windler & Windler 2721 (NCU); ESCAMBIA CO., W of Century, Windler & Windler 2688 (NCU); JACKSON CO., Grand Ridge, Windler & Windler 2712 (NCU); OKALOOSA CO. 4 mi. W of Black- water River Bridge on Fla. Hwy. 4, Windler & Windler 2700 (NCU); SANTA ROSA CO., E of Chumuckla, Windler & Windler 2693 (NCU); S of Milton, Windler & Windler 2697 (NcU). Mississippi: FOREST co., S of Hattiesburg, Windler & Windler 2659 (NCU); HARRISON CO., N of Biloxi, Windler & Windler 2666 (NCU); LAMAR CO., 8.5 mi. SE of Hattiesburg, Cooley 62 (Miss); 7.5 mi. SW of Hattiesburg, Cooper 93 (MISS); Pearl River, 1 mi. N of Picayune, Pullen 641208 (MIss). Crotalaria sagittalis X C. quercetorum MEXICO. Chiapas: Bochil, Windler & Windler 2994 (NCU). Crotalaria rotundifolia X C. purshii UNITED STATES. Alabama: BALDWIN CO. Foley, Windler & Windler 2681 (NCU). Florida: CALHOUN CO., 12 mi. S of Clarksville, Windler & Windler 2425 (NCU); ESCAMBIA CO., 235 miles N of Jet. Fla. Hwy. 97 — US Hwy. 29, Windler & Windler 2686 (NCU); SANTA ROSA CO, S of Milton, Windler & Windler 2699 (NCU). Georgia: GLYNN CO., Brythe Island, Hardin & Duncan 13785 (NCU); MCINTOSH CO., 2.6 mi. W of Cox, Bozeman 1078 (NCU); 1 mi. SE of Ft. Bar- rington, Boseman & Radford 2146 (NCU); THOMAS CO, SW of Thomasville, Windler & Windler 2734 (NCU). Mississippi: HARRISON CO.: Ship Island, Demaree 35028 (LAF); JACKSON CO., Round Island, Demaree 33742 (LAF). ACKNOWLEDGMENTS I am especially grateful to my wife, Bonnie Kay, whose moral support and technical and clerical assistance speeded the completion of this study. Dr. C. Ritchie Bell, who served as research adviser during the study, deserves special thanks for his helpful sugges- 1974] Crotalarias — Windler 203 tions, criticisms, and editorial work on the manuscript. Drs. Albert E. Radford and Clifford R. Parks are also thanked for their helpful suggestions and cooperation during the course of the study and the preparation of the thesis. Dr. Velva Rudd, Smithsonian Institution, is thanked for her critical review of the Systematic Treatment manu- script. I would like to acknowledge my fellow students at the University of North Carolina at Chapel Hill and thank them for their aid, counsel, and criticisms during my years of study at the University. During the extensive collecting trips, help was needed on numerous occasions and was always cheerfully supplied by colleagues at other institutions. Special mention should be made of Dr. Arturo Gomez-Pampa, Universidad Nacional de México; Dr. Sidney McDaniels, Florida State University ; Dr. John Thieret, Southwestern Louisiana State University ; and Dr. D. B. Ward, University of Florida, and their respective staffs that provided plant dryers, and green- house and storage space at various times during collection trips. Dr. Andre Clewell and the staff of the Tall Timber Research Station in Tallahassee, Florida, are also due thanks for assistance and lodging that they provided while collec- tions were being made in the vicinity. Special thanks are extended to the curators of the numerous herbaria (A, ALU, AUA, BM, G, GA, GH, K, LAF, MEXU, MICH, MISS, NCU, NY, P, SIU, TEX, UC, US, USF, WIS) from which specimens were borrowed and to Dr. Norman K. B. Robson, in particular, whose efforts in my behalf at the British Museum (Natural History) in checking some of the early American collections proved invaluable. LITERATURE CITED Baker, E. 1914. The African species of Crotalaria. Journal Lin- nean Society, Botany 42: 241-425. BRITTEN, J. & E. BAKER. 1897. Houstoun’s Central American Legu- minosae. Journal of Botany 35: 225-234. FERNALD, M. L. & B. SCHUBERT. 1948. Studies of American types in British Herbaria, IV. Some species of Thomas Walter. Rhodora 50: 190-208. 204 Rhodora [Vol. 76 GRAUSTEIN, J. 1954. The date of Pursh’s “Flora Americae Septen- trionalis." Rhodora 56: 275. LANJOUW, J. & F. STAFLEU. 1964. Index Herbariorum, Regnum Vegetabile, Vol. 31. 251 pp. Utrecht. POLHILL, R. 1968. Miscellaneous notes on African species of Cro- talaria L., II. Kew Bull. 22: 169-348. SAVAGE, S. 1945. A Catalogue of the Linnaean Herbarium. 225 pages. London. SENN, H. 1939. The North American species of Crotalaria. Rhodora 41: 317-366. SPRAGUE, T. 1939. Analysis of binary combinations published under Anonymos by Walter, Flora Caroliniana (1788). Kew Bull. 1939: 331-334. Warp, D. 1962. The genus Anonymos and its nomenclatural sur- vivors. Rhodora 64: 87-92. WINDLER, D. R. 1970. Systematic studies in Crotalaria sagittalis and related species. Unpub. diss, Univ. of North Carolina at Chapel Hill. 258 pages. 1971. New North American unifoliolate Crotalaria taxa. Phytologia 21: 257-266. 1973. Field and garden studies in Crotalaria sagittalis and related species. Phytologia 26: 289-354. TOWSON STATE COLLEGE BALTIMORE, MARYLAND 21204 THE FLORA OF MOUNT MONADNOCK, NEW HAMPSHIRE HENRY I. BALDWIN Mt. Monadnock (Elevation 3,165 feet) lies in Southern New Hampshire at a distance from other hills of compara- ble height. This mass of granite and gneiss rising from a plain has given rise to the generic term “monadnock” for similar isolated rock peaks. The Grand Monadnock, as it is sometimes called, to distinguish it from other monadnocks, is situated in the towns of Dublin and Jaffrey. Roads sur- round the base of the mountain at elevations of 1,200 to 1,500 feet above sea level. Botanically Mt. Monadnock is of interest because of the presence of a few arctic-alpine plants on the bare rocky summit area, far from other stations for them, and because of the records made by Thoreau on his four visits from 1844 to 1860, and later by other botanists, such as Stone (1885) and Deane (1890) and many later ones, some of whom deposited specimens in New England herbaria. It was my purpose to compare the present flora with that found by Thoreau. The student of land use can read much of its history from the flora, since plant habitat is affected by what has hap- pened to the plants. The history of Mt. Monadnock is of interest because of three distinct periods: (1) the develop- ment of complete forest cover since the time of glacial re- treat, (2) its complete removal on the upper slopes by fire, and partial removal on the lower by clearing for pasture and fire, and (3) the slow renewal of vegetative cover dur- ing the last 150 years. It is generally believed on the basis of geological and pollen evidence and observation of succession on rock and soil recently exposed by the retreat of present day glaciers, that the succession on Mt. Monadnock was probably as fol- lows: the first pioneer plants, following lichens and mosses, may have been shrubs and forbs capable of surviving under 205 206 Rhodora [Vol. 76 severe cold, wind and fluctuating moisture. These may have included species that colonize mineral soil and rock crevices, including some of the arctic-alpines. Later, as climate mod- erated, spruces and pines (possibly Pinus Banksiana) oc- cupied the accumulations of sand and organic matter in depressions. Finally, as humus accumulated, Red Spruce (Picea rubens) came to dominate the flora of the upper part of the mountain where soil development was indefinitely delayed. Mt. Monadnock was completely forested at the coming of the White Man, the summit area covered with Red Spruce to the very summit, and the lower slopes with spruce and hardwood, much as the case of lesser mountains much farther north. All similar rocky peaks below the approxi- mately 4,500 feet climatic timberline in New Hampshire are or were completely forest-clad. Mt. Monadnock is about 1,000 feet lower than climatic timberline and lies almost 100 miles south of the nearest timberline. Tree line is higher at lower latitudes. The bare rocky summits of New Hampshire such as Chocorua, Cardigan, Firescrew, and Percy Peaks all have documented history of fire. Until recent years there were many old stumps and logs near the summit. Thoreau wrote in his Journal (June 1858) : “The whole rocky part of the mountain, except the extreme summit, is strewn with the relics of spruce trees a dozen or fifteen feet long, and long since dead and bleached, so that there is plenty of fuel at hand." The remains near- est the summit were no doubt first consumed by visitors campfires. Visitors were very numerous in Thoreau’s day and before, and in several of his writings he reports having seen fires at the top. Since the area at the extreme summit is quite level, with depressions, it is probable that it was originally as densely forested as the rest of the mountain. Chamberlain (1936) shows a picture (p. 73) of stumps near the top. Obviously, there could not have been such severe fires at the summit had there not been timber to burn. The 1953 fire on the east slope stopped when it reached the scrub zone where there was no fuel. 1974] Flora of Mount Monadnock — Baldwin 207 There are many published reports that confirm the orig- inal forested nature of the summit. Later the practice of clearing the slopes for pasture gave rise to numerous fires. During the celebration of the centennial of the Town of Dublin in 1852, Charles Mason made an address on June 17 (Leonard 1855) including the following remark: “Formerly covered with trees almost to the summit.” At the same celebration in Dublin (Leonard op. cit.) a letter was read from Daniel Elliott, M.D. in which he re- called: “I remember (Monadnock) when clothed with ver- dant foliage to the very summit.” Similar records are found in the Town of Jaffrey: “When the town was first settled the mountain was covered to its summit with forest trees, principally spruce, excepting a small peak southeast of the top, which was called the “Bald Rocks” (Cutter 1881). “Belts of heavy spruce timber on the highest por- tions” were reported by Child (1885). Similar references may be found in Dana (1816) and Ellis (1880). Ecologists recognized that forest had come to occupy the mountain much as peaks farther north. Braun (1950) refers to Monadnock (p. 423) as “covered with Red Spruce from 2,300 feet upward, thus producing a small outlier of the prevailing mountain vegetation in the north.” Spurr (1964) states that “the summit of Mt. Monadnock ... was clothed with a Red Spruce forest until destroyed by fire...” Many fires have occurred during the recorded history of Mt. Monadnock, and it is possible that lightning fires may have burned in earlier times, but it is doubtful if Indians caused any fires on its forbidding slopes. In the same ad- dress referred to above (Leonard, loc. cit.) Charles Mason said: “A great fire spread over a great part of the superior portion of the mountain killing every tree and shrub. Dead trees falling, furnished materials for another conflagration. Some 30 years ago in the latter part of a dry summer, a fire in a clearing made its way up to the higher regions." “Some 30 years ago” would have placed this fire about 1820, al- though the cause of the fire differs from other accounts given below. Also the statement of Dr. Elliott on the same 208 Rhodora [Vol. 76 occasion: “I saw, year after year, the devouring flames climbing his lofty sides", suggests many fires. Thoreau, on his second visit to Monadnock in 1852 (his first was in 1844, the journal report of which has not been preserved) related this incident during his passage through Peterborough: *A man in Peterborough told me that his father told him that Monadnock used to be covered with forest; that fires ran through it and killed the turf; then the trees were blown down, and their roots turned up and formed a dense and impenetrable thicket in which wolves abounded . . . till finally they set fire to this thicket and it made the greatest fire they had ever had in the county.” These and other records fail to give exact dates. As Chamberlain (1936) says (p. 97): “No documentary evi- dence has been found to show exactly when the fires oc- curred.” He also quotes (p. 98) from the New Hampshire Sentinel of September 30, 1825, reporting a great fire of “some years ago”. “This fire consumed only the turf. The trees turned over in every direction, and afforded about three years later fuel for a more tremendous scene.” This places the two “great” fires closer together. The best evi- dence suggests a first fire about 1800, then the 1815 hurri- cane, and the final great fire in 1820, There were later ones, too. Thoreau reported (“A Walk to Wachusett”, July 19, 1842) “A fire blazing on Monadnock that night, which lighted up the whole western horizon.” The first inroads of man occurred on the lower slopes that were cleared for pasture. Later, clearing progressed higher where sheep were grazed. Fire was used both for opening the areas and for keeping them so, and many of the fires that reached the upper levels both before and after the 1820 fire were caused by fire escaping from pasture burn- ing. The repeated burning promoted the growth of blue- berries, which I believe may have spread from lower pastures to the summit after the great fires. At present these are rather scarce. Following abandonment of the pastures, these seeded into Red Spruce, and the process of reclothing the upper reaches 1974] Flora of Mount Monadnock — Baldwin 209 of the mountain progressed rapidly at first, and ever slower on the more exposed parts of the mountain. Here denuda- tion resulted in a worsening of the climate. Wind movement increased over the bare rock, and extremes of temperature and soil moisture retarded growth. Recently a cycle of retarded spread of spruce seems to have set in, since almost no spruce trees less than forty years old can now be found on the upper part of the mountain and a high percentage of the trees display dead tops and reduction in diameter and height growth. The large area laid bare by the fires apparently provided a favorable habitat for a few arctic-alpine plants that may have been present before the fires or had been transported from other sources. In any case they were well-established at the time of Thoreau's first visit, little over twenty years after the great fire. Those recorded by Thoreau (Journal, August 8, 1860, 25-28; 1670-1671) were: Arenaria groenlandica, Vaccinium Vitis-Idaea var. minus and Juncus trifidus. Others with northern or high altitude affinities, not found generally in the lowland, especially in the vicinity of Mt. Monadnock were Potentilla tridentata and Smilacina trifolia. During the present survey Lycopodium Selago var. Selago and var. appressum as well as L. annotinum var. pungens have been added to the list. Thoreau limited his observations to the "upper part of the mountain," and essentially all the taxa reported by him in 1858 “within 200 feet of the summit" were also collected in 1968. Less complete were the lists of Stone (1885), Deane (1890) and Emerson (1869), probably due to the restricted area that they examined. While almost all the taxa recorded earlier are present now (at least those easily discoverable), and a substantial number of new ones have been added, there are doubtless more to be found. There have been changes in distribution and abundance of the plants recorded by the early botanists. The encroachment of Red Spruce and more especially shrubs such as Nemo- panthus mucronata, Rhododendron canadense and Kalmia angustifolia appears to have reduced the habitat available 210 Rhodora [Vol. 76 to the arctic-alpines. Vaccinium Vitis-Idaea can persist a long time under these shrubs, but fruiting is diminished. Some thirty years ago the proprietors of the Half Way House used to collect enough mountain cranberries on the mountain to supply hotel guests, while now only small quantities can be picked. This plant has almost disappeared from the southwest side where the hotel supply was largely obtained. The alpine flora seems to be increasingly re- stricted to the northern side of the peak where a cooler climate prevails. Further, observations indicate that there may be a successional change going on in the mats of Vac- cinium Vitis-Idaea, that are invaded more and more by Potentilla tridentata. Another major change from 100 years ago is the vast reduction in Cladonia lichens. Thoreau remarks frequently how dense they were, crunching under foot, and dry so that fire would spread through them. Cladonia persists around the margins of shrubby hollows, but today does not cover extensive areas. Other lichens mentioned by Thoreau on the summit are now mostly con- fined to moist ravines. Originally it was planned to limit collections to the area studied by Thoreau, “the upper part of the mountain". Be- cause of the difficulty of determining just what this em- braced, the area was extended to include land above 2,000 feet above sea level. (The Half Way House site is at 2,100 feet) Present collections have been supplemented by recent herbarium records kindly supplied by Mr. and Mrs. David E. Boufford of Keene State College. Where several collec- tions of the same taxon were found, only the earliest record is given, in order to save space.* All records that stated Mt. Monadnock as the place of collection were included, although often the specific location was not given. Genera and species are listed in the order of Gray's Manual, 8th Edition (Fernald 1950). The following symbols are used: (*) asterisk: reported by Thoreau 1858-1860. (—) dash: collected 1968-1972. *Complete lists have been deposited in several New England her- baria. 1974] Flora of Mount Monadnock — Baldwin 211 (+) dagger: collected 1968-1972, not previously reported (as far as known) Most of the specimens collected in 1968-1972 were de- posited in the University of New Hampshire and Franklin Pierce College herbaria. Abbreviations for all Herbaria consulted in this study are as follows: HNH Dartmouth College, FPC Franklin Pierce College, KSC Keene State Col- lege, NEBC New England Botanical Club, MASS University of Massachusetts, and NHA University of New Hampshire. LIST OF THE VASCULAR FLORA OF MOUNT MONADNOCK -Lycopodium Selago L. var. Selago L. W. Deane, Aug., 1891 (NEBC). +_L. Selago L. var. appressum Desv. H. I. Baldwin, July. 1968 (FPC). * L.lucidulum Michx. H. I. Baldwin, Aug., 1968 (FPC, NHA). +—L. inundatum L. H. I. Baldwin, Oct., 1968 (FPC). * L.annotinum L. H. D. Thoreau, Aug., 1869 (?) (NEBC). +_L. annotinum L. var. pungens (La Pylaie) Desv. H. I. Baldwin, June, 1971 (FPC, NHA). *_L. clavatum L. T. L. Steiger, Oct., 1935 (NHA). L. clavatum L. var. megastachyon Fern. & Bissell, C. Jenks, Aug. 1883 (NEBC). _L. obscurum L. var. obscurum. A. W. Driggs. Aug., 1916 (NEBC). L. obscurum L. var. dendroideum (Michx.) D. C. Eat. B. L. Robinson, Aug., 1869 (NEBC). *_L. complanatum L. A. W. Driggs, Aug., 1916 (NEBC). Botrychium matricariaefolium A. Br. E. L. Rand, May, 1897 (NEBC). i-Osmunda cinnamomea L. H. I. Baldwin, Aug., 1968 (FPC). Woodsia ilvensis (L.) R. Br. B. L. Robinson, Aug., 1898 (NEBC). —Thelypteris Phegopteris (L.) Slosson. E. F. Williams, June, 1894 (NEBC). —Dryopteris spinulosa (O. F. Muell.) Watt. var. americana (Fisch.) Fernald. E. F. Williams, June, 1894 (NEBC). 212 Rhodora [Vol. 76 —D. marginalis (L.) Gray. D. S. Boufford, June, 1971 (KSC). +—Polystichum acrostichoides (Michx.) Schott. H. I. Bald- win, Aug., 1971 (FPC, NHA). *—Dennstaedtia punctilobula (Michx.) Moore. H. I. Baldwin, Oct., 1968 (FPC). -Pteridium aquilinum ( L.) Kuhn var. latiusculum (Desv.) Underwood. H. I. Baldwin, Aug., 1968 (FPC). *—Polypodium virginianum L. W. Deane, Aug., 1889 (HNH). "—Abies balsamea (L.) Mill, J. R. Churchill, May, 1895 (NEBC). *—Tsuga canadensis (L.) Carr. H. I. Baldwin, July, 1968 (FPC). *—Picea rubens Sarg. E. F. Williams, May, 1898 (HNH). T-Larix laricina (DuRoi) K. Koch. H. I. Baldwin, July, 1968 (FPC). (one tree at 2,900 feet on west side, two in Thoreau Bog). *-Pinus Strobus L. A. R. Hodgdon et al., Aug., 1956 (NHA). Poa compressa L. W. Deane, July, 1884 (HNH). P. pratensis L. E. F. Williams, June, 1894 (NEBC). P. palustris L. J. R. Churchill, July, 1890 (MASS, NEBC). *—_Festuca ovina L. E. L. Dickinson and F. C. Seymour, July, 1930 (Mass). T-F. rubra L. H. I. Baldwin, July, 1968 (FPC). *_Deschampsia flexuosa (L.) Trin. H. D. Thoreau, Aug., 1860 (NEBC). *_Danthonia spicata (L.) Beauv. H. D. Thoreau, Aug., 1860 (NEBC). —D. compressa Aust. A. S. Goodale, July, 1927 (MASS). *Calamagrostis canadensis ( Michx.) Nutt. H. D. Thoreau, Aug., 1860 (NEBC). *—Agrostis scabra Willd. H. D. Thoreau, Aug., 1860 (NEBC). -Scirpus atrovirens Willd. C. W. J enks, Aug., 1883 (NEBC). —S. atrocinctus Fern. A. S. Goodale, July, 1927 (Mass). *—Eriophorum spissum Fern., H. D. Thoreau, Aug., 1860 (?) (NEBC). *E. gracile W. D. J. Koch. A. S. Goodale, July, 1927 (MASS). -E. tenellum Nutt. R. M. Harper, July, 1899 (NEBC). “3,150 feet”. 1974] Flora of Mount Monadnock — Baldwin 213 E. viridi-carinatum (Engelm.) Fern. C. F. Batchelder, Sept., 1909 (NHA). —E. virginicum L. C. W. Jenks and C. W. Swan, Aug., 1883 (NEBC, NHA). Rhynchospora capitellata (Michx.) Vahl. B. L. Robinson, July, 1897 (NEBC). +—R. alba (L.) Vahl. H. I. Baldwin, Oct., 1968 (FPC). +—Carex stipata Muhl. H. I. Baldwin, June, 1969 (FPC). *C. trisperma Dew. H. D. Thoreau, Aug., 1860 (NEBC). “NE swamp”. +-C. trisperma Dew. var. Billingsii Knight, H. I. Baldwin, Oct., 1968 (FPC). * C. canescens L. “E. F. Jr." (?), 1838 (MASS). C. canescens L. var. subloliacea Laested. Sydney Harris, June, 1894 (NEBC). C. brunnescens (Pers.) Poir. W. Deane, July, 1889 (MASS). C. brunnescens (Pers.) Poir. var. sphaerostachya (Tuk- erm.) Kükenth. A. W. Driggs, Aug., 1916 (NEBC). C. cumulata (Bailey) Mackenz. C. W. Jenks, Aug., 1885 (NEBC). —C. communis Bailey. L. Russell, 1898 (NHA). C. novae-angliae Schwein. W. Deane, July, 1891 (HNH, MASS). “Mt. road below house". C. deflexa Hornem. J. R. Churchill, May, 1895 (NEBC). C. umbellata Schkuhr. J. R. Churchill, May, 1896 (NEBC). C. gracillima Schwein. R. J. Eaton, July, 1957 (NHA). C.aestivalis M. A. Curtis. J. R. Churchill, July, 1890 (MASS, NEBC). C. leptonervia Fern. E. F. Williams, June, 1894 (NEBC). C. intumescens Rudge. E. F. Williams, June, 1894 (NEBC). +-C. nigra L. H. I. Baldwin. July, 1971 (FPC, NHA). i-Arisaema triphyllum (L.) Schott. D. W. Allen, May, 1968 (FPC). -Juncus bufonius L. A. W. Driggs, Aug., 1916 (NEBC). *_J. trifidus L. C. W. Swan, Aug., 1883 (NEBC). J.Greenei Oakes and Tuckerm. B. L. Robinson, Aug., 1896 (NEBC). 214 Rhodora [Vol. 76 -J. brevicaudatus (Engelm.) Fern. A. W. Driggs, Aug., 1916 (NEBC). J. articulatus L. C. A. and A. W. Cheever, Sept., 1910 ( NEBC). T-Luzula, multiflora (Retz.) Lejeune. D. W. Allen, May, 1968 (FPC, NHA). *—Uvularia sessilifolia L. D. W. Allen, May, 1968 (FPC). *—Clintonia borealis (Ait.) Raf. T. L. Steiger, May, 1937 (NHA). r-Smilacina racemosa (L.) Desf. H. I. Baldwin, Sept., 1970 (FPC). *—S. trifolia (L.) Desf. Sydney Harris, June, 1894 (NEBC). r-Maianthemwm canadense Desf. D. W. Allen, May, 1968 (FPC). —Streptopus roseus Michx. var. perspectus Fassett. W. Deane, July, 1889 (NEBC). *-Polygonatum pubescens (Willd.) Pursh. H. I. Baldwin, May, 1969 (FPC, NHA). -Medeola virginiana L. D. W. Allen, May, 1968 (FPC). -Trillium erectum L. J. R. Churchill, May, 1883 (Mass). *—T. undulatum Willd. T. L. Steiger, May, 1937 (NHA). r-Sisyrinchium montanum Greene. H. I. Baldwin, July, 1968 (NHA). *-Cypripedium acaule Ait. D. W. Allen, May, 1968 (FPC, NHA). Habenaria Hookeri Torr. Henry A. Purdie, June, 1905 (NEBC). Spiranthes lacera Raf. Hodgdon and Ken Welsh, July, 1957 (NHA). “2,500 feet”. Malaxis uniflora Michx. F. Hunt, Aug., 1946 (NHA). *Salix Bebbiana Sarg. A. R. Hodgdon, June, 1933 (NHA). *—S. discolor Muhl. D. W. Allen, May, 1968 (FPC). *—Populus tremuloides Michx. H. I. Baldwin, Aug., 1968 (FPC). *_Betula alleghaniensis Britton. Hodgdon and Welsh, July, 1957 (NHA). -B. populifolia Marsh. Hodgdon et al., Aug., 1956 (NHA). *—B. papyrifera Marsh. Hattie Merrifield, July, 1879 (NEBC). 1974] Flora of Mount Monadnock — Baldwin 215 —B. papyrifera Marsh. var. cordifolia (Regel) Fern. Hodg- don and Welsh, July, 1957 (NHA). B. papyrifera Marsh. X B. populifolia Marsh. Hodgdon et al., Aug., 1956 (NHA). “2,800 feet". *_Ostrya virginiana (Mill. K. Koch. H. I. Baldwin, Aug., 1971 (FPC). —Fagus grandifolia Ehrh. Hodgdon et al., Aug., 1956 (NHA, NEBC). i-Quercus rubra L. D. W. Allen, May, 1968 (FPC). Polygonum aviculare L. var. angustissimum Meisn. B. L. Robinson, July, 1897 (NEBC). * P. cilinode Michx. W. Deane, Aug., 1889 (HNH). +_P. scandens L. H. I. Baldwin, Aug., 1972 (FPC, NHA). “one plant, Red Spot Trail”. +_Claytonia caroliniana Michx. H. I. Baldwin, June, 1971 (FPC). *_Arenaria groenlandica Du Roi. Thomas Hope, Aug., 1879 (NEBC). Ranunculus abortivus L. Rand and Robinson, June, 1898 (NEBC). +—R. acris L. H. I. Baldwin, July, 1971 (NHA). +_R. bulbosus L. D. W. Allen, June, 1968 (NHA). i-Thalictrum polygamum Muhl. H. I. Baldwin, May, 1969 (NHA). *_Coptis groenlandica (Oeder) Fern. D. W. Allen, May, 1968 (FPC). 3—Actaea pachypoda Ell. H. I. Baldwin, May, 1969 (FPC). *_Aquilegia canadensis L. Rand and Robinson, May, 1897 (NEBC). -Berberis Thunbergii DC. W. H. Manning, Oct., 1913 (NEBC). _Drosera intermedia Hayne. Fred M. Hunt, Aug., 1946 (NHA). +_D. rotundifolia L. H. I. Baldwin, July, 1968 (FPC, NHA). +-Chrysosplenium americanum Schwein. D. E. Boufford, 1971 (KSC). *Ribes cynosbati L. T. Otis Fuller, June, 1883 (NEBC). * R. glandulosum Grauer. J. R. Churchill, May, 1883 (NHA). 216 Rhodora [Vol. 76 R. triste Pall. T. L. Steiger, May, 1937 (NHA). *_Spiraea latifolia (Ait.) Borkh. A. W. Driggs, Aug., 1916 (NEBC). "above timberline". 1-8. tomentosa L. H. I. Baldwin, July, 1968 (FPC). *—Pyrus melanocarpa (Michx.) Willd. C. F. Batchelder. Sept., 1909 (NHA). *—P. americana (Marsh.) DC. J. R. Churchill, July, 1890 (HNH). P. decora, (Sarg.) Hyland. Hodgdon and Kenneth Welsh, July, 1957 (NEBC). ‘2,500 feet". P. americana L. X P. decora (Sarg.) Hyland. Hodgdon and Kenneth Welsh, July, 1957 (NEBC). “2,500 feet". *—Amelanchier laevis Wieg. Rand and Robinson, May, 1897 (NEBC). *_A. Bartramiana (Tausch.) Roemer. J. R. Churchill, May, 1885 (NEBC). A. Bartramiana (Tausch.) Roemer. X A. neglecta (?) Rand and Robinson, May, 1897 (NEBC). T-Crataegus macrosperma Ashe. D. W. Allen, May, 1968 (NHA). T-Fragaria virginiana Duchesne. D. W. Allen, May, 1968 (FPC). Potentilla fruticosa L. J. A. Bates, Aug., 1900 (NEBC). *—P. tridentata Ait. E. S. Hoar, July, 1857 (NEBC). P. norwegica L. A. W. Driggs, Aug., 1916 (NEBC). “sum- mit". T—P. canadensis L. H. I. Baldwin, June, 1969 (FPC, NHA). T—P. simplex Michx. H. I. Baldwin, May, 1970 (FPC). Geum aleppicum Jacq. var. strictum (Ait.) Fern. Henry Purdie, July, 1905 (HNH). Rubus hispidus L. J. P. Poole, July, 1939 (HNH, NHA). R. setosus Bigel. A. W. Driggs, Aug., 1916 (NEBC). *_R. vermontanus Blanch. B. L. Robinson, July, 1897 (NEBC). R. allegheniensis Porter. A. W. Driggs, Aug., 1916 (NEBC). T—R. frondosus Bigel. H. I. Baldwin, July, 1968 (FPC). *—Prunus pensylvanica L. f. C. W. Jenks, Aug., 1883 (NEBC). T—P. serotina Ehrh. H. I. Baldwin, July, 1968 (FPC). t-Trifolium repens L. D. W. Allen, June, 1968 (FPC). 1974] Flora of Mount Monadnock — Baldwin 217 _Oxalis montana Raf. T. Otis Fuller, June, 1883 (NHA). Polygala sanguinea L. R. H. Piper, July, 1927 (MASS). * Nemopanthus mucronata (L.) Trel. Rand and. Robinson, May, 1897 (NEBC). * Acer spicatum Lam. Sydney Harris, June, 1894 (NEBC). * A. pensylvanicum L. C. W. Jenks, Aug., 1883 (NEBC). 1—A. saccharum Marsh. H. I. Baldwin, June, 1968 (FPC). * A. rubrum L. H. I. Baldwin, June, 1968 (FPC, NHA). Hypericum perforatum L. J. P. Poole, July, 1939 (HNH). _H. canadense L. Mary E. Gilbreth, Aug., 1889 (NEBC). (one extremely small plant found in 1971, H. I. Bald- win) +—Viola papilionacea Pursh. H. I. Baldwin, May, 1969 (NHA). V. sororia Willd. E. F. Williams, May, 1898 (NEBC). V. fimbriatula Sm. E. F. Williams, May, 1898 (NEBC). —V. pallens (Banks) Brainerd. A. W. Cheever, Sept., 1910 (HNH). V. blanda Willd. J. R. Churchill, July, 1890 (HNH, NEBC). V. lanceolata L. A. R. Hodgdon, June, 1933 (NHA). 1-V. rotundifolia Michx. H. I. Baldwin, April, 1969 (NHA). V. adunca Sm. Rand and. Robinson, May, 1897 (NEBC). 1—V. conspersa, Reichenb. H. I. Baldwin, May, 1971 (FPC). Circaea alpina L. Edward S. Hoar, 1879 (NEBC). * Epilobium angustifolium L. H. I. Baldwin, Aug., 1971 (FPC). E. leptophyllum Raf. C. W. Jenks, Aug., 1883 (NEBC). Aralia racemosa L. Hattie Merrifield, July, 1879 (NEBC). * A4. hispida Vent. J. P. Poole, July, 1939 (HNH, NHA). _A. nudicaulis L. A. S. Goodale, May, 1926 (MASS). 1—Panaz trifolius L. D. W. Allen, May, 1968 (FPC, NHA). * Cornus canadensis L. C. W. Jenks, Aug., 1883 (NEBC). —C. alternifolia L. f. Hodgdon et al., Aug., 1956 (NHA). Chimaphila umbellata (L.) Bart. var. cisatlantica Blake. J. P. Poole, July, 1939, (HNH, NHA). +-Monotropa uniflora L. H. I. Baldwin, Sept., 1968 (FPC). (rare). * Ledum groenlandicum Oeder. Sydney Harris, June, 1894 (NEBC). 218 Rhodora [Vol. 76 *—Rhododendron canadense (L.) Torr. Sydney Harris, June, 1883 (NEBC). *—-Kalmia angustifolia L. J. P. Poole, July, 1939 (HNH). *-Cassandra calyculata (L.) D. Don. D. W. Allen, May, 1968 (FPC). *—Epigaea repens L. H. I. Baldwin, April, 1969 (FPC). “(one plant only, on cliff) ." *Gaultheria hispidula (L.) Bigel. C. G. Trow, 1871 (MASS). 1T-Calluna vulgaris (L.) Hull. H. I. Baldwin, May, 1965 (NHA). (one isolated colony (introd)). -Gaylussacia baccata (Wand.) K. Koch. A. W. Driggs, Aug., 1916 (NEBC). Vaccinium myrtilloides Michx. Hodgdon et al., Aug., 1956 (NHA). “2,800 feet", *_V. angustifolium Ait. W. Deane, Aug., 1889 (HNH). -V. angustifolium Ait. var. laevifolium House. B. L. Rob- inson, July, 1897 (NEBC). T-V. augustifolium Ait, X V. corymbosum L. H. I. Baldwin, Aug., 1969 (NHA). V. angustifolium Ait. var. nigrum ( Wood) Dole. Fred M. Hunt, Aug., 1946 (NHA). V. corymbosum L. A. S. Goodale, May, 1926 (Mass). *-V. Vitis-Idaea L. var. minus Lodd. C. G. Trow, Aug., 1864 (MASS). T—Asclepias syriaca L. H. I. Baldwin, July, 1971 (FPC). *—Trientalis borealis Raf. D. W. Allen, May, 1968 (FPC). Lysimachia terrestris (L.) BSP. J. P. Poole, July, 1939 (HNH, NHA). T-Fraxinus americana L. H. I. Baldwin, July, 1968 (FPC). Hedeoma pulegioides (L.) Pers. B. L. Robinson, Aug., 1896 (NEBC). f-Veronica longifolia L. Hodgdon and Hehre, Aug., 1969 (NHA). 1—-V. officinalis L. H. I. Baldwin, June, 1969 (FPC, NHA). Gerardia tenuifolia Vahl. B. L. Robinson, Aug., 1898 (NEBC). *“_Melampyrum lineare Desr. var. americanum (Michx.) Beauv. D. W. Allen, June, 1968 (FPC). 1974] Flora of Mount Monadnock — Baldwin 219 1-Mitchella repens L. H. I. Baldwin, Aug., 1968 (FPC, NHA). rare. tT-Houstonia caerulea L. D. W. Allen, May, 1968 (FPC, NHA). *_Diervilla Lonicera Mill. C. W. Jenks, Aug., 1883 (NEBC). *_Viburnum alnifolium Marsh. C. W. Jenks, Aug., 1883 (NEBC). *_V. cassinoides L. C. L. Canfield, June, 1931 (NHA). *_Sambucus pubens Michx. W. Deane, July, 1891 (NEBC). T-Solidago caesia L. H. I. Baldwin, Sept., 1970 (FPC). *_S. macrophylla Pursh. W. Deane, Aug. 1891 (HNH, NEHA, MASS). —S. puberula Nutt. Mary E. Gilbreth, Aug., 1889 (NEBC). —S. Randii (Porter) Britt. C. W. Jenks, Aug., 1883 (NHA). S. Randii (Porter) Britt. var. monticola Porter. C. F. Batchelder, June, 1919 (NHA). S. juncea Ait. Parsons and. Seymour, Aug., 1927 (MASS). —S. nemoralis Ait. Fred M. Hunt, Aug., 1945 (NHA). *—S. rugosa Ait. var. rugosa H. I. Baldwin, Oct., 1968 (NHA). *_S. graminifolia (L.) Salisb. H. I. Baldwin, Sept., 1968 (FPC, NHA). T—Aster divaricatus L. H. I. Baldwin, Sept., 1968 (FPC). *—A. acuminatus Michx. B. L. Robinson, Aug., 1896 (NEBC). T-Erigeron pulchellus Michx. H. I. Baldwin, July, 1968 (FPC). *—Anaphalis margaritacea (L.) B.&H. D. W. Allen, May, 1968 (FPC). Gnaphalium Macounii Greene. C. W. Jenks, Aug., 1883 (NHA). G. obtusifolium L. C. A. Cheever, Sept., 1910 (HNH). G. uliginosum L. W. Deane, Aug. 1891 (HNH). *—Achillea Millefolium L. D. W. Allen, May, 1968 (FPC). t-Chrysanthemum Leucanthemum L. var. pinnatifidum Le- coq and Lamotte. H. I. Baldwin, July, 1971 (FPC, NHA). Taraxacum erythrospermum Andrz. E. F. Williams, May, 1898 (HNH). —T. officinale Weber. E. F. Williams, May, 1898 (NEBC). T-Prenanthes trifoliata (Cass.) Fern. var. nana (Bigel.) Fern. D. W. Allen, May, 1968 (FPC). 1—P. altissima L. H. I. Baldwin, Aug., 1968 (NHA). 220 Rhodora [Vol. 76 Additional taxa reported by Thoreau in his journals of 1858 and 1860. The names, as given by Thoreau, including his question marks are at the left in the following list: Contemporary accepted binomials are listed at the right opposite the name he used if that name is no longer used or if only a common name were given by him. Sensitive fern Black spruce Cinna arundinacea Glyceria elongata Eleocharis tenuis Scirpus Eriophorum Carex straminea C. scoparia ? C. debilis C. Deweyana Juncus acuminatus ? J. parodoxus ? Salix lucida S. humilis Tiarella cordifolia Amelanchier canadensis var Botryapium A. canadensis var oblongifolia Geranium Robertianum Epilobium palustre Water andromeda Checkerberry Chelone glabra Solidago virgaurea Onoclea sensibilis L. No Picea mariana (Mill) BSP. has been found since Tho- reau’s visits; this may be P. rubens Sarg. G. melicaria (Michx.) F. T. Hubbard S. cyperinus (L.) Kunth. C. tenera Desv. J. acuminatus Michx. or J. can- adensis J. Gay perhaps J. canadensis or J. acu- minatus A. arborea (Michx. F.) Fern. A. canadensis (L.) Medic. A. glaucophylla Link or Kalmia polifolia Wang. Gaultheria procumbens L. ? 1974] Flora of Mount Monadnock — Baldwin 221 Three-ribbed goldenrod (not canadensis) S. altissima L. or S. gigantea Ait. Aster thyrsoidea ? Diplopappus umbellatus Aster umbellatus Mill. DISCUSSION The present flora of the upper part of Mt. Monadnock is as noteworthy for the absence of certain plants as by the presence of unusual ones such as the arctic-alpines. Thus Alnus has not been found anywhere on the slopes of the mountain, nor has Picea mariana been recorded from the spruce-lined bogs. Acer saccharum is confined to a small area on the east side. Further there are many instances of lowland adventives that take root and may thrive for a time and then disappear. This may explain why some spe- cies recorded earlier have not now been found (aside from less keen observation). A total of 220 taxa are represented by specimens in herbaria, including 152 taxa collected from 1965 to 1972. Fifty-nine taxa collected during this period were not previously found in herbaria, and may possibly have been collected for the first time on the mountain. Of the 94 taxa reported by Thoreau, 68 of which are in her- baria, 62 were collected during the present survey. Thoreau observed seeds of many lowland plants blowing over the summit, and reflected on the origin of adventives (and perhaps transitory taxa) : “I saw what I took to be a thistle-down going low over the summit, and might have caught it, though I saw no thistle on the mountain top, nor any other plant from which this could have come. (I have no doubt it was a thistle by its appearance and its season.) It had evidently come up from the country below. This shows that it may carry its seeds to higher regions than it inhabits, and it suggests how the seeds of some mountain plants, such as the Solidago thyrsoidea may be conveyed from mountain to mountain, also other solidagos, asters, epilobiums, willows, etc." Tho- reau: Journal August 9, 1860, (p. 50) (page 1677 in the 1962 Dover edition). 222 Rhodora [Vol. 76 It seems obvious that most of the wind-disseminated spe- cies that have restocked the mountain since the 1820 fire, including Red Spruce, have come by the aerial route, mostly probably from the adjacent lowland. Light-seeded arctic- alpines may be included. It is less easy to explain how heavy-seeded plants such as Vaccinium arrived. These may have been present before the fires, surviving the shade of spruce on some steep ledges, or they may have been carried by birds migrating from timberline areas in the White Mountains. According to Pease (1964 p. 15) the following taxa col- lected or reported from Mt. Monadnock “may be listed as essentially alpine": Lycopodium Selago (including var. appressum) Lycopodium annotinum var. pungens Juncus trifidus Arenaria groenlandica Solidago macrophylla var. thyrsoidea Mt. Monadnock appears to be the southern limit for some of these. The relation of fire to the occurrence of arctic-alpine plants needs more investigation. It is curious that they be- came so abundant on Mt. Monadnock 29-30 years after the great fire. Do the ashes from fire favor their establishment (Fernald (1907) thought alkaline soil most unfavorable) or is it open space, exposed rock crevices, or mineral soil alone? Many years ago, searching for Vaccinium Vitis- Idaea on Cape Breton, I could find it only on an area that had been swept by fire about ten years before. There are thus at least two possible explanations for the presence on Mt. Monadnock of taxa usually found above timberline: (1) They have survived since post-glacial times in re- fugia on the mountain itself possibly on lower slopes as pro- posed by Fernald (1907, 1925), and then survived the fires (which were very severe indeed), and then restocked the bare rock and open soil. (2) The summit flora may have arisen from disseminules 1974] Flora of Mount Monadnock — Baldwin 223 carried from timberline areas by wind, animals or birds, or even men. Steele and Hodgdon (1973) reached the same possible explanations for the presence of Carex capitata and Carex Bigelowii on Mt. Cardigan. Regardless of the origin of the present flora, processes of change are at work, reducing the abundance and area occupied by some plants, while others are assuming dom- inance. Many lowland forms may invade and thrive for a time and then disappear. Neither the rate nor direction of succession is uniform. Invasion of spruce on the upper rocks has slowed to a temporary halt, even while the extent of arctic-alpine plants declines. The Problem of the Source and Persistence of Arctic-Alpine Plants How did arctic-alpine plants survive from post-glacial times on lower mountains and escape competition with the forest? One explanation for the presence of arctic-alpine plants below timberline is that they survived in refugia, rock crevices or smal] exposed ledges since post-glacial times, and that these crevices and ledges did not become over- grown by other plants and that the microclimate remained favorable. Then, when more open habitat was exposed by fire, these plants re-invaded the bare summits and increased greatly. Fernald (1907) postulated that alpine plants, “forced to find temporary homes during the ice age on potassic, calcareous or magnesium areas, found in the allu- vial deposits a mixed soil in which they were able to spread. As climate at low altitudes became warmer, these plants found on cold cliffs and exposed mountain summits the only habitats in which they have been able to persist within the temperate regions” It is fairly easy to understand that they were able to persist on alpine tundra above timberline, and indeed on some “cold cliffs’ like those in Dixville Notch, New Hampshire, and possibly elsewhere. Some plants usually restricted to high altitudes and open tundra habitats 224 Rhodora [Vol. 76 are occasionally found below timberline, e.g. on talus and rock piles in Carter Notch (a cool place) and even in some open pastures (where they may be rather temporary resi- dents). They cannot tolerate shade or root competition of lowland plants. A small colony of Mountain Cranberry is found along the Half Way House road on Monadnock (pos- sibly introduced from the summit by berry pickers). However, most steep rocky mountains that, like Monad- nock, do not reach timberline and have not been burned recently, are densely forested with spruce and at higher levels, fir, with a heavy ground cover of moss, ferns and herbs. There are probably hundreds of such mountains in the Adirondacks and New England. Some of them have exposed rocky cliffs, but records showing these to be havens for alpine plants in undisturbed condition are few or want- ing, Pease (op. cit.) reported finding some arctic-alpines on open ledges on secondary mountains in the White Moun- tains, *often caused by the destruction of the peaty humus by fires," but he did not specify just how often these cases were, nor did he give examples of such cases where there was no trace or history of fire, He ventured no theory as to how these plants reached the lowland habitat. If arctic-alpine plants persisted in refugia in lower moun- tains after the latter became forest-covered, it should be possible to find these plants on unburned summits. They have not been found, aside from the cases cited by Pease, which were all close to high mountains supporting stable colonies of these plants, whence invasion of even small burned areas would be relatively easy compared to Mt. Monadnock, which is far more remote. More examples of bona fide refugia from ice-age times need to be found, but even if these can be discovered and dated, it may be impossible to demonstrate that no fires occurred very long ago, or to prove how long the plants have been on the site. Arctic flora of the Pleistocene became adapted to the prevailing habitat conditions (Cf. Fernald, 1925) and it is still largely restricted to areas of open ground and a cool, moist climate. 1974] Flora of Mount Monadnock — Baldwin 225 Did Arctic-alpine Plants Survive Fire and Spread Over the Denuded Areas? Assuming that these plants had succeeded in resisting the competition of shade-tolerant trees and other plants, and had remained in some secluded pocket since post-glacial times, did they then survive the very intense fires that burned these peaks, more destructive than any fires in modern times? Fire destroyed all organic matter, and what little mineral soil remained was then at the mercy of wind and water. It must be remembered that two great fires on Monadnock combined to make a tremendously hot and ex- tensive burn. The 1800 fire consumed the humus and killed the trees. The 1815 hurricane completed toppling them, so that a formidible quantity of dry fuel was present. (It was after a long hot, dry summer.) No fires in recent times have approached it in heat and ferocity. Emerson (1896) remarked that the heat was so great that it “blasted the rock" into the ravines and caused slides. How did Arctic-alpine Plants Come to Occupy Areas Denuded by Fire? Such plants must have arisen from disseminules brought from nearby areas (refugia on the same mountain or in the lowland) or from more distant sources, probably stretches of alpine tundra on Mt. Moosilaukee or on the Presidential Range. Since the lowland and lower slopes around Monad- nock were largely cleared and cultivated or pastured, it is possible that some of these plants had escaped from rocky crevices on the mountain before the fire and had become established on the lower open land. Here they might main- tain themselves until forest succession once more eliminated them. At the time of the fire, this land was still open. Even if restocking of the burned mountain had proceeded from this source, it is hard to see how the spread of heavy-seeded species could have occurred uphill without the aid of birds or animals. The latter carry seeds in burs that cling to animal fur, but animals have not been found to be effective carriers of berries and similar hard seeds. 226 Rhodora [Vol. 76 Were Birds Partly Responsible for the Spread of Plants From Alpine Tundra to Monadnock? Birds are well-known to be active and efficient carriers of seeds. Ridley (1930) noted that birds are attracted by seed color. Red seems to be the most attractive. Vaccinium is a “bird-dispersed genus", he wrote. Forty-three species of birds have been recorded to disperse plants of this genus, and it seems likely that the red fruit of Mountain Cranberry would be very much sought by birds. Taylor (1954) sug- gested that birds are probably responsible for carrying seeds to Macquaric Island, a small sub-antarctic island far from any large land area that was severely glaciated during the Pleistocene. He believed it unlikely that any of the 35 plant species could have survived the ice. Most of these have relatively large seeds, but all have propagules suited to bird transport. Birds are important agents in the natural dissemination of seeds of tree species having fleshy fruits, and presum- ably shrubs as well. Red Cedar, Black Cherry and Black Gum are examples. Seeds of Juniperus that have been eaten by birds are said to germinate more promptly. (Anon. 1948) Indeed such hard-seeded species that display seed coat dormancy may require the acid treatment received during passage through a bird’s intestinal tract in order to germinate. Phillips (1910) made an especially thorough and wide-ranging study of bird dissemination of Juniperus seed, and concluded that birds are responsible for most of the spread of junipers, especially upwards on slopes and up stream courses. It is possible that the most likely candidate for being a vector of seeds from alpine fell fields is the Slate-Colored Junco. Not only did Thoreau remark about its being the most common bird on Monadnock, but it is also abundant there today. Forbush (1929) states that it is probably the only bird that nests above tree limit in the White Moun- tains. (I have found a nest with eggs in late June on Boott Spur.) Forbush (op. cit. p. 90) reported that it eats many 1974] Flora of Mount Monadnock — Baldwin 227 dried berries and seeds. Objections to the theory of bird transport include the questions concerning birds eating be- fore migration, seeds passing undamaged, and limits set by the time to pass through the digestive tract. Egler (1972) questions the explanation “by having far-flying constipated birds eject over distant lands.” While there is thus evidence that some birds do carry seeds, and birds are present on Mt. Monadnock, and that seeds survive the treatment, or may be even benefited by it, it is still not proven that heavy-seeded arctic-alpine plants did actually become established on Monadnock by this means. Since this was written, Steele and Hodgdon (1973) have examined these theories for the arrival on Mt. Cardigan of two alpine species of Carex. They suggest a third possibil- ity, that man may have consciously or unconsciously intro- duced these plants to the bare rocky area. This appears least likely in the case of the plants found on Monadnock, because of the distance. It is also improbable that these plants should appear on Mt. Chocorua and other sites. It would be a very diligent Johnny Appleseed indeed. More important, a prolonged period must be assumed for estab- lishment. Success probably did not come with the first seed. I am very grateful to Dr. Albion R. Hodgdon for checking the identity of specimens collected, and for helpful sugges- tions on the manuscript, also to Mr. David E. Boufford for searching herbaria and listing Monadnock records. LITERATURE CITED ANON. 1948. Woody-Plant Seed Manual. Misc. Pub. 654, U.S. Dept. Agri. 416 p. BRAUN, E. Lucy. 1950. Deciduous Forests of Eastern North Amer- ica. Blakiston, 596 p. CHAMBERLAIN, ALLEN. 1936. Annals of the Grand Monadnock. SPNHF, Concord, N.H. 195 p. CHILD, HAMILTON. 1885. Gazetteer of Cheshire County, N.H. 1736- 1885. Pub. by the Daily Journal, Syracuse, N. Y. 560 p. CUTTER, DANIEL B., M.D. 1881. History of the Town of Jaffrey, N.H. Concord, N.H. 648 p. 228 Rhodora [Vol. 76 DANA, JAMES FREEMAN. 1816. Some Account of the Grand Monad- nock. New England Jour. of Medicine and Surgery 5: 252-257. Boston, Mass. DEANE, WALTER. 1890. Flora of the Summit of Mt. Monadnock, N.H. in July. Bull. Torr. Bot. Club 17: 316-318. EGLER, FRANK E. 1972. (in a review) Ecology 53(5): 988. ELLIS, GEORGE B. 1880. Memoir of Jacob Bigelow. Proc. Mass. Hist. Soc. 17: 383-467. EMERSON, EDWARD W. 1896. The Grand Monadnock. New England Magazine 15: 33-51. FERNALD, M. L. 1907. The Soil Preferences of Certain Alpine and Subalpine Plants. Rhodora 9: 149-193. 1925. Persistence of Plants in Unglaciated Areas of Boreal America. Memoirs Am. Acad Arts & Sciences 15: 235- 342. . 1950. Gray's Manual of Botany. Eighth Edition. Am. Book Co., N.Y. 1632 p. ForBUSH, Epwarp Hower. 1929. Birds of Massachusetts and other New England States. Part III. Mass. Dept. of Agri. Norwood, Mass. 466 p. LEONARD, LEVI W. 1855. History of Dublin, New Hampshire. Bos- ton, Mass. 433 p. PEASE, A. S. 1964. A Flora of Northern New Hampshire. New England Botanical Club. Cambridge, Mass. 278 p. PHILLIPS, FRANK J. 1910. The Dissemination of Junipers by Birds. Forestry Quarterly 8: 60-73. RIDLEY, HENRY N. 1930. The Dispersal of Plants Throughout the World. L. Reeve & Co., Ashford, Kent, England. 744 p. Spurr, S. H. 1964. Forest Ecology. Ronald Press, N.Y. 352 p. STEELE, F. L. and A. R. Hopepon. 1973. Two Interesting Plants on Mt. Cardigan. Rhodora 75: 155-157. STONE, WILLIAM H. 1885. The Flora of Mt. Monadnock. Appalachia 4: 145-150. TAYLOR, R. W. 1954. An Example of Long Distance Dispersal. Ecology 35: 569-572. THOREAU, HENRY D. (1906) 1962. The Journals of Henry D. Tho- reau 1852-1860. Edited by Bradford Torrey and Francis H. Allen. 14 Vol. bound as two. Dover Pub. N.Y. HILLSBORO, NEW HAMPSHIRE 03244. TAXONOMY OF PERITYLE SECTION PERITYLE (COMPOSITAE — PERITYLINAE) * A. MICHAEL POWELL Section PERITYLE, with 27 species and three varieties, is the largest of three sections of the genus Perityle (Powell, 1968a). The distributional center of sect. PERITYLE is in northwestern Mexico and the Baja Californian peninsula, but the total natural range is extended by several taxa that occur in the southwestern United States, Guadalupe Island, the Revillagigedo Islands, and west-central Mexico (Jal- isco). One weedy taxon, P. emoryi, has bicontinental dis- tribution in North America and in Chile and Peru in South America. Separate taxonomic works have been prepared for the other two sections of the genus; Pappothrix (Powell, 1969) and Laphamia (Powell, 1973). The species of sect. PERITYLE comprise a taxon that was recognized as the genus Perityle by Everly (1947) who compiled a taxonomic treatment of the group. In connection with her morphologically oriented investigations of Perityle, Everly also studied the closely related taxa Pappothrix and Laphamia, which she regarded as genera. These taxa are distinguished primarily by somewhat variable pappus and achene characteristies, and several workers have questioned their status as genera. Shinners (1959) merged Pappothrix and Laphamia with Perityle, noting their close morpho- logical relationship and contending that pappus differences were not sufficient to warrant separate generic status. In recognizing Pappothrix and Laphamia as sections of Peri- tyle, I have followed Shinners’ congeneric disposition of the taxa (Powell, 1968a). My systematic judgements were based upon the evaluation of new information from cyto- logical, chromatographic, and hybridization techniques, as well as a re-evaluation of morphological] features. The re- 'Supported by National Science Foundation Grants GB-7740 and GB-20361. 229 230 Rhodora [Vol. 76 sults have suggested that Pappothrix, Laphamia, and Peri- tyle are closely related, essentially natural phylads, and that they are best treated as sections of a single genus. Niles (1970), however, has maintained Perityle and Lapha- mia as distinct genera, while combining Pappothrix with Laphamia. Niles did not conduct a comprehensive study of the sect. PERITYLE species. A more complete review of the taxonomic history concerning Perityle is available elsewhere (Shinners, 1959; Powell, 1968a; Niles, 1970). Perityle is closely allied to Pericome, Amauria, and Eutetras of the subtribe Peritylinae (Helenieae). Rydberg (1914) assigned Perityle and Pericome to the Peritylinae and Amauria and Ewutetras to the subtribe Amaurinae. Turner (1966) and Powell (1972a) have suggested that Amaurinae (in part) should be combined with Peritylinae. Presently the Peritylinae is being re-evaluated with refer- ence to Rydberg's concepts, and the proper tribal position of the subtribe is also being investigated (Powell & Turner, unpublished). CHROMOSOMAL CONSIDERATIONS A discussion of the chromosome numbers of Perityle and related genera is presented elsewhere (Powell, 1968b). Additional counts for the species of sect. PERITYLE are reported in Powell and Sikes (1970) and Powell (1972b). Chromosome numbers have been obtained for 20 of the 27 species of sect. PERITYLE. The counts for specific taxa are included with a list of the species under Phylogenetic Considerations (Table 1), and they are also included with species discussions in the taxonomic portion of this paper. In Powell (1968b), a few changes must be noted with regard to the counts for sect. PERITYLE. When the 1968 paper was compiled, the taxonomic studies of sect. PERITYLE were in early stages. The reports of n — 16 for P. cali- fornica should be attributed to P. cuneata var. marginata. Perityle incompta is now considered as synonymous with P. crassifolia var. robusta, and the counts listed for the former (n = 19, 18) should be attributed to the latter 1974] Perityle — Powell 231 taxon. The diploid P. microglossa (Sikes and Babcock 185 a,b,c,f,) is P. microglossa var. saxosa, and the polyploid P. microglossa (Sikes and Babcock 177 192 a-d, 152, 157 a, 206 a,b,) is var. microglossa. Sikes and Babcock 188 a,b,d,f, and 190 a,b, listed as P. palmeri, are P. cordifolia (n = YT). Perityle cf. spilanthoides is P. turneri (n — 1T) and P. vaseyi (Sikes 116) is P. parryi (n — YT). Most species of sect. PERITYLE are diploid, with counts of n — 19, 18, 17, 16, 13, 12, and 11 having been reported for the various taxa (Powell, 1968b; 1972b). Only P. micro- glossa var. microglossa (n = 34, 51), P. emoryi (n = 32-36, 50-57), and P. incana (n = 50-57) are known to be poly- ploid. The first two taxa are relatively well established as exclusively polyploid, while only one count for the Guada- lupe Island endemic P. incana (Powell, 1972b) has been recorded. Turner and Flyr (1966) reported n = ca. 46 for P. crassifolia var. robusta (Cowan 2265), but I have since identified this collection as P. emoryi. A base number for P. emoryi has not been determined in spite of numerous attempts to obtain exact counts (Powell, 1968b). The ancestral base number for sect. PERITYLE has not been established. Subsequent to an earlier discussion of the subject (Powell, 1968b), all attempts to resolve the question have been unsuccessful, including those which in- volved the analysis of experimenta] hybrids (Powell, un- published). Most chromosomal] and distributional evidence suggest a base of x — 17 for sect. PERITYLE, and the genus as a whole (see Phylogenetic Considerations). Fourteen species of sect PERITYLE are n = 17, and both sect. PAP- POTHRIX (5 spp.) and sect. LAPHAMIA (21 spp.) are x = 17 (Powell, 1969; 1973). The taxa of sect. PERITYLE with chromosome numbers below n = 17, P. californica (n = 13, 12, 11) and P. cuneata (n = 16, 12), are clearly aneuploid derivatives, probably of the » — 17 line. However, the origin of taxa with n = 19 (P. crassifolia and allies) is of prime concern in evaluating the phylogeny of sect. PERI- TYLE. Distributionally, the n = 19 “alliance” ( (southern Baja California peninsula and neighboring islands) is pe- 232 Rhodora [Vol. 76 ripheral to the group where n — 17, which is centered on the mainland of Mexico (see Phylogenetic Considerations). The » — 19 group may be either ancestral to or derived from the » — 17 group (Powell, 1968b), but I now believe that the species with » — 19 and the species with » — 17 both evolved as separate lineages from a common “main- land" ancestor that had a chromosome number of n = 19 (or 18). Consideration of the latter hypothesis in a broader sense is supported by the base numbers of the related genera Pericome (x = 18), Eutetras (x = 18), and Amau- ria (x = 18). ARTIFICIAL HYBRIDIZATIONS Artificial hybridizations have been conducted with 16 species of sect. PERITYLE. Many of the crosses were of intergeneric and intersectiona] nature, although several interspecific combinations have been obtained. The results of most of these crosses are discussed in another paper (Powell, 1972c) where emphasis was placed upon evalua- tion of the intergeneric and intersectional relationships of Perityle. A hybridization program aimed at the evaluation of interspecific relationships within sect. Perityle is still in progress (Powell, unpublished). Where appropriate, some of the preliminary information derived from artificial crosses will be brought into species discussions in the taxo- nomic portion of this paper. Natural hybridization is rare in sect. PERITYLE, and only one such occurrence has been documented (Powell, 1970). Most of the species are geographically isolated, but a few taxa do have overlapping ranges and may occasionally hy- bridize. CHROMATOGRAPHIC CONSIDERATIONS Nineteen species of sect. PERITYLE were analyzed by 2- dimensional paper chromatography of leaf extracts. In general, the techniques outlined by Alston and Turner (1963) were utilized. The procedures for sampling popu- 1974] Perityle — Powell 233 lations and analyzing chromatograms have followed those discussed elsewhere (Powell, 1973). The major components of chromatographic patterns were characterized as flavo- noids (Powell and Tsang, 1966; Powell and Averett, un- published), although the specific identity of only one com- pound is known (Southwick et al., 1972). The use of chromatographic techniques has been a valu- able asset to previous taxonomic studies in the Peritylinae (Powell and Tsang, 1966; Powell, 1968a; 1969; 1973). Intergeneric and infrageneric relationships have been clari- fied through the comparative analysis of pattern data. In previous work I treated the crude chemical information from a chromatographic pattern as a single taxonomic character, and pattern data have not been given more weight than any other chraacter. The results of chromatographic studies in sect. PERITYLE have revealed two strikingly different chemical patterns based upon the presence or absence of yellow compounds as seen under ultraviolet light. The white-flowered species centered in the Sierra Madre Occidental (e.g., P. turneri), and the yellow-flowered species of northwestern Mexico and Baja California (e.g., P. cordifolia and P. californica) have simple patterns consisting of a few dark purple spots. The Baja Californian white-flowered species (e.g. P. crassi- folia), and the Texas yellow-flowered species (e.g., P. par- ryi) have relatively complex patterns that include promi- nent yellow spots. Yellow compounds are considered im- portant because they comprise the basic chromatographic profile of sect. PAPPOTHRIX, sect. LAPHAMIA, and the genera related to Perityle. The actual chromatographic data are not presented here in the form of figures or tables. Instead, where appropriate, the data are included in succeeding dis- cussions concerning relationships of species and species groups. ECOLOGICAL AND REPRODUCTIVE CONSIDERATIONS The species of sect. PERITYLE occur in rock and in soil habitats, unlike the species of sect. PAPPOTHRIX and LAPHA- 234 Rhodora [Vol. 76 MIA, which are exclusively rock-dwelling (Powell, 1969; 1973). Certain species of sect. PERITYLE are obligate, rock- dwelling perennials, while several perennial and annual taxa are soil-obligates. In addition, a few taxa display facultative existence in rock and soil habitats. The rock-dwelling members of sect. PERITYLE occur under a wide range of climatic conditions and at various eleva- tions. About half of the petrophilic taxa are occupants of protected canyons and exposures in relatively xeric desert mountains at altitudes of 1000-4000 feet or less, and the others thrive under relatively mesic conditions at altitudes of 4000-8000 feet. Perityle turneri also occurs under mesic conditions, at 7000-9500 feet in and near Durango, Mexico, but may be rooted primarily in matted soils. The other soil- dwelling taxa occur almost exclusively under desert condi- tions. Perityle is essentially a genus of self-incompatible spe- cies (Powell, 1972c). Only two of the taxa, P. emoryi and P. microglossa var. microglossa, are known to be self-com- patible, and these are members of sect. PERITYLE. Ten spe- cies have not been tested for the compatibility factor, two of sect. LAPHAMIA and eight of sect. PERITYLE, but these taxa are also probably self-incompatible. Both of the self- compatible taxa are widespread polyploids (Powell, 1968b; 1972c) that are “weedy” and annual in habit. Perityle emoryi is the most widespread species of the genus, having bi-continental distribution in North and South America, and P. microglossa var. microglossa probably ranks second in distributional range. Several other species of the genus are polyploid, and a few others are annual, thus suggesting that the capacity for reproductive self-compatibility has allowed for the “weedy” nature of two taxa. PHYLOGENETIC CONSIDERATIONS Speculation concerning phylogeny of the subtribe Perity- linae and the genus Perityle has been discussed generally elsewhere (Powell, 1973), and will not be repezted in detail here. Essentially, it was proposed that Peritylinae origi- 1974] Perityle — Powell 235 nated in the southern Sierra Madre Occidental of Mexico. Perityle is by far the most successful genus of the subtribe, in terms of number of species, with only 2 or 3 species being recognized for each of the other genera (Powell, 1972a; 1972d; and Turner, 1966). Section PERITYLE is assumed to be ancestral to the other two sections of the genus. Based upon distributional, morphological, chromosomal, and other considerations, it is believed that sects. PAPPOTHRIX and LAPHAMIA underwent rapid evolution in late Pliocene or Pleistocene emanating from an ancestral sect. PERITYLE complex (Powell, 1975). Modes of speciation in the subtribe Peritylinae, and Peri- tyle in particular, have been discussed elsewhere (Powell, 1969; 1972c; 1973). In sects. PAPPOTHRIX and LAPHAMIA, speciation is believed to have occurred mostly, if not en- tirely, as a result of geographic isolation. In sect. PERITYLE, speciation probably has occurred as a result of geographic isolation and other, more nearly reproductive phenomena (Powell, 1972c). This subject wil be diseussed further below. In Table 1 the list of sect. PERITYLE species is presented as an aid to discussing phylogenetic considerations. The list has been organized to depict the hypothetical evolution- ary groupings within the section. The arrangements are based upon morphological, chromosomal, experimental, chromatographic, and eco-geographical data. Additional discussions of species relationships, including the evidence upon which such judgements are based, are included in the taxonomic portion of this paper. The total data suggest that sect. PERITYLE comprises at least two distinct evolutionary series. One series, including the first 19 taxa, is marked by white ligules and yellow (or cream-white) disc corollas, with a few exceptions, as indi- cated in Table 1. Perityle incana and the P. jaliscana group are relatively anomalous in the “white-flowered series" and they may comprise separate and equal phyletic entities. A second major series can be delimited by its exclusively yellow flowers. 76 [ Vol. Rhodora 236 ‘SNOGUOLLI Ə OF JYSNOY} ƏV jnq *exej eArjedsei ay}, J04 pojiodet useq eAvY seseyjueded ur s.9quin N.. 9I DYDULB LDU “TBA DYDOUNI “dT “CLG GI DIDIUNI "IBA DIVIUNI "dq “LZ GL ‘SL‘LL DIWAOJ{YDI"*I ‘9% (61) LI 149w]pnd'q “GZ LI 04000) 'd "yc pe1unoo 4ou uphüol]'d ec LI 0$$0]D02d9]'d ZZ LT )050f1p402'd "Ic LI (pioosrp) 08s0]bD'd '0c (6I) LI Wiosna'q ‘61 LI Wia4Dnd, ^q “RT SdüO0HD GQWHHWMO'LI-MOTTMXA pojunoo jou (proosrp) omuoppaí'qd "LI pejunoo 30u DJUOPUOYNA ‘d '9[ p91unoo jou 19804 `d GI LI (proosrp) pounosqp('q ‘I ISI 93IQA-urga4?) pue Se INYM LI “BO LT « (91) LI po1unoo jou SI ‘LT Ic ‘FS peyunod jou peyunod jou LT “89 ‘LT L€-0€ ‘9E-ZE 61 61 6l D1423$19WJ0 “dq “ST nyofidouoLod *q ‘ZT 0401% 'q “TT SUAISIUDI ‘d “OT DSOXDS “IBA DSSO)BOLIIU "q “G6 DSSOPBOLIWUE “IBA VSSOJBOLNU "qp. ` DIVYLIIOLIVNU * 0D90]1.409W1] `A 149W402 *d (proosrp) mwnDow'q (SABI Mo[[oÁ) 02470 “gq LOU ‘A SISUALLOIOS 'q DISNQOL “ABA D10f1$8049 `A “QT DYOJISSDLI “TBA DYOJISSVAI ^q “BT. SIG MOJA pu? SLEY INYM SdNOUD GAYAMOTH-ALIHM e Aled Has SOs ‘sdnois seroeds-peje[or peumnsaid puge 1009 IƏMOY 0} SurpJooo*€ PASI ext, sIequinu eurosouroIq9 orjeure3 jo Áueurums put seroeds AÁ uoreg 'I QLL 1974] Perityle — Powell 237 Several related-species groups are further delimited within both the ‘“white-flowered series" and “yellow- flowered series” (Table 1). The groupings themselves com- prise clearly related taxa, but the phyletic unity of the related-species groups within monophyletic white- or yellow- flowered series is indefinite. For example, origin of the Baja Californian P. crassifolia group has not been con- nected resolutely with the Sierra Madre Occidental P. turneri group, even though both exhibit white ligules. And the anamalous nature of P. incana and the P. jaliscana group is indicated above. Also, there is no certainty that the P. parryi group of Texas and the aneuploid P. cali- fornica group of Baja California have monophyletic ties within a yellow-flowered series. White-Flowered Groups (Table 1). Three subgroups of the white-flowered series are recognized. These are desig- nated as the Perityle crassifolia, P. turneri, and P. jaliscana related-species groups. Distributional information, habitat preferences, and life form habits are given in the following diseussions because of their supposed importance in under- standing speciation in the genus. The Perityle crassifolia group, also including P. socor- rensis, P. emoryi, and P. aurea, is distributed in southern Baja California and nearby islands, except for P. emoryi which is a widespread weed. Also, P. socorrensis is found only in the Revillagigedo Islands, which are over 200 miles south of Baja California. Perityle emoryi (n = 32-36, 50-57) is believed to be a polyploid derivative of P. crassifolia var. robusta. Morpho- logical and chromatographic data strongly suggest that P. aurea belongs with the P. crassifolia (x — 19) group even though this taxon has yellow-ligules and a chromo- some number of n = 17. Both Perityle emoryi and P. aurea are annual in habit, while the other taxa of the P. crassifolia group are essen- tially soil-dwelling perennials. All of the taxa, however, with the exception of P. crassifolia var. crassifolia which seems restricted to sand, might also occur in soil-filled 238 Rhodora [Vol. 76 crevices of rocks especially at sea side. The Guadalupe Island endemic P. incana, is possibly allied with the P. crassifolia group, but evidence as to its true relationship has not been conclusive. The base chromosome number of the Perityle crassifolia group is presumed to be x — 19, a number that might be ancestral in sect. Perityle. On morphological and distribu- tional grounds, however, it is practically impossible to en- vision the other related species groups as having been de- rived from the P. crassifolia line. I believe that the P. crassifolia line was an early offshoot from ancestral Peri- tylinae. The largest related-species group of the white-flowered series is distributionally centered in the Sierra Madre Occidental] of mainland Mexico. Two assemblages of four species each are further distinguishable among this Perityle turneri group, primarily on the basis of floral characters. Perityle turneri, P. microcephala, and P. lineariloba are clearly allied taxa which occur at relatively high altitudes from Durango to central Chihuahua. The fourth member of this assemblage is P. microglossa, which ranges from northern Sonora along the west coast of Mexico south to Colima, into the foothills of the Sierra Madre Occidental, and east to San Luis Potosi and Texas. Habit and distribu- tional considerations (see Taxonomy) strongly suggest that P. microglossa is derived from P. microcephala. I believe that P. turneri is primitive in this assemblage, on the basis of features discussed elsewhere (Powell, 1972e). Perityle coronopifolia, P. ciliata, P. hofmeisteria, and P. canescens are also clearly allied members of the P. turneri group. Perityle coronopifolia extends from northern Chi- huahua into the mountains of southern Arizona and south- ern New Mexico. Perityle ciliata is restricted to central Arizona, while P. hofmeisteria and P. canescens apparently are endemics respectively in Durango and in Sinaloa, Mexico. Most members of the Perityle turneri group are hardy, suffrutescent perennials that live in crevices of rock bluffs. 1974] Perityle — Powell 239 Perityle turneri is a rhizomatous perennial, apparently occurring typically in thin soil covering rocky outcrops, but according to label data, the taxon might also be rock-dwell- ing. Perityle microglossa is a soil-dwelling annual, also found occasionally among rocks, and one collection of P. hofmeisteria (Flyr 332) is said by the collector to be “seem- ingly an annual.” Judging from known distributional data, these species are strictly allopatric, suggesting that they have undergone the type of geographic speciation that is considered typical for sect. PAPPOTHRIX and sect. LAPHAMIA (Powell, 1969; 1973). The white-rayed (or discoid) Perityle jaliscana group with cream-white discs comprises very closely related rock- dwelling perennial taxa that are restricted to Jalisco, Mex- ico. These taxa represent a third distinct assemblage within the white-flowered species of sect. PERITYLE. The three white-flowered subgroups are clearly delimited morphologically, with perhaps the most useful distinguish- ing features being those of the habits, achenes, and pappus (see Taxonomy). The yellow-flowered species are quite different superficially, but the generic unity of both white- and yellow-flowered assemblages is evident. Yellow-Flowered Groups (Table 1). Geographically, the yellow-flowered species are divided into two major assem- blages separated by the Sierra Madre Occidental. One as- semblage, designated as the Perityle parryi group, has a limited distribution in the Big Bend area of western Texas and adjacent Mexico, and the other geographic assemblage, including the P. cordifolia and P. californica groups, is more widely distributed in Sonora, Sinaloa, and Baja California, Mexico. The three species of the Perityle parryi group are closely related perennials, being distinguished on the basis of a few morphological characters. One of the taxa, P. aglossa, apparently is exclusively rock-dwelling in habit, while P. vaseyi seemingly occurs only in desert soils. Perityle parryt, however, is facultative in its existence in rocks and soils. There are interesting biological parallels between the 240 Rhodora [Vol. 76 Perityle parryi group and the P. cordifolia group of west- ern Mexico, and despite their geographic separation, most evidence suggests that both groups had common origin. Both alliances comprise closely related taxa, their vegeta- tive and floral morphologies are strikingly similar, and species of both groups have chromosome numbers of n = 17 with no polyploidy having been discovered in any of the taxa. Additionally, and most remarkably, the evolutionary pattern of the disjunct groups seems to be that soil forms were established when woody perennials “came down out of the rocks and adapted to life in the soil.” Perityle cordi- folia, P. leptoglossa, and P. lloydii seem to be obligately (at least typically) rock-dwelling, while P. palmeri and P. lobata apparently are facultative in their tolerance for existence in rocks or in soils. The presumed “close” phylogenetic relationship between the Perityle parryi and P. cordifolia groups is not supported by their chemical profiles. Members of the P. parryi group produce abundant yellow components, as do P. crassifolia and allies, while chromatographic samples of the P. cordi- folia group have revealed only simple patterns of purplish spots. The chemical indication of affinity between the P. parryt and P. crassifolia groups is complemented by a tenuous morphological feature they have in common. Peri- tyle vaseyi (rarely P. parryi, also) of the former alliance and pappose members of the P. crassifolia group are the only taxa of the genus which have awns with retrorsely- barbed tips. In view of the morphological dissimilarity and geographic distance between these groups, however, I believe that the crude chemical data and pappus structure merely reflect an ancestral connection between the P. crassi- folia and yellow-flowered alliances. If this is true, then the P. cordifolia group must have lost its ability to produce the prominent yellow flavonoids. Both species of the Perityle californica group are soil- dwelling annuals. The taxa are of special systematic inter- est in sect. PERITYLE because of their reduced chromosome numbers. All other yellow-flowered taxa have m — 17. 1974] Perityle — Powell 241 Perityle californica (n = 13,12,11) is distributed in Sonora and Sinaloa of mainland Mexico and along most of the Baja California peninsula to just south of La Paz. Perityle cuneata (n = 16,12) is restricted to the southern tip of Baja California with northern limits to about the latitude of La Paz. Judging from overall morphological similarity and dis- tributional considerations, it is assumed that Perityle cali- fornica and P. cuneata are aneuploid derivatives of the P. cordifolia line. With the exception of the P. parryi group, which seemingly would be ruled out on distributional grounds, only the P. cordifolia group bears any close re- semblance to the aneuploids. Perityle californica occurs sympatrically with P. cordifolia and P. palmeri in Sonora and Sinaloa, apparently without hybridization. Since most species of Perityle are allopatric, but artificially interfertile (Powell, 1972c), it is reasonable to conclude that P. cali- forniea achieved reproductive isolation and speciation through reduction in chromosome number. Summary of Phylogenetic Considerations. Species of the genus Perityle are distributed primarily in the arid to semi- mesic mountains of desert North America, but also in the higher mountains, for example in the Sierra Madre Occi- dental. The majority of the species are obligate rock- dwellers. All five species of sect. PAPPOTHRIX and all 21 species of sect. LAPHAMIA occur as perennials in crevices of rock bluffs, apparently never growing in soil at the base of bluffs. That these species can survive only in such re- strictive habitats surely imposes upon them evolutionary limitations. The basic evolutionary trends of sect. PAPPO- THRIX and LAPHAMIA have followed a pattern of speciation by geographic isolation; thus evolved the groups of closely related, mostly endenmic species, characteristic of the above sections. Greater evolutionary diversity is exhibited among the species of sect. PERITYLE. Indeed, some species of this sec- tion are obligate, rock-dwelling perennials that presumably underwent the type of geographic speciation mentioned 242 Rhodora [Vol. 76 above. But several taxa have adapted to a facultative ex- istence in rocks and in soils, and a few species are exclu- sively soil-dwelling with perennial or annual habits. For certain taxa, the adaptation to life in the soil apparently has been facilitated by two evolutionary mechanisms, poly- ploidy and aneuploidy. Emanating from the white-flowered groups, the widespread weeds Perityle microglossa var. microglossa (n = 34,51) and P. emoryi (n = 32-36, 50-57) exemplify the adaptive success of polyploidy. Perityle cali- fornica (n = 11, 12, 19) and P. cuneata (n = 12, 16) are successful aneuploid species of yellow-flowered lineage. Thus, along with the annual habit displayed by the above species, the phenomena of polyploidy and aneuploidy have independently advanced the adaptive capacity of species belonging to distinct evolutionary units of sect. PERITYLE. ACKNOWLEDGEMENTS I am grateful to the curators of herbaria from which specimens were borrowed. I am also grateful to B. L. Turner who suggested the study of Perityle and provided encouragement and advice throughout the study. I extend thanks to John Averett for his assistance in gathering literature; to Reid Moran for his help in providing vouch- ers, buds, and ecological information for P. incana and other species; to Rogers McVaugh for providing helpful observations and material of P. microglossa and P. fedde- mae; and to Annetta Carter for her assistance in providing specimens and ecological information for several species. Many other people have contributed to my studies of Peri- tyle and related genera; they know of my appreciation. One person who did not know fully of my deep gratitude was the late Lloyd Shinners. His special character will linger to inspire many taxonomists. TAXONOMY PERITYLE Benth. Bot. Sulph. 23. 1844. Perityle section Perityle, see Powell, Sida 3: 277. 1968. 1974] Perityle — Powell 243 Galinsogeopsis Schultz-Bip. in Seem. Bot. Herald. 306. 1856. Nesothamnus Rydb. N. Amer. Fl. 34: 12. 1914. Leptopharynz Rydb. N. Amer. Fl. 34: 21. 1914. (in part). Closia Remy in Gay, Fl. Chile 4: 120. 1849. (in part). Plants shrubby, suffruticose, or herbaceous perennials, or delicate to robust annuals, (2)10-80 cm high, growing in rock crevices or various soils, variously pubescent, to- mentulose to nearly glabrous, frequently glandular-pu- bescent; leaves opposite or alternate, variable in size and shape, 0.7-14 cm long including the petiole, 0.2-7.0 cm wide, ovate, cordate, deltoid, subhastate, reniform, or suborbicu- lar in outline, the margins subentire, serrate to doubly so, or shallow to deeply lobed or divided, frequently 3-5 lobed, pedate, subcruciform, or pinnatifed with narrow segments, nearly glabrous, puberulent, to arachnoid-villous, often glandular-pubescent, thick and semisucculent to thin in texture, viscid, bitter tasting, usually aromatic; capitu- lescence of solitary heads, or heads loosely to tightly aggre- gated in clusters of 3 or more; peduncles short or long; involucres subcylindric to hemispheric; heads 3-12 mm high, 4-15 mm wide; involucres of 2(3) subequal series, bracts ovate or obovate to narrowly oblanceolate, obtuse to at- tenuate at the apex, flattened, ribbed, or keeled, usually spreading or reflexing at maturity; heads radiate or discoid (in 4 spp.), ligules yellow or white, showy or rudimentary ; disc flowers numerous, corollas 4-lobed, yellow or cream- white (in 3-4 spp.), throats tubular to broadly funnelform or narrowly campanulate; style branches, flattened, linear, usually tapering to a fine, minutely pubescent tip; achenes black when mature, 1.3-5.0 mm long, distinctly flattened radially, outer achenes often 3-angled, slightly curved and clasped by bracts, linear, oblong, oblong-elliptic, or oblance- olate, the margins densely ciliate, rarely merely puberulent, prominently calloused or the margins thin and not cal- loused, the faces glabrous and shiny or short-pubescent; pappus of a conspicuous, rarely inconspicuous, crown of laciniate, hyaline, squamellae, and 0-2 (rarely 3-4) delicate or rather stout bristles, 0.5-7.0 mm long, the bristles naked, 244 Rhodora [Vol. 76 barbellate, or subplumose throughout or only at the tips, the barbs antrorse, lateral, or retrorse; base chromosome number, x = 17 or 19. Type: P. californica Benth. Key to the Species 1. Heads discoid. 2 a somos otov t 9 9 n nm m m m rs n, n n n n n. n s s n. d 2. Leaves canescent; Guadalupe Lsland, Mexico. .... TEM 5. P. incana. 2. Leaves not canescent; Texas or Jalisco, Mexico . 3. 3. Pappus bristle 1, 3.7-5.5 mm long; disc corollas yellow; Texas. ................ 20. P. aglossa. 2 Pappus bristles 2(3), 0.5-1 mm long; dise corol- las cream-white or pale yellow; Jalisco. ...... 4. 4. Leaves dissected. ........ 17. P. feddemae. 4. Leaves deltoid to subhalberd. 14. P. jaliscana. 1. Heads radiate. .... ""arzsKAazPe»saerszrresruss D 5. Ray and disc corollas yellow. ................. 6. 6. Plants distinctly annual; pappus bristles 1 or 2 (or rarely absent). ....................... T. 7. Achene margins thin, not calloused; single pappus bristle delicate (rarely absent), and retrorsely barbellate only at the tip. ........ MM 4. P. aurea. 7. Achene margins usually prominently cal- loused; single pappus bristle rather stout and antrorsely subplumose, or 2 delicate bristles. .... 0.0.0.0... .0.0............ 8. 8. Pappus bristle 1, subplumose, 1.5-3.5 mm long. ...... 2 ee. 26. P. californica. 8. Pappus bristles 2 (rarely absent), delicate, 0.5-2.0 mm long. .................... 9. 9. Achenes (2.5) 3-3.8 mm long, obcor- date-cuneate, with broad, callous mar- gins. ... 27a. P. cuneata var. cuneata. 9. Achenes 1.5-2 (2.5) mm long, obovate to subcuneate with prominent callous 1974] Perityle — Powell 245 TOTS suc Se a 27b. P. cuneata var. marginata. 6. Plants perennial, with woody bases, or herba- ceous with fleshy taproots or rather thin branch Toota 2 ee ec ele oe 10. 10. Plants of West Texas and adjacent Mexico. ACTI NS DEUS or weer US sot FP 11. Leaves typically 3-lobed or subcruci- form; taprooted perennials in soil. .. DE AE cod DOT TON 10. P. vaseyt. 11. Leaves typically 3-lobed but not divided; woody-based perennials in rock crevices, or taprooted in soil ..... 18 P. parryi. 10. Plants of Sonora and Sinaloa, Mexico, and Baja California Sur. ..... eoe 12. 19. Plants subshrubs or suffrutescent peren- nidis. eur NEZX C S 13. 13. Leaves densely puberulent and sub- canescent. ..... 22. P. leptoglossa. 13. Leaves pilose and green. ...... 14. 14. Leaves 3.5-11 em long; involu- eral bracts 10-13 mm long. EM _.. 21. P. cordifolia. 14. Leaves 2.5-4.5 cm long; involu- eral bracts 5-7 mm long. .. . ee NICE 23. P. lloydii. 12. Plants herbaceous perennials with rather thin, fleshy roots. ........- 15. 15. Leaf blades 3-5 lobed, cleft, parted or rarely divided, the margins den- tate-lobed and acuminate; Baja California Sur. ..... 24. P. lobata. 15. Leaf blades shallowly or strongly 3.lobed, the margins serrate-den- tate; Sonora and Sinaloa, Mexico. Rl) MN OCA Um 25. P. palmeri. 5. Ray corollas white, disc corollas yellow, except disc corollas cream-white in P. rosea and P. trichodonta. meds cp c a 16. Rhodora [Vol. 76 16. Disc corollas eream-white. ... .. ....... 17. 17. Leaves 1.8-3 cm long, 1-1.5 (2) em wide; ligules 2-2.5 mm long. ....... 15. P. rosei. 17. Leaves 0.7-0.9 em long, 0.25-0.5 cm wide; ligules 1-1.5 mm long. .. 16. P. trichodonta. 16. Disc corollas yellow. .................... 18. 18. 18. Achene margins thin, not calloused ; heads 0.6-1 cm high, 0.6-1.5 cm wide, but possibly smaller in P. emoryi; Baja California Sur and islands, except P. emoryi widespread. DNUS 19. 19. Ligules 6-10 mm long. ......... 20. 20. Leaves usually thick and crisped, arachnoid-villous; Baja California Sur, coastal dune sand. [. . la. P. crassifolia var. crassifolia. 20. Leaves usually rather thin and not crisped, densely short-pubescent to glabrous; Baja California Sur, in various soils. .................. . lb. P. crassifolia var. robusta. 19. Ligules 1.5-4 (6) mm long (rarely ab- sent). ........ eee e eese. 21. 2]. Plants suffruticose perennials; Revillagigedo Islands. .......... ss 2. P. socorrensis. 21. Plants delicate or robust annuals; widespread weed. .. 3. P. emoryi. Achene margins thin or prominently cal- loused; heads 3-7 mm high, 4-8 mm wide, but may be wider in P. turneri; Sierra Madre Occidental, Mexico, and foothills, and Arizona, except P. microglossa wide- spread. ........0.00.0.-0..0.000 004. 22. 22. Achenes 1.8-3.5 mm long; pappus bris- tles 2 (3), 1.5-3 mm long. ....... 23. 23. Leaves deltoid-ovate to ovate- rhombic, the margins serrate, 1974] lS N Perityle — Powell 247 shallow-lobed, or serrate-crenate; central Arizona. ... 11. P. ciliata. Leaves ovate to subspathulate and entire to shallow-lobed, or ovate- cordate and serrate to serrate- lobed, or 2-3-pinnatifid to pedately divided; southern Arizona and Mexico. . ... SUM DNE 24. Leaves entire, shallow-lobed or serrate; Durango, Mexico. ANES 13. P. hofmeisteria. 24. Leaves pinnatifid or pedately divided with spathulate or linear segments; southern Arizona and adjacent Mexico. ow Haani P re er 25. 25. Capitulescence of several heads clustered on short peduncles; achene mar- gins typically long-ciliate. _.. 12. P. coronopifolia. 25. Capitulescence essentially of solitary heads; achene margins merely puberu- lent. 10. P. canescens. Achenes 1.3-2 mm long; pappus bris- tles 2 (or 0-2), 0.5-1.5 mm long. . 26. 26. 26. Leaves pinnately 3-5 divided, the divisions linear or nearly so. — D ! . 7. P. lineariloba. Leaves otherwise. ..... IDEE 27. Heads 0.7-1.4 cm wide. .. MINNS 6. P. turneri. 27. Heads 4-7 mm wide. . 28. 28. Plants suffrutescent per- ennials; leaves densely grayish-puberulent. . 8. P. microcephala. 248 Rhodora [Vol. 76 28. Plants delicate or robust annuals; leaves puberu- lent, glandular-puberu- lent, or glabrous. 29. 29. Ligules 1.5-3.5 mm long; upper pedun- cles usually copi- ously glandular-pu- berulent. .... .. 9a. P. microglossa var. microglossa. 29. Ligules 3.5-4.5 mm long; upper pedun- cles usually sparsely to densely puberu- lent, rarely moder- ately glandular-pu- berulent, . .. 9b. P. microglossa var. saxosa. 1. Perityle crassifolia Brandeg. Proc. Calif. Acad. II, 3: 147. 1891. TYPE: Mexico: BAJA CALIFORNIA: San Jose del Cabo, 6 Oct. 1890, T. S. Brandegee (Holotype, UC; isotypes, GH !, US(2) !). la. Perityle crassifolia var. crassifolia. Plants perennial, 10-75 cm high, younger plants with slender taproots, older plants with fleshy-woody taproots, younger plants erect and branching toward the top, older plants branching at the base, tending to be decumbent, stems pilose-villous to hirsute and glandular-pubescent; leaves mostly alternate, felty to touch, arachnoid-villous to short pilose-hirsute and glandular-pubescent, 2-3 (5) cm long, 1.5-2.0 (3.0) em wide, reniform to cordate in outline, thick and crisped, crenate to 3-5 lobed or dissected ; subses- sile to petiolate, the petioles 0.5-2.0 em long; capitulescence of 1-3 heads borne on peduncles 1-4 (6) em long; heads radiate, ca. 1.0 em high, 1.0-1.5 em wide, involucres hemi- 1974] Perityle — Powell 249 spherical; bracts numerous, ovate-lanceolate; ray flowers 12-16, ligules white, 6-8 mm long often pinkish-tinged, ob- long to elliptic or subspathulate; dise flowers numerous, corollas yellow, 3.0-3.5 (4.0) mm long, throats tubular- funnelform; achenes 2.5-3.0 (4.0) mm long, lanceolate- obovate and curved, callous margins absent or very thin, the margins densely ciliate, ray achenes 3-angled and pubescent on faces, disc achenes 2-angled and glabrous on faces; pappus of a short crown of squamellae and 1 (rarely 2) slender, barbellate bristle, 2-3 mm long, the distal barbs retrorse; chromosome number, n = 19. Apparently restricted to deep coastal sand from Punta Arena to San Jose del Cabo, Cape Region of Baja California Sur, and Isla Coronados. Flowering year around. (Fig. 1). REPRESENTATIVE SPECIMENS: Mexico: BAJA CALIFORNIA SUR: Los Frailes, S of Cabo Pulmo, Arnaud (ps); San Jose del Cabo, Brandegee (DS, GH, NY); SW end of Isla Coronados, Carter 4274 (DS, SD, UC, US); Punta Frailes, Daw- son 1140 (us); 3 mi N of Los Frailes, Hastings and Turner 64-279 (ARIZ) ; sand near shore, Isla Coronados, Moran 9121 (sp, vs); 0.5 mi E of Eureka, Powell and Turner 1849 (SRSC, TEX) ; San Jose del Cabo, Purpus 444 (us) 274, 446 (vc); 0.6 mi S of Buena Vista, Wiggins 14747 (CAS, DS, GH, TEX). This taxon is best distinguished from var. robusta by its arachnoid-villous pubescence, distinct perennial habit with fleshy to woody taproots in older plants, and habitat in coastal sand. Some considerations suggest that var. crassifolia and var. robusta should be accorded specific rank. Plants from one collection of var. crassifolia (Powell and Turner 1849) and several collections of var. robusta have been grown from seed under identical greenhouse conditions. The character- istie morphologies of both taxa persisted under artificial conditions, but var. crassifolia flowered rarely while var. robusta flowered profusely. Vigorous intervarietal hybrids were obtained, but only two flowering heads were produced on only one of several plants. Meiosis was regular and pollen stainability was ca. 3%. The reproductive data are 250 Rhodora [Vol. 76 too meager to allow speculation about relationships. Also, I have seen specimens of var. robusta (among borrowed material) which approach var. crassifolia morphologically. I believe that it is best to retain varietal status for the taxa, following Everly (1947), pending populational study of the habit and habitat of var. crassifolia. 1b. Perityle crassifolia var. robusta (Rydb.) Everly, Con- trib, Dudley Herb. 3: 382. 1947. Perityle robusta Rydb. N. Amer. Fl. 34: 16. 1914. TvPE: Mexico: BAJA CALIFORNIA: Cerralvo Island, 19 April 1911, J. N. Rose 16880 (Holotype, NY!; isotype, US!; isotype fragments UC (2) !). Perityle incompta Brandeg. Univ. Calif. Pub. Bot. 6: 508. 1919. TYPE: Mexico: BAJA CALIFORNIA : Los Dolores, W. E. Bryant (uc!). Perityle macromeres Blake, Proc. Biol. Soc. Wash. 37: 59. 1924. TYPE: Mexico: BAJA CALIFORNIA: La Paz 3 Feb 1906, E. W. Nelson and E. A. Goldman 1483 (us!). Plants perennial or annual, with fleshy taproots to slen- der branching roots, erect with few branches to spreading in large clumps with many branches, stems short-hirsute to glabrous; leaves densely short-hirsute and glandular-pu- bescent to glabrous, 2-10 cm long, 1-7 cm wide, ovate to cordate in outline, rather thick and crisped to thin and not crisped, typically deeply 3-5 lobed with the lobes also in- dented or with somewhat irregularly dissected margins; heads 0.6-1.0 cm high, 0.6-1.5 em wide, involucres hemi- spherical to campanulate; ligules 6-10 mm long; disc corol- las 2-8 (4) mm long; pappus bristle usually 1, rarely 0-3; chromosome number, n = 19, 18. Growing in various soils, including those which are sandy or saline near the sea and occasionally among rocks, rather common in southern Baja California Sur and neighboring islands. Flowering year around. (Fig. 1). REPRESENTATIVE SPECIMENS: Mexico: BAJA CALIFORNIA SUR: Isla Espiritu Santo, Berry (CAS); Isla Magdalena, Brandegee (NY, UC, US); Isla Santa Margarita, 1974] Perityle — Powell a ~ Ln Fig. 1. Distribution of P. crassifolia var. crassifolia (closed cir- cles); P. crassifolia var. robusta (open circles); P. aurea (closed squares); not plotted are P. socorrensis of the Revillagigedo Islands, and P. incana of Guadalupe Island. 251 252 Rhodora [Vol. 76 Brandegee (UC, US); San Jose del Cabo, Brandegee (POM); El Mogote Peninsula, La Paz Bay, Carter 2729 (CAS, DS, GH, UC, us); 45 km N of El Refugio, Carter, Alexander, and Kellogg 2152 (DS, UC, US); Puerto Escondido, Carter and Kellogg 2875 (ARIZ, DS, GH, SD, UC, US); Isla Partida, Collins, Kearney, and Kempton 166 (us); Isla San Francisco, Collins, Kearney, and Kempton 198 (vs); 10 mi W of Comondu, Gentry 4083 (ARIZ, DS, GH, UC); dunes, San Nicholas Bay, Johnston 3720 (CAS, GH, UC, us); Isla Coronados, Johnston 3756 (CAS, DS, GH, NY, UC, US); dunes, Isla Monserrate, Johnston 3865 (CAS, GH, UC, US); beach, Agua Verde Bay, Johnston 3893 (CAS, GH, NY, UC, US) ; dunes, Isla San Francisco, Johnston 3946 (CAS, DS, GH, NY, UC, US) ; Isla Cerralvo, Johnston 4046 (CAS); Isla Espiritu Santo, Johnston 4081 (CAS, GH, NY, UC, US) ; Guadalupe Point, Concepcion Bay, Johnston 4150 (CAS, DS, NY, uc); E base of San Lazaro, Santa Maria Bay, Moram 3537 (ps, UC); Ensenada de los Muertos, Moran 3560 (ps, sD); S end of Isla Cerralvo, Moran 9616 (ns, UC); Isla San Jose, Moran 3751 (ps, vC); Isla San Marcos, Moran 3948 (ps, UC) ; Isla Danzante, Moran 9209 (ps); W side of Isla Catalina, Moran 9329 (sp, UC); NE side of Isla San Jose, Moran 9387 (sp); Isla Las Animas Rock, Moran 9433 (sp, vC, US); Isla San Diego, Moran 9593 (sp); Isla Carmen, Palmer 1 (GH, NY, US); 10 mi N of Loreto, Powell and Sikes 1662 (SRSC, TEX); 2 mi NE of La Paz, Powell and Sikes 1681 (SRSC, TEX); 33 mi W of San Javier, Powell and Turner 1847 (sRSC, TEX); Isla Magdalena, Rose 16319 (Ny, US); Isla Pichilinque, Rose 16529 (GH, NY, US); 8 mi W of San Miguel, Shreve 7125 (ARIZ, DS, GH, us); 17 mi N of La Paz, Sikes and Babcock 270 (SRSC, TEX); 2 mi S of Villa Constitucion, Sikes and Babcock 276 (SRSC, TEX); ca. 29 mi N of Villa Constitucion, Thomas 8403 (CAS, DS, GH, US); 9 mi E of San Ignacio Wiggins 11360 (CAS, DS, GH, UC, US); San Gregoria, 12 mi NW of La Purisima, Wiggins 11467 (ps, GH, UC); 0.6 mi S of Buena Vista, Wiggins 14748 (CAS, DS, TEX, UC); Salino Flat, near S end of Isla Espiritu Santo, Wiggins 15595 (ps); Isla Partida, Wiggins 16160A (Ds). Perityle crassifolia var. robusta is more widespread than is var. crassifolia, and occupies a variety of edaphic condi- tions in coastal, insular, and inland areas in the southern portion of Baja California Sur. Two extreme forms of the morphologically variable var. robusta can be recognized. The plants of coastal habitats (e.g., near La Paz) usually are robust with fleshy, perennial taproots, while plants of the inland forms (e.g., near Commondu) are typically smaller with annual or weak perennial habits. Also, the inland form tends to be less pubescent and has thinner, less 1974] Perityle — Powell 253 crisped leaves than coastal plants. According to my inter- pretation, the “inland form” corresponds to P. incompta which Everly (1947) recognized as a distinct species. The “coastal form” conforms with Everly’s P. crassifolia var. robusta. Although the extremes of coastal and inland forms are evident, the existence of separate taxonomic entities has not been indicated. Instead, examination of exsiccata material and field studies have shown a rather complete morphological intergradation (e.g. Johnston 3946) between the two extremes. Hence P. incompta is treated here as synonymous with var. robusta. Further study is needed to clarify the adaptable habit of var. robusta, i.e., the perennial vs. annual conditions. Also, particular attention should be given to the habitats in which life forms occur. Those plants at La Paz grow in crusty saline soils, while plants of other populations may be in crevices of granite rocks, in rocky soil, or in fine sand. Another aspect of variability in var. robusta is presence or absence of pappus bristles, which seemingly is not taxo- nomically significant. As a generality, however, awnless forms occur on islands while awned forms are peninsular, but there are exceptions. 2. Perityle socorrensis Rose, Bot. Gaz. 15: 118. 1890. TYPE: Mexico: Socorro Island, Revillagigedo Islands, Mar. 1889, C. H. Townsend (Holotype, Us!; isotypes, GH !, NY !, US!). Plants suffruticose perennials, 10-40 cm high, many branched and densely leafy; leaves typically alternate, densely short-hirsute and glandular-pubescent, semisuccu- lent, 2-6 cm long, 0.8-3.0 cm wide, ovate, deltoid-ovate, cor- date to subhastate, 3-5 shallow-lobed and serrate to crenate ; petioles 1.5-3.0 cm long; capitulescence of 1-2 (3-5) heads borne on peduncles 1.0-3.5 (6.0) cm long; heads typically radiate, rarely discoid, ca. 6 mm high, 7-10 mm wide, in- volucres campanulate to narrowly so; bracts lanceolate to oblanceolate or oblong-lanceolate; ray flowers ca. 10 or fewer, ligules white, 2-4 mm long, oblong to oblong-elliptic, rarely with a prominent inner lobelet; disc corollas yellow 254 Rhodora [Vol. 76 (?), 2.0-2.5 mm long, throats tubular-funnelform ; achenes 2.9.8.0 mm long, oblanceolate-obiconical, oblong-oblanceo- late, to narrowly obconical typically curved, callous margins absent or very thin, the margins ciliate, ray achenes sparsely and minutely pubescent on faces, disc achenes glabrous or nearly so; pappus of a short to vestigial crown of squamellae and typically 2 (0-1) bristles, 1.5-2.0 mm long, the bristles retrorse barbellate (rarely lateral or an- trorse) distally; chromosome number, n = 19. Endemic to the Revillagigedo Islands, growing on sea cliffs and in soils near the shore. Flowering mostly in winter and spring. REPRESENTATIVE SPECIMENS: Mexico: REVILLAGIGEDO ISLANDS: Isla San Benedicto: Anthony 372 (CAS, DS, GH, NY, UC, US); Barkelew 175 (ARIZ, DS, GH, NY, POM, UC, us); Mason 1684 (cas, GH, UC, US); Isla Clarion: Anthony 415 (CAS, DS, POM, UC, US); Sulfur Bay, Howell 8347 (CAS, DS, NY, POM, us); W end of island, Mason 1578 (CAS, DS, GH, NY, POM, UC, US); Isla Socorro: Anthony 383 (ARIZ, CAS, DS, GH, POM, SD, UC, US); Barkelew 192 (ARIZ, DS, GH, NY, POM, UC, US); Academy Bay, Carl- quist 368 (CAS, RSA, UC); Brenner's Cove, Howell 8423 (CAS, DS, GH, uc, US); Graysons Cove, Moran 5922 (ARIZ, CAS, DS, GH, NY, RSA, SD, UC, US). Perityle socorrensis is a distinct species closely related to P. crassifolia from which it is distinguished by its typically 2 pappus bristles, reduced squamellae, generally smaller floral features, short ligules (or absent), woody bases, leaf morphology, and its distribution. As discussed by Everly (1947), Perityle socorrensis is notably variable in ligule and pappus features. The ligules, typically short at 2-4 mm, are even shorter and aberrant- looking in many collections, while other specimens are without ligules. According to label data, discoid individuals may occur in populations with radiate forms, and thus no taxonomic significance is attributed to ligule variability. Typically, the number of pappus bristles per achene in P. socorrensis is two, but some individuals or populations may have only one bristle or none. Bristle number is correlated 1974] Perityle — Powell 255 with the island to island distribution: plants with two bris- tles on Socorro and San Benedicto Islands, and awnless forms on Clarion Island. No taxonomic significance is at- tached to bristle variability since exceptions are found on all the islands, and because other features are not corre- lated with the pappus differences. 3. Perityle emoryi Torr. in Emory, Notes Mil. Rec. 142. 1848. TYPE: California: mountains E of San Diego, 29 Nov. 1846, Emory (NY!). Perityle nuda Torr. Pacif. R. Rep. 4: 100. 1857. TYPE: California: Williams’ River, 7 Feb. 1853-4, J. M. Bigelow (NY!). Perityle emoryi var. nuda A. Gray, Bot. Calif. 1: 397. 1876. Perityle emoryi S. Wats. Proc. Amer. Acad. 11: 116. 1876. Not P. emoryi Torr. 1848. Perityle californica A. Gray, Syn. Fl. N. Amer. 1: 321. 1884. Not P. californica Benth. 1844. Perityle californica var. nuda A. Gray, Syn. Fl. N. Amer. 1: 321. 1884. Perityle fitchii var. palmeri A. Gray, Syn. Fl. N. Amer. 1: 321. 1884. Perityle fitehii Green, Bull. Calif. Acad. 2: 403. 1887. Not P. fitchii Torr. 1857. Perityle californica Vasey, Proc. U, S. Nat. Mus. 11: 368. 1889. Not P. californica Benth. 1844. Perityle rothrockii Rose, Bot. Gaz. 15: 114. 1890. TYPE: Nevada: 1872, Wheeler (us!). Perityle greenei Rose, Bot. Gaz. 15: 117. 1890. TYPE: California: Santa Cruz Isl, July-Aug. 1886, E. L. Green (Holotype, ND?; isotypes, DS!, NY!, UC!, US!). Perityle emoryi var. orcuttii Rose, Bot. Gaz. 15: 117. 1890. TYPE: Mexico: BAJA CALIFORNIA. Canyon Cambel- los(?) July, 1884, C. R. Orcutt (Holotype, US!; isotype GH!). Perityle grayi Rose, Bot, Gaz. 15: 118. 1890. TYPE: Mexico: Guadalupe Isl., 1875, E. Palmer 44 (Holotype, Us?; isotypes, GH !, NY!). ~ "T 256 Rhodora [Vol. 7 IN Laphamia nuda Benth & Hook. ex Jacks. Ind. Kew. 30. 1895. Laphamia emoryi Benth & Hook, ex J acks. Ind. Kew. 2: 30. 1895. Partial Synonomy of South American P. emoryi (= Closia). Closia elata Phil Fl. Atac. 31 and Viage Des. Atac. 19, 205. 1860. Type not examined. Closia discoidea Phil. Fl. Atac. 31 and Viage Des. Atac. 205. 1860. Type not examined. Perityle emoryi Torr. var. elata. (Phil.) I. M. Johnston, Contr. Gray Herb. 85: 127. 1929. Perityle discoidea (Phil) I. M. Johnston, Contr. Gray Herb. 85: 128. 1929. Plants delicate or robust annuals, 2-60 cm high, usually herbaceous or the lower stems woody, erect or spreading with few to many stems, puberulent to hirsute and glandu- lar-pubescent; leaves mostly alternate, hirsute to glandular- pubescent, 2-10 cm long 1-5 cm wide, ovate, cordate, sub- orbieular, to triangular in outline, the margins deeply toothed, lobed, cleft, or divided, with the segments also indented to irregularly dissected; petioles 0.3-4.0 cm long ; capitulescence of 1 to many heads borne on peduncles 0.1- 7.0 em long ; heads radiate (rarely discoid), 0.4-1.0 cm high, 0.4-1.0 em wide; involucres hemispherical or campanulate; bracts numerous, lanceolate, oblanceolate, to ovate-lanceo- late; ray flowers 8-12 (14), ligules white, usually oblong, 1.5-4.0 (6.0) mm long, 1.5-3.0 mm wide, rarely rudiment- ary; disc flowers numerous, corollas yellow, 2.0-2.5 (3.8) mm long, throats tubular to tubular-funnelform; achenes (1.5) 2-3 mm long, suboblong, oblanceolate, or subcuneate, the outer ones often curved, margins thin, not calloused, the margins long- or short-pubescent (ciliate), outer achenes 2-3 (4) angled and often pubescent on the faces, inner achenes 2-angled with glabrous (rarely puberulent) faces; pappus of a vestigial or conspicuous crown of squa- mellae and 1 slender bristle, (0.8) 1.0-2.5 (3.5) mm long, antrorse-, lateral-, or retrorse-barbed especially at the tip. 1974] Perityle — Powell 257 Fig. 2. Generalized distribution of P. emoryt (stippled); South American distribution in Chile and Peru not plotted. 258 Rhodora [Vol. 76 or the bristle absent; chromosome number, tetraploid (n = 32-36) or hexaploid (n = 50-58). Widespread weed of desert southwestern United States, Sonora, Mexico, Baja Californian peninsula, and neighbor- ing islands; continental disjunct in Chile and Peru. Fower- ing mostly in winter and spring, but also year around, de- pending upon latitude. (Fig. 2). REPRESENTATIVE SPECIMENS: MEXICO: Baja California: Ensenada, Anthony 180 (CAS, Ds, UC, us); 8.2 mi S of Socorro, Constance 3119 (DS, GH, LL, UC, US); Bahia de los Angeles, Cowan 2265 (TEX); 14 mi S of Santa Rosa- lillita, Powell and Sikes 1647 (SRSC, TEX); 41.6 mi S of Mexicali, Powell and Turner 1710 (SRSC, TEX); Arroyo Calmalli, Purpus 23 (CAS, NY, POM); 14 km NW of Colonia Guerrero, Raven, Lewis, and Thompson 12193 (GH); 8.5 km E of El Rosario, Raven, Mathias, and Turner 12433 (ARIZ, GH); 15.9 mi N of San Felipe, Raven 14775 (ps, uc); N side of El Arco, Sikes and Babcock 295 (SRSC, TEX); 2 mi N of Miission de San Borja, Wiggins and Wiggins 14851 (DS); 9 mi S of Puertocito, Wiggins and Wiggins 15877 (Ds, US). Baja California Sur: Sierra de la Trinidad, Cape Region, Brandegee (UC, US); near Santa Rosalia, Ferris 8702 (Ds); Santo Domingo, Gander 9792 (sp); 10 mi W of Comondu, Gentry 4083 (ARIZ, UC); San Francisquito Bay, Johnston 3566 (CAS, GH, US); Concepcion Bay, Johnston 4150 (GH, US); E base of San Lazaro, Santa Maria Bay, Moran 3537 (sp); 18.6 mi. S of Mulege, Powell and Turner 1836 (SRSC, TEX); ca. 20 mi N of Santa Rosalia, Sikes and Babcock 285 (SRSC, TEX); 8 mi NW of San Ignacio, Sikes and Babcock 288 (SRSC, TEX); San Ignacio, Wiggins 16233 (ps); 3 mi N of El Barril, Wig- gins 16864 (ns); Mulege, Wiggins and Wiggins 18070 (us); 4 mi S of El Arco, Wiggins and Wiggins 18191 (CAS, DS). Islands: ISLA ANGEL DE LA GAURDA: Wiggins 17010 (ps). ISLA CARMEN: Moran 9181 (sp). ISLA CEDROS: Anthony 283 (CAS, DS, GH, NY, POM, UC, US); Haines and Hale (CAS, GH, LL, NY, SD, UC, US); Howell 10684 (CAS, DS, GH, NY, POM, US). ISLA CORONADOS: Moran 6556 (sD); Palmer 16 (GH, NY, US). ISLA GUADALUPE: Anthony 233 (CAS, DS, GH, UC, US); Carlquist 489 (RSA); Mason 1502 (CAs, GH, US); Moran 2900 (DS, GH), 5628 (DS, RSA, SD, UC); Palmer 891 (Ny, UC, US); Rose 16003 (GH, NY, US). ISLA ILDEFONSO: Moran 9062 (sp); Wiggins 18254 (CAS, DS). ISLA NATIVIDAD: Brandegee (UC); Moran 10797 (sp). ISLA PARTIDA: Johnston 3235 (CAS, US); Wiggins 17268 (DS). ISLA PARTOS: Johnston 3240 (CAS, GH, NY, UC, US). ISLAS REVILLAGIGEDO: Mason (CAS, US). ISLA SALSIPUEDES: Moran 8871 (SD, US). ISLA SAN BENITO: Anthony 271 (DS, GH, POM, UC); Palmer 914 (ARIZ, CAS, 1974] Perityle — Powell 259 DS, NY, US). ISLA SAN ESTEBAN: Wiggins 17218 (DS). ISLA SAN LORENZO: Moran 8895 (SD). ISLA SAN MARCOS: Johnston 3622 (CAS, GH, UC, US). ISLA SAN PEDRO MARTIR: Johnston 3148 (CAS, GH, UC, us); Moran 8810 (SD). ISLA TIBURON: Johnston 3268 (CAS, us). ISLA TODOS SANTOS: Moran 16214 (sD). ISLA TORTUGA: Wiggins 17367 (ps). Sonora: Puerto Penasco, Breedlove 1397 (ns); 27.4 mi S of Sonoyta, Breedlove 1389 (Ds, TEX); vicinity of Libertad, Graham 3803 (ps); 3 mi W of Kino, Heckard 1575 (JEPS); 1 mi E of San Carlos Resort, Powell and Sikes 1685 (SRSC, TEX); Empalme, Rose, Standley, and Russell 12636 (GH, NY, us); 5 mi NW of Caborea, Shreve 1532 (ARIZ); 4 mi NW of Caborea, Wiggins 8233 (DS, GH, UC, US). U.S.A.: Arizona: COCONINO CO.: Havasu Canyon, Grand Canyon, Howell 26537 (ARIZ, CAS, SMU); GILA CO.: Roosevelt Dam, Eastwood 6247A (CAS). MARICOPA CO.: Phoenix, Eastwood 6159 (CAS); near Tempe, Gillespie 8808 (DS, GH, POM, UC, us); 2 mi NW of Scottsdale, Russell 10911 (SMU, UC). MOHAVE CO.: 7 mi NW of Alamo, Benson 10074 (pom); 3.4 mi SE of Boulder Dam, Carter and Chisaki 3230 (ARIZ, DS, LL, NY, RSA, SMU, UC, US) ; Toroweap, Grand Canyon, Cottam 13001 (UT); 62 mi N of Wickenburg, Sikes 106 (SRSC). PIMA CO.: 10 mi W of Bates Well, Growler Mts., Benson 9926 (PoM); Gunsight Peak, Fosberg 7865 (POM); ca. 5 mi N of Ajo, Gould and. Macbride 4130 (ARIZ, GH, NY, UC); Alamo Canyon, Ajo Mts, Huey 24351 (cH, sp); Dripping Spring, Organ Cactus Natl. Mon. McClintock 52-37 (CAS). PINAL CO.: Sacaton, Gilman 350 (ARIZ); 10 mi W of Maricopa, Russell 11348 (smu). YUMA CO.: Gila Mts., near US 80, Barr and Lange 64-174 (ARIZ); S end of Castle Dome Mts., Parker, Wright and Lowe 7789 (ARIZ, DS, NY, RSA, us); 15.6 mi E of Yuma, Powell and Turner 1704 (SESC, TEX); S end of Cunningham Pass, Harcuvar Mts., Wiggins 8452 (ps, GH, UC, us). California: IMPERIAL co.: 8 mi from Niland to Blythe, Balls 12921 (RSA); 7 mi N of All American Canal spillway, Wiggins 8613 (DS, NY, POM, UC, US). Inyo co.: Furnace Creek, Death Valley, Carpenter (JEPS) ; Funeral Mts., Coville and Funston 324 (us); Hanaupah Canyon, Coville and Gilman 392 (vs); Darwin Falls, Hitchcock 6219 (uc); Surprise Canyon, Howell 3964 (CAS); Panamint Valley Smith 86 (JEPS). KERN CO.: Last Chance Canyon, El Paso Range, Twisselman 11863 (JEPS). LOS ANGELES CO.: Santa Catalina Isl., Brandegee (UC); Santa Monica Mts., Epling (DS, NY, RSA); Anacapa Isl., Howell 3816 (JEPS); San Clemente Isl., Raven 17345 (RSA, UC). RIVERSIDE CO.: Palm Springs, Abrams 11012 (ps); San Jacinto Range, Benson 4167 (pom): 8 mi NE of Desert Center, Wiggins 9675 (DS, GH. RSA, UC). SAN BERNARDINO CO.: 29 Palms, Alexander and Kellogg 870 (UC); near Parker Dam, Brenckle 51140 (SMU, uc); 89 mi N of Needles, Ferris 7224 (ps); between Kelso and Baker, Jepson 20590 (JEPS). SAN DIEGO CO.: 23.1 mi NW of Coyote Wells, Breedlove 1856 (DS); 260 Rhodora [Vol. 76 Yaqui Wells, Eastwood 2644 (CAS, GH, UC, US); Jacumba, Nelson 11182A (DS, GH, NY, POM, UC); Borego Park, Wolf 8462 (ARIZ, au, NY, RSA). SANTA BARBARA CO.: Santa Cruz Isl., Balls and Blakley 23727 (RSA, UC). VENTURA CO.: Point Mugu, Howell 3733 (cas). Nevada: CLARK COo.: between Las Vegas and Boulder Beach, Cron- quist 9844 (NY, UC); 1 mi below Boulder Dam, Grater 51 (vc); 8 mi SW of Davis Dam, Gullion 258 (Uc). CHILE: Atacama: Chanaral, Beetle 26164 (GH, UC); vicinity of Caldera, Gigoux (GH); below Agua EI Huerto, Johnston 3678 (vs); vicinity of Potrerillos, Johnston 4740 (GH, US); vicinity of Puerto de Chanaral, Johnston 4794 (GH, US); vicinity of Copiapo, Johnston 5028 (GH); Caldera, Johnston 5056 (GH); vicinity of Caleta Pan de Azucar, Johnston 5829 (Us); Vallenar, Werdermann 160 (GH); Tierra Amarilla, Werdermann 406 (GH, NY); Quebrada Paipote, Werdermann 448 (GH, NY). Antofagasta: Taltal, Jaffuel 984 (GH): Tocopilla, Jaffuel 1014 (GH); Antofagasta, Jaffuel 1126 (GH); 6 km N of Puerto Tocopilla, Johnston 3585 (Us). ECUADOR: Galapagos Isles: Cerros Isl, Stewart 40 (cas). PERU: Mts. near Chosica, Lima-Oroya Railroad, Weberbauer 5820 (GH, US). The morphological variation of Perityle emoryi is at- tested to by its considerable synonymy. This widespread annual, polyploid weed exhibits variation in nearly all as- pects of plant form. I have carefully examined the bulky exsiccatae available to me, giving particular attention to plant size, leaf morphology, head size, presence or absence of ligules and pappus bristles, and geographic distributions. None of the variable morphological aspects appear to have populational significance, and thus, in my judgement, do not require taxonomic recognition. I do note, however, that several atypical collections of P. emoryi from Magdalena Island resemble P. crassifolia and Amauria brandegeana, and thus the desirability of further study of Perityle from this locality is indicated. Perityle emoryi is related to and probably derived from the diploid P. crassifolia var. robusta, from which it is delimited by habit, smaller heads, shorter ligules, usually Shorter disc corollas, and chromosome number. With poorly preserved specimens it is often difficult to distinguish these taxa, but I have not had difficulty recognizing them in the field. Perityle emoryi is also remarkably similar to Amauria 1974] Perityle — Powell 261 brandegeana in superficial morphology. A discussion of the later similarity and distinguishing traits is to be found elsewhere (Powell, 1972a). It is assumed that P. emoryi achieved wide distribution as a result of the increased adaptability provided through a combination of polyploidy, reproductive self-compatibility and annual habit. That polyploids often exceed their di- ploids in distribution is well-known (Stebbins, 1950), and the advantage of self-compatibility in the establishment of disjunct colonies is clear. Indeed, Raven (1963) has sug- gested that P. emoryi might have attained bicontinental distribution as a result of long-distance dispersal from the Sonoran Desert, and he indicated the advantage of self- compatibility to such long-distance dispersal. More spe- cifically, I believe that the South American P. emoryt orig- inated from the vicinity of southern Baja California. It is in southern Baja California that the ancestors of P. emoryt are found. Furthermore, geographic variation in length of pollen spines (the spines ornamenting pollen walls) sug- gests that South American and Mexican P. emoryi had common areal origin. In measuring the pollen spines of some 20 populations of P. emoryi, it was found that the spine length of South American populations averaged the same or slightly longer than that of Mexican populations, while the spine length of United States populations was nearly twice as short as the others (Powell & Miller, un- published). From this information it can be postulated that P. emoryi originated and became established in Baja California, migrated to South America by long-distance dispersal in late Pliocene or Pleistocene (Raven, 1963), and subsequently advanced to the north, occupying desert areas in southwestern United States. Further study of the South American Perityle emoryi is needed, particularly chromosomal analyses and living plant comparisons. At least two forms can be recognized among the South American specimens I have seen, and this was discussed to some extent by Johnston (1929). One form (= P. discoidea (Phil.) I. M. Johnst.) deviates from North 262 Rhodora [Vol. 76 American P. emoryi in smaller habit, numerous, smaller and tightly clustered heads with no conspicuous ligules, and a pappus bristle on the achenes. Another form (= P. emoryi var. elata (Phil.) I. M. J ohnst.) is variable in habit, has large and small heads, longer and short ligules, and awnless achenes. I have not been able to make taxonomic distinctions between the South American and North Amer- ican forms, pending further study, and thus have placed Johnston’s combinations in Synonymy. It should be noted that only a partial synonymy of South American P. emoryi (Closia, in part?) is included since I have not seen speci- mens or types upon which several other Closia names are based. A few other collections from South America closely resemble P. crassifolia (e.g., Johnston 5023) or Amauria brandegeana (e.g., Werdermann 160; Morong 93). As mentioned above, these similarities are also seen in a few Mexican specimens. But it is also possible that both P. crassifolia and Amauria brandegeana once made their way to South America even though they have not become estab- lished there. 4. Perityle aurea Rose, Contr. U. S. Nat. Herb. 1: 84. 1890. TYPE: Mexico: BAJA CALIFORNIA: Santa Rosalia, 24 Feb.- 3 Mar. 1889, E. Palmer 185a (Holotype, Us!; isotypes, CAS !, GH !, NY!). Plants annual, 10-60 cm high, erect or spreading, stems of larger plants succulent, reddish, nearly glabrous; leaves alternate, puberulent to glabrous, slightly thick in texture, 1.7-9.0 em long, 1-4 cm wide, subreniform to ovate in out- line, basically 3-lobed with dentate-serrate lobes and mar- gins; petioles 0.6-5.0 em long; capitulescence of 1-3 heads borne on rather short peduncles 1-2 (3) em long; heads radiate, 5-8 mm high, 5-8 mm wide, involucres campanulate to narrowly so; bracts broadly lanceolate; ray flowers ca. 10, ligules yellow, oblong-elliptie to obovate; dise corollas yellow, 2-3 mm long, throats tubular to funnelform; achenes 2-3 mm long, narrowly oblanceolate to suboblong, with thin callous margins, the margins densely white-ciliate, ray achenes pubescent on faces, disc achenes glabrous or nearly 1974] Perityle — Powell 263 so on faces, pappus of a conspicuous crown of squamellae and 1 bristle (rarely 0) 1.5-2.5 mm long, the bristle retrorse barbelate with few barbs at tip; chromosome number, n= M. Restricted in soils and among rocks, near Santa Rosalia of Baja California Sur and a few islands in the Sea of Cortez. Flowering Jan-April. (Fig. i9» SPECIMENS EXAMINED: Mexico: BAJA CALIFORNIA: summit and N slope of high peak near SE corner of Isla San Esteban, Moran 8845 (sD, US); summit of NE peak, Isla San Esteban, Moran 13044 (sb); arroyo on S Isla San Lorenzo, Moran 13060 (SD). BAJA CALIFORNIA SUR: Isla San Marcos, Johnston 3614 (CAS, GH, UC, US); Isla San Marcos, Moran 3965 (ps); Santa Rosalia, Palmer 185a, 185 (CAS, GH, us); 8 mi W of Santa Rosalia, Powell and Turner 1826, 1829 (SRSC, TEX); 9 mi N of Santa Rosalia, Powell and Twrner 22219 (SRSC, TEX); 10 mi N of Santa Rosalia, Reed 6225 (ps, POM) ; ca 12 mi N of Santa Rosalia, Sikes and Babcock 281 (SRSC, TEX). Two growth forms of Perityle aurea are evident in the field, and both have been preserved in existing collections of the species. One form is of small plants, superficially resembling the Baja California populations of P. californ- ica, and the other form is of taller, more robust and succu- lent plants resembling P. crassifolia. The small form is known to occur in roadside soils where dry conditions pre- vail, and the large form occurs in places that receive and retain more water. The growth forms likely are ecologically controlled. I have seen both forms growing only a few paces apart, and when seeds of both plant sizes were grown under identical greenhouse conditions, only the larger, more succulent forms developed. The island specimens of P. aurea differ slightly from the mainland forms, most notably in achene morphology and in being awnless, although awned forms also occur on San Esteban Island. Perityle aurea seemingly is related to P. crassifolia, from which it is readily delimited by yellow ligules, chromosome number (n = 17), and distribution. An affinity of P. aurea with the P. crassifolia alliance is not indicated by its yellow 264 Rhodora [Vol. 76 ligules and chromosome number. All other members of the P. crassifolia group have white ligules and have x — 19. My first impression from superficial morphological exam- inations was that the species was related to P. californica (n = 13, 12, 11) and P. cuneata (n = 16, 12), both annual yellow-rayed taxa with habits similar to the small growth form of P. aurea. But P. aurea exhibits a combination of vegetative and floral features which characterize only the P. crassifolia alliance: robust and rather succulent habit, achenes with callous margins absent or thin, other unique achene morphology, and pappus bristles retrorsely barbed at the tips. Also, chromatographic studies have shown that P. aurea, like members of the P. crassifolia group, pro- duces the abundant yellow flavonoid compounds that have been detected elsewhere in sect. PERITYLE only in the P. parryi group of Texas. I conclude that P. aurea evolved, probably from P. crassifolia var. robusta, through specia- tion that involved adaptation of the annual habit and aneu- ploid reduction in chromosome number. Artifieial hybrid- izations have strongly suggested that the above taxa are reproductively isolated (Powell, 1972c). 5. Perityle incana Gray, Proc. Amer, Acad. 11: 78. 1876. TYPE: Mexico: BAJA CALIFORNIA: Guadalupe Island, Feb.- May, 1875, E. Palmer 43 (Holotype, GH!; isotypes, NY, US!), Nesothamnus incanus (Gray) Rydb. N. Amer. Fl. 34: 12. 1914. Plants shrubby, 10-40 (80) em high, many branched and densely leafy, stems tomentulose-canescent; leaves alter- nate, thickish, tomentulose-canescent, 6-14 em long, 4-7 em wide, deeply 3-divided or palmate, the divisions again deeply lobed or cleft; petioles 1.5-7.0 em long; capitules- cence of many tightly clustered heads (naked corymbs) borne on short peduncles; heads discoid, 5-7 mm high, 4-8 mm wide, involucres campanulate; bracts broadly oblong- lanceolate to broadly linear; disc corollas yellow, 2-3 mm long, throats narrowly campanulate or broadly tubular; 1974] Perityle — Powell 265 achenes 2.0-3.8 mm long, narrowly obdeltoid to narrowly oblanceolate, with prominent callous margins, the margins coarse-ciliate, the faces coarse-pubescent ; pappus crown of 2 broad, laciniate squamellae, 0.5-0.9 mm long, and rarely a narrow, fimbriate scale, 1.0-1.2 mm long; chromosome num- ber, n = ca. 50-57. Endemic to Guadalupe Island and islets, growing on cliffs near the sea. Flowering winter, spring, and early summer. REPRESENTATIVE SPECIMENS: MEXICO: Baja California: ISLA GUADALUPE: Anthony 249 (DS); Carlquist 440 (RSA, uc); Barracks Cove, Copp 157 (DS) ; Franceschi 7 (A, DS, POM, UC); Howell 8172 (CAS); Outer Islet, Lindsay 2621 (sp); Islote Negro, Mason 1519 (ARIZ, CAS, DS, us); Moran 2904 (ps, RSA); Outer Islet, Moran 5682 (CAS, DS, NY, SD, uc); North Twin Canyon, Moran 6133 (sp); N end of island, Moran 6435 (DS, sp); cliffs of the Lower Circus, Moran 12023 (sp); N side of Mt. Augusta, Moran 12042 (DS, SD, uc); SW Oak Canyon, Moran 13792 (SD). This cliff-dwelling Guadalupe Island endemic is perhaps the most distinct species of sect. PERITYLE. Its identifying features include: shrubby habit; tomentulose-canescent pubescence ; clustered, naked capitulescence; coarse pubes- cenct achenes; pappus of broad, laciniate squamellae and rarely à narrow, fimbriate scale. Considering its relatively large, shrubby habit, unique character, and isolated distri- bution, Perityle incana would seem to be a primitive mem- ber of the genus, although this possibility is not strength- ened by its hexaploid chromosome number. According to Reid Moran (personal communication), the taxon is not in danger of extinction, largely because its habitat is out of reach of the goats which have overrun the island for more than a century. It is clear that Perityle incana does not belong with any of the related-species groups, and thus stands as an anoma- lous species. In fact, the species could be recognized as a montotypic genus with little change in the taxonomic nat- uralness of the subtribe. I suspect, however, that the taxon is distantly related to the P. crassifolia group and is best 266 Rhodora [Vol. 76 treated as a member of Sect. PERITYLE. Perityle incana is similar to Pericome in habit and capitulescence characters, and perhaps shares ancestral affinity with that genus. 6. Perityle turneri Powell, Madrofio 21: 456-457. 1972. TYPE: Mexico: DURANGO: 3.4 mi E of Ey Palmito, 2 Apr. 1970, A. M. Powell 1858 (Holotype, TEX ! ; isotype, SRSC!), Plants herbaceous to suffrutescent perennials, decumbent, prostrate, or semierect, stems Spreading 20-45 em long, often purplish, lower stems often rooting at the nodes, upper stems puberulent to densely so, short-pilose, or sub- tomentose; leaves mostly opposite, puberulent to densely so, short-pilose to densely so, or subtomentose, often purplish, 3.0-4.5 (8.5) em long, 1.5-3.0 (4.0) em wide, ovate, deltoid, or subcordate, the apexes acute or attenuate, the Margins serrate, serrate-crenate, Serrate-lobed to doubly so, the ser- rations or lobes acute or acuminate; petioles 1.0-1.5 (3.0) cm long; capitulescence of 1-3 (4) heads on peduncles (1.0) 2-6 cm long; heads radiate, subglobose, 5-7 mm high, 0.7-1.4 cm wide, involucres hemispherical to broadly campanulate; receptacles conical; ray flowers ca. 12-18, ligules white, (3) 6-9 mm long, oblong; disc corollas yellow, (1.8) 2.0-2.8 (3.0) mm long, throats short-campanulate, short-funnel- form, rarely narrowly tubular-funnelform ; achenes 1.5-2.0 mm long, obovate to oblong-ovate, with thin or prominent callous margins, the margins ciliate, the faces puberulent in the center; pappus of a prominent crown of squamellae and 2 (rarely 1) unequal bristles, the longest 0.6-1.5 mm long; chromosome numbers n — 17,n — 17 II 4 I. Moist seeps and canyons, higher elevations to ca. 9500 feet, southern Chihuahua and Durango in the Sierra Madre Occidental. Flowering spring and fall. (Fig. 3). REPRESENTATIVE SPECIMENS: Mexico: CHIHUAHUA, 4 mi SW of Villa Matamoros, Correll and Gentry 22819 (LL); near La Rocha, NE slope of Sierra Mohinora, Correll and Gentry 23109 (LL); Burro Canyon near Parral, Pringle 18650 (ARIZ, Cas, GH, SMU, TEX, UC, US). Durango. Quebrada San Juan, ca. 50 mi W of Durango and 23 mi NW of Los Coyctes RR, 1974] Perityle — Powell 267 Fig. 3. Distribution of P. turneri (closed squares) ; P. lineariloba (closed star); P. microcephala (closed triangles) ; P. microglossa var. microglossa (closed circles); P. microglossa var. saxosa (open tri- angles); P. ciliata (open squares); P. coronopifolia (open circles) ; P. hofmeisteria (open star); not plotted is P. canescens. 268 Rhodora [Vol. 76 Cronquist 9568 (NY, SMU, TEX, US); 2 km S of El Espinozo del Diablo, and 65 km SW of El Salto, Cronquist and Fay 10786 (NY); 6 mi W of La Ciudad, Flyr 273 (TEX); ca. 35 mi W of El Salto, Gentry and Arguelles 18210 (Us) ; San Ramon, Palmer 60 (GH, NY, UC, US); 2.2 mi E of El Palmito, Powell and Turner 1857 (SRSC, TEX); 12 mi W of La Ciudad, Sikes and Babcock 380 (SRSC, TEX). SINALOA. 0.9 mi W of El Palmita, ca. 47 mi E of Concordia, Breed- love 1721 (Ds). Plants of this remarkable species were rarely collected until part of its high-altitude distributional range was made accessible by the road cut across the Sierra Madre Occiden- tal from Durango to Mazatlan. At the type locality near Mex. 40, Perityle turneri is abundant in wet mats formed by dense growths of Selaginella and small ferns. In overall aspect, at least at the type locality, the taxon shows striking resemblance to members of the tribe Astereae. Perityle turneri is related to P. lineariloba, from which it is easily delimited by leaf shape, and to P. microcephala, from which it can be distinguished by its capitulescence and head size. Further discussion concerning the taxonomy of this species is available elsewhere (Powell, 1972e). 7. Perityle lineariloba Rydb. N. Amer. Fl. 34: 18. 1914. TYPE: Mexico: DURANGO: San Ramon, 21 Apr.-18 May 1906, E. Palmer 89 (Holotype, NY!; isotypes, GH!, UC!, US!). Plants seemingly perennial, with semi-annual roots, stems lignescent near the base, decumbent, ca. 30 cm long, grayish- puberulent above; leaves mostly opposite, finely puberulent, 2.5-6.0 em long, 1.5-3.5 cm wide, pinnately 3-5 divided, the divisions linear or nearly so and entire or cleft into linear segments; petioles 0.7-2.0 em long; capitulescence of solitary heads on long peduncles, 4-10 cm long; heads radiate, 5-6 mm high, 5-7 mm wide, involucres broadly campanulate; ray flowers ca. 10-15, ligules white, 4-7 mm long, oblong ; disc corollas yellow (?), 1.8-2.0 mm long, throats narrowly campanulate-funnelform; achenes 1.3-1.5 mm long, nar- rowly obovate to oblong-obovate, with thin or prominent callous margins, the margins and faces short-pubescent ; pappus of a conspicuous crown of squamellae and 1 (rarely 1974] Perityle — Powell 269 none) delicate bristle, ca. 0.5 mm long; chromosome num- ber, unknown. Known only from the type collection. (Fig. 3). The distinguishing characteristics of Perityle lineariloba include its pinnately divided leaves, long peduncles, and rel- atively large heads with long ligules. Most of its floral features are like those of P. turneri, to which the species is related. 8. Perityle microcephala A, Gray, Proc. Amer. Acad. 21: 391. 1886, TYPE: Mexico: CHIHUAHUA: rocky hills near Chihuahua, 8 Oct. 1885, C. G. Pringle 571 (Lectotype, GH ! ; isolectotype, US!). LECTOPARATYPE: Chihuahua: mountains above Batopilas, Oct, 1885, E. Palmer 268 (GH) ; isolecto- paratypes, NY! US !). Plants suffrutescent perennials, 20-40 cm high, many- stemmed and densely leafy, erect or decumbent, densely grayish-puberulent to pilosulous; leaves mostly opposite, densely grayish-puberulent, 1.5-3.5 (5.0) em long, 0.8-2.0 (3.0) em wide, deltoid-ovate, ovate, to subcordate, the mar- gins serrate-crenate to doubly so, or shallow-lobed; petioles 0.3-1.0 (1.8) cm long; capitulescence of severa] heads tight- ly clustered on short peduncles; heads radiate, 3-4 mm high, 4-5 mm wide, involucres short-campanulate; ray flowers ca. 8-12, ligules white, 2-5 mm long, oblong; disc corollas yellow, often purple-tinged, 1.5-2.0 mm long, throats nar- rowly campanulate-funnelform; achenes 1.5-1.8 mm long, narrowly obovate to oblong-ovate, with callous margins, the margins and faces short-pubescent; pappus of a prominent crown of squamellae and 2 (rarely 1) unequal, delicate bristles, the longest 0.5-1.0 mm long; chromosome number, unknown. Sierra Madre Occidental of Chihuahua and Sonora. Flowering Aug-Nov, and probably in the spring also. (Fig. 3). REPRESENTATIVE SPECIMENS: Mexico: CHIHUAHUA. Damon s.n. (UC); Guicorichi, Rio Mayo, 2'(0 Rhodora [Vol. 76 Gentry 1973 (ARIZ, UC, US); near Batopilas, Goldman 215 (GH, NY, us); Madrono Mine, ca. 5 km NW of Pinos Altos, Hewitt 171 (GH); Guayanopa Canyon, Sierra Madre, Jones s.n. (POM, US); Santa Clara Mts., LeSueur 340 (ARIZ, CAS, GH, LL, TEX, UC); Mapula Mts., Pringle 764 (LL, NY, RSA, UC, US). SONORA. La Mesa Colorado, Gentry 541, 541m (ARIZ, DS); Sierra de Papas, Gentry 630 (Ds). Everly (1947) treated Perityle microcephala as a syn- onym of P. spilanthoides. I have found, however, that P. microcephala represents a valid species, while P. spilan- thoides is synonymous with the weedy variety of P. micro- glossa. Perityle microcephala belongs with the group of white- rayed species which includes P. turneri, P. lineariloba, and P. microglossa. Allof these taxa are quite similar in achene, pappus, and most other floral characters, but P. micro- cephala is clearly delimited from P. turneri and P. lineari- loba by its capitulescence of small, tightly clustered heads, grayish pubescence, and is easily separated from P. micro- glossa by its pubescence and perennial habit. Overall morphological similarity and distributional considerations (Fig. 3) suggest that P. microcephala is most closely re- lated to P. microglossa. Pringle 764 and Gentry 630, here ascribed to P. microcephala, approach P. microglossa in vegetative characteristics. 9. Perityle microglossa Benth. Bot. Sulph. 119. 1844. TYPE: Nicaragua (?): Realejo, Nicaragua (?), 1837, R. B. Hinds (K!). 9a. Perityle microglossa var. microglossa Perityle acmella Harv. & Gray, Mem. Amer. Acad. II. 4: 77. 1849. TYPE: Mexico: “California”, Coulter 278 (Holo- type, GH!; isotypes, NY!). Galinsogeopsis spilanthoides Schultz-Bip. in Seem, Bot. Herald 307. 1856. TYPE: Mexico: SINALOA: Sierra Madre, 1849, B. Seeman 1982 (Lectotype, K!). LECTOPARATYPE: same data, B. Seeman 1981 (K). Pericome spilanthoides Benth. & Hook. ex. Hemsl. Biol. Centr. Amer. Bot. 2: 215. 1881. Perityle microglossa var. effusa A. Gray, Syn. Fl. N. 1974] Perityle — Powell 271 Amer. 1: 332. 1884. TYPE: Arizona: Santa Catalina Mts., 5 June 1882, C. G. Pringle (Holotype, GH!; fragment, us!; isotype NY!). Perityle effusa Rose, Contr. U.S. Nat. Herb. 1: 104. 1891. TYPE: Mexico: SONORA: Alamos, 26 Mar.-8 Apr. 1890, E. Palmer 350 (Lectotype, US!; isolectotypes GH !, NY !). LECTO- PARATYPE: same data, E. Palmer 377 (US!) ; isolectopara- types, (GH, NY !). Perityle spilanthoides (Schultz-Bip.) Rydb. N. Amer. FI. 34: 17. 1914. Plants weedy, herbaceous annuals, 20-60 cm high, erect or decumbent, mostly branching above the base, stems few or many, upper portions, especially peduncles, copiously glandular-puberulent, rarely merely puberulent with few glandular hairs; leaves opposite or alternate, mostly oppo- site, puberulent, glandular-puberulent or glabrous, often turning purplish, (2.0) 3.0-7.0 (10.0) em long, (1.5) 2.0- 5.0 (9.0) em wide, extremely variable in size and shape, cordate, ovate, broadly ovate-cordate to subreniform, or subdeltoid, the margins merely singly or doubly crenate- even or irregular, to strongly 3-lobed or cleft, pedately divided, or subhastate; petioles 0.5-3.0 (4.0) em long; capit- ulescence of 1-3 or many heads clustered on relatively short peduncles 0.6-3.5 (6.0) em long; heads radiate, 3.5- 4.5 (6.0) mm high, 4.0-6.5 (7.0) mm wide, involucres cam- panulate; ray flowers 6-12, ligules white, 1.5-3.5 mm long, oblong; disc corollas yellow, 1.2-2.2 mm long, throats tubu- lar-funnelform; achenes 1.5-2.0 mm long, linear-oblong to linear-elliptic, rarely very narrow and short obovate, with thin to prominent callous margins, the margins ciliate, the faces glabrous or puberulent; pappus of a crown of squa- mellae and 2 unequal bristles, the longest 0.8-1.2 mm long; chromosome numbers, n = 34, 51. Rather widespread weed of northwestern Mexico, most common in Sonora, Sinaloa, and Baja California Sur, rang- ing south to Colima, seemingly introduced in northeastern Mexico and extreme south Texas. Flowering probably year around. (Fig. 3). 272 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS: MEXICO: Baja California Sur. San Jose del Cabo, Anthony 326 (CAS, DS, GH, POM, UC, US) ; ca. 6 km NW of Mira Flores, Carter 2663 (ns, UC, US) ; Arroyo de los Pozos, Moran 6886 (sp); La Paz, Palmer 92 (CAS, GH, NY, US); Boca de la Sierra, Powell and Sikes 1672 (SRSC, TEX); 1 mi N of Pescadero, Powell and Sikes 1676 (SRSC, TEX); vicinity of San Jose del Cabo, Wiggins 5689 (DS, NY, UC, US). Chihuahua. Tres Hermanos, SW of Batopilas, Hewitt 36 (GH). Colima. Colima, Oreutt 4551 (ps). Nayarit. Acaponeta, Jones s.n. (PoM, UC); San Blas, Maltby 22 (us); Maria Magdalena Islands, Maltby 175 (Ny, US); Tepic, Palmer s.n. (us). San Luis Potosi. near waterfall at El Salto, King 3877 (NY, TEX, UC, us); from San Luis Potosi to Tampico, Palmer 1093 (GH, NY, US). Sinaloa. Culiacan, Brandegee s.n. (UC); ca. 64 mi S of Culiacan, Breedlove 1545 (bs); Imala, Gentry 5455 (ARIZ, DS, NY, UC); San Blas, Jones, s.n. (CAS, NY, POM, UC); Lodiego, Palmer 1614 (GH, NY); 56 mi NW of Mazatlan, Powell and Sikes 1682 (sRsc, TEX); Rio del Fuerte, near El Fuerte, Rose, Standley and Russell 13587 (NY, US); vicinity of Villa Union, Rose, Standley and Russell 13932 (NY, US); ca. 5 mi E of Costa Rica, Sikes and Babcock 192 (SRSC, TEX); 70 mi S of Mazatlan, Sikes and Babcock 206 (SRSC, TEX). Sonora. 15 mi NE of Alamitos, Rio San Miguel, Abrams 13357 (ps); Hermosillo, Brandegee s.n. (DS, GH); 12 mi W of Navojoa, Gentry 7950 (uc, US); Alamos, Palmer 673 (us); Rio Mayo, Navojoa, Sikes and Babcock 177 (SRSC, TEX); 2.4 mi N of Soyopa, Sikes and Babcock 157 (SRSC, TEX); 28 mi E of Navojoa, Sikes and Babcock 185 (SRSC, TEX). Zacatecas. near San Juan Capistrano, Rose 2427 (Ny, US). U.S.A.: TEXAS: Came- ron Co. Rio Hondo, Chandler 7007 (us); E of Rio Hondo, Clover 1742 (ARIZ, TEX); Laguna Atacosa Refuge, Fleetwood 6069 (TEX). Willacy Co. Sauz Ranch, Johnston 53, 280 (TEX). The type locality originally given for Perityle m icroglossa, Realejo, Nicaragua, is probably in error. The southern-most eollection I have seen was from Colima, Mexico (Fig. 3), and it seems likely that the species would have been col- lected again in Nicaragua or in intermediate localities if it were there. The type locality probably was San Blas, Naya- rit, which also was visited during the Voyage of the Sulphur and where P. microglossa is known to occur. According to Rogers MeVaugh (personal communication), the localities of many other Sulphur collections were mixed up in similar fashion. The variable nature of Perityle microglossa is evident 1974] Perityle — Powell 213 from the synonymy and from Everly's (1947) confused treatment of the taxon. The current studies have led to the recognition of two biological entities within P. microglossa. One is var. microglossa (Fig. 3), a widespread, weedy poly- ploid (n = 34, 51), and the other is var. saxosa (Fig. 3), a more restricted "native" diploid (n — 17). Greenhouse tests utilizing numerous collections of var. microglossa have shown that it is self-compatible and that it reproduces prolifically by seed. Only a few such tests have been conducted with var. saxosa, but all have indicated self-in- compatibility for the diploid taxon. The morphological char- acters which distinguish the two varieties are not entirely consistent, and several morphological intermediates, for which chromosome numbers are not known, are found among existing collections. Variety microglossa typically exhibits shorter ligules, glandular-pubescent and shorter peduncles, tighter capitulescences, and a weedy habit, as compared to var. saxosa that usually displays longer ligules, rather smooth and longer peduncles, more lax capitules- cences, and a “native” habit. Morphologically, Perityle microglossa (particularly var. saxosa) is close to P. microcephala. Considering the evi- dence presented for these two species, it can be postulated that P. microglossa evolved from the montain-dwelling P. microcephala through derivation of annual habit and adap- tation to soil habitats. Variety microglossa must have evolved in response to polyploidization and development of reproductive self-compatibility which allowed for its “weedy” proliferation. Additional studies of Perityle microglossa var. saxosa, as well as P. microcephala, are most desirable. If var. saxosa proves to be a “native,” diploid, and self-incompatible pro- genitor of var. microglossa, then perhaps specific status should be accorded the latter taxon. 9b. Perityle microglossa var. saxosa (Brandeg.) Powell, comb. nov. Perityle saxosa Brandeg. Zoe 5: 225. 1905. TYPE: Mexico: SINALOA: vicinity of Culiacan, Cerro Colorado, 5 Nov. 1904, 274 Rhodora [Vol. 76 Brandegee (Holotype, UC!; isotypes, GH!, POM!, US). Perityle urticifolia Rydb. N. Amer. Fl. 34: 15. 1914. TYPE: Mexico: SINALOA: Tepic, 5 Jan.-6 Feb. 1892, E. Palmer 1960 (Holotype, NY !; isotypes, GH !, UC!, Us!). Plants herbaceous annuals, possibly rarely weak peren- nials, 15-50 cm high, erect, upper stems and peduncles sparsely to densely puberulent, rarely moderately glandular- puberulent; leaves opposite or alternate, mostly opposite, glabrous to puberulent, rarely glandular-puberulent, 2-6 (8) cm long, (1.0) 1.5-3.5 (5.0) em wide; capitulescence usually open, of 1-3 heads on relatively long peduncles, 2-7 em long; ligules 3.5-4.5 mm long; dise corollas 1.7-2.0 mm long; achenes 1.0-1.6 mm long; pappus bristles 2, unequal, the longest 1.0-2.0 mm long; chromosome numbers n = 17, 18. Mostly in foothills and lower elevations, both sides of the Sierra Madre Occidental, perhaps most common in Chihua- hua, Sonora, and Sinaloa, Flowering probably year around. (Fig. 3). REPRESENTATIVE SPECIMENS: MEXICO: Chihuahua. Temores, Flyr 60 (TEX); near Descanso, Flyr 77 (TEX); Palmer 238 (GH, NY, Us). Durango. Tamazula, Gentry 5234 (ARIZ, DS, GH, NY, UC). Nayarit, 4 mi E of Jalcocotan on road to Tepic, MeVaugh 12147 (Us). Sinaloa. 68 mi N of Ma- zatlan, Flyr 128 (TEX); 4 mi S of La Cruz junction, on MEX 15, Flyr 122 (TEX); Cerro Colorado, Gentry 5468 (ARIZ, DS, NY, UC, US); Canyon de Tarahumare, Sierra Surotato, Gentry 7315 (GH, NY, US); road from Las Flechas to La Rastra, Goldman 323 (GH, NY, US); 30.4 mi E of Villa Union, Powell and Turner 1853 (SRSC, TEX). Sonora. just S of Jecori, Drouet, Richards, and Lockhart 3692 (ps, GH, NY); 18 mi NW of Campas, Drouet, Richards, and Lockhart 3704 (Ds, GH); San Bernardo, Rio Mayo, Gentry 1255 (ARIZ, GH, UC); near Alamos, Gentry 4825 (ARIZ, DS, GH, NY); 12 mi SE of Colonia Oaxaca, Hastings and Turner 65-43 (ARIZ, DS, SD); Petaquilla Canyon, White 3320 (ARIZ, GH); La Vega Azul, SW of Cononia Morelos, White 4852 (ARIZ, GH, NY, US); 35 mi NE of Cajeme, on road to Tesopaco, Wiggins 6408 (Ds, US). The Hastings and Turner and the White collections from NE Sonora are tentatively assigned to var. saxosa, but their 1974] Perityle — Powell 275 somewhat doubtful affinity with this taxon should be noted. The specimens have elongated achenes, approaching the fruit characteristics of Perityle ciliata and P. coronopofolia, and they are perennial. In vegatative features they are more like P. microglossa. Perhaps the unusual collections are from a relict population which is transitional between the shorter-achened (e.g., P. microglossa) and the longer- achened (e.g., P. ciliata) members of the white-rayed al- liance. Again, however, as is the case with most of the poorly collected Sierra Madre taxa, further study is re- quired. 10. Perityle canescens Everly, Contrib. Dudley Herb. 3: 393. 1949. TvPE: Mexico: SINALOA: Capadero, Sierra Tacuicha- mona, 12 Feb. 1940, H. S. Gentry 5588 (Holotype, DS!; iso- types ARIZ!, GH !, NY !, UC], Us!). Plants low, suffruticose perennials, 6-15 cm high, densely leafed upper stems pilose; leaves opposite or alternate, pilose-hirsute to villous, 1.2-1.7 em long, 0.5-0.9 cm wide, 3-pinnatifid with linear segments, ovate-cordate in outline, crisped; petioles 5-9 mm long; capitulescence essentially of solitary heads on short peduncles; heads radiate, 5-7 mm high, 6-8 mm wide, involueres broad campanulate; ray flowers ca. 10, ligules white, 2.5-3.5 mm long, oblong to broadly so; disc corollas yellow, becoming purple-tinged 2.5.3.0 ram long, throats broad tubular; achenes 2.0-3.5 mm long, oblong-oblanceolate and nearly truncate on both ends, with thick callous margins, the margins merely puberulent, faces evenly puberulent; pappus of a prominent crown of united, erose squamellae, and 2 very unequal, rather stout, bristles, the longest 1.5-2.5 mm long; chromosome number, unknown. Known only from the type collection. Perityle canescens is related to P. coronopifolia from which it is distinguished by habit; dense indument; tend- ency for solitary heads; leaf morphology ; thickly calloused achenes with short-pubescent margins; strong, more or less 276 Rhodora [Vol. 76 united crown of pappus squamellae; stout pappus bristles: and distribution. Two other species, Perityle ciliata and P. hofmeisteria are grouped with the above taxa in the white-rayed alliance, largely on the basis of their similar floral characteristics (particularly achene and pappus). In habit and achene shape, P. canescens resembles P. lemmoni (sect. Laphamia: Powell, 1973), but the former taxon obviously belongs with sect. Perityle on the basis of its pappus crown. 11. Perityle ciliata (L. H. Dewey) Rydb. N. Amer. Fl. 34: 17. 1914. Laphamia ciliata L. H. Dewey, Bot. Gaz. 20: 425. 1895. TYPE: Arizona: rocks along Pine Creek near Pine, Ariz., 26 Aug. 1891, D. T. MacDougal 676 (Holotype, US!; iso- type, US!). Plants suffruticose perennials, 15-30 cm high, many- stemmed, erect to pendulous, densely short-pubescent above; leaves opposite or alternate, densely gray-pubescent under- neath, pubescent to puberulent on upper surfaces, 0.8-2.0 (3.0) em long, 0.6-1.3 (2.4) cm wide, deltoid-ovate to ovate- rhombic, the marigns serrate, shallow-lobed, or serrate- crenate; capitulescence of several heads clustered on short peduncles; heads radiate, 5-7 mm high, 5.5-7.0 mm wide, involucres campanulate; ray flowers ca. 6-10, ligules white, 3-6 mm long, broadly oblong to oblong-elliptic; dise corollas yellow, often purple tinged, 2.0-2.5 (3.0) mm long, throats tubular to tubular-funnelform; achenes 2.0-2.8 mm long, linear-oblong, with prominent callous margins, the margins long-ciliate, the faces slightly angled and pubescent; pappus of a crown of squamellae and 2 (rarely 3) slender bristles, 1.5-2.0 (2.5) mm long; chromosome number, n — 17. Restricted in distribution to the mountains of central Arizona; rock-dwelling. Flowering spring, summer, and fall. (Fig. 3). REPRESENTATIVE SPECIMENS: Arizona: APACHE C0.: Springerville, MeGinnies (ARIZ). COCONINO CO.: Long Valley, Coconino Natl. Forest, Coconino Rec. Crew 492 1974] Perityle — Powell 277 (us); Oak Creek, W of Troutdale, Goodding 198-47 (ARIZ, SMU); 4 mi W of Strawberry, Sikes 99 (SRSC, TEX). GILA CO.: Barnhart Pass, Matzatzal Mts., Collom 102 (GH, NY, US); Fossil Creek Hill, Collom 594 (us); Parker Creek Canyon, Sierra Ancha Mts., Gould 8632 (ARIZ, CAS, UC); Workman Creek Falls, Sierra Ancha, Johnson (ARIZ); Peterson Ranch, Sierra Ancha, Pase 1306 (ARIZ); 5.8 mi W of Strawberry, Sikes and Patterson 427 (SRSC, TEX). MOHAVE CO.: Hualapai Mts., Braem (ns); trail to Potato Patch, Hualapai Mts., Braem 875 (Ds). YAVAPAI CO.: near Senator Mine, along road from Prescott, Eastwood 16721 (CAS); between Prescott and Ash Fork, Eastwood 16770 (cas); Groom Creek, mts. near Prescott, Kearney and Peebles 9750 (ARIZ, uc); near Granite Dells, vicinity of Prescott, Kearney and Peebles 12780 (ARIZ, GH, NY, us); Prescott, Peebles, Harrison, and Kearney 2645 (ARIZ, US). This species is clearly related to P. coronopifolia from which it is conveniently delimited by leaf shape. 12. Perityle coronopifolia A. Gray, Pl. Wright. 2: 82. 1853. TYPE: New Mexico: GRANT CO.: copper mines of Santa Rita del Cobre, Sept., 1851, C. Wright 1196 ( Holotype, GH!; isotypes, NY!, US!). Laphamia coronopifolia (A. Gray) Hemsl. Biol. Centr. Amer. Bot. 2: 210. 1881. Laphamia scopulorum M. E. Jones, Contr. West. Bot. 12: 48. 1908. TYPE: Mexico: CHIHUAHUA: Colonia Juarez, 6000 ft., 12 Sept. 1903, M. E. Jones (Holotype, POM!; fragment, Us!). Plants low, suffruticose perennials, 6-36 cm high, erect or pendulous, many-stemmed, very leafy, grayish-pubescent ; leaves opposite or alternate, puberulent to densely so, or nearly glabrous, 0.7-2.5 (3.5) cm long, 0.5-1.2 (2.0) cm wide, 3-palmate and lobed with slender-spathulate seg- ments; pedately divided, or delicately 2,3-pinnatifid with linear-filiform segments; petioles 2-8 (12) mm long; capi- tulescence of several heads clustered on short peduncles; heads radiate, 5.0-6.5 mm high, 5-6 mm wide, involucres campanulate; ray flowers ca. 8-12, ligules white, 3-7 mm long, broadly oblong, oblong-elliptic, to subspathulate; disc corolas yellow, often purple tinged, 2.0-2.8 mm long, throats tubular, tubular-funnelform, or tubular-campanu- 278 Rhodora [Vol. 76 late; achenes 1.8-2.5 mm long, linear-oblong to narrowly oblanceolate, with thin or prominent callous margins, the margins typically long-white ciliate, rarely short ciliate, the faces slightly angled and glabrous or pubescent; pappus of a crown of squamellae and 2 (rarely 3) slender bristles, 1.5-2.5 mm long; chromosome number, n = 17. Rather widsepread and common in southern New Mexico- Arizona, and rare in northern Chihuahua, Mexico ; rock dwelling. Flowering spring, summer, and fall. (Fig. 3). REPRESENTATIVE SPECIMENS: MEXICO: Chihuahua. near Colonia Juarez, 6000 ft, Sierra Madres, Jones (POM, US). U.S.A.: Arizona: COCHISE co.: Miller Canyon, Hauchuca Mts., Barneby 5172 (CAS, NY); Ramsey Canyon, Huachuca Mts., Goodding 758 (GH, NY, RSA). GRAHAM co.: Fry Canyon, Pinaleno Mts., Shreve 1365 (ARIZ); Graham Mt., Thronber and Shreve 7783 (ARIZ). GREEN- LEE CO.: Mts. back of Clifton, Greene (NY); San Francisco Mts., Greene (POM); 0.5 mi N of Metcalf, Maguire, Richards and Moeller 11822 (ARIZ, GH, NY, US). PIMA CO.: Spud Ranch, Rincon Mts., Blumer 3331 (ARIZ, DS, GH, UC) ; north slope Baboquivari Peak, Clark 12565 (GH); San Pedro Vista, Santa Catalina Mts, Parker 8096 (ARIZ, NY, RSA, US); below Goose-Head Rock, Mt. Lemmon, Sikes 95 (SRSC). SANTA CRUZ co.: Flux Canyon, Patagonia Mts., Kearney and Peebles 10193 (ARIZ, US); Nogales to Ruby, Kearney and Peebles 14961 (ARIZ, GH, NY). New Mexico: DOÑA ANA CO0.: W side, Organ Mts., Dunn 7274 (NMC, RSA); E peaks, Organ Mts., Powell 1393 (SRSC); Dripping Springs Canyon, Organ Mts., Sikes 108 (SRSC). OTERO CO.: Dry Canyon, Rehn and Viereck (US). SOCORRO co.: Mogollon Creek, Mogollon Mts., Metcalf 323 (NMC, NY, UC, US); Gila Hot Springs, Mogollon Mts., Metcalf 828 (POM, UC, US). Perityle coronopifolia is closely related to P. ciliata, from which it is best distinguished by leaf Shape. The species also has affinity with P. canescens. Both P. coronopifolia and P. canescens have lobed leaves which are technically distinguishable, and they are further delimited by the long white-ciliate achene margins, clustered heads, pubescence, and distribution. The principal distribution of Perityle coronopifolia is in southern Arizona and New Mexico. The one known Mexican collection, originally described as Laphamia scopulorum, is 1974] Perityle — Powell 279 considered to be a morphological variant. The Jones speci- mens from near Colonia Juarez are both of poor quality but show delicate pinnatifid leaves, short-pubescent achene margins, and inconspicuous pappus squamellae. These two fruit characteristics indeed approach those of Laphamian taxa (Powell, 1973), but the strong indication of its rela- tionship with the Peritylean P. coronopifolia suggests its correct placement with the latter species. 13. Perityle hofmeisteria Rydb. N. Amer. Fl. 34: 18. 1914. TyPE: Mexico: DURANGO: vicinity of Durango, Apr.-Nov., 1896, E. Palmer 28 (Holotype, NY !; isotypes, GH !, US!). Plants suffrutescent to herbaceous perennials (possibly also annuals), 12-25 cm high, puberulent; leaves opposite or alternate, puberulent, 1.0-5.5 em long, 0.2-2.0 em wide, ovate to subspathulate and entire to shallow-lobed, or ovate- cordate and serrate to serrate-lobed ; petioles 0.5-2.0 (3.0) em long; capitulescence of 1-3 heads borne on relatively short peduncles, 1.5-2.8 cm long; heads radiate, 6-7 mm high, 4-6 mm wide, involucres turbinate-campanulate; ray flowers ca. 10-12, ligules white, 4-5 mm long, oblong; disc corollas yellow, 2.5-3.0 mm long, throats tubular-funnel- form; achenes 2.5-3.0 mm long, linear-oblong to narrowly obovate, with thin or thick callous margins, the margins prominently ciliate, the faces puberulent especially in the centers; pappus of a crown of squamellae and 2 subequal bristles, 2-3 mm long; chromosome number, n= 16 =+ 1. Seemingly endemic in Durango, Mexico; probably rock- dwelling. Flowering spring-fall. (Fig. 3). SPECIMEN EXAMINED: Mexico: DURANGO: 3 mi N of Rodeo, Flyr 332 (TEX). The Flyr 332 specimen, said by the collector to be *seem- ingly an annual," conforms to the types of Perityle hofmeis- teria in floral but not in vegetative features. The main dif- ferences are in habit appearance and leaf shape. The spe- cies is poorly understood at present, but its affinity is with 280 Rhodora [Vol. 76 P. ciliata and P. coronopifolia from which it is readily distinguished by leaf morphology and distribution (Fig. 3). 14. Perityle jaliscana A. Gray in S. Wats. Proc. Amer. Acad. 22: 431. 1887. TYPE: Mexico: JALISCO: Rio Blanco, ca. 10 mi NW of Guadalajara, 17-23 Sept. 1886, E. Palmer 554 (Holotype, GH ! ; isotypes, NY !, Us (2) !). Plants low, suffruticose perennials, 5-17 cm high, stems Spreading, densely puberulent; leaves opposite or alternate, densely puberulent to nearly glabrous, 2.0-3.5 em long, 0.8- 1.8 (2.0) em wide, deltoid, deltoid-rhombic, deltoid-ovate, or subhastate to subhalberd, usually 2-4 lobed proximally, rarely serrate-lobed; petioles 0.7-1.8 (1.7) em long; capit- ulescence of 1-several heads clustered on short, slender peduncles; heads discoid, 5.0-6.5 mm high, 4-5 mm wide, involucres funnelform-campanulate; disc corollas cream- white and purple-tinged, 2.0-2.5 (3.0) mm long, throats tubular-campanulate; achenes 1.8-2.2 (2.8) mm long, ob- long or nearly so, slightly tapering to the base, truncate, with prominent callous margins, rarely with 3-callous mar- £ins, angled on both faces, the margins and faces short- pubescent; pappus of a vestigial erown of squamellae, and 2 (3) slender bristles, 0.5-1.0 mm long, the bristles rarely flattened ; chromosome number, n = 17. Rock-dwelling endemic in Jalisco, Mexico. Flowering spring-fall. (Fig. 4). SPECIMENS EXAMINED: Mexico: JALISCO: ca. 40 km N of Guadalajara, road to San Cristo- bal de la Barranca, MeVaugh 22114 (NY, sD); Sierra San Esteban. near Guadalajara, Pringle 2352 (GH, UC, US) ; Sierra de San Esteban, Pringle 156834 ( ARIZ, CAS, LL, SMU, US); Rose and Painter 7476 (NY, US); 5 mi SW, by road, from San Cristobal de la Barranca, Sikes and Babcock 383 (SRSC, TEX). Although relatively few collections of Perityle jaliscana are known, it is the best-collected of four closely related endemic species of Jalisco, Mexico. The others, P. rosei, P. trichodonta, and the recently described P. feddemae, are known only from type collections. 1974] Perityle — Powell 281 jaliscana (open circles); P. rosei (open square); P. feddemae (closed Fig. 4. Distribution of P. (closed square); P. trichodonta circle). 282 Rhodora [Vol. 76 Perityle jaliscana and P. feddemae are the most distinct of the four species. I have followed Everly (1947) in recog- nizing P. rosei and P. trichodonta, even though future col- lections might indicate that they should be merged as a single species. Furthermore, both P. rosei and P. tricho- donta might simply be ecological variants of P. jaliscana. I recognize the four Species at this time mainly to preserve taxonomic consistency. In all three sections of the genus Perityle there are groups of closely related species that are no more distinctive than are the Jalisco endemics. In Jalisco, P. feddemae, with its dissected leaves, seems to be quite distinct, and it was recently discovered in a locality not far from the other species (Fig. 4). Strict endemism is common among the rock-dwelling taxa of Perityle, and lacking evidence to the contrary, I think it possible that the taxa in question are endemic species. Perityle jaliscana is most closely related to P. rosei. Most of its distinctive features, such as discoid heads, smaller, oblong achenes, shorter pappus bristles and squamellae, subhastate leaves, and short, dense pubescence, appear as if they could have been derived from P. rosei. 15. Perityle rosei Greenm. Proc, Amer. Acad. 40: 45. 1905. TYPE: Mexico: JALISCO: in the Sierra Madre W of Bolanos, 15-17 Sept. 1897, J. N. Rose 2947 (Holotype, Us!; isotypes, GH !, NY!). Plants suffruticose perennials, 10-20 em high, stems erect or spreading, densely short-pilose; leaves opposite or alter- nate, short-pilose to puberulent, 1.8-3.0 em long, 1.0-1.5 (2.0) em wide, deltoid to deltoid-ovate or deltoid-rhombic, rarely subhastate, the margins entire or nearly so; petioles 5-9 mm long; capitulescence of 1.-several heads on short peduncles; heads radiate, ca. 5 mm high, 4-5 mm wide, involucres campanulate; ray flowers ca. 13, ligules white, 2.0-2.5 mm long; dise corollas cream-white and purple- tinged, 2.0-2.3 mm long, throats tubular-campanulate ; achenes 2.0-2.5 mm long, narrowly obdeltoid, with promi- nent callous margins, slightly angled on both faces, the 1974] Perityle — Powell 283 margins and faces puberulent to subglabrous; pappus of a short crown of squamellae, more or less united, and 2 (3-4) slender bristles, 1.5-2.0 mm long, the 3-4 bristles, if present, reduced in length; chromosome number, unknown. Known only from the type collection. (Fig. 4). Perityle rosei is best delimited from P. jaliscana by its radiate heads, nearly obconical achenes, longer crown of pappus squamellae and longer bristles, nearly deltoid, sub- entire leaves, and longer, less dense pubescence. 16. Perityle trichodonta Blake, Proc. Biol. Soc. Wash. 37: 60. 1924. TYPE: Mexico: JALISCO: Sierra Madre W of Bo- lanos, 15-17 Sept. 1897, J. N. Rose 2978 (Holotype, us!; isotype, GH !). Plants low, suffruticose perennials, ca. 10 cm high, many- stemmed and spreading, densely gray-pubescent above; leaves mostly opposite, densely hirtellous-pilose and gray- ish, 7-9 mm long, 2.5-5.0 mm wide, ovate to ovate-rhombic and subhastate; petioles 2-4 mm long; capitulescence of 1-several heads clustered on short peduncles; heads radiate, ca. 4.5 mm high, ca. 4 mm wide, involucres funnelform- campanulate; ray flowers ca. 8, ligules white, 1.0-1.5 mm long; disc corollas cream-white and purple-tinged, 2.0-2.5 mm long, throats tubular-campanulate; achenes 1.5-2.2 mm long, narrowly obdeltoid, with prominent callous margins, slightly angled on both faces, the margins and faces puberu- lent to subglabrous; pappus of a short crown of squamellae, more or less united, and 2 slender bristles, 1.5-2.2 mm long, chromosome number, unknown. Known only from the type collection. (Fig. 4). Perityle trichodonta looks as though it could be a small- leafed, more densely pubescent, shorter liguled form of P. rosei, but it does differ in the characters listed. Future col- lections might reveal that P. trichodonta should be merged with P. rosei. 17. Perityle feddemae McVaugh, Contr. Univ. Mich. Herb. 284 Rhodora [Vol. 76 9: 431-433. 1972. TvPE: Mexico: JALISCO: Rancho Viejo, ca. 10 km ENE of Huejuquilla el Alto, rock-crevices, open oak-woodland, elev. 1900 m, 31: Oct. 1963, Feddema 2278 (Holotype, MICH ; isotype, SRSC!). Plants suffruticose perennials, 10-25 cm high, stems spreading, puberulent; leaves mostly opposite, puberulent, glandular-dotted, 1.7-3.5 em long including petioles, dis- sected-pinnatifid with linear-spatulate segments; capitu- lescence of 1-3 heads on short, slender peduncles; heads discoid, 5-6 mm high, 3-5 mm wide, involucres funnelform- campanulate; disc corollas pale yellow, 2.3-2.8 mm long, throats tubular-campanulate; achenes 2-3 mm long, oblong- oblanceolate, with prominent callous margins, angled on both surfaces, the margins and faces short-pubescent; pap- pus of a short crown of squamellae and 2 slender, unequal bristles, ca. 1.0 mm long; chromosome number, unknown. Known only from the type collection. (Fig. 4). This taxon possibly is most closely related to Perityle jaliscana that also has discoid heads, but its dissected leaves mark it as the most distinctive of the Jalisco endemics. 18. Perityle parryi Gray, Pl. Wright. 2: 106. 1853. TYPE: Texas: Canyon of the Rio Grande, Aug. 1852, C. C. Parry 521 (Isotype, NY!). Laphamia parryi (Gray) Benth. & Hook. ex Hemsl. Biol. Centr. Amer. Bot. 2: 210. 1881. Leptopharyna trisecta Rydb. N. Amer. Fl. 34: 23. 1914. TYPE: Mexico: CHIHUAHUA: dry cliffs, rocky hills near Chihuahua, 20 Mar. 1885, C. G. Pringle 183 (Holotype, NY l; isotypes, DS!, GH !, UC!, US!). Plants low, woody-based perennials in crevices of rocks or tall, spreading, herbaceous to suffruticose perennials in soil, 10-75 em high; leaves opposite or alternate, glandular- pubescent, 2-6 (9) cm long, 0.8-4.0 (6.0) cm wide, cordate to subreniform in outline, margins irregularly lobed, den- tate, or laciniate, but typically shallow to deeply 3-lobed, and in some specimens even subcruciform; petioles 1.2- 1974] Perityle — Powell 285 Fig. 5. Distribution of P. parryi (open triangles); P. vaseyi (closed triangles); P. aglossa (open squares); P. cordifolia (closed circles); P. leptoglossa (closed squares); P. lloydii (open star); P. lobata (closed stars) ; P. palmeri (open circles). 286 Rhodora [Vol. 76 4.0 em long; capitulescence of 1-3 heads borne on pe- duncles 1-7 cm long; heads radiate (rarely discoid), 0.8-1.0 em high, 0.7-1.4 em wide, involucres hemispherical to campa- nulate; bracts numerous, lanceolate to linear, rarely ob- lanceolate, acute to attenuate at the apex; ray flowers ca. 12-16 (rarely 1-6), ligules yellow, oblong, showy; disc flow- ers numerous, corollas yellow, (3.2) 4-6 mm long, throats tubular; achenes (2) 3-4 mm long, linear-elliptic to ob- conical-elliptic, with thin to prominent callous margins, the margins short-pubescent to dense-pilose, faces pubescent ; pappus of a crown of squamellae, and 1 (rarely 0 or 2) barbellate bristle, (1.0-2.5) 3-6 mm long, typically tapering to a point, the barbs typically antrorse throughout, rarely becoming lateral or retrorse toward the tip; chromosome number, n = 17. Western Big Bend area of Texas and adjacent Mexico; occurring in rocks and soils. Flowering essentially year around. (Fig. 5). REPRESENTATIVE SPECIMENS: MEXICO: Chihuahua. 11 mi NE of Camargo, Johnston 7920 (GH); road to San Carlos Mines, Johnston and Muller 39 (GH); 12 mi S of Ojinaga, Johnston and Muller 1443 (GH); near new lake on Rio Conchos, Powell, Turner, and Magill 2030 (SRSC, TEX); rocky hills near Chihuahua, Pringle 183 (Ds, GH, NY, UC, US); 1 mi E of km 100, Ojinaga-Camargo road, Sikes and Patterson 397 (SRSC, TEX) ; 20 km N of Chihuahua, Stewart and Johnston 2119 (GH). U.S.A.: Texas: Brewster Co. Lost Mine Peak, Chisos Mts, Correll and Johnston 24540 (LL); Green Valley Ranch, Correll and Wasshausen 27825 (LL); Agua Fria Spring, Cory 18638 (GH); Santa Helena Canyon, Cory 26475 (us); N side Whirlwind Spring area, 02 Ranch, Fletcher 1143 (srsc); Packsaddle Mt., McAfee 261 (SRSC); Chisos Mts., Muller 8247 (GH, NY, US, TEX); lower Window Trail, Big Bend Park, Powell and Sikes 1528 (sRSC, TEX); Basin, Chisos Mts., Sikes 32 (SRSC, TEX); Boot Springs Trail, Sikes 116 (SRSC, TEX); Nine- Point Mesa, Warnock and Johnston 17670 (srsC); Rosillos Mts., Warnock 20603 (sRsc). Presidio Co. S edge of Presidio, Correll and Johnston 21867 (LL); 10 mi SE of Redford, Correll and Johnston 21898 (LL); Casa Piedra road, 20 mi S of Marfa, Cox s.n. (SRSC) ; Capote Creek, Havard s.n. (US); near Cottonwood Springs, Sierra Tierra Vieja, Hinckley 1850 (GH, NY, US); ca. 9 mi N of Lajitas, Powell 1243 (sRSC) ; 5 mi S of Redford, Powell, Sikes, and Watson 1974] Perityle — Powell 287 1396 (sRsc); 12 mi S of Redford, Powell, Sikes, and Watson 1398 (SRSC); 3 mi E of Ruidosa, Powell and Brey 1513 (sRsc); Capote Creek, ca. 1 mi below Capote Falls, Powell 1518 (sRsC). The three yellow-flowered species of west Texas and ad- jacent Mexico (Fig. 5), Perityle parryi, P. vaseyi, and P. aglossa, are morphologically similar. Perityle parryi and P. vaseyi are particularly close and somewhat confusing taxonomically, primarily because of intermediate leaf forms and uncertain distributional integrity. Perityle parryi occurs as a rock-dwelling subshrub and as a soil-dwelling perennial, typically with leaves that are merely 3-lobed. Perityle vaseyi is a soil-dwelling, taprooted perennial, with deeply trisected leaves. However, some Mexican populations of P. parryi (— L. trisecta) have deeply trisected leaves, as do some soil-dwelling Texas forms, and it is not always possible to distinguish the taxa by leaf morphology. Perityle parryi is further delimited by long-tapering pappus bristles (one per achene) which are antrorsely barbellate, bracts usually lanceolate or linear and somewhat attenuate, and disc corollas with usually narrowly tubular throats, as opposed to P. vaseyi with shorter pappus bristles that are retrorsely barbellate at the tips, bracts usually oblanceolate and acute, and more broadly tubular dise corollas. It is suspected that the soil-dwelling forms of Perityle parryi and P. vaseyi may occasionally hybridize. Any hy- bridization, however, would be difficult to detect morpho- logically. It is not known whether the leaf variability of P. parryi is intrinsic or whether it might be the result of hybridization with P. vaseyi. In spite of marginal sympatry (Fig. 5), mixed populations of the two species have not been found, and no clear-cut hybrids have been detected. The subject is under experimental study. The populations of Perityle parryi near Ruidosa, Texas, in the Sierra Vieja and one collection 12 mi S of Ojinaga (Johnston and Muller 1443) are notably anomalous. Their pappus bristles are short and have retrorsely barbed tips, resembling those of P. vaseyi. In addition, the Sierra Vieja 288 Rhodora [Vol. 76 populations may be nearly rayless or entirely discoid. In other morphological characters the anomalous populations conform with P. parryi. The discoid forms should not be confused with P. aglossa, which is always rayless. 19. Perityle vaseyi Coult. Contr. U. S, Nat. Herb. 1: 42. 1890. TYPE: Texas: BREWSTER CO. Chisos Mts., 1889, G. C. Nealley (245?), (Holotype, Us!). Plants perennial with fleshy taproots, (10) 15-75 cm high, herbaceous to suffruticose, stems erect or spreading; leaves opposite or alternate, glandular-pubescent, 2.5-6.5 em long, 2-5 em wide, typically palmately divided into 3 lobes or subcruciform, with the lobes also dissected, cleft, or parted, rarely the leaves are strongly 3-lobed, cleft, or parted; petioles 1.4-3.5 cm long; capitulescence of 1-3 heads borne on peduncles 1-8 cm long; heads radiate, 0.8-1.0 cm high, 1.0-1.3 em wide, involucres hemispherical to campanu- late; bracts numerous, oblanceolate to linear-lanceolate, typically acute, less often alternate at the apex; ray flowers ca. 14-16, ligules yellow, oblong, showy; disc flowers numer- ous, corollas yellow, 3.2-5 (6) mm long, throats tubular to broadly so; achenes (2.0) 3.0-4.0 mm long, linear-elliptic to obconical-elliptic, with thin to prominent callous margins, the margins short-pubescent to dense pilose, faces pubes- cent; pappus of a crown of squamellae, and 1 (rarely 0 or 2) barbellate bristle, 1.5-4.0 mm long, typically truncate, the barbs retrorse or lateral toward the tip; chromosome number, n — 17. Desert soils, especially gypsiferous clay, southwestern Big Bend area of Texas and adjacent Mexico. Flowering essentially year around. (Fig. 5). REPRESENTATIVE SPECIMENS: MEXICO: Chihuahua. 6.5 mi S of Ojinaga, Powell, Turner, and Magill 2002 (sRsC, TEX); 5 mi S of Ojinaga, Powell, Turner, and Magill 2079 (sRsc, TEX). Coahuila. Picachos Colorados, Johnston and Muller 141 (GH). U.S.A.: Texas: BREWSTER CO. petrified forest, Cory 2706 (GH); Hot Springs, Cory 6477 (POM); Glenn Springs, Cory 44014 (TEX); Terlingua Creek on 02 Ranch, Fletcher 917 1974] Perityle — Powell 289 (SRSC); 2-8 mi E of Terlingua, Flyr 16 (SMU, SRSC, TEX); 8 mi E of Lajitas, Flyr 19 (SMU, TEX); 1 mi SE of Terlingua, Flyr 180 (SMU, TEX) ; 2 mi SE of Hen Egg Mt., Johnston and Warnock 3656 (SRSC); top third of Packsaddle Mt., McAfee 184 (snsc); 70 mi S of Alpine, Powell and Sikes 1530 (sRSC, TEX); 18 mi SE of Castolon, Powell, Averett, and Watson 1539 (sRsC, TEX) ; 0.4 mi N of Castolon, Powell, Averett, and Watson 1544 (SRSC, TEX); near Big Bend Tun- nel, Sikes 39 (SRSC, TEX) ; near hwy. 118 entrance to Big Bend Park, Sikes 40 (SRSC, TEX); 7 mi E of Castolon, Sikes and Averett 330 (SRSC, TEX); road to Fisk Canyon, Warnock 1148 (sRsc); Avary Canyon, Big Bend Park, Warnock 9139 (sRsC); Big Bend Park, Warnock and Wallmo 12211 (sRSC); San Vicente, near the Rio Grande, Young s.n. (TEX). As discused previously under Phylogenetic Considera- tions, an ancestral connection is hypothesized for Perityle vaseyi and its allies with the P. crassifolia group of Baja California. The primary evidence for this possible relation- ship are the similar habits and pappus bristles (with re- trorsely barbed tips) of P. vaseyi and P. crassifolia and the chromatographic profiles with yellow compounds, common to both groups. Morphologically, Perityle vaseyi and its relatives are closer to the P. cordifolia group of western Mexico. The distribution of the Texas species (Fig. 5) could have been achieved by migration from a Sierra Madre Occidental origin. The headwaters of the Rio Conchos are in the Sierra Madre near Creel, just across the divide from P. cordifolia country. The distribution of P. parryi near the Rio Conchos in eastern Chihuahua and the location of P. vaseyi and P. aglossa near the Rio Grande, suggest the possibility of eastern migration along the river systems. 20. Perityle aglossa Gray, Pl. Wright 2: 107. 1853. TYPE: Texas: Rio Grande below Presidio del Norte, Aug. 1852, C. C. Parry (Lectotype, GH !). Laphamia aglossa (Gray) Benth. & Hook. ex Hemsl. Biol. Centr. Amer. Bot. 2: 210. 1881. Leptopharynx aglossa (Gray) Rydb. N. Amer. Fl. 34: 24. 1914. Plants woody-stemmed perennials in rock crevices, 15-45 290 Rhodora [Vol. 76 cm high; leaves opposite or alternate, glandular-pubescent, 2.4 em long, 0.6-2.3 cm wide, ovate, subeordate or subdel- toid, irregularly lobed, laciniate, or serrate-dentate, often shallow-3-lobed; petioles 0.8-2.3 em long; capitulescence of 1-3 heads borne on peduncles 1.0-2.5 em long; heads discoid, 8-11 mm high, 4-8 mm wide, involucres campanulate; bracts linear to linear-lanceolate, attenuated at the apex; disc flowers numerous, corollas yellow, typically pink- or purple- tinged, 4.2-6.0 mm long, throats tubular; achenes (2.0) 2.5.3.2 mm long, oblong-obconical, with prominent callous margins, the margins short-pubescent to dense-pilose, faces pubescent; pappus of a crown of squamellae and 1 (rarely 2) barbellate bristle, 3.7-5.5 mm long, tapering to a point, the barbs antrorse throughout; chromosome number, n = 17. Restricted in southern Brewster Co. and Terrell Co., Texas, along and near the Rio Grande; crevices of lime- stone. Spring-fall. (Fig. 5). REPRESENTATIVE SPECIMENS: Texas: BREWSTER CO.: mouth of Regan Canyon on Rio Grande, Hinckley and Warnock 3737 (US, SRSC); near Boquillas, Marsh. 306 (GH); near Big Bend Tunnel, Sikes 38 (sRsC); 4 mi W of Hot Springs, Big Bend Park, Warnock and Turner 8291 (sRsC); San Vicente Canyon, Big Bend Park, Warnock and Parks 864 (LL, SMU, SRSC); limestone hills, Terlingua area, Warnock 13980 (SRSC) ; Heath Canyon, Black Gap Refuge, Warnock 20615 (sRSC); mouth of Regan Canyon, Warnock and Hinckley 461050 (SMU, SRSC, TEX). TERRELL CO.: mouth of San Francisco Canyon, Surratt 133 (SRSC) ; mouth of San Francisco Canyon, Warnock and Surratt 9817 (LL, SMU, SRSC) ; 9 mi E of Sanderson, Warnock 13155 (sRsC) ; along Rio Grande, between Regan Canyon and Sanderson Canyon, Warnock 15854 (sRsC). Although only one type specimen is cited for Perityle aglossa (Parry, Aug. 1852), I have seen another specimen which probably came from Parry's original collection. At NY a sheet labeled *P. aglossa, Arroyo San Carlos, April 1852," displays specimens of both P. aglossa and P. parryi. Judging from present-day distributions of the species, I believe that the specimens were probably mixed after 1974] Perityle — Powell 291 Parry's collections were made. Furthermore, the P. aglossa specimen on the NY sheet probably was from the original colleetion and perhaps should be recognized as an isolecto- type. The P. parryi specimen is discoid and probably came from the vicinity of Ruidosa above Presidio. Perityle aglossa is a limestone-dwelling subshrub with good distributional integrity. In addition to habit and dis- coid heads, the species is distinguished from P. parryi, its closest relative, by subtle characters of the leaves, involu- eral bracts, and disc corollas. 21. Perityle cordifolia (Rydb.) Blake, Contr. U. S. Nat. Herb. 23: 1602. 1962. Leptopharynx cordifolia Rydb. N. Amer. Fl. 34: 22. 1914. TYPE: Mexico: SINALOA: vicinity of Topolobampo, 23 Mar. 1910, J. N. Rose, P. C. Standley, and P. G. Russell 13294 (Holotype, NY!?; fragment of type, UC!; isotype, GH!). Plants suffrutescent perennials, rather low and spreading or semi-pendulent, 10-20 cm high, stems strong but herba- ceous except at the bases, pilose; leaves mostly alternate, bright green, pilose, 3.5-11.0 cm long, 1.5-9.0 cm wide, cor- date to broadly so, veins prominent, the margins doubly serrate-dentate, rarely crenate; petioles 1.5-4.5 mm long; capitulescence of solitary heads borne on rather stout pe- duncles, 2.5-6.0 em long; heads radiate, rarely cernuous, 10-12 mm high, 9-15 mm wide, involucres hemispherical] to campanulate, typically truncate at the base; bracts 10-13 mm long, linear and attenuate; ray flowers numerous, lig- ules yellow, 9-15 mm long, oblorg-elliptic; disc flowers numerous, corollas yellow, 5-7 mm long, throats tubular ; achenes 2.5-3.0 mm long, narrowly oblong, with thin or prominent callous margins, the margins typically densely ciliate, rarely puberulent, faces glabrous or puberulent in the center; pappus of a crown of squamellae, ca. 0.5 mm or less, and a single bristle, 4-7 mm long, the bristle tapering from base, a fine tip and densely barbellate; chromosome number, n = 17. 292 Rhodora [Vol. 76 A petrophilic, cliff-dwelling species, possibly also rooted in soils of shaded bluffs, Sonora and Sinaloa, Mexico. Flow- ering probably year around. (Fig. 5). REPRESENTATIVE SPECIMENS: Mexico: SINALOA: hill near Topolobampo, Flyr 63 (TEX); N of Topolobampo, Flyr 76 (TEX); road to Los Animas beach from Los Mochis, Flyr 105 (TEX); 23 mi N of Culiacan, Flyr 111B (TEX); Cerro Tecomate, W of Pericos, Gentry 5743 (ARIZ, DS, GH, NY, UC); ca. 1 mi E of Topolobampo, Sikes and Babcock 188 (SRSC, TEX); bluffs W of Topolobampo, Sikes and Babcock 190 (SRSC, TEX). SONORA: 32 mi N of Navojoa, Flyr 103 (TEx); Canyon Sapopa, Rio Mayo, Gentry 1282 (ARIZ, GH, NY, UC); Aquibiquichi, near Bata- cosa, Gentry 3003 (ARIZ, CAS, GH, UC, US); Cerro de Bayajori, 12 mi W of Navojoa, Gentry 7949 (uc, US); 23 km E of Navojoa, Moran 4032 (DS, sD, UC); 1-2 mi N of San Carlos Resort, Powell and Sikes 1689 (SRSC, TEX); 10 mi N of Guaymas, Randle and Werner 1 (ARIZ). Perityle cordifolia belongs with a group of four other morphologically similar species; P. palmeri, P. leptoglossa, and P. lloydii also of northwestern (mainland) Mexico, and P. lobata of Baja California. In spite of their similarities, the taxa are rather clearly delimited by vegetative features, excepting P. lloydii, which has characters of the other three mainland species. I have grown all of the taxa except P. lloydii under identical conditions, and each did maintain its characteristic morphology. Everly (1947) treated Perityle cordifolia as synonymous with P. palmeri. I recognize these taxa as related but cer- tainly distinct species. In fact, the soil-dwelling P. palmeri could have evolved from the rock-dwelling P. cordifolia, but on morphological grounds, P. palmeri shares even closer relationship with P. lobata. Perityle cordifolia is readily delimited from P. palmeri by its woody habit; bright green, pilose, cordate leaves with prominent veins and doubly ser- rate margins; long ligules; long attenuate involucral bracts; long pappus bristles; and long style branches. 22. Perityle leptoglossa Harv. & Gray in A. Gray, Mem. Amer. Acad. II. 4: 77. 1849. TYPE: Mexico: SONORA: “vi- cinity of Hermosillo,” 1829-1830, Coulter 277 (Holotype, GH!; fragment, Us!). 1974] Perityle — Powell 293 Leptopharynx leptoglossa (Harv. & Gray) Rydb. N. Amer. Fl]. 34: 22. 1914. Plants shrubby, spreading or erect, 10-60 cm high, stems densely grayish-puberulent; leaves mostly alternate, densely puberulent, gray-green, 3.0-5.5 em long, 1.3-3.5 cm wide, cordate to cordate-ovate, the margins shallow-lobed and crenate to serrate, or merely crenate-serrate; petioles 1.3- 2.5 cm long; capitulescence of solitary heads borne on rela- tively stout peduncles, 2-5 em long; heads radiate, rarely cernuous, 8-10 mm high, 7-12 mm wide, involucres cam- panulate-cylindroidal and truncate at the base; bracts 5-8 mm long, linear to linear-lanceolate and attenuate to sub- attenuate; ray flowers numerous, ligules yellow, 6-10 mm long, oblong to oblong-elliptic; disc flowers numerous, corol- las yellow, 4-5 mm long, throats tubular; achenes (2.2) 2.5-3.0 mm long, narrowly oblong, with thin or prominent callous margins, the margins long- or short-ciliate, the faces glabrous or minutely pubescent at the center; pappus of a crown of squamellae less than 0.5 mm long, and a single bristle, 2.5-4.5 mm long, the bristle tapering from base to apex and densely barbellate; chromosome number, n = 17. Strictly rock-dwelling, in the vicinity of Hermosillo, So- nora, Mexico. Flowering probably year around. (Fig. 5). REPRESENTATIVE SPECIMENS: Mexico: SONORA: 3 mi S of Magdalena, Aleramo 13216 (ps); rocky sides of El Cerro de la Campana, Drouet, Richards and Alva- rado 3387 (DS, GH, NY); 6 km N of Hermosillo, Drouet and Richards 3757 (ps); San Miguel de Horcasitas, Eisen (UC); 41 mi N of Hermosillo, Flyr 96 (TEX) ; 26 mi S of Hermosillo, Gentry 45'12 (UC) ; ca. 28 mi N of Hermosillo, Powell and Averett 1520 (SRSC, TEX) ; ca. 28 mi N of Hermosillo, Powell and Sikes 1683 (SRSC, TEX); near Hermosillo, Ripley 14309 (CAS) ; vicinity of Hermosillo, Rose, Standley, and Russell 12346 (NY, US); near Magdalena, Shreve (NY); near Estacion Torres, Shreve 6070 (ARIZ); 8 mi W of Hermosillo, Wiggins and Rollins 96 (ns, GH, NY); ca. 1 mi N of Torres, Wiggins 6266 (DS, GH, POM, US); 12 mi from Ures on road to Babiacari, Wiggins 7359 (DS, TEX, US). The most distinctive features of Perityle leptoglossa are: 294 Rhodora [Vol. 76 shrubby habit; leaf margins shallow-lobed or merely cre- nate-serrate; densely puberulent upper stems and leaves; small heads and short ligules; short bracts; achene margins typically short-ciliate; pappus bristles typically slender at bases. At one locality ca. 28 mi N of Hermosillo, I observed a few plants that were 214 ft. high, the largest woody plants in the genus. The species is related to P. palmeri and P. lloydii. 23. Perityle lloydii Rob. & Fern. Proc. Amer. Acad. 30: 118. 1894. TvPE: Mexico: SONORA. Badehuachi, 2 Dec. 1890, C. E. Lloyd 400 (Holotype, GH ! ; isotypes, NY !, UC!, US(3)!). Leptopharynz lloydii (Rob. & Fern.) Rydb. N. Amer. Fl. 34: 22. 1914. Plants semishrubby, erect, 10-40 cm high, stems woody to herbaceous, pilose; leaves mostly opposite, pilose, 2.5- 4.5 em long, 1.5-2.5 em wide, cordate to cordate-ovate, the margins doubly serrate; petioles 1.5-3.0 em long; capitu- lescence of solitary heads borne on peduncles 2.0-3.5 cm long; heads radiate, rarely cernuous, 8-9 mm high, 6-7 mm wide, involucres campanulate-cylindroidal and truncate at the base; bracts 5-7 mm long, linear and attenuate; ray flowers numerous, ligules yellow, 5-8 mm long, oblong to oblong-elliptic; disc flowers numerous, corollas yellow, ca. 4 mm long, throats tubular; achenes ca. 3 mm long, nar- rowly oblong, the callous margins short-pubescent; pappus of a minute crown of squamellae (ca. 0.2 mm long), and a single bristle ca. 3 mm long, the bristle slightly tapering from base to apex and densely barbellate; chromosome number unknown. Known only from the type collection. (Fig. 5). Perityle lloydii is poorly delimited from the other main- land taxa of the P. cordifolia species-group. In leaf shape, pubescence, and achene morphology (short-pubescent mar- gins and reduced pappus squamellae) P. lloydii resembles P. cordifolia. In shrubby habit, small heads, short ligules, and other floral characters, the taxon approaches P. lepto- 1974] Perityle — Powell 295 glossa. Its pubescence is like that of P. palmeri. Overall, P. lloydii possibly is closer to P. leptoglossa, and further collections might well provide evidence for merging the taxa. 24. Perityle lobata (Rydb.) I. M, Johnston, Proc. Calif. Acad. IV. 12: 1205. 1924. Leptopharynz lobata Rydb. N. Amer. Fl 34: 23. 1914. TYPE: Mexico: BAJA CALIFORNIA: Commondu, 19 Feb. 1889, T. S. Brandegee (Holotype, GH!; isotype, uc!). Plants fleshy-rooted perennials, possibly also annuals, low and spreading, semierect, or semipendulent, 10-40 cm high, stems weak, herbaceous, and rather brittle, usually purplish, glabrous to sparse-pilose ; leaves mostly alternate, bright green, turning purplish, glabrous to sparse-pilose, 3-8 (12) cm long, 1.8-6.0 (8) cm wide, cordate, cordate- orbiculate, or subreniform, strongly 3-lobed, cleft, or parted, rarely divided, the segments further shallow-lobed or the blades even 5-cleft, the margins dentate-lobed and acumi- nate, overall the blades appear regularly laciniate; capitu- lescence of solitary heads borne on weak peduncles 3-8 (15) em long; heads radiate, cernuous, 10-12 mm high, (5) 7-9 mm wide, involucres cylindroidal to campanulate, truncate at the base; bracts 5.0-9.5 mm long, linear-lanceolate to lanceolate-ovate, subattenuate, often purplish; ray flowers 8-12, ligules yellow, often turning purplish when dry, 2.5- 9.0 mm long, oblong-elliptic; disc flowers numerous, corollas yellow, often purple-tinged, 3.8-5.0 mm long, throats tubu- lar-funnelform; achenes (3) 3.5-5.0 mm long, narrowly oblong to oblong-elliptic, with prominent callous margins, the margins densely ciliate, faces glabrous to puberulent at the center; pappus of a crown of squamellae, 0.7-1.0 mm lonz, and a single bristle 3.5-6.0 mm long, the bristle very stout at the base, tapering to a point, densely barbellate ; chromosome number, n = 17. Restricted to Baja California Sur in the area of Sierra de la Giganta. Flowering year around. (Fig. 5). 296 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS: MEXICO: BAJA CALIFORNIA SUR. Arroyo Carrizal, E of Rancho El Horno, NE of San Javier, Carter and Ferris 3811 (UC, US) ; Arroyo Santo Domingo, Rancho El Horno, Carter and Ferris 3867 (DS, SD, UC); vicinity of Portezuelo E of La Victoria, Carter and Ferris 3911 (ns, UC); NW of Rancho El Horno, on steep escarpment of Mesa de San Alejo, W of San Javier, Carter and Sharsmith 4199 (Uc); Purisima, Gentry 4213 (ARIZ, Ds, GH, UC); Puerto Escondido, Johnston 4115 (CAS, GH, NY, UC, US); ca. 10 mi W of Loreto, Powell and Sikes 1663 (SRSC, TEX); ca. 10 mi S of Mission Dolores, Wiggins, Carter, Ernst 287 (ps, vs); arroyo Quisapol, E of La Presa, trail to Laguna Caquihui, Sierra de la Giganta, Wiggins 15544 (ps). Perityle lobata has perhaps been separated from main- land Mexico and its relatives there for about four million years. At least according to Moore and Buffington (1968), this is how long the Baja California peninsula has been drifting westward from the mainland. Perityle lobata is clearly related to P. palmeri. Distributional considerations suggest that the two present-day taxa once had a common range (Fig. 5), or else proximal distributions before the Sea of Cortez became a barrier of its present width. It is possible that Perityle lobata has been introduced to the mainland and has hybridized there, accounting for some of the variability in mainland taxa such as P. palmeri, In fact, it might be postulated that P. palmeri has evolved from P. lobata instead of the other way around as previ- ously assumed. However, if the latter conjecture were ac- curate, one might expect that P. palmeri would be more like P. lobata of Baja California. I have not found any evidence that P. lobata has been introduced recently to the mainland, As opposed to Perityle palmeri, P. lobata can be recog- nized by its leaf architecture, its purplish stems, pubescence, ligule length and number, bract width, achene length, and its stout bristles. 25. Perityle palmeri S. Wats. Proc, Amer. Acad. 24: 57. 1889. TYPE: Mexico: SONORA: Guaymas, Oct., 1887, E. Palmer 308 (Holotype, aH !; isotypes, NY!, uc!, us (3) !). 1974] Perityle — Powell 297 Leptopharynx palmeri (S. Wats.) Rydb. N. Amer. Fl. 34: 23. 1914. Plants fleshy-rooted perennials, low and spreading, semi- erect, or semipendulent, 10-25 cm high, stems herbaceous and rather brittle, often purplish, short or long pilose; leaves mostly alternate, short or long pilose, grayish-green, 6-10 em long, 3.0-5.5 cm wide, cordate, cordate-ovate, to cordate-orbiculate, the margins acutely shallow-lobed and serrate-dentate, the lobes several, or the blades strongly 3-lobed, overall the blades may appear irregularly laciniate; capitulescence of solitary heads borne on peduncles 2-6 (8) cm long; heads radiate, often cernuous at maturity, 8-11 mm high, 6-13 mm wide, involucres hemispherical to cam- panulate and truncate at the base; bracts 7-10 mm long, linear to linear-lanceolate and attenuate; ray flowers nu- merous, ligules yellow, 7-8 (10) mm long, oblong-elliptic ; disc flowers numerous, corollas yellow, 4-6 mm long, throats tubular; achenes 2.5-3.5 mm long, narrowly oblong, with prominent callous margins, the margins densely long-ciliate, faces glabrous or puberulent; pappus of a crown of squa- mellae, 0.5-0.9 mm long, and a single bristle, 3-5 mm long, the bristle stout, tapering from base to apex, and densely barbellate; chromosome number, n = 17. Primarily, if not entirely, soil-dwelling, most common in the vicinity of Guaymas, Sonora, but also in northern Sina- loa, Mexico. Flowering probably year around. (Fig. 5). REPRESENTATIVE SPECIMENS: Mexico: SINALOA: Topolobampo, Blakey (ARIZ.) SONORA: Guay- mas, Brandegee (POM, UC); Plam Canyon, W side of Sierra Libre, 35 mi S of Hermosillo, Carter (ARIZ); San Pedro Bay, Craig 691 (POM); Guaymas, Dawson 1003 (ps, US); cliffs, E side of Guaymas, Drouet and Richards 3846 (ps); 2.8 mi N of Guaymas, Lewis 5334 (sMU); 36 mi S of Hermosillo, Lindsay 1125 (ps); near Guaymas, Palmer 1218 (us); Sahuaral Bay, Powell and Averett 1523 (SRSC, TEX); 1 mi E of San Carlos Resort, Powell and Sikes 1687 (SRSC, TEX); 6 km N of Empalme, Ripley 14307 (CAS); 1 mi E of San Carlos Resort, Sikes and Babcock 169 (sRSC, TEX); 1 mi N of Naval Base, Guaymas Bay, Sikes and Babcock 176 (SRSC, TEX). 298 Rhodora [Vol. 76 Perityle palmeri is best distinguished from P. lobata, perhaps its closest relative, and from P. cordifolia by the following characters: fleshy roots, weak perennial habit in soils, lobing of the leaves, ligule length, bract length, and pilose pubescence. 26. Perityle californica Benth, Bot. Sulph. 23. 1844. TYPE: Mexico: BAJA CALIFORNIA: Magdalena Bay, 1837?, R. B. Hinds (K!). Perityle plumigera Harv. & Gray, Mem. Amer. Acad, II. 4: 77. 1849. TYPE: Mexico: SONORA: “California” (vicinity of Hermosillo), Coulter 279 (Holotype, GH!; fragment, uc!, us!). Perityle deltoidea S. Wats. Proc. Amer. Acad. 24: 57. 1889. TYPE: Mexico: BAJA CALIFORNIA: Los Angeles Bay, 1887, E. Palmer 568 (Holotype, GH!; isotypes, NY!, UC! US!), Plants delicate or robust annuals, 6-35 cm high, erect or decumbent, sparsely or densely leaved; leaves opposite or alternate, glandular-puberulent to short pilose, 1-6 (10) em long, 0.5-2.5 (6.0) cm wide, often purplish when mature, deltoid, ovate, or cordate, 3-lobed to subcruciform with the segments serrate-crenate or shallow-lobed, or the margins merely shallow-lobed to serrate; petioles 0.3-1.5 (4.0) em long; capitulescence of 1-5 heads borne on delicate pe- duncles; heads radiate, 3.5-7.5 mm high, 3.0-5.0 (7.5) mm wide, involucres narrowly to broadly campanulate; bracts linear, oblanceolate, or obovate; ray flowers 10-15, ligules yellow, oblong-elliptic; disc corollas yellow, 2.0-3.0 (3.5) mm long, throats tubular to subfunnelform; achenes 1.5-2.5 (3.0) mm long, narrowly obovate, oblanceolate, to oblong- oblanceolate, with thin or prominent callous margins, the margins short or long ciliate, faces puberulent to densely so; pappus of a crown of squamellae and a single bristle, 1.5-3.0 (3.5) mm long, the bristle barbellate to subplumose; chromosome numbers, n — 13, 12, 11. Common in desert soils, Sonora and Sinaloa of mainland 1974] Perityle — Powell 299 Mexico, and along most of the Baja California peninsula to just S of La Paz. Flowering Dec.-June. (Fig. 6). REPRESENTATIVE SPECIMENS: MEXICO: BAJA CALIFORNIA. San Juan Mine, Moran 8117 (DS, Sb, uc); Los Angeles Bay, Palmer 568 (GH, NY, UC, US); 4 mi S of Rancho Rosarito, Powell and Turner 1808 (sRSC, TEX); El Arco, N end of town, Sikes and Babcock 297 (SRSC, TEX); mesa S of Rancho Mesquital, Wiggins 11326 (DS, GH, UC). BAJA CALIFORNIA SUR. Isla Magdalena, Brandegee s.n. (GH, UC, US); 20 km E of San Ignacio, Carter, Alexander, and Kellogg 1972 (ps, UC, US); 26 km NW of San Ignacio, Carter, Alexander, and Kellogg 2521 (DS, GH, UC, US); Arroyo del Cajon de Tecomaja, SW of Puerto Escondido, Carter and Kellogg 2906 (uc); Rancho El Horno, NE of San Javier, Carter and Ferris 3789 (uc); SE of La Soledad, Carter 5418 (uc); Arroyo, 4 mi above La Purisima, Constance 3146 (DS, GH, LL, US); Isla San Jose, Moran 3785 (DS, SD, UC); Isla Santa Catalina, Moran 3867 (DS, sp); Isla San Marcos, Moran 8983 (sp, US); Isla. Carmen, Moran 9133 (sp); Isla Danzante, Moran 9253 (sp); Santa Rosalia, Palmer 185 (CAS,GH, US); ca. 19 mi N of La Paz, Porter 414 (CAS, Ds, Uc); 20 mi S of El Arco, Powell and Sikes 1650 (SRSC, TEX); 0.5 mi S of San Ignacio, Powell and Sikes 1653 (SRSC, TEX); ca. 8 mi S of Santa Rosalia, Powell and Sikes 1657 (SRSC, TEX); 33 mi S of Santa Rosalia, Powell and Sikes 1659 (SRSC, TEX); Bahia. de la Concepcion, Powell and Sikes 1661 (sRSC, TEX); 10 mi W of Loreto, Powell and Sikes 1664 (SRSC, TEX); 13.6 mi S of Mulege, Powell and Turner 1835 (SRSC, TEX); 11.5 mi W of Loreto, Powell and Turner 1837 (SRSC, TEX); 15 mi W of San Javier, Powell and Turner 1845 (SRSC, TEX) ; 3 mi E of La Paz, Sikes and Babcock 216 (SRSC, TEX); near km 40 on hwy between La Paz and Santo Domingo, Thomas 8455 (CAS) ; Isla Partida, Wiggins, Carter, and Ernst 412 (DS); Commondu, Wiggins 5502 (CAS, DS, GH, NY, UC, US); 22 mi N of San Ignacio, Wiggins 7888 (DS, GH, TEX, UC, US); 20 mi S of Mulege, Wiggins 11408 (CAs, ps, GH, UC, US); 6 mi W of Canipole, Wiggins 11489 (CAS, DS, GH, Uc, US); Llano Caquihui, W of Los Dolores, Wiggins 15516 (ps, US); 16 mi E of La Paz, Wiggins 15625 (Ds, us); Isla Carmen, Wiggins 17497 (ps). SINALOA. N of Topolobampo, Flyr 72 (TEX); vicinity of San Blas, Rose, Standley, and Russell 13244 (NY, US); vicinity of Fuerte, Rose, Standley and Russell 13472 (NY, us); W side of Topolobampo Harbor, Sikes and Babcock 189 (SRSC, TEX). SONORA. 8 mi N of Hermosillo, Breedlove 1758 (ns); San Pedro Bay, Craig 662 (PoM); E side of Guaymas, Drouet and Richards 3844 (CAS, DS, GH, US); 73 mi S of Hermosillo, Flyr 98A (TEX); 47 mi N of Hermosillo, Frye and Frye 2305 (GH, us); 1 mi E of San Carlos Resort, Powell and Sikes 1686 (SRSC, TEX); 1-2 mi 300 Rhodora [Vol. 76 Fig. 6. Distribution of P. californica (closed circles); the bars indicate populations from which chromosome numbers were taken, with (1) n — 11 II, (2) n = 12 II, (3) n — 13 II, and (4) n —10 II + 1 III; all mainland counts have been n= 13. P. cuneata var. cuneata (open circles) ; P. cuneata. var. marginata, (closed triangles). 1974] Perityle — Powell 301 N of San Carlos Resort, Powell and Sikes 1691 (SRSC, TEX) ; vicinity of Navojoa, Rose, Standley, and Russell 13140 (Ny, US); 110 mi E of Hermosillo, Sikes and Babcock 151 (SRSC, TEX); 32 mi N of Hermosillo, Shreve 10053 (LL); 10 mi N of Hermosillo, Wiggins 6250 (ns, POM, US); 5 mi SE of Torres, Wiggins 6273 (DS, GH, POM, US). In her discussion of P. californica, Everly (1947) em- phasized the variable habit of this taxon. The species oc- curs in Baja California and in Sinaloa-Sonora of mainland Mexico. Most of its morphological variability is expressed in plant size, number of leaves, and leaf shape. Floral char- acteristics are essentially the same throughout the distribu- tional range. Chromosomal variability also exists in Perityle californica where numbers of n — 13, 12, 11, and 10 II + 1 III have been reported (Powell, 1968; 1972b). Some distributional and morphological correlation with chromosome number has been observed, although no taxonomic status is war- ranted for the cytotypes. Plants of the mainland popula- tions are n = 13, as so far determined, and tend to be rather tall and robust with deeply lobed (often trilobed) leaves. In wet years P. californica is a common weed in some parts of Sonora and Sinaloa. Chromosome numbers of n = 13, 12, 11, and 10 II + 1 III have been found in Baja California populations (Fig. 6), and the plants are usually small with leaves not deeply lobed (mostly ser- rately-lobed). I have found the Baja California populations to be invariably small and scattered, but I have not seen the peninsula in a wet year. Populations with » — 11 are most common in Baja California and probably extend throughout the range of the species (Fig. 6). One popula- tion with n = 12 has been found, and those with n = 10 II + III presumably arose through hybridization of cyto- types with n = 12 and n = 11 (Fig. 6). The one peninsula collection with » — 13 (Fig. 6) could have been introduced from the mainland, or seemingly less likely at present, could represent the presence of native n — 13's on the peninsula. It seem reasonable to conclude that P. californica in Baja California evolved by aneuploid reduction in chromosome number from mainland cytotypes. 302 Rhodora [Vol. 76 Perityle californica does not share close or obvious rela- tionship with any species, and its origin as a low-numbered aneuploid is not clear. The species is superficially similar to P. cuneata (n = 16, 12), but these taxa differ consider- ably in floral characters, particularly of the achenes and pappus. Indeed, the floral] differences between these two aneuploid species are significant enough to cast considerable doubt upon their common origin. Perityle californica has narrow oblanceolate achenes with a single, long, subplumose pappus bristle, while P. cuneata has broad, obcordate or obovate achenes with two delicate pappus bristles. The morphology of P. californica (including flower color and achene and pappus features) resembles that of the P. cordi- folia group (x = 17), and possibly originated therefrom by aneuploid reduction. Certain aspects of the Perityle californica synonymy are somewhat confusing, but a good explanation of the histori- cal situations was given by Everly (1947). 27. Perityle cuneata Brandeg. Zoe 1: 54. 1890. TYPE: Mexico: BAJA CALIFORNIA: Sierra de Laguna, near Todos Santos, 21 Jan. 1890, T. S. Brandegee (Holotype, UC!; iso- type, US!). 27a. Perityle cuneata var. cuneata. Plants robust annuals, 25-50 (80) cm high, erect or de- cumbent, sparsely to densely leaved; leaves opposite or alternate, glandular-puberulent to nearly glabrous, 3-8 (15) em long, 1.5-4.5 (11.0) cm wide, often purplish when ma- ture, cordate to broadly so, ovate, or subhastate, the mar- gins serrate-lobed to serrate-crenate; petioles 0.6-2.0 (5.0) em long; capitulescence of 1-3 heads borne on rather stout peduncles; heads radiate, 5-8 (10) mm high, 6-10 (12) mm wide, involucres campanulate to hemispherical; bracts ob- lanceolate to obovate and attenuate; ray flowers 10-15, ligules yellow, oblong-elliptie; disc corollas yellow, 3.0-3.5 mm long, throats tubular; achenes (2.5) 3.0-3.8 mm long, obcordate-cuneate to oblanceolate, with broad callous mar- 1974] Perityle — Powell 303 gins, often purplish, the margins puberulent, faces glabrous to puberulent; pappus of a crown of squamellae and 2 (rarely 0-2) delicate bristles, 0.5-1.2 (2.0) mm long; chro- mosome number, n = 12. Infrequent in soils, Cape Region of Baja California Sur to just S of La Paz, most common on eastern side. Fower- ing Jan.-Sept. (Fig. 6). REPRESENTATIVE SPECIMENS: Mexico: BAJA CALIFORNIA SUR: San Jose del Cabo, Brandegee (DS, GH, US) ; Sierra San Francisquito and La Chuparosa, E side of Sierra de la Victoria, Carter and Ferris 3386 (DS, SD, UC); ca. 3 km above Boca de la Sierra, Moran 7091 (CAS, DS, GH, SD, TEX); Potrero de Almenta, S fork of Canyon San Pedro, Moran 7387 (CAS, DS, SD) ; ca. 5 mi S of San Antonio, Powell and Sikes 1667 (SRSC, TEX) ; Boca de la Sierra, Powell and Sikes 1671 (sRSC, TEX) ; 2 mi S of San Antonio, Sikes and Babcock 233 (SRSC, TEX); ca. 2 mi N of San Antonio, Sikes and Babcock 265 (sRSC, TEX); ca. 6 mi SW from Santiago, Thomas 7715 (CAS, DS, SD, US); ca. 5 mi SW of La Palma and ca. 8 mi NW from Santa Anita, Thomas 7738 (DS, SD, UC). The two varieties of Perityle cuneata are morphologi- cally similar, although extreme forms of these taxa do ex- hibit considerable differences. Everly (1947) recognized P. cuneata and P. cuneata, var. marginata as conspecific. In addition to the morphological differences, I have recog- nized var. cuneata, (n = 12) and var. marginata (n = 16) because of their chromosome numbers, Indeed, the chromo- somal differences suggest that further study might demand the elevation of var. marginata to specific rank. Besides its chromosome number, Perityle cuneata var. cuneata is characterized by robust habit, large cordate leaves, large heads, tubular disc corollas, obcordate-cuneate achenes, and attenuate involucral bracts. The plants of var. marginata are usually smaller with smaller leaves, smaller heads and other floral characters, campanulate-funnelform dise corollas, obovate achenes, and acute involucral bracts. In addition, var. cuneata usually occurs in higher hills and canyons of the Cape Region while var. marginata occurs mostly in sandy places not far above sea level. 304 Rhodora [Vol. 76 Perityle cuneata does not exhibit a close morphological relationship with any other species. Superficially, P. cu- neata most resembles P. californica, but it possibly evolved independently of the latter taxon by aneuploid reduction from the P. cordifolia line. 27b. Perityle cuneata var. marginata (Rydb.) I. M. Johns- ton, Proc. Calif. Acad. IV, 12: 1204. 1924. Perityle marginata Rydb. N. Amer. Fl. 34: 14. 1914. TYPE: Mexico: BAJA CALIFORNIA: San Jose del Cabo, 25 Mar. 1911, J. N. Rose 16459 (Holotype, NY!; fragment, UC!; isotypes, GH !, US!). Plants delicate or robust annuals, 6-40 cm high, erect or decumbent; leaves 1.5-5.0 (10) em long, 0.9-3.0 (5.0) cm wide, cordate, ovate, or deltoid, the margins doubly serrate- crenate to shallow-lobed; petioles 0.2-1.8 (4.0) em long; heads radiate, 4-7 mm high, 4-7 mm wide; bracts subovate, oblanceolate, or linear-lanceolate and acute; ligules yellow; dise corollas yellow, 2 (2.5) mm long, throats narrowly campanulate, campanulate-funnelform, rarely subtubular; achenes 1.5-2.0 (2.5) mm long, obovate, subcuneate, rarely almost oblong, with prominent callous margins, the margins short or long-ciliate; pappus of a crown of squamellae and 2 (rarely none) delicate bristles, 0.5-1.0 (1.5) mm long; chromosome number, n = 16. Infrequent in desert soils, Cape Region of Baja California Sur and N to near La Paz, most common on southern and western sides. Flowering probably year around. (Fig. 6). REPRESENTATIVE SPECIMENS: Mexico: BAJA CALIFORNIA SUR. San Jose del Cabo, Jones s.n. (NY, POM, SD, UC); Arroyo de los Pozos, Moran 6886 (CAS, DS, GH, TEX) ; 2.5 km NE of Cabo San Lucus, Moran 7049 (CAS, DS, GH, SD) ; 6.4 mi N of Todos Santos, Porter 79 (CAS, DS, UC); 1.4 mi W of El Coyote, Porter 118 (CAS, DS, UC); ca. 17 mi S of San Antonio, Powell and Sikes 1668 (SRSC, TEX); ca. 9 mi S of San Bartolo, Powell and Sikes 1670 (SRSC, TEX); 26 mi NE of Todos Santos, Powell and Sikes 1680 (SRSC, TEX); 2 mi NE of Eureka, Powell and Turner 1848 (SRSC, TEX); ca. 1 mi N of Pescadero, Sikes and Babcock 255 (SRSC, TEX) ; near km 40 between La Paz and Santo Domingo, Thomas 8455 (Ds, 1974] Perityle — Powell 305 GH); 5 mi W of San Jose del Cabo, Wiggins 14370 (CAS, DS, TEX, UC) ; 8 mi E of Cabo San Lucas, Wiggins 14645 (CAS, DS, GH, TEX, UC). The NY (holotype) and UC specimens of the type collec- tion (Rose 16459) are typical of var. marginata, while the US and GH specimens (isotypes) approach var. cuneata. It is possible that the type collection may be mixed, containing plants of both varieties. Some of the small forms of var. marginata are so like P. californica superficially that the two taxa can not be told apart except by microscopic examination of the achenes and pappus. The Thomas 8455 collection includes plants of both var. marginata and P. californica, but I have not otherwise noted sympatric occurrence of the taxa. No interspecific hybridization has been detected. LITERATURE CITED ALSTON, R. E., & B. L. TURNER. 1963. Natural hybridization among four species of Baptisia (Leguminosae). Am. Jour. Bot. 50: 159-173. EvERLY, M. L. 1947. A taxonomic study of the genus Perityle and related genera. Contrib. Dudley Herb. 3: 375-396. JOHNSTON, I. M. 1929. In: Papers on the Flora of Northern Chile. Contrib. Gray Herb. 85: 127-128. Moore, D. G., & E. C. BUFFINGTON. 1968. Transform faulting and growth of the Gulf of California since the late Pliocene. Science 161: 1238-1241. NiLES, W. E. 1970. Taxonomic investigations in the genera Perityle and Laphamia (Compositae). Mem. N. Y. Bot. Gard. 21: 1-82. PowELL, A. M. 1968a. Additional discussions pertaining to the con- generic status of Perityle and Laphamia (Compositae). Sida 3: 270-278. 1968b. Chromosome numbers in Perityle and related genera (Peritylinae — Compositae). Am. Jour. Bot. 55: 820-828. 1969. Taxonomy of Perityle section Pappothrix (Compositae — Peritylinae). Rhodora 71: 58-93. . 1970. Natural intersectional hybridization in Peri- tyle (Compositae). Brittonia 22: 3-10. 1972a. Taxonomy of Amauria (Compositae — Peri- tylinae). Madroño 21: 516-525. 1972b. Peritylinae, In IOPB Chromosome Number Reports. Taxon 21: 344. 306 Rhodora [Vol. 76 1972c. Artificial hybridizations in the subtribe Peri- tylinae (Compositae — Helenieae). Am. Jour. Bot. 59: 760-768. 1972d. Taxonomy of Pericome (Compositae — Peri- tylinae). Southwest. Nat. 18: 335-339. 1972e. New species of Perityle and Amauria (Com- positae). Madroño 21: 456-458. 1973. Taxonomy of Perityle section Laphamia (Compositae — Peritylinae). Sida 5: 61-128. , & S. Sikes. 1970. Chromosome numbers of some Chihuahuan Desert Compositae. Southwest. Nat. 15: 175-186. ,& H. S. Tsang. 1966. Preliminary chromatographic studies of Laphamia and related genera of Compositae. South- west. Nat. 11: 190-195. RAVEN, P. H. 1963. Amphitropical relationships in the floras of North and South America. Quart. Rev. Biol. 38: 151-177. RYDBERG, P. A. 1914. Carduaceae — Helenieae — Peritylinae. N. Am. Fl. 34: 11-27. SHINNERS, L. H. 1959. Species of Laphamia transferred to Perityle (Compositae — Helenieae). Southwest. Nat. 4: 204-209. SOUTHWICK, L., T. J. MABRY, J. AVERETT, & A. M. POWELL. 1972. Penduletin 4'-0-Methyl Ether, a new flavonoid from Perityle vaseyi (Compositae). Phytochemistry 11: 2351. STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press, New York. TURNER, B. L. 1966. Taxonomy of Eutetras (Compositae — Heleni- eae). Southwest. Nat. 11: 118-122. TURNER, B. L. & D. FLYR. 1966. Chromosome numbers in the Com- positae. X. North American species, Am. Jour. Bot. 53: 24-33. DEPARTMENT OF BIOLOGY SUL ROSS STATE UNIVERSITY ALPINE, TEXAS 7983 CAREX EXILIS, GEOCAULON LIVIDUM AND OTHER PLANTS OF INTEREST IN PITTSBURG, NEW HAMPSHIRE. South Bay Bog at the southern end of First Connecticut Lake in Pittsburg, Coos County, New Hamp- shire apparently was not visited by A. S. Pease, S. K. Harris and other botanists who collected extensively in Coós County. The bog is very extensive, much of it open but with scattered clumps of stunted Black Spruce and Tamarack. Lying between 1600 and 1700 feet elevation, it appears in every way to be boreal in character with the following species of shrubs in abundance, Ledwm groenlandicum, Rhododendron canadense, Andromeda glau- cophylla, Kalmia polifolia, K. angustifolia, Cassandra. caly- culata, Pyrus melanocarpa, Viburnum cassinoides and Vaccinium angustifolium. Bog Branch of Cedar Stream flows through or along the northern edge of South Bay Bog and near it, in sites where some drainage appears to run off the adjacent upland, there are scattered trees of Thuja occidentalis L., some up to 30 inches d.b.h. These represent remnants of the primeval forest since, along with dying trees of large size, there are fallen ones in all stages of decay. Several species of orchids grew in the immediate vicinity of trees of Arbor Vitae, the rarest of which was Listera convallarioides (Liv.) Nutt. A plant here of perhaps greater interest was Epilobium palustre L. var. palustre of which Seymour (1969) states “Often with Thuja,” but of which he cites only one locality in New Hampshire, “Mt. Washington." Pease (1964) states its habitat as “Boggy places, especially on the high moun- tains,” but he gives two localities for it at lower elevations both probably with some “Cedar” associated, “Stewarts- town, Cedar bog” and “Columbia, near Lime Pond.” In the more open part of South Bay Bog dominated by sphagnum moss and the previously mentioned shrubs and dwarf conifers there are plants of still greater interest to a taxonomist, one being Carex exilis Dewey and the other Geocaulon lividum (Richards.) Fern. Carex exilis is a sedge of local but wide distribution in New England, it being 307 308 Rhodora [Vol. 76 found occasionally in all States except Connecticut. Fer- nald (1950) omits mention of New Hampshire and Vermont in his statement of range for the species though he does mention Massachusetts, while Seymour (1969) gives but one township for it in New Hampshire, “Tamworth” where it was collected by F. L. Steele in a bog near Chocorua Lake and where he has shown it to me. Specimens of it are in the herbaria of the New England Botanical Club (NEBC) and the University of New Hampshire (NHA). However also at NHA are two other collections of fairly long stand- ing, the earlier being from Windham in Rockingham County collected on June 14, 1897 by W. S. Harris. This record was included in the list published by Hodgdon and Fried- lander (1949). The second collection of Carex exilis at NHA is from a bog in Pittsburg known as “Moose Pasture” lying along East Inlet about two miles northeast of Second Connecticut Lake. This was collected by me in company with Peter Allen on July 23, 1960 and bears my collection num- ber 11636. Pease (1964) did not include C. exilis in his "Flora" and it is quite possible that this specimen of it was not available for inspection at the time that Pease was gleaning additional records for Coós County and Harris was carefully checking the manuscript before publication. The specimens of C. exilis observed at South Bay Bog on our visit of June 8, 1973 were completely dioecious, the clones or colonies of female individuals being widely separated from the males. The other species of outstanding interest at South Bay Bog was Geocaulon lividum. I have been quite familiar with the Northern Comandra from the frequency of its occurrence in bogs on Campobello Island, New Brunswick and its presence on the Wolf Islands in the Bay of Fundy where R. B. Pike and I have observed it. It seems strange that a species inhabiting bogs at low elevations in coastal eastern Maine and the Maritime Provinces should be con- fined to montane areas inland though the same relationship seems to hold for Rubus chamaemorus L. and Lycopodium annotinum L. var. pungens (LaPyliae) Desv. Harris (1965) 1974] Pittsburg, N. H. Plants — Hodgdon 309 discussed the distribution of Geocaulon lividum in a critical way, pointing out that its habitat in stations that he visited was mostly damp but not boggy sites and that the lowest elevation for it was at 2500 ft. on the Basin Rim in Coos County. The station at South Bay Bog is in no way mon- tane and the elevation is between 1600 and 1700 feet. It is entirely possible that botanists have concentrated their collecting more in the mountains than in the more prosaic terrain of bogs that often lie between the mountains, thus passing by an interesting intermediate type of habitat for certain species that are supposed to occur only in coastal areas or near the tops of mountains. In any event it is evident that Geocaulon lividum should be searched for widely as Stuart Harris pointed out, not only as he sug- gested, in montane situations but also in bogs at inter- mediate elevations in Northern New England. Specimens of Carex exilis and Geocaulon lividum have been deposited in the herbaria of the New England Bo- tanical Club (NEBC) and the University of New Hampshire (NHA). These discoveries would not have been possible except for the expert guidance of Dr. Peter Allen. In addition I have borrowed freely in preparing this article from notes that he made or that we recorded jointly on the occasion of our visit to South Bay Bog on June 8, 1973. LITERATURE CITED FERNALD, M. L. 1950. Gray’s Manual of Botany, eighth edition, American Book Company, N. Y. 1682 p. Harris, S. K. 1965. Geocaulon lividum in New England. Rhodora 67: 407-409. HopGDoN, A. R. and HERBERT FRIEDLANDER. 1949. Additions to the Harris “Flora of Windham, New Hampshire.” Rhodora 51: 110. PEASE, A. S. 1964. A Flora of Northern New Hampshire, New England Botanical Club Inc. Cambridge, Mass. 278 p. SEYMOUR, F. C. 1969. The Flora of New England. The Charles Tuttle Co., Rutland, Vermont. 596 p. A. R. HODGDON DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE, DURHAM 03824 NEW STATION FOR NUPHAR MICROPHYLLUM IN DURHAM, NEW HAMPSHIRE. Pittsfield, N. H. (Mer- rimack County) is no longer the southernmost collection point in New Hampshire for Nuphar microphyllum (Pers.) Fern. as reported in my article *Some Morphological Aids in Distinguishing Nuphar microphyllum from Similar Aquatics” (Rhodora 75: 65-74, March 1973). On October 8, 1973 while A. R. Hodgdon and I were scouting the Lamprey River in Durham as a possible field trip site for his course in aquatic plants at the University of New Hampshire, I discovered a colony of N. microphyllum growing with N. variegatum and Nymphoides cordata in about two feet of water in a small cove near the upper end of Moat Island. Perhaps closer observation may reveal some plants of the hybrid, Nuphar X rubrodiscum. Undoubtedly other sta- tions for N. microphyllum exist but its relative rarity in southern New Hampshire has made it unnoticeable to the casual observer. The two specimens collected late in the season are a bit ragged and will be kept as vouchers in the NHA Herbarium. An attempt will be made to secure more presentable material for the New England Botanical Club Herbarium. JOHONET C. WICKS DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, N. H. 03824 310 TYPIFICATION OF CHAMAESARACHA CORONOPUS The type of Chamaesaracha coronopus Was erroneously listed as Berlandier 1494 (= 234) (Rhodora 75: 339). The section referring to the type material should read: Chamaesaracha coronopus (Dunal) Gray, Bot. Calif. 1: 540. 1876. Holotype (G!), [Texas, Bexar Co.], Rio Medina, between Laredo and Bejar [San Antonio]. 1828. Berlandier 1513 (= 253). Isotypes: GH! & K!. The locality indicated on the holotype and original de- scription is only between Laredo and San Antonio, then Bejar, Mexico. An isotype at Kew, however, has Rio Medina on the label. The Medina River flows southeast from Bandera County into Medina County, then east into Bexar County. The route taken by Berlandier likely erossed the Medina River in Bexar County, about 12 miles south of San Antonio. JOHN E. AVERETT DEPARTMENT OF BIOLOGY UNIVERSITY OF MISSOURI-ST. LOUIS RANGE EXTENSION OF WOODWARDIA AREO- LATA. While I was walking through a swampy area close to Spruce Hole Bog in Durham, N.H. on October 20, 1973, a fern frond came to my attention that from afar resembled an aberrant form of sensitive fern, Onoclea sensibilis. Upon further investigation, however, what was found turned out to be the fertile frond of Woodwardia areolata (L.) Moore, the netted-chain fern, so named for its long sori arranged in a row on each side of the secondary midrib. The discovery of this fern is noteworthy when one real- izes that only two stations for it have ever been recorded from New Hampshire, both in Rockingham County: Rye Beach by Walter Deane in 1886 (N.E.B.C.) and Seabrook by A. A. Eaton in 1896 (N.E.B.C.) In the seventy-seven years since those recorded findings, Woodwardia areolata has not been found in New Hampshire. In addition, this new station extends the range of this coastal plain species by approximately twenty miles. Shaded by two bushes of Vaccinium corymbosum, a total of six plants have been located within an area of three feet. The plants are growing on rotting stump matter and are surrounded most of the year by a foot of water or more. Such a find always leads one to speculate as to how this fern came to occur at this location, how long it has been there, and why it was not discovered long before this. A voucher specimen of the Woodwardia areolata is de- posited in the University of New Hampshire herbarium. AMINTA KITFIELD DEPT. OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 EDITORIAL ANNOUNCEMENT With this number the present editor-in-chief with twelve and a half years of service in that capacity, is completing his tenure. Also Dr. Radcliffe B. Pike, as a valued associate editor during this entire period, is completing his editorial connection. We trust that the younger group of editors who are now in charge will find the journal to be prospering. Many changes in Rhodora have been made during the past twelve years; we express the hope that the journal can continue to be molded to fit changing needs. The new editor-in-chief is Dr. Alfred Linn Bogle, Associate Professor of Botany at the University of New Hampshire. Please send all manuscripts and communica- tions to Dr. A. Linn Bogle, Department of Botany and Plant Pathology, University of New Hampshire, Durham, New Hampshire 03824. A. R. HODGDON DEPARTMENT OF BOTANY AND PLANT PATHOLOGY UNIVERSITY OF NEW HAMPSHIRE, DURHAM, N. H. 03824 313 INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA Manuscripts must be double-spaced or preferably triple- spaced (not on corrasable bond), and a list of legends for figures and maps provided on a separate page. Footnotes should be used sparingly, as they are usually not necessary. Do not indicate the style of type through the use of capitals or underscoring, partieularly in the citations of specimens, except that the names of species and genera may be under- lined to indicate italics in discussions. Specimen citations should be selected critically especially for common species of broad distribution. Systematic revisions and similar papers should be prepared in the format of “The System- aties and Ecology of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- larly with reference to the indentation of keys and syno- nyms. Papers of a floristic nature should follow, as far as possible, the format of “Contribution to the Fungus Flora of Northeastern North America. V.," H. E. Bigelow & M. E. Barr, Rhodora 71: 177-203. 1969. For bibliographie cita- tions, a recommended list of standard journal abbreviations is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300. 1958. Volume 76, No. s06, including pages 151-314, was issued July 3, 1924. CONTENTS: — continued New Station for Nuphar microphyllum in Durham, New Hampshire Johonet C. Wicks eese eee eene eene erano peonia ei sinit 310 Typification of Chamaesaracha coronopus John B: Averell .cictcocesdacsncsisccanscnectsiescscavencecusscesepsatstgseesteduccsese 811 Range Extension of Woodwardia areolata Aminta Kit field ............. 4. eee eere eese eese eee esee estan nost tatto a ah taa a 312 Editorial Announcement ................ eere eene eene nnne ennas 813 Instructions for Contributors .............. eene 314 ena Hovova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALFRED LINN BOGLE, Editor-in-Chief ROLLA MILTON TRYON STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY RICHARD EDWIN WEAVER Associate Editors Vol. 76 September, 1974 No. 807 CONTENTS: A Systematic Study of the Genus Kalmia (Ericaceae) TORN. E DIIP aoieanna aia E e 315 Marine Algae of Rhode Island R. D. Wood and Martine Villalard-Bohnsack .................. 399 A Biosystematic Revision of Lupinus lyallii 20) 08 OT es Ts canioniusnndctsvsensereaseesanssise A E 422 The Role of Some Haloragaceae in Algal Ecology L, C. Colt, Jr., and C. Barre Hellquist ................... 446 Illinois Field and Herbarium Studies Robert H. Mohlenbrock and Dan K. Evans ......... 460 (Continued on Inside Back Cover) Che Nef England Botanical Club, Ine. Botanical Museum. Oxford St.. Cambridge, Mass. 02138 RHODORA. — A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price $10.00 per year, net, postpaid, in funds payable at par in the United States currency at Boston. Some back volumes, and single copies are available. For information and prices write RHODORA at address given below. 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 submitted in duplicate, and 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. Ex- tracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Dept. of Botany, University of New Hampshire, Durham, New Hampshire 03824 Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to RHODORA, Botanical Museum, Oxford Street. Cambridge, Mass. 02138. In order to receive the next number of RHODORA, changes of address must be received prior to the first day of March, June, September or December. Second Class Postage Paid at Boston, Mass. MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 76 September, 1974 No. 807 A SYSTEMATIC STUDY OF THE GENUS KALMIA (ERICACEAE) JoHN E. EBINGER A study of the cytology and genetics of the genus Kalmia was started in 1961 by Dr. Richard A. Jaynes of the Con- necticut Agricultural Experiment Station. At that time little was known about the genetics of this group although a few of the species are important ornamental shrubs. Dr. Jaynes has now accumulated many living representatives of most of the species due to the efforts of numerous bota- nists and friends. As a result of his studies (Jaynes 1968a, 1968b, 1969, 1971b) information is now available concern- ing cross compatibility, speciation, and possible evolution- ary relationships within the genus. As treated here, the genus is composed of seven species, six varieties and numerous forms, both natural and culti- vated. The genus is restricted to North America and Cuba, being particularly abundant along the east and west coasts and at higher elevations and latitudes. No monographic study of the genus has been undertaken, but the species have been adequately described in many floristic works, and Small (1914) treated all of the species that are presently recognized. Since one of the species is commonly cultivated and all are probably poisonous, this taxon has also been reviewed in horticultural and agricultural literature. This present study is an attempt to bring together most of this 315 316 Rhodora [Vol. 76 information in the light of what is now know about the geneties and cytology of the genus. ACKNOWLEDGEMENTS The author is grateful to the many people who helped in the completion of this study. Dr. Richard A. Jaynes pro- vided access to his herbarium specimens and living material as well as his notes and extensive knowledge concerning the cytology and genetics of the genus. The Connecticut Agricultural Experiment Station provided space and equip- ment for 4 months to examine and collect material. Also, many individuals at Eastern Illinois University provided helpful criticism and advice in the preparation of the manu- script. The author is also indebted to the many curators who sent specimens for examination. Over 8,000 specimens were examined from 75 herbaria from the United States, Canada, and Europe. In the lists of representative speci- mens these herbaria are indicated by their abbreviations following Lanjouw and Stafleu (1964). HISTORY OF THE GENUS References to mountain laurel, Kalmia latifolia, appear in the early literature of the American colonies. The Cape Cod “Rose-trees” of Henry Hudson's log (Purchas, 1625) may have been this species (Holmes, 1956), and Smith (1624) mentions that laurel occurs in Virginia. This spe- cies was first described and figured by Plukenet (1696) under the name Cistus sempervirens, described by Grono- vius (1739) who called it “The Common Laurel/vulgarly called Ivy," while the first color print was. figured by Catesby (1743) under the name Chamaedaphne. Because of their potential as ornamentals, some of the species were introduced into Europe. According to Aiton (1789) Kalmia latifolia and K. angustifolia were intro- duced into England in 1734 and 1736 by Peter Collinson, an English merchant and fellow of the English and Swedish Royal Societies. A third species, K. polifolia, was intro- 1974] Genus Kalmia — Ebinger 317 duced in 1767. Catesby (1743) reported that K. latifolia from Pennsylvania flowered in his garden at Fulham in 1740. Numerous horticultural forms have been developed as a result of breeding and selection in Europe for the past 200 years. Probably the first detailed account of the toxic character- istics of the genus occur in Peter Kalm's journal (Benson, 1937) in which he described in detail the poisonous proper- ties of the “Laurel Trees" (Kalmia latifolia). He also described both K. angustifolia and K. latifolia as to their characteristics, economic importance, habitat requirements and general distribution. The generic name Kalmia was first proposed by Chenon (1751) in a doctoral thesis, and was based on specimens that Peter Kalm had collected during his trip to the New World. Kalmia latifolia and K. angustifolia are described in this thesis, and Linnaeus (1753) included both in his Species Plantarum. Later, Linnaeus (1754) described the genus and placed it in his first order (Monogynia) of his tenth class (Decandria). The genus is now regarded as a relatively primitive mem- ber of the Ericaceae. Drude ( 1889) placed it in the Tribe Phyllodoceae Drude of the subfamily Rhododendroideae Drude, and its position has not been changed by most subsequent authors. The members of this tribe are gen- erally evergreen plants, usually with buxoid or ericoid leaves, with regular flowers, and with seeds that usually lack appendages. Two microscopic characteristics of the anthers have also been used to characterize the tribe: the presence of resorption tissue involving the epidermis of the anther and the absence of filaments among the pollen tetrads. Copeland (1943) discussed in detail the morphol- ogy and anatomy of the tribe Phyllodoceae and considered the genera Rhodothamnus Reichenb., Kalmia, and Phyllo- doce Salisb. to be relatively primitive, an opinion also ex- pressed by Wood (1961). Studies of wood anatomy by Cox (1948) indicate that the genus Kalmia is closely related to 318 Rhodora [Vol. 76 Rhodothamnus and that these two genera represent one of the three lines of evolution in the Phyllodoceae. There have been few nomenclatorial complications in the genus, and only two other generic names have been pro- posed. Kuntze (1891) used Catesby's pre-Linnean name of Chamaedaphne and made wholesale transfers to it, while Small (1914) divided Kalmia into two genera. He proposed the genus Kalmiella for Kalmia hirsuta and its related species that differ by having a deciduous calyx and flowers borne singly in the axils of ordinary or somewhat reduced leaves. In all other morphological and anatomical character- istics these species do not differ from the other members of the genus, and there is no real justifieation for their removal. Except for Alain (1957), all subsequent authors have treated these species as members of the genus Kalmia. ECONOMIC IMPORTANCE OF KALMIA The genus Kalmia is economically important mainly be- cause a few of the species are commonly grown as orna- mentals. By far the most important species is the moun- tain laurel, Kalmia latifolia, which is commonly planted as an ornamental throughout its natural range as well as in other parts of North America and Europe where climate and soil are satisfactory. This species varies widely in flower color, foliage, and size, and many cultivars have been named and described (Holmes, 1956; Dudley, 1967). Culti- vars of mountain laurel had been named by 1840, and at the present time at least 90 names have been applied to different forms of the species. This species has been sug- gested as a possible choice for the national flower of the United States (Van Rensselaer, 1889), and presently it is the state flower of Connecticut and Pennsylvania. The sheep laurel, Kalmia angustifolia, is also becoming an important cultivated plant. Unlike the mountain laurel, this species can be propagated by cuttings. A number of cultivated forms are available, and selections should become available as soon as nurserymen become familiar with hand- 1974] Genus Kalmia — Ebinger 319 ling them. This species is not important for its wood as few specimens exceed 3 feet in height, but it is economically important as a weed. Smith, Hodgdon and Eggert (1947) considered it an important weed in low-bush blueberry fields and tried fire, continued cutting, and lime to control it. Later, Trevett (1956) reported on the effects of 2,4-D and 24,5-T in controlling sheep laurel in blueberry fields. Porcher (1869) found that the leaves and twigs of this species will dye cotton a fine drab color if a copper mordant is used. The foliage of Kalmia latifolia is commonly gathered in eastern United States for floral displays as well as for Christmas decorations and other festivities, continuing a tradition started in colonial days. As early as 1913, Britton suggested that this species should probably be protected since it was becoming rare in some areas due to indiscrimi- nate collecting. Buttrick (1924), in an extensive article concerning growth and cultivation of mountain laurel in Connecticut, estimated that the amount of foliage of this species used in New York City probably exceeded 1,000 tons per year and that the maximum amount used in the United States for decorative purposes probably exceeded 10,000 tons per year. It is still commonly found in decora- tions, but no figures are presently available concerning the amount used. The foliage of mountain laure] can also be used to dye cloth. Nicholson and Clovis (1967) found that the boiled leaves of this species produced a yellow tan color on wool if a chrome mordant was used. In the past, the wood of the mountain laurel was occa- sionally used to make small items. Peter Kalm in his journal (Benson, 1937) mentions that the strong wood is carved into weaver's shuttles, pulleys, spoons and trowels. Browne (1857) considered the wood to be similar to that of the European Box and that it was sometimes used for the handles of light tools, for screws, boxes, and that it was supposed to make good clarinets. He also mentioned that the American Indians used the wood for making small dishes, spoons, and other domestic utensils, and according 320 Rhodora [Vol. 76 to Benson (1937), this was the source of the common name "spoonwood." Other uses include the making of rustic furniture and as a substitute for brier pipes (Britton, 1913). Due to the small size of most mountain laurel shrubs, the wood is rarely used at the present time except locally for tool handles and novelties. Kalmia latifolia, as a common understory shrub, is im- portant in preventing water runoff and soil erosion. Studies by Johnson and Kovner (1956) in the southern Appalachian Mountains show that the cutting of dense stands of laurel and rhododendron in hardwood stands decreased the evapo- transpiration rate and correspondingly increased the annual streamflow on the average of 2 inches per year. Extracts from the leaves of most species of Kalmia were at one time considered important as medicine, Barton (1802, 1804) mentions that a saturated tincture prepared from the leaves of Kalmia latifolia in a proof spirit could be used for certain fevers. According to Rafinesque (1830) preparations of this species were used as a narcotic, errhine, antisyphilitic and antiherpetic, while Griffith (1847) con- sidered it useful for fever, itching and in cases of hemor- rhaging and dysentery. Wood and Bache (1851) considered it useful mainly for cutaneous affections when applied as an ointment or decoction. During the latter part of the 19th century the leaves of K. latifolia were still commonly found in drug stores. According to Paschkis (1881), they were dried, rolled into a pipe-form and used principally as a remedy for diarrhea. The leaves of K. hirsuta were used in the south as a cure for itching and for mange in dogs (Porcher, 1869), while Holmes (1884) reports that K. angustifolia was commonly used by the Cree Indians of the Hudson Bay region for bowel complaints and as a tonic. PROPAGATION OF KALMIA Numerous cultivars of Kalmia latifolia have been named; no large scale nursery production of any of these forms exists, however, because they have been considered difficult 1974] Genus Kalmia — Ebinger 321 to root, hard to graft, and are untrue from seed. As a result, one common source of this species for ornamental purposes has been native woodland stands. Recent studies, however, indicate that it is possible to root cuttings of this species and that selected types do breed true. According to Jaynes (1971d) most Kalmia species can be started from cuttings. He found that greenwood cuttings of Kalmia angustifolia, K. hirsuta, K. polifolia, and K. microphylla root readily under mist or a plastic tent. By contrast, K. cuneata cuttings were difficult to root, while K. latifolia was highly variable in its rooting ability. In this latter species the ability to root is related to the age of the stock plant, since 89% of the cuttings from 1-year-old plants rooted, while only 21% of those from 4 or more year-old plants rooted. Jaynes also reported that most of the sterile hybrids can be propagated readily from cuttings, even when K. latifolia is one of the parents. Jaynes (1972) found that clones of K. latifolia vary in their ability to root from cuttings and that cuttings of cuttings or young grafts root more readily than cuttings from the original stock plant. Few attempts have been made to graft any of the species of Kalmia. Trumpy (1893) had some success using side grafts with current year’s wood of Kalmia latifolia, while Jaynes (1971d, 1972) grafted current year’s growth of K. latifolia onto stock of the previous year using cleft or side-veneer grafts. The grafts were made in mid-June, kept in plastic tents for 6 weeks, hardened for 2 weeks, and overwintered in a lath house. The first report on the suc- cess of layering K. angustifolia was probably by Curtis (1795). The propagation of Kalmia from seeds has been exten- sively studied. Nichols (1934) found that Kalmia polifolia seeds germinated only after stratification, while K. angusti- folia, and K. latifolia seeds had a germination rate 5 times higher after stratification for 12 weeks. In contrast, Jaynes (1971c) reported that of these 3 species, only K. latifolia required stratification, the others germinating without any 322 Rhodora [Vol. 76 pregermination treatment. Jaynes also found that K. cu- neata, as well as K. latifolia, required stratification, but that soaking the seeds in gibberellin for 24 hours created the same response as stratification. The most difficult species to propagate appears to be K. hirsuta, which germinated best after a heat-humidity treatment (heating the seeds in a moist chamber at 80°C for 30-60 minutes). Jaynes (1968c) suggested that this response to high temperature may have adaptive value and could have evolved as a result of recurring ground fires. He also suggests that K. ericoides, a closely related species that is native to sandy pine areas of Cuba, may require a similar pregermination treatment. Kalmia microphylla is similar to K. polifolia in that it needs no pregermination treatment. The studies of Flemer (1949) and Jaynes (1971c) indicate that all species of Kalmia germinate best at 22°C under light in a mixture of sand, ground sphagnum, and peat moss that has a pH of 4.2. A mycorrhizal association, without which growth will not occur, has been demonstrated for Kalmia latifolia, and it is probably necessary for all species of the genus. Flemer (1949) found that mountain laurel seedlings grew very poorly in sterile culture without the associated endophyte, but when the seedlings were combined with the fungal associate they rapidly outgrew the controls. Microscopic examination revealed hyphae sheathing the younger por- tions of the roots and frequent hyphal connections with mycorrhizal coils in the cortical cells. No attempt was made to identify the fungus involved. GENETICS OF THE GENUS KALMIA Many species of Kalmia are sympatric, and most have similar habitat requirements. Even so, no natural hybrids have been reported, and recent studies suggest that such hybrids are unlikely to occur. Jaynes (1968a, 1971d) made all possible crosses, including reciprocals, between 6 species of the genus (omitting only Kalmia ericoides) and found that the barriers to gene flow are well developed. Most of 1974] Genus Kalmia — Ebinger 323 the interspecific crosses were difficult to complete since no seed was set, the resulting seed failed to germinate, or the seed produced albino or yellow-green seedlings. Stylar inhi- bition of pollen tube growth appeared to be the most com- mon reason for the lack of seed set. Only 9 combinations yielded viable seedlings and all were sterile. By contrast, intra-specific crosses between plants differing in flower color and growth habit within and between geographie sources showed no evidence of genetic barriers. Little is known about the inheritance of traits in the genus; the only detailed study involves flower color in Kalmia angustifolia. In this species, Jaynes (1971b) found that the presence of anthocyanin pigment is determined by a single dominant gene, with the homozygous recessive lacking red pigment in the leaves and stem and having white flowers. This trait is carried in the population as a heterozygote, but is rarely expressed. These white forms are at a selective disadvantage in the wild as the seedlings are weak, especially when small. Inbreeding depression and self incompatibility have been observed in some species of Kalmia. Jaynes (1968b) found that self pollinated Kalmia latifolia and K. angustifolia set 85-90% less seed than cross pollinated plants, while K. poli- folia, in contrast, showed no self incompatibility. The seed- lings of all three species, however, showed inbreeding de- pression. As measured by height growth, the vigor of the outcrossed seedlings was generally twice that of the selfs. Jaynes (1971a) also noticed inbreeding depression when sibling red-budded mountain laurel was crossed. It is possi- ble that compact seedlings of Kalmia species of potential ornamental value can be obtained by selfing. Few cytological studies of the genus have been under- taken. Hagerup (1928) in an extensive cytological study of the bicornes reported a chromosome number of n = 12 for Kalmia latifolia and n = 24 for K. glauca (K. poli- folia). He found that the chromosomes of K. polifolia were about half as large as those of K. latifolia, so that the amount of chromatin in the two species seemed to be about 324 Rhodora [Vol. 76 the same. Later, Callan (1941) reported a chromosome number of 2n — 44 for K. polifolia, which in light of recent studies is an error. Jaynes (1969) studied cytologically all of the taxa except for the Cuban species. He found that with the exception of K. polifolia (n — 24), all had a chro- mosome number of » — 12. No differences were found in the size of the chromosomes, and their smallness prevented a karyotype analysis. POLLINATION IN KALMIA The most distinctive feature of the genus is its pollen- discharge mechanism. Near the middle of the corolla are 10 sacs that form small lobes on each keel of the flower bud. Just before anthesis the anthers are pushed upward into these sacs by the elongating filaments. As the corolla opens the anthers are held in the sacs and are carried down and outward, the elastic filaments being bent backward under tension. When the flower is disturbed by a large insect, one or more of the anthers are released from their sacs. The filament, which is under tension, snaps the anther upward, showering the insect with pollen. This mechanism was first described by Sprengel (1793) and later by Hasskar] (1863) for Kalmia latifolia, while Rothrock (1867) observed the same mechanism in K. angustifolia, Studies by Penhallow (1882) showed that the pollen of K. latifolia could be thrown up to 14 cm. from the flower while in K. angusti- folia it was thrown up to 8 cm. away. These authors sug- gested that this was a method of insuring self-pollination since the pollen was thrown toward the stigma of the flower. Beal (1867) was the first to describe the way in which the flowers were cross-pollinated by insects. He observed that while a bumble bee was searching around the base of the flower, the proboscis liberated the stamens which projected their pollen onto the underside of the insect’s body. This pollen was then rubbed onto the stigma of other flowers visited. This process was also discussed by Gray (1876, 1879). 1974] Genus Kalmia — Ebinger 325 When an anther is released, the pollen tetrads are dis- charged as a fine powder in some species, and in others as a sticky net caused by the presence of viscid threads that hold the tetrads together. According to Matthews and Knox (1926) these extremely fine, non-cellular threads are prob- ably derived from small quantities of microspore mother cell protoplasm which are excluded from the pollen tetrads. They also suggest that this feature is an adaptation that facilitates pollination in relatively large, upright flowers like those of the genus Rhododendron. These threads were found in Kalmia latifolia by Ikuse (1954) and have been observed by the present author in K. hirsuta and K. eri- coides. Their presence or absence was used by Copeland (1943) as a major characteristic in separating the tribes of the subfamily Rhododendroideae, with the tribe Phyllodo- ceae being the only one that consistently lacked these threads. According to Wood (1961) it is now apparent that viscid threads do occur in some species of Kalmia, Phyllo- doce and Loiseleuria, all members of the Phyllodoceae. Because of this, the usefulness of this characteristic to distinguish tribes is doubtful. Only a few species of insects have been observed polli- nating Kalmia species. Beal (1867) listed bumble bees and other Hymenoptera, but mentions no species. The only de- tailed account was by Lovell and Lovell (1934) who col- lected and identified the pollinators found on a population of Kalmia angustifolia near Waldoboro, Maine. Their ob- servations show that K. angustifolia is effectively cross- pollinated by bees and that a total of 14 species of insects is capable of springing the stamens. Of these, Bombus ternarius and Andrena vicina were the most common visi- tors to the flowers. During the present study, populations of K. angustifolia and K. latifolia were examined in south- ern Connecticut, and all insects found to release the stamens were collected and identified. These studies were made for 2 hour periods, at various times each day, from June 15 to June 25, 1970. In most instances the insect did not release the stamen by just landing on the flower. Usually the sta- 326 Rhodora [Vol. 76 mens were sprung by the proboscis while searching for nectar, but sometimes the legs set the stamens free. No honey bees were found on the flowers, and none were re- ported by Lovell and Lovell (1934). Recently bees have been used by Jaynes (1971a) to polli- nate caged plants of Kalmia latifolia and K. angustifolia for the large scale production of seed. In these experiments selected plants were caged before the flower buds opened. Honey bees (Apis spp.) or bumble bees (Bumbus spp.) were introduced into the cages when the flowers started to open. The plants that were in cages with bumble bees set large quantities of seeds (58 to 95% of the flowers forming capsules), while plants in cages with honey bees set fewer seeds (2 to 31% of the flowers forming capsules). Plants that were in cages with no bees rarely set seed (less than 1% of the flowers forming capsules). CHEMISTRY OF KALMIA Catesby (1743) was one of the first to report on the poisonous properties of members of this taxon. He found that when cattle and sheep were deprived of better food during severe winters, many died from eating the leaves of Kalmia latifolia (as Chamaedaphne). An extensive report by Kalm (Benson, 1937) mentions that young sheep die immediately or become very sick after eating leaves of Kalmia latifolia, but calves “were cured by giving them gunpowder and other medicines.” According to Kalm, larger domestic animals are also affected by the leaves. Most of the literature on livestock poisoning by Kalmia latifolia and K. angustifolia has been reviewed by Crawford (1908), Muenscher (1957) and Kingsbury (1964). Some experimental work has been done to determine the symp- toms, dosage, and treatment of the poisoning by these spe- cies. One of the first studies was made by Wood (1883), who fed a decoction of K. angustifolia leaves to a sheep. The symptoms were similar to those observed by Pritchard (1956) who studied K. angustifolia poisoning in a flock of 1974] Genus Kalmia — Ebinger 327 Minnesota sheep. Experiments with K. latifolia by Cary and Matthews (1903), Crawford (1908) and Marsh and Clawson (1930) showed that all livestock tested were poi- soned by this species. The western species, K. microphylla, is also poisonous but, because of its habitat, is seldom encountered by livestock. Fleming (1920) fed this species to some calves and sheep, most of which became sick, while Clawson (1933) found that the symptoms exhibited by sheep and goats were similar to those observed in animals fed K. latifolia and K. angustifolia. No information is available concerning the poisonous properties of the other members of the genus. Catesby (1743), Kalm (Benson, 1937), Bigelow (1818), and other early authors mentioned that deer can eat the leaves of mountain laurel without suffering ill effects. Forbes and Bechdel (1930, 1931) indicate that laurel is eaten in considerable quantity by deer, especially in time of food shortage. When restricted to a laurel diet, however, deer become thin and weak and usually develop a mild con- dition of rickets. Death results when the animals are fed in excess of 1.29% of their live weight of laurel leaves. All species of Kalmia are probably poisonous to humans (Hardin and Arena, 1969), although no deaths have been attributed directly to this genus. Bigelow (1818) and Grif- fith (1847) have reported instances of poisoning in humans. In these cases, a decoction made from the leaves of Kalmia latifolia produced toxic symptoms. The tough leaves and their bitter taste make it unlikely that anyone would eat them. Also, the quantity required to produce illness is greater than what a person would probably consume. Ac- cording to Barton (1802) the Delaware Indians used the leaves of K. latifolia for suicide. It has been suggested by various authors that the honey produced from Kalmia flowers is poisonous, but no definite proof is available. Barton (1802) reported that the honey from Kalmia angustifolia, K. latifolia, and K. hirsuta is poisonous to humans and listed a few instances of suspected poisoning by honey of these species. Miller (1895) and 328 Rhodora [Vol. 76 Pellett (1920), in contrast, doubted that mountain laurel was responsible for poisonous honey. They felt that due to the abundance of this species it would seem that the cases of poisoning would be reported much more frequently if there was good reason to suspect the honey from this source. Some doubt is also expressed by Lovell (1926, 1951) since bees are rarely found on mountain laurel. Howes (1949), as well as Hardin and Arena (1969), indicate that Kalmia latifolia honey is an extremely distressing narcotic, varying in its effect in proportion to the quantity eaten. In none of the reports is the source of the poisonous honey known for certain, the genus Kalmia being suspected because other parts of these plants are known to be poisonous. Andromedotoxin, the poisonous compound found in the genus Kalmia, is known to occur also in other genera of the Ericaceae. Numerous attempts have been made to isolate and determine the chemical structure of this compound. Lasché (1889) reported the occurrence of andromedotoxin in the leaves and twigs of Kalmia angustifolia and also in the fruits of K. latifolia, while Matusow (1897) found what he considered andromedotoxin in the roots of K. latifolia. More recently Jacobs and Lloyd (1939) and Waud (1940) isolated andromedotoxin from K. angustifolia. The chemical structure of andromedotoxin has been studied by Wood, Stromberg, Keresztesy and Horning (1954) and more re- cently by Tallent, Riethof and Horning (1957). The latter authors renamed the substance acetylandromedol and de- veloped a procedure for detecting its presence in plant extracts through the use of paper electrophoresis. Though the chemical structure of andromedotoxin is still in doubt, its empirical formula is probably C..H,,0.. FOSSILS OF KALMIA Only a few fossil species of Kalmia have been named. Most are described from leaf impressions, and except for size, shape, and probable coriaceous texture, there is little reason to consider them members of the genus Kalmia, or 1974] Genus Kalmia — Ebinger 329 members of the family Ericaceae. One living species is also known from fossil remains. Kalmia brittoniana Hollick (1892). This fossil was first described from clay beds of the Raritan formation, near Kerischerville, Staten Island, New York and is Upper Cretaceous in age (Museum, Staten Island Association of Arts and Science). The material il- lustrated consists of parts of three oblong, entire, probably coriaceous leaves, 0.7 X 2.7 cm., with prominent midribs and no secondary veins. Hollick (1906) mentions that the absence of secondary nervation, indicating a leaf of thick, coriaceous texture, was what largely influenced his refer- ring the leaf to the genus Kalmia. This species has also been reported from the Upper Cretaceous deposits of North Carolina (Berry, 1907, 1920) and Alabama (Berry, 1913, 1919). When he described Kalmia brittoniana from the Tuscaloosa formation of Alabama, Berry (1919) suggested that there is no reason for its placement in the genus Kalmia, or even in the Ericaceae. He further sugested that the fossil probably represents some Upper Cretaceous spe- cies of the Proteaceae or the Myrtaceae. Kalmia ? elongata Ashlee (1932). This species was found in the Latah formation, Elk River, Idaho, and is Upper Miocene in age (University of Idaho collection No. 303). The specimen consists of two narrowly elliptical, subcoriaceous leaves, 0.9 X 1.7 cm., with entire margins and a petiole 2-4 mm. long. The leaves are super- ficially similar to Kalmia microphylla, but since the midrib is grooved and the secondary and tertiary veins are present, this fossil must be considered a doubtful member of the genus. Kalmia elliptica Brown (1962). This species was found in the Fort Union formation (Ilo Post Office, Wyoming) and the Denver formation (Golden, Colorado) and is Paleocene in age. The specimen illustrated by Brown (1962) consists of a single elliptic leaf with entire margins and secondary and tertiary veins that ir- 330 Rhodora [Vol. 76 regularly fork and loop near the margins. The specimen is similar in outline to a typical leaf of Kalmia latifolia but is about twice as large (7 X 17 cm.). The presence of the well developed and large number of secondary and tertiary veins, as well as their angle to the midrib, is not typical of any present day species of Kalmia. It must therefore be considered a doubtful member of the genus. Kalmia saxonica Litke (1968). This species was found at Brandis, northwest Saxony, Germany, and is Lower Miocene in age. It consists of pieces of cuticle with the upper epidermis occasionally attached. 'These fragments indicate that the leaf was entire margined, 1-1.3 em. wide, and had an obvious midvein. The cells are polygonal with wavy margins, the stomata are round, 18u wide, surrounded by a double cuticular ring and occasionally by radiating cuticular striations. The basal portion of glan- dular hairs are also present. The structure and arrange- ment of the cells is similar to that found in Kalmia latifolia, and it is possible that this fossil represents a member of this genus or a closely related genus of the Ericaceae. Similar cuticle remains were placed in the form genus Kalmiophyllum by Kráusel and Weyland (1959) when they described the species Kalmiophyllum marcodurense. Kalmia polifolia, the Bog Laurel, has also been found in fossi] form. It was first reported by Berry and Johnston (1922, as K. glauca Ait.) from interglacial deposits at Point Grey near Vancouver, British Columbia. The fossil specimen illustrated is of the same shape and size as those of living specimens of K. polifolia. Since the other fossil plants found clearly indicate a bog habitat, it is very pos- sible that this fossil is a species of Kalmia. Kalmia polifolia has also been reported from Pleistocene lake deposits of the upper Connecticut River Valley in northern New Hamp- shire (Hollick, 1931). These fossils are post-glacial in age and according to Hollick are representative of the flora that migrated northward in the wake of the retreating Wiscon- sin ice sheet. The other fossil species found in the same deposits indicate a habitat and climate similar to that pres- 1974] Genus Kalmia — Ebinger 331 ently prevailing in the area. The plates in this article show excellent examples of leaves very similar in shape and size to those of living K. polifolia. As mentioned by Hollick, however, positive identification would “merely represent a matter of personal opinion" since the leaves of K. polifolia, Andromeda polifolia, and Ledum groenlandicum appear al- most identical in outline. TAXONOMIC TREATMENT KALMIA Linnaeus, Sp. Pl. 1: 391. 1753. Kalmiella Small, Fl. Southeast. U. S. 886. 1908. Chamaedaphne Catesby ex Kuntze, Rev. Gen. Pl. 2: 388. 1891, non Moench. Evergreen or rarely deciduous shrubs or small trees with pubescent to occasionally glabrous, terete or 2-edged twigs. Lateral buds very small, flattened, with 2 exposed valvate to overlapping scales. Leaves alternate, opposite, or in whorls of 3, exstipulate, usually coriaceous, petioled to sub- sessile, margins entire. Flowers solitary in the axils of leaves, or in terminal or axillary corymbiform racemes, or in terminal compound corymbs. Pedicels slender, terete, usually pubescent and with stalked glandular hairs, sub- tended by a pair of bractlets in the axil of a small leaf-like bract. Flowers 5-merous, actinomorphic, with hypogynous insertion. Calyx synsepalous, usually green and persistent in fruit; lobes 5, longer than the tube. Corolla sympetalous, white to pink, red or purple, with a short cylindrical tube extending into a rotate limb that is shallowly 5-lobed and commonly 10-keeled, the limb with 10 saccate depressions in which the anthers are held under tension. Stamens 10, shorter than the corolla; filaments usually filiform, glabrous or pubescent, dorsifixed; anthers 2-celled, unappendaged, short, ovoid, opening by large terminal slits; pollen dis- charged in tetrads, with or without viscid strands. Ovary superior, 5-celled, placentation axile, placentae massive with numerous ovules; style slender, terete, nearly straight; stigma unexpanded, slightly grooved. Fruit a subglobose 332 Rhodora [Vol. 76 to globose, 5-valved, septicidal capsule usually subtended by the persistent calyx. Seeds numerous, small and light- weight, testa tight or loose and extending well past the ends of the remainder of the seed. A North American genus extending from Alaska south in the mountains to California and Utah, eastward through Canada to the At- lantic Ocean and southward through eastern United States to Florida and Cuba. Lectotype Species: Kalmia latifolia L. (cf. Britton 1908). KEY TO THE SPECIES OF KALMIA A. Leaves opposite; inflorescence a terminal corymbiform raceme. B. Midrib of leaves lacking stalked glands; seeds less than 1.5 mm long . .. ..... 1. K. microphylla. B. Midrib of leaves with stalked glands on both sur- faces; seeds more than 1.5 mm long. . 2. K. polifolia. A. Leaves alternate or in whorls; inflorescence various. C. Inflorescence a terminal compound corymb; leaves usually more than 2 em broad. ..... 3. K. latifolia. C. Inflorescence a simple corymbiform raceme or the flowers solitary in the axils of the leaves; leaves usually less than 2 cm broad. D. Leaves more than 1.5 em long; inflorescence a corymbiform raceme from the axils of the leaves. E. Leaves in whorls of 3, evergreen. .......... IDEM ... 4. K. angustifolia. E. Leaves alternate, deciduous. .. 5. K. cuneata. D. Leaves less than 1.5 cm long; flowers mostly soli- tary in the axils of the leaves. F. Leaves broad, flat, the margins only slightly revolute; flowers scattered along the stem in the leaf axils. .............. 6. K. hirsuta. F. Leaves narrow, strongly revolute; flowers clustered toward the ends of the twig giving the appearance of a terminal corymbiform raceme. ........... ls 7. K. ericoides. 1974] Genus Kalmia — Ebinger 333 1. Kalmia microphylla (Hooker) Heller, Bull. Torrey Bot. Club 25: 581. 1898. Low growing alpine plant to sparingly branched shrub to 6 dm tall; branchlets terete to slightly 2-edged below the leaves, glabrous to lightly pubescent; leaves opposite, sub- sessile, the broad petiole to 2 mm long, the blades coriaceous, ovate to oval to narrowly lanceolate, to 4 em long, 0.4-1.1 cm broad, whitened beneath with small hairs, a band of white hairs usually present on the upper surface on either side of the midrib, midrib lacking stipitate glands, margins flat to slightly revolute; inflorescence a few-flowered, terminal, racemose cluster, each flower on a glabrous pedicel 1-3 cm long, each pedicel subtended by a pair of bractlets in the axis of a leaf-like bract; calyx pink to light green, 5-10 mm broad, the lobes obtuse, margins ciliate; corolla rose-purple (rarely white), 0.8-2.0 cm broad; anthers purple, 0.6-1.6 mm long; fruit subglobose, 4-7 mm broad, 3.5-6 mm long; seeds 0.8-1.4 mm long, the testa extending past the ends of the seed. Chromosome number n — 12. Kalmia microphylla has rarely been considered specifically distinct from K. polifolia, Hall (1912), when he transferred the taxon to a variety of K. polifolia mentioned that “No characters upon which the variety might be raised to spe- cific rank have as yet been pointed out, except the variable one of size in plant and leaf." Most present floristic works maintain this taxon as a variety or subspecies of K. poli- folia, though it has been considered a separate species by Small (1914), Rydberg (1954), Hultén (1948) and Abrams (1951). Hultén (1948) mentioned that it was with some hesitation that he accepted K. microphylla as being specifi- cally different from K. polifolia, but that in Alaska, there does not seem to be any intermediate types. However, in 1968 he considered K. microphylla a subspecies of K. poli- folia. When these taxa are considered distinct the characters of plant and leaf size are the most commonly used for separation. Kalmia polifolia is a large shrub, 3 or more dm 384 Rhodora [Vol. 76 tall, the leaves are 2-4 cm long, 214 to 3 times longer than broad, and the margins are usually strongly revolute. Kalmia microphylla, in contrast, is a much smaller plant, usually less than 1.5 dm tall, the leaves are usually less than 1.5 em long, less than 215 times as long as broad, and the margins are little if at all revolute. In most instances, these characters are sufficient to separate the two taxa; however, variation and overlapping of characters do occur. Other characters that have previously been overlooked are much more reliable in separating the two species. In K. polifolia clavate glands are present on the upper and lower surface of the leaf midrib, and the seeds are 1.5 to 2.2 mm long, while in K. microphylla these glands are absent, and the seeds are usually less than 1.3 mm long. Kalmia microphylla is the only species of the genus found west of the Rocky Mountains. It extends from central California north to Alaska, and east through the Yukon and Northwest Territories to the extreme northwest corner of Manitoba. It is primarily a low-growing alpine shrub rarely exceeding 1 dm in height, but in bogs at lower eleva- tions it may reach a height of 6 dm. The species consists of two varieties, one restricted to alpine and tundra areas, and one found in coastal lowland regions. KEY TO VARIETIES 1. Leaves oval to broadly elliptic, usually less than 2 cm long and less than 214 times as long as broad; calyx less than 7 mm broad. .......... la. var. microphylla. 1. Leaves lanceolate, usually 2.5 cm long or longer, 21% to 4 times as long as broad; calyx over 7 mm broad. .... gaa new wea nEy ecaey eenerhu names 1b. var. occidentalis. 1a. Kalmia microphylla (Hooker) Heller var. microphylla. Kalmia glauca Ait. y microphylla Hook. Fl. Bor.-Amer. 2: 41. 1834. TYPE: swamps in the Rocky Mountains, Drum- mond (K). Kalmia polifolia var. microphylla Rehder in Bailey, Cycl. Am. Hort. 2: 854. 1900. (without basionym). Kalmia polifolia var. microphylla (Hooker) Hall, Univ. 1974] Genus Kalmia — Ebinger 335 California Publ. Bot. 4: 201. 1912. Kalmia polifolia ssp. microphylla (Hooker) Calder & Taylor, Canad. Jour. Bot. 43: 1398. 1965. Sparingly branched alpine shrub usually less than 1 dm tall (to 3 dm tall at lower elevations) ; nodes usually less than 1 em apart; leaves ovate to oval to broadly elliptic, to 1.5 (rarely 2.5) cm long, less than 214 times as long as broad; calyx pink to light green, 5-7 mm broad; corolla rose-purple 0.8-1.2 (rarely 1.5) cm broad; fruit 4-5 mm broad, 3.5-4.5 mm long; seeds 0.8-1.3 mm long. Distribution: Mountainous regions of western North America from California, Nevada, Utah and Colorado north through the Rocky Mountains to the Yukon and Northwest Territories. Common name: Alpine Laurel, Small-leaved Kalmia. REPRESENTATIVE SPECIMENS UNITED STATES: Alaska: Skagway, Anderson 1608 (1sc), Eastwood (GH). California: EL DORADO CO.: Echo Lake, Beatle & Beatle 3959 (ARIZ, DUKE, NY). FRESNO CO.: Marie Lake, Kruckeberg 3487 (RM, WS). HUMBOLDT CO.: Trinity Summit, Davy & Blasdale 5823 (Us). INYO CO.: Pine Creek, Duran 3338 (GH, MICH, US). MADERA Co.: Shadow Creek, Raven (NY). MARIPOSA CO.: Crescent Lake, Congdon (MIN, WS). MODOC CO.: Dismal Swamp, Weatherby 1598 (NY, RM). MONO CO.: Treble Lake, Munz 19998 (NY). NEVADA co.: Frog Lake, Sonne 438 (MU, PH, US). PLACER CO.: Squaw Valley, Eggleston 21607 (GH, NY, US, WS). PLUMAS C0.: Long Lake, Hall 9325 (GH, RM, US). SHASTA C0.: Lassen Nat Park, Ferris & Lorraine 10459 (GH, RM, US). SISKIYOU CO.: Mt. Shasta, Cooke 11296 (GH, osc), 17654 (WS, WIU). TULARE CO.: Sky Blue Lake, Howell 25555 (US). TUOLUMNE CO.: Unicorn Creek, Mason 698 (GH). Colorado: BOULDER C0.: Brainard Lake, Green 279 (PENN). CLEAR CREEK CO.: Loveland Pass, Robbins 732 (ARIZ). GILPIN C0.: Forest Lake, Roberts (CM). JACKSON CO.: Cameron Pass, Beaman & Erbisch 1362 (MSC, RM). JEFFERSON CO.: Geneva Creek, Livingston 298 (DUKE). LARIMER co.: Estes Park, Cooper 116 (RM), 157 (ARIZ, MIN, NMC, RM). ROUTT CO.: Routt Natl Forest, Weber 6991 (ws). Idaho: BLAINE co.: Sawtooth Peaks, Macbride & Payson 3707 (GH, RM, US). BOISE CO.: Middle Spangle Lake, Hitchcock & Muhlick 10130 (GH, NY, RM). BONNER CO.: Priest Lake, MacDougal 292 (NY). BOUNDARY co.: Mt. Roothaan, Daubenmire 44289 (WTU). CLEARWATER CO.: Lolo Trail Rd., Joslyn 953 (1D), 954 (ID). FREMONT CO.: Henry Lake, 336 Rhodora [Vol. 76 Payson & Payson 2047 (cH, NY, RM). IDAHO CO.: Big Fog Mt. Baker 14762 (1D). SHOSHONE co.: Little Lost Lake, Wilson 318 (GH, IDS, WS). VALLEY CO.: Boulder Lake, Boone 177 (1D). Montana: BEAVERHEAD CO.: Odell Lake, Hitchcock & Muhlick 14915 (NY, WS, WTU). CARBON CO.: Beartooth Mts., Cronquist 8068 (GH, NY, ws, WTU). DEER LODGE CO.: Wate (MONT). FLATHEAD CO.: Johns Lake, Standley 18513 (US). GALLATIN CO.: Spanish Peaks, Vogel (cu, MONT, US). GLACIER CO.: Hidden Lake, Hitchcock 1905 (MONT, RM). GRANITE CO.: Helena, Kelsey (MU, NY). LAKE CO.: McDonald Lake, Hitchcock 18343 (RM, ws, WTU). MADISON C0.: Old Hollowtop, Rydberg & Bessy 4654 (MONT). MISSOULA CO.: Lace Lake, Rossbach & Rossbach 184 (WTU). PARK CO.: Lady of the Lake, Witt 1233 (MINN, NY, WS, WTU). POWELL CO.: Upper Tineup Creek, Trask 117 (MONT). RAVALLI CO.: St. Mary's Peak, Hitchcock & Muhlick 15332 (NY, WS, WTU). STILLWATER CO.: Stillwater, Hawkins 35502 (MONT). SWEET GRASS CO.: Rainbow Lake, Hitchcock 16501 (CAN, IDS, NY, WS, WTU). TETON C0.: Henry Mts., Williams (GH). Nevada: ELKO Co.: Lamoille Lake, Holmgren 1404 (BRY, NY). WASHOE Co.: Mt. Rose, Archer 6425 (ARIZ, GH, NY, RENO). Oregon: BENTON Co.: Three Sisters, Vechten 212 (osc). CLACKAMAS CO.: Clackamas Lake, Bart- lett & Grayson 791 (NY). CROOK COo.: Des Chutes Nat'l. Forest, Ferris & Duthie 488 (RM). DESCHUTES CO.: Sparks Lake, Bellinger (WILLU). DOUGLAS CO.: Umpqua, Ingram 1515 (ORE). JACKSON CO.: Mt. Pitt, Peck 5184 (WILLU). KLAMATH CO.: Crater Lake Applegate 319 (US, wILLU), 345 (Us). LANE Co.: McKenzie Pass, Henderson 16315 (ORE). LINN C0.: Hensley Meadows, Coville & Applegate 1152 (US). MARION CO.: Russell Lake, Lee 44 (osc). UNION co.: Head 1298 (osc). WALLOWA CO.: Mirror Lake, Peck 1777 (NY, WILLU). Utah: DUCHESNE CO.: Mt. Agassiz, Cottam & Biddulph 3703 (BRY). SUMIT CO.: Motte Peak, Payson & Payson 5139 (CHRB, GH, PH, RM, US, WS). UINTAH CO.: Leidy Peak, Maguire 17679 (RM). UTAH CO.: Grandaddy Lakes Basin, Stutz 224 (BRY). Washington: CHELAN CO.: Ice Lake, Morrill 332 ( MONT, ND, WTU). KITTITAS CO.: Mt. Stuart, Elmer 1103 (GH, K, MIN, US, WS). KLICKITAT CO.: Falcon Valley, Suksdorf (PH, ws, WTU). LEWIS CO.: Reflection Lake, Townsend (WS). OKANOGAN CO.: Slate Peak, Ownbey & Meyer 2302 (an, IDs, ISC, MIN, NY, OSC, US). PEND OREILLE CO.: Sullivan Lake Dist., Layser 866 (WS). PIERCE CO.: Mt. Rainier, Allen 96 (CAN, GH, K, MIN, MSC, NY, US, WS). SKAMANIA CO.: Chiquash Mts., Suksdorf (WS). WHATCOM CO.: Mt. Baker, Muenscher 8077 (ws). YAKIMA co.: Bird Creek Meadows, Zuberbuhler (ws). Wyoming: ALBANY co.: Brooklyn Lake, Nelson 9226 (GH, MIN, RM). BIG HORN CO.: Upper Crater Lake, Lofgren 13 (MIN, NY, RM). CARBON CO.: Silver Lake, Henry (CM). FREMONT CO.: Gannett Creek, Jozwik 386 (Gu, RM). JOHNSON CO.: Powder R. Pass, Stephens 40827 (VDB). LARIMER 1974] Genus Kalmia — Ebinger 337 co.: North Park Range, Goodding 1839 (PENN). PARK CO.: Island Lake, Truman 54298 (WTU). SHERIDAN CO.: Dome Lake, Pammel & Stanton 62 (MIN). TETON CO.: Taggart Lake, Williams 1119 (ID, MIN, OSC, RM). YELLOWSTONE NAT'L PARK: Artists Paint Pots, Nelson & Nelson 6170 (CM, ISC, MIN, GH, NY, RM, US). CANADA: Alberta: Banff Natl. Park: Bow Pass, Saunders 11 (rRT); Lake Louise, Brown 578 (GH, PH, US), McCalla 2162 (ALTA, NY, US); Waterfowl Lake, Walker 2443 (PENN). Jasper Nat. Park: Athabasca Falls, Moss 2752C (ALTA, GH); Maligne Lake, Scamman 2664 (GH); Amethyst Lake, MacFadden 3246 (No). Waterton Lakes Nat! Park: Summit Lake, Moss 975 (GH, NY, US); Upper Twin Lake, Blais & Nagy 1793 (CAN). Fairfax Lake, Damais & Anderson 2335 (ALTA); Mercoal, Woollven 6 (ALTA); Viewpoint, Damais & Kempinsky 5683 (ALTA); Virginia Hills, Moss 12469 (ALTA), 12505 (ALTA, CAN). British Columbia: Ernest C. Manning Prov. Park: Timberline Valley, Hardy 18501 (usc, v); Three Brother Mts., Underhill 782 (v). Garibaldi Prov. Park: Castle Towers, Davidson (usc). Glacier Nat'l. Park: Bald Mt., Fogg, Jr. 1273 (PENN, UBC); Eagle Mt., Rosendahl 944 (MIN). Kokanee Glacier Prov. Park: Thompson 14447 (CAN, GH, MICH, NY, PH, US, WTU). Kootenay Natl. Park: Ross Lake, McCalla 9603 (uBC, v); Emerald Lake, Brown (TRT). Mt. Assiniboine Prov. Park: Magog Lake, Green 551 (PENN). Mt. Revelstroke Nat. Park: Calder & Savile 10861 (Ny, US), 8884 (Nv, US). Tweedsmuir Prov. Park: Trew 17177 (v). Wells Gray Prov. Park: Battle Mt., Ahti & Ahti 6975 (v, wTU). Yoho Nat'l Park: Emerald Lake, Beer (cM); Selkirk, Shaw 41 (GH, MIN, NY, PENN, US); Yoho Lake, Pease 22371 (MASS). Vancouver Island: Mt. Arrowsmith, Taylor 164 (TRT); Mt. Albert Edward, Hett 352 (v); Moat Lake, Underhill 299 (wTU). Atlin, Szezawinski (UBC, V). Caribou Pass, Henry 608 (GH, ISC); Ft. Steele, Johnstone (UBC, v); Lillooet, Beamish & Vrugtman 610710 (vBC); Mt. Hazel, Slocomb (v); Penticton, Calder & Savile 10751 (NY, RM, UBC, V, WS) ; Prince George, Holland (v); Sikanni Chief R., Henry 743 (PH); Summit Lake Landing, Weber 2589 (GH, Ws); Skagit Valley, Macoun 72406 (ND, NY); Tenquille Lake, Beamish & Vrugtman 61010 (1D, UBC, WTU). Manitoba: Churchill, Beckett M-45-5 (MIN), Ellis 1287 (Us), Gillett 1832 (Isc, us), Gillis 3244 (msc), Voss 8999 (MICH), Walker 5090 (WIN); Tidal, Brown 178 (CAN, SASK). Northwest Territories: Aberdeen Lake, Macpherson 327 (CAN); Beverly Lake cabin, Kuyt 107 (CAN); Contwoyto Lake, Thieret 9415 (GH, US); Ennadai Lake, Brown 1320 (CAN): Gordon Lake, Henderson 19 (CAN); Great Bear Lake, Porsild 17117 (cAN), Porsild & Porsild 3429 (CAN), 3509 (CAN, US); Great Slave Lake, Pike (K); Mac- Millan Pass, Porsild & Breitung 11258 (CAN, GH, US) ; Maguse Lake, Gussow 33 (CAN); Nueltin Lake, Harper & Porsild 2211 (CAN, PH), 338 Rhodora [Vol. 76 2246 (CAN), 2271 (MICH, MIN, PH), 2554 (MICH, MIN, NY); Small Tree Lake, Maini & Swan 506 (RM, SASK); Thelon Game Sanctuary, Carroll 65 (CAN); Yellowknife Highway, Thieret & Reich 7742 (CAN, MIN), 8121 (Us). Yukon: Granite Mt., Porsild & Breitung 10576 (CAN, ISC, GH, US, WTU); Mayo Dist., Bostock 102 (CAN); McQueston Area, Campbell 493 (CAN); Mt. Sheldon, Porsild & Brei- tung 11094 (CAN, GH, US, WTU), 11485 (CAN, GH); pass between Teslin & Nisutlin R., Porsild & Breitung 11045 (CAN, Ny), 11148 (CAN); St. Elias Mt., Pearson & Porsild 67-166A (CAN); Yukon- Mackenzie border, Calder & Kukkonen 27664 (Isc, NY); White Pass Summit, Bolton (us). Plant and leaf size in Kalmia microphylla var. micro- phylla is controlled to some extent by the environment. In the typical alpine form the leaves are extremely small, usually less than 1.2 em long. The type specimen, which represents this extreme, has leaves less than 1 cm long and is extremely short, less than 1 dm tall. At lower elevations the leaves are not uncommonly 2 em long and the plant 2 dm tall. Rarely taller plants are found with leaves to 2.6 cm long. These larger leaved forms are superficially similar to K. polifolia and probably account for the variation in tax- onomic treatment of K. microphylla in the past. Kalmia microphylla and K. polifolia probably never come into contact with each other. In the Northwest Territories, however, the northern part of both their ranges, it is possi- ble that the two taxa may occur together. The author has seen very few specimens of K. polifolia from the Northwest Territories, but this species is common in northern Alberta, Saskatchewan, and Manitoba. A few specimens of K. micro- phylla have been examined from the Northwest Territories, and this species extends all the way to the northwestern corner of Manitoba. No specimens, however, have been observed from Alberta or Saskatchewan. Sterile hybrids have been produced by crossing these two taxa (Jaynes, 1968a), but no specimens from northern Canada have been found that are morphologically similar to these hybrids. The only other taxon with which Kalmia microphylla var. microphylla might be confused is K. microphylla var. occi- dentalis. Both taxa occur in western North America and 1974] Genus Kalmia — Ebinger 339 their ranges overlap. The two are easily separated, how- ever, since var. occidentalis has lanceolate leaves that are usually more than 2.5 cm long and 214-4 times as long as broad, while var. microphylla has oval to eliptical leaves that are smaller and less than 215 times as long as broad. Also, var. occidentalis has larger flowers, with the corolla more than 1.5 em broad and the calyx 7-10 mm wide. Vari- ety microphylla, by contrast, has smaller flowers, with the corolla less than 1.5 cm broad (usually less than 1.2 cm broad) and the calyx less than 7 mm wide. Although both varieties are found in marshes, bogs, and other open wet areas, it appears that they rarely come into contact with one another. Of the var. microphylla speci- mens examined, most were collected at relatively high elevations aecording to the information available on the herbarium labels. The elevation at which this taxon was collected varied from an average altitude of 8,500 feet above sea level in California (5,200 to 11,500), to about 10,000 in Colorado, Nevada, Utah and Wyoming (7,000 to 12,500), to about 6,500 in Oregon (5,800 to 7,700), and to about 6,000 in Washington, Alberta and British Columbia (3,000 to 7,300). The altitudes in parentheses indicate the mini- mum and maximum elevations recorded from the labels. The information on the labels of specimens of var. occi- dentalis indicates that this species is found at lower eleva- tions. Most of the specimens were from at or near sea level, but a few had been collected as high as 3,000 feet. Kalmia microphylla var. microphylla and K. microphylla var. occidentalis apparently hybridize when they come into contact. A few specimens from Vancouver Island and the Cascade Mountains of Washington appear to be intermedi- ate morphologically. These intermediate individuals are robust plants, 2-4 dm tall, similar in habit to the large- leaved var. microphylla. The leaves are 1.8-2.5 cm long, usually more ovate than lanceolate, and about twice as long as broad. The flowers are large, more typical of var. occi- dentalis. A few hybrids have been produced by crossing these varieties and they are highly fertile and set large 340 Rhodora [Vol. 76 quantities of seed (Jaynes, personal communication). Of the few suspected hybrids found, the pollen stainability varied from 10-90% normal pollen, indicating that some sterility occasionally results. 1b. Kalmia microphylla var. occidentalis (Small) Ebinger, coinb. et stat. nov. Kalmia occidentalis Small, N. Am. Fl. 29: 53. 1914. TvPE: Washington: foothills of Mt. Rainier, 1885, Mrs. Bailey Willis (NY). Kalmia polifolia subsp. occidentalis (Small) Abrams, Ill. Fl. Pacific St. 3: 302. 1951. Sparingly branched shrub to 6 dm tall; leaves lanceolate, flat to slightly revolute margins, 2-4 cm long, some of the leaves always over 2 cm long and 215-4 times as long as broad; calyx light pink, 7-10 mm broad; corolla usually rose-purple (rarely white), 1.5-2.0 cm broad; fruit 5-7 mm broad, 4.5-6 mm long; seeds 1-1.4 mm long. Distribution: Along the western coast of North America from southern Alaska south to Oregon. Common name: Western Swamp Kalmia; Western Bog Laurel. REPRESENTATIVE SPECIMENS UNITED STATES: Alaska: Admiralty Isl., Hibber 236 (UNM); Annette Isl, Coville & Kearney 381 (vs); Douglas Isl, Thorne 10280 (UWM); Gravina Isl., Anderson 5352 (1sc), 24599 (CAN, ISC); Kuiu Isl, Rigg 17 (wTU); Metlakatlan Isl, Emgstrom (WTU); Mitkof Isl, Brown (ALA); Pleasant Isl, Reiners (HNH); Prince of Wales Isl., Flett 1976 (vs); Revillagigedo Isl., McCabe 8590 (WTU); Woronkofski Isl., Klein 486 (ALA); Yakobi Isl., Anderson 1389 (US); Juneau, Anderson 6338 (ALA, BRY, CAN, GH, ISC, PH, RM, V) ; Menden- hall, Anderson 88 (CHRB); Petersburg, Scamman 515 (ALA, GH); Sitka, Anderson 160 (1sc, US); Wrangell, Walker & Walker 638 (CM, GH, NY, RM, US, ws); Yes Bay, Gorman 21 (K, NO, NY, US). Oregon: CLATSOP CO.: Gearhart, Adair (osc); Knappa, Bouch (WTU); Delmoor, Cooke 10828 (osc). Washington: GRAY HARBOUR co.: Lake Quinault, Thompson 11429 (ALTA, GH, MONT, NY, PH, US, WS, WTU). ISLAND CO.: Whidbey Isl, Smith 480 (ws, WTU). KING CO.: Green Lake, Benson 1474 (WTU). MASON CO.: Lost Lake, Jones 6527 (PH, V, WTU). PACIFIC CO.: Ilwaco, Foster 865 (ORE, US). PIERCE C0.: Tacoma, Sloan (WTU). SAN JUAN CO.: Mt. Constitution, Zeller & Zeller 1193 (CM, GH, MIN, NY, US). SKAGIT CO.: Mt. 1974] Genus Kalmia — Ebinger 341 Higgins, Thompson 9639 (ND, US, WTU). SNOHOMISH CO.: Marys- ville, Grant (ARIZ, OAC, MU, RM, US). THURSTON CO.: Upper Valley of the Nisqually, Allen 4 (CAN, K, MIN, MSC, NY, US, WS). WHATCOM co.: Custer, Muencher & Muencher 6007 (PH, Us, WS). CANADA: British Columbia: Aristazabal Isl, McCabe 3424 (wTU); Dewdney Isl, Schofield 70-27 (UBC); Goose Isl, Martin & Guiquet (v); Hope Isl., Calder & Mackay 31240 (cM, MIN); Lulu Isl., Beamish & Vrugtman 60519 (Ny, osc, UBC); Graham Isl., Calder & Savile 21349 (osc, TRT, ws); Kunghit Isl., Foster & Bigg (UBC); Langara Isl., Beebe (v); Moresby Isl., Calder & Taylor 35771 (UBC); Prince Royal Isl, McCabe 3358 (wTU); Saltspring Isl, Ashlee (UBC, V); Vancouver Isl.: Comox, Anderson 483 (v); McCoy Lake, Carter 473 (us); Lost Lake, Macoun 618 (CAN, US); Port Hardy, Racey (UBC); Trout Lake, Davidson 6192 (usc); Alice Arm, Szczawinski (V); Alouette R., Garman (v); Garibaldi Prov. Park, Dow D-2A-C (v); Haney, Taylor 1002 (UBC); Hope-Prince Hwy., Brayshaw 49470 (uBC); Kelsey Bay, Calder & MacKay 29204 (WTU); Kitsault, Fisher (UBC); New Westminster, McCalla 2944 (ALTA); Prince Rupert, Mendel 78 (v); Prudhomme Lake, Taylor & Lewis 618 (UBC); Terrace, Szczawinski (UBC, V). Kalmia microphylla var. occidentalis has been considered specifically distinct only by Small (1914). In all floristic works of the west coast of North America this taxon has been considered a part of the eastern K. polifolia, although it nas been rarely considered as a separate subspecies (Abrams, 1951). Hultén (1948) mentions that he was not able to separate the Alaskan specimens from those of East- ern America. He indicates, however, that a slight racial difference might exist, but that a large body of material would be necessary for a definite statement. Kalmia microphylla var. occidentalis and K. polifolia are strikingly similar. Both have the same general habit and size, and except for a slight difference in flower size these structures are also the same. Under close morphological examination, however, these two species are easily sepa- rated. In K. polifolia, the leaf margins are usually revolute, and well developed stalked glands are present on both sides of the midrib. In K. microphylla var. occidentalis, by con- trast, the leaves are not revolute, or only slightly so, and the stalked glands are absent. The two taxa also differ in seed size and calyx width. In K. polifolia, the seeds are long 342 Rhodora [Vol. 76 and narrow, 1.5-2.1 mm long, while the calyx varies in width from 5-7 mm. In K. microphylla var. occidentalis, the seeds are shorter and thicker, 1-1.4 mm long, and the calyx varies from 7-10 mm broad. The geographic distribution of the two taxa also differs, and they never come into contact with one another. Kalmia polifolia is mainly an eastern species, occurring as far west as Saskatchewan and northeastern Alberta, while K. micro- phylla var. occidentalis occurs along the west coast from southern Alaska, south through British Columbia and from Washington to Oregon. Most of the specimens of var. occi- dentalis examined were from marshes, bogs and wet open areas at lower elevations from coastal areas and the islands off the coast of Alaska and British Columbia. Hultén (1968), who did not distinguish between these two species, shows this difference in distribution and also has an excel- lent drawing of K. microphylla var. ? occidentalis (as K. polifolia subsp. polifolia) . Cytologically, Kalmia microphylla var. occidentalis (n = 12) is more closely related to K. microphylla var. micro- phylla (n = 12) than it is to K. polifolia (n = 24). Crosses between var. occidentalis and K. polifolia result in triploid F, hybrids that produce less than 10% stainable pollen and no seeds (Jaynes, 1968a). In contrast, crosses between K. microphylla var. microphylla and var. occidentalis were highly fertile with the F,’s being intermediate between the two parents, producing viable seed and pollen that looked normal and fertile (Jaynes, persona] communication). Kalmia microphylla var. occidentalis f. alba Ebinger, forma nov. Forma typicae habita statura etc. similis, floribus omni- bus albis. This white flowering form, which is similar to that found in some other species of Kalmia, has been collected at two different localities. Alaska: near Wrangell, Mrs. J. C. Dart, 1922. (us #1,118,961, Holotype). British Columbia: Lulu Island, peat bog between N. Westminster Highway and No. 4 road, H. Rhodes, et al., 29 May 1954 (UBC). 1974] Genus Kalmia — Ebinger 343 29. Kalmia polifolia Wang., Schr. Berlin Ges. Naturf. Freunde 8: 130. 1788. TYPE: unknown. Kalmia polifolia Ludwig, Neu. Wilde Baumz. 25. 1783. (nom. nud). Kalmia glauca Loddiges ex Ludwig, Neu. Wilde Baumz. 25. 1783. (nom. nud.), Ait. Hort. Kew. 2: 64. 1789. TYPE: Plant growing in the Kew Gardens, (K). Chamaedaphne glauca | ( Ait.) Kuntze, Rev. Gen. Pl. 2: 388. 1891. Kalmia glauca var. praecox Lavallée Arbor. Segrez. 159. 1877. (nom. nud.). Kalmia glauca var. stricta Jager, Ziergeholze 279. 1865. Kalmia glauca var. superba Makoy ex Bosse, Vollst. Handb. Blumengart. 2: 348. 1841. (nom. nud.), Jager & Beissner, Ziergeholze 194. 1884. Kalmia superba K. Koch, Dendrologie 2(1): 154. 1872. (nom. nud.). Kalmia rosmarinifolia Dum.-Cours. Bot. Cult. 2: 250. 1802. TYPE: unknown. Kalmia glauca p rosmarinifolia (Dum.-Cours.) Pursh, Fl. Am. Sept. 296. 1814. Kalmia polifolia var. rosmarinifolia (Dum.-Cours.) . Rehd. in Bailey, Cycl. Amer. Hort. 2: 854. 1900. Kalmia oleaefolia Dum.-Cours. Bot. Cult. ed. 2. 3: 322. 1811. (as synonym). Sparingly branched shrub to 1 m tall; branchlets slightly flattened, 2-edged, glabrous or puberulent, commonly with decurrent ciliate lines from the petiole base; leaves oppo- site, subsessile with a broad petiole to 2 mm long, coriaceous, linear to oblong, the margins usually revolute, 1.5-4 cm long, 0.3-1.3 em broad, 214 to 4 times as long as broad, whitened beneath with small hairs, the midrib covered with purple clavate hairs on both surfaces; inflorescence a few- flowered, terminal corymbiform raceme, each flower on a glabrous pedicel 1.5-3 cm long, each pedicel subtended by a pair of bractlets in the axis of a leaf-like bract; calyx deeply 5-lobed, colorless to light pink, 5-7 mm broad, the lobes obtuse, margins ciliate ; corolla rose-purple, 1.2-1.8 cm broad; anthers purple, 1-1.5 mm long; fruit subglobose, 344 Rhodora [Vol. 76 4-7 mm broad, 4-6 mm long; seeds 1.5-2.2 mm long, the testa extending past the ends of the seed. Chromosome number n — 24, Distribution: Bogs, swamps, and other wet, open areas from northeastern Alberta east to the Atlantic Ocean and south to the Great Lakes, Pennsylvania, New York and New Jersey. Common name: Bog Laurel, Swamp Laurel, Pale Laurel, Gold Withy. REPRESENTATIVE SPECIMENS UNITED STATES: Connecticut: HARTFORD CO.: Suffield, Harger 6524 (ARIZ), Blewitt 3490 (PH). LITCHFIELD CO.: Bingham Pond, Weber 1489 (Isc). NEW HAVEN CO.: Middlebury, Shepard (conn). Maine: ANDROSCOGGIN CO.: Livermore, Gordon 657 (MAINE). AROOS- TOOK CO.: Knoles Corner, Neal 1908 (MAINE), True 6068 (PENN). CUMBERLAND CO.: Standish, Rossbach 5599 (ACAD, MAINE). FRANKLIN co.: Mt. Abraham, Rossbach 5389 (MAINE). HANCOCK CO.: Mt. Desert Isl, Redfield 4266 (US). KENNEBEC CO.: Monmouth, Hyland 90 (MAINE). KNOX CO.: Rockland, Rossbach 1429 (ACAD). OXFORD co.: Old Spec, Richards 6032 (MAINE). PENOBSCOT CO.: Orono, Fernald 349 (GH, Mass, MIN, MSC, ND, NY, PH, US). PISCATAQUIS CO.: Greenville, True 4701 (PAC), 4814 (PENN). SOMERSET CO.: Jackman, True 4668 (PENN). WALDO CO.: Northport, Rossbach 2499 (ACAD), 5349 (MAINE). WASHINGTON CO.: Jonesport, Steinmetz 844 (MAINE). YORK CO.: Sunken Pond, True 214 (GH, PENN). Massachusetts: BERKSHIRE C0.: Wolf Swamp, Walters (NHA). ESSEX CO.: Andover, Steele (MU). FRANKLIN CO.: Hawley, Forbes 24 (MASS). HAMPDEN co.: Ludlow, Seymour 557 (DUKE, GH, MASS, NY). HAMPSHIRE CO.: Belchertown, Torrey (MASS). MIDDLESEX CO.: Concord, Eaton (MAINE). WORCESTER C0.: Worcester, Dowling 818 (NY). Michigan: ALCONA CO.: Hall & Hagenah 447 (BLH). ALPENA co.: Hatt (BLH). BARAGA CO.: Baraga, Richards 1783 (MICH, NY), 2956 (MAINE, MICH). CHARLEVOIX CO.: Beaver Isl, Reis 906 (BLH, MSC). CHEBOYGAN CO.: Mud Lake, Gleason, Jr. 665 (DUKE, GH). CHIPPEWA CO.: Sugar Isl., McVaugh 8745 (BLH, CAN, MICH). CLARE CO.: Lake Station, Dreis- bach 5650 (MICH). CRAWFORD Co.: Hartwick, Bingham (BLH). DELTA CO.: Bull Run Creek, Hagenah 6723 (BLH). EMMET CO.: Carp Lake, Gates & Gates 10334 (RM). GOGEBIC CO.: Watersmeet, Voss 7846 (MICH), 12472 (MICH). GRAND TRAVERSE CO.: "Traverse City, Dieterle 1299 (cM, MICH). GRATIOT C0.: Alma, Davis (MONT). HOUGHTON CO.: Laurium, Herman 7591 (NY, US). IRON CO.: Golden Lake, Voss 4122 (MICH), KALKASKA CO.: Carlisle, Hagenah 4602 (BLH). KEWEENAW CO.: Mott Isl, Bailey & Bailey 4131 (MICH). 1974] Genus Kalmia — Ebinger 345 LEELANAU CO.: Maple City, Thompson L-1693 (BLH). LENAWEE CO.: Clinton, Wood & Wood (us). LUCE CO.: Spring Lake, Hagenah 1913 (BLH). MACKINAC CO.: Hagenah 4876 (BLH). MANISTEE CO.: Manis- tee, Daniels 4267 (MONT). MARQUETTE CO.: Turin, Barlow (NY). MIDLAND CO.: Sanford Game Preserve, Dreisbach 7941 (MICH). MONTMORENCY CO.: Barding Swamp, Case (MICH). OAKLAND CO.: Lakeville, Farwell 4905 (BLH). OGEMAW CO.: Clear Lake, Case, Jr. (MICH). ONTONAGON CO.: Berglund, Beaman 1848 (MS). OSCEOLA co.: Evart, Monaghan (MSC). OSCODA CO.: Zimmerman 416 (MICH). OTSEGO COo.: Otsego Lake Village, Voss 10847 (MICH). ROSCOMMON co.: Higgins Lake, Denton 1038 (MSC). ST. CLAIR CO.: Capae, Dodge (MU, TENN). SANILAC CO.: Minden City, Voss 8341 (MICH). SCHOOL- CRAFT CO.: Long Lake, Voss 2657 (MICH). WEXFORD CO.: Cadillae, House 4660 (Us). Minnesota: BELTRAMI CO.: Star Isl, Butters & Rosendahl 6584 (MIN). CARLTON CO.: Atkinson, Lakela 3550 (DUL, MIN). CASS CO.: Lake Kilpatrick, Ballard (ORE, RM, US, WS). CLEAR- WATER CO.: Long Lake, Morley 1106 (MIN). COOK CO.: Sea Gull Lake, Lakela 3669 (DUL, MIN). CROW WING CO.: Bay Lake, Sheldon 2172 (MIN). ITASCA CO.: Sand Lake, Johnson 2042 (PH), 2207 (GH, US). KANABEC C0.: Mora, Moore & Tryon, Jr. 17504 (MIN). KOOCHI- CHING Co.: Rainy Lake, Moore & Moore 11793 (ISC, MIN). LAKE co.: Basswood Lake, Lakela 8898 (DUL). PINE CO.: Bruno, Jerabek 66 (MIN). ST. LOUIS CO.: Palo, Nickelson 2432 (ISC, NY, ws). New Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon & Steele 9125 (NHA). CARROLL CO.: Church Pond, Gale (NHA). CHESHIRE CO.: Keene, Gilbert (NY, PH). coos Co.: Mt. Washington, Hartmann 207 (NY); Moore 260 (Gv), 1081 (GY). GRAFTON CO.: Mt. Lafayette, Fernald & Smiley 11839 (GH). HILLSBORO CO.: Peterborough, Batch- elder 1891 (KY). ROCKINGHAM CO.: Derry, Batchelder (NHA). STRAFFORD CO.: Barrington, Hodgdon & White 11935 (NHA). SULLI- VAN Co.: Springfield, Hodgdon & Steele 9128 (NHA). New Jersey: MORRIS CO.: Mt. Hope, Edwards & Rusling (CHRB). SUSSEX CO.: Kittatinney Ridge, Griscom 9562 (GH). New York: CHAUTAUQUA CO.: Petit, Thurber (GH). COLUMBIA CO.: Knickerbocker Lake, McVaugh 1013 (PENN). ERIE CO.: Protection, Miller 3764 (MSC). ESSEX CO.: Whiteface Mt., Rudd 824 (US). FRANKLIN C0.: Upper Savanac Lake, Guest 182 (PAC, PENN). HAMILTON CO.: Bern Bog, Henry (CM). HERKIMER CO.: Twin Lake, VanPelt & Long (PH). MADISON CO.: Recksport, House 6930 (US). ONEIDA CO.: White Lake, True 38 (PENN). ONONDAGA CO.: Labrador Pond, Clausen 2129 ANY, PAC, PENN). OSWEGO CO.: Mud Pond, Wiegand 13764 (GH). ST. LAWRENCE co.: Norfolk, Phelps 757 (GH, NY, US). SARATOGA CO.: Saratoga Lake, House 27663 (PENN). SCHENECTADY CO.: Princetown, Svenson (GH). SULLIVAN CO.: Neversink, Jones, Phillips & Stebbins 21 (GH). TIOGA CO.: Owego, Mapes 4767 (GH). ULSTER CO.: Rockland, Camp 3036 (NY). WASHINGTON CO.: Salem, Stewart 63 (NY). Pennsylvania: 346 Rhodora [Vol. 76 CARBON CO.: Albrightsville, Pretz 13321 (PH). LACKAWANNA CO.: Elmhurst, Glowenke 4820 (PENN). MONROE CO.: Tannerville, Leeds 2237 (MIN, PH). PIKE CO.: Shohola Falls, Byhouwer & Kobuski 226 (GH). SUSQUEHANNA Co.: Ararat, Glowenke 184 (PENN). WAYNE co.: Lehigh Pond, Adams 3699 (GH). Rhode Island: PROVIDENCE CO.: Glocester, Collins (CONN, GH, MASS, NHA, NY, PENN, PH). Vermont: ADDISON CO.: Ripton, Eggleston 280 (GH). CALEDONIA CO.: Peacham, Blanchard (PH, US). CHITTENDEN CO.: Colchester, Pringle (BRY). LAMOILLE CO.: Mt. Mansfield, Moseley (MASS). ORLEANS CO.: Lake Willoughby, Stevens (US). RUTLAND CO.: Mt. Tabor, Seymour 24407 (BRY). WINDHAM CO.: Tyson, Moldenke & Moldenke 10263 (Ny). Wisconsin: BAYFIELD C0.: Sand Bay, Cheney 6371 (UWM). DOOR CO.: Bailey’s Harbor Ridge, Shinners (UWM). FLORENCE CO.: Spruce Lake, Mau (UWM). FOREST CO.: Crandon, Palmer 27776 (GY). IRON co.: Mercer, Salamun 228 (UWM). JACKSON CO.: Ketchum, Hartley & Morrissey 9158 (ISC). KEWAUNEE CO.: Gregor (OS). LINCOLN CO.: Tomahawk, Engel (UWM). MARINETTE CO.: Walsh, Throne (UWM). MONROE CO.: La Grange Twp., Hartley 7884 (MIN). ONEIDA CO.: Three Lakes, Wadmond 2758 (MIN). RUSK CO0.: Ladysmith, Goessl 2162 (UWM). SHAWANO CO.: Camp Tivol, Keefe (UWM). TAYLOR co.: Buch Lake Area, Piehl (UWM). viLAS co.: Eagle River, Hollister 334 (US). WASHBURN CO.: Spooner, Cheney 6135 (UWM). CANADA: Alberta: Lake Athabaska: Smith Landing, Preble & Preble 99a (US); Alta, Raupp & Abbe 4529 (ALTA, CAN, MIN, PH). Gregoire Lake, Dumais & Anderson 2693 (ALTA); Richardson Lake, Whitehorn & Barber 185 (SASK). Labrador: Anatolak, Sewall 469 (GH, US), 471 (GH, US); Ashuanipi Lake, Harper 3877 (CAN, MIN, us); Attikamagen Lake, Harper 3501 (CAN, ISC, MIN, PENN); Battle Harbour, Koelz 36 (MICH, US); Cape St. Charles, Potter & Brierly 3410 (GH, NMC); Cartwright, Hitchcock 23818 (vs); Gannet Isl, Potter & Brierly 3411 (GH); Goose Bay, Gillett & Findley 4931 (ACAD, NY); Gready Isl., Bishop 474 (GH), 475 (GH); Hopedale, Walker 370 (PENN); Mealy Mts., Gillett & Findley 5273 (ACAD, NY, TRT, US); Mills Harbour, Cook 88 (GH, US); Petty Harbour, Bishop 476 (CAN, GH); Spotted Isl, Wallace 42 (CONN); Sandwich Bay, Woodworth 329 (GH), 330 (GH); Windy Tickle, Potter & Brierly 3407 (MASS). Manitoba: Baralzon Lake, Scoggan & Baldwin 8216 (GH, MIN, WIN); Cranberry Portage, Freedmann (WIN); Churchill Dist., Schofield 1001 (DUL, GH, MIN, WS); Duck Lake, Scoggan & Baldwin 8245 (ALTA, CAN, GH); Gillam, Punter 97 (WIN); Goldesand Lake, Keleher 107 (WIN); Heming Lake, Keleher 159 (WIN); Herch- nier, Core 215 (WVA); Lake Winnipeg, Lowe (WIN); Lake Winni- pegosis, Scoggan & Baldwin 8152 (CAN); Nelson River, Scoggan 3016 (ALTA, CAN, MIN, WIN); Nueltin Lake, Baldwin 2180 (CAN, TRT, WIN, WS); Oxford Lake, Scoggan 5390 (CAN); Reindeer Lake, Bald- 1974] Genus Kalmia — Ebinger 347 win 2028 (CAN, TRT, WIN); Seal River, Ritchie 1794 (CAN, MIN); Third Cranberry Lake, Freedman (WIN); Wabowden, Gillett 2790 (us); Whiteshell Forest Reserve, Scoggan 8781 (ALTA, WIN); York Factory, Punter 134 (WIN). New Brunswick: ALBERT CO.: Grindstone Isl, Christie 1576 (NBM). CHARLOTTE CO.: Campobello Isl., Malte 955/29 (CAN). GLOUCESTER CO.: Bathurst, Malte 13 (CAN). KENT co.: Little Forks Brook, Squires & Squires (NBM). MADAWASKA CO.: St. Leonard, Maite 605 (CAN). QUEENS CO.: Castaway, Gorham & Squires (NBM). SAINT JOHN CO.: Portland, Matthew (NBM). WEST- MORELAND CO.: Sackville, Fowler (QUK). YORK CO.: Fredericton, Dore & Gorham 4569 (ACAD). Newfoundland: BONAVISTA SOUTH DIST.: Lethbridge, Rouleau 8363 (NFLD). BURGEO-LA POILE DIST.: Grandy Brook, Fernald, Long & Fogg 359 (GH, PENN, PH). FERRYLAND DIST.: Witless Bay, Rouleau 5391 (CAN, NFLD, US). FOGO DIST.: Fogo Isl., Sornborger (GH). FORTUNE BAY-HERMITAGE DIST.: Hermitage Bay, Palmer 1353 (GH, US). GRAND FALLS DIST.: Bishop Falls, Fernald & Wiegand 6023 (CAN, GH, K, NY, PH). HUMBER DIST.: Eagle's Nest Brook, Rouleau 3069 (NFLD). ST. BARBE DIST.: Trout R., Rouleau 3515 (NFLD). ST. MARY'S DIST.: Colinet, Rouleau 4175 (NFLD). ST. JOHN’S DIST.: Doctor Hill, Fernald, Long & Fogg 1916 (GH, PENN, PH, US). TRINITY NORTH DIST.: George’s Brook, Rouleau 6078 (NFLD). WHITE BAY DIST.: Twin Lakes, Rouleau 6525 (NFLD, US). Northwest Territories: Baralzo Lake, Scoggan & Baldwin 8335 (CAN), 8379 (CAN); Gagnon Lake, Scotter 3162 (CAN); Ingalls Lake, S.C.Z. (CAFB). Nova Scotia: ANNAPOLIS CO.: Karsdale, Johnson (ACAD). CAPE BRETON CO.: Scatari Isl, Ershine 51-1167 (QUK). COLCHESTER co.: Folleigh Lake, Perry (ACAD). CUMBERLAND CO.: Sand R., Schofield & Webster 5592 (ACAD). DIGBY CO.: Central Grove, Fernald & Long 24285 (GH). GUYSBOROUGH CO.: Tor Bay, Eshbaugh 522 (IND). HALIFAX CO.: Fernald, et al. 24284 (GH, PH). HANTS CO.: Mt. Uniacke, Jack 3230 (GH), 3231 (GH). INVERNESS CO.: Mackenzie Mt., Rossbach 4235 (ACAD). KINGS CO.: Caribou Bog, Erskine (ACAD). LUNENBURG CO.: Deep Cove, Smith, et al. 19604 (ACAD, QUK). PICTOU co.: New Glasgow, Dore & Gorham 45-389 (ACAD). QUEENS C0.: Eagle Head, Donly 1009 (ACAD). RICHMOND CO.: Grand R., Smith, et. al. 10110 (ACAD, TRT). SHELBURNE CO.: Barrington, Fernald, Long & Linder 22154 (CAN, GH). VICTORIA CO.: Cape Breton Nat’l. Park, Armstrong 452 (QUK). YARMOUTH CO.: Yarmouth, Pease & Long 22152 (GH, PENN). Ontario: ALGOMA DIST.: Michi- picoten Harbour, Hosie, Harrison & Hughes 1223 (CAN, TRT). CARLETON CO.: Mer Blue, Porsild 7743 (CAN). COCHRANE DIST.: Fraserdale, Kirk 75 (TRT): Timmins, Baldwin 4761 (CAN, TRT). DURHAM C0.: Newtonville, Krug & Purchase 473 (TRT). FRONTENAC co.: Kingston, Beschel 10128 (QUK). GREY CO.: Euphrasia Twp., Jackson (TRT). HALIBURTON CO.: Dorset, Walshe 12 (CAN, TRT). HURON C0.: Wingham, Anderson 724 (TRT). KENORA DIST.: Rich- 348 Rhodora [Vol. 76 mond Gulf, Abbe & Abbe 3383 (CAN, MIN); Las Seul, Baldwin 8739 (CAN, MIN, QUK, TRT); Big Trout Lake, Moir 13 (MINN), 21 (CAN, MIN), 1744 (MIN); Swan Lake, Moir 156 (MIN); Fort Severn, Moir 1337 (CAN, MIN); Black Duck R., Moir 1831 (MIN), 2037 (CAN, MIN, TRT) ; Windigo R., Cringan P-26 (ACAD, TRT). MANITOULIN DIST.: Great Duck Isl, Grassel 6028 (NY). MIDDLESEX CO.: London, Baker 1638 (OAC). MUSKOKA DIST.: Huntsville, Soper & Shields 4927 (TRT), 4967 (TRT); Margaret Lake, Chow 6415 (TRT). NIPISSING DIST.: Algonquin Park, Watson 3967 (TRT). PARRY SOUND DIST.: Axe Lake, Shields & Shields 1640 (TRT). PEEL CO.: Heart Lake, Catling & Sparling (TRT). RUSSELL CO.: Bourget, Heimburger 2612 (TRT). SUDBURY DIST.: Skead, Winterhalder 66114 (CAN). THUNDER BAY DIST.: Bear Trap Lake, Voss 10361 (MICH, TRT). TIMISKAMING DIST.: Hilliard Twp., Baldwin & Breitung 2564 (CAN, SASK, TRT). WATERLOO CO.: Baden, Montgomery 997 (au, OAC). WELLAND CO.: Humberstone, Soper & Dale 3776 (GH, MIN, TRT, US). WELLINGTON Co.: Puslinch, Stroud (TRT). YORK co.: Vandorf, Robb (TRT). Prince Edward Island: KING Co.: Bothwell, Fernald & St. John 11148 (CAN, GH, US, WS). PRINCE CO.: North Point, Erskine & Dore 1080 (ACAD, NY). QUEENS CO.: Brackley Point Rd., Fernald, Long & St. John 7901 (CAN, GH, PH, US, WS). Quebec: ABITIBI CO.: Watson Lake Area, Bentley 58145 (TRT), 5837 (CAN, TRT). BERTHIER CO.: Lanoraie, Scoggan 1538 (CAN). BONAVENTURE CO.: Nouvelle, Collins & Fernald (GY). CHAMBLY CO.: Saint-Bruno, M.-Victoria & R.- Germain 43884 (cM, GH, ND, PH). CHICOUTIMI CO.: Lac Merlac, Brisson 63124 (CAN). GASPE WEST CO.: Mt. Albert, Scoggan 1086 (CAN). GATINEAU CO.: Eardley, Cody & van Rens 11167 (DUKE, GH, MSC, NY, TENN). HULL CO.: King Mt., Zavitz 1107 (TRT). HUNT- INGTON CO.: Covey Hill, M.-Victoria & R.-Germain 43300 (GH). LABELLE C0.: Nominingue, Roy 1167 (wvA), 3650 (ACAD, CONN, NY, RM, US). LAKE ST. JOHN WEST CO.: Peribonka, M.-Victorin 15849 (GH). LEVIS CO.: St. Nicolas, Cayouette & Cayouette 54-89 (TRT). MAGDALEN ISLANDS CO.: Grindstone Isl, Fernald, et. al. 7900 (CAN, PH, WS). MATANE CO.: Montagne Lake, Boivin 787 (PENN). MATA- PEDIA CO.: Sayabee, Scoggan 13600 (CAN). MEGANTIC CO.: Black Lake, Fernald & Jackson 12137 (GH). MONTCALM CO.: Lac Tapani, Pennell 16686 (PH). MONTMAGNY CO.: Lake Monroe, R.-Germain & Courtemanche 454 (US). PAPINEAU CO.: Notre Dame du Laus, C.-Joseph 8508 (ACAD). PONTIAC COo.: Danford Lake, Dore 19994 (TRT). QUEBEC CO0.: Montmorency Fall, Macoun 68560 (am). RIMOUSKI CO.: Father Point, Bartram & Long 581 (PH). SAGUENAY co.: Fort Chimo, Porsild 21873 (CAN); Knob Lake, Harper 3350 (HNH, QUK); Cairn Isl., Abbe, et. al. 3067 (RM): Seven Isl., Robinson 868 (CAN, GH, NY); Mutton Bay, Lewis 582 (CAN). ST. JEAN CO.: Villeneuve, M.-Victorin, et. al. 43047 (MU). SHERBROOKE CO.: Orford, Forest 16058 (CAN, CM, QUK, SASK). TERREBONNE CO.: St. Janvier, 1974] Genus Kalmia — Ebinger 349 Scoggan 314 (CAN). TWO MOUNTAINS CO.: Oka, M.-Victorin 8228 (NY, US). WOLFE C0.: Lac Aylmer, Hamel 13217 (CAN). Saskatche- wan: Athabaska Lake, Lang 188 (Us); Raupp 6175 (CAN, GH, NY), 6657 (GH), 6731 (GH, NY); Bittern Cr., Wheaton (CAFB); Candle Lake, Boivin & Breitung 6157 (NY); Cree Lake, Maini 21 (RM, SASK), 235 (SASK), 236 (CAN, RM, SASK); Crooked R., Breitung 70 (SASK); Cumberland House, Argus 4083 (SASK); Hatchet Lake, Maini 91 (OAC, RM, SASK); Nipawin Prov. Park, Argus 4122 (SASK); Orley, Breitung 268 (ALTA, CAN, NY, US); Prince Albert Nat’l. Park, Fraser (PAC, SASK); Stoney Rapids, Maini 205 (RM, SASK), 275 (SASK), 462 (RM, SASK); White Fox, Fraser (SASK) ; Windrum Lake, Bryenton 166 (CAN). The name Kalmia polifolia was first used by Ludwig (1783), who gave it the common name Canada Laurel or Poley Leaved Laurel, but did not describe it. This name first appears with a valid description a few years later (Wangenheim, 1788). He gives its distribution as extend- ing northward from 45 degrees latitude in America, Nova Scotia and the colder parts of Canada. His description, illustration, and the distribution leave no doubt as to the taxon referred to. Recently, Boivin (1967) suggested that the name be spelled poliifolia to conform to article 73 of the International Code of Botanical Nomenclature (Lanjouw, 1966). The name K. glauca was first used as a synonym of K. polifolia by Ludwig (1783) and later described by Aiton (1789). The type was based on a plant from New- foundland growing at Kew. A specimen in the Kew Her- barium has written on the label “Kalmia glauca Hort. Kews. sp. nov. 1784” and probably represents the type of this name. Kalmia rosmarinifolia was used by Dumont de Courset (1802) to distinguish individuals that have narrow leaves with strongly revolute margins. Later, he listed the name as a synonym of K. glauca (Dumont de Courset, 1811). Kalmia polifolia is found in bogs, swamps, and other wet places, commonly forming a border around ponds and lakes. In bogs, its roots usually form dense mats that extend out over the water. It is also found at higher ele- vations in the mountains of northeastern United States and Canada. This species is the most widely distributed member 350 Rhodora [Vol. 76 of the genus, extending from northeastern Alberta, across Canada to the east coast and south to New Jersey. Its entire range, however, is hard to determine with certainty. There are two reports of its occurrence as far north as the Arctic Circle in the Northwest Territories (Gussow, 1933, and Thieret, 1963). Both of these (Maguse Lake, Gussow 33; Contwoyto Lake, Thieret 9415) are specimens of K. microphylla var. microphylla. The only specimens of K. polifolia seen from the Northwest Territories are from the extreme southern part (Scotter, 1966). In the Great Lakes region this taxon extends as far south as northern Illinois (Fuller, 1944), and along the east coast to New Jersey (Griscom, 1931). This species has also become naturalized in Europe since Scott (1962) reported that it has been col- lected from Flanders Moss, Perthshire, Scotland. Besides leaf size and shape, the character most commonly used by previous authors to separate Kalmia polifolia from K. microphylla is revolute leaf margins. In K. microphylla, the leaf margins are not revolute, or only slightly so, while in K. polifolia the margins are usually strongly revolute. This characteristic is somewhat variable. In about 80% of the specimens examined, the leaf margins were strongly revolute, resulting in a narrow, linear leaf. In the remain- ing specimens, some lacked revolute margins entirely, while in others the younger leaves were revolute while older leaves were not. The variability of this characteristic is responsible, in part, for the confused taxonomic status of K. microphylla. The major distinguishing vegetative character that sep- arates Kalmia polifolia from the related species, K. micro- phylla, is the presence of purple clavate hairs on the leaf midrib. Boivin (1968) appears to be the first to have made use of this character when he separated variety microphylla from variety polifolia. He mentions, however, that west of Alberta the distinction of the two varieties becomes mean- ingless, as most of the British Columbia material is inter- mediate, the leaves being long and narrow like variety poli- folia, but glandless on the midrib. It appears that he was 1974] Genus Kalmia — Ebinger 851 confusing K. polifolia with the western K. microphylla var. occidentalis, which is similar vegetatively, except for the glandless midribs. Clavate glands were found on all speci- mens of K. polifolia examined. These glands vary in size, shape and color, and are more plentiful on some specimens than on others. The glands average about 0.3 mm long, are colorless near the base and become deep purple to black toward the swollen apex. In some specimens, however, they are nearly colorless throughout, and in others barely ex- panded above. Also, their distribution varies, being more common on some leaves than on others. On older leaves, it is not uncommon to find that many of the glands have broken off. The glands on the upper side of the midrib are commonly shorter, darker colored and not as obvious as those on the under side of the leaf. For determining the presence of these glands it is best to look at the developing leaves. Morphologically and genetically Kalmia polifolia (n = 24) is most closely related to K. microphylla (n = 12). The F, of a cross between these two species is a highly viable triploid which has a pollen stainability of 0-15% (Jaynes, 1968a). These similarities suggest that these two taxa have probably become specifically distinct as a result of the last ice age and the resulting development of a tetraploid taxon from a wide ranging ancestral diploid species. Jaynes (1969) suggested that these taxa should be considered sep- arate species based on his cytological and genetic studies. Kalmia polifolia can also be crossed with other members of the genus, but few viable offspring are produced. Kalmia polifolia forma leucantha Schofield & Smith, Canad. Field Nat. 67: 94. 1953. TYPE: Newfoundland: near Hodgewater Line, Trinity South, E. E. Smith and A. C. Smith 1055 (ACAD). i This white flowering form, which is similar to that found in some of the other species of Kalmia, is known from one collection. This single specimen was found in a bog where the typical form was very common. 352 Rhodora [Vol. 76 3. Kalmia latifolia Linnaeus, Sp. Pl. 1: 391. 1753. TYPE: “Habitat in Marilandia, Virginia, Pennsylvania" (LINN 560.1, lectotype, specimen at the top right). Chamaedaphne latifolia (L.) Kuntze, Rev. Gen. Pl. 2: 388. 1891. Kalmia latifolia var. acuminata Raf. Medical Fl. 2: 17. 1830. (nom. nud.). Kalmia latifolia var. alba Raf. Medical Fl. 2: 17. 1830. Kalmia latifolia f. alba (Mouillefert) Rehd. Rhodora 12: 2. 1910. Kalmia latifolia var. arborea Raf. Medical Fl. 2: 17. 1830 (nom. nud.). Kalmia latifolia var. pavarti André, Revue Hort. 60: 541. 1888. Kalmia latifolia f. rubra K. Koch, Dendrologie 2(1) : 152. 1872. Kalmia latifolia var. rubra (K. Koch) Rehd. in Bailey, Cycl. Amer. Hort. 2: 854. 1900. Kalmia latifolia var. ternata Raf. Medical Fl. 2: 17. 1830. Kalmia latifolia var. laevipes Fern. Rhodora 42: 53. 1940. TYPE: Virginia: NORFOLK CO.: dry woods of a *hammock", Great Dismal Swamp, west of Yadkin, Fernald & Long 11101 (GH). Kalmia lucida K. Koch, Dendrologie 2(1): 152. 1872. (as synonym). Kalmia nitida Forbes, Hort, Woburn. 93. 1833. TYPE: unknown, probably a living plant. Much branched shrub to 4 m tall (rarely a small tree) ; branchlets terete, reddish brown, becoming gray, stipitate- glandular to glabrous; leaves alternate (appearing nearly whorled on slow growing twigs), flat, coriaceous, elliptic to elliptic-lanceolate, 4-12 cm long, 1.5-4 (rarely 5) em wide, dark green and glabrous above, light green to reddish brown and with short glandular hairs beneath, apex acute, base narrowly cuneate, midrib puberulent above; petiole 1-3 em long, glabrous to puberulent, sometimes glandular-stipitate; inflorescence a terminal, compound corymb; pedicels 2-4 cm long, puberulent to stipitate-glandular, often floccose, viscid 1974] Genus Kalmia — Ebinger S00 (rarely glabrous), subtended by 2 bracteoles in the axil of a puberulent, lanceolate bract, 1-5 mm long; major inflor- escence branches puberulent, usually stipitate-glandular, to 4 cm long; calyx glabrous to stipitate glandular, green to reddish, 5-8 mm broad, the lobes usually oblong, acute; corolla usually light pink (white to deep red) with purple spots around each anther pocket, usually lightly stipitate- glandular outside, puberulent inside, 1.5-3 cm broad; style 1-1.8 cm long; anthers dark purple to brown, 1.2-1.7 mm long; filaments puberulent; capsule depressed globose, stipi- tate-glandular, 4-7 mm broad, 3-5 mm long; seeds .7-1 mm long, the testa loose and extending slightly past the ends of the seed. Distribution: Mostly rocky or sandy hardwood forests in eastern United States from southern Maine, west through southern New York to central Ohio, south to southern Mis- sissippi, Alabama, and Georgia and northwestern Florida. Common Name: Mountain Laurel, Broad-leaved Laurel, Calico-bush, Spoonwood, Ivy, Mt. Ivy, Big-leaved Ivy, Laurel-leaves, Calmoun. REPRESENTATIVE SPECIMENS Alabama: CALHOUN CO.: Pendergrass (TENN). CHEROKEE CO.: Centre, Clonts 95 (1sC). CHOCTAW CO.: Silas, McDaniel 4218 (VDB). CLARKE CO.: Tallahatta Creek, Kral 19749 (VDB). CLEBURNE CO.: Kral 31843 (VDB). CONECUH CO.: Castleberry, Howell 760 (us). CULLMAN CO.: Cullman, Kral 28911 (VDB). DALE CO.: Newton, God- frey 55477 (TENN). DE KALB CO.: Mentone, Demaree 50521 (TENN). ESCAMBIA CO.: Little Escambia River, Ahles 7229 (MISSA). FRANK- LIN CO.: Russellville, Kral 26321 (VDB). HENRY co.: Headland, Wiegand & Manning 2370 (CU). JACKSON co.: Flat Rock, Kral 34948 (VDB). LEE CO.: Chewacla, Barnes 119 (VDB). MARION CO.: Hackelberg, Kral 20024 (VDB). MARSHALL CO.: Albertville, Harbison 15887 (TENN). MOBILE CO.: Mt. Vernon, Burnham 84 (VDB). PIKE co.: Troy, McDaniel 7830 (vpB). TUSCALOOSA Co.: Lake Nichols, Williams 104 (VDB). WASHINGTON CO.: Bates Creek, Segars 191 (WVA). WINSTON cCOo.: Haleyville, Kral 29476 (vpB). Connecticut: FAIRFIELD CO.: Wilton, Eames 10426 (CONN). HARTFORD CO.: Wind- sor, Ahles 65351 (MASS). LITCHFIELD CO.: Preston, Lumsden (MU). MIDDLESEX CO.: Millers Pond, North 51 (MICH, US). NEW HAVEN CO.: Milford, Fames 2813 (CONN). NEW LONDON CO.: Lantern Hill, Glea- 354 Rhodora [Vol. 76 son (DUKE). TOLLAND CO.: Andover, Travis 2408 (CONN). WINDHAM co.: Crystal Lake, Peterson (cU). Delaware: NEW CASTLE CO.: Arden, Travis 1086 (PENN). SUSSEX CO.: Gumboro, Beaven 241 (DUKE). Florida: ESCAMBIA CO.: McDavid, Goodale (MASS). GADSDEN co.: Sycamore, Godfrey 61925 (Msc). HOLMES CO.: Darlington, Godfrey 55498 (TENN). LIBERTY CO.: Greensboro, Karl 1916 (TENN). OKALOOSA CO.: Laurel Hill, Godfrey 61284 (US). WALTON CO.: De Funiak Springs, Perkins & Hall 2510 (cv). Georgia: CLARK CO.: Middle Oconee River, Duncan & Duncan 4358 (ID, IND, ISC, PENN, RENO, TENN, US). CLAY CO.: Cemochechahee Creek, Thorne 3453 (CU). DADE CO.: Lookout Mt., MeVaugh 9023 (MICH), EARLY CO.: Sawhatchee Creek, Duncan & Thorne 1778 (CU, TENN). EFFINGHAM co.: Savannah River, Harper 1814 (US). GWINNETT CO.: McGuires Mill, Small (US). MORGAN CO.: Hard Labor Creek St. Park, Jones 15456 (VDB). PICKENS CO.: Jasper, Cronquist 4483 (NO, US). RABUN cc.: Darling Spring, J.H.M. E1525 (WVA). RANDOLPH CO.: Cuthbert, ‘Harbison 13729 (TENN). SCREVEN CO.: Newington, Cronquist 5023 (MICH, US, WS). SUMTER CO.: Americus, Harper 523 (US). UNION co.: Vogel St. Park, McVaugh & Pyron 841 (ORE). UPSON co.: Flint River, Browne & Browne, Jr. 6587 (KY). WALKER CO.: Pigeon Mt., Wilson 176 (US). WALTON Co.: Athens, Cronquist 4468 (MICH, NO, US). WHITE CO.: Cleveland, Lovett (DUKE). WHITFIELD CO.: Dalton, Cronquist 5045 (MICH, US, WS). Indiana: CLARK CO.: Borden, Deam 24495 (IND, MIN). CRAWFORD CO.: Riddle, Deam 16497 (IND). PERRY co.: Derby, Ramsey 10876 (IND). Kentucky: BELL co.: Pike Mt. St. Park, Collins 180 (KY). BREATHITT CO.: Robinson Forest, Smith 1726 (KY). CARTER CO.: Cascade Cavern, Gilbert & McCoy 93513 (us). CASEY CO.: Liberty, Wharton 4366 (KY). EDMONSON CO.: Mammoth Cave Nat’l. Park, Pittillo 2452 (Kv). ESTILL CO.: South Irvine, Wharton 2298b (Kv, MICH). FLEMING CO.: Plummer's Mill, Wharton 3873 (KY). FLOYD CO.: Hueysville, Centers & Blair TT (KY). GRAYSON CO.: Clifty Creek, Browne & Browne, Jr. 6494 (KY). HARLAN CO.: Poor Fork, Kearney 182 (ISC, MSC, OS, US). JESSAMINE co.: Camp Daniel Boone, Webb 104 (Ky). KNOTT CO.: Littcarr, Smith 59 (KY). LETCHER CO.: Browne & Browne, Jr. 5400 (KY). LEWIS CO.: Vanceburg, Wharton 5012 (KY, MICH). LINCOLN CO.: Ottenheim, Wharton 4866 (KY, MICH). MCCREARY CO.: Cumberland Falls, Rogers 55 (DUKE, IND, MIN, PENN, TENN, US, WVA). MADISON co.: Berea, Nelson (MICH, MIN, MU). MENIFEE CO.: Frenchburg, MelInteer 2302 (KY). MORGAN CO.: Ezel, Oldfield 57 (KY). POWELL co.: Stamping Ground, Singer (BRY). PULASKI CO.: Mt. Victory, Warden 129 (Kv). TODD CO.: Clifty, McCoy (WVA). TRIMBLE CO.: Bedford, Born (KY). WOLFE CO.: Sky Bridge, McInteer 1055 (KY). Louisiana: WASHINGTON PARISH: Bogalusa, Cocks (NO); Varnodo, Thieret 23481 (DUKE). Maine: CUMBERLAND CO.: Richville, Hyland 444 (MAINE). OXFORD 1974] Genus Kalmia — Ebinger 355 co.: Fryeburg, Hyland 446 (MAINE). PENOBSCOT CO.: Carmel, Harvey 622 (US). SAGADAHOC CO.: Phippsburg, Gordon 592 (MAINE). WASHINGTON CO.: Cherryfield, Hyland 760a (MAINE). YORK CO.: Kennebunk, Hyland 252 (MAINE). Maryland: ALLEGANY CO.: Cum- berland, Richards 128 (MAINE). ANNE ARUNDEL CO.: Camp Whip- poorwil, Grimes (WVA). BALTIMORE C0.: Abingdon, Nieuwland (ND). CAROLINE CO.: Newton, Quidas 37 (MAINE, PENN). CECIL CO.: Elkton, Randolph & Randolph 124 (CU). FREDERICK Co.: Hunting Creek, Earle 1872 (PENN). GARRETT CO.: Grantsville, Stone 14111 (PENN). KENT CO.: Chestertown, Vanatta (CHRB). PRINCE GEORGE co.: Suitland, Meyer 9178 (ISC, WVA). QUEEN ANNES CO.: Chester River, Thatcher 5 (MIN). ST. MARY'S CO.: Patuxent River, Grether 8276 (MICH). TALBOT CO.: Easton, Earle 810 (PENN). WASHINGTON co.: Boonsboro, Tidestrom 724a (US). WICOMICO CO.: Sharptown, Tidestrom 12228 (MICH). WORCESTER CO.: Ferry Creek, Redmond 225 (MICH). Massachusetts: BERKSHIRE CO.: Williamstown, Ames (MICH). BRISTOL CO.: New Bedford, Bartlett 319 (IND, MICH). ESSEX CO.: Newburyport, Mock (NHA). FRANKLIN CO.: Mt. Toby, Elwell (MASS). HAMPDEN CO.: Granville, Seymour 197 (MASS). HAMPSHIRE CO.: Amherst, Elwell (MASS). MIDDLESEX CO.: Concord, Mann (HNH). NORFOLK CO.: Dedham, Greenman 2371 (MIN). PLYMOUTH CO.: Plymouth, Morong 1184 (US). WORCESTER CO.: Brookfield, Markert (MASS). Mississippi: CLARKE CO.: Burstone Cuesta, Jones 12050 (VDB). FORREST CO.: McCallum, Ray, Jr. 6045 (MISSA). GEORGE CO.: Cedar Creek, Demaree 35222 (VDB). HARRISON co.: Gulfport, Bomhard 33 (NO). HINDS CO.: Natchez Trace Park- way, Price (MISSA). JACKSON CO.: Van Cleave, Demaree 82454 (DUKE). JASPER CO.: Rose Hill, Reynolds 96 (MISSA). LAUDERDALE co.: Meridian, Hayes 20 (MISSA). LINCOLN CO.: Boque Chitto, Ray, Jr. 7830a (MISSA). PEARL RIVER Co.: Picayune, Ray, Jr. 6519 (MISSA). STONE CO.: Perkinson, Ray, Jr. 7713 (MISSA). TISHOMINGO co.: Fisherman Ford, Ray, Jr. 7507 (MISSA, VDB). WAYNE CO.: Clara, Graham 118 (MISSA). New Hampshire: BELKNAP CO.: Center Harbor, Hodgdon, et. al. 9507 (NHA). CARROLL CO.: Albany, Pease 25358 (MAINE). CHESHIRE CO.: Fitzwilliam, Hodgdon 7069 (HNA). GRAFTON co.: Hanover, Drew (HNH). HILLSBORO Co.: Manchester, Hodgdon & Kent 7070 (NHA). MERRIMACK CO.: Hooksett, Hall (NHA). ROCK- INGHAM CO.: Windham Center, Harris 430 (NHA). STRAFFORD CO.: Farmington, Lord (NHA). New Jersey: ATLANTIC CO.: Alsion, Crow 126 (MSC). BERGEN CO.: Palisades, Gershoy (CU). BURLINGTON CO.: Pemberton, Palmer 89752 (MICH, MIN). CAMDEN CO.: Gibbsboro, Hynes (PENN). CAPE MAY CO.: Dennisville, Fender 5049 (PENN). CUMBERLAND CO.: Gouldtown, Long 575 (ARIZ, CAN, CU, IND, ISC, KY, NHA, NO, TENN, UARK, WVA). ESSEX CO.: Millburn, Rusby 1710 (MICH). GLOUCESTER CO.: Mantua, Adams 258 (PENN). HUNTERDON co.: Fairmount, Good (CHRB). MIDDLESEX CO.: Spotswood, Laugh- 356 Rhodora [Vol. 76 ridge & Chrysler (CHRB). MONMOUTH CO.: Clarksburg, Fogg, Jr. 14129 (PENN). MORRIS CO.: Longwood, Davidson 764 (CHRB). OCEAN C0.: South Lakewood, MacKenzie 3116 (US). PASSAIC co.: Utter- town Bog, Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6606 (CHRB, PENN). SOMERSET CO.: Willowood estate, Blackburn (CHRB). SUSSEX CO.: Kittatinny Ridge, Davidson 1509 (CHRB). WARREN CO.: Sand Pond, Large & Clausen 1280 (CU). New York: BRONX CO.: Bronxdale, Burnham 428 (CU). CATTARAUGUS CO.: Rock City Park, Muenscher 15841 (CU). CHEMUNG CO.: Susquehana River, Lucy 6616 (MU). COLUMBIA CO.: Ghent, McVaugh 881 (PENN). DELAWARE CO.: Deposit, Muenscher 16244 (cu). DUTCHESS CO.: Clove, Standley & Bollman 12138 (US). GREENE CO.: Winter Clove Mt., Gershoy (CU). NASSAU CO.: Woodbury, Stewart (CU). ONEIDA CO.: New London, House 27859 (CU). RICHMOND CoO.:. Haverstraw, Fosberg & Fosberg 14897 (PENN). ROCKLAND CO.: Ramapo, Gershoy (CU). SCHUYLER CO.: Cayuta Lake, Muenscher & Brown 21703 (cU). STEUBEN CO.: Painted Post, Muenscher 15842 (CU). SUFFOLK CO.: Southampton, St. John 2852 (CU). SULLIVAN CO0.: Camp Ranachqua, Weber 272 (isc). TIOGA CO.: Spencer, Hames & McDaniels 4765 (CU). TOMPKINS co.: Enfield, Palmer 928 (IND). ULSTER C0.: Lake Minnewaska, Gillis 3090 (MSC). WESTCHESTER CO.: Ossining, Gershoy (CU, MU). North Carolina: ALEXANDER CO.: Keever 21 (DUKE). BUNCOMBE co.: Asheville, McCarthy (US). BURKE CO.: Shortoff Mt., Wilbur 7004 (VDB). CALDWELL CO.: Granite Falls, Randolph & Randolph 1085 (CU). CHATHAM CO.: Costello 14 (DUKE). CLAY CO.: Perry Gap Road, Wilbur 1275 (MIN, DUKE). CUMBERLAND CO.: Rockfish Creek, Correll & Rankin 9053 (DUKE). DURHAM CO.: Catsburg, Wilbur 12230 (DUKE). EDGECOMBE CO.: Tar River, Fox & Whitford 1746 (DUKE). FORSYTH CO.: Schallert 1092 (DUKE). GASTON CO.: Kings Mt., Oosting 1646 (DUKE). GRAHAM CO.: Thunderhind Mt., Beardslee & Kofoid (MU). GRANVILLE CO.: Tar River, Correll 516 (DUKE, MASS). HARNETT CO.: Bunnlevel, Laing 1104 (MIN). HAY- woop co.: Waynesville, Standley 5401 (US). HENDERSON CO.: Hen- dersonville, Cottam 11140 (UT). JACKSON CO.: Wesner Bald, Ramseur 171 (MAINE). JOHNSTON CO.: Neuse River, Fox & Whitford 1293 (DUKE, WS). LEE CO.: Deep River, Beard 405 (KY). MCDOWELL CO.: Rogers 8358 (DUKE, US). MACON CO.: Mt. Satula, Harbison 1848 (TENN). MADISON C0.: Hot Springs, Wehmeyer 533 (MICH). MIT- CHELL CO.: Roan Mt., Cannon 143 (CU). MOORE CO.: Carthage, Harriot (NO). ORANGE Co.: Hillsboro, Correll 325 (DUKE). POLK co.: Columbus, Townsend 236 (CU). ROCKINGHAM. CO.: Smith River, Correll & McDowell 10695 (DUKE). SCOTLAND CO.: Laurel Hill, Wiegand & Manning 2369 (cu). STANLY CO.: Badin, Fox, Godfrey & Boyce 1427 (DUKE, TENN, US, WS). STOKES CO.: Sauratown Mt., Radford 34665 (1sc). SURRY co.: Pilot Mt., Williams 296 (DUKE). SWAIN CO.: Andrews Bald, Bruhn 4 (TENN). TRANSYLVANIA CO.: 1974] Genus Kalmia — Ebinger 357 Rosman, Morley 891 (18C). UNION CO.: Rocky River, Ahles & Haesloop 31521 (UARK). WAKE CO0.: Raleigh, Godfrey 3765 (US). WATAUGA co.: Boone, Ahles & Duke 43728 (PAC). WAYNE CO.: Mt. Olive, Rose 36 (MASS). WILKES CO.: Wyatt 832 (ARIZ, BRY, NHA, VDB). Ohio: ADAMS CO.: Mineral Springs, Herrick (0S). ATHENS CO.: Athens, Kellerman (0S). COLUMBIANA CO.: Sanders (OS). COSHOCTON co.: New Castle Twp., Selby (os). CUYAHOGA Co.: Bayville, Biebel (OS). FAIRFIELD C0.: Sugar Grove, Potzger 2969 (ND). GALLIA CO.: Jones (0S). GUERNSEY CO.: Liberty Twp., Thomas (0S). HOCKING co.: Clear Creek, Stuckey 2912 (0s). HOLMES CO.: Killbuck Twp., Herrick (08S). JACKSON CO.: Liberty Twp., Bartley & Pontius (os). JEFFERSON C0.: Amsterdam, Cusick (08). KNOX co.: Pipesville, Moldenke 12186 (0S). LAWRENCE CO.: Hanging Rock, Werner (os). LICKING C0.: Blackland Gorge, Stuckey 6603 (os). MEIGS CO.: Columbia Twp., Jones (0S). MONROE CO.: Adams Twp., Thomas (OS). MORGAN CO.: Bur Oak St. Park, ‘Herrich (0S). MUSKINGUM co.: Laurel Hill, Reed (0s). PERRY CO.: Jonathan Creek, Hicks & Dambach (os). PIKE CO.: Chimney Rocks, Crowl (0S). ROSS CO.: Tar Hollow, Crowl (os). SCIOTA CO.: Cooper Head Fire Tower, Demaree 10664 (os). TUSCARAWAS CO.: Wayne Twp., Brown (os). VINTON C0.: Lake Hope St. Park, Stuckey 6407 (0S). WASHINGTON co.: Vincent, Walpole 1129 (BLH). Pennsylvania: ADAMS CO: Birch River, Tanger 4434 (PENN, WS). ALLEGHENY C0.: Tom's Run Rd., Henry & Buker (PENN). ARMSTRONG CO.: Sherrett, Henry (PENN). BEDFORD CO.: Tatesville, Berkheimer 982 (PENN). BERKS CO.: Bech- telsville, Fender 1022 (PENN). BLAIR CO.: Blue Knob, Westerfeld 16177 (PAC). BRADFORD Co.: Canton, Westerfeld 907 (PAC, PENN). BUCKS CO.: Janney, Long 76067 (PENN). BUTLER CO.: Clintonville, Brubaker 2537 (PENN). CAMBRIA CO.: Johnstown, Tinan (os). CAMERON CO.: Sinnemahoning Creek. Fogg, Jr. 11555 (PENN). CARBON CO.: Little Gap, Fogg, Jr. 11900 (MIN, PENN). CENTRE CO.: Yarnell, Keener 2581 (PAC). CHESTER CO.: Harmonvville, Fender 1401 (PENN). CLARION CO.: Callensbure, Wood. Jr. 2170 (PENN). CLEARFIELD CO.: Troutville, Ehrle 2475 (PAC). CLINTON CO.: Boone- ville, Wahl 3198 (Pac). COLUMBIA co.: Mill Grove, Pohl 2554 (PENN). CUMBERLAND CO.: Shipnensbere, Saxe, Jr. 36 (PENN). DAUPHIN Co.: Pillow. Berkheimer 12883 (PAC, PENN). DELAWARE co: Darby, Baker (Pac). ELK CO.: Medix Run, McClelland (PENN). FAYETTE CO.: Wvymps Gan, Bwinger (PENN). FOREST CO.: Brooks- ton. Shields D-2157 (PAC). FRANKLIN CO.: Mercersburg, Kriner 104 (PAC). FULTON CO.: Deneen Gap, Westerfeld 7798 (Pac. PENN). GREENE CO.: Jefferson. Wherry & Bell (PENN). HUNTINGDON CO.: Maddensville, Westerfeld 9823 (PAC). JEFFERSON CO.: Sieel, Wahl 10944 (PAC). JUNIATA CO.: Cross Keys, Westerfeld 14535 (PAC). LACKAWANNA CO.: Montdale, Adams 3542 (PENN). LANCASTER CO.: Collins, Brubaker 1521 (PENN). LAWRENCE CO.: McConnells Mills, 358 Rhodora [Vol. 76 Russell 1582 (PENN), LEBANON CO.: Cold Spring, Travis 1170 (PENN). LUZERNE CO.: Mountaintop, Adams 3484 (PAC), LYCOMING co.: Cedar Run, Wahl 19261 (PAC). MIFFLIN CO.: Standing Stone Creek, Westerfeld 3961 (PAC). MONROE CO.: Scotrun, Niering 1214 (PAC). MONTGOMERY CO.: Audubon, Wherry (PENN). NORTHAMPTON co.: Wind Gap, Schaeffer, Jr. 18252 (PENN). NORTHUMBERLAND C0.: Sunbury, Wagner 8987 (PENN). PERRY CO.: Bloomfield, Adams & Thebes 1232 (PENN). PIKE CO.: Porter’s Lake, DePue 136 (PAC). SCHUYLKILL C0.: Delano, Wagner 3227 (PENN). SNYDER CO.: Paxton- ville, Wade 783 (PENN). SOMERSET CO.: Jennerstown, Fogg, Jr. 15258 (PENN). SULLIVAN CO.: Laporte, Westerfeld 381 (PAC, PENN). SUSQUEHANNA CO.: North Knob, Adams 3612 (PENN). TIOGA CO.: Mainesburg, Fender 1537 (PENN). UNION CO.: Laurelton, Ludwig 1521 (PAC). VENANGO CO.: Franklin, Wahl 12633 (PENN). WARREN co.: Irvington, Moldenke 17479 (ND). WAYNE CO.: White Mills, Adams 4624 (PENN). WESTMORELAND CO.: Ligonier, Matthews 2399 (PAC). WYOMING CO.: Crane Swamp, Stone (PENN). YORK CO.: Dillsburg, Hoover 3078 (PAC). Rhode Island: NEWPORT CO.: Tiverton, Mearns 211 (US). PROVI- DENCE CO.: Pascoag, Myers 350 (WVA). WASHINGTON CO.: Kingston Hill, Reynolds 649 (HNH). South Carolina: ANDERSON CO.: Carpen- ter’s Mill, Davis (MICH, MIN, UT). GREENVILLE CO.: Saluda River, Freckman 1239 (ISC). LEXINGTON CO.: Batesburg, McGregor 297 (US). MARLBORO CO.: Little Pee Dee River, Radford 12494 (TENN). OCONEE CO.: Whitewater, Davis 392 (MASS). PICKENS CO.: Table Rock, Rodgers 540 (DUKE). SPARTANBURG CO.: Campobello, Erdman 973 (BRY). SUMTER CO.: Poinsette St. Park, Holdaway (DUKE). Tennessee: ANDERSON CO.: Coal Creek, Kearney (MIN, OS, US). BLOUT CO.: Cades Cove, Wallace & Jennison 1621 (TENN). CAMP- BELL CO.: Jellico, Lamb & Williams (TENN). CHEATHAM CO.: Mar- rowbone Creek, De Selm 1258 (TENN). CLAIBORNE CO.: Lone Mt. Ridge, Kelley 2459 (TENN). COFFEE CO.: Old Stone Fort, Blum 3732 (VDB). CUMBERLAND CO.: Ozone, Demaree 47496 (KY, NO, VDB). DAVIDSON CO.: Nashville, Hubbard 9112 (IND). DE KALB CO.: Smith- ville, De Selme 173 (TENN). DICKSON CO.: Sycamore School, Jobe (TENN). FENTRESS CO.: Clear Fork, Sharp 25648 (TENN). FRANKLIN co.: Elk River, Sharp & Adams 10785 (TENN). GRAINGER CO.: Blaine, Hodges 20 (TENN). GREENE CO.: Horse Creek Rd., Sherman & Sharp 22280 (TENN). GRUNDY CO.: Mont Eagle, Stifler (CU). HAMILTON co.: Lookout Mt., Scribner (US). HARDIN CO.: Kral 36161 (VDB). HAWKINS CO.: Clinch Mt., Wolfe 19180 (TENN). HOUSTON CO.: Erin, Palmer 17610 (vs). JOHNSON CO.: Barclay 12B (TENN). KNOX CO.: New Hopewell, Sharp 40454 (TENN). LAUDERDALE CO.: Summertown, Postal (IND). LAWRENCE C0.: Shoal Creek, Sharp, Adams & Felix 11058 (TENN). LEWIS CO.: Trace Creek, Sharp, Adams & Felix 10065 (TENN). MARION CO.: Fosters Falls, Webb 368 (TENN). 1974] Genus Kalmia — Ebinger 359 MONROE CO.: Skona River, Galyon & Sharp 21603 (TENN). MORGAN CO.: University Forest, Shanks 3141 (TENN). POLK CO.: Ocoee River, Wilson 3726 (TENN). PUTNAM CO.: Monterey, Norris & Shanks 2475 (TENN). RHEA CO.: Spring City, James (08). ROANE C0.: Caney Creek, Shanks 22078 (TENN). ROBERTSON CO0.: Pleasant View, Shanks, Woods, & Cooley 14776 (TENN). SEVIER CO.: Pittman Center, Jen- nison 3350 (TENN). UNICOI CO.: Rich Mt., Price 658 (DUKE). UNION co.: Norris Lake, Kelley (TENN). VAN BUREN CO.: Spencer, Shanks 1288 (TENN). WASHINGTON CO.: Cherokee Mt., Sharp 3819 (TENN). WAYNE CO.: Butler Creek, Shanks, Woods & Cooley 9156 (TENN). WHITE CO0.: Caney Fork, Channell 8956 (VDB). WILLIAMSON CO.: Ferndale, Major 6 (vpB). Vermont: BENNINGTON C0.: Pownal, Eggleston (HNH, US). Virginia: ALBEMARLE CO.: Miller School, Tinsley (NMC). AUGUSTA CO.: Stuarts Draft, Churchill (Msc). BLANO CO.: Suiter, Kral 10453 (VDB). CRAIG CO.: Newcastle, Fogg, Jr. 13211 (PENN). FAIRFAX CO.: Great Falls, Bartlet 1783 (IND). GILES CO.: Salt Pond Mt., Fogg, Jr. 12688 (PENN). GREENE CO.: Bear Fence Mt., Fosberg 17205 (PENN). ISLE OF WIGHT CO.: Smith- field. Wise (KY). JAMES CITY C0.: Gordon Creek, Fernald & Long 11393 (DUKE). KING GEORGE CO.: King George, Muller 3787 (ND). LOUDOUN CO.: Ashburn, Pratt (MU). MADISON CO.: Old Ray Mt., Fosberg 16909 (PENN). MONTGOMERY CO.: Blacksburg, Musselman 645 (UWM). NANSEMOND CO.: Nansemond River, Fernald & Long 13719 (US). NELSON CO.: Hall 3755 (VDB). NORFOLK CO.: Great Dismal Swamn. Fernald & Long 12147 (TENN). PITTSYLVANLA CO.: Smith Mt., Zobel (DUKE). ROANOKE CO.: Surarloaf Mt., Ebinger 4041 (MU). ROCKBRIDGE CO.: G. Washington Nat’l. Forest, Hall 3716 (VDB). ROCKINGHAM CO.: Shenandow Forest, Eggleston 18624 (us). RUSSELL CO.: Clinch Mt., Kral 23850 (VDB). SMYTH CO.: Pond Mt., Small (msc, US). WISE CO.: Norton, Pennell 11813 (Isc). West Virginia: BARBOUR CO.: Nestorville, Core 5922 (WVA). BERKELEY co.: Back Creek. Core 5847 (WVA). BRAXTON CO.: Little Birch, Harris (WVA). CALHOUN CO.: Pink, Harris (WVA). CLAY CO.: Clay, Core 6372 (WVA). DODDRIDGE CO.: West Union, Bartholomew (WVA). GRANT CO.: Dolly Sods Mt., Davis & Davis 7129 (WS). GREENBRIER co.: Muddy Creek Mt., Fox (WVA). HAMPSHIRE CO.: Ice Mt., Frye 1269 (WVA). HARDY CO.: North Mt. Core & Ludwia 399 (WVA). HARRISON CO.: Duck Creek, Martin 327 (WVA). JEFFERSON CO.: Charlestown, Harper (CU). KANAWHA CO.: Nitro. Greenlee (WVA). MCDOWELL CO.: Panther, Music (WVA). MARION CO.: Pinesville, Bodola (WVA). MARSHALL CO.: Calis, Bartholomew 1941-217 (WVA). MERCER CO.: Pinnacle Rock, Fox (WVA). MINERAL CO.: Patterson Creek, Brown (WVA). MINGO CO.: Chattaroy, Bowen (WVA). MONONGALIA CO.: Cassville, Watkins (WVA). MONROE CO.: Wayside, Fox (WVA). MORGAN CO.: Langent, Core 5739 (WVA). NICHOLAS CO.: Calvin, Creasy (WVA). PENDLETON CO: Seneca Rocks, Clarkson 2001 360 Rhodora [Vol. 76 (WVA). POCAHONTAS CO.: Droop Mt., Ludwig 77 (WVA). PRESTON co.: Pisqah Rd., Meyer 247 (HNH, PENN, TENN, WVA). RANDOLPH co.: Roaring Plains, Core 3171 (WVA). RITCHIE CO.: Cairo, Good- win 251 (WVA). SUMMERS C0.: Barger Springs, Boone (WVA). TAYLOR CO.: Wickwire Run, Core 5475 (WVA). TUCKER CO.: Black- bird Knob, Gibson 1039 (WVA). UPSHUR C0.: Bucklin, Pollock (MIN, US). WAYNE CO.: Hurricane Creek, Lycan 59 (WVA). WEBSTER CO.: Camp Caesar, Fox (WVA). WETZEL CO.: Smithfield, Wagner (wva). WIRT CO.: Standingstone Creek, Bartholomew W-5007 (WvA). Mountain laurel commonly forms dense thickets in rocky and sandy forests throughout most of its range, particularly where there are openings in the canopy. It is also found in pastures and open fields and commonly forms thickets at the edge of roads. A detailed map of the known range of Kalmia latifolia is given by Kurmes (1967) who considers the report of this species from Ontario and New Brunswick to be an error, as is the report from the valley of the Red River in Arkansas (Britton, 1908). The report of this spe- cies from Canada probably originates from Pursh (1814), and it has been perpetuated by Brunet (1867), Sargent (1893), and more recently by Munns (1938). There is no evidence that this species is native to any part of Canada. It is possible that these early reports were based on culti- vated plants or large-leaved specimens of K. augustifolia (Macoun, 1884). Presently the species is known from south- ern and southeastern Maine (Fernald, 1911; Lamson- Scribner, 1891; Norton, 1913, 1935; and Perkins, 1935), southern New Hampshire (Hodgdon, 1955) and southern Vermont (Deane, 1899). The ecological requirements of Kalmia latifolia in the deciduous forests of eastern United States have been exam- ined by a few authors. Kurmes (1961) studied the ecology of mountain laurel in southern New England and deter- mined its exact range from herbarium material and field observations. The relationship of this species to water run-off and the water-holding capacity of the soil was studied by Colvin and Eisenmenger (1943) and Johnson and Kovner (1956). Studies have also been made of the rate of water-vapor loss from its leaves (Meyer, 1927), the 1974] Genus Kalmia — Ebinger 361 seasonal variation in the moisture content of its leaves (Reifsnyder, 1961) and seasonal changes in cold resistance (Parker, 1961). The competition of mountain laurel with tree seedlings and sprouts has also been studied. Wahlen- berg and Doolittle (1950) found that under normal condi- tions in the southern Appalachian mountains K. latifolia forms dense thickets that preclude any natural regeneration of timber trees. Their studies indicate that partial clearing of these thickets followed by planting of desirable tree spe- cies is an effective method of forest regeneration. They also found that mountain laurel has a mean annual height growth of 0.4 feet and a lateral growth of 0.3 feet. Kalmia latifolia is usually a tall, spreading shrub that throughout most of its range rarely exceeds a height of 12 feet. In the fertile valleys between the Blue Ridge and the Alleghany Mountains, however, this species may attain the size of a small tree. Engelmann, Canby, and Gray (1877) observed a number of large individuals growing in the bottom of a dell, back of Caesar’s Head, on the ex- treme western border of South Carolina. One of the trunks measured 4 feet 11/4, inches in circumference about 1 foot above the ground while another in the same area was 3 feet 4 inches above the first fork, and at 1 foot above the ground was 4 feet 4 inches in circumference. Later, Sargent (1893) records its maximum size as 30-40 feet tall with a diameter of 18-20 inches. The largest specimen on record, as listed by the American Forestry Association (1955), is a plant from the Great Smoky Mountains National Park with a circumference of 3 feet 6 inches, a height of 25 feet, and a spread of 45 feet. This specimen is probably located at Forney Ridge, Swain County, Tennessee (Barksdale & Jen- nison 1285). Besides the numerous studies concerned with the poison- ous properties of mountain laurel, some early attempts were undertaken to determine what other substances were pres- ent. These early studies reported the presence of arbutin, gums, and tannins (Kennedy, 1875), bark tannins (De- Graffe, 1896) and fat deposits (Stanley, 1931). More re- 962 Rhodora [Vol. 76 cently the effects of herbicides on Kalmia latifolia have been studied. Egler (1947, 1948) found that 2,4-D has no permanent effect, causing only temporary stunting and cur- vature of the foliage. Later, Egler (1949, 1950) found that a weak solution (.25%) of a 2,4-D and 2,4,5-T combination causes temporary cessation of growth, but a strong solution (596) is necessary to ensure death. The synonomy of mountain laurel is not very extensive. Most of the names proposed are for forms and varieties which are the result of the extreme variation that exists in flower color, leaf shape and size, plant size and pubescence. In 1940, Fernald described a variety (laevipes) of Kalmia latifolia from the coastal plain on southeastern United States. He considered this variety in its extreme develop- ment as having pedicels that are “quite smooth or with the merest suggestion of sessile glandular spots". In typical K. latifolia, in contrast, the pedicels are densely floccose and viscid with stalked glandular hairs. Many individuals with few or no glands on the pedicels are found in Indiana and Ohio, as well as on the coastal plain from Virginia to Louisiana. In some populations specimens with both gland- less and glandular pedicels are found. Also, Radford, Ahles, and Bell (1968) mention that this condition is very variable, with a tendency for pedicels to be floccose on plants in the mountains and less glandular on the coastal plain. Because of this variation it appears best to consider these individuals as an extreme that occurs in the southern and western parts of the range of K. latifolia. A total of 7 forms of Kalmia latifolia have been recog- nized by Rehder (1910, 1945, 1949). Many of these were originally described as varieties, but that classification is not appropriate for these taxa in either the horticultural or botanical sense. It is possible that these taxa could be treated as cultivars as done by Holmes (1956) and Dudley (1967), but most are better treated as forms as defined by Davis and Heywood (1963). The genetic evidence indicates that most of these taxa are not unique clones, but rather are sporadic variants distinguished by a single or a few 1974] Genus Kalmia — Ebinger 363 linked characters, without a distinct distribution. Also, they do not represent the normal extremes of most popula- tions, but are the result of the repeated association of re- cessive genes that are of a limited occurrence in some popu- lations. Of the 7 forms recognized by Rehder, 5 fit the criteria of a form as described above. The remainder, both variations in flower color (alba and rubra), unlike the forms, are not aberrant or disjunct, but are connected to the typical light pink flower by intermediate color types. Kalmia latifolia f. angustata Rehd. Jour. Arnold Arb. 26: 481. 1945. TYPE: New Jersey: CAPE MAY CO.: Dennis town- ship, H. A. Scribner (GH). Kalmia latifolia B. salicifolia Forbes, Hort. Woburn. 93. 1833. (nom. nud.). This very rare foliage form has narrowly oblanceolate to linear leaves that are 4-8 cm long and 4-10 mm wide. Liv- ing material of the type is cultivated at the Arnold Arbore- tum (Rehder, 1945). The only specimens of f. angustata seen by the author are cultivated plants. Common name: Willow-leaved Mountain Laurel. Kalmia latifolia f. fuscata (Rehd.) Rehd. Rhodora 12: 2. 1910. Kalmia latifolia var. fuscata Rehd. Moller's Deutsch. Gart. Zeit. 18: 578. 1903. TvPE: New York: Canaan Four Corners, Dr. Childs (GH). Kalmia latifolia var. coronata Gray, Amer. Nat. 2: 324. 1868. TYPE: Massachusetts: woods near Framingham, James Parker (GH). Kalmia latifolia var. maculata Raf. Medical Flora 2: 17. 1830. TvPE: unknown. This form is distinguished by white to pink flowers that have a heavily pigmented, usually continuous, brownish- purple or cinnamon band on the inside of the corolla at the level of the anther pockets. This band breaks up into brownish dots toward the base and the margin of the corolla. It also shows through to the outside of the bud giving it a muddy appearance. 364 Rhodora [Vol. 76 Some variation does exist in the size, shape and color of the band with an interrupted band existing in some of the specimens examined. According to Jaynes (personal com- munication) controlled crosses with specimens showing this banded characteristic strongly suggests that the character is determined by a single dominant gene. Common name: Banded Mountain Laurel, Crowned Moun- tain Laurel. SPECIMENS EXAMINED Alabama: near Citronella, Milligan (US). Connecticut: near Chaplin, Jaynes (EIU). Massachusetts: near Framingham, Parker (GH); Wales, Clark (MASS); Guterman (MASS). New York: Brew- ster, Jaynes 240 (EIU). North Carolina: Highlands, Anderson (US). Ohio: Holden Arboretum, Mentor, Jaynes 94 (EIU). Pennsylvania: nursery in Stroudsburg, Jaynes 168 (EIU). Kalmia latifolia f. myrtifolia (Bosse) K. Koch, Dendrol- ogie 2(1) : 153. 1872. Kalmia latifolia var. myrtifolia Bosse, Vollst. Handb. Blumengart. 2: 348. 1841. Kalmia myrtifolia Andre, Revue Hort. 55: 10. 1883. TYPE: unknown, probably a living plant. Kalmia latifolia f. minor K. Koch, Dendrologie 2(1) : 153. 1872. TYPE: unknown, probably a living plant. Kalmia latifolia var. nana Lavallée, Arbor. Segrez. 159. 1877. (nom, nud.); Mouillefert, Traité Arb. Arbrisse 2: 1027. 1897. TYPE: unknown, probably a living plant. This form is in all respects a miniature mountain laurel, being compact and slow-growing, rarely exceeding a height of 3 feet. In all specimens examined the leaves are much smaller than in typical Kalmia latifolia, averaging 2-4 cm long and about 1 cm wide. Also, flower size and length of the internodes are approximately 1/3 to 1/2 normal. Recent evidence (Jaynes, personal communication) suggests that this dwarf condition is controlled by a single recessive gene. According to Dudley (1967) this form has been grown since 1840 and is occasionally found in gardens where space is limited. Though occasionally mentioned in the early horticulture literature of Europe, the first reference to this form in North American literature appears to be by 1974] Genus Kalmia — Ebinger 365 Sprague (1871). The only report of wild plants of f. myrti- folia appeared in 1895 (Anonymus) which, according to the author, were identical to the cultivated dwarf form. Common name: Miniature Mountain Laurel, Dwarf Mountain Laurel. SPECIMENS EXAMINED Connecticut: Portland, Jaynes 25 (EIU). Massachusetts: Arnold Arboretum, Hornby (MASS). Kalmia latifolia f. obtusata (Rehd.) Rehd. Rhodora 12: 2. 1910. Kalmia latifolia var. obtusata Rehd. Móller's Deutsch. Gart. Zeit. 18: 577. 1903. TYPE: Connecticut: near Pom- fret, Bowditch. Kalmia latifolia var. ovatifolia Raf. Medical Flora 2: 17. 1830. (nom. nud.). This foliage form has leaves that are oval to oblong- obovate and rounded at both ends. In the specimens exam- ined the leaves average 4-7 cm long and 2.5-4 cm broad, are apiculate, and have short petioles that rarely exceed 1 cm in length. According to Rehder (1910) this form is slow growing, forms a compact bush, and has been cultivated at the Arnold Arboretum since 1886 when it was propagated from the type plant. Dudley (1967) suggested that the cultivar ‘Ovata’ which is available from some European sources is a synonym of this variant. Common name: Hedge Mountain Laurel. SPECIMENS EXAMINED Connecticut: Riverton, Jaynes 291 (EIU). Massachusetts: 1 mi. east of Ashby, Ahles 65185 (MASS); Arnold Arboretum, Wyman 2854 (MSC), Jaynes 238 (EIU). Kalmia latifolia f. polypetala (Nickolson) Beissner, Schelle, & Zabel, Handb. Laubholz-Benennung 386. 1908. Kalmia latifolia var. polypetala Nicholson, Hand-list Trees Shrubs Arb. 2: 49. 1896. Kalmia latifolia f. polype- tala (Nicholson) Rehd. Rhodora 12: 1. 1910. TYPE: Mas- sachusetts: near South Deerfield, Miss M. Bryant (GH). Kalmia latifolia var. monstruosa Mouillefert, Traité Arb. 366 Rhodora [Vol. 76 Arbrisse 2: 1027. 1897. TYPE: Unknown, probably a living plant. This form has the corolla deeply divided into 5 narrow to fairly broad, strap-shaped petals. In some specimens these petals are extremely narrow and thread-like due to the revolution of the margins. Gray (1871) reports that these narrow petals resemble filaments and are sometimes tipped with imperfect anthers. More commonly the petals are broader, and a few specimens have been found in which the individual flowers appear much like apple blossoms. Other variations include an apetalous type which has been observed in some nursery stock from Massachusetts, and a cultivar named ‘Bettina’ (Dudley, 1967) in which the corolla is deeply lobed. According to Jaynes (personal communica- tion) the inheritance data fits the hypothesis that the genetic control of the polypetalous condition is by a single recessive gene. This form was first described by Gray (1871) and later by Sargent (1890), while Bean (1897) mentioned that this unusual form flowers each year at the Kew Gardens. It was given the name var. polypetala by Nicholson (1896) and the name var. monstruosa by Mouillefert (1897), both based on Sargent's description of 1890. This taxon was first considered as a form by Beissner, Schelle, and Zabel (1903) who use the name polypetala and cite Sargent as the author. This form appears to occur most commonly in Massachu- setts. Stone (1909) reported it growing at the edge of a road in Leverett, not far from Mt. Toby, while Britton (1925) reports that Dr. Stone has found it from three dif- ferent stations, all on or near Mt. Toby. Mt. Toby is close to South Deerfield, the type locality of this form. Common name: Feather Petal Mountain Laurel. SPECIMENS EXAMINED Massachusetts: Arnold Arboretum, Wyman 2458 (Msc); Leverett, Mt. Toby, Henry (MASS), Osmun (MASS), Stone (MASS), 67570 (Mass). North Carolina: Danbury, Hanging Rock State Park, Rogers (EIU). 1974] Genus Kalmia — Ebinger 367 4. Kalmia angustifolia Linnaeus, Sp, Pl. 1: 391. 1753. Type: “Habitat in Pensylvania, Nova Caesarea, Nove- boraco" (LINN 560.2, lectotype, specimen on the right). Chamaedaphne angustifolia (L.) Kuntze, Rev. Gen. PLA 388. 1891. Much branched shrub to 1.5 m tall; branchlets terete, reddish-brown, glabrous to pubescent and stipitate-glandu- lar, usually ascending; leaves in whorls of 3 (rarely a few alternate or opposite), slightly coriaceous, evergreen, flat, oblong to elliptic-lanceolate, 2-5 (rarely 7) cm long, 0.5-2 cm wide, usually lightly puberulent above with short white hairs, sometimes glabrate with age, midrib puberulent above, glabrous to puberulent beneath, sometimes stipitate- glandular; petioles to 1 (rarely 1.5) cm long, usually pu- berulent; inflorescence of numerous corymbiform racemes from the axils of last year’s leaves; pedicels 0.5-2 cm long, puberulent and stipitate-glandular, subtended by 2 bracte- oles in the axil of a lanceolate bract, 1-6 mm long; main axis of the inflorescence less than 1.5 cm long; calyx pu- berulent outside, glabrous within, usually green, sometimes the tip reddish, or reddish throughout, 3-6 mm broad, the lobes usually acuminate; corolla reddish-purple to pink (rarely white), puberulent outside, puberulent toward the base within, 6-13 mm broad; anthers purplish-brown, .5-1 mm long; filaments puberulent on the lower half; capsule depressed globose, 3-5 mm broad, 2-3.5 mm long, puberu- lent and stipitate-glandular; seed stramineous, 0.6-1 mm long, and testa loose and extending past the ends of the seed. Chromosome number n — 12. Kalmia angustifolia is a common species in eastern North America. It is found from northern Ontario, Quebec and Labrador, through the Great Lakes region, and south along the east coast of the United States to the Carolinas. It is primarily a plant of bogs and other wet, open areas, but is also found in open woods and as a weed in open areas, some- times forming dense thickets. Genetically this species is most closely related to Kalmia latifolia. When K. angustifolia is the male parent the cross 368 Rhodora [Vol. 76 is not successful because the sheep laurel pollen will not grow down the style of mountain laurel (Jaynes, 19682). The reciprocal cross, however, resulted in about 18% of the flowers producing capsules which contained some viable seed. These resulting interspecific hybrids were intermedi- ate to the parental types, although leaf arrangement was primarily alternate, as in K. latifolia. Jaynes (1968a) also reported that K. angustifolia could be crossed with both K. hirsuta and K. polifolia. Both crosses produced mostly weak, yellow-green individuals that died within the year. A few individuals lived longer, and some produced flowers with sterile pollen. Small (1903, 1914) considered this taxon as two separate species, a procedure not followed by subsequent authors. Both Fernald (1950) and Gleason (1952) considered this complex as being one species with two fairly distinct vari- eties, a view that has recently been followed by Radford, Ahles, and Bell (1968). Considering the genetic and mor- phological similarities of these taxa it is more realistic to follow the latter view and consider it as a single species with two fairly distinct varieties. KEY TO VARIETIES l. Calyx densely stipitate-glandular; mature leaves not puberulent beneath. ............ 4a. var. angustifolia. l. Calyx lacking stipitate-glands, or nearly so; mature leaves densely puberulent beneath. . 4b. var. caroliniana. 4a. Kalmia angustifolia L. var. angustifolia. Kalmia angustifolia B carnea Ait. Hort. Kew. 2: 64. 1789. Kalmia angustifolia var. hirsuta Jager, Ziergeholze 278. 1865. Kalmia angustifolia f. hirsuta Voss, Blumengart. 1: 591. 1894. (without basonym). Kalmia angustifolia var. lucida Lavalée, Arbor. Segrez. 159. 1877. (nom. nud.). Kalmia angustifolia f. lucida Beissner, Schelle, & Zabel, Handb. Laubholz-Benennung 386. 1903. (nom. nud.). 1974] Genus Kalmia — Ebinger 369 Kalmia angustifolia var. minima Bosse, Vollst. Handb. Blumengart. 2: 347. 1841. Kalmia angustifolia var. nana Bosse, Vollst. Handb. Blumengart. 2: 347. 1841. Kalmia angustifolia f. nana Voss, Blumengart. 1: 591. 1894. (as snyonym). Kalmia angustifolia B ovata Pursh, Fl. Amer. Sept. 296. 1814. Kalmia angustifolia f. procumbens Beissner, Schelle, & Zabel, Handb. Laubholz-Benennung 386. 1903. (nom. nud.). Kalmia angustifolia, var. rosea Forbes, Hort. Woburn. 93. 1833. Kalmia angustifolia f. rosea Beissner, Schelle, & Zabel. Handb. Laubholz-Benennung 386. 1903. (nom. nud.). Kalmia angustifolia var. variegata Forbes, Hort. Woburn. 93. 1833. Kalmia elliptica Raf. Autikon Bot. 87. 1840. (as syno- nym). Kalmia ferruginea Raf. Autikon Bot. 86. 1840. TYPE: unknown. Kalmia media K. Koch, Dendrologie II; 1: 153. 1872. Kalmia angustifolia f. media Beissner, Schelle, & Zabel, Handb. Laudholz-Benennung 386. 1903. (nom. nud.) TYPE: unknown. Kalmia intermedia Lange Ind. Sem. Hort. Hauniensi Nom. emendata 3. 1873. Kalmia glauca B intermedia Lange Ind. Sem. Hort. Hauniensi Nom. emendata 3. 1873. (as synonym). TYPE: a herbarium specimen in the Botanical Museum of Copenhagen (C) labeled Kalmia intermedia Lge ad int. Hort. bot. Haun. 16. Juni 1873, sem. ex hort. Paris s.n. Kalmia angustifolia B oleifolia Pers. Syn. Pl. 1: 4T". 1805. Kalmia oleifolia (Pers.) Bosse, Vollst. Handb. Blu- mengart. 2: 347. 1841. (as synonym). TYPE: unknown. Kalmia angustifolia var. pumila Forbes, Hort. Woburn. 93. 1833. Kalmia pumila Bosse, Vollst. Handb. Blumengart. 2:347. 1841. (as synonym). 370 Rhodora [Vol. 76 Kalmia pumila rubra Bosse, Vollst. Handb. Blumengart. 2:347. 1841. (as synonym). Kalmia pumila serotina Bosse, Vollst. Handb. Blumengart. 2: 347. 1841. (as synonym). Kalmia angustifolia f. pumila Voss, Blumengart. 1: 591, 1894. (without basonym). TYPE: unknown. Kalmia angustifolia « rubra Ait. Hort. Kew 2: 64. 1789. Kalmia angustifolia var. rubra (Ait.) Loddiges, Bot. Cab. 6: 502. 1821. Kalmia rubra (Loddiges) Bosse, Vollst. Handb. Blumengart. 2: 347. 1841. (as synonym). Kalmia angustifolia f. rubra Voss, Blumengart. 1: 591. 1894. (without basonym). TYPE: unknown. Kalmia angustifolia var. serotina Bosse, Vollst. Handb. Blumengart. 2: 347. 1841. Kalmia serotina Bosse, Vollst. Handb, Blumengart. 2: 347. 1841. (as synonym). TYPE: unknown. Mature leaves glabrous or with scattered stalked glandu- lar hairs beneath, usually not puberulent; calyx puberulent and strongly stipitate-glandular; corolla glabrous or lightly puberulent on the outside. Distribution: Bogs, Swamps, and other wet open areas from Ontario east to the Atlantic Ocean and southeast through the Great Lakes to Pennsylvania and south along the coast to Virginia. Common name: Sheep Laurel, Lambkill, Sheepkill, Wicky, Narrow-leaved Kalmia, Dwarf Laurel, Pig Laurel. REPRESENTATIVE SPECIMENS UNITED STATES: Connecticut: FAIRFIELD CO.: Trumbull, Eames 2809 (CONN). HARTFORD CO.: Kensington, Cowles (MU). LITCHFIELD CO.: Salisbury, Ebinger 697 (MU). NEW HAVEN co.: New Haven, Safford 163 (US). NEW LONDON CO.: Groton, Jausson (CONN, UT). TOLLAND CO.: Donovans Bog, Travis 1959 (PENN). WINDHAM CO.: Hasting’s Woods, Petersen (VDB). Delaware: NEW CASTLE CO.: Hog- swamp Road, Tatnall 4858 (GH). SUSSEX CO.: Canby (Us). Maine: ANDROSCOGGIN CO.: Lewiston, Bean 14793 (VDB). AROOSTOOK CO.: Knoles Corner, Neal 1907 (MAINE). CUMBERLAND CO.: Cape Eliza- beth, Gayle 766 (Us). HANCOCK CO.: Bar Harbor, Rossbach 2757 (ACAD, NCU). KENNEBEC CO.: Clinton, Bean (MAINE). KNOX CO.: 1974] Genus Kalmia — Ebinger 371 Glencove, F'riesner 7721 (MU, RENO, UT, WS, WVA). LINCOLN CO.: Ocean Point, Fassett 3946 (DUKE). OXFORD CO.: White Mt. Nat. Forest, Richards 6066 (MAINE). PENOBSCOT CO.: Old Town, Davis (MAINE). PISCATAQUIS CO.: Sangerville, Fernald 127 (MIN). SAGA- DAHOC CO.: Five Islands, Rehder 1018 (GH). SOMERSET CO.: Baker Brook, St. John & Nichols 2431 (CAN, NY). WALDO CO.: Searsmont, Rossbach 1932 (ACAD, MAINE). WASHINGTON CO.: Joe Dyer's Point, Walker 3039 (MAINE). YORK CO.: Sand Pond, Richards 5951 (MAINE). Maryland: ANNE ARUNDEL C0.: Severn Run, Adams & Wherry 3200 (GH). cECIL co.: North East, Long 54422 (GH). CHARLES CO.: Waldorf, Niewwland (ND). PRINCE GEORGES CO.: Airport Bog, Her- man 11546 (NY). Wicomico CO.: Sharptown, Tidestrom 12229 (GH). WORCESTER CO.: Furnace, Redmond 237 (MICH). Massachusetts: BARNSTABLE CO.: Falmouth, Greenman 3043 (MIN). BERKSHIRE CO.: Great Barrington, Eggleston 21413 (MIN, NY). ESSEX CO.: Marble- head, Nelson 341 (CAN). FRANKLIN CO.: Mt. Toby, Seymour 2566 (DUKE, MASS). HAMPDEN CO.: Granville, Seymowr 137 (MASS, MIN, NY). HAMPSHIRE CO.: Amherst, Torrey (MASS). MIDDLESEX CO.: Townsend, Ahles 65188 (MASS). NANTUCKET CO.: Polpis, Mackeever 504 (PENN). NORFOLK CO.: Medfield, Batchelder 234 (NCU). PLYMOUTH CO.: Ellisville, Erdman 4223 (BRY). WORCESTER CO.: Mount Wachusett, Palmer 43196 (MIN). Michigan: ALCONA CO.: Lincoln, Voss 4583 (MICH). ALPENA CO.: Alpena, McVaugh 11405 (MICH). | ARENAC CO.: Omer, Voss 6794 (MICH). CHARLEVOIX CO.: Garden Isl, Cain 149 (TRT). CHEBOYGAN CO.: Wolverine, Gleason & Gleason, Jr. 248 (GH, IND, ISC, NY, WVA). CRAWFORD CO.: Grayling, Voss 3098 (MICH). 10sCO CO.: Sand Lake, Dreisbach 7243 (CONN). KALKASKA CO.: Zavitz 1168 (QUK). MIS- SAUKEE CO.: Moorestown, Voss 3087 (MICH, OS). MONTMORENCY CO.: Hagenah 4936 (BLH). oscopa Co.: Rhodes Lake, Voss 3578 (MICH). OTSEGO CO.: Hardwood Lake, MeVaugh 10888 (BLH, CAN, MICH, MO). PRESQUE ISLE CO.: Emma, Clover 26193 (MICH). ROSCOMMON CO.: Roscommon, Voss 4258 (MICH). SAGINAW CO.: Sage Lake, Case, Jr. (MICH). New Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon & Steele 9124 (NHA). CARROLL CO.: Albany, Gale (NHA). CHESHIRE co.: Winchester Manning (BRY). coos Co.: Randolph, Pease 4696 (GH, MICH). GRAFTON CO.: Pike, Mathias 163 (MIN). HILLSBORO co.: Mt. Monadnock, Andrews 895 (ORE). MERRIMACK CO.: Wilmot, Hodgdon & Steele 9949 (NHA). ROCKINGHAM CO.: Harris 224 (NHA). STRAFFORD CO.: Rochester, Barrett & Hodgdon 15522 (NHA). SULLI- VAN CO.: Springfield, Hodgdon & Steele 9129 (NHA). New Jersey: ATLANTIC CO.: Hammonton, Keefe 123 (UWM). BERGEN CO.: Carl- stadt, Niederer (NY). BURLINGTON CO.: Rancocas River, Leonard 6314 (US). CAMDEN CO.: Atco, Painter 684 (MIN, US). CAPE MAY co.: Belleplain, Fender 4916 (PAC, PENN), CUMBERLAND CO.: Mill- ville, Adams 324 (GH, PENN). ESSEX CO.: Franklin, Rusby (CHRB). 372 Rhodora [Vol. 76 GLOUCESTER CO.: Repaupo, Long 16065 (GH). MIDDLESEX CO.: Mill- town, House (US). MONMOUTH CO.: Farmingdale, Long & Brown 9557 (PENN). MORRIS CO.: Mt. Hope, Britton (CHRB). OCEAN CO.: Lakewood, Mackenzie 3112 (Us). PASSAIC co.: Uttertown Bog, Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6616 (PENN). SOMERSET CO.: Sourland Mts., Fogg, Jr. 12406 (PENN). SUSSEX CO.: Cranberry Lake, Mackenzie 717 (ARIZ, IND). New York: ALBANY CO.: Karmer, House 7850 (CAN). CHENANGO co.: Plymouth Pond, Wiegand 6976 (GH). CLINTON CO.: Rudd 816 (US). COLUMBIA CO.: Ghent, McVaugh 362 (PENN). DUTCHESS CO.: Susan Court, Ahles 64464 (MASS). ESSEX CO.: Lake Harris, House 8072 (CAN). FRANKLIN CO.: Upper Saranac Lake, Muenscher, et. al. 476 (NY). FULTON CO.: Gloversville, Haught (WVA). GREENE CO.: Coville (GH). HAMILTON CO.: Little Tupper Lake, Fosberg 16101 (PENN). JEFFERSON CO.: Perch Lake, Beschel 17144 (QUK). LEWIS co.: Martinsburg, ‘Hotchkiss 2654 (GH). NASSAU CO.: Massapequa St. Park, Cain 93 (MO, PENN, TENN). ONEIDA CO.: Sylvan Beach, Maxon (US). ORANGE CO.: Sutherland Pond, Mearns 125 (US). RENSSELAER CO.: West Sand Lake, Brown 652 (DUKE). RICHMOND co.: Staten Isl, Dowell & Davis 2224 (GH). ST. LAWRENCE CO.: Norfolk, Phelps 756 (GH, NY, US). SARATOGA CO.: Corinth, Whitney 4124 (PENN). SUFFOLK CO.: South Haven, Moldenke 18685 (NY). SULLIVAN CO.: Wurtshore, Hess 1267 (ARIZ). ULSTER CO.: Highland, Chrysler (CHRB). WARREN CO.: Luzerne, Fogg, Jr. 20309 (PENN). WASHINGTON CO.: Lake Nebo, Burnham 2756 (MIN). Pennsylvania: BERKS C0.: Hamburg, Berkheimer 1734 (PENN). BRADFORD CO.: Troy, Beardslee (MU). BUCKS CO.: Bristol, Adams 4514 (MAINE, PENN). CARBON CO.: Little Gap, Adams 3331 (GH, ISC, PENN). CHESTER CO.: Warwick, Wherry (GH). COLUMBIA CO.: Centralia, Fogg, Jr. 14256 (NY, PENN). CUMBERLAND CO.: Sand Springs, Ward (PAC), DELA- WARE CO.: Tinicum, Baker (PENN). FRANKLIN CO.: Caledonia St. Park, Overhots (PAC). LACKAWANNA CO.: West Mt., Glowenke 6528 (MIN, NY). LANCASTER CO.: Smithville, Small (US). LEHIGH CO.: Allentown, Dowell 1115 (Us). LUZERNE CO.: Freeland, Orton (PAC). LYCOMING CO.: Bald Eagle Mt., Keener 2561 (PAC). MONROE CO.: Tannersville, Swartley 791 (MAINE). NORTHAMPTON CO.: Daniels- ville, Schaeffer, Jr. 6184 (PAC). NORTHUMBERLAND CO.: Montandon, Reed 235 (CHRB). PIKE CO.: Twin Lakes, De Pue 878 (PENN). SCHUYLKILL CO.: Tamaqua, Kriebel 8586 (ND). SOMERSET CO.: Salisbury, Caffrey (PAC). SULLIVAN CO.: Double Run, Fogg, Jr. 14437 (PENN). WAYNE CO.: Big Spring Pond, Glowenke 2434 (PENN). WYOMING CO.: Falls, Glowenke 2023 (PENN). YORK CO.: McCall’s Ferry, Heller & Halbach (MIN). Rhode Island: KENT CO.: Warwick, Congdon (08). NEWPORT CO.: Newport, Mearns 254 (GH). PROVIDENCE CO.: Slaterville, Palmer 43348 (ISC, MIN). WASHINGTON co.: South Kingstown, Collins (CONN). Vermont: ADDISON CO.: 1974] Genus Kalmia — Ebinger 373 Bristol Pond, Eggleston 21453 (US). BENNINGTON CO.: Manchester, Day 267 (GH). CALEDONIA CO.: Barnet, Blanchard (HNH, MO). CHITTENDEN CO.: S. Burlington Twp., Charette 2190 (BRY, MIN). FRANKLIN CO.: East Berkshire, Clausen (MO, US). ORANGE CO.: Fairlee, Grover (MU). RUTLAND CO.: Middletown Springs, Carpenter (NCU). WASHINGTON CO.: Roxbury, Countryman 820 (HNH, NHA). WINDHAM CO.: Westminster, Robinson 30 (GH). WINDSOR CO.: Reading, Kittredge (NY). Virginia: ISLE OF WIGHT Co.: Franklin, Heller 1124 (GH, MIN, MO, PENN, US, WS); Lee’s Mill, Fernald & Long 12146 (GH, MO, NY, US), Harvill 15946 (NCU); Zuni, Fernald, Griscom & Long 6665 (GH, NY), 6666 (GH, PENN). NANSEMOND CO.: Cherry Grove, Fernald & Long 11392 (DUKE, GH, US); County Rd. 665, Duke 1067 (NCU); South Quay, Fernald & Long 10366 (GH, NY). CANADA: Labrador: Assizes Isl., Potter & Brierly 3424 (GH); Cape St. Charles, Potter & Brierly 3427 (GH); Cartwright, Brown (GH); Forteau, Long 28832 (GH); Goose Bay, Gillett & Findley 5352 (MIN, MO, TRT, WS), 5584 (ACAD, NY, US); Kenamu River, Gillett & McKay 5831 (ACAD, GH); St. Clary’s River, Wallace 23 (CONN). New Brunswick: ALBERTA CO.: Grindstone Isl. Christie 1575 (NBM). CARLETON CO.: East Wolf Isl, Hodgdon & Pike (NCU). CHARLOTTE co.: Deer Isl., Squires & Squires (NBM), GLOUCESTER CO.: Four Roads, Roberts & Pugh 65-4358 (CAN). KENT CO.: Richibucto, Scoggan 12135 (CAN). KINGS CO.: Havelock, Perry (ACAD). MADA- WASKA CO.: St. Leonard, Malte 294 (CAN). NORTHUMBERLAND CO.: Mt. Carleton, Smith & Clattenburg 20066 (ACAD, QUK). QUEENS CO.: Mill Brook, Squires & Christie (NBM). RESTIGOUCHE CO.: Dalhousie, Scoggan 11721 (ACAD, CAN). ST. JOHN CO.; Mispec, Crawford & Patterson (ACAD). SUNBURY CO.: Mill Settlement, Christie 245 (NBM). WESTMORLAND CO.: Johnson Lake, Beschel 5508 (QUK). Newfound- land: BONAVISTA NORTH DIST.: Glovertown, Smith, Squires, & Smith 116 (ACAD), 135 (ACAD). BONAVISTA SOUTH DIST.: Lethbridge, Rouleau 8370 (NFLD). BURGEO-LA POILE DIST.: Osmond, Rouleau 7375 (NFLD, US). CARBONEAR DIST.: Old Perlican, Torrey 25 (CAN, GH). FERRYLAND DIST.: Witless Bay, Rouleau 5364 (NFLD). FOGO DIST.: Stag Harbour, Ludlow 125 (ACAD). FORTUNE BAY-HERMITAGE DIST.: Wallace Cove, Rouleau 8089 (NFLD). GANDER DIST.: Gander, Rouleau 6133 (CAN, NFLD, US). GRAND FALL DIST.: Overflow Pond, Rouleaw 7961 (NFLD). HUMBER DIST.: Corner Brook, Rouleau 3960 (ACAD, CAN, GH, NFLD, NY, US). ST. BARBE DIST.: Bonne Bay, Bishop 471 (GH, CAN). ST. JOHN’s DIST.: St. John's, Robinson & Schrenck 41 (GH, CAN, MIN, MO, NY, P). TRINITY SOUTH DIST.: Whitbourne, Rouleaw 6069 (NFLD). TWILLINGATE DIST.: Loon Bay, Rouleau 4866 (NFLD). WHITE BAY DIST.: St. Anthony, Savile & Vaillancourt 2561 (NCU, NFLD, NY). Nova Scotia: ANNAPOLIS CO.: Belleisle, Bent 12723 (ACAD). ANTIGONISH co.: Linwood, Smith, et. al. 10765 374 Rhodora [Vol. 76 (MAINE). CAPE BRETON CO.: Frenchvale, Smith, et. al. 15094 (ACAD). COLCHESTER CO.: Truro, Malte, (CAN, QUK). CUMBERLAND CO.: Wharton, Schofield 3197 (ACAD). DIGBY CO.: Corberrie, Jack 3121 (MAINE, NY, US). GUYSBOROUGH CO.: Canso, Rousseau 35450 (CAN). HALIFAX C0.: Halifax, Jack 3235 (NY, TENN). HANTS CO.: Bishop- ville Rd., Wellwood 12662 (ACAD). INVERNESS CO.: North Mt., Smith, et. al. 11275 (ACAD). KING CO.: Wolfville, Anderson (ACAD). LUNENBURG C0.: Beck Lake, Smith, et. al. 18551 (ACAD, NCU). PICTOU co.: Westville, Sampson 28 (ACAD). QUEENS CO.: Tobeatic Lake, Martin 5210 (CAN). RICHMOND CO.: Grand Lake, Ershine 51993 (QUK). SHELBURN C0.: Barrington Passage, Macoun 81371 (CAN). VICTORIA CO.: Cape Breton, Senn 1553 (MIN, US). YARMOUTH CO.: Seal Isl, Erskine 54.977 (ACAD). Ontario: BRUCE CO.: Big Bay, Krotkov 9295 (NY). CARLETON CO.: Wright’s Cove, Minshall 236 (MIN, NY). COCHRANE CO.: Abitibi, Pritchard 208A (MASS). DUNDAS co.: Elma, House 23357 (PENN). FRONTENAC CO.: Kingston, Fowler (US). HALIBURTON CO.: Minden Twp., Roberts (TRT). KENORA DIST.: Attawapiskat River, Porsild, et. al. 19986 (CAN). LENNOX CO.: Skoota- matta, Jackson (TRT). MANITOULIN DIST.: Honora, Soper & Bell 9819 (CAN, QUK, TRT). MUSKOKA DIST.: Sparrow Lake, Walden 110 (TRT). NIPISSING DIST.: Stuarts, Ross 2 (TRT). ONTARIO CO.: Cannington, Thompson 516 (TRT). PARRY SOUND DIST.: Ahmic Lake, Soper 5225 (CAN, TRT). PRESCOTT CO.: Alfred Twp., Calder & Cody 922 (PENN). RENFREW CO.: Renfrew, Stroud (TRT). STORMONT CO.: Newington, Jenkins 8297 (NCU). SUDBURY DIST.: Sudbury, Ostenfeld 613 (CAN). THUNDER BAY DIST.: Longlac, Baldwin & Breitung 3693 (CAN, TRT). TIMISKAMING DIST.: Cattle Isl., Moss 77 (ALTA). WELLAND C0.: Niagara, Zenkert (OAC). Prince Edward Island: KING CO.: Dundee, Fernald, Long, & St. John 7899 (GH). QUEENS C0.: Brackley Beach, Grandtner 18518 (CAN). Quebec: ABITIBI CO.: Amos, Morton 11035 (US). ARGENTEUIL CO.: Lachutem, Porsild 6337 (CAN). BERTHIER co.: Lanoraie, Scoggan 370 (CAN). BONAVENTURE CO.: New Rich- mond, Chrysler 1228 (CHRB). BROME CO.: Bolton, Knowlton (GH). CHICOUTIMI CO.: Lac Kenogami, Lavoie, Doyon, & Lord 600706-01 (ACAD, TRT). GASPE WEST CO.: Mt. Albert, Cottam 11852 (UT). GATINEAU CO.: Wilson Corners, Mitchell 1750 (TENN). KAMOURASKA co.: Ouelle, Gauthier 65-207 (QUK). LABELLE CO.: Nomininque, Roy 1053 (ACAD). LEVIS CO.: Chaudiere, Forbes (CAN). MAGDALEN ISLAND co.: Amherst Isl, Potter & Brierly 3425 (NMC, UT). MASKINONGE co.: Metis Beach, DuBoulay & DuBoulay 2589 (QUK). MATANE CO.: Lake Montagne, Boivin 786 (PENN). MATAPEDIA CO.: Saint Zenon, Blain 321 (CAN). MEGANTIC CO.: Black Lake, Blais, Deshaies, & Forest 10379 (CAN, QUK, SASK). MISSISQUOI CO.: Farnham, Maycock 1863 (QUK). MONTCALM CO.: Lac Tapani, Pennell 16678 (NY). MONTMORENCY CO.: Boischatel, Cauchon, et. al. 69-111 (MU, TRT). PORTNEUF CO.: Marie-Victorin 9437 (WS). RICHELIEU CO.: Saint-ours, 1974] Genus Kalmia — Ebinger 975 Rolland-Germain 48518 (GH). RIMOUSKI CO.: Cap Enrage, Rousseau 26877 (GH). RIVIERE DU LOUP CO.: Soper 3670 (TRT). SAGUENAY CO.: Mingan, Townsend (GH); Natashquan, Abbe 1188 (GH); Romaine, St. John 90652 (GH); Round Lake, Gillett 5208 (GH); Seven Isl., Harper 3309 (CAN, HNH, PENN, US), Robinson 769 (CAN, GH, NY). SHERBROOKE CO.: Orford, Martineau 35 (CAN). TEMISCOUATA CO.: Trois Pistoles, Fox (TRT). TERREBONNE CO.: St. Sophe, Wahl 19419 (PAC). TIMISKAMING CO.: Ottawa River, Baldwin 5823 (CAN, SASK). TWO MOUNTAINS CO.: La Trappe, Louis-Marie 10 (GH). WOLFE CO.: Lake Nicolet, Brisson & Hamel] 11295 (ACAD, OAC, SASK, TRT). The synonomy of this taxon is fairly extensive. Numerous varietal and forma names have been proposed for the vari- ation that exists, and some of these names have been ele- vated to specifie rank. Many of these lack descriptions or are listed as synonyms under a variety or form name. Ex- cept for these subspecific categories only a few other names have been applied to this taxon. One of these was when Kuntze (1891) transferred the specific epithet angustifolia to another genus while the others were proposed by Rafin- esque (1840). Most of the subspecific categories that have been pro- posed for Kalmia angustifolia var. angustifolia are for variation and extremes in flower color (carnea, rosea, rubra), variation in size and habit of the plant (media, minima, nana, procumbens, pumila) and variation in shape and color of the leaves (lucida, oleifolia, ovata, variegata) . In general, the differences in flower color appears to repre- sent natural variation within populations and the above are better treated as cultivars than as forms. This is also true of the variation in leaf shape and color, and, in part, for variation in plant height. Holmes (1956) discusses some of these cultivars. The dwarf condition reported for Kalmia angustifolia var. angustifolia is either the result of selection of more northern or high altitude sources or because of inbreeding depression. Jaynes (19682) found that seedlings from selfed plants average about half the height of seedlings from outcrossed plants. He also found that seed set from self-pollinated plants is 85-90% less than that from cross 376 Rhodora [Vol. 76 pollination and that there is a marked reduction in survival of the seedlings from selfed plants. This indicates that this species is only weakly self compatible and that outcrossing usually occurs under natural conditions. Kalmia angustifolia var. angustifolia commonly occurs in bogs, swamps, and other wet places, usually forming dense thickets around ponds and lakes. It is also found in open woods, as a weed in fields, and in moist openings of pine savannas. It is distributed from the northeastern part of the Michigan peninsula and the eastern half of Ontario, east through Quebec, the Maritime Provinces, and New- foundland; north to at least the Attawapiskat River (Ken- ora District) in Ontario and to Goose Bay and Cartwright in Labrador; and south in eastern United States through New England and eastern New York, eastern Pennsylvania and Maryland to the coast in New Jersey and Delaware and the southeastern tip of Virginia. One specimen (Davis & Davis, 4691) has been seen which is reputed to have come from Randolph Co., West Virginia, but this collection should be considered doubtful. This species has also been reported from Indiana, but according to Deam (1940) this is an error. It is possible that extracts from sheep laurel may inhibit the growth of other plants, particularly coniferous trees, thereby maintaining open areas for extended periods of time. Peterson (1965) found that the leaves of this species contain a substance that hinders the primary root develop- ment of black spruce by destruction of the epidermal and cortical cells. He further suggests that this may be the reason for the abnormally poor growth of trees associated with Kalmia angustifolia on upland sites in the boreal region of eastern Canada. Studies by Jaynes (personal communication) reveal that some incompatibility exists between the two varieties of Kalmia angustifolia. This incompatibility is highly variable, depending upon the parents used, with the resulting off- spring varying from completely sterile to highly fertile. Many of these hybrids produced a large quantity of viable 1974] Genus Kalmia — Ebinger 377 seed while in others the capsules remained small and lacked seeds. From some crosses, both sterile and fertile individu- als were produced. Also, among the offspring that pro- duced no seeds, stainable pollen varied from 64% to 2%. All artificially produced hybrids between these two taxa are similar to Kalmia angustifolia var. angustifolia, and except for the lack of seed set in some, could not be dis- tinguished from it. All individuals have glandular calyces and most have leaves that are not puberulent beneath, both characteristics of typical var. angustifolia. Only 2 hybrids were found that had leaves which were puberulent beneath. More studies are needed to clarify the status of these taxa, but until more information is available, it seems best to maintain them as varieties. Kalmia angustifolia L. var. angustifolia f. candida Fern. Rhodora 15: 151. 1913. Kalmia angustifolia var. candida (Fern.) Rehd. In: Bailey, Stand. Cycl. Hort. 3: 1734. 1915. TYPE: Newfound- land: damp mossy or turfy hollows on hill south of St. John’s, Fernald & Wiegand 6019 (GH). According to Jaynes (1971a, 1971b) the presence of pig- ment in Kalmia angustifolia is determined by a single dominant gene, and the true-breeding recessive is white flowered. These white flowered individuals are anthocyan- inless with green stems, as contrasted to the normal wild types with reddish stems. This characteristic is particularly obvious in seedlings. It is possible that the recessive genes for the anthocyan- inless condition are farily common in populations of this species, but are rarely expressed. In a population of Kalmia angustifolia from Madison, Connecticut, Jaynes (1971b) found that about 24% of the population was heterozygous for the anthocyaninless condition, but no homozygous re- cessive plants were found. It appears that the anthocyanin- less plants are at a selective disadvantage in the wild. SPECIMENS EXAMINED UNITED STATES: Maine: ANDROSCOGGIN CO.: Gordon 407 378 Rhodora [Vol. 76 (MAINE). LINCOLN CO.: Hyland 69 (MAINE). Massachusetts: Sher- born, Loomis (GH). CANADA: Newfoundland: St. Johns, Fernald & Wiegand 6019 (GH). 4b. Kalmia angustifolia L. var. caroliniana (Small) Fern. Rhodora 39: 476, 498. 1937. Kalmia caroliniana Small, Fl. Southeast. U.S. 1336. 1903. TYPE: North Carolina: Flat Rock, L. R. Gibbs (NY). Kalmia carolina Small, Fl. Southeast. U. S. 886. 1903. Kalmia angustifolia var. carolina (Small) Fern. Rhodora 39: 437. 1937. Mature leaves permanently puberulent beneath with a dense covering of whitish, simple hairs, scattered stalked glandular hairs also present; calyx strongly puberulent, lacking stalked glandular hairs; corolla usually strongly puberulent on the outside. Distribution: Open woods and shrubby bogs in the moun- tains and sandy woods, pocosins, savannas and bogs on the coastal plains. Extending from extreme southern Virginia through North Carolina to eastern South Carolina and west to eastern Tennessee and northeastern Georgia. Common name: Sheep Laurel, Lambkill. REPRESENTATIVE SPECIMENS Georgia: RABUN C0.: Oakey Mt., Duncan 3281 (GH, ID, ISC, MO, NY, TENN, US), 4312 (Isc); Tom's Swamp, Bookout & McDowell 65 (GH). North Carolina: ALLEGHANY CO.: Edmonds, Radford 38191 (NCU). AVERY CO.: Pineola Bog, Cain & Duncan 251-3 (IND, ND). BEAUFORT CO.: Chocowinity, Radford 33396 (NCU). BLADEN CO.: White Lake, Kral 27210 (VDB). BUNCOMBE CO.: Asheville, McGarthy (US). CARTERET CO.: Beaufort, Bentley (VDB). CHOWAN CO.: Tyner, Ahles 39907 (NCU). COLUMBUS C0.: Chadbourn, Wilbur 5315 (DUKE). CRAVEN C0.: New Bern, Harbison 14080 (NcU), 14932 (NCU). CUMBERLAND CO.: Stedman, Wilbur 5169 (DUKE). DARE CO.: Manns Harbor, Radford 8197 (NCU). DUPLIN Co.: Charity, Fogg, Jr. 5481 (GH, PENN). GATES CO.: Gatesville, Godfrey 5214 (DUKE, GH). GREENE CO.: Walstonburg, Radford 36508 (NCU). HARNETT CO.: Blomquist 4486 (DUKE). HENDERSON CO.: Muddy Creek, Smith 221 (180, US). HOKE CO.: Timberland, Ahles 29514 (IND). HYDE CO.: Radford & Stewart 803 (NCU). JACKSON CO.: Cullowhee, Biltmore 1974] Genus Kalmia — Ebinger 379 Herb. 1344e (ALTA, GH, US). JOHNSTON CO.: Benson, Smith (IND, NCU). JONES CO.: Hargetts Store, Radford 36956 (NCU). LENOIR co.: Pine Hill, Radford 31651 (NCU). MACON co.: Highlands, Wilson 1854 (TENN). MARTIN CO.: Robersonville, Radford 35364 (Isc). MONTGOMERY CO.: Pekin, Radford 14595 (NCU). NEW HAN- OVER CO.: Wilmington, Newber 165 (UARK). ONSLOW co.: Dixon, Moldenke 1242 (DUKE, MIN, MO, NY, PENN, US). PASQUOTANK CO.: Elizabeth City, Ashe (NCU). PENDER CO.: Wards Corner, Ahles & Bell 7856 (MISSA, MSC). PERQUIMANS CO.: Beach Spring, Radford 5470 (NCU). PITT CO.: Staton, Radford 34940 (NCU). RICHMOND co.: Rockingham, Coker (NCU). ROBESON CO.: Allenton, Ahles & Ramseur 23770 (ND, PAC). SAMPSON CO.: Tomahawk, Fox & Boyce 3792 (KY). SCOTLAND CO.: Wagram, Correll 1181 (DUKE). TRAN- SYLVANIA CO.: Cedar Mt., Freeman 58130 (NCU). TYRRELL CO.: Cross Landing, Radford 33820 (NCU). WASHINGTON CO.: Hoke, Radford 35264 (NCU). WATAUGA CO.: Deep Gap, Radford 6239 (NCU). WAYNE CO.: Goldsboro, Radford 22006 (NCU). South Carolina: DARLINGTON CO.: between Darlington & Society Hill, Smith 1068 (NCU). HORRY CO.: Radford & Stewart 979 (NCU, NY). ORANGEBURG co.: Edisto Swamp, Ravenel (GH). SUMTER CO.: Poinsette St. Park. Holdaway (DUKE). Tennessee: JOHNSON CO.: Cole’s Bog, Shanks & Sharp 1115 (PENN, TENN); Shady Valley, Barclay (TENN), Sharp & Underwood 1252 (vps), 1552 (NCU, NY), 3161 (NY). KNOX CO.: Kingston Pike, Jennison 717 (TENN). Virginia: CARROL CO.: Wood- lawn, Massey 5058 (GH). NANSEMOND CO.: Whaleyville, Fernald & Long 7564 (GH, NY, US). NORFOLK CO.: Dismal Swamp, Coville & Kearney, Jr. 91 (08, US), Portsmouth Ditch, Grimes 4517 (MIN, NY). SOUTHAMPTON CO.: Franklin, Fernald & Long 10002 (GH). The name first applied to this taxon was Kalmia carolina when Small (1903) separated the plants with puberulent leaves from typical K. angustifolia. Later in a list of genera and species in the same publication (page 1336) the name was changed to K. caroliniana. It is very possible that this later name is a printing error, particularly since the name K. carolina was used by the author in a later publication (Small, 1914). Fernald (1937) transferred the epithet carolina to a variety of K. angustifolia (page 437), but later in the same publication (page 476, 498) changed the name to variety caroliniana, stating that this spelling, “since it accompanied the citation of the type, should prevail.” Kalmia angustifolia var. caroliniana has a fairly re- stricted range. Most of the specimens examined are from 380 Rhodora [Vol. 76 the coastal plain and the mountains of North Carolina with the species being extremely common on the pocosins and in bogs. This taxon also has a sporadic occurrence in the coastal plain of South Carolina (Radford, Ahles, and Bell, 1968) and in southern Virginia (Fernald, 1938, 1939), while it has been reported from a few localities in the moun- tains of extreme eastern Tennessee by Jennison (1935) and Shanks (1952) and from two mountain bogs in the Blue Ridge of northeastern Georgia by Duncan (1948). The characteristics used to separate Kalmia angustifolia var. caroiimiana are very reliable, but occasionally some overlap does occur. Usually, var. angustifolia has glandular calyces and leaves that are not puberulent beneath while in var. caroliniana the calyces are eglandular and the leaves are strongly puberulent beneath. All but two specimens of var. caroliniana examined had strongly puberulent leaves and even those were lightly puberulent on the undersurface of the leaf. This was also noticed by Radford, Ahles, and Bell (1968) who mentioned that a few specimens of var. caroliniana with glabrate leaves were found in the Caro- linas. A few specimens of var. angustifolia from Nantucket Island and the coast of Massachusetts and Connecticut have leaves that are strongly puberulent beneath. All of these, however, have strongly glandular calyces, typical of var. angustifolia, and are referred to that taxon. The best char- acter to separate the two varieties is the presence or absence of stalked glandular hairs on the calyx. Close examination, however, reveals that on some specimens of var. caroliniana a few glandular hairs are present on the calyces. The glands are not very numerous, however, and mostly restricted to the margins of the lobes or to the base of the calyx. Even on the holotype a few of these stalked glandular hairs are found. Because of this variability it is probably best to maintain these taxa as varieties of one species rather than as separate species. A white flowered, anthocyaninless form of var. carolin- iana also exists. It is similar genetically to Kalmia angusti- folia var. angustifolia f. candida in that it is a homozygous 1974] Genus Kalmia — Ebinger 381 recessive plant in which flower color is determined by a single pair of genes (Jaynes, 1971b). A specimen of this form was obtained by Dr. Jaynes from the Garden-in-the- Woods, in Sudbury, Massachusetts. No wild individuals have been found and its origin is not known. The best crossing results obtained by Jaynes (1968a) between K. angustifolia and other species of the genus were when this white flowered form was used as the female parent (Jaynes, 1971b). 5. Kalmia cuneata Michaux, Fl. Bor.-Amer. 1: 257. 1803. Chamaedaphne cuneata (Michx.) Kuntze, Rev. Gen. Pl. 2: 388. 1891. TYPE: In Carolina, Michaux s.n., (GY). Much branched rhizomatous shrub to 1.5 m tall; branch- lets terete, reddish brown, usually adscending, stipitate- glandular and puberulent, becoming glabrous and light gray; leaves alternate, deciduous, flat, oblanceolate to ellip- tic-lanceolate, 2-6 cm long, 0.5-1.5 cm wide, lightly stipitate glandular beneath, glabrous above, apex obtuse to acute, apiculate, base narrowly cuneate, midrib puberulent above, stipitate-glandular beneath; petiole to 4 mm long; inflor- escence a reduced corymbiform raceme or fascicle of 3-10 flowers in the upper axils of the last season’s growth, pedi- cels 1-3 mm long, glabrous to lightly stipitate-glandular, recurved in fruit, subtended by 2 bracteoles in the axil of a small lanceolate bract about 1 mm long, main axis of the inflorescence less than 3 mm long; calyx glabrous to lightly stipitate-glandular, 6-7.5 mm broad, the lobes oblong, apex acute; corolla creamy white with a red band within, lightly stipitate-glandular outside, puberulent near the base within, 1.3-1.8 mm broad; anthers purplish brown, 1-1.3 mm long; filaments puberulent toward the base; style 7-10 mm long; ovary stipitate-glandular; capsule depressed globose, 4-6 mm broad, 2.5-3.5 mm long, stipitate-glandular; seeds light brown, 0.6-0.9 mm long, the testa not loose or extended past the ends of the seed. Chromosome number n — 12. Distribution: Wet thickets and shrub bogs in southeastern 382 Rhodora [Vol. 76 North Carolina and adjacent South Carolina. Common name: White Wicky. REPRESENTATIVE SPECIMENS North Carolina: BLADEN CO.: Colly Creek, Ashe (PH); 10 miles north of Elizabethtown, Ebinger 11271 (EIU); Jerome Pocosin, Buell & West 1740 (MIN). CUMBERLAND C0.: Hope Mill, Alexander (NY). MOORE CO.: Pinehurst, Wicker (NY). PENDER CO.: Squalling Bluff Rd., Coville (MICH, NY, US); Burgaw, Chrysler 2262 (CHRB). RICH- MOND CO.: Hamlet, Wiegand & Manning 2365 (GH). South Carolina: DARLINGTON CO.: Hartsville, Smith (NY), Coker (NY). A small fragment of the type of Kalmia cuneata is at the Gray Herbarium. It consists of one fascicle of 5 flowers and a small stem with a few leaves attached. On the label is written “Ex Herb. Michaux Kalmia cuneata." At the pres- ent time the rest of the type has not been located. Kalmia cuneata is very distinct. The only species that it could possibly be confused with is K. angustifolia since their general habit and leaf size are similar. Kalmia cune- ata, however, is the only deciduous member of the genus and the only one in which the flowers are borne in fascicles from the axils of last season’s growth. Other distinguishing characteristics are the creamy-white flowers with a red band within, the oblanceolate leaves, and the recurved fruit- ing pedicels. Genetically, this species appears to be most closely re- lated to Kalmia hirsuta, another alternate leaved species of the southeastern United States. Jaynes (1968a) reported that K. cuneata set no seed when used as the female parent, but as the male parent crossed with both K. angustifolia and K. hirsuta. The cross with K. angustifolia produced only inviable hybrids. Most of the hybrids with K. hirsuta were also inviable, producing sectors of albino or yellow- green tissue, but one hybrid flowered and was pollen sterile. This species is one of the rarest shrubs in North America and few early reports exist concerning its distribution. Pursh (1814) reports that it occurs on the mountains of Carolina, while Nuttall collected it between Camden, South Carolina and Statesville, North Carolina (Wood, 1961). 1974] Genus Kalmia — Ebinger 383 Curtis (1835) reported its occurrence about 30 miles NE of Wilmington, North Carolina. Present studies indicate that it grows only in the sandhills and bay margins of the pocosin ecotones of the Carolinas. It now is known from 8 counties in this region (Radford, Ahles & Bell, 1968). 6. Kalmia hirsuta Walt. Fl. Carol. 138. 1788. Chamaedaphne hirsuta (Walt.) Kuntze, Rev. Gen. Pl. 2: 388. 1891. Kalmiella hirsuta (Walt.) Small, Fl. Southeast. U.S. 886. 1903. TYPE: unknown. Kalmia ciliata Bartram, Trav. N. & S. Carol. v, 7, 94, 171, 242,1791. (nom. nud.). Low, sparingly branched shrub, to 6 dm tall with a thick basal caudex; branchlets terete, reddish brown, puberulent and densely hispid; leaves alternate, sessile or with a petiole to 1 mm long, elliptic to ovate, 5-14 mm long, 1.5-8 mm broad, the margins usually slightly revolute, usually pu- berulent and hispid on both surfaces, small stipitate glands usually present beneath, apex acute, base broadly cuneate to rounded; flowers solitary (rarely in fascicles of 2-5 or in compact racemes) in the axils of the leaves of new growth; pedicels to 1.5 cm long, stipitate-glandular and hispid, subtended by a pair of bracts; calyx green, the lobes foliaceous, lanceolate, 3-8 mm long, hispid, tardily deciduous in fruit; corolla pink (rarely white), marked with red around the anther pockets and a red ring near the base, 1-1.5 em broad, usually lightly hirsute and stipitate-glandu- lar on the keels outside, puberulent at the base within; anthers purplish brown, 0.7-1.3 mm long; filaments puberu- lent toward the base; ovary usually stipitate glandular; style 5-7 mm long; fruit 2-4 mm broad, 2-3.5 mm long, subglobose to ovoid, thinly stipitate-glandular; seeds light brown, 0.3-0.5 mm long, the testa not loose or extended past the ends of the seed. Chromosome number n = 12. Distribution: Low, sandy pine savanna, sand hills, dunes and flat pine woods on the coastal plains of southern Ala- 384 Rhodora [Vol. 76 bama and Georgia, northern Florida and the southern tip of South Carolina. Common name: Sandhill Laurel, Calico-bush. REPRESENTATIVE SPECIMENS Alabama: BALDWIN CO.: Gulf State Park, Webster & Wilbur 3539 (GH, NY, US). MOBILE C0.: Theodore, Pennell 4513 (MICH, NY, PENN, PH). Florida: ALACHUA CO.: Cross Creek, Ford 2250 (GH, TENN). BAY CO.: Lynn Haven, Billington (MicH, us); Lazor 5006 (Pac). CLAY CO.: Kingsley Lake, Henry 3828 (PH). COLUMBIA CO.: Lake City, Nash 2193 (GH, MASS, MICH, MIN, MSC, ND, NY, OS, PH, US). DIXIE C0.: Old Town, West & Arnold (PH). DUVAL CO.: Jacksonville, Curtis 4359 (ARIZ, ND, NY, US), 4921 (GH, ISC, MIN, MSC, NY, US). ESCAMBIA CO.: Pensacola, Webster & Wilbur 3572 (GH, NY, US). FLAGLER CO.: Bunnell, Ames (GH). FRANKLIN CO.: Carrabelle, Mosier 884 (MIN, NY, PENN, US); Moldenke 1134 (MIN, NY, PENN, US). GADSDEN CO.: Quincy, Sargent (MIN). GILCHRIST CO.: Trenton, Ward, West & Will 2651 (GH, US). GULF CO.: Port St. Joe, Ward & Ward 1460 (GH). HAMILTON CO.: Bowers & Wofford 71-550 (TENN). HOLMES CO.: Bonifay, Blanton 6582 (US). JACKSON CO.: Compass Lake, Knight & Arnold (PAC). JEFFERSON Co.: Wacissa, Godfrey 53552 (GH). LAFAYETTE CO.: Smith & Myint 189 (GH). LEON CO.: Silver Lake, Godfrey 53660 (GH, MISSA, NY). LEVY CO.: Bronson, Godfrey & Lindsey 56989 (GH). LIBERTY CO.: Bristol, Small, Small & DeWinkeler 11448 (Gn, NY, PENN, TENN). MADISON CO.: Green- ville, Ramsey 106 (MSC). MARION CO.: Ocala, Beardslee, Jr. (MU). OKALOOSA CO.: Crestview, McArthur (GH). PUTNAM CO.: Interlachen, Fox 5703 (GH, MICH, MIN, NY, TENN). ST. JOHNS CO.: Anastasia Island, Crawford (NY, PH). ST. LUCIE CO.: Fort Walton, Henry 1087 (PH). SANTA ROSA CO.: Munson, Ford & Arnold 4190 (us). WAKULLA CO.: St. Marks Wildlife Refuge, Godfrey 57860 (GH). WALTON CO.: Santa Rosa, Ripley & Barneby 10128 (NY). Georgia: APPLING CO.: Baxley, Cottam 11060 (UT); Ebinger 11204 (E10). BRANTLEY CO.: Waynesville, Ebinger 11232 (EIU). BULLOCH CO.: Portal, Henry 7311 (PH). CAMDEN CO.: Waverly, Ebinger 11228 (EIU). CANDLER CO.: Metter, Henry 7277 (PH); Hermann 10122 (GH, US). CHARLTON CO.: Folkston, Harbison 5816 (GH). COFFEE co.: Harper 687 (GH, US). EFFINGHAM CO.: Harper 961 (GH). GLYNN C0.: Emanuel Church Rd., Ebinger 11231 (EIU). JEFF DAVIS cc.: Hazlehurst, Ebinger 11195 (EIU). LANIER CO0.: Adams 50 (Ky). LIBERTY CO.: Altamaha River, Small (GH, MIN, NY, PH). LOWNDES co.: Valdosta, Beardslee, Jr. (MU). MCINTOSH CO.: Darien, Cron- quist 5366 (GH, MICH, NY, PH). TATTNALL CO.: Reidsville. Cronquist 5840 (GH, IND, MICH, NY, PH, US). THOMAS CO.: Thomasville, Taylor (GH, MIN): TIFT CO.: Tifton, Hardin & Duncan 13649 (US). WARE 1974] Genus Kalmia — Ebinger 385 co.: Suwannee Lake, Harper 48 (PH), 298 (GH, NY, PH, US). WAYNE co:. Grangerville, Ebinger 11233 (EIU). WORTH CO.: Poulan, Pollard & Maxon 563 (NY, Us). South Carolina: BEAUFORT CO.: Ferry Landing, Bell 3805 (NY). HAMPTON CO.: Hampton, Leonard & Rad- ford 1662 (ARIZ, BRY, CHRB, NHA). JASPER CO.: Ridgeland, Ebinger 11242 (EIU). Except for transfers of the specific epithet, hirsuta, to other genera, only one other name has been proposed for this taxon. Bartram (1791) used Kalmia ciliata for a plant he observed on his travels through the southeastern United States. He used this name a number of times but did not include a description (Merrill, 1945). He did, however, describe “a diminutive but elegant Kalmia" (page 18) which undoubtedly refers to K. ciliata. This description leaves no doubt that his K. ciliata is the same as K. hirsuta. The first time the name K. ciliata appears with a descrip- tion is in the work of Dumont de Courset (1811) who attributes the name to Bartram. Kalmia hirsuta has a relatively limited distribution, oc- curring along the coastal plain in southeastern United States. Small (1903, 1914) indicated that it extended from Virginia to Florida, perpetuating an error started by Gray (1856). Its occurrence in Virginia is extremely doubtful (Fernald, 1947) and is probably due to a few herbarium specimens with incomplete or erroneous collecting data. Two such specimens have been found during the present study. Both were collected during the middle of the last century and beside the collector’s name and the year col- lected include the name Virginia. Most of the specimens of Kalmia hirsuta examined are hispid on the stems, leaves, and pedicels. A few specimens were found, however, that were completely glabrous, lack- ing the long coarse hairs as well as the stipitate-glands and smaller hairs commonly found on most parts of the plant. Variation also occurs in the arrangement of the flowers on the plant. Usually the flowers are solitary in the axils of the leaves, but occasionally the flowers occur in fascicles of 2-5, while a few specimens were found in which short 386 Rhodora [Vol. 76 racemes occur in the axils of the leaves. In other specimens the flowers appeared as a terminal raceme due to a shorten- ing of the internodes and a reduction in the size of the sub- tending leaves. The pedicels also vary extensively in length. The flowers of some plants are nearly sessile, while in others the pedicels are 1.5 cm long with the two basal bracts scattered along the stalk. Genetically Kalmia hirsuta appears to be closely related to K. latifolia with which it can be crossed to produce viable hybrids with 0-10% pollen stainability. Jaynes (1968a) also reported that it can be crossed with both K. angustifolia and K. cuneata producing viable but sterile hybrids. Based on habit and morphology K. hirsuta is most closely related to K. ericoides. These species were placed in the genus Kalmiella by Small (1914) since they have a deciduous calyx and their flowers are borne singly in the axils of the leaves. Unfortunately the Cuban member of the genus is not available for genetic studies. 7. Kalmia ericoides Wright ex Griseb. Cat. Pl. Cubensium 51. 1866. Chamaedaphne ericoides (Wright ex Griseb.) Kuntze, Rev. Gen. Pl. 2: 388. 1891. Kalmiella ericoides (Wright ex Griseb.) Small, North Amer. Fl. 29: 54. 1914. TYPE: Cuba occ. in distr. Guane, in pinetis arenosis pr. La Grifa, C. Wright 2199 (GH, NY, US - Isotypes). A sparingly branched, erect to spreading shrub, to 10 dm tall with a thick basal caudex; branchlets light gray to reddish-gray, puberulent and commonly densely hispid and stipitate-glandular (rarely glabrous) ; leaves alternate, per- sistant, thick, leathery, subsessile or with a puberulent petiole to 1 mm long, linear, 4-13 mm long, 0.6-2 mm broad, the margins strongly revolute, usually lightly hispid and stipitate-glandular on both surfaces, sometimes strongly puberulent (rarely glabrous); flowers solitary (rarely in fascicles or in compact racemes) in the axils of the leaves at the end of the branches, forming a terminal cluster; pedicels to 13 mm long, commonly hispid and stipitate- 1974] Genus Kalmia — Ebinger 387 glandular, subtended by a pair of puberulent bracts 1-3 mm long; calyx green, the lobes foliaceous, lanceolate, 3-6 mm long, usually hispid and stipitate-glandular, tardily decidu- ous in fruit; corolla light pink to white, 10-17 mm broad, usually lightly hirsute and stipitate-glandular on the keels outside, lightly puberulent at the base within; anthers pur- plish brown, 0.8-1.2 mm long; filaments puberulent toward the base; ovary usually stipitate-glandular; style 4-7 mm long; fruit 2-4 mm broad, 2-3 mm long, subglobose to ovoid, lightly stipitate-glandular ; seeds reddish-brown, 0.4-0.7 mm long, the testa not loose or extending past the ends of the seed. Kalmia ericoides is the only member of the genus that has not been studied cytologically. Morphologically, how- ever, it is most closely related to K. hirsuta, a species re- stricted to southeastern United States. Both have relatively small leaves (less than 1.5 em long), that are usually hispid, their flowers are borne singly in the axils of the leaves, and the calyx is tardily deciduous. The two are easily separated, however, since the leaves of K. ericoides are thick and leathery with revolute margins while in K. hirsuta the leaves are thin and lack a strong revolute margin. Also, the flowers are borne scattered along the stem in the axils of the leaves in K. hirsuta while in K. ericoides, they are solitary in the leaf axils, but clustered toward the ends of the stems giving the appearance of a terminal corymbiform raceme. Kalmia ericoides is endemic to savannas and pine barrens of western Cuba. Although it has a very limited distribu- tion, the variation that exists in pubescence and compact- ness of the inflorescence has resulted in a division of this complex into three species by Alain (1957), while others considered it a single highly variable species (Wood, 1961). Present studies indicate that the compactness of the inflor- escence is not completely reliable in separating the taxa. The variation in leaf pubescence, however, is relatively stable. As a result, the following two varieties are recog- nized. 888 Rhodora [Vol. 76 KEY TO VARIETIES 1. Leaves glabrous or lightly hispid and with a few scat- tered stalked glandular hairs on the upper surface. .... rr rr Ta. var. ericoides. 1. Leaves densely puberulent and stipitate-glandular on the upper surface. .............. 7b. var. aggregata. Ta. Kalmia ericoides Wright ex. Griseb. var. ericoides. Kalmiella simulata Britt. & Wils. Mem. Torrey Bot. Club 16: 93. 1920. TYPE: Cuba: vicinity of Los Indios, Isle of Pines, Britton, Britton, & Wilson 14205 (NY). Shrub with ascending, hispid, stipitate-glandular and sometimes puberulent branchlets; leaves scattered, usually only 5-12 per cm of branchlet, glabrous to lightly hispid and stipitate-glandular above, not puberulent, 4-9 mm long, 0.6-1.5 mm wide; pedicels usually longer than the subtend- ing leaves, giving the inflorescence an open appearance; calyx lobes 3-4.5 mm long, hispid and lightly puberulent on the outside, usually glabrous within. Distribution: Low, white-sand savanna and pine barrens on the Isle of Pines and in the Province of Pinar del Rio, Cuba. REPRESENTATIVE SPECIMENS Cuba: PINAR DEL RIO: La Grifa, Ekman 18165 (NY, Us), Wright 2199 (GH, NY, US); Laguna de Alcatraz Grande, León & Victorin 18706 (GH, US), 20474 (GH, Us); El Sabalo, Alain 1326 (GH, US). ISLE OF PINES: Los Indios, Britton Britton, & Wilson 14205 (GH, NY), Killip 42882 (GH, NY, US), León & Victorin 17853 (GH), Vic- torin & Alain TTa (US). Kalmia ericoides var. ericoides has a very restricted range and few herbarium specimens are available. It appears to be a fairly stable variety that varies mostly in the extent of leaf pubescence. All of the specimens seen from the prov- ince of Pinar del Rio are typical of the type specimen which has a few scattered glandular hairs and is lightly hispid on the upper leaf surface. In contrast, most of the specimens of this variety from the Isle of Pines are glabrous on the 1974] Genus Kalmia — Ebinger 389 upper leaf surface. These glabrous specimens were sepa- rated as Kalmiella simulata by Britton and Wilson (Britton, 1920). These glabrous specimens all have small, narrow leaves (less than 9 mm long and 0.6-1.2 mm wide), pedicels that are much longer than the subtending leaves, and calyx lobes that are fairly short (less than 4.5 mm long) all char- acteristics of variety ericoides. 7b. Kalmia ericoides var. aggregata (Small) Ebinger, comb. nov. Kalmiella aggregata Small, North Amer. FI. 29: 54. 1914. Kalmia aggregata (Small) Copeland, Amer. Midl. Nat. 30: 571. 1943. TYPE: Cuba: pinelands, Los Indios, Isle of Pines, Jennings 324 (NY). Shrub with adscending, lightly hispid and strongly pu- berulent branchlets; leaves crowded, usually 15-25 per cm of branchlet, densely puberulent and stipitate-glandular above, 8-13 mm long, 1-2 mm wide; pedicels usually as short as the subtending leaves, giving the inflorescence a compact appearance; calyx lobes 3.5-5.5 mm long, hispid and densely puberulent on the outside, densely puberulent within. Distribution: Low, white-sand savanna and pine barrens on the Isle of Pines, Cuba. REPRESENTATIVE SPECIMENS Cuba: ISLE OF PINES: Beach at Siquanea, Morton 10098 (us); between Mina de Oro and Playa del Soldado, Killip 43905 (us), 45385 (GH); Lama Daquille, Ekman 72492 (NY); Los Indios, Jen- nings 324 (GH, NY, US), León & Victorin 18857 (GH, US), Victorin & Alain 77 (GH, NY, US), Playa Roja, Killip 48001 (GH, NY, US); San Pedro, Britton, Britton, & Wilson 14146 (GH, NY, US), Killip 43698 (US), 45566 (vs), Morton 10018 (Us), 10053 (US); Siquanea region, Killip 44043 (us), 44044 (us), 44753 (us), 44755 (US). This variety appears to be more robust than the preced- ing. Specimens to 13 dm have been reported, and the leaves and flowers are, in general, slightly larger than in variety ericoides. Besides this difference in size, all of the speci- mens of variety aggregata examined are strongly puberu- lent throughout. In particular the leaves are strongly 390 Rhodora [Vol. 76 puberulent on both surfaces, but this is also characteristic of the branches, pedicels and sepals. UNCERTAIN OR EXCLUDED NAMES Kalmia lanceolata Raf. Autikon Bot. 86. 1840. This name should probably be referred to Kalmia polifolia but the description could apply to other taxa. No specimen to represent this name has been found. Kalmia serotina Hoffmgg. Verz. Pflanzenkulturen 70. 1824. Nomen nudum. Kalmia spuria Bartr. Trav. N. & S. Carol. 308. 1791. Nomen nudum. LITERATURE CITED ABRAMS, L. 1951. Illustrated Flora of the Pacific States: Wash- ington, Oregon, and California, Vol. 3. Stanford. AITON, W. 1789. Hortus Kewensis. Vol. 2. London. ALAIN, BR. H. 1957. Ericaceae. In: LEON, BR. H. & BR. H. ALAIN, Flora de Cuba. Vol. 4. Havana. AMERICAN FORESTRY ASSOCIATION. 1955. These are the Champs. Am. Forests 61(9): 31-40. ANONYMUs. 1895. New or little-known plants. Garden Forest 8: 315. ASHLEE, T. R. 1932. 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Relationships of natu- ral vegetation to the water holding capacity of the soils in New England. Soil Sci. 55: 433-446. CoPELAND, H. F. 1943. A study, anatomical and taxonomic, of the genera of Rhododendroideae. Am. Midl. Nat. 30: 533-625. Cox, H. T. 1948. Studies in the comparative anatomy of the Ericales. I. Ericaceae — Subfamily Rhododendroideae. Am. Midl. Nat. 39: 220-245. CRAWFORD, A. C. 1908. Mountain laurel, a poisonous plant. U. S. Dept. Agr. Bur. Pl. Ind. Bull. 121: 21-35. Curtis, M. A. 1835. Enumeraticn of plants growing spontaneously around Wilmington, North Carolina, with remarks on some new and obscure species. Boston Jour. Nat. Hist. 1: 82-141. CurTis, W. 1795. Kalmia angustifolia. Narrow-leaved Kalmia. Bot. Mag. 9: 331. Davis, P. H., & V. H. HEvwoop. 1963. Principles of Angiosperm taxonomy. Princeton. DEAM, C. C. 1940. Flora of Indiana. Indianapolis. DEANE, W. 1899. Kalmia latifolia in Vermont. Rhodora 1: 136. DEGRAFFE, B. L. 1896. The tannins of some Ericaceae. Am. Jour. Pharm. 68: 313-321. DRUDE, O. 1889. Ericaceae. In: ENGLER, A., & K. PRANTL, Die naturlichen Pflanzenfamilien. IV(1): 15-65. DuprEY, T. R. 1967. Ornamental mountain-laurel and a new cul- tivar: Kalmia latifolia ‘Bettina’. Am. Hort. Mag. 46: 245-248. DUMONT DE CoursET, G. L. M. 1802. Le botaniste cultivateur. Vol. 2. Paris; 1811. Ed. 2, Vol. 3. Paris. Duncan. W. H. 1948. Preliminary reports on the flora of Georgia. I. The distribution in Georgia of Spermatophytes new to or rare in the state. Castanea 13: 70-83. EGLER, F. E. 1947. Effects of 2,4-D on woody plants in Connecti- cut. Jour. Forest. 45: 449-452. —. 1948. 2.4-D effects in Connecticut vegetation, 1947. Ecology 29: 382-386. 1949. Herbicide effects in Connecticut vegetation, 1948. Ecology 30: 248-256. . 1950. Herbicide effects in Connecticut vegetation, 1949. Bot. Gaz. 112: 76-85. ENGELMANN. G.. W. M. CANBY. & A. GRAY. 1877. Large trunks of Kalmia latifol'a. Amer. Nat. 11: 175. FERN^'Dp. M. L. 1911. 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Breaking seed dormancy of Kalmia hirsuta with high temperatures. Ecology 49: 1196-1198. 1969. Chromosome counts of Kalmia species and re- valuation of K. polifolia var. microphylla. Rhodora 71: 280-284. 1971a. Laurel selections from seed. True breeding red- budded mountain laurel. Cire. Conn. Agr. Exp. Sta. 240: 1-10. 1971b. A gene controlling pigmentation in sheep laurel. Jour. Hered. 62: 201-203. 1971c. Seed germination of six Kalmia species. Jour. Am. Soc. Hort. Sci. 96: 668-672. . 1971d. The Kalmias and their hybrids, Quart. Bull. Am. Rhododendron Soc. 25: 160-164. . 1972. Selection and propagation of improved Kalmia latifolia cultivars. Int. Pl. Prop. Soc. 21: 366-373. JENNISON, H. M. 1935. Notes on some plants of Tennessee. Rhodora 37: 309-323. JOHNSON, E. A., & J. L. KovNER. 1956. Effects on stream flow of cutting a forest understory. Forest Sci. 2: 82-91. 1974] Genus Kalmia — Ebinger 395 KENNEDY, G. W. 1875. Arbutin in Kalmia latifolia, Lin. Am. Jour. Pharm. 47: 5-6. 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Stockholm. LirkE, R. 1968. Pflanzenreste aus dem untermiozán in Nordwest- sachsen. Palaeontographica Abt. B. 123: 173-183. LovELL, J. H. 1926. Honey plants of North America. Medina. , & H. B. LovELL. 1934. The pollination of Kalmia angustifolia. Rhodora 36: 25-28. Lovett, H. B. 1951. The mechanics of pollination. Bees 5: 18-23. Lupwic, C. F. 1783. Die neuere wilde baumzucht in einem alpha- betischen und systematischen verzeichisse. Leipzig. MACOUN, J. 1884. Catalogue of Canadian Plants. Part II. Gamo- petalae. Montreal. MansH, C. D., & A. B. CLAWSON. 1930. Mountain-laurel ( Kalmia latifolia) and sheep laurel (Kalmia angustifolia) as stock- poisoning plants. U. S. Dept. Agr. Tech. Bull. 219: 1-22. MATTHEWS, J. R., & E. M. KNox. 1926. The comparative morphology of the stamens in the Ericaceae. Trans. Proc. Bot. Soc. Edinb. 29: 243-281. Matusow, H. 1897. Analysis of the root of Kalmia latifolia. Am. Jour. Pharm. 69: 341-343. MERRILL, E. D. 1945. In defense of the validity of William Bar- tram's binomials. Bartonia 23: 10-35. MEYER, B. S. 1927. The measurement of the rate of water-vapor loss from leaves under standard conditions. Am. Jour. Bot. 14: 582-591. 396 Rhodora [Vol. 76 MILLER, C. C. 1895. Honey from mountain laurel. Am. Bee Jour. 35: 825-826. MOUILLEFERT, P. 1897. Traite des Arbres & Arbrisse. Vol. II. Paris. MUENSCHER, W. C. 1957. Poisonous plants of the United States. Revised ed. New York. Munns, E. N. 1938. The distribution of important forest trees of the United States. U. S. Dept. Agr. Misc. Publ. 287: 1-176. NICHOLS, G. E. 1934. The influence of exposure to winter tempera- ture upon seed germination in various native American plants. Ecology 15: 364-373. NICHOLSON, G. 1896. Hand-list of trees and shrubs grown in Arbo- retum. Part II. Gamopetalae to Monocotyledons. Royal Gardens, Kew. NICHOLSON, S. M, & J. F. CrLovis. 1967. Dye plants and dye methods in West Virginia. Castanea 32: 111-116. NORTON, A. H. 1913. 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J. 1788. Beschreibung der Polenblattrigen Kalmia und der gelbbluhenden Rosskastanie. Schr. Berlin Ges. Naturf. Freunde 8: 129-133. Waup, R. A. 1940. The action of Kalmia angustifolia (Lambkill). Jour. Pharm. Exp. Therap. 69: 103-111. Woop, C. E. JR. 1961. The genera of Ericaceae in the South- eastern United States. Jour. Arnold Arb. 42: 10-80. Woop, G. B., & F. BACHE. 1851. The Dispensatory of the United States of America. Philadelphia. Woop, H. B., V. L. STROMBERG, J. C. KERESZTESY, & E. C. HORNING. 1954. Andromedotoxin. A potent hypotensive agent from Rhodo- dendron maximum. Jour.“Am. Chem. Soc. 76: 5689-5692. Woop, T. F. 1883. Is sheep laurel poisonous to sheep? Am. Agr. 42: 66. EASTERN ILLINOIS UNIVERSITY CHARLESTON, ILLINOIS 61920 MARINE ALGAE OF RHODE ISLAND R. D. Woop AND MARTINE VILLALARD-BOHNSACK With increasing marine research activity in the Narra- gansett Bay area, the need for information about local marine algae has become urgent. Therefore, the available information has been compiled (Wood, in press) and a list of species known to occur in the State is presented. The earliest serious collecting of algae in Rhode Island seems to have been by S. T. Olney and his associates, and considerable effort was made in the years of 1846-1849. But, also, during the summer of 1846, J. W. Bailey visited Rhode Island and collected with Olney. Bailey, working independently, obtained specimens from Europe and worked up the collections systematically and included the Rhode Island records in his publication of 1847. Olney, on the other hand, sent his material to W. H. Harvey of Dublin, who identified them. The records were published by Olney in 1847, the same year that Bailey's appeared. Whose work appeared first is not clear, but Olney was the first Rhode Islander to publish on the marine algae of the State, while Bailey was probably the first worker to both study and publish on algae of Rhode Island. Subsequent records are those by Harvey (1852-1858), Olney (1871), Farlow (1873, 1875, 1876, and 1881), and Bennett (1888), and the several reports by Collins between 1882 and 1896 which were largely summarized in his works of 1900 and 1909-1918. More recent reports are those by Wood & Palmatier (1954), the dissertation by Villalard (1971), and thesis by Vaughan (1971). In addition, three collections of concern are the *Algae Rhodiaceae" by Olney (1871) and the two large exsiccata sets in which Rhode Island algae occur — Farlow, Anderson & Eaton’s (1877-1889) “Algae Exsiccatae Ameri- cae-Borealis” and Collins, Holden & Setchell’s (1895-1919) “Phycotheca Boreali-Americana.” The senior author (rdw) prepared a list of species re- corded from Rhode Island by Taylor (1957), and then 399 400 Rhodora [Vol. 76 searched the literature for other early records. The sup- porting specimens were sought and examined, especially those at Brown University (courtesy of G. L. Church and W. H. Snell) and at the Farlow Herbarium at Harvard University (courtesy of I. M. Lamb). The complex of iden- tifications and synonyms required careful tracing, but veri- fication of most early names by examination of the actual collections was possible. The background information is to appear elsewhere (Wood, in press). The final list was checked by the junior author (mvb), and the latter also compiled the currently suggested synonyms that appear parenthetically in the list. Lastly, an evaluation of the frequency of occurrence of each taxon, based upon the authors’ personal observations, was included. Where identi- fications were still uncertain, Dr. Wm, Randolph Taylor graciously aided. The final manuscript was examined and edited by Urda K. Wood. It is hoped that this list will provide a background for future work on the algae of Rhode Island waters. LIST OF MARINE ALGAE KNOWN FOR RHODE ISLAND The nomenclature below follows Taylor (1957), but other suggested names and references are indicated parentheti- cally. However, author citations have been checked, and those involving “in” and “ex” entries include both authors. Species reported for the first time in Rhode Island are desig- nated by an asterisk (*). The earliest known published rec- ord for each species in Rhode Island is cited in the second column, the abbreviations being: B — Bailey (1847), BENN. = Bennett (1888), B,o = Bailey (1847) and Olney (1847), c = Collins (1900), F = Farlow (1873), o = Olney (1847), OAR [for Olney’s “Algae Rhodiaceae"] = Olney (1871), PBA = in Collins, Holden & Setchell (1895-1919), T = Taylor (1937), (VA) = thesis by Vaughan (1971), (VI) = disserta- tion by Villalard (1971), W = Wood (1962), w&P = Wood & Palmatier (1954). Species which should be expected in the state, based on 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 401 Taylors (1957) range records, but for which no adequate report is yet available, are summarized at the end of the list. The frequency of occurrence of each species based on our combined personal observations is indicated in the third column as: t = trace, rare, + = local and uncommon, ++ = locally common, +++ = general and quite common. Where a rare taxon has been seen by only one of us, it is indicated as by “rdw” or “mvb” for R. D. Wood or M. Villalard-Bohnsack, respectively. CHLOROPHYTA Ulotrichaceae 1. Stichococcus marinus (Wille) Hazen T + 2. Ulothrix flacca (Dillw.) Thuret in Le Jol. C dar Chaetophoraceae 3. Pseudendoclonium submarinum Wille T Gomontiaceae 4. Gomontia polyrhiza (Lagerh.) Born. et Flah. T Ulvaceae * 5. Capsosiphon fulvescens (C.Ag.) Setch. et Gard. (VI) * * 6. [Enteromorpha ahlneriana Blid.] (VI) t 7. E. clathrata (Roth) J. Ag. OAR + (E. clathrata (Roth) Grev. — Parke & Dixon, 1964) 8. E. compressa (L.) Grev. B,O + (ecotype of E. intestinalis (L.) Link — DaSilva & Burrows in Norton, 1972, p. 385) 9. E. cruciata Collins OAR 402 10 Rhodora . E. erecta, (Lyngb.) J. Ag. (E. flexuosa ssp. paradoxa (Dillw.) Blid. — Bliding, 1963) * 11. E. groenlandica (J.Ag.) Setch. 12. 13. 14. 15. 16. 17. 18. 20. 21. 23. 24. et Gard. E. intestinalis (L.) Link E. linza (L.) J. Ag. E. marginata J. Ag. (Blidingia marginata (J. Ag.) Dang. — Dangeard, 1958) E. minima Nag. (Blidingia minima (Nag. ex Kitz.) Kylin — Kylin, 1947) E. prolifera (Müll.) J. Ag. Monostroma fuscum (Post. et Rupr.) Witt. (Ulvaria obscura (Kütz.) Gayr. — Gayral, 1965) M. grevillei (Thuret) Witt. (Ulvopsis grevillei (Thuret) Gayr. — Gayral, 1965) . M. leptodermum Kjellm. (Kornmannia leptoderma (Kjellm.) Blid. — Bliding, 1968) M. oxyspermum (Kütz.) Doty (Ulvaria oxyspermum (Kiitz.) Blid. — Bliding, 1968) M. pulchrum Farlow . Percursaria percursa (C. Ag.) Rosenv. Ulva lactuca L. var. latissima (L.) DC. Prasiolaceae Prasiola stipitata Suhr. in Jess. B,O (VI) B,O B [Vol. 76 +++ ++ 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 403 Cladophoraceae 25. Chaetomorpha aerea (Dillw.) Kütz. B + (Ch. linum (Müll. Kütz. — Christensen, 1957) 26. Ch. linum (Miill.) Kütz. B ++ 27. Ch. melagonium (Web. et Mohr) Kiitz. t(rdw) 28. Cladophora albida (Huds.) Kiitz. F tT 29. Cl. flexuosa (Dillw.) Harv. C + (Cl. serica, (Huds.) Kütz. — Hoek, 1963) f. densa Collins PBA 30. Cl. glaucescens (Harv.) Harv. F (Cl. albida (Huds.) Kutz. — Hoek, 1963) 31. Cl. gracilis (Griff. ex Harv.) Kütz. [o rain (Cl. serica (Huds.) Kütz. — Hoek, 1963) 32. Cl. hutchinsiae (Dillw.) Kütz. var. distans (C. Ag.) Kütz. T (Cl. hutchinsiae (Dillw.) Kütz. — Hoek, 1963) * 83. Cl. laetevirens (Dillw.) Kütz. (VI) m 34. Cl. magdalenae Harv. F (Cl. aibida (Huds.) Kütz. — Hoek, 1963) 35. Cl. refracta (Roth) Kütz. O (Cl. albida (Huds.) Kütz. — Hoek, 1963) 36. Cl. rudolphiana (C. Ag.) Harv. C (Cl. serica (Huds.) Kütz. — Hoek, 1963) 37. Cl. rupestris (L.) Kütz. OAR zu 38. Rhizoclonium riparium (Roth) Harv. C t 404 * 89. 40. * A1. * 42. 43. 44. 45. 46. Rhodora var. implexum (Dillw.) Rosenv. var. polyrhizum (Lyngb.) Rosenv. R. tortuosum Kütz. (R. riparium (Roth) Harv. — Scagel, 1966) Spongomorpha arcta (Dillw.) Kütz. (Acrosiphonia arcta (Dillw.) J. Ag. — Kornmann, 1962) S. lanosa (Roth) Kütz. (S. aeruginosa (L.) Hoek — Hoek, 1963) S. spinescens Kütz. (Acrosiphonia arcta (Dillw.) J. Ag. — Kornmann, 1962) Urospora penicilliformis (Roth) Aresch. Bryopsidaceae Bryopsis plumosa (Huds.) C. Ag. Codiaceae Codium fragile (Sur.) Hariot ssp. tomentosoides (van Goor) Silva PHAEOPHYTA Ectocarpaceae Ectocarpus confervoides (Roth) Le Jol. (E. siliculosus (Dillw.) Lyngb. — South & Cardinal, 1970) var. pygmaeus (Aresch.) Kjellm. B,O (VI) 0?,c [Vol. 76 ++ t(mvb) +++ ++ 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 405 4T. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. E. dasycarpus Kuck. E. fasciculatus Harv. E. penicillatus (C. Ag.) Kjellm. E. siliculosus (Dillw.) Lyngb. E. tomentosus (Huds.) Lyngb. (Spongonema tomentosum (Huds.) Kütz. — Kuckuck, 1960) Giffordia granulosa (Smith) Hamel G. mitchellae (Harv.) Hamel (G. mitchelliae (Harv.) Hamel — Earle, 1969) G. ovata (Kjellm.) Kylin G. secunda (Kiitz.) Batt. Pylaiella littoralis (L.) Kjellm. (Pilayella littoralis (L.) Kjellm. — Dixon & Russell, 1964) Sorocarpus micromorus (Bory) Silva Streblonema aecidioides (Rosenv.) Fosl. (Entonema aecidioides (Rosenv.) Kjellm. — Edelstein & McLachlan, 1968) . S. chordariae (Farl.) De Toni Sphacelariaceae . Cladostephus verticillatus (Lightf.) Lyngb. . Sphacelaria cirrosa (Roth) C. Ag. . S. radicans (Dillw.) C. Ag. Myrionemataceae . Ascocyclus orbicularis Magnus (Myrionema magnusii (Sauv.) Lois. — Loiseaux, 1967) B,O ++ + (rdw) 406 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 14. 75. 76. 77. Rhodora Hecatonema terminalis (Kiitz.) Kylin Myrionema corunnae Sauv. M. foecundum (Stromf.) Fosl. (Hecatonema foecundum (Strómf.) Lois. — Parke & Dixon, 1968) M. globosum (Reinke) Fosl. (Microsporangium globosum Reinke — Parke & Dixon, 1964) M. strangulans Grev. Ralfsiaceae Ralfsia borneti Kuck. R. clavata (Harv. in Hook.) Crouan frat. R. pusilla (Strómf.) Batt. R. verrucosa (Aresch.) J. Ag. Elachisteaceae (Elachistaceae — Parke & Dixon (1964) ) PBA Elachistea chondri Aresch. (Elachista chondri Aresch. — Fritsch, 1945; Parke & Dixon, 1964) E. fucicola (Vell.) Aresch. ( Elaschista fucicola Aresch. — Fritsch, 1945; Parke & Dixon, 1964) Halothrix lumbricalis (Kütz.) Reinke Chordariaceae Chordaria flagelliformis (Müll.) C. Ag. var. densa Farl. Eudesme virescens (Carm. ex Harv. in Hook.) J. Ag. Q [Vol. 76 ++ ++ ++ 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 407 78. Leathesia difformis (L.) Aresch. OAR ++ 79. Sphaerotrichia divaricata (C. Ag.) Kylin OAR + Stilophoraceae 80. Stilophora rhizodes (Turn.) J. Ag. OAR Desmarestiaceae 81. Arthrocladia villosa (Huds.) Duby C t 82. Desmarestia aculeata (L.) Lamour. B + 83. D. viridis (Müll. Lamour. B "n Striariaceae 84. Striaria attenuata (Grev.) Grev. C Punctariaceae * 85. Asperococcus echinatus (Mert.) Grev. (VI) Tom (A. fistulosus (Huds.) Hook. — Edelstein, Wynne & McLachlan, 1970) 86. Desmotrichum undulatum (J. Ag.) Reinke B,O 87. Litosiphon filiformis (Reinke) Batt. PBA 88. Myriotrichia clavaeformis Harv. C 89. M. filiformis Harv. F 90. Petalonia fascia (Müll. Kuntze BENN. ++ 91. Punctaria latifolia Grev. BENN. + 92. P. plantaginea (Roth) Grev. O ule ce 93. Rhadinocladia cylindrica Schuh T 94. R. farlowii Schuh T 408 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. *106. 107. 108. 109. Rhodora Scytosiphon lomentaria (Lyngb.) Link (S. lomentarius (Lyngb.) Link — Farlow, 1881) Dictyosiphonaceae Dictyosiphon chordaria Aresch. (as Gobia) D. foeniculaceus (Huds.) Grev. Laminariaceae Alaria esculenta (L.) Grev. Chorda filum (L.) Lamour. C. tomentosa Lyngb. Laminaria agardhii Kjellm. (L. saccharina (L.) Lamour. — Wilce, 1965) L. digitata (Huds.) Lamour. L. intermedia Fosl. (L. digitata (Huds.) Lamour. — Sundene, 1958) L. longicruris De la Pyl. L. saccharina (L.) Lamour. Fucaceae Ascophyllum mackaii (Turn.) Holmes et Batt. (A. nodosum (L.) Le Jol. — Gibb, 1957) A. nodosum (L.) Le Jol. Fucus edentatus De la Pyl. (F. distichus L. ssp. edentatus (De la Pyl.) Pow. — Powell, 1963) F. evanescens C. Ag. (F. distichus L. ssp. evanescens (C. Ag.) Pow. — Powell, 1963) B,O B,O [Vol. 76 ++ +++ +++ ++ + (rdw) ++, +++ TG +? 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 409 110. F. spiralis L. B res var. limitaneus Montagn. T 111. F. vesiculosus L. B,O ro Sargassaceae 112. Sargassum filipendula C. Ag. B,O + *113. S. natans (L.) J. Meyen (VA) t(drift) RHODOPHYTA Bangiaceae 114. Bangia ciliaris Carm. ex Harv. C t (Erythrotrichia ciliaris (Carm. ex Harv.) Thur. in Le Jol. — Parke & Dixon, 1964) 115. B. fuscopurpurea (Dillw.) Lyngb. B,O FT 116. Erythrotrichia carnea (Dillw.) J. Ag. C 117. Goniotrichum alsidit (Zanard.) Howe C t 118. Porphyra leucosticta Thuret in Le Jol. t *119. P. miniata (C. Ag.) C. Ag. (VI) x 120. P. umbilicalis (L.) J. Ag. B,O dT Acrochaetiaceae 121. Acrochaetium daviesti (Dillw.) Nag. C 122. A. flexuosum Vickers T 123. Kylinia secundatum (Lyngb.) Papenf. C (Acrochaetium secundatum (Lyngb.) Nig. — Parke & Dixon, 1968) 124. K. virgatula (Harv.) Papenf. C (Acrochaetium virgatula (Harv.) J. Ag. — Parke & Dixon, 1968) 410 125. 126. 127. 128. *129. 131. 132. 133. Rhodora Rhodochorton penicilliforme (Kjellm.) Rosenv. R. purpureum (Lightf.) Rosenv. Helminthocladiaceae Nemalion multifidum (Web. et Mohr) J. Ag. (N. helminthoides (Vell. in With.) Batt. — Parke & Dixon, 1964) Chaetangiaceae Scinaia furcellata (Turn.) Bivona Bonnemaisoniaceae Asparagopsis hamifera (Hariot) Okam. (VI), (VA) (Bonnemaisonia hamifera Har. — Feldmann & Feldmann, 1942) Gelidiaceae . Gelidium crinale (Turn.) Lamour. Dumontiaceae Dumontia incrassata (Miill.) Lamour. Rhizophyllidaceae Polyides caprinus (Gunn.) Papenf. (P. rotundus (Huds.) Grev. — Parke, 1953) Squamariaceae Hildenbrandia prototypus Nardo [Vol. 76 ++ + (mvb) ++ ++ +++ 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 411 134. Rhododermis georgii (Batt.) Collins T (Rhodophysema georgii Batt. —Dixon, 1964) Corallinaceae 135. Corallina officinalis L. BENN. mtm 136. Fosliella farinosa (Lamour.) Howe e (Melobesia farinosa Lamour. — Hamel & Lemoine, 1952) 137. F.lejolisii (Rosan.) Howe C + (Melobesia lejolisit Rosan. — Hamel & Lemoine, 1952) 138. Lithophyllum corallinae (Crouan frat.) Heydr. PBA 139. L. macrocarpum (Rosan.) Fosl. T (Dermatolithon pustulatum (Lamour.) Fos]. — Edelstein & McLachlan, 1968) 140. L. pustulatum (Lamour.) Fosl. OAR *141. L. lenormandi (Aresch. in J. Ag.) Fosl. (VI) oe (Phymatolithon lenormandii (Aresch. in J. Ag.) Adey — Adey, 1966) 142. Melobesia membranacea (Esper) Lamour. C (Epilithon membranaceum (Esper) Heydr. — Parke & Dixon, 1964) 143. Phymatolithon laevigatum (Fosl.) Fosl. T ++ (Lithothamnion laevigatum Fos]. — Parke & Dixon, 1964) 412 144 145 146. 147. 148. 149. 150. 151. Rhodora P. polymorphum (L.) Fosl. C (Lithothamnion polymorphum (L.) Aresch. in J. Ag. — Parke & Dixon, 1964) Gloiosiphoniaceae Gloiosiphonia capillaris (Huds.) Carm. ex Berk. C Kallymeniaceae Euthora cristata (C. Ag.) J. Ag. F Choreocolacaceae Choreocolax polysiphoniae Reinsch C Harveyella mirabilis (Reinsch) Reinke T Solieriaceae Agardhiella tenera (J. Ag.) Schmitz B (as used by Taylor (1957), but designated A. baileyi (Harv. ex Kütz.) Taylor in Taylor & Rhyne (1970: 13), and since changed to Neoagardhiella baileyi (Harv. ex Kütz.) Wynne & Taylor, Wynne & Taylor, 1973, p. 101) Rhodophyllidaceae Cystoclonium purpureum (Huds.) Batt. B,O var. cirrhosum Harv. C Hypneaceae Hypnea musciformis (Wulf.) Lamour. BENN. [Vol. 76 t—+ (periodic) t(drift?) ab ++ +++ 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 413 Gracilariaceae 152. Gracilaria foliifera (Forssk.) Borg. (0) t 153. G. verrucosa (Huds.) Papenf. W&P +++ Phyllophoraceae 154. Ahnfeltia plicata (Huds.) Fries B,O T 155. Gymnogongrus griffithsiae (Turn.) Mart. C t(mvb) 156. Phyllophora brodiaei (Turn.) Endl. F + (P. truncata (Pallas) Newr. et Taylor — Newroth & Taylor, 1971) var. catenata (C. Ag.) Collins T 157. P. membranifolia (Good. et Woodw.) J. Ag. 0 mo (P. pseudoceranoides (Gmel.) Newr. et Taylor — Newroth & Taylor, 1971) 158. P. traillii Holmes ex Batt. C Gigartinaceae 159. Chondrus crispus Stackh. B,O ttt 160. Gigartina stellata (Stackh.) Batt. G M (Palmaria palmata (L.) Stackhouse — Guiry, 1974, p. 219.) Rhodymeniaceae 161. Rhodymenia palmata (L.) Grev. B,O TARERE Champiaceae 162. Champia parvula (C. Ag.) Harv. B,O bad a 163. Lomentaria baileyana (Harv.) Farl. OAR TRT 414 *164. 165 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. Rhodora [L. clavellosa (Turn.) Gail.] L. orcadensis (Harv.) Coll. ex Taylor Ceramiaceae Antithamnion americanum (Harv.) Farl. in Kjellm. A. cruciatum (C. Ag.) Nag. A. plumula (Ellis) Thuret in Le Jol. A. pylaisaei (Mont.) Kjellm. Callithamnion baileyi Harv. var. laxum Farlow C. byssoides Arnott ex Harv. in Hook. C. corymbosum (Smith) Lyngb. C. roseum (Roth) Lyngb. C. tetragonum (With.) C. Ag. Ceramium deslongchampii Chauv. in Duby C. diaphanum (Lightf.) Roth C. fastigiatum Harv. C. rubriforme Kylin (prox.) C. rubrum (Huds.) C. Ag. var. pedicellatum Duby (C. pedicellatum DC. — Parke & Dixon, 1964) var. squarrosum Harv. C. strictum Harv. Griffithsia globulifera Harv. G. tenuis C. Ag. Pleonosporium borreri (Smith) Näg. ex Hauck Plumaria elegans (Bonnem.) Schmitz Ptilota serrata Kütz. [Vol. 76 + (mvb) OAR C BENN. + F C OAR t T OAR B =t- C + O O B,O +,+ + OAR ++ (VA) t? B,O +++ C OAR C + C + O0 t OAR F Tte B?,F t 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 415 186. 187. 188. *189. 190. 191. 193. 194. 195. 196. 197. 198 199 Seirospora griffithsiana Harv. (S. seirosperma (Harv.) Dixon — Dixon, 1964) Spermothamnion turneri (Mert.) Aresch. (S. repens (Dillw.) Rosenv. — Rosenvinge, 1924) Spyridia filamentosa (Wulf.) Harv. in Hook. T railliella, intricata (J. Ag.) Batt. (sporophyte of Asparagopsis hamifera — Taylor, 1957, p. 291) Delesseriaceae Grinnellia americana (C. Ag.) Harv. Phycodrys rubens (L.) Batt. Dasyaceae . Dasya pedicellata (C. Ag.) C. Ag. (D. baillouviana (Gmel.) Mont. — Dixon & Irvine, 1970) Rhodomelaceae Chondria baileyana (Mont.) Harv. C. dasyphylla (Woodw.) C. Ag. C. sedifolia Harv. C. tenuissima (Good. et Wooaw.) C. Ag. Polysiphonia denudata (Dillw.) Grev. ex Harv. in Hook. . P. elongata Harv. in Hook. . P. fibrillosa (Dillw.) Spreng. OAR (VI) B,O ++ ++ ++ ob ++ 416 Rhodora [Vol. 76 200. P. harveyi Bail. 0 tt var. olneyi (Harv.) Collins O 201. P. lanosa (L.) Tandy B,O ++, +++ 202. P. nigra (Huds.) Batt. F + 203. P. nigrescens (Huds.) Grev. B + 204. P. novae-angliae Taylor B 205. P. subtilissima Mont. F + 206. P. urceolata (Lightf. ex Dillw.) Grev. C ++ 207. Rhodomela confervoides (Huds.) Silva B,O +++ 208. R. virgata Kjellm. C SPECIES TO BE EXPECTED IN RHODE ISLAND, AND REPORTED FROM ADJACENT STATES BY TAYLOR (1957) CHLOROPHYTA Chlorococcaceae Codiolum gregarium A. Br. Ulotrichaceae Ulothrix implexa Kütz. Chaetophoraceae Bolbocoleon piliferum Pringsh. Entocladia flustrae (Reinke) Batt. (Epicladia flustrae Reinke — Parke & Dixon, 1968) E. wittrockii Wille (Ectochaete wittrockii (Wille) Kylin — Parke & Dixon, 1968) Pringsheimiella scutata (Reinke) Marchew. Protoderma marinum Reinke (Pseudendoclonium marinum (Reinke) Aleem et Schulz — Loiseaux, 1967) Ulvaceae Enteromorpha plumosa Kütz. (E. flexuosa ssp. paradoxa (Dillw.) Blid. — Bliding, 1963) Cladophoraceae Chaetomorpha atrovirens Taylor Cladophora crystallina (Roth) Kütz. (Cl. vagabunda (L.) Hoek; Cl. glomerata (L.) Kütz. — Hoek, 1963) Cl. expansa (Mert.) Kütz. (Cl. vagabunda (L.) Hoek — Hoek, 1963) 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack Cl. flavescens (Roth) Kütz. (a doubtful name — Hoek, 1963) Derbesiaceae Derbesia vaucheriaeformis (Harv.) J. Ag. Bryopsidaceae Bryopsis hypnoides Lamour. Phyllosiphonaceae Ostreobium quekettii Born. et Flah. XANTHOPHYTA Vaucheriaceae Vaucheria litorea C. Ag. V. thuretii Woronin PHAEOPHYTA Ectocarpaceae Ectocarpus subcorymbosus Farl. (Farlowiella onusta (Kütz.) Kuck. — Kuckuck, 1956) Elachisteaceae (Elachistaceae — Parke & Dixon, 1964) Myriactula minor (Farl. Taylor Chordariaceae Acrothrix novae-angliae Taylor Eudesme zosterae (Lyngb.) Kylin (Cladosiphon zosterae (J. Ag.) Kylin — Parke, 1953) Striariaceae Stictyosiphon subsimplex Holden Punctariaceae Desmotrichum balticum Kiitz. (D. undulatum (J. Ag.) Reinke — Kylin, 1947a) Sargassaceae Sargassum fluitans Borg. RHODOPHYTA Bangiaceae Asterocytis ramosa (Thwaites in Harv.) Gobi ex Schmitz Acrochaetiaceae Audouinella membranacea (Magnus) Papenf. 417 418 Rhodora [Vol. 76 Corallinaceae Lithothamnium laeve (Strómf.) Fosl. (Leptophytum laeve (Strómf.) Adey — Adey, 1966) Ceramiaceae Callithamnion circinatum (Kütz.) J. Ag. Delesseriaceae Membranoptera denticulata (Mont.) Kylin Rhodomelaceae Bostrychia rivularis Harv. Polysiphonia flexicaulis (Harv.) Collins LITERATURE CITED ADEY, W. H. 1966. The genera Lithothamnion, Leptophytum (nov. gen.) and Phymatolithon in the Gulf of Maine. Hydrobiologia 28(3-4): 321-370. BAILEY, J. W. 1847. Notes on the algae of the United States, Am. J. Sci. & Arts 2: 80-85, 399-403. BENNETT, J. L. 1888. Plants of Rhode Island, being an enumeration of plants growing without cultivation in the State of Rhode Island. Published by the author, Providence Press Co., Provi- dence. 128 pp. BLIDING, C. 1963. A critical survey of European taxa in Ulvales, I, Capsosiphon, Percursaria, Blidingia, Enteromorpha. Opera Bot. 8(3): 1-160. 1968. A critical survey of European taxa in Ulvales, II, Ulva, Ulvaria, Monostroma, Kornmannia, Bot. Not. 121: 535-629. CHRISTENSEN, T. 1957. Chaetomorpha linum in the attached state. Bot. Tidsskr. 53: 311-316. CoLLINS, F. S. 1900. Preliminary lists of New England plants, V. Marine algae. Rhodora 2: 41-52. 1909-1918. The green algae of North America. Tufts Coll. St. 2(3): 79-480; First Suppl, ibid. 3(2): 69-109, 1912; Second Suppl., ibid. 4(7): 1-106, 1918. CoLLINS, F. S., I. HOLDEN, & W. A. SETCHELL. 1895-1919. Phyco- theca Boreali-Americana. (Exsiccata) Fasc. 1-46 + A-E. Malden. DANGEARD, P. 1958. La reproduction et le développement de l'Enteromorpha marginata J. Ag. et le rattachement de cette espéce au genre Blidingia. C. R. Acad. Sci., Paris 246: 347-351. DixoN, P. S. 1964. Taxonomic and nomenclatural notes on the Florideae IV. Bot. Not. 117: 56-78. 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 419 Drxon, P. S., & L. M. IRVINE. 1970. Miscellaneous notes on algal taxonomy and nomenclature, III. Bot. Not. 123(4): 474-487. DixoN, P. S, & G. RUSSELL. 1964. Miscellaneous notes on algal taxonomy and nomenclature, I. Bot. Not. 117(3): 279-284. EARLE, S. A. 1969. Phaeophyta of the eastern Gulf of Mexico. Phycologia 7(2): 71-254. EDELSTEIN, T., & J. MCLACHLAN. 1968. Investigations of the marine algae of Nova Scotia, V. Additional species new or rare to Nova Scotia. Can. J. Bot. 46: 993-1003. EDELSTEIN, T., M. J. WYNNE, & J. MCLACHLAN. 1970. Melanosiphon intestinalis (Saund.) Wynne, a new record for the Atlantic. Phycologia 9(1): 5-9. Fartow, W. G. 1873. List of seaweeds or marine algae of the south coast of New England. Rep. U. S. Comm. Fish & Fisheries 1: 281-294, 1875. List of the marine algae of the United States with notes of new or imperfectly known species. Proc. Am. Acad. Arts & Sci. 10: 351-380. . 1876. List of the marine algae of the United States. Rep. U. S. Comm. Fish & Fisheries 5: 691-718. 1881. The marine algae of New England. Rep. U. S. Comm. Fish & Fisheries for 1879, Append. A-1: 1-210. FARLOW, W. G., C. L. ANDERSON, & D. C. EATON. 1877-1889. Algae Exsiccatae Americae-Borealis. (Exsiccata) Fasc. 1-5. Boston. FELDMANN, J., & G. FELDMANN. 1942. Recherches sur les Bonne- maisoniacées et leur alternance de générations. Ann. Sci. Nat. Bot., Ser. II, 3: 75-175. FRITSCH, F. E. 1945. The structure and reproduction of the algae. Vol. II. The Univ. Press, Cambridge. 939 pp. GAYRAL, P. 1965. Monostroma Thuret, Ulvaria Rupr., emend. Gayral, Ulvopsis Gayral. (Chlorophycées, Ulotrichales): struc- ture, reproduction, cycles, position systematique. Rev. Gén. Bot. 72 (857): 627-638. GIBB, D. C. 1957. The free-living forms of Ascophyllum nodosum (L.) Le Jol. J. Ecol. 45(1): 49-83. Guiry, M. D. 1974. A reappraisal of the genus Palmaria Stack- house. Brit. Phyc. J. 9(2): 219. : HAMEL, G., & P. LEMOINE. 1952. Corallinacées de France et d'Afrique du Nord. Arch. Mus. Natn. Hist. Nat., Paris 7(1): 17-136. Harvey, W. H. 1852-1858. Nereis boreali-americana. Part I. Smithsonian Contrib. Knowledge 3(4): 1-150; Part II, ibid., 5(5): 1-258, 1853; Part III, tbid., 10: 1-140, 1858 (including supplements). HokK, C. VAN DEN. 1963. Revision of the European species of Cladophora. E. J. Brill, Leiden. 248 pp. 420 Rhodora [Vol. 76 KORNMANN, P. 1962. Eine Revision der Gattung Acrosiphonia. Helgolànd. Wiss. Meeres. 8: 219-242. Kuckuck, P. 1956. Ectocarpaceen-Studien IV. Herponema, Kutz- ingiella nov. gen., Farlowiella nov. gen. herausgegeben von Peter Kornmann. Helgolànd. Wiss. Meeres. 5: 292-325. 1960. Eectocarpaceen-Studien VI. Spongonema. Helgo- lànd. Wiss. Meeres. 7: 93-113. KvLIN, H. 1947. Ueber die Fortpflanzungsverhaltnisse in der Ord- nung Ulvales. K. Fys. Sallsk. Lund Fórh. 17: 174-182. 1947a. Die Phaeophyceen der Schwedischen Westküste. Lunds Univ. Arsskr. N. F., Avd. 2, 43(4) : 1-99. LorsEAUX, S. 1967. Morphologie et cytologie des Myrionémacées. Critères taxonomiques. Rev. Gén. Bot. 74(878): 329-947. Lunp, S. 1959. The marine algae of East Greenland, I. Taxo- nomical part. Medd. Gronland 156(1): 1-248. NEwRoTH, P. R., & A. R. A. TAYLOR. 1971. The nomenclature of the North American species of Phyllophora Greville. Phycologia 10(1): 93-97. Norton, T. A. 1972. The marine algae of Lewis and Harris in the Outer Hebrides. British Phycol. J. 7(3): 375-385, OLNEY, S. T. 1847. Rhode Island plants, 1846, or additions to the published lists of the Providence Franklin Society. Proc. Provi- dence Franklin Soc. 1(2): 25-42. 1871. Algae Rhodiaceae. A list of Rhode Island algae, collected and prepared by Stephen T. Olney, in the years 1846- 1849, now distributed from his own herbarium. Published by the author, printed by Hammond, Angell & Co., Providence. 13 pp. (Exsiccata, few sets). PARKE, M. 1953. A preliminary check-list of British marine algae. J. Mar. Biol. Ass. U. K. 32: 497-520. PARKE, M., & P. S. Dixon. 1964. A revised check-list of British marine algae. J. Mar. Biol. Ass. U. K. 44: 499-542. 1968. Check-list of British marine algae — second re- vision. J. Mar. Biol. Ass. U. K. 48: 783-832. PowELL, H. T. 1963. Speciation in the genus Fucus L. and related genera. In: J. P. Harding & N. Tebble (ed.), Speciation in the sea. Syst. Ass. Publ. 5: 63-77. RosENvINGE, K. L. 1924. The marine algae of Denmark. Vol. III. Rhodophyceae. III. (Ceramiales). K. Danske Vidensk. Selsk., Ser. Nat. 7: 287-486. ScaGeL, R. F. 1966. Marine algae of British Columbia and northern Washington. Part I: Chlorophyceae (green algae). Nat. Mus. Can., Biol. Ser. No. 74, Bull. 207: 1-257. SOUTH, R., & A. CARDINAL. 1970. A check list of marine algae of eastern Canada. Can. J. Bot. 48: 2077-2095. 1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 421 SUNDENE, O. 1958. Interfertility between forms of Laminaria digi- tata. Nytt. Mag. Bot. 6: 121-128. TAYLOR, W. R. 1937. Marine algae of the northeastern coast of North America. Univ. Michigan Press, Ann Arbor. 427 pp. 1957. Ibid., 2nd ed. 509 pp. TAYLOR, W. R., & C. F. RHYNE. 1970. Marine algae of Dominica. Smithsonian Contrib. Botany 3: 1-16. VAUGHAN, C. F. 1971. An ecological survey of the macroscopic algae of Point Judith Pond (Salt Pond), Rhode Island. M. S. Thesis, University of Rhode Island. 179 pp. VILLALARD, M. 1971. Phytosociology of intertidal algae of Conani- cut Island, Rhode Island. Ph.D. Thesis. University of Rhode Island. 227 pp. WiLCE, R. T. 1965. Studies in the genus Laminaria. III. A re- vision of the North Atlantic species of the Simplices section of Laminaria. Bot. Goteborg. 3: 247-256. Woop, R. D. 1962. Codium is carried to Cape Cod. Bull. Torrey Bot. Club 89: 178-180. ————— ————. (in press). Information for a list of marine algae of Rhode Island. New England Marine Resources Information Pro- gram, University of Rhode Island, Kingston. Woop, R. D, & E. A. PALMATIER. 1954. Macroscopic algae of the coastal ponds of Rhode Island. Am. J. Bot. 41(2): 135-142. WYNNE, M. J., & W. R. TAYLOR. 1973. The status of Agardhiella tenera and Agardhiella baileyi (Rhodophyta, Gigartinales). Hydrobiologia 43(15): 93-107. R. D. WOOD DEPT. OF BOTANY UNIVERSITY OF RHODE ISLAND KINGSTON, RHODE ISLAND 02881 M. VILLALARD-BOHNSACK DEPT. OF BIOLOGY ROGER WILLIAMS COLLEGE BRISTOL, RHODE ISLAND 02809 A BIOSYSTEMATIC REVISION OF LUPINUS LYALLII B. J. Cox! Systematics of the genus Lupinus is often viewed as one of the most perplexing studies ever attempted. This is attested by the diverse taxonomic treatments of various floras and special publications. While several students of the lupines have recognized many entities, others have found these plants in such confusing mosaics that enormous synonymic reductions were made; neither approach has solved the taxonomic problems formed by lupine diversity. In a biosystematic revision of the Lupinus lepidus-L. caespitosus complex of the Sierra Nevada, Cascade and Rocky Mountains, various techniques of study were em- ployed in an effort to show the degree of relationship be- tween the taxa (Cox 1972a). Over 75 epithets have been applied to various members of this complex since 1828 when L. lepidus was described, and these epithets have been considered by some as referable to a single species. In order to present the concept of the L. lepidus-L. caes- pitosus complex in a digestible and economically feasible form, a taxonomic treatment of only Lupinus lyallii, its five subspecies and six varieties will be offered here. More complete distribution maps, cytological and chemotaxonomic data, photographs of typical specimens, and illustrations of typical leaflet and floral parts are given elsewhere (Cox 1972a, 1972b, 1973a, 1973b). Chromosome numbers: Taxa within Lupinus lyallii have chromosome number reports of 2n — 48 (subsp. lyallii, var. lyallii and var. villosus, and subsp. minutifolius) by Cox (1972b). The chromosomes are extremely small and counts are difficult to obtain, undoubtedly explaining the lack of previous reports. Although Phillips (1957) reported a 'Studies conducted at the Division of Biological Sciences, University of Missouri, Columbia, Missouri. 422 1974] Genus Lupinus — Cox 423 chromosome count for L. lyallii of n = 24, his study plant was L. minimus. Chromosome counts have offered little in determining the genetic affinities within the L. lepidus-L. caespitosus complex since all reported numbers are 2n — 48 (Cox 1972b). Chemosystematic data. Seed proteins: Comparisons were made of seed proteins of four of the L. lyallii taxa by poly- acrylamide gel electrophoresis (Cox 1972a, 1973b). Con- siderable variation was found in several polymorphic popu- lations; largely, the electrophoresis disc protein data closely resemble those data from the morphological study in the degree of variation found. Although no protein band oc- curred in common with all taxa studied, likewise no pro- tein band was found unique to any one taxon. Each popu- lation was found to have a distinct combination of proteins, resulting in low percent similarity values between the taxa. Seed Alkaloids: The seed alkaloids of all of the sub- species and varieties of Lupinus lyallii were compared by thin-layer chromatography (TLC) separation by Cox (1973a). Although some interpopulational variation was found, the alkaloids were generally quite consistent. Corre- lations were made between alkaloid shapes, R: values, posi- tive color reactions with two different visualizing agents, and comparisons with standards. Percent similarity values were high among the interspecific taxa, and identical alka- loid combinations were shared among a few, these differing only in relative amounts of individual alkaloids. Morphological and. distributional considerations: Field and herbarium-laboratory studies were conducted over a three year period on the Lupinus lepidus-L. caespitosus complex. Techniques and preparations of specimens for measurements in the descriptions are given by Cox (1972a). The following herbaria are acknowledged for the use of over 1000 specimens of the taxa of Lupinus lyallii: AEC, ARIZ, BRY, CAN, DS, GH, ISC, JEPS, MIN, MO, NMC, NY, POM, ORE, OSC, RENO, RM, RSA, SMU, UBC, UC, UMO, US, UTC, V, WILLU, WS. 424 Rhodora [Vol. 76 Lupinus lyallii A, Gray Plants perennial, herbaceous, caespitose or mat-like, less than 18 cm. tall; leaflets few (5-9), oblanceolate, the largest less than 2 cm. long; banners glabrous, usually re- flexed below the midpoint. Distribution: The taxa are primarily found in the high Sierra Nevada and Cascade Mountains with a few popula- tions radiating out into isolated localities (Fig. 1). KEY TO THE SUBSPECIES AND VARIETIES OF LUPINUS LYALLII A. Plants with spreading villous-hirsute to shaggy pubes- CENCE. aooaa lee. B. B. Plants lees than 10 cm. tall; flowers 6-7 mm. long; largest leaflets 4-6 mm. long; Sierra Nevada, alpine, dwarf. ...... 1b. L. lyallii subsp. lyallii var. danaus. B. Plants usually taller; flowers over 7 mm. long; larg- est leaflets over 6 mm. long. ............ . C. C. Plants decumbent, mat-like; leaflets 6- T, largest 10-13 mm. long; Sierra Nevada, with a few sta- tions in Klamath Co., Ore. . lf. L. lyallii subsp. lyallii var. villosus. C. Plants erect to semi-decumbent, branching, 10-18 cm. tall; leaflets 5-6, largest 9-21 mm. long; Washoe Co., Nevada. ...... , ZEE . 9. L. lyallii subsp. washoensis. A. Plants with appressed to subappressed pubescence. .. D. D. Plants erect in densely branching clumps; 12-18 cm. tall; Rogue River-Crater Lake region, Oregon. .. E. E. Branching stems mostly woody, less than 1.5 mm. in diameter; leaflets 5-6; racemes 1.3-1.4 cm. in diameter ; flowers 6-7.5 mm. long. . 1c. L. lyallii subsp. lyallii var. fructiculosus. E. Branching stems mostly herbaceous, more robust; leaflets 7-9; racemes 1.9-2.5 cm. in diameter; flowers 7-9.5 mm. long. ................. s. e le. L. lyallii subsp. lyallii var. roguensis. D. Plants decumbent or erect and densely caespitose. 1974] Genus Lupinus — Cox 425 F. Plants mat-like, usually less than 10 cm. tall; flowers 6-7 mm. long; largest leaflets 4-6 mm. long; alpine, Sierra Nevada Range. .........- Mero Rd 1b. L. lyallii subsp. lyallii var. danaus. F. Plants usually larger or flowers longer. ..... G. G. Plants 4-6 cm. tall; flowers 7.5-10 mm. long; largest leaflets 6-9 mm. long, upper surface glabrous; Steens Mts, Harney Co., Oregon. TM IO ay e «= 3. L. lyallii subsp. minutifolius. G. Plants mostly taller; flowers various; largest leaflets various lengths, mostly pubescent (becoming somewhat glabrate in subsp. alcis- temporis). occ ccc ecw cece gien p hh n H. H. Plants mostly decumbent; Clearwater Co., Idaho. .. 2. L. lyallii subsp. alcis-temporis. H. Plants erect or mat-like, not decumbent. I Leaves all basal, erect, in small tufts; petioles 3-9 em. long; largest leaflets 9-22 mm. long; SW Idaho & NE Ne- vada. .. 4. L. lyallii subsp. subpandens. I. Leaves cauline, basal, or both, generally ascending or more or less spreading, general not in small tufts; petioles usually 2-5.5 em. long; largest leaflets CEPEOGOEG o ni PTT J. J. Flowers 10-12.5 mm. long; racemes 2.5-4.5 (5.5) cm. long, 2.5-3.0 cm. in diameter ; leaflets 6-9; high alpine areas of western Cascade Range, Öre & Wash UU b e d nih s 1d. L. lyallii subsp. lyallii var. macroflorus. J. Flowers 7.5-10.5 mm. long; racemes 1.5-3 em. long, 2.0-2.3 cm. in di- ameter; leaflets 5-7; alpine and sub- alpine, wide distribution, Sierra Ne- vada Range, Calif. & Cascade Range, 426 Rhodora [Vol. 76 Ore., Wash. & B. C. .......... la. L. lyallii subsp. lyallii var. lyallii. la. Lupinus lyallii subsp. lyallii var. lyallii Lupinus lyallii A. Gray, Proc. Amer. Acad. 7: 334. 1868. Type: “Summit of the Cascade Mountains, lat 49°, elev. 7500 ft.”, Lyall, Jul 1860 (Holotype: K; Isotypes: BM, GH, POM). Photos: (RSA, UC, UMO, US). L. lepidus subsp. lyallii (Gray) Detl., Amer. Mid. Nat. 45: 490-491. 1951. L. alpinus Heller, Muhlenbergia 6: 22. 1910. Type: NEVADA: WASHOE CO.: Mt Rose, Heller 9914a (Holo- type: Nev. Agr.; Isotypes: GH, ISC, MIN, NMC, NY, POM, RSA, UC, US, WS). Photos: (RSA, UMO). L. paulimus Greene, Leafl. Bot. Obs. & Crit. 2: 234. 1912. Type: OREGON: DESCHUTES CO.: Paulina Lake, W end, Leiberg 550 (Holotype: US; Isotypes: GH, UC). L. rubro-soli Eastw., Leafl. West. Bot. 3: 203. 1941. Type: CALIFORNIA: MENDOCINO CO.: Red Mt., Eastwood & Howell 4674 (Holotype: CAS; Isotype: US). Photos: (RSA, UMO). Plants perennial, caespitose, forming mat-like radicle clumps, 15-25 cm. in diameter, from a thick, woody caudex which often branches in older plants, 6-17 cm. tall; foliage 2-7 cm. tall; stems poorly developed, less than 1.5 mm. in diameter, semi-prostrate, some sprawling-decumbent, some erect, subappressed silky sericeous throughout, hairs usually less than 1 mm. long; cauline nodes 0-2; petioles slender, 2-5(7.5) cm. long; stipules generally inconspicuous; leaf- lets 5-7, oblanceolate-linear to elliptic-obovate, apex acute to short mucronate, largest 5-12(18) mm. long, av. 9.95 mm., 1.5-4.5 mm. wide, length to width ratio 2.17-4.0(6.0), av. 3.18, appressed silky-sericeous on both surfaces; peduncles usually much exceeding the foliage, 3-8(10) cm. long; racemes capitate to subcapitate, 1.5-3 em. long, occasionally to 4.0 cm. at anthesis, 20-23 mm. in diameter, rarely 2X as long as wide; flowers dense, blue, 7.5-10.5 mm. long, av. 9.44; bracts persistent to caducous, 3-6.5 mm. long, subu- 1974] Genus Lupinus — Cox 427 late to lance-linear; pedicels 1.5-2.5 mm. long; calyx bi- labiate, the upper-lip 3.5-6.0 mm. long, bilobed, the notch 1-2.5 mm. deep, the lower-lip 3.8-6.8 mm. long, apex serrate to tridentate, the center tooth to 0.5 mm. long, lips connate 1-2 mm., bracteoles usually present but inconspicuous, to 0.5 mm. long, attached at the sinus, sericeous; banner ellip- tic-oblong to suborbicular, glabrous, 7-10 mm. long, 5.1-8.0 mm. wide, length to width ratio 1.10-1.57, av. 1.29, reflexed below or near the midpoint, the reflexed to appressed ratio 1-1.3, av. 1.14, angle 120-150°; wings 6.5-9.8 mm. long, 3.5- 5.5 mm. wide, length to width ratio 1.56-2.25 av. 1.90, claws 1.0-2.2 mm. long, glabrous; keel 2.5-3.0 mm. wide at the widest point, angle 105-111°, ciliated along the upper mar- gin near the acumen; ovules 3-5; pods 10-15 mm. long, 4-5 mm. wide, short sericeous; seeds 2-4, 1.8-3.0 mm. X 2.6-3.5 mm., 1-1.5 mm. thick, tan, beige, brown and green mottled. This variety is primarily alpine to subalpine, ranging latitudinally in its distribution from the high Sierra Nevada Mountains of California through the Cascade Range of Oregon and Washington into British Columbia. These plants may be found in several soil types, such as volcanic ash, disintegrated granite, sandy, gravelly rock of several compositions to shale. They usually occur on slopes associ- ated most frequently with Pinus contorta, P. murrayana, P. albicaulis and Tsuga mertensiana. Altitudinally, the plants are found above timberline at 3660 m. down to about 2439 m., or in the more northerly latitudes to about 1830 m. Flowering occurs between June and September; the plants flowering during the latter part of the season usually are found in sites of lingering snowbanks. Representative specimens: CANADA: British Columbia: Bomford Mt, Ashnola Dist, Barr 9351 (UBC); Chileotin, Copley 472 (v); Lillooet, Anderson, Jul 1916 (v); Manning Park, Hardy, Aug 1945 (v); Mt McLean, Anderson 7987 (v); Mt Moor, Bird 3436 (UBC) ; Nicoamen River head, Dawson 51833 (CAN); Noaxe Lake, Brink, Aug 1957 (UBC); Yalakom River, Sugden 118 (v). USA: California: ALPINE CO.: Carson Pass, Yates 3980 (UC). EL DORADO CO.: Desolation Valley, Abrams 12740 (GH, NY, POM, UC). FRESNO CO.: Humphreys 428 Rhodora [Vol. 76 Basin, headwaters Piute Creek, Quibell 5118 (UC). LASSEN CO.: Lassen Volcanic Natl. Park, Rickett, Aug 1968 (UMO). MADERA CO.: Mt Lyell, at Moraine Meadows, Akey 318 (UC). MARIPOSA CO.: Mt Connes, Tioga Pass, Ramsey & Ramsey 2496 (POM). MODOC CO.: Antelope Valley, Applegate 847 (US). MONO CO.: Bodie, 2.2 mi N-NW, Graham 274 (UC). NEVADA CO.: Mt Stanford, Sonne 54 (MO). PLACER CO.: Snow Mt, Bolt 103 (UC). SHASTA CO.: Bolam Road, near the S.P. RR, Cooke 15003 (uc). SISKIYOU CO.: Bolam Creek Canyon, opposite Coquette Falls, Mt Shasta. Cooke 25691 (UC). TEHAMA CO0.: King Creek Meadow, Lassen Volcanic Natl. Park, Heller 15426 (Mo, NY). TRINITY CO.: Fisher Ridge, E of Black Rock, North Yolla Bolly Mts, Munz 16743 (RSA). TULARE CO.: Golden Trout Creek, Hall & Hall 8417 (uc). TUOLUMNE CO.: Bond Pass, Winkelman 276 (Uc). Nevada: WASHOE CO.: Mt Rose, Ken- nedy 1148 (NY, UC, US). Oregon: CURRY CO.: Bear Camp Ranger Station, Leach 1692 (ORE). DESCHUTES CO.: Bend, Farnham & Beach 97 (OSC). DOUGLAS CO.: Bailey Mt, Applegate 4121 (uc). JACKSON co.: Ashland Peak. Thompson 12383 (MO, NY, UC). JFEFFFPSON CO.: Hoodoo, Big Lake Turnoff, US Hwy 20, Isely & Isely 11299 (ISC, UMO). JOSEPHINE CO.: Free and Easy Trail, head of Babyfoot, Leach 2661 (ORE). KLAMATH CO.: btn Fort Klamath & Summit, Furlong et al, 1901 (UC). CRATER LAKE NATL. PARK: Summit Lake, Henderson 9634 (ws). LAKE CO.: Drews Valley. Austin 1649 (Us). LANE CO.: McKinzie Pass, Peck 5467 (WILLU). LINN CO.: Bie Lake, Hawley & Anderson, Jul 1927 (osc). WALLOWA CO.: Wallowa Mts, Aneroid, 2 mi S, Peck 17985 (uc). Washington: CHELAN CO.: Blewett Pass region, Grant, Aug 1929 (UC). CLALLAM co: Hurricane Ridge, Thompson 14226 (GH, NY, US). JEFFERSON CO.: Tron Mt, Thompson 11049 (MO. NY, POM. US). OKANOGAN CO.: Bunker Hill summit, Edwards 32 (ws). PIERCE co.: Mt Rainier Natl. Park: Flett’s Glacier, Flett 3124 (ws). wHATCOM co.: Mt Baker. Heliotrope Ridge, Muenscher 8201 (GH, UC. US. WS). YAKIMA CO.: Hell Rearing Canyon rim, Langdon 116 (osc). The populations from the Wallowa Mountains (Wallowa Co., Oregon) are isolated from other Lupinus lyallii subsp. lyallii populations by 322 km.; the subsp. alcis-temporis occupies areas along the Clearwater and Selway Rivers in Idaho, but it is quite distinctive. The growth habit of the Wallowa populations appears to be typically that of var. lyallii specimens in the normal distribution, with short vegetation to 7 cm. tall, capitate inflorescences 2-2.5 cm. long, small flowers 6-8 mm. long and short leaves 7-10 mm. long. But, these plants hold in common with L. minimus a 1974] Genus Lupinus — Cox 429 strong woody branching caudex at above ground level, many stems in a clump with leaves appearing basal, not more than 2-3 cauline leaves, and silky pubescence. The specimens of Lupinus lyallii subsp. lyallii var. lyallii in Deschutes Co., Oregon (Leiberg 550 — GH, UC, US), de- scribed as L. paulinus by Greene, are somewhat larger than other L. lyallii plants of the immediate region but closely resemble several populations found in California. The racemes of some plants range from 3-4 cm. to 7.5 cm. long and from 16 mm. to 22 mm, in diameter, with the leaflets 11-15 mm. long, the upper surface of the leaflets sparsely pubescent, flowers approaching 9 mm. long and peduncles 6-7 cm. long. Several plants resemble L. sellulus var. lobbii with the flowers scattered in verticils somewhat like L. sellulus var. medius (Cox 1973c). However, the population sample indicates that these variants are more closely related to L. lyallii subsp. lyallii var. lyallii than to any other taxon and not distinctive enough to warrant a sepa- rate epithet. The material found on Mt. Rose in Nevada, which Heller called Lupinus alpinus, flowers from late July to late Sep- tember and occurs at elevations of 2927-3110 m. The plant habit is low and spreading mats, much like that on Mt. Rainier in Washington. Colonies occur in grassy open meadows, in the alpine belt. Although the plants may be somewhat atypical of the California L. lyallii subsp. lyallii var. lyallii specimens, they are much like those of Oregon and Washington. 1b. Lupinus lyallii subsp. lyallii var. danaus (A. Gray) S. Wats., Proc. Amer. Acad. 8: 534. 1873. L. danaus Gray, Proc. Amer. Acad. Arts & Sci. 7: 335. 1868. Type: CALIFORNIA: TUOLUMNE CO.: Mt Dana, Bolander 5087 (Holotype: GH; Isotype: UC, US). L. danaus var. bicolor Eastw., Leafl. West. Bot. 2: 201. 1939. Holotype: CALIFORNIA: TUOLUMNE C0.: Mt Dana, Howell 14539 (CAS). Photos: (RSA, UMO). L. oreocharis Eastw., Leafl. West. Bot. 3: 171. 1942. 430 Rhodora [Vol. 76 Holotype: CALIFORNIA: TULARE CO.: Little Five Lakes Basin, Howell 17392 (CAS). Photos: (RSA, UMO). Plants differing from L. lyallit subsp. lyallii var. lyallii by matted vegetative growth, 1-4 cm. tall; plant overall height usually less than 10 cm. tall; pubescence frequently sparse, spreading; largest leaflets mostly 4-6 mm. long, rarely to 10 mm. long; peduncles 1.5-3 em. long, rarely to 8 em. long; racemes capitate, 1.5-2.0 cm. long; flowers fre- quently more loosely arranged than in var. lyallii, smaller, 6-7 mm. long. Representative specimens: California: FRESNO C0.: Humphreys Basin, headwaters Piute Creek, W of Mt Humphreys, Sharsmith 3136 (uc). INYO co.: Piute Pass, E slope, Peirson 2552 (RSA). MADERA co.: Isberg Pk, 1 mi SW, Akey 332 (UC). MARIPOSA CO.: Mt Dana, Congdon, Aug 1898 (Uc). MODOC CO.: Mt Warren, Payne 307 (JEPS). MONO C0.: San Juaquin Peak, Wright K3 (POM). TULARE co.: Mt Whitney, Clemens, Jul 1910 (POM, UMO). TUOLUMNE CO.: Gaylor Lakes, Howell 20401 (ws). The depauperate forms of L. lyallii subsp. lyallii are interpreted here as being ecological variants. Under a par- ticular set of environmental conditions populations of L. lyallii dwarfs to the matted form of var. danaus. However, frequently within the same population typical var. lyallii may occur in the larger form. Both varieties are found in high alpine rocky, sunny areas. Although this depau- perate or ecological variant of L. lyallii appears to be morphologically blending with the larger variety, it is considered here as a distinct taxon at the varietal level. A variant of similar vegetative morphology was observed among the Lupinus lyallii specimens of Canada and the higher alpine areas of Washington and Oregon, but the reproductive structures differ from that of var. danaus in California. Lupinus oreocharis is reduced to synonymy under var. danaus. Although some of these plants range to 14 cm. tall and resemble var. villosus in pubescence, and some may differ from var. danaus by having slightly elongated stems 1974] Genus Lupinus — Cox 431 which may have 1-2 cauline leaves, the available evidence does not indicate the warrant of a special epithet. 1c. Lupinus lyallii subsp. lyallii var. fruticulosus (Greene) C. P. Sm., Bull. Torrey Bot. Club 51: 303. 1924. L. fruticulosus Greene, Muhlenbergia 8: 117. 1912. Type: OREGON: KLAMATH CO.: Anna Creek Valley, Austin 1650 (Holotype: US; Isotypes: NY, POM). Photos: (RSA, UMO, US). L. perditorum Greene, Muhlenbergia 8: 117. 1912. Holotype: OREGON: JACKSON CO.: Upper Rogue River, Applegate 2636 (US). Photo: (US). Plants in dense semi-shrubby clumps, branching pro- fusely, erect to semi-decumbent; stems 19-17 em: tall, usually less than 1.5 mm. in diameter; leaves cauline; petioles slender, 2-5 cm. long; leaflets 5-6, elliptic-oblanceo- late, largest 5-12 mm. long, av. 8.4, 1.8-3.5 mm. wide, length to width ratio 2.78-4.0, av. 3.42, subappressed, strigose- pilose on both surfaces; peduncles 2.5-5.5 em. long; racemes numerous, 1.5-4.3 em. long, 13-14 mm. in diameter, capitate and dense to somewhat scattered; flowers blue, 6.0-7.5 (8.0) mm. long, av. 6.9 mm.; bracts 1.5-3.0 mm. long; pedicels 0.5-1.5 mm. long; banner elliptic to suborbicular, reflexed below the midpoint; pods 7.5-10 mm. long, 3.3-4 mm. wide; seeds 2-3. The greatest development of var. fruticulosus is along the flood plain of the Rogue River in Klamath Co., Oregon. Plants are found at elevations of 1067-1372 m., flowering in July. Representative specimens: Oregon: JACKSON CO.: Ashland, 9.7 km S, Sherwood 842 (WILLU). KLAMATH CO.: Crater Lake Natl. Park, Evinger 608 (osc). LINN CO.: Hunts Cove, Mt Jefferson area, Leach 4485 (ORE). Variety fruticulosus differs morphologically from other variants of Lupinus lyallii by its semi-shrubby base, pro- fusely branching clump habit and smaller flowers. A few L. lyallii specimens from Washington (mainly var. macro- 432 Rhodora [Vol. 76 florus) have thick, woody branching bases, but these are part of the underground caudex system and differ by having distinctively larger inflorescence structures. The Washing- ton material is more robust than fruticulosus, Variety fruticulosus is one of the most distinctive variations of L. lyallii subsp. lyallii. Variety fruticulosus is morphologically similar to var. roguensis, of the same general geographical region, and to subsp. alcis-temporis of the Clearwater and Selway Rivers in Idaho. It may be separated from these two taxa on the basis of its flower morphology, more scattered flower arrangement on the racemes and smaller size, as well as differences in vegetative traits. These plants exhibit a more erect woody branching habit with more filiform petioles and stems than any closely related entities. They are generally not as leafy, and the leaflets are fewer (5-6 per leaf), smaller, and arranged in the upper portion of the branching clump. 1d. Lupinus lyallii subsp. lyallii var. macroflorus B. J. Cox, var. nov. Type: WASHINGTON: PIERCE CO.: Mt Rainier, Yakima Park, Thompson 11086 (Holotype: MO; Isotypes: NY, POM-in part, US). Photo of holotype: (Cox 1972 a). Plantae perennes, humiles; caules ex caudice profuse ramificanti lignoso, 5-17 cm. alti, 1-15 mm. diametro, nodis 0-2(8); petioli 3-5.5 em. longi; foliola 6-9, oblanceolata, maxima 5.5-11.3 mm. longa, 2.5-5.0 mm. lata; pedunculi 3.0-5.5 em. longi, supra folia 1-2 em, extensi; racemi 2.5-5.5 cm. longi, 2.5-3.0 cm. diametro, capitati usque subcapitati, floribus 5-17 in 1-5 verticillis; flores caerulei 10-12.5 mm. longi; calyx gibbosus; vexillum suborbiculatum usque obo- vatum ; ovula 3-5; legumina 13-15 mm. longa, 4-5.5 mm. lata. Plants perennial, in dense low mats; stems from a pro- fusely branching woody base, 5-17 cm. tall, 1-1.5 mm. in diameter, pubescence appressed to ascending, some spread- ing, cauline nodes 0-2(8); petioles 3-5.5 em. long; leaflets 6-9, oblanceolate, largest 5.5-11.3 mm. long, av. 8.29 mm., 2.5-5.0 mm, wide, av. 2.56 mm. ; peduncles 3.0-5.5 cm. long, 1974] Genus Lupinus — Cox 433 exceeding the foliage by 1-2 cm. or more; racemes 2.5-5.5 em. long, 2.5-3.0 cm. in diameter, capitate to subcapitate, flowers 5-17 in 1-5 verticils; flowers blue, 10-12.5 mm. long, av. 11.2 mm. long; calyx gibbous at the base; banner sub- orbiculate to obovate, glabrous; keel densely ciliated near the acumen to glabrous in some; ovules 3-5; legumes 13-15 mm. long, 4-5.5 mm. wide. Populations of var. macroflorus are found on the western side of the Cascade Mountain Range on both sides of the Columbia River in Washington and Oregon. They are found on Mary’s Peak in Benton Co., on Mt. Hood of Clackamus and Hood River Counties, Santiam Pass in Linn Co., Ore- gon, and Mt. St. Hellens of Skamania Co., and Mt. Rainier of Pierce Co., Washington. Like var. lyallii, var. macro- florus occurs at the upper vegetational limit between tim- berline and snowline at elevations of 1524-2378 m. Plant communities commonly associated include the species: Pinus contorta, P. albicaulis, Abies lasiocarpa, Tsuga. mer- tensiana and less frequently Pseudotsuga menziesii. Representative specimens: Oregon: BENTON CO.: Corvallis, W at Mary's Peak, Gilbert 108 (osc, US). CLACKAMUS CO.: Mt Hood, Jct of Hwys 26 & 35, Downey 4B (UMO, UTC). HOOD RIVER CO.: Mt Hood, Abrams 9445 (MO, POM, RM, WILLU). LINN CO.: Santiam Pass, Cronquist 7853 (GH, NY, RSA, UC, WS). Washington: PIERCE CO.: Mt Rainier Natl. Park, Burrough’s Mt & Frozen Lake, Flett 3121 (WS). SKAMANIA CO.: Mt St Hellens, Buzzettii, Aug 1848 (osc). Variety macroflorus is distinct primarily on the basis of its larger flower size (10-12.5 mm. long) from var. lyallii flower size (7.5-10.5 mm. long). There are several inter- mediates between the variations of L. lyallii, but in general var. macroflorus is the extreme in flower morphological deviation. le. Lupinus lyallii subsp. lyallii var, roguensis B. J. Cox, var. nov. Type: OREGON: JACKSON co.: Farewell Bend Forest Camp, 1.6 km N of Union Cr, on Rogue River flood plain, Hitchcock & Martin 4997 (Holotype: RSA; Isotypes: MO, NY, UC, WS). Photo of isotype: (Cox 1972a). 434 Rhodora [Vol. 76 Plantae perennes, herbaceae, caespites densos ramificantes formantes, 2.5-3.5 dm. diametro, 12-18 cm. altae; caules humifusi usque erecti, 1-3 mm. diametro, ex caudice dense ramificanti lignoso, nodis 6-10, ad nodos ramificantes ; foliola 7-9, maxima 8-13 mm. longa, 2.5-3.0 mm. lata; racemi capi- tati usque subcapitati 1.5-3.5 mm. longi, 4.5 mm. longi fructiferi, 1.9-2.5 mm. diametro; flores 7-9.5 mm. longi; vexillum et alae glabrae. Plants perennial, herbaceous, in dense branching clumps, 2.5-3.5 dm. in diameter, 12-18 cm. tall; stems procumbent to erect, 1-3 mm. in diameter, from densely branching woody base, nodes 6-10, branching at the nodes; leaflets 7-9, largest 8-13 mm. long, 2.5-3.0 mm. wide; racemes capi- tate to subeapitate, 1.5-3.5 mm. long, 4.5 mm. long in fruit, 1.9-2.5 mm. in diameter; flowers 7-9.5 mm. long; banner and wings glabrous; keel ciliated along the upper margin near the acumen. Variety roguensis is found at Diamond Lake, Huckleberry Mt., Davis Lake, and on the flood plain of the Rogue River, in Jackson and Klamath Counties, Oregon, These plants occur in sandy soil in pine forests at elevations of 915-1067 m. Flowering is from mid-June to mid-July. Representative specimens: Oregon: JACKSON CO.: Huckleberry Mt, Rogue River Natl. Forest, Thompson 130835 (MO, RSA, UC, WILLU, WS). KLAMATH CO.: Davis Lake, W shore, Hitchcock & Martin 4935 (RSA). Morphologically, these plants are most closely related to var. fructiculosus, but they differ in several significant characteristics. They are more robust, have larger branch- ing procumbent to erect stems, have larger flowers that are more densely arranged in racemes, and have more leafy foliage. The leaves are more numerous and are scat- tered throughout the clump, and the leaflets are larger with more leaflets per leaf. The manner of branching re- sembles that of L. aridus, especially subspecies lenorensis (Cox 1972a) found along the Clearwater and Selway Rivers in Idaho, Although the general branching pattern is often 1974] Genus Lupinus — Cox 435 difficult to detect, careful scrutiny finds verticillate spread- ing branching from an erect shoot; these branches become somewhat decumbent, but secondary and tertiary branches arise from them either adaxially or twisting around the decumbent stems and growing upward forming erect clumps. Several color variations are found in this segment of Lupinus lyallit. Many plants of var. roguensis have albino flowers (white to cream color — Maguire & Holmgren 26493 — GH, NY, UC, US) ; however, others within the same popu- lation are intermediate in flower color to the albino and typical blue form (white to cream colored petals except for the purple banner and keel tip — Maguire & Holmgren 26494 — GH, MO, NY, UC, US, WS). 1f. Lupinus lyallii subsp. lyallii var. villosus Jeps., Jepson, Fl. Calif. 2: 267. 1936. Holotype: CALIFORNIA: ALPINE CO.: Carson Pass, Jepson 8100 (JEPS). L. hypoleucus Greene, Leafl. Bot. Obs. & Crit. 1: 74. 1904. Type: CALIFORNIA: White Chief Peak, Culbert- son 16, Jul 1904 (distributed by Baker 4416 — specimen not located.) Paratype: CALIFORNIA: Mt, Goddard, Hall & Chandler 707 (US). Plants densely caespitose; stems decumbent to semi-erect, from a thick branching woody caudex, sericeous with as- cending to long spreading white hairs turning reddish- brown with age; petioles 4-5 cm. long; leaflets 6-7, elliptic- oblanceolate, apex acute-mucronate, largest 10-13 mm. long, 3-5 mm. wide, villous on both surfaces; peduncles 3-7 cm. long; racemes densely verticilate 2.5-5 cm. long; bracts persistent, 4-6 mm. long; pedicels 1-1.5 mm. long; banner and wings glabrous; keel ciliate near the acumen; pods 10-11 mm. long, 4 mm. wide. This variety occurs scattered throughout much of the distributional range of L. lyallii in several counties of California in the Sierra Nevada northward to several popu- lations in Klamath Co., Oregon. 436 Rhodora [Vol. 76 Representative specimens: California: ALPINE CO.: Carson Pass, Higgins 1752 (BRY). FRESNO CO.: Bubbs Creek, Peirson 531 (RSA). HUMBOLDT CO.: Grouse Mt, back of Clear Lake, Tracy 16672 (UC). LASSEN C0.: Grass Valley, Fisher & Johnson F203 (UC). MADERA co.: Garnet Lake, Howell 16421 (RSA, UC, US). MARIPOSA CO.: Donohue Pass, Schreiber 1767 (UC). MENDOCINO CO.: South Yolla Bolly, Jepson 57t (JEPS). MONO CO.: San Juaquin Mt, Howell 16875 (Ws). PLACER CO.: Mt Lincoln, Howell 18670 (US). SHASTA Yolla Bolly, Jepson 57t (JEPS). MONO C0.: San Juaquin Mt, Howell 16875 (ws). PLACIER CO.: Mt Lincoln, Howell 18670 (US). SHASTA co.: Squaw Valley Creek Meadows, Cooke 13651 (NY). TRINITY CO.: Trinity Center, 3 mi S, E of Scott Ranch, Cantelow, Aug 1936 (RSA). TUOLUMNE C0.: Gaylor Lakes, Howell 20418 (GH). Oregon: KLA- MATH CO.: Llao Rock, Baker 6245 (NY, OSC, UC, WS). Variety villosus is recognized primarily on the basis of its dense pubescence. In L. lyallii, pubescence ranges from puberulent-sericeous to woolly-villous approaching spread- ing hirsute. Several populations exhibit different forms of pubescence within a single colony; this is especially true of those specimens viewed from the Carson Pass vicinity (the type locality of var. villosus), Alpine Co., California. 2. Lupinus lyallii subsp. alcis-temporis (C. P. Sm.) B. J. Cox, comb. nov. Lupinus alcis-temporis C. P. Sm., Sp. Lup. 558. 1946. Type: IDAHO: CLEARWATER CO.: Pierce, 17.7 km. E, Crystal Creek Grade, Christ 14634 (Holotype: DS; Iso- types: NY, US). Photos: (RSA, UMO). Plants sprawling caespitose, branching profusely from a slender woody base, in clumps 11-30 cm. in diameter; stems subdecumbent, 8-15 em. long, strigose-sericeous with dense coarse ascending hairs, nodes 4-6; cauline petioles 2-4 em. long; stipules 7-9 mm. long, connate to the petioles 3-4.5 mm., filiform; leaflets 5-7, elliptic-oblanceolate, apex acute to short mucronate, longest leaflets 2-18 mm., av. 10 mm. long, 0.8-5 mm. wide, length to width ratio 2.33- 4.33, av. 3.18, strigose-pilose to villous on both surfaces, some more sparsely so on the upper surface to glabrous; peduncles 1.5-3.5 cm. long; racemes capitate, 2-3.5(6) cm. long, densely verticillate; flowers blue with yellow sulcus, 1974] Genus Lupinus — Cox 437 7-8.2 mm. long; bracts 5-9 mm. long, subulate-attenuate, sericeous to villous; pedicels 1.2-1.8 mm. long; calyx upper- lip 4.0-5.0 mm. long, bilobed, notch 2-3.5 mm. deep, lower- lip 5.0-6.0 mm. long, apex tridentate, center tooth 0.2-0.5 mm. long, lips connate 1.0-1.5 mm. ; bracteoles 0.8-1.5 mm. long, attached at the sinus, villous ; banner elliptic-oblanceo- late, 7.0-8.0 mm. long, 4-4.5 mm. wide, length to width 1.56- 1.88, av. 1.74, reflexed to appressed ratio 1.25-1.50, av. 1.37, angle 148-158°; wings 6.0-7.3 mm. long, 3.0-3.4 mm. wide, length to width ratio 1.97-2.85, av. 2.16; keel 2 mm. wide at the widest point, angle 111-122°, ciliated sparsely near the acumen ; ovules 2-3; pods 8.5-9.5 mm. long, 3.4-4.0 mm. wide, densely short woolly; seeds 2-3, 1.8-2.0 mm. X 2.0- 2.2 mm., 0.9-1.1 mm. thick, tan color. Subspecies alcis-temporis is found in sandy and gravelly slopes on road cuts, generally in otherwise barren areas. One population occurs in a Pinus monticola forest; the Christ & Smith collection (15385 — NY) was found on “forest soil bordering beds of Boykinia major; the Dauben- mire collection (47150 — WS) was made in a Spruce-Fir Zone. Plants oecur at 915-1220 m. elevation. Representative specimens: Idaho: CLEARWATER CO.: Pierce, 17.7 km E at the confluence of Big Breakfast Creek and Crystal Creek, Christ & Smith 15385 (NY); Washington and Lodge Creeks, S31, T39N, R7E, Lingenfelter 466 (NY, RSA, UC, WS). This taxon was first described by C. P. Smith from a Christ “little number picked up in Clearwater County (Idaho) during elk season in October.” Hence the name *Christ's Elk-Season Lupine”, Lupinus alcis-temporis in Smith’s Latinum utilizes the genitive for both elements of the compound name (alces — elk and tempus — season). Both Christ and Smith were deeply interested in the fact that a lupine in anthesis could be found in Clearwater Co. in the middle of October “without a trace of fruit or seed, and at only 4,000 feet elevation." Several stages of development are found in more ex- tensive study of other collections of these lupines, all of 438 Rhodora [Vol. 76 which have been collected on July 10th or July 11th. Small clusters of young herbaceous-stemmed, procumbent plants may be found flowering the same time as older plants with semi-woody branching stems, sprawling 30-45 cm. in di- ameter. The isotype material viewed from NY and US appears merely to be atypically late in reaching anthesis. It is in anthesis with some fruit in the initial stages of enlargement, and all branches are woody including second- ary and tertiary branches; only the leaves and inflorescences remain herbaceous at this late seasonal date. 3. Lupinus lyallii subsp. minutifolius (Eastw.) B. J. Cox, comb. & stat. nov. Lupinus minutifolius Eastw. Leafl. West. Bot. 2: 267. 1940. Holotype: OREGON: HARNEY CO0.: Fish Lake, Steens Mts., Henderson 8132 (CAS). Photos: (RSA, UMO). Plants acaulescent; foliage to 4.5 cm. tall; stems 4-6 cm. tall, appressed hispidulose with more coarse scaberose hairs; petioles filiform, 2.5-4 cm. long; leaflets 5-7, elliptical, glabrous above to sparse pubescence on the margins, short coarse appressed hairs below, largest 6-9 mm., av. 6.8 mm., 2.9.8 mm. wide, av. 2.45 mm., length to width ratio 2.4-3.0, av. 2.75; peduncles 6-8 cm. long or only 3 cm. long in im- mature plants; racemes 1.5-3.0 em. long, 3-4 compact verti- cils, ascending or erect from nearly prostrate stems; flowers 7.5-10 mm. long, mostly blue with nearly white banners and purple tipped keels; bracts caducous, 4.5-6 mm. long, subulate; pedicels 1.0-2.0 mm. long; calyx upper lip 4-5 mm. long, bilobed, notched 2.5-2.7 mm. deep, lower-lip 4.5-5.0 mm. long, apex entire-serrate to tridentate with the center tooth approaching 0.7 mm. long, lips connate 1.2-1.3 mm., bracteoles 0.2 mm. long, attached at the sinus, strigose to short ascending pilose; banner elliptic-oblong, 8-8.5 mm. long, 4.2-5.5 mm. wide, length to width ratio 1.45-1.90, av. 1.65, reflexed to appressed ratio 1.08-1.57, av. 1.34, angle 150-160°; wings 8.8-9.0 mm. long, 3.8-4.2 mm. wide, length to width ratio 2.10-2.57, av. 2.34, claws 1.0-1.8 mm. long, glabrous; keel 2.0-2.2 mm, wide at the widest point, angle 1974] Genus Lupinus — Cox 439 118-1235, ciliated along the outer half of the upper margin, densest toward the acumen; ovules 3; pods 9.0-10.5 mm. long, 3.5-4.2 mm. wide, short appressed; seeds 2-4. Subspecies minutifolius is found in the Steens Mountains of Harney Co. Oregon, geographically replacing subsp. lyallii. These plants are found on rocky, gravelly, gentle basaltic slopes but frequently occur in alpine meadows on hillside areas with grasses and sedges in sloping rocky habitats. Several specimens have been collected in beds of lingering snowbanks where snow has accumulated and remained during the winter and spring. Anthesis generally occurs during the latter part of July. Plants are found at elevations of 2439-2896 m. Representative specimens: Oregon: HARNEY CO.: Big Fish Creek Canyon, Maguire & Holmgren 26789 (GH, NY, UC, US); Frenchglen, 31.4 km due E-SE, Hansen 490 (osc). Although some populations of subsp. minutifolius re- semble subsp. lyallii var. danaus, they have several distinct morphological characteristics. The flowers of subsp. minu- tifolius generally have more oblong shaped banners, more linear wings, and therefore a greater length to width petal ratio than subsp. lyallii. Although the vegetative mor- phology of subsp. minutifolius resembles the alpine var. danaus and other northern forms of subsp. lyallii, it shows distinction in the filiform petioles from the basal clumps of less than 4.5 cm. tall, and frequently by the fewer (4-7, av. 5) leaflets that are glabrous on the upper surface. The plants of the Cronquist collection. (8694 — NY, OSC, RM, RSA, UC, WS) are morphologically larger than normal for the taxon. The foliage of these specimens approach 5 cm. tall, overall plant height approaches 9.5 cm.; flowers are an intermediate 8.5 mm. long ; the leaflets are larger 10 mm. x 8.5 mm.; peduncles are larger 6.5-7 mm. long, and racemes approach 4 em. long. 4. Lupinus lyallii subsp. subpandens C. P. Sm. ex Dunn, El Aliso 3: 171. 1955. Type: NEVADA: ELKO CO.: Elko, 440 Rhodora [Vol. 76 113 km. NW, Holmgren, Jun 1938 (Holotype: UTC; Iso- types: NA, NEV, AGR.). Photos: (RSA, UMO). Plants with many small tufts from a profusely branch- ing woody caudex, erect; foliage 4.5-9.0 cm. in height, mostly less than 6 cm. tall; stems 7-18 cm. tall, usually less than 1 mm. in diameter, appressed strigose-pilose pubes- cence less than 2 mm. long ; leaves in basal clusters ; petioles slender, 3-9 cm. long; stipules inconspicuous, 6-10 mm. long, connate to the petioles 4-5 mm, subulate; leaflets 5-6, linear- oblanceolate, apex acute, largest 9-22 mm. long, 2-5 mm. wide, length to width ratio 3.5-6.3, av. 4.8, sericeous-strigose below, more sparsely so above with glabrous apex and mid- rib; peduncle 5-9 cm. long, usually exceeding the foliage by 2-5 cm.; racemes capitate to subcapitate, 1.5-4 cm. long; flowers dense, blue, 9.5-11.5 mm. long; bracts 4-8.5 mm. long, appressed to ascending pilose; pedicels 1-2 mm. long; calyx upper-lip 4.2-5.5 mm, long, notch 1.3-3.2 mm. deep, lower-lip 4.5-5.5 mm. long, apex deeply serrate to tridentate, center tooth to 0.5 mm. long, lips connate 1.0-1.5 mm., bracteoles slender, inconspicuous, 0.5-1.0 mm, long, attached to the sinus, strigose-sericeous, hairs to 2 mm. long; banner elliptic, 8-9.5 mm. long, 5-6.5 mm. wide, length to width ratio 1.45-1.90, av. 1.56, reflexed to appressed ratio 1.13-1.71, av. 1.44, angle 146-150°; wings 8.2-9.8 mm. long, 3.4-4.8 mm. wide, length to width ratio 2.04-2.65, av. 2.36, claws 1.5-2.0 mm. long, glabrous; keel 2.2-3.0 mm. wide at the widest point, angle 108-120°, ciliated along the outer half of the upper margin, most densely near the acumen, cilia to 0.7 mm. long; ovules 3-5; pods 11-14 mm. long, 5-5.5 mm. wide, dense strigose to short villous; seeds 2-3, 2.0-2.2 mm. X 2.8-3.0 mm. 1.3 + 0.1 mm. in thickness, beige to tan with darker brown mottling. These plants are found in dry, rocky creek beds and open well-drained draws in northeastern Nevada, in Elko Co., and southwestern Idaho, in Camas and Owyhee Coun- ties. The most common associate is Artemisia sp. Plants flower from mid-July to the first part of August. Altitudi- 1974] Genus Lupinus — Cox 441 nally, they are found at 1677-1982 m., with one population found on Alleghany Creek at 2439 m. Representative specimens: Idaho: CAMAS CO.: Fairfield, 16 km. SE, Ripley & Barneby 10671 (NY). OWYHEE Co.: Mud Flat, 4.8 km. SW, on road to Juniper Mt, Maguire & Holmgren 26305 (GH, POM, UC, US, UTC, WS). Nevada: ELKO Co.: O’Neil, 16 km. S, Holmgren, Cooper & Cain 181 (GH, osc, UTC) ; Wells, 48 km. N, Jones, Jul 1901 (POM). Dunn (1955) interpreted Lupinus lyallii subsp. sub- pandens as an acquisition of a combination of the char- acters of L. aridus with those of L. lyallii. The pubescence type and system of branching from the caudex, as well as certain flower morphology, is characteristic of L. aridus, whereas the small caespitose habit with elongated peduncles and capitate inflorescences with small flowers is more in- dicative of L. lyallii. Dunn further suggests that the plant’s occupancy of the flatland Artemisia habitat of 1677-1982 m. elevation indicates a considerable amount of the L. aridus physiological adaptations, contrasting the higher altitudinal L. lyallii habitat. This study concurs with Dunn's interpretation of the affinity of subsp. subpandens with L. lyallii. These plants inhabit a geographically distinct region, also having both morphological and physiological distinction. Although each subsp. subpandens population has aequired a unique com- bination of characteristics, the plants can usually be recog- nized by the erect habit of both flowering culms and petioles as well as by the glabrous apex and midrib of the upper leaflet surfaces. The specimens from Camas Co., Idaho (Ripley & Barneby 10671 — NY) have larger flowers than the taxon mean, with banners only slightly reflexed and frequently with one node on the flowering culm above 2 cm. from the basal cluster of leaves, and racemes of 3-5 cm. long, being about 2 cm. in diameter. The specimens collected south of O'Neil, Nevada (Holm- gren, Cooper & Cain 181 — GH, OSC, UTC) are more slight in habit, having shorter petioles with smaller leaflets that 442 Rhodora [Vol. 76 are more glabrous on the upper surface, and shorter, more capitate racemes with the banners less reflexed. The Mud Flat, Owyhee Co., Idaho specimens (Maguire & Holmgren 26305 — GH, POM, UC, US, UTC, WS) have fewer leaflets than other populations, and the petioles are more elongated and slender. 5. Lupinus lyallii subsp, washoensis (Heller) B. J. Cox, comb & stat. nov. L. washoensis Heller, Muhlenbergia 6: 72. 1910. based on L. pinetorum Heller, Muhlenbergia 6: 25. 1910, not L. pinetorum Jones, Contr. West. Bot. 8: 25. 1898. Type: NEVADA: WASHOE CO.: Ridge S of Alum Creek, Heller 9748 (Holotype: Nev. Agr.; Isotypes: DS, GH, ISC, MO, NY, ORE, POM, UC, US, WS). Photos: (RSA, UMO). L. aridus var. washoensis (Heller) C. P. Sm,, Bull. Torrey Bot. Club 51: 3083. 1924. Plants 10-18 cm. tall, in dense caespitose clumps, 15-30 cm. in diameter, from woody vertical caudex branching at ground level, with 10-18 racemes flowering simultaneously; stems procumbent to erect, 1-2.5 mm. in diameter, villous to hirsute-shaggy white pubescence 2-4 mm. long; 2-3 nodes; cauline petioles 4.5-8 cm. long, pubescence as on the stems; basal petioles on immature plants 4.5-7 cm. long; stipules 6-10 mm. long, connate to petioles 4-6 mm., densely white bearded pubescence; leaflets 5-6, oblanceolate to ellip- tic-oblong, apex short mucronate, largest 9-21 mm. long, av. 13.7 mm., 3-7 mm. wide, length to width ration 1.86- 3.27, av. 2.71, silvery appressed to ascending pubescence below, more sparsely so above, upper surface appearing more greenish; peduncles 3-5 cm. long, generally exceeding the leaves; racemes 3-6 cm. long, 1.6-1.9 cm. in diameter, densely subverticillate; flowers blue to violet, banner with a white sulcus, 8-9 mm. long; bracts 3-5.5 mm. long, subu- late-attenuate, pilose-sericeous; pedicels 1.5-2.0 mm, long; calyx upper-lip 3.5-4.5 mm. long, bilobed, the notch 0.8-1.2 mm. deep, lower-lip 4-5.5 mm, long, apex serrate, lips con- nate 1-2 mm., bracteoles 0.5-1.0 mm. long, attached at the 1974] Genus Lupinus — Cox 443 sinus, villous; banner elliptic-oblong, 7.8-8.0 mm. long, 5.0- 6.0 mm. wide, length to width ratio 1.33-1.67, av. 1.52, reflexed to appressed ratio 1.25-1.35, angle 140-150? ; wings 7.5-8.0 mm. long, 4-4.5 mm. wide, length to width ratio 1.74-2.00, av. 1.87, claws 1.5-2.0, glabrous; keel 2.5-3.0 mm. wide at the widest point, angle 118-125°, ciliated along the upper margin near the acumen; ovules 2-3; pods 9-13 mm. long, 5-6 mm. wide, villous to ascending woolly; seeds 1-3, 1.8-2.2 mm. X 2.4-2.7 mm., 0.9-1.2 mm. thick, tan with brown mottling. Subspecies washoensis has a restricted habitat on the eastern slopes of the Sierra Nevada in the southern part of Washoe Co., SW of Reno, Nevada. Plants occur on gravelly montane open pine slopes at elevations of 1524- 2134 m., flowering in June. Representative specimens: Nevada: WASHOE CO.: Dinsmore Camp, Hunter Creek Canyon, Kennedy 1614 (Mo, Ny, UC); Reno, Hillman (POM); 10-13 km. SW, Hunter’s Creek Road, Archer 6090 (NY). The general appearance of some of these plants indicates relationship with Lupinus aridus, a portion of the racemes being exceeded in height by the foliage. Affinity is also indicated with L. sellulus var. lobbii in general morpho- logical structure (Cox 1972a, 1973c). Considering the total conformation — narrow, short subcapitate or longer racemes, and the procumbent, branching, sprawling vege- tative habit — this study concludes that the washoensis affinity lies most closely with L. lyallii. These plants re- place L. lyallii subsp. lyallii geographically and ecologically, aso being recognizably distinct by somewhat larger stature, having longer racemes, larger leaves, more yellow-green color and spreading silvery stem and petiole pubescence. The Dinsmore population (Kennedy 1614 — MO, NY, UC), which is only a few miles north of the type locality in Hunter Creek Canyon, has been used in sheep feeding ex- periment (Nev. Agr. Exp. Sta. Bull. 71: 30. 1909). The plants are said to be relished by the sheep, proving to be a good forage plant. Rhodora [Vol. 76 ` oo N » 0O Q BRITISH COLUMBIA N ———— \ o% E a |! WASHINGTON i Y N |o M LUPINUS LYALLII is we RN i ( * & i / 1 , \ OREGON C Lees / IDAHO | ^ | PO q—. de. | : Ap ——— | AA! ^| NEVADA ' b | subspecies: variety: A | odes. | o-lyallii ——— lyallii 33 N j +- lyallii ————— fruticulosus 3 x EAT X | *- | yallii villosus (] Et i i e - | yallii roguensis oo N M go NOM x—lyallii danaus ii .. CALIFORNIA N v - lyallii macroflorus A - washoensis a- minutifolius x - alcis- temporis A-subpandens Figure 1. Distribution of Lupinus lyallii. 1974] Genus Lupinus — Cox 445 LITERATURE CITED Cox, B. J. 1972a. Biosystematies of Lupinus lepidus-L. caespitosus complex. Ph.D. dissertation. Univ. of Missouri, Columbia. 343 p. 1972b. IOPB Chromosome number reports XXXVIII. Taxon 21: 680-681. 1973a. A chemosystematic comparison of the Lupinus lepidus-L. caespitosus complex. Bull. Torrey Bot. Club 100: 12-17. 1973b. Protein relationships among the perennial cae- spitose lupines. Bull. Torrey Bot. Club (in press). 1973c. Taxonomy of Lupinus sellulus (Papilionaceae). Amer. Mid. Natur. (in press). Dunn, D. B. 1955. A new subspecies of Lupinus lyallii. El Aliso $2 2171-172. PHILLIPS, L. L. 1957. Chromosome numbers in Lupinus. Madroño 14: 30-36. INDUSTRIAL BIO-TEST LABORATORIES NORTHBROOK, ILLINOIS 60062 THE ROLE OF SOME HALORAGACEAE IN ALGAL ECOLOGY L. C. COLT, JR., AND C. BARRE HELLQUIST While collecting vascular aquatie plants in several ponds in east-central Massachusetts we found that there was a considerable accumulation of algae upon some of the vascu- lar species, but very little on others. The present paper is the result of our futher observations on and examination of the relationship between the algae and the vascular plants upon whose surfaces they were found. There is little published data elaborating the substrate preferences of fresh-water algae. Water chemistry, light, temperature, water density, pH, and a wide variety of metabolic processes have been treated frequently (see Jack- son, 1964, and Smith, 1950) as environmental factors in algal growth. Reference texts (e.g., Prescott, 1962) indi- cate that some species of algae are epilithic, endophytic, or epizooic, or that many are epiphytic upon vascular plants or larger algae. Other than Chapman (1962) there seems to be no treatment of situations in which the algae have the opportunity to grow on or in association with a variety of substrates and in which clearly some means of preference for one another is established. Tiffany (1951), for exam- ple, suggested that the amount of substrate available to the spores for fixation is of major importance to algal growth, but he fails to include any discussion of the character of the substrate itself, or of the various environmental factors that might be operative in the selection and utilization of a particular substrate. Chapman suggests that, (1) the age and nature of the substrate flora are important, (2) diatoms are more likely to be on older leaves, (3) epiphytes are most numerous on well lighted surfaces, (4) epiphytes are most numerous on submerged material, (5) depressions between cells of the host surface seem to provide better substrate for attachment, and (6) ponds with muddy bottoms have a 446 1974] Algae on Haloragaceae — Colt & Hellquist 447 reduced number of epiphytes. With few epiphytes occur- ring in our collections we tend to agree with the last prem- ise, but we have insufficient evidence to either support or refute the other ideas. We might generally suggest, how- ever, that rather similar principles may operate for both tychoplankters and epiphytes. Our examination of material from different ponds clearly showed that algae grew profusely upon some vascular plants but sparsely or rarely on others. We also found that both the number of species and the types of tychoplankter algae varied with similar plants in different environments. In order to identify, if possible, the operative environmental factors, we selected two sites in which the algal habitats might be contrasted. In both instances, (1) a small leaved aquatic plant was the primary algal substrate, (2) the sub- strate plant grew in the shallower portions of the pond, (3) both ponds and collecting sites were in virtually iden- tical locations beside highways, and (4) there was no shad- ing of the collecting stations by macrovegetation on or near the banks of the pond. THE POND STATIONS 1. Pratt Pond. This pond is located at the junction of Grove Street, Hopkinton Road, and Westboro Road in the town of Upton, Worcester County, Massachusetts. All collections were made in the western sector of Pratt, from water varying between 1.5 and 2.0 meters in depth. Most of the remainder of the pond is sufficiently deep so as to permit swimming and diving. The bottom of the pond is sandy-mud throughout. The pH averaged 6.5 and the alka- linity was 3 ppm. The dominant vascular aquatic in the collection area of Pratt was Myriophyllum heterophyllum Michx., occurring over more than one-third of the center of the bay. In so far as we were able to determine during the several months of field work, no other vascular plant grew in or near the stand of Myriophyllum. This resulted in a clear zone around the stand extending to within several meters of the pond’s edge. 448 Rhodora [Vol. 76 In this shallower water a few other rooted aquatics grew sparsely. Of these Nymphaea odorata Ait., Pontederia cor- data L., and Potamogeton spirillus Tuckerm., were the most common. 2. Icehouse Pond. This pond is beside West Main Street, 0.9 kilometers east of Interstate Route 495, in the town of Hopkinton, Middlesex County, Massachusetts. Samples were collected from water up to 0.5 meters in depth along the southern edge of the pond. The deepest portion of this pond was approximately 1.5 meters, the bottom being mud and gravel. Water loss from this pond during the late summer months left some of the westerly edges as exposed mud and rocks. The pH averaged 6.1, and the alkalinity was 3.5 ppm. The shallow waters of this pond were densely populated with a variety of rooted aquatic plants. Algae were asso- ciated most profusely with Proserpinaca palustris L., which grew in scattered stands of a few plants each. Other com- mon vascular plants in the collection area were Potamogeton pulcher Tuckerm., P. capillaceus Poir., Eleocharis acicularis (L.) R. & S., Pontederia cordata f. taenia Fassett, Nym- phaea odorata Ait., Nuphar variegatum Engelm., Brasenia sehreberi Gmel., Ludwigia palustris var. americana (DC.) Fern. & Grisc., Callitriche verna L., Myriophyllum humile (Raf.) Morong, and Utricularia purpurea Walt. PROCEDURE samples of the vascular plants were collected from the two pond stations and the species determination made using Fassett (1966) and Fernald (1950). Representative speci- mens were pressed and are in the herbarium at Boston State College. samples of the primary substrate plants were collected, some portions retained fresh for immediate scrutiny, and the remainder preserved in a solution of 3% formalin. De- termination of the associated algal species was made from Smith (1924), Irenee-Marie (1939), Prescott (1964), and Patrick & Reimer (1966). Microscope slides were made of 1974] Algae on Haloragaceae — Colt & Hellquist 449 some algae using the Kaiser Glycerin-Gelatin method in order to facilitate drawing by camera lucida. All other samples were placed in Transeau's Fluid for permanent storage. DISCUSSION Twenty-one species of algae were found on or in asso- ciation with Myriophyllum heterophyllum in Pratt Pond. Sixty-one algal taxa were discovered in various degrees of association with Proserpinaca palustris in Icehouse Pond. Seven species were common to each pond. The wide dis- parity in numbers despite virtually simultaneous collection times strongly suggests that Proserpinaca offers a more favorable habitat to the algae than does Myriophyllum. In- asmuch as a difference in surface characteristies might lead to the difference in habitats, it was the first possibility examined. We were not able to substantiate such a premise on any physical basis. One of the problems, however, is that only five genera of epiphytes were recorded during the study. Stigeoclonium and Characium each had one species present infrequently in Pratt Pond, while Bulbochaete and Gloeotrichia each had one commonly found species in Ice- house Pond, and Oedogonium with one common species and one infrequently encountered, both in Icehouse Pond. The pattern of occurrence of these was generally the same as for the tychoplankters and appeared to be due to the overall differences in the habitats rather than individual differences between the substrate plants. Thus it seemed likely that there was a combination of factors operating in each situation, with sufficient variance between the two habitats such as to account for the differ- ence in algal populations. This view is strengthened by the difference in tychoplankter populations, Icehouse Pond hav- ing approximately twice as many taxa. Inasmuch as the tychoplankters do not adhere to a particular substrate, but live in association with the substrate, they would not neces- sarily be affected by physical characteristics. Some previous work gave direction to our scrutiny. An- 450 Rhodora [Vol. 76 derson & Walker (1920) concluded from an investigation of some shallow Nebraska lakes that light and mechanical support are of primary influence to algal growth. They also suggested that shallow water and wind protection give rise to as nearly uniform conditions as possible. Needham & Lloyd (1916) showed that in shallow waters with abundant vegetation currents are reduced or eliminated and there is little possibility of wind disturbance. The authors' own experience has been that tychoplankter populations are par- ticularly rich among the wet sphagnum of New England bogs. Comparison of the two sites on these bases yielded some rather interesting results. It has already been stated that sunlight was readily and equally available to the water in which both sets of substrate plants grew. The collecting area in Icehouse Pond, however, was much shallower than that of Pratt Pond. This difference was negated by the position of the tychoplankter algae in Pratt Pond; they were all within 14 meter of the surface of the water. The only living algae below that level in Pratt Pond were dia- toms, and they were usually obscured by the detritus on the lower branches of plants of Myriophyllum. Physically, Icehouse Pond is protected from surface winds by the mixed deciduous-coniferous forest around it. Al- though its total surface area approximates that of the sec- tion of Pratt Pond from which we collected, we were unable to observe more than mere surface ripples on Icehouse Pond during any visit to the site. Part of the reason for this is the dense vegetation within the water itself. At Pratt Pond, however, we were able to note considerable surface disturb- ance by even relatively light wind currents. We were unable to observe any current within Icehouse Pond. This is largely due to the dense aquatic vegetation. Water movement could easily be discerned in Pratt Pond. As we collected samples in Pratt Pond, broken pieces of Myriophyllum would rise within the stand of plants then drift “downwind.” Our observation of this phenomenon showed that the drag of the wind on the surface of the 1974] Algae on Haloragaceae — Colt & Hellquist 451 water caused the upper portion of the water to move, cre- ating an effective pull on the lower water body. Water then rose through the plants and downwind. Davis (1955) sug- gested that tychoplankters would be unable to adapt evolu- tionarily to such current conditions, On the basis of our limited work we concur, finding no evidence of specialized attachment structures or polymorphism which would give any species an advantage in maintaining its position. It appears to us that the lack of water movement within Ice- house Pond is conducive to an optimum interchange of chemical nutrients. On the other hand, the flow within Pratt Pond apparently reduces a similar interchange, thus inhibiting population growth to some degree. Our final consideration was the growth habit of the small leaved algal substrates and the associated macroflora. In Pratt Pond Myriophyllum produced vertical axes greater than the depth of the water. Near the surface of the water the distal portions of the plants would bend with the drag of the surface water. This appeared to produce a matting effect at the surface which would effectively reduce the light available to lower leveis. The Proserpinaca in Icehouse Pond grew only to the surface of the water and distal flex- ing was rare and then of a minor nature, Light could pass easily through the finely dissected leaves to lower levels. Moreover, the densely crowded vegetation of Icehouse Pond effectively reduced lateral “waving” by the plant. Pro- serpinaca occurred in larger clumps than any of the other small leaved plants in Icehouse Pond (e.g., Myriophyllum, Utricularia). Examination of the broader-leaved aquatics in each pond did not reveal any substantial amount of algae growing on or associated with them. The most frequent species were diatoms, Tabellaria, for example, being the most common. SUMMARY From our observations we conclude that for the optimum growth of tychoplankter algae there are at least three basic requirements. Not necessarily in order of importance they 452 Rhodora [Vol. 76 are: (1) a substrate which provides both maximum surface area and exposure to the available light; (2) an absence of strong or prolonged water currents; and (3) the presence of a varied flora which provides a broad spectrum of nutri- ents within the system. Based on our observations at both ponds we suggest that small leaved plants will most often provide the optimum habitat for tychoplankter algae. We are not convinced, however, that monospecific stands provide the optimum en- vironment for tychoplankters, although such stands may be the best of the available substrates. We have found com- parable situations in bogs, as it has been our experience that the sphagnum bogs with the greatest variety of macro- flora usually yield the largest number of algal species. We further suggest that pH and water temperatures are primarily useful only in contrasting regions with geograph- ical and/or geological differences. It can be reasonably inferred that evolution has already adapted local algal species to pH and water temperature as they occur in the algal habitat. Finally, we concur with previous workers who have sug- gested that quiet, stable and uniform water conditions are best for the growth of tychoplankter algal populations. The list of algae below follows the sequence of Smith (1950). Nomenclature of the Bacillariophyceae is according to Patrick & Reimer (1966). Location and frequency are indicated by the following: IP — Icehouse Pond, PP — Pratt Pond, R — rare (less than 5 collected in all samples), F — frequent (averaged up to 10 in all samples), C — com- mon (averaged more than 10 in all samples), E — epiphyte, T — tychoplankter. CHLOROPHYTA Stigeoclonium subsecundum Kuetzing PP-R-E Bulbochaete Furberae Collins IP-C-E Oedogonium michiganense Tiffany IP-C-E O. pseudoplenum Tiffany IP-C-E Characium ornithocephalum A. Braun PP-R-E 1974] Algae on Haloragaceae — Colt & Hellquist 453 Pediastrum Boryanwm (Turp.) Meneghini IP-R-T P. duplex var. cohaerens Bohlin P. tetras (Ehrenberg) Ralfs P. tetras var. tetraodon (Corda) Rabenhorst Nephrocytium lunatum W. West Kirchneriella contorta (Schmidle) G. M. Smith Scenedesmus abundans var. brevicauda G. M. Smith S. bijuga (Turp.) Lagerheim S. dimorphus (Turp.) Lagerheim S. quadricauda var. maxima West & West - Actinastrum Hantzschii Lagerheim two easily separable species Spirogyra spp. IP-R-T IP, PP-R-T IP-C-T PP-R-T PP-R-T IP-R-T IP, PP-C-T IP-R-T IP, PP-C-T IP, PP-C-T were common in the vegeta- tive state in Icehouse Pond. Gonatozygon pilosum Wolle Triploceras gracile Bailey Closterium Jenneri Ralfs Cl. parvulum Naegeli Cl. Ralfsii var. hybridum Rabenhorst Cl. setaceum Ehrenberg Pleurotaenium coronatum (Breb.) Rabenhorst P. subcoronulatum var. detum West & West P. Trabecula (Ehrenb.) Naegeli P. Trabecula var. rectum (Delp.) W. West P. Trochiscum var. tuberculatum G. M. Smith Euastrum abruptum f. minor West & West E. binale (Turp.) Ehrenberg E. ciastonii Raciborski E. gemmatum Brebisson Micrasterias Nordstedtiana, Wolle IP-C-T, PP-R-T IP-R-T IP-R-T IP-R-T IP-F-T IP-F-T IP-F-T IP-R-T IP-C-T IP-R-T IP-R-T pr IP-R-T IP-R-T IP-R-T IP-R-T 454 Rhodora Cosmarium Blytii Wille C. difficile var. dilatatwm Borge C. furcatospermum West & West C. granatum Brebisson C. isthmium W. West C. ornatum Ralfs C. portianum Archer C. pseudoconnatum Nordstedt C. pseudopyramidatum Lundell C. pyramidatum var. transitorium Heimer] C. tumidum Lundell Staurastrum dejectum Brebisson S. gracile var. nanum Wille S. gracile var. tenuissima Ralfs S. Leptocladum Nordstedt S. Ravenelii var. spinulosum Irenee-Marie Spondylosium planum (Wolle) West & West Desmidium Aptogonum Brebisson D. Baileyi (Ralfs) Nordstedt Xanthidium cristatum Brebisson CHRYSOPHYTA Tabellaria fenestrata (Lyngb.) Kuetzing T. flocculosa (Roth) Kuetzing T. quadrisepta Knudson Meridion circulare (Grev.) Agardh Eunotia elegans Ost. E. formica Ehrenberg E. pectinalis (O.F.M.) Rabenhorst E. pectinalis var. ventricosa Grunow E. praerupta var. bidens (Ehr.) Grunow E. tenella (Grun.) Cleve E. serra var. diadema (Ehrenb.) Patrick Hannaea arcus Patrick Pinnularia flexuosa Cleve P. formica (Ehrenb.) Patrick Frustulia rhomboides (Ehrenb.) de Toni IP-R-T IP-R-T IP-R-T IP-R-T IP-F-T IP-F-T IP-F-T IP-F-T PP-R-T PP-R-T IP-R-T IP-F-T IP-R-T IP-F-T IP-R-T IP-R-T IP-C-T IP-R-T IP-F-T IP-F-T IP-C-T IP-C-T PP-C-T [Vol. 76 IP, PP-C-T IP-C-T IP-F-T PP-F-T PP-F-T IP-F-T PP-F-T IP-F-T PP-R-T IP-F-T IP-R-T IP-C-T 1974] Algae on Haloragaceae — Colt & Hellquist 455 E. vulgaris Thwaites IP-F-T Gomphonema acuminatum var. coronatum (Ehrenb.) W. Smith IP, PP-C-T G. augur Ehrenberg PP-C-T CYANOPHYTA Microchaete tenera (Thuret) de Toni PP-F-T Hapalosiphon intricatus West & West IP-C-T Gloeotrichia Pisum (C. A. Agardh) Thuret IP-F-E LITERATURE CITED ANDERSON, E. N., & E. R. WALKER. 1920. An ecological study of the algae of some sandhill lakes. Trans. Amer. Micros. Soc. 39: 51-85. CHAPMAN, V. J. 1962. The Algae. MacMillan & Co., N. Y. Davis, C. C. 1955. The marine and fresh-water plankton. Mich. State Univ. Press, East Lansing, Mich. Fassett, NoRMAN C. 1966. A manual of aquatic plants. Univ. Wisc. Press, Madison, Wisc. FERNALD, M. L. 1950. Gray's Manual of Botany. American Book Co., New York, N. Y. IRENEE-MARIE, FRERE. 1939. Flore Desmidiale de la Region de Montreal. Laprairie, P. Q. JACKSON, DANIEL F. 1964. Algae and Man. D. F. Jackson, Ed. Plenum Press, New York, N. Y. NEEDHAM, JAMES G., & J. T. LLOYD. 1916. The life of inland waters. Comstock Publishing Co., Ithaca, N. Y. PATRICK, R., & C. W. REIMER. 1966. The Diatoms of the United States exclusive of Alaska and Hawaii. Monograph No. 13, Acad. Nat. Sci. Phila. Prescott, G. W. 1962. Algae of the Western Great Lakes Area. Wm. C. Brown Co., Dubuque, Iowa. 1964. How to know the fresh-water Algae. Wm. C. Brown Co., Dubuque, Iowa. SMITH, G. M. 1924. Phytoplankton of the Inland Lakes of Wiscon- sin. Bull Univ. Wisc. Press, Serial 1048. 1950. Fresh-water Algae of the United States. Mc- Graw-Hill Book Co., N. Y., N. Y. TIFFANY, L. H. 1951. Ecology of fresh water Algae. In: Manual of Phycology, Chronica Botanica, Waltham, Mass. BOSTON STATE COLLEGE BOSTON, MASS. 02115 [Vol. 76 Rhodora 456 Plate 1 1974] Algae on Haloragaceae — Colt & Hellquist 457 Plate 1: a) Eunotia formica Ehrenberg b) E. elegans Ost. c) Staurastrum leptocladum Nordstedt (semicell) d) Gomphonema acuminatum var. coronata (Ehr.) W. Smith e) G. auger Ehrenberg f) Spondylosium planum (Wolle) West & West g) Pinnularia formica, (Ehr.) Patrick h) Pediastrum duplex var. cohaerens Bohlin i) Desmidium Baileyi (Ralfs) Nordstedt j) Gloeotrichia Pisum (C. A. Agardh) Thuret k) Seytonema crispum (C. A. Agardh) Bornet 1) Cosmarium isthmium W. West m) Tabellaria flocculosa (Roth) Kuetzing n) Eunotia praerupta var. bidens (Ehr.) Grunow o) Scenedesmus bijuga (Turp.) Lagerheim p) Characium ornithocephalum A. Braun q) Euastrum gemmatum Brebisson (semicell) r) Scenedesmus abundans var. brevicauda G. M. Smith 458 Rhodora [Vol. 76 "75w A, 50w J, "Sue bl, H24 C-i, km-p Plate 2 1974] Algae on Haloragaceae — Colt & Hellquist 459 Plate 2: a) b) c) d) e) f) g) h) i) j) k) 1) m) n) o) p) Oedogonium pseudoplenum Tiffany Oe. michiganense Tiffany Scenedesmus dimorphus (Turp.) Kuetzing Staurastrum dejectum Brebisson Eunotia serra var. diadema (Ehr.) Patrick Scenedesmus quadricauda var. maxima West & West Desmidium aptogomum Brebisson Staurastrum curvatum var. elongatum G. M. Smith Eunotia pectinalis (O.F.M.) Rabenhorst Pleurotaenium subcoronulatum var. detum West & West Euastrum abruptum fa. minor West & West Bulbochaete furberae Collins Pediastrum tetras var. tetraodon (Corda) Rabenhorst Euastrum ciastonii Rabenhorst Xanthidium cristatum Brebisson (semicell) Nephrocytium lunatum W. West ILLINOIS FIELD AND HERBARIUM STUDIES RoBERT H. MOHLENBROCK AND DAN K. EVANS Further field and herbarium studies of Illinois plants have added several new taxa to the state flora. Several are ad- ventive species from the Chicago area, while others repre- sent native species of taxonomically difficult groups and are treated in some detail. Appreciation is accorded to Mr. Floyd Swink and his colleagues at the Morton Arboretum who kindly made available some of their recent collections. All records cited are in the herbarium of Southern Illinois University (siU), the Morton Arboretum (MOR), or the Missouri Botanical Garden (MO). Poa bulbosa L. The bulbous blue grass, native to Eurasia, is previously unreported from Illinois. Collections have been made of this from cultivated beds at the Morton Ar- boretum. DU PAGE CO.: Morton Arboretum, within the shrub collection in Lacy Land, May 20, 1971, F. Swink (MOR). Phleum pratense L. f. viviparum (S. F. Gray) Louis- Marie. Vivipary, whereby plantlets or vegetative buds re- place the flowers in part or all of the inflorescence, may be encountered in several genera. The phenomonon seems most frequent in the Gramineae, especially such cool season grasses as Poa and Festuca. Vivipary in Scirpus atrovirens (Cyperaceae) is also commonly encountered. Because of the apparent rare occurrence of this type of apomictic re- production in Phleum pratense, this report seems worth- while. DU PAGE CO.: In the Morton Arboretum, July 27, 1971, F. Swink (MOR). Fuirena scirpoidea Michx. The previously known range of this species was along the coastal plain of Georgia, Florida, Alabama, and Louisiana. Its discovery in Illinois at the Dolan Lake Conservation area marks a range exten- tion of more than 500 miles. One colony was found in shallow water growing beside Eleocharis obtusa. Svenson 460 1974] Illinois Studies — Mohlenbrock & Evans 461 (1957), the most recent monographer of the genus, com- bines F. pumila Torrey and F. breviseta Coville with F. squarrosa Michx. The three species now attributed to the Gray's Manual region may be identified in the following key. 1. Leaves reduced to sheaths without blades. F. scirpoidea. 1. Leaf blades well developed. ...................... 2. 2. Achene dark brown with the beak pubescent; scale awn terminal. ............ ET . F. squarrosa. 2. Achene yellow-brown with the beak 2 scale awn arising below the apex. ......... F. simplex. Although F. squarrosa (F. pumila) has been attributed to Illinois (Fernald, 1950; Jones 1950, 1963; Patterson, 1876), a thorough search by the senior author has failed to reveal any such specimens from this state. HAMILTON CO. : Shallow water, Dolan Lake shore in Dolan Conservation area, June 13, 1970, N. Tracy (SIU). Salix X myricoides Muhl. This taxon is a reputed hybrid between S. rigida Muhl. and S. sericea Marsh., although it appears to have more of the characteristics of S. sericea. It differs from S. sericea by its canescent twigs and thinly sericeous capsules, while S. sericea has glabrous or glabrate twigs and densely sericeous capsules. We have recently collected it from Jackson County for its first known locality in Illinois, but it undoubtedly occurs elsewhere in the state. JACKSON CO.: Along rocky stream, Giant City State Park, August 30, 1971, R. H. Mohlenbrock (SIU). Populus X smithii Boivin. This is the hybrid between P. grandidentata Michx. and P. tremuloides Michx., first described from Michigan by Wagner in 1970. It is inter- mediate in several characters between the parent species, the most obvious being the number of dentations along the leaf margins. The following key will serve to distinguish this hybrid from its parents. 1. Margin of leaf dentate, with 5-25 teeth (averaging 10- 20)? budà pubescent. ............... ce wee ces 2. 462 Rhodora [Vol. 76 2. Margin of leaf with 5-15 teeth (averaging 10); petiole 5-10 em long (averaging 7 em). .......... SNNT? P. grandidentata. 2. Margin of leaf with 12-25 teeth (averaging 20); petiole 3-6 cm long (averaging 5.5 em). .......... EEDUVUOOÓO nul P. X smithü. 1. Margin of leaf finely crenate, with 20 or more teeth (averaging 31) ; buds glabrous or nearly so. ...... ... VN P. tremuloides. LA SALLE CO.: Starved Rock State Park, June 16, 1943, G. N. Jones 15791 (MO). PEORIA CO.: Horse Shoe Bottom, July 27, 1919, V. H. Chase 3217 (Mo). Carya pallida (Ashe) Engl. & Graebn. Correspondence with Dr. Wayne Manning, noted authority on the genus Carya, suggested the possibility of C. pallida in Illinois, since it occurs in western Kentucky and southern Indiana. Consequently, the senior author has kept a close vigil for this species in southern Illinois and was rewarded with the discovery of a specimen from Union County referable to C. pallida. The distinction between C. pallida and C. texana Buckl. is difficult. Both have yellow-lepidote buds, and the fruits are not different, although the fruits in C. pallida are usually smaller. Carya pallida usually has at least a few leaves with tomentose rachises, while C. texana usually does not except in var. villosa (Sarg.) Little. In the spring, C. texana has distinctive red hairs, while C. pallida does not. UNION CO.: Dry, acidic woods, along Grapevine Trail, May 21, 1972, R. H. Mohlenbrock (SIU). Quercus veiutina Lam. f. missouriensis (Sarg.) Trel. This form of Black Oak has the lower surface of the leaves densely and permanently pubescent. In this respect, f. mis- souriensis resembles Q. falcata Michx., but differs from this species in leaf shape and bud and acorn characteristics. SALINE CO.: Edge of dry woods, Old Stone Face, July 3, 1971, R. H. Mohlenbrock (SIU). Chenopodium. While preparing the treatment of Cheno- 1974] Illinois Studies — Mohlenbrock & Evans 463 podium for The Illustrated Flora of Ilinois series, the senior author has studied many collections of the genus and has re-evaluated the species previously known from the state. He has followed closely the treatment by Steyermark (1963) in Flora of Missouri in order to make more uniform the treatment of this genus in the central midwest. Three additional recognizable species from Illinois emerged from the study. Chenopodium pallescens Standl. In general, all Illinois Chenopodium with white-mealy, narrow, entire leaves have been designated either C. leptophyllum Nutt. (Fernald, 1950; Gleason, 1952) or C. pratericola Rydb. (Jones, 1963). Illinois material can be separated reliably into two taxa, one to be called C. pallescens Standl, the other C. desic- catum A. Nels. var. leptophylloides (Murr.) Wahl. The following key serves to separate these taxa. 1. Leaves 1-nerved, linear; pericarp firmly attached to readily removed from seed. ©... 1 eee eee nn nn C. desiccatum var. leptophylloides. Chenopodium pallescens is found in rocky ground and in waste areas, primarily in the north-central counties of the state. Chenopodium dissectum var. leptophylloides is almost always in waste ground in the northern half of Illinois. Chenopodium bushianum Aellen. Although Jones (1963) combined this species with C. album L., the conspicuous honeycombed surface of the seeds of C. bushianum makes it a readily recognizable species. It is found occasionally throughout the state in fields, woods, and waste places. Chenopodium strictum Roth var. glaucophyllum (Aellen) Wahl. This taxon is related to C. album L. and C. mis- souriense Aellen. It differs from C. album by its smaller seeds (0.8-1.2 mm broad), and from C. missouriense by its coarsely toothed lower leaves and its calyx lobes which do 464 Rhodora [Vol. 76 not cover the fruit. JACKSON CO.: Waste ground, one mile north of Carbondale, September 3, 1971, R. H. Mohlenbrock (SIU). Draba verna L. var. boerhaavii Van Hall. This variety differs from typical var. verna by its fruits that are never twice as long as broad and by the seeds fewer than 40 per fruit. It is naturalized from Europe and adventive in a mowed field in Illinois. JACKSON CO.: Field, Southern Illi- nois University campus, Carbondale, April 4, 1972, R. H. Mohlenbrock (SIU). Draba reptans (Lam.) Fern. var. micrantha (Nutt.) Fern. This variety, unreported previously from Illinois, occurs occasionally in northern Illinois where it sometimes is found growing with var. reptans. Variety micrantha differs by its hispidulous fruits. Suaeda depressa (Pursh) S. Wats. This species is nor- mally found in the naturally occurring saline soils of south- ern Canada, through the plains states, and south to Texas. In an apparent response to salting by the highway depart- ment, the plant grew along the highway median in the Chicago area. COOK CO.: Median strip, one mile north of Illinois 22, one mile south of Everett Pond, October 7, 1972, R. Schulenberg (MOR). Spergu'aria media (L.) C. Presl. Another adventive ele- ment new to the Illinois flora is this member of the pink family. The previous range of this species appears to be the natural saline soils of central and coastal New York as well as coastal California and Oregon. Illinois collections were made near Elgin along a tollway where salt is fre- quently used by the highway department. KANE CO.: 314 miles west of the Elgin toll booth on the Northwest Tollway right-of-way bank, in sterile clay, September 3, 1972, R. Read. (MOR). Pyrus pyrifolia (Burm. f.) Nakai. The Chinese Pear is an occasional cultivated plant in Illinois. A specimen about 1974] Illinois Studies — Mohlenbrock & Evans 465 thirty-five feet tall has been found growing along a rocky stream in Giant City State Park. This species differs from the more common P. communis L. by its rounder fruits, slightly larger flowers, and sharply toothed leaves. JACKSON CO.: Along Stonefort Creek, Giant City State Park, Septem- ber 11, 1970, R. H. Mohlenbrock (SIU). Crataegus marshallii Egglest. This is one of the few spe- cies of Crataegus in Illinois which has the veins of the leaf running to the sinuses as well as to the points of the lobes. It differs from C. phaenopyrum (L. f.) Medic. by its petioles much longer in relation to the length of the blade, its fewer stamens with reddish anthers, and by its usually two nut- lets per fruit. Its habitat in swampy woods is also dis- tinctive. JACKSON CO.: Swampy woods, Greentree Reser- voir, 2144 miles southwest of Gorham, R. Anderson (SIU). Triadenum virginicum (L.) Raf. The discovery of this species in Lake County brings to four the number of spe- cies of Triadenum known from Illinois. The following key separates these four species. 1. Leaves without punctations. . ....... T. tubulosum. 1. Leaves punctate, at least on the lower surface. .... . 2. 2. Leaves petiolate. ................ ... T. walteri. 2. Leaves sessile. ....... DEMNM o re 9. 3. Sepals obtuse, up to 5 mm long; styles up to 1.5 mm Jong, ........... aa REA T. fraseri. 3. Sepals acute, 5-8 mm long ; styles 2-3 mm long. E. laeua gy ax v7 12 JE NEN T. virginicum. LAKE CO.: Boggy ground, Illinois Beach State Park, Au- gust 3, 1972, R. H. Mohlenbrock (SIU). Mentzelia decapetala (Pursh) Urban & Gilg. This species is native to dry prairies and plains west of the Mississippi River. In Grundy county, apparently its easternmost site, it was collected along a railway where it occurred with other such adventives as Kochia scoparia, Setaria viridis, Helianthus annuus, and Grindelia lanceolata f. latifolia. 466 Rhodora [Vol. 76 GRUNDY CO.: Cinder and gravel ballast, at Eileen, along right-of-way of Santa Fe RR, August 13, 1972, R. Schulen- berg, D. Kropp, & D. Ladd (MOR). Polypremum procumbens L. This species was expected to occur in southern Illinois since it is known from Kentucky to the south and five counties in nearby southeastern Mis- souri. Collections were made from a sandy field in the Horseshoe Lake Conservation area. The inconspicuous corolla, procumbent habit, and slender leaves probably have resulted in this species being overlooked in other localities. ALEXANDER CO.: Low, sandy field, Horseshoe Lake, August 11, 1971, J. Huston 920 (SIU). Jacquemontia tamnifolia (L.) Griseb. This species is na- tive to the southeastern United States and represents a genus new to the Illinois flora. This probable adventive was collected from a disturbed habitat where it grew in associa- tion with Agropyron repens, Ipomoea purpurea, and Eu- phorbia dentata, The following key serves to distinguish Jacquemontia from Ipomoea, Convolvulus, and Calystegia in Illinois. 1. Stigma 1, capitate or with 2 to 3 lobes. ...... Ipomoea. 1. Stigmas 2, noncapitate, without lobes. ............ 2. 2. Stigmas elliptic, oblong or flattened. . Jacquemontia. 2. Stigmas filiform or subulate. ... ........... . 9. 3. Calyx not concealed by large bracts; fruit 2-locu- lar. ....... see Convolvulus. 3. Calyx concealed by 2 large bracts; fruit 1-locular. BENNETT UU Calystegia. GRUNDY CO.: Railroad tracks at Gardner, October 15, 1972, R. Schulenberg & E. Lace (MOR). Vinca major L. The large-flowered periwinkle, an intro- duced European cultivar, was previously known in the wild in the coastal states from North Carolina south to Missis- sippi. In Illinois the species was collected around an old homestead in Pope County where it grew without cultiva- tion. POPE CO.: Randolph Farm, V. Randolph (SIU). 1974] Illinois Studies — Mohlenbrock & Evans 467 Matelea decipiens (Alex.) Woodson. This climbing mem- ber of the milkweed family is widely distributed in central and southern Missouri, but the record cited below is ap- parently the first from Illinois. All three Illinois repre- sentatives of this genus are restricted to the southernmost counties. They may be distinguished by the following key. 1. Flowers greenish-yellow; pedicels glabrous; follicles smooth, angular. ......... 0 .........., M. gonocarpa. 1. Flowers rose, maroon, or rarely cream; pedicels pubes- 2. Petals 1.5-2.5 mm wide, rose or rarely cream. ..... CETERAE id en vue 39 oa A RE yo G's UR tw deen M. obliqua. WILLIAMSON CO.: Low floodplain woods, natural levee of Big Muddy River, north of Colp, May 28, 1972, M. Swayne & J. Swayne (SIU). Physalis macrophysa Rydb. This plant, treated variously as a species, variety, or form, differs from P. subglabrata Mack. & Bush by its translucent leaves and its large, pyra- midal calyces when in fruit. The calyx becomes 3-6 cm long during fruiting. Illinois specimens, previously labelled P. subglabrata, have been seen from Champaign and Peoria counties. Antirrhinum orontium L, The often cultivated lesser snapdragon has been collected from a cultivated oat field. Although it has probably escaped elsewhere in Illinois, this is the first report of such an occurrence. DU PAGE CO.: Oat field, National Accelerator Laboratory near Warrenville, September 28, 1972, F. Swink (MOR). Penstemon cobaea Nutt, This sometimes cultivated beards-tongue was previously known from southern Mis- souri northwest to Nebraska, south to Texas, and east to Arkansas. Within this range the typical habitats are prai- ries, glades, and bluffs. In Illinois this species was collected from a dry meadow where it grew in association with such 468 Rhodora [Vol. 76 prairie elements as Asclepias verticillata, Poa compressa, and Physalis subglabrata. Although this species is not known to be native to northeastern Illinois, the plant asso- ciates as well as the prairie habitat might suggest an indig- enous condition. KANE CO.: Flat, dry meadow, west of Montgomery, near Kendall County line, June 25, 1972, J. Phillips, D. Young, & R. Schulenberg (MOR). Lagenaria siceraria Standl. This highly variable gourd is frequently planted in Illinois and occasionally escapes from cultivation. An early collection from Hancock County and a recent one from Jackson County verify its existence in Illinois. Cucurbita pepo L. var. ovifera (L.) Alef. This variety of the common Field Pumpkin has extremely variable fruits, many of them the source of interesting ornamental gourds such as the Pear, the Bell, the Apple, the Egg, and the Orange. A single collection has been made for this plant in Illinois. UNION CO. : in disturbed soil along a stream, 2 miles north of McClure, July 31, 1971, R. H. Mohlenbrock (SIU). Sherardia arvensis L. This new element of the Illinois flora is adventive but well established in areas of the SIU campus at Carbondale. It may be confused with Galiwm but is distinguished by its long, funnel-form, blue corolla and narrow leaf-like involucres that subtend the flowers. In addition to our location, the species is local in Nova Scotia, southeastern Canada, Missouri, Tennessee, and North Caro- lina. JACKSON CO.: Lawn, SIU campus by the Forest Ser- vice Laboratory, April 30, 1971, D. Evans 11408 (SIU). Campanula rotundifolia L. var. velutina A. DC. The oc- currence of typical C. rotundifolia from the rocky environs of northeastern Illinois is not uncommon. However, variety velutina is heretofore reported only from a single collection by E. J. Hill from Cheboygan County in northern Michigan. We have this taxon from near Apple River Canyon State Park. Here the plant grows in the crevices of an east-facing limestone wall which borders Apple River. Associated spe- 1974] Illinois Studies — Mohlenbrock & Evans 469 cies include the typical form, Aquilegia canadensis, Carex granularis, and the cliff fern Cryptogramma stelleri, Con- sidering the close habitat association with the markedly different typical form, it seems unlikely that this pubescent variety is a mere ecological variant as claimed by some. JO DAVIESS CO.: East-facing limestone wall bordering Apple River, 14 mile northeast of Apple River Canyon State Park, June 13, 1972, D. Evans 1156 (SIU). Grindelia lanceolata Nutt, f. latifolia Steyerm. Prior to this report from Grundy County, f. latifolia was unknown in Illinois. The typical form remains unreported from the state. The closest occurrence of this new taxon to the Il- linois site is the type locality in Stone County, Missouri. Collections are also known from Oklahoma. GRUNDY CO.: At Eileen, in N 14 of Sect. 36, T33N, R6E, August 13, 1972, R. Schulenberg & D. Ladd (MOR). Senecio jacobaea L. Collections from railways and road- sides often produce rare or unusual adventive species. This waif recently has been collected from a railway in the Chi- cago area. DU PAGE CO.: Great Western RR at Euclid Ave- nue, Villa Park, August 17, 1972, F. Swink (MOR). Thelesperma gracile (Torr.) Gray. The natura] habitat and range of this unusual composite are the prairies and plains from Nebraska to Wyoming, south to Arizona, and east to Texas and Oklahoma; also Mexico and South Amer- ica. In addition, Steyermark (1963) reports two adventive sites in Missouri. This roadside collection from Kane county represents the easternmost extension of its range in North America. KANE CO.: Along Ancutt Road, west of Montgom- ery, July 4, 1972, F. Swink (MOR). LITERATURE CITED FERNALD, M. L. 1950. Gray's Manual of Botany, ed. 8. The Ameri- can Book Company, New York. 1632 pp. GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of Northeastern United States and Adjacent Canada. New York Botanical Garden, New York. 3 Vol. 1762 pp. 470 Rhodora [Vol. 76 Jones, G. N. 1950. Flora of Illinois, ed. 2. Am. Midl. Nat. Monogr. 5: 1-368. University of Notre Dame Press, Notre Dame, Indiana. 1963. Flora of Illinois, ed. 3. Am. Midl. Nat. Monogr. 7: 1-401. University of Notre Dame Press, Notre Dame, Indiana. PATTERSON, H. N. 1867. Catalogue of phaenogamous and vascular eryptogamous plants of Illinois. Oquawka, Illinois. 54 pp. STEYERMARK, J. A. 1963. Flora of Missouri. The Iowa State Uni- versity Press, Ames, Iowa. 1725 pp. SvENSON, H. K. 1957. Fuirena. In: North American Flora 18(9): 505-507. SOUTHERN ILLINOIS UNIVERSITY CARBONDALE, ILLINOIS 62901 TWO NEW COLOR-FORMS FROM SOUTHERN MAINE While collecting in the Ocean Park section of Old Orchard Beach, York County, Maine, in recent summers, I have noticed certain plants of Spiraea latifolia, Meadowsweet, with deep-pink flowers. Some of these plants have pubes- cent panicles; others are glabrous. I have found this form growing both in open wet spots and in shaded woodland stations. Spiraea latifolia (Ait.) Borkh. forma rosea T. W. Wells, forma nova. TYPE: Maine: YORK co.: Old Orchard Beach, 11 October, 1969, T. W. WELLS (NEBC). Differt a forma latifolia floribus valde roseis. While collecting in salt-marshes along Goose Fare Creek, which is the southern boundary of Ocean Park, I found plants of Polygonum ramosissimum which differ from the typical in that the sepals have bright-white margins. Polygonum ramosissimum Michx. forma alba T. W. Wells, forma nova. TYPE: Maine: YORK co.: Old Orchard Beach, 5 September, 1970 T. W. WELLS (NEBC). Differt a forma ramosissimum sepalis albomarginatis. THEODORE W. WELLS MILTON ACADEMY MILTON, MASSACHUSETTS 02186 RANGE EXTENSIONS AND NEW RECORDS FOR THE BAHAMA FLORA STEVEN R. HILL In recent years the flora of the Bahama Islands has again aroused the interest of several taxonomists, who have found The Bahama Flora of N. L. Britton and C. F. Millspaugh (1920) to be inadequate in many respects. Not only have numerous nomenclatural changes been proposed, but also since that time collections of species new to the islands have been made (Howard, 1950; Howard & Dunbar, 1964; Lewis, 1971). At the present time The Fairchild Tropical Garden is undertaking the much-needed revision of the outdated work. This writer has made two collecting trips to the Bahamas; the first from April 28 to May 16, 1970, on New Providence Island, Long Island, and Inagua Island, and the second from May 18 to June 3, 1972, on Long Island alone. Both trips were as an undergraduate at Bates College, Lewiston, Maine, and were under the guidance and direction of Dr. Harold E. Hackett of the Department of Biology at that institution. Four collections were from New Providence, 147 from Long Island, and 51 from Inagua. It was noted that the flora of Long Island, a flora which seems to re- semble that of the eastern coast of Cuba, was particularly poorly known. Long Island is an “out island" nearly one- hundred miles long and four miles wide, trending north- south, and bisected by the Tropic of Cancer. Since the flora was poorly known, Walter Cerbin, SSJ, of Clarence Town, Long Island agreed to collect in his area, with the process- ing of specimens to be done at the New York Botanical Garden. To date Father Cerbin's collections number 76 (nos. 102-167 from Long Island, and 1001-1010 from Inagua). These are deposited in this writer's personal herbarium (designated as SRH) housed at the New York Botanical Garden, with duplicates at New York and the Arnold Ar- boretum. 471 472 Rhodora [Vol. 76 According to Britton and Millspaugh (1920), five col- lectors had gathered specimens on Long Island up to the time of their work, and of their collections only 173 num- bers were cited (nos. 495-520, 524 of William C. Coker, 1903; and nos. 6225-6307 of Britton and Millspaugh, 1907). The two others considered to have collected on the island were Swainson (1830-42), whose collections were not known to the authors of The Bahama Flora, and H. F. A. von Eggers (1888), whose collections were apparently not suf- ficiently well-known to be cited. Among those who have touched upon Long Island since that time are Dr. Harold E. Hackett of Bates College (53 numbers in the Duke Uni- versity Herbarium), Father Walter Cerbin, and this writer. The results of the recent collections include several ncw records: 49 species newly reported for Long Island, 4 new to Inagua, and 8 new to the Bahama Islands. The collections also include the rediscovery of a species formerly known only from the type, Zamia lucayana Britton. It is interest- ing to note that many of these new records are of rather common species in the West Indies, reflecting the paucity of the collections from the area to date. Cultivated plants have been included since they commonly escape on the islands and become a part of the local flora. The list of taxa is divided into four sections: 1) species newly reported for Long Island, 2) species newly reported for Inagua, 3) species newly reported for the Bahamas, and 4) a rediscovery. The citations after the collection number refer to those institutions in which the specimen is housed, and the abbreviations are as in the Index Herbariorum (Lanjouw and Stafleu, 1964). The author wishes to thank Dr. William T. Gillis for his help in reviewing the manuscript, Dr. Harold E. Hackett for his guidance and encouragement while in the Bahamas and at Bates College, Drs. Arthur and Noel Holmgren, Dr. Gerrit Davidse, and Mr. Kenneth Becker for their help with certain determinations, and Dr. John T. Mickel, his advisor at the New York Botanical Garden. 1974] Bahama Flora — Hill 473 I. Species Newly Reported for Long Island. A. Native or long established species. POLYPODIACEAE: Acrostichum danaeifolium Langsd. & Fisch.: Hill 840 (sRH) sinkhole, Village Road, S. Clarence Town. Asplenium dentatum L.: Hill 834 (A, SRH) sink- hole, Deadman’s Cay. Pteris longifolia L.: Hill 819b (SRH) moist de- pression, Mangrove Bush. Thelypteris normalis (C. Chr.) Moxley: Hill 125, 835 (SRH) sinkholes, Village Road, S. Clarence Town. GRAMINEAE: Chloris petraea Sw.: Hill 467 (SRH) Clarence Town. Det. G. Davidse. Distichlis spicata (L.) Greene: Hill 123 (A, SRH) Harbor Point, Clarence Town. Det. A. Holmgren. Uniola paniculata L.: Hill 826 (A, SRH) Harbor Point, Clarence Town. AMARYLLIDACEAE: Agave sisalama (Engelm.) Per- rine: Hill 824 (sRH) Clarence Town. Hymenocallis declinata (Jacq.) M. Roem.: Hill 829 (A, SRH) Turtle Cove. BROMELIACEAE: Tillandsia usneoides L.: Hill 670b (US, A, SRH) Galloway's Cave, South Clarence Town. Tillandsia utriculata L.: Hill 529 (sRH) Village Rd., S. Clarence Town. COMMELINACEAE: Rhoeo spathacea (Sw.) Stearn: Hill 244 (sRH) Clarence Town. ORCHIDACEAE: Encyclia bahamensis (Griseb.) Brit- ton: Hill 532 (US, A, SRH) 4 mi. W. of Clarence Town. POLYGONACEAE: Antigonum leptopus H. & A.: Cer- bin 120 (NY, A, SRH) Clarence Town. Coccoloba swartzii Meisner: Hill 471, 841 (SRH) Clarence Town. NYCTAGINACEAE: Commicarpus scandens — (L.) Standley: Cerbin 104 (NY, A, SRH) Turtle Cove. 474 Rhodora [Vol. 76 Guapira bracei (Britt.) Little: Hill 848 (SRH) Turtle Cove. BATIDACEAE: Batis maritima L.: Hill 531 (SRH) Clarence Town. MIMOSACEAE: Acacia farnesiana (L.) Willd. : Cerbin 144 (NY, A, SRH) Village Road, South Clarence Town. Albizia lebbeck (L.) Benth.: Hill 540 (SRH) Clarence Town. CAESALPINIACEAE: Cassia biflora L.: Cerbin 116 (NY, A, SRH). ZYGOPHYLLACEAE: Kallstroemia maxima (L.) T. & G.: Hill 952 (SRH) Clarence Town. EUPHORBIACEAE: Pedilanthus tithymaloides (L.) Poit.: Hill 139 (SRH); Cerbin 139 (NY, A, SRH) Clarence Town. Ricinus communis L.: Hill 542 (SRH) common escape in Clarence Town. MALVACEAE: Abutilon trisulcatum (Jacq.) Urban: Cerbin 160 (NY, A, SRH) Clarence Town. Cienfuegosia yucatanensis Millspaugh: Hill 1015 (SRH) moist depression between Hamilton's and Buckley's. Identified by Dr. Paul Fryxell. Thespesia populnea (L.) Soland.: Hill 129 (a, SRH) Clarence Town. TURNERACEAE: Turnera diffusa Willd.: Hill 845 (SRH) Harbor Point, Clarence Town. PASSIFLORACEAE: Passiflora bahamensis Britton: Hill 515b (A, SRH); Cerbin 166 (NY, A, SRH) Clar- ence Town. CARICACEAE: Carica papaya L.: Hill 142 (A, SRH) occasional in Clarence Town. SAPOTACEAE: Mastichodendron foetidissimum (Jaeq.) Cronq.: Hill 839 (A, sRH) Turtle Cove. APOCYNACEAE: Plumeria obtusa L.: Hill 831 (SRH) Turtle Cove. ASCLEPIADACEAE: Cryptostegia grandiflora (Roxb.) 1974] Bahama Flora — Hill 475 R. Br.: Hill 128 (SRH); Cerbin 153 (NY, SRH) Clarence Town outskirts. CONVOLVULACEAE: Evolvulus squamosus Britton: Cerbin 113 (NY, A, SRH) Mangrove Bush. Merremia dissecta (Jacq.) Hall: Cerbin 117 (Ny, A, SRH) Morrisville. SCROPHULARIACEAE: Stemodia maritima L.: Hill 947 (SRH); Cerbin 119 (NY, A, SRH) 4 miles west of Clarence Town. Determined by Dr. Noel Holmgren. ACANTHACEAE: Blechum brownei Juss.: Cerbin 126 (A, NY, SRH) Clements. Ruellia tuberosa L.: Hill 846 (SRH) Clarence Town. RUBIACEAE: Casasia clustifolia (Jacq.) Hitch.: Cer- bin 141, 152 (NY, A, SRH) Salt Pond. ASTERACEAE: Bidens pilosa L.: Cerbin 111 (NY, A, SRH) Mangrove Bush. Wedelia trilobata (L.) Hitche.: Cerbin 136 (Ny, A, SRH) Salt Pond. Verbesina encelioides (Cav.) Benth. & Hook.: Cerbin 159 (NY, A, SRH) Clarence Town. Deter- mined by K. Becker. B. Cultivated or escaped. AMARYLLIDACEAE: Hippeastrum puniceum (Lam.) Urban: Hill 1017 (sRH) Hamiltons. FABACEAE: Sesbania grandiflora (L.) Pers.: Cerbin 102 (NY, A, SRH) Turtle Cove. EUPHORBIACEAE: Codiaeum variegatum (L.) Blume: Cerbin 122 (NY, A, SRH) Clarence Town. ARALIACEAE: Polyscias guilfoylei (Bull. Bailey: Cerbin 114 (NY, A, SRH) Morrisville. APOCYNACEAE: Thevetia peruviana (Pers.) K. Schum.: Hill (A, SRH) Clarence Town. OLEACEAE: Jasminum fluminense Vell.: Cerbin 140 (NY, A, SRH) Salt Pond. SCROPHULARIACEAE: Russellia equisetiformis Schl. & Cham.: Cerbin 133 (NY, A, SRH) Deadman’s Cay. 476 II. III. Rhodora [Vol. 76 CUCURBITACEAE: Momordica charantia L.: Cerbin 125 (NY, A, SRH) Clements. Species Newly Reported for Inagua. LILIACEAE : Sansevieria hyacinthoides (L.) Druce: Cer- bin 1001 (NY, A, SRH) Along road to Northwest Point. PASSIFLORACEAE: Passiflora bahamensis Britton: Hill 515 (sRH) Union Creek. APOCYNACEAE: Nerium oleander L.: Hill 488a & b. (SRH) Matthew Town. CONVOLVULACEAE: Merremia dissecta (Jacq.) Hall: Hill 489 (SRH) Matthew Town. Species Newly Reported for the Bahama Islands. AMARYLLIDACEAE: Crinum asiaticum L.: Cerbin 124 (NY, A, SRH) Clarence Town, Long Island. Cultivated and possibly escaped in moist places. Zephyranthes citrina Baker: Hill 827 (A, SRH) Clar- ence Town, Long Island. Zephyranthes puertoricensis Traub: Hill 836 (SRH) Hamilton’s, Long Island. Also in cultivation from this population at NY. AMARANTHACEAE: Alternanthera brasiliana (L.) Kuntze: Cerbin 127 (NY, A, SRH) Clements, Long Is- land. Determined by Mr. Kenneth Becker, NYBG. CAESALPINIACEAE: Bauhinia variegata L.: Hill 566 (SRH) Nassau, New Providence. Commonly cultivated and escaped. MALPIGHIACEAE: Malpighia glabra L.: Cerbin 1005 (NY, A, SRH) Matthew Town, Inagua. Cult. Cultivated and possibly escaped. CACTACEAE: Opuntia rubescens (Salm-Dyck) Lemains: Hill 476 (sRH) Matthew Town, Inagua. Dooryard. SOLANACEAE: Datura inoxia Mill.: Hill 850 (SRH) Clar- ence Town, Long Island. Roadside. Also Nash and Taylor 1357, formerly determined as D. metel L. but re-identified by H. N. Moldenke. Inagua. IV. Rediscovery. CYCADACEAE: Zamia lucayana Britton: Hill 828 (Mo, 1974] Bahama Flora — Hill 477 K, US, A, F, BM, TRIN, SRH, NY). Found in limestone and sand 100 yards from the sea, about five miles north of the type locality; Turtle Cove, Long Island. Formerly known only from the type collection “In a sandy coastal thicket Clarence Town, Long Island, only one plant found after a long search (Britton and Millspaugh 6271)” (Britton, 1907). This writer discovered an extensive colony with many old plants as well as a thriving understory of seedlings and hermit crabs which were feeding on the fleshy seed coats. Plants in the shade generally exhibited much longer leaves than those in sun, those specimens in shade agreeing more closely with the type. Male and female cones were collected, and living material is in cultivation at NYBG, Harvard, and the Fairchild Tropical Garden. LITERATURE CITED BRITTON, N. L. 1907. Contributions to the Flora of the Bahama Islands. IV. Bull. N. Y. Bot. Gard. 5: 311. BRITTON, N. L. & C. F. MiLLsPAUGH. 1920. The Bahama Flora. Hafner Publishing Co., Inc. New York, Howanp, R A. 1950. Vegetation of the Bimini Island Group. Ecol. Monog. 20: 317-349. , & H. F. DUNBAR. 1964. Additions to the Flora of Inagua, the Bahamas. Rhodora 66: 6-15. LANJOUW, J. & F. A. STAFLEU. 1964. Index Herbariorum. 5th. ed. Part I. The Herbaria of the World. Reg. Veg. 31. Utrecht. LEWIS, W. H. 1971. Additions to the Flora of the Bahama Islands. Rhodora 73: 46-50. NEW YORK BOTANICAL GARDEN BRONX, NEW YORK 10458 A NEW SPECIES OF LECHEA (CISTACEAE) FROM PENINSULAR FLORIDA ROBERT L. WILBUR! A dozen years ago a strikingly different Lechea was noted (Rhodora 63: 117. 1961.) among collections from southern Florida. It then seemed unwise, however, to de- scribe the population that they represent as a distinct new taxon since the specimens available were not as complete or mature as would be desired in proposing a new entity in such a perplexing and technical genus. However, additional collections from southern Florida and the low-keyed but persistent hints from Dr. Olga Lakela that something wasn’t quite satisfactory with available treatments finally prodded me into taking another look. Her own collections have largely eliminated one of my reservations about for- mally proposing a new taxon, although additional collections from throughout the growing season are still very much to be desired. Practically nothing is known of the plants ex- cept in the fruiting stage and although that is the most important stage for the accurate identification of members of this genus, it still would be most desirable to know something more about the vegetative features of this spe- cies. Leaves in this species are almost completely lacking by the time fruits are ripening. Still it seems probable that more of a stimulus for additional collections of this taxon and of the other highly distinctive representatives of the genus in peninsular Florida will be forthcoming if the taxon is formally proposed now than if we wait for more col- lections. KEY TO THE PENINSULAR FLORIDIAN SPECIES OF LECHEA 1. Pubescence of aerial stems mostly strongly divergent and spreading. ......ssse e nn 2. "This study was supported by NSF GB-13815, for which grateful acknowledgment is made. 478 1974] Genus Lechea — Wilbur 479 2. Internal sepals conspicuously V- or U-shaped in cross-section with thin scarious margins and a roughened, often sparsely pilose keel (the sepal otherwise glabrous) ; leaves often over 1.5 cm. long; capsule thin-walled, readily splitting into 3 valves at maturity, subglobose, about equaling the calyx in length. .... 1. L. mucronata Raf. (L. villosa Ell.). Internal sepals but slightly bowed in cross-section with texture appearing uniform and pubescence scat- tered across surface; leaves all less than 1 cm. long; capsule thick-walled, indehiscent, ellipsoid or some- what rounded- barrell-shaped, exserted from the calyx for at least 1/3-1/2 its length. .............. ems Lis 2. L. divaricata Shuttlew. ex Britt. 1. Pubescence of aerial stems mostly closely appressed or THOME ea iio ae on an a e oo owe ees ale ee 3. 3. External sepals equaling or exceeding the internal sepals in length. ........... 00. ce eee n n n 4. 4. External sepals at least one-fifth longer than the inner sepals and usually equaling or exceeding the capsule in length; capsule equaling or exceed- ing the inner sepals by not more than one-fifth its length; cauline leaves usually elliptic to elliptic- oblong, usually less than 5 times as long as wide, those below the inflorescence commonly appear- ing whorled and often more than 2 mm. wide. ck ct acs eo tie eet zu. De dA minor L. 4. External sepals about equaling the inner sepals and never equalling the capsule in length; capsule usually exceeding the inner sepals by about 1/3- 1/2 its length; cauline leaves narrowly oblong to linear, usually 6 times (or more) as long as wide, those below the inflorescent alternate and less than 2 mm. wide. ..........-..- rnm n HERR 4. L. sessiliflora Raf. (L. patula Legg.). 3. External sepals shorter than the internal sepals. . 5. 5. Leaves pubescent on both upper and lower sur- 480 Rhodora [Vol. 76 faces (at least those of the basal shoots conspicu- ously pilose above and below while the cauline and rameal leaves are usually inconspicuously pubescent over entire surface); flowers or fruits mostly clustered in 2's or 3's; capsule wall thick- ened and indurate. ........ 5. L. cernua Small. Leaves variously pubescent below but glabrous on upper surface; flowers or fruits not appearing fascicled (but obviously attached separately) ; capsule wall thin or at least not conspicuously indurate. 2.0.0.0... e 6. 6. Aerial stems perennial, suffruticose, clearly woody at base, with wiry woody branches; capsule exserted from the often spreading calyx by 1/3-1/2 its length; calyx sparingly short-pubescent to glabrous. ... ...... ... Dee eee eee eee ees 6. L. deckertii Small. 6. Aerial stems annual, herbaceous, dying to the base each year; capsule equaling the calyx or exserted not more than 1/5 its length from the closely enveloping sepals; calyx moder- ately to densely pilose. ............. sus. 7. 7. Leaves abruptly tapering at apex into a hardened, shiny, conical callosity about 0.25 mm. long; inner sepal clearly 3-nerved (often best demonstrated by moistening) ; pedicels averaging over 1.5 mm. long; capsule exceeding the sepals by about 1/5 its length; seeds mostly 2. ... 7. L. pul- chella Raf. (= L. leggettii Britt. & Holl.). 7. Leaves pointed but not differentiated into a callosity; inner sepal 1-nerved; pedicels averaging less than 1.5 mm. long; capsule almost completely enveloped by the sepals; seeds mostly 3-6. ...............00055 8. 8. Calyces densely appressed pilose; stems and under surface of the leaves 1974] Genus Lechea — Wilbur 481 or at least the midvein and often the margins appressed pilose; capsules equaling or slightly shorter than the closely enveloping inner sepals. ...... PW ga EE 8. L. torreyi Legg. ex Britt. 8. Calyces glabrous; stems and leaves completely glabrous; capsules very slightly exceeding the inner sepals at Maturity l.c vs vs 9. L. lakelae. Lechea lakelae Wilbur, sp, nov. Caudex simplex vel paucis aliquotve ramis; folia ramealia peranguste linearia-elliptica, acuta, 0.6-1.2 cm. longa et 0.3-1.0 mm. lata, subtus costa et margine glabra; pedicelli glabri, (0.8) 1.0(1.2) mm. longi; calyx fructifer obovoideus, 1.5-1.9 mm. longus et 1.2-1.6 mm. latus; sepala interiora glabra et leviter carinata, 1.5-1.9 mm. longa et 1.2-1.6 mm. lata; sepala exteriora glabra linearia vel anguste lanceolata 1/2 vel 2/3 plo breviora quam sepala interiora; capsula c. 1.2-1.6 mm. longa; semina 3, c. 0.8 mm. longa. Caudex simple or with few to several branches; basal resting shoots unknown; aerial (i.e. flowering and fruiting) stems 1 to several, 2-3 (4) dm. tall, branching mostly above the middle and forming a rather compact, wiry-stemmed, bushy top with completely glabrous axes; cauline leaves lacking on all specimens seen; rameal leaves very narrowly linear-elliptic, tapering to both the apex and base, 0.6-1.2 em. long and 0.3-1 mm. wide, completely glabrous above and below with the midvein somewhat elevated beneath; fruit- ing branches and pedicels glabrous, the pedicels (0.8)1.0 (1.2) mm. long; fruiting calyx obovoid, broadest above the middle and tapering into the broadly obpyramidal, coria- ceous base, drying dark reddish brown, completely glabrous; inner sepals about 1.5-1.9 mm. long, including the indurate, obpyramidal base, and 1.2-1.6 mm. broad, slightly exceeded by the mature capsule, very broadly elliptic to almost or- bicular, apically broadly rounded, only the strongly elevated 482 Rhodora [Vol. 76 or slightly keeled midvein clearly visible; outer sepals linear to narrowly lanceolate, about 1/2-2/3 as long as the inner; capsules broadly barrel-shaped, cylindrical, about 1.2-1.6 mm. long and 1-1.2 mm. in diameter, the valves firm, in- durate, tardily dehiscing about 1/2-2/3 the distance from apex to base; seeds 3, equilateral, dorsiventrally compressed, about 0.8 mm. long and 0.4 mm. wide, blackish. TYPE: Collier Co., Florida: Marco Island, moist level grassy area, higher beach of recently made lake. 7 August 1967. Lakela 30953 (Holotype, DUKE; Isotype, USF). OTHER SPECIMENS EXAMINED: Collier Co.: coastal strand, Serenoa- Ceratiola scrub, growing with Cyperus in white sand, Lakela 31879 (DUKE); Marco Island, coastal strand — Pinus elliottii association, Lakela & Almeda 31567 (DUKE); Marco Island with Indigofera, Poly- gonella, in Ceratiola-Quercus scrub, on U.S. 92, Lakela 27852A (usF); lagoon head, upper beach, Lakela 31673 (DUKE). My recollection and rather brief notes made more than a dozen years ago indicate that the species also occurs or occurred on Florida’s eastern coast in Broward Co.: sandy scrub above Fort Lauderdale, Buswell 24 July 1936 (Bus!). The apparent nearest relative of the new species, Lechea lakelae, is L. torreyi. The most obviously distinctive fea- tures are indicated in the key, but the two taxa may be readily distinguished at a glance. Obviously I feel that both merit recognition at specific rank. In the mind of some there still may remain a nagging suspicion that the striking- ly distinctive plants of the newly described taxon are merely a glabrous form of L. torreyi. I feel, however, that the ap- parently consistently longer capsules when compared to the length of the inner sepals suggest that L. lakelae is more than that. This species is named in recognition of Dr. Olga Lakela upon whose keen eye and careful collections our present knowledge of this taxon is largely based. It is humbling to recall that her recent significant contributions to south- eastern botany were made in her “retirement” years after an active career as a teacher and researcher in Minnesota. 1974] Genus Lechea — Wilbur 483 Not many botanists are destined to write floras of as distant regions as one of Minnesota's northeastern counties and Florida's southern tip. DEPARTMENT OF BOTANY DUKE UNIVERSITY DURHAM, N.C. 27706 PIMPINELLA SAXIFRAGA L. (UMBELLIFERAE) IN WISCONSIN: This species, adventive from Eurasia, has heretofore been reported to extend from Newfoundland and New Brunswick to Delaware and the District of Columbia, west to Pennsylvania, Ohio, and Indiana (Mathias & Con- stance, 1944. Umbelliferae, in North American Flora 28B (1): p. 133; Fernald, Gray’s Manual, 1950; Gleason, Illus- trated Flora, 1952; Gleason & Cronquist, Manual of Vascu- lar Plants, 1963). Jones (Flora of Illinois, 1950) and Steyermark (Flora of Missouri, 1963) do not mention the plant, so it appears not to have spread westward. I report here its discovery in Wisconsin: WINNEBAGO CO.: roadside at junc. Country Club Road and Black Wolf Point Road, sect. 24, TI7N, R16E, 24 September 1967, Below 146; in roadside ditch ca. 144 mile from the junc. of county trunks YY and N, on N, sect. 20, T17N, R16E, 26 September 1972, Rhyner 044. The two specimens are deposited in the herbarium here at Oshkosh. The curators of the herbaria at MIL, UWM, and WIS have no records of the plant from Wisconsin; Iltis at WIS has kindly confirmed the identifications. The plant's occurrence at two sites four miles apart and spanning a five-year period warrants its inclusion as an element in the Wisconsin flora. NEIL A. HARRIMAN BIOLOGY DEPARTMENT U. OF WISCONSIN — OSHKOSH 54901 A SECOND REPORT OF THE PROTHALLIA OF LYCOPODIUM INUNDATUM IN NORTH AMERICA' Davip M. LANE AND A. LINN BOGLE Gametophytes of Lycopodium inundatum L., first re- corded for North America in 1972 (Bruce, 1972) from two localities in Michigan, were found growing at the edges of shallow wet depressions along the margins of a sandy access road on the shore of an inactive and flooded sandpit (Figs. 1, 2) in Somersworth, Strafford Co., New Hampshire. The mature sporophytes of L. inundatum var. inundatum (compare Gillespie, 1962) are abundant along the shoreline, often forming dense mats (Fig. 3). Samples of sandy soil containing the smallest visible sporophytes were collected in October and examined accord- ing to Bruce’s technique. Some of the young sporophytes were still attached to living gametophytes, as adjudged by the green color, turgidity, and characteristic lobing of the latter. Gametophytes were also found which had not yet produced sporophytes. Gametophytes ranged in size from less than one mm. in diameter for one which bore no sporophyte, up to about three mm. in diameter among those bearing sporophytes. Included in the latter category were one gametophyte bear- ing a sporophyte which had produced about ten microphylls, and two gametophytes each bearing two young sporophytes (Fig. 4). In contrast, however, some very young sporo- phytes were found to have no attached gametophytes, the latter apparently having rotted away, even though the sporophytes bore only one or two microphylls. This condi- tion appeared particularly prevalent where the soil particles were overgrown with moss protonemata and algae. In addition to variation in size discussed by Bruce, vari- ations in occurrence and condition, as well as in form (Bold, 1973), of the gametophyte in relation to differences in the ‘Published with the approval of the Director of the University of New Hampshire Agricultural Experiment Station as Scientific Con- tribution No. 756. 484 1974] Lycopodium inundatum — Lane & Bogle 485 microhabitat suggest the need for microecological studies in the field and laboratory. These would complement lab- oratory study of the culture and development of gameto- phytes of other species (Freeberg, 1962; Freeberg and Wet- more, 1957) and of the embryogeny of the sporophyte of Lycopodium inundatum (Bruce, 1972). The fact that spores of Lycopodium inundatum germinate readily and produce green photosynthetic gametophytes which live on the surface of the soil is of potential value to biology teachers (Bierhorst, 1964) in areas where the spe- cies occurs in New England (Fig. 5; for cireumpolar range map see Hultén, 1968, p. 26), for the gametophytes might be grown on artificially denuded patches of soil next to the sporophytes. Although gametophytes of mosses and ferns are readily available and commonly used for study in biology classes, few students ever see the live, fleshy gametophytes of Lycopodium. It is possible that careful searching in suit- able locations near colonies of mature sporophytes would produce numerous live gametophytes for class use. LITERATURE CITED BrERHORST, D. W. 1964. Suggestions and Comments on Teaching Materials of the Non-Seed Bearing Plants. In: Vestal, P. A. et al. 1964. The use of living material in the teaching of botany. Amer. Biol. Teacher. 26: 89-113. BorLp, H. C. 1973. Morphology of Plants, Ed. 3. Harper & Row, New York. 688 p. BRUCE, J. G. 1972. Observations on the Occurrence of the Pro- thallia of Lycopodium inundatum. Amer. Fern Jour. 62: 82-87. FREEBERG, J. A. 1962. Lycopodium prothalli and their endophytic fungi as studied in vitro. Amer. Jour. Bot. 49: 530-535. , and R. H. WETMORE. 1958 (dated 1957). Gametophytes of Lycopodium as grown in vitro. Phytomorphology 7: 204-217. GILLESPIE, J. P. 1962. A Theory of Relationships in the Lyco- podium inundatum Complex. Amer. Fern Jour. 52: 19-26. HuLTÉN, E. 1968. Flora of Alaska and Neighboring Territories. Stanford Univ. Press. 1008 p. Seymour, F. C. 1969. Flora of New England. Charles E. Tuttle Co., Rutland, Vermont. 596 p. DEPARTMENT OF BOTANY AND PLANT PATHOLOGY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 486 Rhodora [Vol. 76 Figure 1. Typical macrohabitat with abundant mature sporo- phytes along brook (to left) and pond (to right) of access road (center). Note area of Fig. 2 (box). Figure 2. Microhabitat of gametophytes at edges of depression (left of center) darkened by growth of moss protonemata and algae. 1974] Lycopodium inundatum — Lane & Bogle 487 — Figure 3. Vertical view of dense mat of mature sporophytes (each branch less than one cm. thick). Figure 4. Gametophyte bearing two young sporophytes (upper right dark area was green and minutely lobed, the remainder of the gametophyte was colorless). 488 Rhodora [Vol. 76 Figure 5. County distribution of Lycopodium inundatum varieties in New England (plotted from Seymour, 1969): var. inundatum (dots), var. Bigelovii Tuckerm. (circles), and var. robustum R. J. Eat. (triangles). Note site of gametophytes (X) discussed here. NEW AND INTERESTING PLANTS FROM THE CENTRAL PLAINS STATES LAWRENCE K. MAGRATH AND RONALD R. WEEDON As a result of field and herbarium studies, a number of new or interesting records of plants have been found for Texas, Oklahoma, Kansas, and Nebraska. All specimens cited, unless indicated otherwise, are located at KANU. Corallorhiza wisteriana Conrad. Nebraska: DAWES CO.: Chadron State Park, open ponderosa pine wooded hillside, pine needle litter, clay loam soil, two stems growing in ravine, in fruit, 14 July 1972, Magrath & Weedon 7620 (KANU). This collection represents an extension of the range of the coralroot southward from the Black Hills of South Dakota into the Pine Ridge of northwestern Nebraska. Holosteum umbellatum L. Texas: COLLINGSWORTH CO.: 0.3 mi E of the Salt Fork of the Red River on T 203, growing in sandy soil in roadside ditch, scattered and uncommon, 10 April 1973, Magrath & Richardson 7936 (KANU). KEN- EDY CO.: 18.3 mi S of Armstrong on US 77, growing in railroad right-of-way, 20 March 1972, Magrath, Rice & Johnston 7225 (KANU, OKLA). This collection represents an extension of the range of the species westward from Oklahoma into the Texas Panhandle and southward from Oklahoma to the Gulf Coastal area of southern Texas. Scleranthus annuus L. Oklahoma: GRADY CO.: Oklahoma College of Liberal Arts Campus (17th and Grand Avenue) in Chickasha; growing in flower bed on E side of Austin Hall; sandy soil; rare; associated with Arenaria, Bromus and Holosteum, flowers greenish-white, apetalous; 24 May 1973; Magrath 8032 (KANU). This collection represents an extension of the range of this species southwestward from Missouri and eastern Kansas. 489 490 Rhodora [Vol. 76 Ranunculus testiculatus Crantz. Nebraska: DAWES CO.: Chadron State Park, 8 mi S of Chadron on Hwy #385, in campground in waste areas in shade of deciduous trees, weedy areas dominated by Taraxacum officinale, sandy soil, abundant, 6 June 1972, Weedon & Gates 7011 (KANU); Doris Gates, 29 April 1970. The range of this species is now known to be extended eastward into northwestern Nebraska. The above collection indicates the probable establishment of the species in abundance in this area. Thlaspi perfoliatum L. Oklahoma: KAY CO.: rest area on I-35, 2.8 mi N of Blackwell-Medford Exit; clay loam soil, somewhat sandy; small area near moist drainage ditch; only one colony seen, apparently introduced; associated with Bromus, Holosteum and Rumex; 6 May 1973; Magrath 7969 (OKL). This collection represents an extension of the range of the species southwestward from Missouri and Kansas. Eryngium prostratum Nutt. var. prostratum. Kansas: CHEROKEE Co.: 1.5 mi S of Galena on K 26, small cut-over and disturbed marshy area in Ozarkian oak-hickory woods, gravelly soil, associated with Juncus and sedges, apparently rare — only one plant found, prostrate vine rooting at the nodes, flowers bluish-purple, 22 October 1972, Magrath 7855 (KANU). This collection represents an extension of the range of the taxon westward from the eastern Ozarks of Missouri, and northward from southeastern Oklahoma. Lactuca saligna L. f. ruppiana (Wallr.) G. Beck. Oklahoma: OKLAHOMA C0.: ca. 3 mi N of the Jct of I-44 and I-35 on I-35, a few scattered plants growing in road-cut, sandy loam soil, 5 August 1972, Magrath 7700 (KANU, OKL, OKLA). This collection represents an extension of the range of this taxon southwestward from Missouri and Kansas. LAWRENCE K. MAGRATH RONALD R. WEEDON DEPARTMENT OF BIOLOGY DEPARTMENT OF BIOLOGY OK. COLLEGE OF LIBERAL ARTS CHADRON STATE COLLEGE CHICKASHA, OK. 73018 CHADRON, NEB. 69337 A NEW DROSERA FROM VENEZUELA J. A. STEYERMARK AND L. B. SMITH Shortly after entering the Grand Savanna of Venezuela on the northern side we left the road to examine a sandy area with scattered small shrubs. On the patches of white sand between was a Drosera so small that it could easily be covered by a quarter and with so short a peduncle that the one or two flowers appeared practically sessile. This new species was such a happy augury for the Smiths' intro- duction to the Guayana Highland that we have christened it as follows: Drosera felix Steyermark & Smith, sp. nov. Fig. 1. Plantae solitariae; caulibus curtissimis; foliis dense rosu- latis, patentibus; stipulis ad petiolum 1/3 adnatis, ad me- dium 3-laciniatis, membranaceis, 2.5-3 mm longis, basi vaginantibus, solidis, 1 mm latis, segmentis laciniatis an- guste triangulari-lanceolatis, longe attenuatis, 1.7-2.2 mm longis, 0.2-0.3 mm latis, glabris; petiolis 5-5.5 mm longis, 0.7 mm crassis, supra minute papilloso-verruculosis et pilis patentibus paucicellularibus acicularibus laxissime vestitis, subtus pilis multis adpressis simplicibus acicularibus onus- tis; laminis suborbicularibus, apice late rotundatis, 3 mm longis, 2.5 mm latis, castaneo-purpureis, supra margine glandulis 2-4 stipitatis et centro glandulis subsessilibus vestitis, subtus pilis eis petioli similibus; pedunculo nullo vel haud ultra 2.5 mm longo, eglanduloso; bractea lineari- lanceolata, acuminata, 1.5 mm longa, 0.1 mm lata, ciliata; floribus 1-2; pedicello 1-5 mm longo, pilis adpresso-adscen- dentibus simplicibus et glandulis remotis sessilibus atro- purpureis vestito; hypanthio cupulato-subcampanulato, 0.8 mm longo, 1.2 mm lato, pilis patentibus remotis vestito; sepalis 5, lanceolatis, 1.5-1.8 mm longis, sub medium 0.8 mm latis, e pilis laxis ascendentibus simplicibus paucicellularibus hirsutulis; petalis 5, patentibus, albis, obovatis, apice ro- tundatis, basi cuneatis, 2.5 mm longis, 1.5 mm latis; stami- 491 492 Rhodora [Vol. 76 Fig. 1, a-h, Drosera felix: a, habit; b, single leaf (upper side); c, single leaf (lower side); d, hair detached from petiole; e, calyx and pistil with pedicel; f, stamen; g, petal; h, seed, ventral and dorsal views from left to right. 1974] Venezuela Drosera — Steyermark & Smith 493 nibus 5, quam petalis brevioribus; filamentis 1 mm longis, glabris; antheris subhemisphaericis, 0.5 mm longis, 0.4-0.5 mm latis; ovario subgloboso, 1 mm longo, 1 mm lato, glabro; stylis 3, ad basim bipartitis, ramis 1.7 mm longis, glabris. Venezuela: BOLÍVAR: Carretera El Dorado to Santa Elena de Uairen south of El Dorado, alt 1200-1400 m, 19 Feb 1972, Julian A. Steyermark, Lyman & Ruth Smith, G. C. K. & E. Dunsterville 105468 (holotype VEN, isotypes NY, US). INSTITUTO BOTANICO CARACAS, VENEZUELA AND DEPARTMENT OF BOTANY U. S. NATIONAL MUSEUM WASHINGTON, D. C. 20560 INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA Manuscripts should be submitted in duplicate and should be double-spaced or preferably triple-spaced (not on cor- rasable bond), and a list of legends for figures and maps provided on a separate page. Footnotes should be used sparingly, as they are usually not necessary. Do not indi- cate the style of type through the use of capitals or under- scoring, particularly in the citations of specimens, except that the names of species and genera may be underlined to indicate italics in discussions. Specimen citations should be selected critically especially for common species of broad distribution. Systematic revisions and similar papers should be prepared in the format of “The Systematics and Ecology of Poison-Ivy and the Poison-Oaks,” W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particularly with reference to the indentation of keys and synonyms. Papers of a floristic nature should follow, as far as possible, the format of “Contribution to the Fungus Flora of Northeastern North America. V.,” H. E. Bigelow & M. E. Barr, Rhodora 71: 177- 203. 1969. For bibliographic citations, a recommended list of standard journal abbreviations is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300, 1958. Volume 76, No. 807, including pages 315-495, was issued Nov. 15, 1974. 495 CONTENTS: — continued Two New Color-forms from Southern Maine Theodore W. Wells .......... eren nnne ener Range Extensions and New Records for the Bahama Flora Steven R. Hill 1... eese eene tornato tne eo osten atteso ases e tenet otto A New Species of Lechea (Cistaceae) from Peninsular Florida Robert L, Wilbur ccccccccccccsscccccsccssecssecesseesnseceeeseeseessseeenseneeeeaeees Pimpinella Saxifraga L. (Umbelliferae) in Wisconsin Neil A. Harriman ....... eere eene enne eet tnnt neenon seno eno A Second Report of the Prothallia of Lycopodium inundatum in North America David M. Lane and A. Linn Bogle ............ ee New and Interesting Plants from the Central Plains States Lawrence K. Magrath and Ronald R. Weedon .................. A New Drosera from Venezuela J. A. Steyermark and L. B. Smith ........ eee Instructions to Contributors ............ eene 470 471 478 483 484 ———— Dodota JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALFRED LINN BOGLE, Editor-in-Chief ROLLA MILTON TRYON STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY RICHARD EDWIN WEAVER Associate Editors Vol. 76 December, 1974 No. 808 CONTENTS: Phenology and Physiognomy of the Hydrophyte Community in Otsego Lake, N. Y. Willard N. Hammam eene nnne nennt nter Panicum ensifolium Baldw. — New for Mexico Monn EONIA a iedera a REED Significant New Distributional Records for the Genus Sphagnum in the Northeastern United States Richard E. Andrus ersessostastootestlüesesastssecesrepesosevveseveeevecectéasesoaoseneven Environmental Control of Needle Characteristics in Sub- alpine Black Spruce Brian F. Chabot eeseshssesssthsósssstsosshseshpseesseosttsesereeeveeceooccessausnsecnasne (Continued on Inside Back Cover) 497 509 511 The Nem England Botanical Club, Jne. Botanical Museum. Oxford St.. Cambridge. Mass. 02138 RHODORA. — A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price $20.00 per year, net, postpaid, in funds payable at par in the United States currency at Boston, Some back volumes, and single copies are available. For information and prices write RHODORA at address given below. 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 submitted in duplicate, and 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. Ex- tracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Dept. of Botany, University of New Hampshire, Durham, New Hampshire 03824 Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to RHODORA, Botanical Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive the next number of RHODORA, changes of address must be received prior to the first day of March, June, September or December. Second Class Postage Paid at Boston, Mass. MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 76 December, 1974 No. 808 PHENOLOGY AND PHYSIOGNOMY OF THE HYDROPHYTE COMMUNITY IN OTSEGO LAKE, N.Y. WILLARD N. HARMAN As part of a biological survey conducted in 1969, the species and distribution of aquatic macrophytes in Otsego Lake, Otsego Co., N.Y., were determined (Harman 1970). The 1969 information was compared with data collected by Muencher (1936) in 1935 (Harman & Doane 1970). Emer- gent plants were more abundant in 1969 than in 1935. The number of species present in the lake had decreased between 1935 and 1969 and an introduced species, Potamogeton erispus L., that was not present in 1935, was the most obvious plant in the lake in early summer of 1969. The reduction in species in this lake is similar, although not as pronounced, as changes observed over 50 years in Oneida Lake, N.Y. (Harman & Forney 1970) and over 70 years in Put-in-bay Harbor, Lake Erie, Ohio (Stuckey 1971). During the summer of 1969 it was noted that the zooben- thos and zoo-meroplankton populations underwent drastic fluctuations that correlated with the phenology of the aqua- tic macrophytes and the seasonal changes in the physiog- nomy of the plant community. A study was initiated to more accurately determine the composition of the flora at various times during the year so that we could quantita- tively ascertain its effects on the associated fauna. The purpose of this report is to describe the seasonal changes in 497 498 Rhodora [Vol. 76 the appearance of the littoral zone in Otsego Lake at Rat Cove, at the SUNY Oneonta Biological Field Station at Cooperstown, New York. METHODS In 1970, a transect about 500 m long was plotted from the end of our docks through the littoral zone, to the deeper water beyond. Plants growing along this line were meas- ured by divers each week, from 1 June through 2 Septem- ber, to determine the emergence time of the spring cohorts, greatest maximum heights attained, times of flowering, and times of death and decomposition of each species (Harman 1971). Because all growth represented even-aged stands, sufficient data were obtained by measuring the tallest in- dividual plants of each species each week. It was impossible to measure the same individuals throughout their existence because the act of measurement disrupted, if not destroyed these fragile organisms. Also, movements along the tran- sect disturbed the sediments so that by the end of Septem- ber we had to move farther and farther away from the original line in order to measure plants in an undisturbed environment. Secchi transparency and surface water tem- peratures in Rat Cove were recorded from 15 June through 25 August. In 1971 the same procedures were followed with these changes and additions. Samples were taken through the ice on 5 February. Weekly observations were initiated Shortly after the ice breakup (22 April) and continued until 16 December after all plants had completed their seasonal growth. 'The transect was marked with an an- chored hand line for divers to follow in order to alleviate the problems of getting lost in the often turbid water as had happened a few times in 1970. Secchi transparency and surface water temperatures in Rat Cove were recorded weekly from 10 May 71 until 3 January 72. Samples of benthic organisms were taken near the tran- sect at depths of 1-2 m, 3-5 m, and 6-8 m each week, to 1974] Otsego Lake — Harman 499 determine and correlate seasonal changes of the macro- benthos with those of the aquatic plants. Likewise, num- ber 20 plankton nets were carried by divers through the macrophytes occurring at 1-2 m, 3-5 m, and 6-8 m depths each week, to correlate the successional changes in the zoo- plankton community associated with them. The data result- ing from these observations will be reported in a manuscript now in preparation. Surface temperatures were essentially the same for both 1970 and 1971 (high = 24°C; mean for June, July, and August = 21.0 and 21.2°C, respectively). Mean Secchi transparency for June, July, and August was 4.4 m in 1970 and 4.3 in 1971. One appreciable difference in transparency was noted in June, the time that many of the plants were growing most actively. In 1970 the mean Secchi transpar- ency was 5.3 m, in 1971, 4.2 m (Harman 1972). PHENOLOGY AND PHYSIOGNOMY The zonation of hydrophytes according to water depth, ie. emergents, floating leaved varieties, and submergents (e.g., Sculthorpe 1967), is common knowledge. The com- ponents of the submergent macrophyte flora also occur in similar patterns, presumably because of the varying com- pensation points of each species relative to decreasing light intensity in deeper water, or to their growth in the shade of other species, and to the severe competition for space on eutrophic littoral substrates. In any one area along the Rat Cove transect, between 50 em and 550 cm in depth, several species of hydrophytes usually occur. This often results in a complex physiognomy exhibiting an overstory (usually of low density), a moder- ately dense intermediate stratum, and a low, very dense understory, although up to 7 distinct strata may be present. Vertical stratification of the community is continually changing during the growing season as the various species of macrophytes emerge from the substrate, attain maximum height and density, and then decompose. 500 Rhodora [Vol. 76 The shallowest waters along the transect are barren be- cause of the action of the waves on the shore. Nuphar variegatum Engelm., the yellow water lily, grows in scat- tered clones in water from 20 to 60 cm in depth in associa- tion with Elodea canadensis Michx. and Megalodonta Beckii (Torr.) Greene. Water from 50 em to 500 cm in depth maintains all the remaining macrophytes studied except for Nitella flexilis (L.) C. A. Agardh and Potamogeton crispus which occur in water from 500 to 700 em in depth. The growth of macrophytes in 1971 was similar to that observed in 1970 with the following exceptions: Potamoge- ton crispus, P. Richardsonii (Ar. Benn.) Rydb., and P. illinoensis Morong reached heights of approximately 300 cm in 1970, but attained less than 250 cm in 1971 before de- composition began. In 1969 and 1970 Potamogeton crispus reached the surface in some areas of the lake forming dense beds that hindered navigation, but did not reach the surface anywhere in 1971. After the spring cohort of the pond weeds mentioned died back in 1970, comparatively few scattered plants were observed during the rest of the sum- mer. In 1971, after the first cohort decomposed, many individuals resumed growth and died back sporadically until October. The following are descriptions of the phenology and phy- siognomy of the plants at 4 sites (0.5 m, 2.0 m, 4.0 m, and 5.5 m in depth) along the transect (Fig. 1). WATER DEPTH 0.5 M: Nuphar variegatum emerged from the substrate the first week in May; by 15 June, 1971 the leaves had reached the surface. They remained there until decomposed in early September. Elodea canadensis and Megalodonta Beckii started their seasonal growth in late April and the middle of May respectively. They maintained a maximum height of about 30 cm beneath the Nuphar during August, By 15 September M. Beckii had collapsed to the bottom and was decomposing. At the same time, E. canadensis started to decompose from the base and the entire plants became very brittle, By 15 November the 501 Otsego Lake — Harman 1974] "Ádouso əsuəp = :Ádouvo əsnyıp = - - — -'sejep pejose[es ye sejAudoupAu juepunqe jo sjudreQ Jurwous ‘YLE o3osj() 940) yey UI yydep ur ur G'Q uumn[OO I93€A gZ eun3rj LZ 19quie^oN GL LL 19quieides GL LZ 15nBny GL LL Aine GL LZ eunr SL ejuopoyebay eapo|3 e3uopojeBa|A "9 eapoj[3 ?epo|3 ?apo|3 9 e3uopojebejA| 1eudnwN seydny seydny ejensqng woi wo 'LL6T ‘GT Á[nf uo uorjejeZeA gueurwop Jo syry ejvorpur seul[ [fjuozHOH 'gG-G sein3g jo Suoreoo[ Burmoys j?esut1j 940) yey JO e[goid orjeunue?dSer[ ‘T ANB L4 ie:0niiqng —] = 9e Asqne —____ aoeyins L- - 31vHisg8ns WOHud SHu313W + L a10ys 502 Rhodora [Vol. 76 winter condition was reached. Megalodonta Beckii and E. canadensis were prone on the bottom with winter buds 1 to 2 em long developing at the nodes. During July and August, two obvious layers of vegetation were present, Nuphar variegatum at the water's surface and Elodea canadensis and Megalodonta Beckii below. From autumn into the winter there were no actively growing macrophytes at this site. Figure 2 illustrates these changes at five selected dates during the year. WATER DEPTH 2.0 M: This site was always dominated by dense stands of Chara vulgaris L. Only in June was Megalo- donta Beckii able to emerge above this species. Elodea canadensis occurred only slightly higher than the Chara in June, July, and August before dying in the fall. In late August and early September Najas flexilis (Willd.) Rostk. & Schmidt was present, but never grew higher than the Chara. During August and September there were two definite vegetational strata present. A diffuse overstory of Vallisneria americana Michx. and Myriophyllum exalbes- cens Fern. occurred above C. vulgaris. With the decompo- sition of V. americana and M. exalbescens in November, only Chara remained. The first week in May dense stands of Chara vulgaris were at their lowest height of the entire year. Chara grew steadily until the middle of November, when the plants started to decrease in size, presumably from basal decom- position, since the apices remained healthy all winter. Megalodonta Beckii and Elodea canadensis started growth at the same time as in the shallower water but attained somewhat greater heights. Megalodonta Beckii reached ap- proximately 35 cm in height in August, decomposing in Sep- tember. Elodea canadensis attained about 75 cm in height in early September and then began decomposing. Myrio- phyllum exalbescens appeared in early May, grew steadily until September and then fell to the bottom with new shoots appearing at the nodes. Vallisneria americana emerged from the substrate the middle of June; grew more rapidly 503 Otsego Lake — Harman 1974] ayeaysqng wos) wo 00L SCL osL SLL ooz "'Ádougo esuep = quepunge jo sjugreu Sutmoys 'exe'[ o3esj() ‘ea0p Fey ur ujdep ur ur (rg uum[oo 1976M '$ əng :Kdoueo esngrip = - — — 'sə}ep papəps ye sejAudoipáu LZ 1equia^oN GL LL 19quiejdeg GL LL 15nBny GL LL Ainp GL LZ eunf SL p ET e euopoebew| | | [seeN Marco eloh RESCUE NEMENES f | winyjAydouAy dips seleN eueusieA eapo|3 ç eapo|3 y eyg Errr anar E E NU gae sefeN L— — euopogBew| [t | esuopojebayl| DUPPEPTTPETTTRETIS ese 8 supose eu) winyAydouAW eeyo — se ee ee mee |) eapo[3 enaus je A me s — winyjAydouAW eey x S umpAydouAw | Ruin) AqdouAw — —À IFETTHTIETN E ——— o o o o -— m eIJ8USI] [EA 504 Rhodora [Vol. 76 than its competitors, attaining its maximum height of about 125 cm in September, and then rapidly decomposed. Najas flexilis emerged from the substrate in the middle of July, grew rapidly to about 60 cm high in early September and disappeared. Figure 3 shows the strata in 2.0 m of water at various times during 1971. WATER DEPTH 4.0 M: As at the 2 m site Chara vulgaris grew in dense beds at a consistent rate throughout the summer. Elodea canadensis was present until August as before. Early in May Potamogeton pusillus L., P. zosteri- formis Fern., P. Richardsonii, and P. illinoensis emerged from the substrate. By 15 July P. Richardsonii and P. illinoensis attained about 200 cm in height while P. pusillus and P. zosteriformis reached only about 100 cm heights. All of the species of Potamogeton were decomposing by August, although a second cohort of P. Richardsonii re- mained until early September. In late May, Heteranthera dubia (Jacq. MacM. emerged from the substrate and maintained steady growth until late September when it collapsed to the substrate. In June the morphology of the plant community was very complex with 7 species of actively growing hydrophytes all at different heights in the water column. By J uly 3 obvious Strata were present: a diffuse overstory composed of Pota- mogeton Richardsonii and P. illinoensis, a discontinuous stratum of intermediate height composed of P. pusillus and P. zosteriformis, and a dense understory of Chara, Elodea and Heteranthera. In September this same area was en- tirely different in appearance with 2 major strata present. There was a rather dense overstory of Heteranthera dubia. underlain by Chara vulgaris. By November the Chara was all that remained, Figure 4 illustrates the physiognomy at this depth at several selected dates. WATER DEPTH 5.5M: In these deeper waters Nitella flexilis replaced Chara vulgaris. Dense beds attaining 95 em occurred in this area. The only macrophyte asso- ciated with this alga was Potamogeton crispus, which ap- 505 Otsego Lake — Harman 1974] ajeusqng uio4j w3 "'Ádoueo əsuəp = :Adouvd osngtrp = - — — ‘sazep pəpəjəs e sayAydorpAy juepunge jo sju3reu 3urMous IYE 03ƏS7O ‘ea0g yey ut ujdep ur ur (yp uum[0o I939AA `$ 9In3lLq : LZ 18quie^O0N GL LZ 1equieideg GL LL 1snBny GL LZ Aine GL LZ eunf GL E: vi ADD & iuosp1euatH 'd eee "di PAIJETUJSU ?8po|3 snjjisnd ^q s d: S6 4 sisuaout ^d T ix ay x^ eyg ejaqyuejejeH| JfePOIl27o 0 0 os - eeu eueyd BS ae v^ S1WI404}143}S0Z "d J e9pol3 —y PAPPI3| fuuospueuoni “dw GZ - -— [epo 7 I1uospjeuotH "d, eyg 00L eJou1ue1839H snyisnd d ^ cy ——— | | "X "siünojaaisoz “g] GZL 4 Sisuaout| `d pdec ci qM oa eee | ccc cc cud dpt M IIUOSpJEU2IH ‘d em £J9Q1UPJ4919H SISUBOUIII! ^d. ~N ooz 4 [^ R^ muospieuol "d l IR | eus ed CN NN WoW 506 Rhodora [Vol. 76 peared from the substrate the first week in May. The latter species grew extremely rapidly, attaining about 250 cm in height by the middle of July. It then collapsed to the sub- strate and decomposed. In June and July two definite layers of vegetation were present at these depths. Potamogeton crispus formed a diffuse overstory while Nitella flexilis composed a dense understory. From August on into the winter N. flexilis was the only species present at the site (Fig. 5). The macrophytes studied can be separated into four groups according to their morphology during the winter season. These characterize the appearance of the substrate during this period and provide food and cover for the local benthos. Group 1. Chara vulgaris, Nitella flexilis. Once maturity is reached the oldest parts of these algae are con- tinually decomposing. During the winter period entire stands decrease in height as the rate of decomposition greatly exceeds the rate of growth. The apical meristems remain in a healthy condition throughout the year. In early spring much new growth is added from germinating oospores that cannot be distinguished from the older plants remaining during the same time periods. Group 2. Myrio- phyllum exalbescens, Elodea canadensis. When these or- ganisms decompose (late September and late August, re- spectively), the old growth lays on the substrate and winter buds, 2-3 cm in height, appear at once from the nodes. These grow extremely slowly over the winter reaching 4-6 cm by early May. At that time the roots have become estab- lished, internodal tissues from the parent plants have rotted away and the new plants grow rapidly until the fall when the process is repeated. Group 3. Heteranthera dubia. Like the organisms in Group 2, this plant collapses to the bottom in a living condition (in mid-October) but the new shoots do not appear at the nodes until late May or early June of the next year. They grow rapidly unti] fall and the cycle begins again. The parent stems remain intact throughout much of the growing season. Group 4. Potamogeton cris- pus, P. Richardsonii, P. illinoensis, P. zosteriformis, P. 507 Otsego Lake — Harman 1974] jugpunqe jo SJT Surmoys faye] o3esj() ‘eao0g FEY UI u3dep ur ur g'g uumn[oo 137e M ejensqns wos uio ‘Adouvs esuep = LL saquianon GL LZ 1equieydes GL :Ádoueo esnyip = - - — ‘sazep pejoe[es ye sazAydorpAy LL 1snBny GL LZ Aine GL 'G ONS LZ eunr SL 06 SL 71 ela}! N ella}! N ool 4 Sil = pem sndsi42 `d elə} N S S ella} N PION sndsio 'd sndsiJ2 ‘d NS = i 35 S Y 508 Rhodora [Vol. 76 pusillus, Vallisneria americana, Najas flexilis, Megalodonta Becki, Nuphar variegatum. This group contains all the remaining plants studied. These macrophytes grow rapidly from buried winter buds or underground stems or root- stocks and then decompose, shoots not appearing above the substrate until the next growing season. LITERATURE CITED HARMAN, W. N. 1970. Aquatic biology studies, pp. 15-16 + Ap- pendix pp. 1-142. In: Second annual report (1969-70) SUNY Oneonta Biological Field Station, SUNY, Oneonta (mimeo). 1971. Biological studies — Otsego Lake, pp. 1-28. Im: Third annual report (1971) SUNY Oneonta Biological Field Station, SUNY, Oneonta (mimeo). 1972. Aquatic biology studies, pp. 4-15. Im: Fourth annual report (1971) SUNY Oneonta Biological Field Station, SUNY, Oneonta (mimeo). , & T. R. DOANE. 1970. Changes in the aquatic flora of Otsego Lake, New York between 1935 and 1969. N. Y. Fish Game Jour. 17: 121-123. , & J. L. Forney. 1970. Changes in the molluscan com- munity in Oneida Lake, N.Y. between 1917 and 1967. Limn. Ocean. 15: 454-460. MUENSCHER, W. C. 1936. Aquatic vegetation of the Susquehanna and Delaware areas. pp. 205-221. In: A biological survey of the Delaware and Susquehanna watersheds. N. Y. S. Cons. Dept., Albany. SCHULTHORPE, C. D. 1967. The biology of aquatic vascular plants. 610 pp. St. Martin's Press, New York. Stuckey, R. L. 1971. Changes of vascular aquatic flowering plants during 70 years in Put-in-bay Harbor, Lake Erie, Ohio. Ohio Jour. Sci. 71: 321-342. BIOLOGY DEPARTMENT STATE UNIVERSITY COLLEGE ONEONTA, NEW YORK 13820 PANICUM ENSIFOLIUM BALDW. — NEW FOR MEXICO MELVIN L. CONRAD During the summer of 1972 a survey of grasses ( Conrad, et al., 1972) was conducted for the Food Agriculture Or- ganization of the United Nations at the Instituto Tecno- lógico y de Estudios Superiores de Monterrey which is developing a range management experiment station near Tempoal, Veracruz. On July 19th the group of investiga- tors accompanied the coordinators of the program, Dr. Donald Huss and Ing. Edmundo L. Aguirre, on an excur- sion south of Tempoal toward Molango, Hidalgo, to observe the cloud forest vegetation (Leopold, 1950) near the crest of the Sierra Madre Oriental. At a point 95.2 km. from Tempoal, before reaching Molango, the caravan of two automobiles stopped at a small parking area on the steep mountain slope to turn around due to heavy fog, rain, and an increasing number of fallen boulders on the highway. In the parking area I was observing a tree fern (Cyathea sp.) that grew beside the bluff from nearly 10 meters below; and after a moment, I realized that I was standing in a shallow wet depression about 5 meters in diameter that was filled with a Panicum different from several species that had been collected at lower elevations. Collections were taken. In the laboratory the grass from the parking area on the mountain was identified as Panicum ensifolium Baldw. Intermixed with it was P. cordovense Fourn., (Hitchcock and Chase, 1910). Other species associated with these were Juncus dudleyi Wieg., Carex festucacea Schkuhr, Agrostis perennans (Walt.) Tuckerm., and Sporobolus poirettii (Roem. & Schult.) Hitche. It was first hypothesized that this population of P. ensi- folium had been introduced, due to the considerable distance separating this station from its previously reported range 509 510 Rhodora [Vol. 76 in the southeastern United States, (Hitchcock, 1950). How- ever, it is likely to be native, since reports by Sharp (1946) of disjunction in other taxa indicate similar affinities be- tween the floras of these two regions. The assistance of Dr. A. A. Beetle at the University of Wyoming for verification of the identification, and the con- firmation by both Dr. Beetle and Dr. Arturo Gómez-Pompa at the Instituto de Biologia in Mexico City that P. ensi- folium is new for Mexico is hereby acknowledged with deep appreciation. Thanks are also due Dr. David B. Dunn for the use of the herbarium at the University of Missouri. Voucher specimens of P. ensifolium Baldw. (No. FAO Z-101) are deposited in herbaria at the Departamento Zootécnia, I.T.E.S.M., in Monterrey, México, Northeast Missouri State University, University of Missouri at Colum- bia (UMO), and The University of Wyoming (RM). LITERATURE CITED CoNRAD, M. L., W. K. MEDLER, S. NEFF, & C. PEPKING. 1972. 'Some Grasses (Gramineae), Sedges (Cyperaceae), and Rushes (Junca- ceae) of the Huasteca Zone in Veracruz and Hidalgo, Mexico. Instituto Technológico y de Estudios Superiores de Monterrey. Monterrey, México. Unpublished report. Hitcucock, A. S. 1950. Manual of the Grasses of the United States, U. S. Dept. Agric. Misc. Pub. No. 200. Revised by Agnes Chase. , & A. CHASE. 1910. The North American Species of Panicum. Contrib. U. S. Natl. Herb. 15: 1-396. LEOPOLD, A. S. 1950. Vegetation Zones in Mexico. Ecol. 31: 507- 518. SHARP, A. J. 1946. Informe preliminár sobre algunos estudios fitogeográficos efectuados en México y Guatemala. Rev. Soc. Mex. Hist. Nat. 7: 35-40. DIVISION OF SCIENCE NORTHEAST MISSOURI STATE UNIVERSITY KIRKSVILLE, MISSOURI 63501 SIGNIFICANT NEW DISTRIBUTIONAL RECORDS FOR THE GENUS SPHAGNUM IN THE NORTHEASTERN UNITED STATES RICHARD E. ANDRUS Recent collecting and herbarium studies reveal interest- ing new distributional records for several species of Sphag- num in the northeastern United States. Most of the records are from New York, and these are presented on the accom- panying maps. Collection numbers, unless otherwise indi- cated, are those of the author. Duplicates for most stations collected by the author are at NYS, and many are also at NY and SU-CF (=S.U.N.Y. College of Environmental Science and Forestry). Sphagnum henryense Warnst. A more difficult taxon than the literature would indicate, S. henryense has frequently been mistaken for S. palustre L., S. papillosum Lindb., or S. imbricatum Russow. Forms of S. henryense lacking the characteristic worm-like ridges on the hyaline cell walls where they overlie chlorophyll cells are especially difficult to identify. Such forms can usually be separated from the very similar S. palustre by the pore characteristics of the convex surface of the branch leaf. In S. henryense, the hyaline cells have numerous small round pores along the commissures, often 10 to 20 per cell, and usually lack large membrane gaps near the leaf apex. S. palustre, on the other hand, has hyaline cells with fewer and more elliptic pores as well as membrane gaps near the leaf apex. Macro- scopically, well developed forms of S. henryense may be recognized by their reddish-brown pigmentation (often visible in the capitulum) and a flattened capitulum with small pointed head branches. In contrast, S. palustre is browner and typically has a rounded capitulum with the head branches blunter and more obese. Sphagnum henryense is fairly widespread near the coast from Long Island, N.Y. (Ketchledge 1957) south to Florida 511 [Vol. 76 Rhodora 512 pulchrum S. A = S, henryense subfulvus O=s. torreyanua1 O-s. [0] = S, tenellum * D jensenii = 8. A-s. strictun 1974] Sphagnum — Andrus 513 and westward to Texas. The Massachusetts station reported here represents a northward extension of this coastal range. The inland stations extending into western New York and Pennsylvania are more surprising, however, but can be considered, perhaps, as analogous to similar vascular plant distributions noted by Fernald (1937). On this basis, one might expect to find Sphagnum henryense in other Great Lakes states such as Ohio and Michigan as well as southern Ontario. In comparing Sphagnum henryense and S. palustre from inland stations, one particularly interesting pattern emerges. Where it is found, S. henryense is normally an obvious and abundant member of its plant community. S. palustre, on the other hand, although found at many more inland locations, typically occurs as a minor vege- tational element, usually as a few scattered hummocks. Limited observations on Long Island indicate the same pattern may also hold in coastal areas. The factors re- sponsible for this difference are unknown. New York: ERIE C0.: Collins, Glowny 10850, 10848 (NYS). GENESEE co.: Bergen Swamp, Durand 3642, 3624 (cu), Winne 1590 (cu), S. J. Smith 47742, 47982 (NYS). CHEMUNG CO.: Winter Bog, 1594, 2168, 2170, Winne 2094, 2083 (NYS). ONONDAGA CO.: Cicero Swamp, 2852. SCHENECTADY CO.: Featherstonhaugh Lake, 2142, 2146. GREENE co.: Grapeville, S. J. Smith 48750 (NYS). ORANGE CO.: Sterling Forest, G. L. Smith 2666 (NY). CATTARAUGUS CO.: St. Bonaventure Campus, Boehner, Aug. 1, 1940 (St. Bonaventure Coll.. Massachu- setts: SUFFOLK CO.: Milton, Eaton & Faxon, Sphagna Boreali-Ameri- cana #159. Pennsylvania: MCKEAN CO.: Smethport, Glowny 12007. Sphagnum portoricense Sull. The previously known northern limit for S. portoricense was on Long Island, N.Y. (Mapes 1962). The northern range extreme is now extended to Mt. Desert Island, Maine, where a collection was made at a site strikingly different from those at the stations formerly thought to be the most northerly. On Long Island and in New Jersey (Andrews 1912) S. por- toricense is aquatic in nature, being confined to shallow sandy-bottomed ponds. On Mt. Desert Island it was clearly 514 Rhodora [Vol. 76 non-aquatic, with the plants being found at the margin of Big Heath, a large open bog, where they formed a carpet well above the water table along a small water course in the shade of black spruce. One possible explanation may lie in the high humidity characteristic of Big Heath, which is situated very near the coast and often shrouded in fog. S. portoricense also forms non-aquatic carpets in the very humid environment of the high-elevation cloud forests of- Puerto Rico. Maine: Mt. Desert Island, Big Heath, 2030. Sphagnum strictum Sull. The discovery of S. strictum in New York was not unexpected because of its occurrence northward and southward along the Atlantic coast (Maass 1966). The record from the Shawangunk Mountains of upstate New York is, however, unusual in view of the close affinity for the coastline that S. strictum exhibits in the northern part of its range (Maass 1966). Sphagnum strictum, like the closely related S. compactum DC., is a pioneer species. It is found on more organic and shaded substrates than the latter species. New York: ULSTER CO.: Lake Minnewaska, S. J. Smith 48953 (NYS). SUFFOLK CO.: Riverhead, Third Pond, Latham 34419 (cu, NYS); Sandy Pond, 2251, 2205, 2193, Mapes 1435 (Nvs) ; Swan Pond, S. J. Smith 48158 (Nvs); Manorville, Latham 33936 (NYS, CU). Sphagnum torreyanum Sull. (S. cuspidatum var. torreyi (Sull.) Braithw.). Sphagnum torreyanum is reported here for the first time from upstate New York. Like S. henry- ense, it was previously known only from coastal locations, in this case from Newfoundland to North Carolina (An- drews 1938). In upstate New York, as on Long Island, it occurs as an aquatic in poor fens, often intermixed with S. cuspidatum Hoffm. New York: osWEGO CO.: Kibbe Lake, 1521, 1610. SARATOGA CO.: Mulleyville Pond, S. J. Smith 46881, 46882 (Nvs); Luther's Woods, 2163. 1974] Sphagnum — Andrus 515 Sphagnum riparium Angstr. The range of S. riparium, previously reported southward to New Hampshire, Michi- gan, Wisconsin, and Washington (Andrus & Layser 1971), is now further extended into New York. A robust northern species, it typically grows in moderately minerotrophic sites near the margins of poor fens associated with such Sphagnum species as S. angustifolium (Russow) C. Jens., S. fallax (Klinggr.) Klinggr. and S. fimbriatum Wils. Present collecting data indicates that S. riparium is a very minor element in New York mire vegetation. New York: ESSEX co.: Raybrook Bog, 1661; Peninsula Nature Trails, 2152. FRANKLIN CO., Upper St. Regis Lake, 1707; 0.6 mi. E. of Corey’s, 2816. HAMILTON CO.: 4 mi. S. of Long Lake, 1969; Johnny Mack Bk. fen, 2846. oSWEGO co.: Kibbe Lake, 1528, 1529. CORTLAND co.: Little York Lake, 2795. MADISON CO.: Nelson Swamp, 2778. SCHENECTADY CO.: Featherstonhaugh Lake 2134. RENSSELAER CO.: Round Pond outlet, 1884; !4 mi. NE. cf Round Lake, 1914. Sphagnum pulchrum (Braithw.) Warnst. A subocean- ically distributed species, in eastern North America S. pul- chrum is known from the arctic (Persson & Sjórs 1960) south along the coast to New Jersey (Andrews 1938) and inland around the Great Lakes to Michigan (Crum & Miller 1969) and Wisconsin (Andrews 1938). The only previous New York report (Ketchledge 1957) is from Big Moose Station, Herkimer Co. (Peck £45, NYS). The numerous New York records are not unexpected and fill in the known distributional pattern. Sphagnum pulchrum is a characteristic poor fen species and occurs in both mire wide and mire edge vegetation, often in abundance. Typical associated Sphagnum species include S. papillosum, S. majus, S. fallax, S. angustifolium and S. rubellum. New York: CLINTON CO.: Cannon Flatrock, 1997a. FRANKLIN CO.: Upper St. Regis Lake, 1808, 1811, 1346, 1725. ST. LAWRENCE CO.: Sevey Pond, 1978; Wanakena, 1481; Grass River Flow, 1990; 1 mi. E. of Sevey Corners, 1967, 1968. ESSEX CO.: Algonquin Peak, 1245, 1246, 1247, 1248, 1249; Raybrook Bog, 1283, 1288, 1671, 1282, 1821; Lower Cherry Patch Pond, 1389; Whiteface Mt., 1215; Lake Tear, 516 Rhodora [Vol. 76 1333; between Scott and Wallface Ponds, 1351; near Scott Pond, 2425; Elk Lake, Phelps 100 (NYS). HERKIMER CO.: Big Moose Sta., Peck 45 (NYS). RENSSELAER CO.: Sand Lake, Peck 43 (CU, NYS); 1.5 mi. E. of Taborton, 2175, 2176, 2177, 2178; Bucks Corners, 1922. SUFFOLK CO.: Napeague Beach, 2357; 1 mi. E. of Manorville, 2867; Cedar Pond, N. G. Miller 6865 (NCH). Sphagnum jensenii H. Lindb. (S. annulatum var. poro- sum (Schlieph. & Warnst.) Maass & Isov.). A far northern species, S. jensenii is noted for the first time for the U.S. east of Michigan (Maass 1967b). The collection site was at the margin of a large poor fen where it was growing in a shallow pool in the bog mat associated with S. pulchrum, S. papillosum, S. subsecundum Nees, S. flexuosum Dozy & Molk and S. rubellum Wils. Maass (1967b) gives an excel- lent account of the separation of S. jensenii from closely allied species. New York: ST. LAWRENCE CO.: Sevey Pond, 2829, 2832, 2833. Sphagnum tenellum (Brid.) Brid. Sphagnum tenellum was collected among sand dunes in a small bog pocket where S. pulchrum was also found. As in the case of S. strictum, this report fills a coastal distribution gap between Maine and New Jersey. Wet hollows in ombrotrophic mires are the most frequent habitat of Sphagnum tenellum. Although such sites are found elsewhere on Long Island, they are devoid of the species. Lack of suitable habitat, therefore, is an unlikely explanation for its rarity on Long Island. Climatic factors are probably responsible, since on Long Island S. tenellum is very near the southern extreme of its eastern North American coastal distribution — this extreme being in New Jersey. New York: SUFFOLK CO.: Napeague Beach, 2238. Sphagnum subfulvum Sjórs. The single previous con- terminous U.S. report for S. subfulvum is from Michigan (Crum & Miller 1969). It undoubtedly reaches its southern 1974] Sphagnum — Andrus 517 range limit in New York. The only other published records for North America (Maass 1967a) give a range of “Green- land to Bay of Fundy and N. Ontario and from Alaska into western Yukon." The collection sites were medium to rich fens, uncommon mire types in New York, and this may account for its ex- treme rarity in the state. A plant that typically forms low to medium hummocks, Sphagnum subfulvum was found associated with S. warnstorfii Russow, S. contortum K. F. Schultz, S. centrale C. Jens., Scorpidium scorpioides (Hedw.) Limpr., Calliergonella cuspidata (Hedw.) Loeske and Cam- pylium stellatum (Hedw.) C. Jens. — all common rich mire species. Sjórs' (1944) original species description contains an excellent discussion on the distinction of Sphagnum sub- fulvum from the similar S. subnitens Russow & Warnst. and S. flavicomans (Card.) Warnst. New York: ESSEX CO.: Lost Pond, 1475. WARREN CO0.: Rush Pond, 1860, 1865; Glen Lake, 1854, 1885, 1889, 1886, 1890; Jenck's Swamp, 2815. BIBLIOGRAPH Y ANDREWS, A. L. 1912. Notes on North American Sphagnum, II. The Subgenus Inophloea Russow. Bryologist 15: 1-9. 1913. Sphagnaceae. N. Am. Fl. 15: 1-31. 1938. The North American Atlantic species of Sphag- mum. Ann. Bryol. 11: 15-20. ANDRUS, R. E. and E. F. LAYSER. 1971. Sphagnum riparium Angstr., a new record for the western United States. Bryologist 74: 211. Crum, H. and N. G. MILLER. 1969. Bryophytes new to Michigan. II. Michigan Bot. 8: 28-29. FERNALD, M. L. 1937. Local plants of the inner Coastal Plain of southeastern Virginia. Rhodora 34: 465-491. IsovirTA, P. 1966. Studies on Sphagnum L. I. Nomenclatural re- vision of the European taxa. Ann. Bot. Fenn. 3: 199-264. KETCHLEDGE, E. H. 1957. Checklist of the mosses of New York State. N. Y. S. Mus. Bull. 363: 55 pp. Albany, N. Y. Maass, W. S. G. 1966. Untersuchungen über die Taxonomie und 518 Rhodora [Vol. 76 Verbreitung von Sphagnum VI. Sphagnum pylaesii Brid. und das boreo-atlantische Florenelement unter den Torfmoosen in Südamerika. Nova Hedw. 12: 81-105 + 5 pls. 1967a. Studies on the taxonomy and distribution of Sphagnum II. Sphagnum angermanicum Melin in North Ameri- ca and its relation to allied species. Nova Hedw. 13: 449-465 + pls. 116-128. 1967b. Studies on the taxonomy and distribution of Sphagnum IV. Sphagnum majus, Sphagnum annulatum, Sphag- num mendocimum and Sphagnum obtusum in North America. Nova Hedw. 14: 187-214 + pls. 69-81. Mares, H. M. 1962. Sphagnum portoricense and Discelium nudum in New York. Bryologist. 65: 68. PERSSON, H., and H. SJörs. 1960. Some bryophytes from the Hud- son Bay Lowland of Ontario. Sv. Bot. Tidskr. 54: 247-268. SJORs, H. 1944. Sphagnum subfulvum Sjórs and its relations to S. flavicomans (Card.) Warnst. and S. plumulosum Warnst. P.P. Sv. Bot. Tidskr. 38: 403-427 + pls. 7, 8. DEPARTMENT OF FOREST BOTANY AND PATHOLOGY STATE UNIVERSITY OF NEW YORK COLLEGE OF ENVIRONMENTAL SCIENCE AND FORESTRY SYRACUSE, NEW YORK 13210 ENVIRONMENTAL CONTROL OF NEEDLE CHARACTERISTICS IN SUBALPINE BLACK SPRUCE BRIAN F. CHABOT'” Two distinct forms of Picea mariana, one upright and the other more or less prostrate, exist in the vicinity of treeline on several New England mountains. Characters distinguishing the two types were discussed by Teeri (1969) in establishing the varietal status of Picea mariana (Mill) BSP. var. semiprostrata (Peck) Teeri. Individuals of this taxon are recognized by their prostrate growth form, lack of a dominant upright axis, and short, glaucous needles that are slightly adaxially curved. The most prom- inent character other than the growth form is the needle length. Needles average 4 mm. (2.2 to 5.5 mm.) in length in the prostrate form as compared with upright individuals that have needles averaging 7.5 to 8.5 mm. The study reported here indicates that needle characteristics and, perhaps, other features of the prostrate variety are subject to eonsiderable modification by the environment. Plants were collected during late August and September 1972 from Lion's Head (1450 m.) and Cape Horn (1280 m.) on Mt. Washington, N.H., and in Bear Swamp, Wolcott, Vt. (340 m.). Both upright and prostrate forms were collected at the treeline sites on Mt. Washington. Only the typical form was obtained from Wolcott. Individual plants from all populations were transferred to a commercial potting mixture (Jiffy Mix) and grown in a cool greenhouse (10 to 20°C). All individuals were watered daily and fertilized weekly with a 20-20-20 nutrient solution (Universal Chem- 1Present address: Section of Ecology and Systematics, Langmuir Laboratory, Cornell University, Ithaca, New York 14850. *Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution Number 685. 519 520 Rhodora [Vol. 76 ical Co.). Buds expanded during April, and both the new needles and the previous year's needles produced under natural conditions were analyzed in early July. Fifty needles were selected from each indicated age class on three separate individuals from each of the populations studied. The samples were pooled according to age class and source population. A final sample of 100 needles was taken from each pooled sample for actual measurement. Length measurements were taken to the nearest 0.1 mm. Photographs of sample individuals were also taken. New growth needles of the short needle var. semipro- strata contrasted dramatically in length compared with growth which had occurred under natural conditions (Fig. 1). The new growth needles were not only longer by about 5 mm. (Table 1), but they were darker green in color and had no pronounced curvature. New needles of the other populations of Picea mariana also had a healthier appear- ance than the naturally produced needles, but, otherwise, differences in needle length were not at all pronounced (Fig. 1, Table 1). Average needle lengths along with standard deviations are presented in Table 1. Tests for significance were ap- plied using the analysis of variance technique. The natural needle length (short form) of var. semiprostrata was sig- nificantly different (5% level) from the other sample lengths including the needles produced in this variety under cultivation. The needle lengths resulting from culti- vation of all three populations were not significantly dif- ferent. Table 1. Average needle length and standard deviation for three populations of Picea mariana. Source Elevation (m.) Needle Length (mm.) Natural Cultivated A) 1450 (var. semiprostrata) 4.5 + 0.54 9.4 + 1.2 B) 1450 and 1280 (var. mariana) 8.5 + 0.97 8.1 + 1.2 C) 340 (var. mariana) 6.4 + 0.73 8.5 + 1.8 1974] Black Spruce — Chabot 521 Figure 1. Branches of Picea mariana with needles produced naturally (below arrow) and under cultivation (above arrow). Sources of the original populations are: A) 1450 m., Mt. Wash- ington, var. semiprostrata; B) 1450 m., Mt. Washington, long-needle form; and C) 340 m., Bear Swamp. The results indicate that needle length in var. semipro- strata is under some degree of environmental control. Needle curvature is also capable of being modified. Curva- ture appears to be related to needle length in that the Shorter needles (less than 5 mm.) of all populations had varying degrees of adaxial bending, while the longest needles were most frequently straight. Modified needle growth in black spruce under cultivation has previously been observed, but not fully reported (Kozlowski, 1971; 522 Rhodora [Vol. 76 R. Pike, pers. comm.). The precise environmental factors which operate here have not been isolated, but they cer- tainly include both the improved nutrient and thermal regimes of the greenhouse. The nutrient regime may be particularly important in that the bog population, from an environment which has been characterized as having low nutrient availability (Small, 1972), also tended to have shorter needles. Additional information regarding the genetic structure of var. semiprostrata can be obtained from observations concerning the prostrate form itself and the fact that both short- and long-needle types exist in close proximity to each other at treeline (Teeri, 1969). The latter observation may be taken as presumptive evidence of genetic distinctiveness assuming that the forms are not actually distributed in different microenvironments. The environmental situation must be studied carefully as personal observation and the studies of Tiffney (1972) indicate that slight differences in microtopography, slope, and degree of exposure can make considerable difference in the success of individual plants at treeline. The genetic nature of the prostrate growth habit is dif- fieult to demonstrate experimentally because of the long period of time necessary to develop this form. There is a tendency for new twigs on the uppermost branches to occur vertically. Frequently the preceding year's needles on these protruding branches have been removed on the windward side. Thus we may be witnessing an example of wind trim- ming as has been observed in other climates (Thomas, 1973). Additionally, the work of Jaffe (1973) suggests that the stunting of individuals could be an aspect of thigmo- morphogenesis resulting from the mechanical action of wind. The contiguous natural distribution of characteristic forms of both varieties as well as character variation which is essentially clinal in nature (Teeri, 1968) perhaps has resulted from disruptive selection similar to processes described by Antonovics and Bradshaw (1970) and Snay- 1974] Black Spruce — Chabot 523 don (1970). Phenotypic variation which has little im- portance in other parts of the range of Picea mariana may assume considerable significance at alpine treeline. Several morphological features of var. semiprostrata make it par- ticularly well adapted to the alpine environment (Teeri, 1969). Additionally, it has frequently been observed that growth rates, such as needle elongation, and cold-hardiness are inversely related (Alden and Hermann, 1971). Thus, at the distributional limit for a species, a uniquely adapted genotype could exist sympatrically with the more common genotype even in a homogeneous environment. Some degree of reproductive isolation exists between the varieties due to the infrequency of cone production at treeline (Teeri, 1968). It is not the intention of this study to reorganize the taxonomy of black spruce. The variety semiprostrata as defined from natural specimens is sufficiently unique to merit distinction. However, it should be recognized that the genetic basis for this taxon lies not in a narrowly specified phenotype, but in its plasticity and the sensitivity of the genotype to interaction with the environment. ACKNOWLEDGMENT Appreciation is expressed to the Superintendent, White Mountain National Forest, and Peter Marchand for their assistance in certain phases of this study. LITERATURE CITED ALDEN, J., & R. K. HERMANN. 1971. Aspects of the cold-hardiness mechanism in plants. Bot. Rev. 37: 37-142. ANTONOVICS, J., & A. D. BRADSHAW. 1970. Evolution in closely adjacent plant populations. VIII. Clinal patterns at a mine boundary. Heredity 25: 349-362. JAFFE, M. J. 1973. Thigmomorphogenesis: the response of plant growth and development to mechanical stimulation. Planta 114: 143-157. KozLowsKr, T. T. 1971. Growth and development of trees. Vol. 2. Academic Press, N.Y. 333 pp. 524 Rhodora [Vol. 76 SMALL, E. 1972. Ecological significance of four critical elements in plants of raised sphagnum peat bogs. Ecology 53: 498-503. SNAYDON, R. W. 1970. Rapid population differentiation in a mosaic environment, I. Response of Amthoxanthum odoratum popula- tions to soils, Evolution 24: 257-269. TEERI, J. A. 1968. The ecology of subalpine black spruce in New England. M.S. Thesis, Univ. of New Hampshire. 62 pp. 1969. The phytogeography of subalpine black spruce in New England. Rhodora 71: 1-6. Tuomas, T. M. 1973. Tree deformation by wind in Wales. Weather 28: 46-58. TIFFNEY, W. N., JR. 1972. Snow cover relationships of White Mountain alpine plants. Ph.D. Thesis, Univ. of New Hampshire. 75 pp. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 LINDERA MELISSAEFOLIUM IN ARKANSAS. An excellent description of Lindera melissaefolium (Walter) Blume (Lauraceae) and a detailed account of its known distribution in 1949 were given by Steyermark (Rhodora 51:153-162). He suggested that the shrub might some day be found in Arkansas because of its occurrence in adjacent Ripley County, Missouri. The discovery of an Arkansas locality in 1972 is not unexpected, therefore, but is here recorded in view of the extreme rarity of the species. Several colonies representing hundreds of plants are now known from the northern part of Clay County, Arkansas. The area is a part of the Mississippi Embayment region and is characterized locally by a series of low woods in which swampy depressions are surrounded by sandy knolls. The shrub occurs in the water of these depressions, where it forms dense stands which average 2-4 feet in height. Some of the colonies occurring at the woods margins grow in close association with the rare and interesting Corkwood, Leitneria floridana Chapman. Specific collection data for specimens from the Arkansas locality are as follows: CLAY COUNTY: Low woods having depressions with standing water surrounded by sandy knolls, Sect. 2, T21N, R4E, 10 June 1972 (with R. Davis), Tucker 9642 (UARK, NCU, APC). G. E. TUCKER DEPARTMENT OF BIOLOGICAL SCIENCES ARKANSAS POLYTECHNIC COLLEGE RUSSELLVILLE, ARKANSAS 72801 SPIRANTHES CASEI, A NEW SPECIES FROM NORTHEASTERN NORTH AMERICA PAUL M. CATLING AND JAMES E. CRUISE In 1923 Henry Mousley made a collection of Spiranthes orchids from the vicinity of Hatley (45° 11' N., 71° 56’ W.), Stanstead Co., Quebec, He reported these plants in 1924, under the name of Spiranthes cernua (L.) Rich. var. ochro- leuca (Rydberg) Ames (Can. Field-Nat. 38: 61-63, 86-88, p. 88). About 17 years later he sent these collections to Dr. D. S. Correll at Harvard University, who was then engaged in a study of the genus. Correll reported back to Mousley that the plants from Hatley represented a mixed collection of S. cernua (L.) Rich. var. odorata (Nutt.) Correll and S. vernalis Engelm. & Gray. This was reported by Mousley in 1941 (Can. Field-Nat. 55: 19-80, pl. 1-2), and by Correll in 1950 (p. 227). Another station of S. vernalis was discovered by Mousley during 1941 near Ste. Dorothée (45° 32’ N., 73° 39' W.), Quebec, and described in some detail by him in 1942 (Can. Field-Nat. 56: 1-2, pl. 1-2). Gleason and Cronquist (1963) reported south- eastern Quebec to be within the range of S. vernalis, a decision probably based on the report of Correll (1950). In recent floristic work concerning eastern Canada (e.g. Marie- Victorin, 1964; Roland & Smith, 1963-64; Boivin, 1967) no further mention has been made of S. vernalis, and surpris- ingly the reports of Mousley and Correll have not been repeated. However, Case ( 1964) discussed the occurrence of plants referable to S. vernalis in the western Great Lakes region, and Voss (1972, p. 461) has mapped the distribution of S. vernalis in Michigan, although both of these authors assigned their plants to S. vernalis with some hesitation. Over the past three years a species of Spiranthes that compared favorably with Mousley's specimens from Quebec was found to be frequent and locally abundant in some parts of the Canadian Shield region of southern Ontario, and it 526 1974] Spiranthes — Catling & Cruise 521 was listed for the province as S. vernalis by Whiting and Catling (1971). A survey of herbarium collections in east- ern North America revealed many new localities for this «northern S. vernalis" in Ontario, several new stations in Quebec, Michigan, and New England, and two localities in Nova Scotia where it was previously unknown. As specimens were examined over this wide range, it be- came clear that the northern plants were quite distinct and easily separated from Spiranthes vernalis as it occurs in the southeastern states. A series of specimens from the Coastal Plain and the gulf states, including the type of S. vernalis Engelm. and Gray (AMES 82967), differed consistently from the northern plants in exhibiting a light-colored and fre- quently more dense, non-glandular pubescence on the rachis, and a denser, less robust spike with a larger number of narrower and slightly longer flowers. Plants collected from Massachusetts, including the types of Spiranthes X inter- media Ames (AMES 2246) and Spiranthes neglecta Ames (AMES 2518) and the justifying specimens for Spiranthes cernua X gracilis Ames (AMES 17 391), are comparable with the “southern S. vernalis.” In his enumeration of North American orchids, Ames (1924) included S. X intermedia, S. neglecta and S. cernua X gracilis in synonomy with his x S. vernalis. A general northern limit for the *southern vernalis" is approximated by a line joining Massachusetts, Kentucky, and northern Missouri. The northern plants re- ferred to vernalis are distributed in a broad band from Michigan to northern New England and the Canadian maritime provinces. As these northern plants are quite distinct and different from the southern plants (including the type of S. vernalis), and as they appear to have a dis- crete distribution pattern we are herewith describing them as a new species. Spiranthes casei Catling et Cruise sp. nov. Differt a S. vernalis pubescentia septata et rubri-glandu- losa in inflorescentia, spica robustiore et minus densa, flori- bus paucioribus, latioribus, brevioribus. 528 Rhodora [Vol. & Splranthes vernalis Engeln, and Gray rooms Ont = = MN a fes * Simcoe "ese Moist adge cf aspen woods, a Lot 4, eone, X, l k Innlafil tpe ma 30 Aug. 1370 FX" T, Razniosk 1470 Soul, bel 410 sm T. Reantook - s Thera pee" a ‘come ot s 40 of thes^ overlockei orchids in this area. Figure 1. A typical herbarium specimen of Spiranthes casei. The elongate spike with flowers loosely arranged in a single spiral, the erect oblanceolate leaves, and the robust nature of the plant are characteristic. (T. Reznicek 470, Simcoe County Museum). i Figure 2. Part cf a spike of Spiranthes casei from Dorset, Ontario. (Photograph by Dr. Erich Haber). 530 Rhodora [Vol. 76 Plantae (13-) 27-33 (-43) cm. altae sub anthesi. Folia omnia laevia, saepe glauca, usque ad tertiam caulis partem assurgentia; infima ad anthesin non permanentia, ovato- lanceolata, 1-2 cm. lata, 7 cm. longa; superiora ad et non- numquam per anthesin permanentia, oblanceolata vel lin- eari-lanceolata, usque ad 15 vel 20 cm. longa, plerumque minus quam 1 cm. lata, brevia, sub inflorescentia ad vaginas sine laminis reducta. Caulis basaliter glaber, in rachidi pilis septatis rubri-glandulosis 0.10-0.30 mm. longis pubescens. Spica plerumque 6-10 cm. interdum ad 15 cm. longa, floribus laxe et saepe in spiram unicam dispositis. Bracteae florales ovatae vel ovato-lanceolatae, longi-acuminatae, 7-12 mm. longae, basaliter plus minusve pubescentes. Flores cremei. Ovarium sub anthesi 4-6 mm. longum, supra oblique tumi- dum. Pubescentia in ovarii, sepali dorsalis, sepalorum lat- eralium superficie pubescentiae rachidis similis. Sepala lateralia 5-7 mm. longa; sepalum petalaque dorsalia sursum vix curvata, 5-7 mm. longa; petala dorsalia pagina externa papillosa. Labellum florum inferiorum 6-6.5 mm., superi- orum 5-6 mm. longum; calli basales crassi, 0.8-1.0 mm. longi, incurvi; labelli inferior superiorque centralis pagina et callorum regio brevi-pubescens vel papillosa. Different from S. vernalis in its septate, reddish-glandu- lar pubescence in the inflorescence, and its less dense, more robust spike with fewer, wider and shorter flowers. Plants (13-) 27-33 (-43) cm. tall at anthesis. Leaves glab- rous, often glaucous, ascending up to 1/3 the length of the stem; lowest leaves, not persisting unti] anthesis, ovate- lanceolate, 1-2 cm. wide, 7 cm. long; upper leaves, persisting until and sometimes throughout flowering, oblanceolate or linear-lanceolate, to 15 or 20 cm. long, and usually less than 1 cm. wide, reduced to bladeless sheaths below the inflores- cence. Stem glabrous basally, becoming pubescent in the rachis with septate, reddish-glandular hairs 0.10-0.80 mm. long. Spike usually 6-10 cm. long, occasionally to 15 cm. long, with the cream-colored flowers arranged loosely and often in a single spiral. Floral bracts ovate or ovate-lanceo- late, long-acuminate, 7-12 mm. long, more or less pubescent 1974] Spiranthes — Catling & Cruise 531 basally. Ovary 4-6 mm. long at anthesis, obliquely swollen on the upper side. Pubescence on the surface of the ovary, dorsal sepal, and lateral sepals, similar to that of the rachis. Lateral sepals 5-7 mm. long. Dorsal sepal and petals barely upcurved, 5-7 mm. long, the dorsal petals papillose on the outer surface. The lip 6-6.5 mm. long in the lower flowers, 5-6 mm. long in the upper flowers. Basal calli of the lip stout, 0.8-1.0 mm. long, incurved. The lower surface and central upper surface of the lip, and region of the calli short pubescent or papillose. TYPE: CANADA: Ontario: MUSKOKA DISTRICT: ca. 6 mi. w. of Bracebridge along highway 118 in Monck township, (45° 02’ N., 79° 29’ W.), on dry hillside in full sun, Catling & Whiting (TRT 169205). REPRESENTATIVE SPECIMENS CANADA: Nova Scotia: SHELBURNE C0.: Hope's Lot Barrens, Clyde River, 7 Sept. 1921, M. L. Fernald & B. Long (AMES 85418). Ontario: FRONTENAC CO.: Salmon Lake, Frontenac Park, Bedford Tp., 28 Aug. 1968, R. Hainault & I. MacDonald (QK 97607). SIMCOE co.: lot 4, cone. 10, Innisfill Tp., 30 Aug. 1970, T. A. Reznicek (Simcoe Co. Museum 470) Quebec: STANSTEAD CO.: Hatley, 11 Sept. 1924, Mousley (QFA 42012, AMES 63699). UNITED STATES: Michi- Zan: MARQUETTE CO.: summit of Huron Mountain, 1-9 Sept. 1916, C. K. Dodge (MicH). New Hampshire: coos Co.: Colebrook, 21 Aug. 1942, A. S. Pease 29802 (NEBC). Vermont: ORLEANS CO.: Sutton Rd., Willoughby, 13 Sept. 1899 (Mrs. Bruche) (NEBC). We dedicate this species to Mr. Frederick W. Case II, whose book on orchids of the western Great Lakes region has been enjoyed and admired by amateurs and profession- als alike. Mr. Case has indicated that the northern plants placed with Spiranthes vernalis differed from the southern plants, and he repeated Ames' suggestion of a hybrid origin for the northern plants (Spiranthes lacera X Spiranthes cernua) here described as S. casei. While such a hybrid origin is quite possible, several char- acteristics of this plant suggest that it is worthy of specific status. All plants in populations studied have been found to be fertile, and many maintain themselves in the absence of one or both putative parents. There is no evidence of back- 532 Rhodora [Vol. 76 cross-swamping of the distinctness of Spiranthes casei. Although S. casei is intermediate between S. cernua and S. lacera in many characters, certain other characters are hard to rationalize in a hybrid; for example, the flowers of S. casei are creamy-colored but white in both S. cernua and S. lacera. Spiranthes casei (Figs. 1 and 2) has been much confused with other species of Spiranthes growing within its range, particularly with S. cernua (L.) Rich. and S. lacera (Raf.) Raf., but also with S. romanzoffiana Cham. Spiranthes romanzoffiana, which also has creamy or pale yellowish-green flowers, is readily distinguished by the pandurate lip that is sharply constricted beyond the middle (distally), and also by the connivent nature of the sepals and petals which form the hood. Further, the flowers of S. romanzoffiana are usually larger (perianth 8-10 mm. long) and arranged in two or more rows in a compact spike. Where S. casei and S. romanzoffiana occur sympatrically, the latter reaches its peak of flowering in late July and early August, while the former is in full flower in late August and early September. Spiranthes lacera has a singly-spiralled elongate spike like that of S. casei, but it differs in being less robust, in its large, oval, widely spreading leaves in a basal rosette, and its smaller flowers (perianth usually ca. 4-5 mm. long) with green colouration in the lip. S. lacera reaches its peak of flowering in Ontario in late July and early August. We have noticed that some plants of S. lacera have gone to seed by late August when the first flowers of S. casei are open- ing. Spiranthes lacera frequently occupies the same habi- tats as S. casei, and both species grow in close proximity. Spiranthes cernua may be distinguished from S. casei by its shorter stature (20-30 cm.), usually short and compact Spike (ca. 5-7 cm. long), and its larger, pure white flowers (perianth usually 9-11 mm. long). In early September we have encountered S. casei growing on dry sandy roadside banks only a few feet removed from moist ditches with abundant plants of S. cernua. On several occasions the 1974] Spiranthes — Catling & Cruise 533 plants of S. casei were noted to be more advanced in flower- ing, with all of the flowers open and the lower flowers fading. The nearby plants of S. cernua were less advanced with the lower flowers fresh and many of the upper flowers still in bud. Although this may be a result of microclimate, it has nevertheless been a consistent observation. Spiranthes casei shares some characteristies with S. cer- nua (L.) Rich. var. ochroleuca (Rydb.) Ames, particularly the cream-coloured flowers and a preference for an upland habitat. The var. ochroleuca is stated to differ from the typical form principally in yellowish-white flower colour, longer floral bracts, monoembryonic seeds, and preference for an upland habitat. Ames (1921, p. 78) has indicated that the only reliable means of distinguishing var. ochro- leuca is through microscopic examination of the seeds. Apparently in other respects it is very close to typical S. cernua, as is obvious from the plate provided by Ames (loc. cit. pl. 127, Figs. 12-13), which was subsequently used in “The Orchids of North America” (Correll, 1950, pl. 70, 12-13). Clearly, this drawing does not illustrate S. casei, and the distinctive features of each make the separation of S. casei from either variety of S. cernua relatively simple. DISTRIBUTION AND ECOLOGY The range of Spiranthes casei extends from Michigan through Ontario and southern Quebec to the New England states and Canadian maritime provinces (Fig. 3). It is fairly frequent in the Muskoka-Haliburton region of On- tario, where there are 34 stations (and probably many more) separated from one another by distances of at least one mile. Here it is usually found in dry, open locations growing in coarse sandy soil in areas that have been dis- turbed but are not continually disturbed. Associated plants in such habitats include Danthonia spicata (L.) Beauv., Pteridium aquilinum (L.) Kuhn., and Polytrichum spp. The barren tops of ridges (Fig. 4) are the only naturally occurring habitats in which this orchid has been frequently encountered. 534 Rhodora [Vol. 76 D: = 4 | | J Ex ^] ( f JN o 700 300 ` Pa k MEES m— — —Á M Figure 3. Distribution of Spiranthes casei based on specimens examined in the following herbaria: ACAD, AMES, CAN, DAO, HAM, LKHD, MICH, MTMG, NBM, NEBC, NFLD, OAC, QFA, QK, TRT, UWO. Figure 4. Barren ridge-top on the Precambrian Shield in Matche- dash Township, Simcoe County, Ontario. Spiranthes casei was quite frequent on this ridge, growing in coarse shallow soil in open loca- tions. 1974] Spiranthes — Catling & Cruise 535 The earliest collection of Spiranthes casei from Ontario was made in 1904. Prior to 1965 it had been collected only eight times in the province. However, during the period from 1965-1972 more than 25 specimens have been added to local herbaria. It is true that some of this represents our own field work, but we cannot overlook the possibility that this orchid has become more abundant in Ontario due to human activity, such as land-clearing and road-building. Of 14 collections made since 1970, 10 were from formerly disturbed habitats such as roadsides, sand pits, abandoned fields, and hydro-line transects. Spiranthes casei becomes apparent in these sites several years after the initial dis- turbance. As with many other plants that quickly colonize habitats created by human activity, Spiranthes casei has proved to be at least partially apomictic. Seeds obtained from flowers, from which pollinators were excluded with cloth bags dem- onstrated 10-2596 germination. ACKNOWLEDGEMENTS The authors wish to thank Dr. B. Boivin (Central Ex- perimental Farm, Ottawa) for his helpful criticism of this manuscript and for suggesting the new name. Dr. L. A. Garay (Harvard University) was of considerable help dur- ing a visit to examine types in the Oakes Ames Orchid Herbarium, and also kindly criticized our work, Miss S. M. McKay, Mr. T. A. Reznicek and Mr. R. E. Whiting assisted with field work. We are also indebted to the herbarium curators who made specimens available for study: Dr. J. F. Alex; Dr. D. S. Christie; Dr. L. Cinq-Mars; Mr. W. J. Cody; Mr. C. E. Garton; Dr. A. E. Garwood; Dr. J. M. Gillet; Dr. O. A. Olson; Dr. J. B. Phipps; Dr. J. S. Pringle; Mr. T. A. Reznicek; Dr. E. A. Shaw; Mr. D. Strickland; Dr. S. Van der Kloet; Dr. E. G. Voss; and Dr. D. W. Woodland. Financial assistance has been provided by the National Research Council of Canada through grant no. A-2363. 536 Rhodora [Vol. 76 REFERENCES AMES, O. 1903. Natural Hybrids in Spiranthes and Habenaria. Rhodora 5: 261-264, pl. 47. 1904. Spiranthes neglecta. Rhodora 6: 27-81, pl. 51. 1905. Contributions toward a monograph of the Ameri- can species of Spiranthes. Orchidaceae, fasc. 1: 113-156. . 1921. Notes on New England Orchids, — I. Spiranthes. Rhodora 23: 3-85, pls. 127-129. 1924. An enumeration of the orchids of the United States and Canada. Am. Orchid Soc., Boston, 120 pp. BoiVIN, B. 1967. Enumeration des plantes du Canada. Naturaliste Canadien 94: 131-157. CasE, F. W. 1964. Orchids of the Western Great Lakes region. Cranbrook Inst. Sci. 147 pp. CORRELL, D. S. 1950. Native orchids of North America north of Mexico. Chronica Botanica Co., Waltham, Mass. 399 pp. GLEASON, H. A., & A. CRONQUIST. 1967. A manual of vascular plants of the northeastern United States and adjacent Canada. Van Nostrand Co., Princeton, New Jersey. 810 pp. MARIE-VICTORIN, FRERE. 1964. Flore Laurentienne. Les presses de l'universitie de Montreal, 925 pp. Mousey, H. 1924. Further notes on the orchids of Hatley, Stan- stead County, Quebec, 1923. Canadian Field-Nat. 38: 62-63, 86-88. ——————. 1941. Two orchids new to the Province of Quebec and the Dominion of Canada, Spiranthes vernalis Engelm. & Gray and Spiranthes cernua (L.) L. C. Rich. var. odorata (Nutt.) Correll. Canadian Field-Nat. 55: 79-80, + 2 pl. 1942. The spring ladies’ tresses (Spiranthes vernalis Engelm. & Gray) in Canada. Canadian Field-Nat. 56: 1-2, + 2 pl. ROLAND, A. E., & E. C. SMITH. 1963-1964. The flora of Nova Scotia, Part 1. Proc. Nova Scotian Inst. Sci. 26(2): 1-238. Voss, E. G. 1972. Michigan Flora — Part I. Gymnosperms and Monocots. Cranbrook Inst. Sci. & Univ. Mich. 488 pp. WHITING, R. E, & P. M. CATLING. 1971. Checklist of Ontario orchids. Fed. Ontario Nat., Newsletter 12(3): 9-10. DEPARTMENT OF BOTANY UNIVERSITY OF TORONTO TORONTO, CANADA MDS 1A1 INVESTIGATIONS OF NEW ENGLAND MARINE ALGAE VI: DISTRIBUTION OF MARINE ALGAE NEAR CAPE COD, MASSACHUSETTS? DOUGLAS C. COLEMAN AND ARTHUR C. MATHIESON Cape Cod is a major phytogeographic boundary on the northeast coast of North America, delineating a distinctive northern and southern marine flora (Chapman, 1964; Humm, 1969; Setchell, 1922; Stephenson & Stephenson, 1949). Harvey (1852-1858) and Farlow (1870, 1882) first recognized its importance as a phytogeographic boundary. Additional information regarding the uniqueness of the Cape Cod flora was contributed by Collins (1900), Davis (1913a, b) and Taylor (1937, 1957). Although the marine flora of Cape Cod has received con- siderable attention since the time of Harvey, no one has ever conducted simultaneous year round studies of the algae on both sides of the Cape. In addition, nothing is known about the seasonal changes that occur in the Canal, which joins Cape Cod Bay and Buzzards Bay. Conover (1958) and Sears (1971) conducted seasonal studies of the algae in southern Cape Cod, but they gave no consideration to the Canal. The purpose of the present investigation was to atudy the horizontal and vertical distribution of seaweeds at seven locations from Scituate to Woods Hole, Massachu- setts, and reproduction including the Cape Cod Canal. The seasonal occurrence of seaweeds at the same sites will be discussed in another paper. Monthly collections of all the conspicuous algae at seven stations (Fig. 1) were made from January to December, 1969. Severe winter conditions (heavy ice and surf) ex- isted from December to March, and they restricted some "Published with the approval of the Director of the University of New Hampshire Agriculture Experiment Station as Scientific Con- tribution Number 656. 537 538 Rhodora [Vol. 76 collections. Specimens were collected on foot in the inter- tidal zone and by SCUBA diving to a depth of 60 feet in the subtidal zone. Diving in the Canal was done at slack water (Anon. 1969a). The collections were brought to the Uni- versity of New Hampshire for identification and processing. Taylor (1937, 1957, 1960) was the primary source of identification and nomenclature. A number of additional references were consulted for the identification of species and the determination of recent geographical records along the Atlantic Coast of North America subsequent to Taylor's (1960) summary. These include: Adey, 1964, 1965, 1966; Bel & McFarlane, 1933a,b; Blomquist & Humm, 1946; Cardinal, 1964, 1965, 1966, 1967a,b,c, 1968; Collins, 1909; Edelstein, et al., 1967, 1969, 1970; Edelstein & McLachlan 1966, 1967a,b, 1968a,b, 1969; Fritsch, 1935, 1945; Hoek, 1964; Hoyt, 1920; Lamb & Zimmermann, 1964; Lee, 1968, 1969; Lewis, 1914; MacFarlane & Bell, 1933; McFarlane & Milligan, 1965; Mathieson & Fuller, 1969; Mathieson, Dawes & Humm, 1969; Rhodes, 1970; South & Cardinal, 1970; Stone, et al., 1970; Wilce, 1959, 1965; Williams, 1948, 1949; Wulf, et al., 1968; Zaneveld & Barnes, 1965; Zaneveld, 1965, 1966a,b, 1972. The nomenclature of Parke & Dixon (1968) was applied in most cases. Surface water temperature and salinity information was recorded when each station was investigated. Temperature was determined with a laboratory grade mercury thermom- eter; salinity was measured with a set of hydrometers ( G. M. Mfg., Co.). In addition, hourly and daily surface water temperature data were obtained for each end of the Canal from thermographs monitored by the Fisheries Division of the State of Massachusetts. Additional (daily) temperature and salinity information was supplied by the Woods Hole Oceanographic Institute, Falmouth, Massachusetts (per- sonal communication with Dean Bumpus and Joseph Chase). Figure 1 illustrates the location of the seven study sites. One station (Scituate) was located north of the east end of the Canal, four were in the Canal proper and two (Woods 1974] Marine Algae — Coleman and Mathieson 539 LI LI BS SCITUATE 42° L` 7 CAPE COD BAY Ü x » 4147 en CAPE COD CANAL 7 V a L/^O * v CAPE COD À GS ECK g y 4 f á | u d f <2 ( f ay o PI WOODS HOLE 70°30' 70510 Figure 1. Map of Cape Cod, Massachusetts, showing the seven stations. Hole and Wings Neck) were located south of the west end of the Canal. The land-cut of the Canal cuts across the Cape in an east-west direction from the town of Buzzards Bay (approximately 41°44’ N Latitude, 70°37’ W Longi- tude) to Sandwich (approximately 41°46’ N Latitude, 70°30’ W Longitude). It is seven miles in length, with an additional seven miles of approach channels. Several differences are obvious between the two sides of the Cape. To the north, the substrate consists of granite outcrops, boulders and cobbles, while to the south the sub- 540 Rhodora [Vol. 76 22 TaN Í N 201 7 N 18] 16) . 14 O o 12 [99] c 10! O A 8 6} 4 Jg | 4 i STATION 1 —— 2 Da STATION 3 -——- £g; STATION 5 —-—:- 0} cy 2 STATION 7 ---- PI TFITMITATMI I TI TATSTOINTI DI MONTHS Figure 2. Mean monthly variation of surface water temperatures at stations 1, 3, 5 and 7 during 1969. strate is largely sand and mud with fewer boulders. The Cape is also the dividing line between the Labrador Current to the north and the Gulf Stream to the south. Thus, a marked difference in water temperatures occurs during the summer. The yearly temperature range south of the Cape was approximately 22°C., while that to the north was about 17°C. (Fig. 2). The salinity ranges south of the Cape were usually higher than those to the north, particularly during spring and summer (Fig. 3). The tidal amplitude is much greater on the north than the south side. Thus, a three foot tidal amplitude exists in Buzzards Bay, while it is approxi- mately ten feet at Scituate (Anon. 1969a). The waters of Cape Cod Bay are over 100 feet deep while those in Buz- zards Bay are shallower. 541 Marine Algae — Coleman and Mathieson 1974] ‘6961 Sutanp L pue G ‘g ‘g suorj93s ye ÁAjrui[es iej3eA ezans jo UuorgLieA ÁA[qquour ULIW '6 əng SHLNOW Eg: p EO Tsepv EP ECINI V. Ew iE] -——- 4 NOILVIS 06g -.—.- 6 NOILYLS | ——- € NOILW1S |" OG —— | NOI1V1S - oc» A got? NP E E T$ "T = lC u— UOCE NL Sze 542 Rhodora [Vol. 76 STATION 1 —— STATION 3 ——— STATION 5 -:—.- 25; STATION 7 -——- ee 20: 151 o e Ome — see — + — —— -— ——— e — -~ oe e— — o ee eo co ooo -————-—— 9 e- — — — —— -— — — — e DEGREES C' 104 JULY JANUARY LES .- —e es er me e.o rr dm ^ N [99] N Aa ho Cn NO Oo DAYS Figure 4. Mean daily variation of surface water temperatures at stations 1, 3, 5 and 7 during January and July, 1969. 1974] Marine Algae — Coleman and Mathieson 543 The Canal has a limited amount of solid substrate, no wave action and a reduced intertidal zone. The seasonal temperature ranges are intermediate between the north and south sides of the Cape, but daily fluctuations during the summer are greater than on either side of the Cape (Fig. 4). The summer temperatures in the Canal are somewhat lower in the east than the west end (Fig.2). Seasonal and spatial variations of salinity are minimal (Fig. 3). The tidal amplitude in the Canal ranges from four feet in the west end to ten feet in the east end (Anon. 1969b). There is a difference of about two hours between the tides at the two ends (Anon. 1969b). The current reaches a peak of about six knots, three hours after slack water. Slack water occurs uniformly throughout the Canal, and it does not coincide with either high or low water (Anon. 1969b). The substrate at each site in the Canal grades vertically from granitic boulders to sand-silt. In addition, there is a general decrease in stable substrate towards the east end. The Canal is maintained by the Army Corps of Engineers. A private access road along the banks was used to reach the stations. Telephone poles are located at 100 foot inter- vals along the Canal; they are numbered beginning at the east end. Stations 2-5 were located in the Canal and eor- respond to poles 10, 45, 245 and 385 respectively. Station 3 (pole 45) is located in front of the Cape Cod Canal Power Plant. A detailed description of the Canal stations, plus the Scituate, Wings Neck and Woods Hole sites is summarized in the Appendix. SPECIES COMPOSITION A total of 106 taxa was collected at the seven stations, including 52 Rhodophyta, 35 Phaeophyta and 19 Chloro- phyta. Table I summarizes the species composition at each station. Scituate and Woods Hole showed the greatest diversity of species, while stations 2-6 showed lower num- bers. No subtidal collections were made at station 2. It is of interest to note that the relative percentages of Rhodo- phyta, Phaeophyta and Chlorophyta were about the same at each station. 544 Rhodora [Vol. 76 60: NORTHERN ---- | SOUTHERN -= COSMOPOLITAN 50; 40: NUMBER OF TAXA/ GROUP w Q \ \ \ 7 204 \ ; \ \ 7 \ m ^ MEMBRE d 104 V Ll 4 7 P — : =. =. -e ] æ. —. Á 0 : | 2 3 4 5 6 7 STATIONS Figure 5. Number of northern, southern and cosmopolitan species at each station. 1974] Marine Algae — Coleman and Mathieson 545 A listing of the known geographical distribution of each taxon along the Atlantic Coast of North America is also designated numerically in Table I. The species can be grouped into three components (i.e., northern, southern and cosmopolitan) according to their major centers of distribu- tion. The northern species are found most commonly north of the Cape, and they include groups 1, 2, 3, 4 and 15. The southern species are more common south of the Cape; they include groups 7, 11, 12 and 13. Cosmopolitan species are found on each side of the Cape, and they include groups 5, 6, 8, 9, 10, 14 and 16. Figure 5 summarizes the number of northern, southern and eosmopolitan species at each station. Most of the spe- cies were cosmopolitan. The northern component declined towards the south, while the southern component increased towards the south, reaching a maximum at Wings Neck and Woods Hole. Cosmopolitan species included Chaetomorpha melagonium, Rhizoclonium tortuosum, Ulva lactuca, Asco- phyllum nodosum, Petalonia fascia, Ahnfeltia plicata and Ceramium rubrum, while conspicuous northern components included Codiolum petrocelidis, Monostroma spp., Chorda filum, Fucus spiralis, Laminaria digitata, Choreocolaz poly- siphoniae, and Porphyra umbilicalis. Representative species more common to the south of the Cape included Codium fragile subsp. tomentosoides, Sargassum filipendula, Agard- hiella tenera, Champia parvula and Hypnea musciformis. Scituate had a species composition typical of northern New England, while Wings Neck and Woods Hole were more typical of warmer water locations. Thus, Laminaria spp., Gigartina stellata, Ralfsia fungiformis, Chaetomorpha atrovirens, Urospora collabens, and Fucus vesiculosus were common at the former station, while Sargassum filipendula, Agardhiella tenera, Callithamnion baileyi, and Codium frag- ile subsp. tomentosoides were common at the latter stations. The Canal stations, 3-5, included representatives from both sides of the Cape (e.g. Laminaria saccharina, Chondrus crispus, Rhodymenia palmata, Agardhiella tenera, Codium fragile subsp. tomentosoides and Sargassum filipendula. 546 Rhodora [Vol. 76 VERTICAL DISTRIBUTION Figure 6 summarizes the vertical distribution of the con- spicuous species at each station. The distributions were recorded as follows: 1) species restricted to the intertidal zone; 2) species restricted to the subtidal zone; 3) species found in both the intertidal and subtidal zones. Most of the plants at Scituate were collected from the intertidal and subtidal zones, including tide pools. Species that were common to both zones included Chondrus crispus, Chorda spp., Ectocarpus siliculosus, Petalonia fascia, Ceramium rubrum, Corallina officinalis, Dumontia incrassata, and Chaetomorpha linum. Few species (e.g. Chaetomorpha atrovirens, Cladophora flexuosa, Polyides rotundus, Aspero- coccus echinatus, and Laminaria spp.) were restricted to the subtidal zone or intertidal zone (e.g. Codiolum spp., Enteromorpha intestinalis, Rhizoclonium tortuosum, Ulo- thrix flacca, Choreocolax polysiphoniae, and Fucus spiralis). Subtidal substrate was lacking at station 2, and the lowest number of species was found here. The Canal stations (2-5) had the highest subtidal com- ponent. Species restricted to the subtidal zone in the Canal were also found in the intertidal zone at other stations. The most common subtidal species in the Canal included Lami- naria spp., Chondrus crispus, Agardhiella tenera, Chorda spp., Chordaria flagelliformis, Ulva lactuca, Petalonia fas- cia, and Scytosiphon lomentarius. Species restricted to the intertidal zone included Urospora penicilliformis, Ascophyl- lum nodosum, Fucus spiralis, and Choreocolax polysiphon- iae; those common to both zones included Ectocarpus sili- culosus, Fucus vesiculosus, and Sphacelaria cirrosa. The majority of species collected at Wings Neck and Woods Hole was found in the subtidal zone and the inter- tidal-subtidal zones. Few species were restricted to the intertidal zone. Species restricted to the subtidal zone in- cluded Agardhiella tenera, Sargassum filipendula, Callitham- nion roseum, Dasya pedicellata, Seirospora griffithsiana, Leathesia difformis, Chordaria flagelliformis, and Clado- 1974] Marine Algae — Coleman and Mathieson 547 INTERTIDAL INTER-SUBTIDAL SUBTIDAL NUMBER OF SPECIES STATIONS Figure 6. Vertical distribution of species at each station. phora spp. Species found in both zones included Chondrus crispus, Melobesia lejolisii, Ectocarpus siliculosus, Fucus vesiculosus var. spaerocarpus, and Chaetomorpha linum, 548 Rhodora [Vol. 76 while those restricted to the intertidal zone included Bangia fuscopurpurea, Porphyra wmbilicalis, Fucus spiralis, and Ulothrix flacca. DISCUSSION Setchell (1917) was one of the first workers to emphasize the role of temperature in determining the geographical distribution of seaweeds. He divided the oceans into 5°C. intervals or isotherms, according to the maximum summer water temperatures. Hutchins (1947) also confirmed that 5°C. intervals either favored or inhibited growth and re- population of marine organisms. Williams (1948), Parr (1933), Well and Gray (1960), and Humm (1969) have reported a similar relationship between temperature and species composition. As suggested by Setchell (1917) Cape Cod is a dividing line between the 15° and 20°C. maximum summer isotherms. Hence, it is not surprising that it is a major phytogeographic boundary, with distinct floras oc- curring north and south of the Cape. Several factors are responsible for the marked difference in summer temperatures between the north and south side of Cape Cod. A southern extension of the Labrador Current carries cold water as far south as the Cape, where it re- mains throughout the year. In addition, the deep waters of Cape Cod Bay warm up slowly during the summer. Davis (1913a, b) described Cape Cod Bay as a “holding pocket” of cold water. He further described the islands of Martha’s Vineyard and Nantucket as barriers protecting Buzzards Bay from the intrusion of cold waters off Gay Head. The high surface water temperatures in Buzzards Bay result from a northern extension of the Gulf Stream into the shallow confines of Buzzards Bay. During the winter, cold air lowers the temperature on both sides of the Cape and the sharp differential of temperature disappears. Winter temperatures in Buzzards Bay, however, may be somewhat lower than in Cape Cod Bay, because of the shallowness of the water of the former location. 1974] Marine Algae — Coleman and Mathieson 549 The transitional nature of the Cape Cod Canal is docu- mented by its species composition. The Canal is dominated by cosmopolitan species; northern species increase toward the east end of the Canal, while southern components in- crease toward the west end. Setchell (1922) also recognized a cosmopolitan group of plants common to both sides of the Cape — in contrast to more northern and southern elements. The completion of the Canal in 1914 presumably provided a direct route for spores to pass from one side to the other. Stephenson (1944) has shown a similar transition of flora and fauna near the Cape of Good Hope in Africa; on the west coast the waters are relatively cold, while on the east coast the shore is bathed by the warm waters of the Indian Ocean. Seven major components are evident near the Cape of Good Hope; the major components are the cosmopolitan, warm- and cold-water elements. Physical factors, such as tidal amplitude, wave action and availability of solid substrate, determine the local distribu- tion of species at the seven sites. Salinity was not con- sidered a major factor in the areas studied because of the small range recorded. Scituate had the highest number of species; it is the most exposed site having ample substrate. The lack of wave action, lower tidal amplitude, and reduced substrate in the Canal were probably responsible for the low number of species at stations 2-5. Woods Hole had the second highest number of species, even though tidal ampli- tude, wave action, and substrate were reduced south of the Cape. The abrasive action of sand at Wings Neck resulted in low species numbers. A variety of physical factors, such as temperature, salin- ity, light intensity, substrate and exposure, determine the vertical distribution of seaweeds (Stephenson & Stephen- son, 1949; Chapman, 1964). Most algae at Scituate were collected from the intertidal and subtidal zones, while in the Canal and south of the Cape the majority of species was restricted to the subtidal zone. The former location (Scitu- ate) is characterized by greater tidal amplitude and wave action, both of which are necessary for the development of 550 Rhodora [Vol. 76 an extensive intertidal flora — assuming substrate is not limiting. The vertical position of species varied from sta- tion to station. Species found in the intertidal and subtidal zones at Scituate were often restricted to the subtidal zone in the Canal or south of the Canal. ACKNOWLEDGEMENTS We would like to thank Dr. A. Hodgdon for his critical review of the manuscript. In addition we express our grati- tude to Dr. R. Fralick, Dr. R. Turner, and Mr. John Hans- man for their assistance with diving and collection of specimens, to the Army Corps of Engineers at Cape Cod for diving access to the Cape Cod Canal, and to Dean Bumpus and Joseph Chase for the use of unpublished tem- perature and salinity data. APPENDIX DESCRIPTION OF STATIONS Scituate (station 1) is located about 30 miles north of the east end of the Canal at approximately 42°12’ N latitude and 70°43’ W longitude (Fig. 1). It is a semi-exposed site, consisting of massive granite outcroppings. There are several tide pools and a well developed intertidal zone. The substrate in the subtidal zone consists of large outcroppings, boulders, cobbles and sand. The surface water temperatures ranged from approximately 2-16°C. The highest tempera- tures occurred in August, while the lowest occurred in March. The daily temperature range was about 2°C. during the summer and 3°C. during the winter. The average monthly summer temperatures at Scituate were the lowest of all the stations. Salinity values remained relatively con- stant throughout the year, ranging from a low of 30.6°/,, in the summer to a high of 32.5°/,, during the winter. Station 2 is located at the east end of the Canal at the Scusset Breakwater (Fig. 2). Collections were restricted to the intertidal zone because of the lack of subtidal sub- strate. An extensive intertidal zone is present. The water 1974] Marine Algae — Coleman and Mathieson 551 temperatures ranged from 12°C. to 19°C. The tidal ampli- tude is 10 feet (Anon. 1969a). Station 3 is located at the power plant (Fig. 2). Rip-rap extends to about 6 feet below M.L.W., while a shelf extends out 30-40 feet and to a depth of 15 feet. During operation, the power plant discharges water of over 23.9°C. Substrate is limited and a small intertidal zone is present. The tidal amplitude is approximately 10 feet. Station 4 is approximately 2.5 miles east of the west end of the Canal proper (Fig. 2). The study site was on the south bank of the Canal. The substrate grades from rocks (boulders) in the upper shore to sand-silt in the lower sub- tidal zone. The tidal amplitude is about 6 feet (Anon. 1969a). A limited intertidal zone is exposed during low tide. Station 5 is located at the Engineer’s station at the west end of the Canal. Collections were made in the vicinity of the pier. Rip-rap extends to about 12 feet below M.L.W., while a shelf extends out to about 75 feet and slopes to about 20 feet below M.L.W. The tidal amplitude is about 4 feet (Anon. 1969a). Wings Neck (station 6) is located at approximately 41°31’ N latitude and 70°40’ W longitude in the town of Bourne, Massachusetts (Fig. 1). It is a semi-exposed area subject to southwest winds during the summer. A limited intertidal zone is present; it is composed of small cobbles and a few large boulders. A few shallow tide pools are evident. The tidal amplitude is about 4 feet (Anon. 1969a). The range of water temperatures was 0.2-22.19C. The mean monthly winter temperature (2.2°C.) was lowest in Janu- ary. At that time the temperature was lower than at Scitu- ate, the northernmost station. The area is subject to severe ice scouring during the winter. The salinity remained rela- tively constant throughout the year, with a range of about 0.9°/oo (Fig. 4). Woods Hole (station 7) is located at approximately 41°31’ N latitude and 70°40’ W longitude in the town of Falmouth, Massachusetts (Fig. 1). Collections were made at the jetty behind the U. S. Fisheries Building. The bottom 552 Rhodora [Vol. 76 is approximately 20 feet below M.L.W., at the deepest point, and the bottom substrate is composed of sand and silt. Some small rocks were present at one side of the jetty. The tidal amplitude is about 2 feet (Anon. 19692). The temperature ranged from a low of 0.2°C. in January to 22.5°C. in Au- gust (Fig. 3). Daily temperature ranges were small during the year (Fig. 5). The salinity remained relatively con- stant throughout the year with a range of only 0.89/, (Fig 4). Footnotes to Table I ! Geographical distribution zones relating to Table I, right hand column. T Northern Massachusetts to Newfoundland and north. Northern Massachusetts to Nova Scotia. Southern New England-Long Island Sound to Gaspé and Labrador. Southern New England-Long Island to Newfoundland and north. New Jersey-Maryland to Newfoundland and north. New Jersey-Maryland to Gaspé and Labrador. New Jersey-Maryland to ‘Cape Cod. North Carolina to Newfoundland and north. North Carolina to Gaspé. South Carolina to Newfoundland and north. . South Carolina to northern Massachusetts. Tropies to southern New England-Cape Cod. . Tropies to northern Massachusetts. Tropics to Newfoundland and north. . Southern New England-Long Island Sound to northern New Eng- land-Nova Scotia. 16. Tropies to northern New England-Nova Scotia. | 9l st Ree RH HM NC CET ? — Neoagardhiella baileyi (Harvey ex Kützing) Wynne et Taylor. (Wynne and Taylor, 1937). 3 — Phyllophora truncata (Pallas) Newroth et Taylor. (Newroth and Taylor, 1971). * — Phyllophora pseudoceranoides (Gmelin) Newroth et Taylor. (Newroth and Taylor, 1971). 553 Marine Algae — Coleman and Mathieson 1974] Sl IL 6 9 FP SI SI Y Eo x X X X I X Eo X X X X X X5 8 X X x I X X g kna X X 8 X X Y X X X X y X X X X uox X X X wT xX X X X xX x x L X X X X X Z X g X x PE.xx X FÉ 8X X xX x X g x x X b na ph Xx X € Y x x v X [mde ) 9 G $$ 8 € I -08D; uonedg uonnqrujstq SHIOSdS JO [e303qn$ Snogosely (YOY) swwuofyprouad v4odso. f] 81933€eg 2? SOWOH (u1p1e3y 2) suaqnvjjoo 040ds04[] "[ DINJIN) VMN PINYL (UAM) VIY x144701N SUZY $uoosouids vydsowmobuods SUZMA (UAM) 079040 vud.40woDuodg: SUIZINY wnsonj402 WUWMUOJIOZIY Y MOJICA WnLYIINd 0u,042801,0 Jg X201331uUA, (39o4nu[) 227122240 01404280u0]g Yplesy ‘f (7T) 02ui1] vydsowosaquy xXurT (I) $0u22s21u2 vd40140423u5] BAIS (4004) ULA) $api0803u2ut02 ‘dsqns Joey (CNS) 2516044 wnrpog yonyony svpyaso.zad wn]01p07) unelg "v Wniunbasb wn]opo;) SUIZINY (AvAIvH xo syylay) 229016 0401d0p7]7) LAICH (IANN) vsonxay n401d0pv]/) SUIZINY (IYON 22 1eqo M) wnruobyjau vydsowojavyy SUZMA (IAW) UNUN vydwowojanyy) IOJAB L $u3.240420 VYALOWOJIVY J uoxt[ e3&udoao[u;) NOLLOSIHLISIG ‘I W'ISVIL [Vol. 76 Rhodora 554 8 X X X X V X X X X X X X [4 X X X X X g X X X T X G X X X X X X VI X X X X X X X G X X X 6 X X 8 X X X G X X X X X X g X X X X X g X X X X X X X € X X X X X X X G X X X X X X X [2 X X X X X X 8 X X X X X X X jeorydeis L 9 G P $8 Z I -0991 u01}8}S uonnqrnsiq "[ snsojnaisaa SNIN "[ sypuids SNIN H [jemog (Yypresy *D) $uaosounao 'dsqus 'pueure "T $704/02282p snony [[9oq (upresy 1) $n302u2pu2 'dsqus 'PUIWƏ "T $7/22281p SNIN Å [moq (upre3y 7) $ni[21281p ‘dsqns *pueuro "T $70/22282p SNIN 3nouosedy (LAPA) 0j0010nj 028112117] aAqsuAT (UAMTIIG) Snsojnayis Snd.020225] Ər (uospng) $12320]mo1u20jf uowdasofi201(q exured (upieSy f) wn30mmpun wnwy214201482(] SUIZINY UNIVA wm1[214200482(] XNOINOWP'T (2o[[n]J) SPLA 71282401489 (T xnoJnouire'] (I) 0309]n290 0128947]S2(] upae3y D) (ABN) 520440/2]7200 v140p-1017) OLATUA 0$02u21402 0p401//) esnogxoejs (I) UMY vp4oty;) 9[[IA9J42) (SUBSTI) $nj0w129 $n2200042d8 Pp SOf IT (T) wnsopou wnjjfidoosy UoxXeL v14udoeeuq uoo I W'IS VL 955 Marine Algae — Coleman and Mathieson 1974] 62 TZ 9% IZ $2 8I 92 G > Cae aap Y Y XX X X t x PON UN X X XX X 2 ee s X M q E CY X Yxx.EX I xx X q > A i g X X Xx X q X X X X X X X wT xX X X X X¥ X X wT xX X X X X X 8 X X X X X X G X X X X X X v X X gc x q X X X 9 X X e "ue E. wma »)9 Sg $ 8 Z I -085): uonejg uonnqruasuq [e3o03qn$ UAY (Up4e3y Q) 9??»214Da01p 012143049 D1(d Sy upiesy “OQ (YOY) 08504419 0140]9201d S JUTI (oAqSuA'T) sni403u2140] uoydisozhag upaes3y ‘O »jpuodapj wunsspDADS upies3y 'e (snoyosealy) »soon.A420 MS {DY Joup.e2 22 042499 (1euuns) szuofibunj visfqoa] IAIA (YOY) 02u:003w0]d 0140909und 9[[IA9.12) 2220/220] 014029unqd ueu[[oÉw (T) $27040322] Daiva 9zjunw (1e[[h]N) 02280/ 01u0]022d 9[[IAo.15 SUVINDUDLS 0W2WoOtL pA 3nouosedy (J) snuaoffip 01821[2097] XNOINOWPT (J) 0w140)12090$ 01.501140] xnoainoure (uospng) 220320?p Manur $103jeq (SUIZA) vpunoos Mpo fW |eureH (yug) vsomuvab mpo fw MOAB $270./2ds “ABA SNSOMDNSIA SNIN H upies2y 'fe SNAALDIOLIVYALS ‘AeA SOSOMNSIA SNIN H uoxe |, ‘quod ej&qdoeeuq — I W'IHViL [Vol. 76 Rhodora 556 £ X X X X X X XNOINOWeT (uop[n]y) VIVSSVLIU? 013uowwn(T GT X X X ero, (xnoanoureT) wnznjnjsnd uo12120?0w.t0(] IIT X X X X Qpie3y ‘O (upae3y O) 020j221pod visvg ST X X X X X X DAICH WNSOYLL9 "IBA $J193j3€q (uospng) wnaindind wniwogo03sfi) tT xX X X X X X I sypupygo vu1]]040;) I X ƏUSOF (3[9gyur9.13g) wn3da4oswmno419 WUNYALOWOLYI DIJ L4 X X X X X qosureq aviwoydishjod 13:0]0209401/7) G X X X X X X X osnouxoe3g sndsio0 Sn4pwuoty 7) eI X X AVAIVH 71]0/1p98 D14pW01;) eI X X X AoAIeH (upadeSy 9) nynaund vidaum[;) 9I X X X X K9AJA€H 4/1221428 4n] 0497) PI X X X X X X X yplesy 7) (uospnH) wmnaqna wniuwupAo) 9I X X X X eAq3uÁ'[ (YOY) wnoso.4 wowwn31]v7) S X X X ADAICH IVD uowump1110;) GI X OLIV Diafiuny D40810042WuOg FI X X X X X oAqsduA'] (u&aA[[IT) 224nd4ndoosnj mbung G X MOJICA (&oA1eH) ?!wnupnot40wD uowwnygyiquy S X X X X X X X seri (uospng) vood miyafuyy GI X X X X X X Zjgnuqog (upiedy '() vua} vjjovp4oBy Iworydeigs L 9 S p 8 Gc I UOX", -099r uone3g ByAydopoyy uonnqrijsiq juo» I W'IHV.L 557 Marine Algae — Coleman and Mathieson 1974] P oM MX PA p< Ko OM pI v x p< PAPA P Pd PA PS PS A P4 PS PS MPO FA PA P jeorydeis L 9 9 v? & 6 -09*); uonejg I UorjnqHastq zjtuuog (uosreureuuog) SUNANI vi4Amwn]d Kepy (Snoyosely) wupvw.oua) 0121020 ufa q upae3y "f (preMpooM xə q3nouepoor) vyofruniquow nioydonhyd .qorpum (19UIN L) 220:p04qQ vnuoydonhyd ueui[[e£X (1g»e1dngp) ?/4opuoppuu 52904472 d yoursoy 2:$120/2] 01820012 W Ao[Ae] Va sut[[o?) (&oA1eH) SIsUapDILO 0140]W9'WtO'T Ao[re, (LIAIGH) 0wnfio]imq MAVJUIWO'T ueui[[9£w 21012018 WNIUUMDY LOY WT uonip&eH (UBN0IQ) ovun. wy hydoy WT XNOANOWLT (UIF[NAA) s2u40ft98nwu voudhH opaeN snd/igogo..d o:pun4quappH LIAIGH (upiedy Q) VUVNALIUD 01]]19WWAMT) ypresy O $2uo3 MSYPPLÐ ƏMOH (rurtpreueZ) 1?p19]0 UNYINAJOUOD Ko[|ox1og xa PPLYITUILQ (UOSpnH) s?40]j1109 01401,d18010]7) sie3jeg (esnouxoejg) n2vj22$ DUAD uoxer 'juoo vyAydopoyy — I d'Id ViL [Vol. 76 Rhodora 558 08 S9 99 69 Ay GP IS 8e 08 IE Zg 02 FI LE e X X X X X L X e X X X X X X G X X X X X v X X Ol X XX X X X X I X X X 6 X X X q X X X X X X q X X X X O0 X XX X X X X G X X X X X X X Oo XX X X X v X X X 9T X X X X X v X X X X X X [ondei L 9 G p 8g ZB I -092 uonejg uonnqtjstq "IV.LOLL 18403440N $103jeg (upiesy `f) n370142u2 DINDA I, uoxIg (LƏAILH) vunisydjfi4b DLodsouag e[[IA9.12) (7T) Voua niuamhpoyy vA[IS (uospnH) səproasə fuos mawopoyy e3urAuesoq (JOOFLYSIT) Pwo fyporuəd uoztoyIopoyy yplesy ^f CT) suvowuiqum v4fiqdaoq Ypresy ‘O (upiedy 9) omuru duhyd.og 1e1nug, 2222250912] n4fia[d.0q AAD (ULMA LA 70074431T) 727]0224n v1uoudisfijoq J0 AeL 20:28u0-20a0w muoydishjod Əə (uospng) su22s2401w viwo1disfijoq Apue[ (T) 2s0u»] nuoydishjog Kepreq ?/iaa40u o1uowdisfijoq Jesueidg (uospnpg) 2326u0]? Diuoydishjog JexooH ut AoA1eH x2 o[[IA9.12) (Usma) 9?vpnuop viuowudisfijoq Ər (uospnH) snpungo4 saprhjog uoxu] ‘quod e3&uydopoud — I a TAV iL 1974] Marine Algae — Coleman and Mathieson 559 REFERENCES ADEY, W. H. 1964. The genus Phymatolithon in the Gulf of Maine. Hydrobiologia, 24: 377-420. 1965. The genus Clathromorphum (Corallinaceae) in the Gulf of Maine. Ibid. 26: 539-573. 1966. The genus Lithothamnion, Leptophytum (nov. gen.) and Phymatolithon in the Gulf of Maine. Ibid. 28: 321- 370. ANONYMOUS. 1969a. Tide tables, high and low water predictions for 1969, North and South America including Greenland. En- vironmental Sciences Services Administration, Coast and Geo- detic Survey. 280 pp. -.—————, 1969b. Tide tables, Cape Cod Canal. U. S. Army Engineer Division, New England Corps of Engineers. 12 pp. BELL, H. P., & C. MACFARLANE. 1933a. The marine algae of the maritime provinces of Canada, I. List of species with their distribution and prevalence. Can. Jour. Res. 9: 265-279. 1933b. The marine algae of the maritime provinces of Canada, II. A study of their ecology. Ibid. 9: 280-293. BLoMQUIST, H. L., & H. J. HUMM. 1946. Some marine algae new to Beaufort, North Carolina. Jour. Elisha Mitchell Sci. Soc. 62: 1-8. CARDINAL, A. 1964. Étude sur les Ectocarpacées de la Manche. Beihefte zur Nova Hedwigia. 15. 86 pp. 1965. Liste préliminaire des algues benthiques de la baie des Chaleurs. Rapp. Ann. 1964, Sta. Biol. mar. Grande Riviere: 41-51. 1966. Additions a la liste benthiques de la baie des Chaleurs. Rapp. Ann. 1965, Sta. Biol. mar. Grande Riviere: 85-43. 1967a. Inventaire des algues marines benthiques de la baie des Chaleurs et de la baie de Gaspé (Québec), I. Pheo- phycees. Nat. Can. 94: 233-271. 1967b. Ibid. II. Chlorophycees. Ibid. 94: 4471-469. 1967c. Ibid. III. Rhodophycees. Ibid. 94: 735-760. 1968. Répertoire des algues marines benthiques de l'est du Canada. Cah. Inf., St. Biol. mar. Grande Riviere, Que. No. 48. 213 pp. CHAPMAN, V. J. 1964. The Algae. Macmillan and Company Ltd., London. 472 pp. CoLLINS, F. S. 1900. Preliminary lists of New England plants V. Marine algae. Rhodora 2: 41-52. 1909. The green algae of North America. Tufts Col- lege Studies. (Sci. Ser.) 2: 79-480. 560 Rhodora [Vol. 76 CONOVER, J. T. 1958. Seasonal growth of benthic marine plants as related to environmental factors in an estuary. Publ. Inst. Mar. Sci. 5: 97-147. Davis, B. M. 1913a. General characteristics of the algal vegeta- tion of Buzzards Bay and Vineyard Sound in the vicinity of Woods Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries 31: 443-544. 1913b. A catalogue of the marine flora of Woods Hole and vicinity. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries 31: 795-833. EDELSTEIN, T. L. CHEN, & J. MCLACHLAN. 1970. Investigations of the marine algae of Nova Scotia, VIII. The flora of Digby Neck Peninsula, Bay of Fundy. Can. Jour. Bot. 48: 621-629. EDELSTEIN, T., & J. MCLACHLAN. 1966. Investigations of the marine algae of Nova Scotia I. Winter flora of the Atlantic Coast. Can. Jour. Bot. 44: 1035-1055. & . 1967a. Ibid. III. Species of Phaeophy- ceae new or rare to Nova Scotia. Ibid. 45: 203-9210. & . 1967b. Ibid. IV. Species of Chlorophy- ceae new or rare to Nova Scotia. Ibid. 45: 211-214. & . 1968a. Ibid. V. Additional species new or rare to Nova Scotia. Ibid. 46: 993-1003. & . 1968b. Ibid. VI. Some species new to North America. Ibid. 47: 555-560. & . 1969. Petroderma maculiforme on the coast of Nova Scotia. Ibid. 47: 561-563. ; , & J. S. CRAIGE. 1967. Investigations of the marine algae of Nova Scotia II. Species of Rhodophyceae new or rare to Nova Scotia. Ibid. 45: 193-202. , .& . 1969. Preliminary survey of the sublittoral flora of Halifax County. J. Fish. Res. Br. Canada 26: 2703-2713. FanLow, W. G. 1870. XVII. List of sea-weeds or marine algae of the south coast of New England pp. 281-294. In: U. S. Comm. of Fish and Fisheries. Commissioner’s Report 1871-1872. Wash- ington. Government Printing Office. . 1882. I. The marine algae of New England. Ibid., report 1879. pp. 1-210. FRITSCH, F. E. 1935. The structure and reproduction of the algae, I. xviii + 791 pp. Cambridge Univ. Press. 1945. Ibid., II. xiv + 939 pp. Cambridge Univ. Press. Harvey, W. H. 1852-58. Nereis Boreali-Americana. I. Melano- spermae. Smithsonian Contrib. Knowl., 3: 1-150, pl. 1-12. 1852; II. Rhodospermae. Ibid. 5: 1-258, pl 13-36. 1853; III. Chlorospermae. Ibid. 10: ii + 1-140, pl. 37-50. 1858. 1974] Marine Algae — Coleman and Mathieson 561 HoEK, C. VAN DEN. 1963. Revision of the European species of Cladophora. xi + 248 pp. Leiden. Hovr, W. D. 1920. Marine algae of Beaufort, N. C. and adjacent regions. Bull. U. S. Bur. Fish. 36: 367-556. HuMM, H. J. 1969. Distribution of marine algae along the Atlan- tic Coast of North America. Phycologia 7: 43-53. Hutcuins, L. W. 1947. The bases for temperature zonation in geographical distribution. Ecol. Monographs 17: 325-335. LAMB, M. & H. ZIMMERMANN. 1964. Marine vegetation of Cape Ann, Essex County, Massachusetts. Rhodora 66: 217-254. LEE, R. K. S. 1968. A collection of marine algae from Newfound- land, I. Introduction and Phaeophyta. Nat. Can. 95: 957-978. 1969. Ibid., II. Chlorophyta and Rhodophyta. Ibid. 96: 123-145. Lewis, I. F. 1914. The seasonal life-cycle of some red algae at Woods Hole. Plant World 17: 31-35. MACFARLANE, C., & H. P. BELL. 1933. Observation of the seasonal changes in the marine algae in the vicinity of Halifax with particular reference to winter conditions. Proc. Nova Scotian Inst. Sci. 18: 134-176. , & G. M. MILLIGAN. 1965. Marine algae of the Maritime Provinces of Canada. A preliminary checklist. Nova Scotia Res. Found., Seaweed division. Halifax. Mimeographed, 24 pp. MATHIESON, A. C., & S. FULLER. 1969. A preliminary investigation of the benthonic marine algae of the Chesapeake Bay Region. Rhodora 71: 524-534. , C. Dawes, & H. J. HvMM. 1969. Contributions to the marine algae of Newfoundland. Rhodora 71: 110-159. NEwRoTH, P. R., & A. R. A. TAYLOR. 1971. The nomenclature of the North Atlantic species of Phyllophora Greville. Phycologia 10: 93-97. PARKE, M., & P. S. Drxon. 1968. Checklist of British marine algae — second revision. J. Mar. Biol. Assoc. U. K. 48: 783-832. Parr, A. E. 1933. A geographic-ecological analysis of the sea- sonal chenges in temperature conditions in the shallow water along the Atlantic Coast of the U. S. Bull. Bingham Oceanogr. Coll., 4. 90 pp. Ruopes, R. G. 1970. Seasonal occurrence of marine algae on an oyster reef in Burston's Bay, Virginia. Chesapeake Sci. 2: 61-63. Sears, J. R. 1971. Morphology, systematics and descriptive ecology of the sublittoral benthic marine algae of southern Cape Cod and adjacent islands. Ph.D. Thesis, 295 pp. University of Massa- chusetts. 562 Rhodora [Vol. 76 SETCHELL, W. A. 1917. Geographical distribution of the marine algae. Science 45: 197-204. . 1920. Stenothermy and zone-invasion. Am. Nat. 54: 385-397. . 1922. Cape Cod in its relation to the marine flora of New England. Rhodora 24: 1-11. SovrH, G. R., & A. CARDINAL. 1970. A checklist of marine algae of eastern Canada. Can. J. Bot. 48: 2077-2095. STEPHENSON, T. A. 1944. The constitution of the intertidal fauna and flora of South Africa. Part II. Ann. Natal. Mus. 10: 261-358. , & A. STEPHENSON. 1949. The universal features of zonation between tide-marks on rocky coasts. Jour. Ecol. 3T: 289-305. STONE, R. A., E. HEHRE, J. CONWAY, & A. C. MATHIESON. 1970. A preliminary checklist of the marine algae of Campobello Island, New Brunswick, Canada. Rhodora 72: 314-338. TAYLOR, W. R. 1937. The marine algae of the northeastern coast of North America. 405 pp. Univ. of Michigan Press, Ann Arbor. 1957. Marine algae of the northeast coast of North America. 509 pp. Ann Arbor Press. 1960. Marine algae of the eastern tropical and sub- tropical coasts of the Americas. 870 pp. Ann Arbor Press. WELL, H. W., & I. E. Gray. 1960. Summer upwelling off the northeast coast of North Carolina. Limnol. Oceanogr. 5: 433- 468. WiLCE, R. T. 1959. The marine algae of the Labrador Peninsula and northwest Newfoundland (ecology and distribution). Nat. Mus. Canada 158: 1-103. 1965. Studies in the genus Laminaria III. A revision of the North Atlantic species of the simplices section of Lami- naria. Botanica Gothoburgensia III. pp. 247-256. WILLIAMS, L. G. 1948. Seasonal alternation of marine floras at Cape Lookout, North Carolina. Am. Jour. Bot. 35: 682-695. 1949. Marine algal ecology at Cape Lookout, North Carolina. Bull. Furman Univ. 31: 1-21. WULFF, B. L., E. M. T. Wurrr, B. H. RoBINSON, J. K. Lowry, & H. J. HuMM. 1968. Summer marine algae of the jetty at Ocean City, Maryland. Chesapeake Sci. 9: 56-60. WYNNE, M. J., & W. R. TAvLoR. 1973. The status of Agardhiella tenera and Agardhiella baileya (Rhodophyta, Gigartinales). Hydrobiologica 43: 93-107. ZANEVELD, J. S., & W. D. BARNES. 1965. Reproductive periodicities of some benthic algae in lower Chesapeake Bay. Chesapeake Sci. 6: 17-32. 1974] Marine Algae — Coleman and Mathieson 563 ZANEVELD, J. S. 1965. The benthic marine algae of Virginia. Vir- ginia Jour. Sci. 1965: 346. 1966a. The marine algae of the American coast be- tween Cape May, N. J. and Cape Hatteras, N. C., I. The Cyano- phyta. Bot. Mar. 9: 101-128. 1966b. The benthic algae of Delaware. A preliminary checklist. 35 pp. Scientific Series, Publ. no. 2, Instit. of Ocean- ogr., Old Dominion College, Norfolk, Virginia. . 1972. The benthic marine algae of Delaware, U.S.A. Chesapeake Sci. 13: 120-138. JACKSON ESTUARINE LABORATORY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 A REVISION OF NORTH AMERICAN POTAMOGETON SUBSECTION PUSILLI (POTAMOGETONACEAE)' RoBERT R. HAYNES The importance of the aquatic environment is becoming more evident with our present ecological concerns. Many aquatic plants have been shown to be indicators of water quality and of changes in the aquatic environment (Volker & Smith, 1965; Lind & Cottam, 1969; Harman & Doane, 1970; Stuckey, 1971; Wentz & Stuckey, 1971). Certain species, e.g. Najas minor, Potamogeton foliosus, P. pusillus, and Vallisneria americana, are tolerant of polluted waters. Other species, e.g. Najas flexilis, Potamogeton amplifolius, P. friesii, and P. strictifolius are intolerant. Early workers, e.g. Pond (1905) and Shelford (1918, 1937), considered aquatic vascular plants of little value in the food chain of aquatic animals. Later workers, e.g. Moore (1915), Moyle and Hotchkiss (1945), and Welch (1935, 1952), however, considered the aquatic vascular plants an important source of food for aquatic animals. In a survey of literature on the significance of aquatic vascular plants in the nutrition of animals, Gaevskaya (1966) con- sidered the aquatic vascular plants as a primary source of food for animals of the aquatic environment. According to Martin (1951), the pondweeds (Potamoge- ton) are probably more valuable to ducks than any other group of plants in the United States. Trautman (1957) indicates that the distribution in Ohio of certain species of fish is contingent upon that of associated pondweeds. Gaevskaya (1966) lists 124 species of animals (89 inverte- brates and 35 vertebrates) that feed upon Potamogeton. 'Contribution from the Botany Department, The Ohio State Uni- versity (Paper No. 867). Portion of a Ph.D. dissertation written at The Ohio State University. Field work for two summers was supported by The University of Michigan Biological Station and for one summer in part by a Research Grant from the Ohio Academy of Science and Sigma Xi Grant-in-Aid for Research. 564 1974] Potamogeton — Haynes 565 The plants also serve as shelter for panfish and young fish of other species. Many aquatic insects have at least one stage of their life cycle, if not their entire life cycle, upon the vegetative structures of the pondweeds (Welch, 1952). These insects are, in turn, preyed upon by fish and some birds. Müller-Liebenau (1956) found 294 invertebrates in beds of five species of Potamogeton in the lakes of eastern Holstein. The pondweeds are, therefore, a major factor in the food cycle of the aquatic environment. In fact, among the submerged and floating-leaved species of aquatic vas- cular plants, Gaeyskaya (1966) states that as regards distribution, productivity, and trophic relations of aquatic animals, Potamogeton is the most important genus. Potamogeton Linnaeus is a cosmopolitan genus with ap- proximately 100 species of submerged and floating-leaved aquatic plants. The genus has been separated into two subgenera (Raunkiaer, 1896) and numerous sections and subsections (Graebner, 1907; Hagstrom, 1916). The North American representatives of one of the latter, Potamogeton subsection Pusilli Graebner (narrow-leaved pondweeds), are the subject of this paper. The Pusilli can be separated from the other taxa of the genus by their 3-7(-9)-nerved leaves that are 2-5 mm broad, by their stipules free from the blade, and by their small (to 4.0 mm long) fruits that normally are produced terminally. Although the Pusilli have been included in at least four revisionary treatments (Morong, 1893; Ascherson, 1907; Hagstrom, 1916; Fernald, 1932), uncertainty still exists as to the number of taxa to be included in the group and at which rank these taxa should be recognized. Taylor (1909), for example, accepted only six taxa; Ogden (1957) divided the group into 18 taxa; Muenscher (1944) recognized only 12 taxa; Fernald (1932) acknowledged 21 taxa. The Pusilli have been considered difficult taxonomically for many years (Fassett, 1940) and, in fact, many of the recently collected specimens from North America have been extremely difficult, if not impossible, to identify. The reasons for these difficulties may have been: 1) the Pusilli 566 Rhodora [Vol. 76 are morphologically small and, thus, the characters are difficult to observe; 2) vegetatively (the condition in which much of the material is encountered) the species super- fieially resemble each other; 3) ecological variability has not been fully understood; 4) the nomenclature has been confused. Many authors in the past have not recognized the vegetative variability of the species. As a result, more than 60 names have been proposed for the taxa of North American Pusilli. Because of the apparent economic importance of the group, it is essential to understand the taxonomic relation- ships within the genus. As stated above, this understanding has not previously been accomplished. Therefore, I con- sidered that the subsection Pusilli was in need of a thorough revision. It is understood that problems still exist within the Pusilli, but it is hoped this re-evaluation will clarify some of the confusion within the group. TAXONOMIC HISTORY Potamogeton was first described by Tournefort (1719) who recognized 13 species. Linnaeus (1753), basing his treatment upon that of Tournefort, accepted 12 species. Since that time, a proliferation of names has occurred, both at the generic and infra-generic levels, resulting in con- siderable chaos of the nomenclature. However, because of the nomenclatural confusion and the fact that I am treating only the Pusilli, a brief outline of the major events is pre- sented rather than a detailed discussion of the history. 1788. Walter applied Potamogeton to two species of Myriophyllum (Haloragaceae). 1790. Loureiro named Hydrogeton, based on P. hetero- phyllum. 1819. Peltopsis was created by Rafinesque, based on Potamogeton perfoliatus. 1837. Koch divided the genus into five sections: Chloe- phylli, Coleophylli, Enantiophylli, Heterophylla, Homophylla. 1974] Potamogeton — Haynes 567 1845. Reichenbach recognized five major groups, with- out giving these groups formal rank: Antiphyl- logeton, Chloegeton, Coleogeton, Heterophylloge- ton, Homophyllogeton. 1845. Spirillus was named by Gay, without any species having been listed. 1858. Irmish divided the Homophylla of Koch into two sections, naming the new one Batrachoseris. 1896. Raunkiaer placed all Danish species into two sub- genera: Coleogeton and Hupotamogeton. 1903. Raunkiaer divided the genus into 16 groups, which were not given any formal taxonomic rank. 1907. Graebner (in Ascherson & Graebner) gave formal rank to most of Raunkiaer’s groups (mostly at the subsectional level) and named several other sub- sections. 1913. Nieuwland transferred 20 species of Potamogeton to Spirillus. 1916. Hagstrom named several new sections and numer- ous subsections. Potamogeton subsection Pusilli was first recognized as a natural unit by Raunkiaer (1903). He gave no formal name to the group, but referred to it as “P. pusillus-group.” Graebner in Ascherson and Graebner (1907) combined two of Raunkiaer’s groups — P. pusillus-group and P. confer- voides-group — into the subsection Pusilli. Based upon characters of the stem anatomy, Hagstrom (1916) removed P. confervoides from the subsection. Hagstrom's concept of the group was the one followed by Fernald (1932; 1950) and accepted in the present work. MORPHOLOGY HABIT: The Pusilli are rooted linear-leaved obligate aquatic plants that grow totally submerged except for the spikes, which are usually extended above the surface of the water. When fruiting, the spikes are normally withdrawn below the surface so that the plant is again totally sub- 568 Rhodora [Vol. 76 merged. The mature plants, at least in the temperate re- gions, are mostly functionally annuals, which overwinter as seeds, winter buds, or turions. The winter buds are produced in such abundance that Fernald (1932) and Hagstrom (1916) were of the opinion that the plants rarely reproduced by seeds, but instead reproduced by winter buds. These conclusions were based upon a high percentage of plants having been collected with the winter buds still attached. Muenscher (1938), however, has shown that for Potamogeton pusillus [var. tenuissimus], at least, and possibly for other Pusilli, the young seedlings are very similar to a winter bud and these could easily be mistaken for old winter buds. STEM: The stems of the Pusilli are erect to decumbent or prostrate, depending upon the water depth and rate of water flow, and are round to compressed in cross section. Ogden (1972) has suggested that no species of the Pusilli from New York possess a rhizome. This statement implies that the stem arises from a winter bud or a similar struc- ture formed from the seedling as illustrated by Muenscher (1936). This situation probably represents the majority of the cases. However, some taxa, such as Potamogeton hillii, may produce a rhizome if the stem becomes decum- bent and subsequently becomes covered by debris. Rooting at the nodes may then occur, and a rhizome-like structure several centimeters in length would be formed. Ogden (1966), in fact, illustrated a rhizome of P. foliosus from Texas. Slight variation in the shape of the stem in cross-section and in the number and sizes of ridges along the stem has been observed. Certain species, e.g. Potamogeton foliosus and P. friesii, may have compressed stems, whereas other species, e.g. P. pusillus and P. clystocarpus, usually have terete stems. However, at certain times the stem of all species may be slightly compressed. Therefore, stem char- acters used for identification are helpful only as supple- mental evidence. 1974] Potamogeton — Haynes 569 Raunkiaer (1903), Hagstrom (1916), and Ogden (1943) have utilized stem anatomy for distinguishing certain spe- cies of other sections of Potamogeton. However, the stem anatomy of the various species of the Pusilli is so similar that this character is of little or no use to separate the species. At the nodes of some species, a pair of translucent oil glands may be present; in other species, they are extremely uncommon. The glands range in color from white, cream, gold, brown, or green, and in diameter from 0.1-1.0 mm. When present, by variations in color and size, the glands can be useful supplemental evidence. As with the lacunae, their presence (or absence) should not be relied upon abso- lutely, as they are apparently a physiological response to unknown ecological factors. LEAF BLADES: The blades of the Pusilli are rufes- cent, green, pellucid, to nearly black, setaceous to linear, non-lunate, sessile, stipulate, and acute to obtuse or apicu- late. Venation is parallel with 1-9 veins (nerves). The lateral nerves join the midrib at or just below the apex. In some species, one to several rows of lacunae parallel the midrib. To my knowledge, no one has studied the develop- ment or structure of the lacunae of Potamogeton. Singh (1964) studied the vegetative structures of various species of Potamogetonaceae, including P. berchtoldii [P. pusillus var. tenuissimus], which is characterized by having lacunae bordering the midrib. If Singh noticed any lacunae, he made no mention of them. Fernald (1932) suggested they are rows of nearly empty and colorless cells, presumably giving buoyance to the leaves. To my knowledge, the origin and functions of the lacunae are not known. However, regardless of their origin, the lacunae appear as light lines on each side of the midrib. The individual rows can be observed with 10X magnification, which would indicate that each row is several cells in width, rather than one cell wide. The lacunae are useful for identification, as some taxa rarely possess them, whereas others commonly have 1-. 570 Rhodora [Vol. 76 several rows on each side of the midrib. This character usu- ally serves as good supplemental evidence when used in con- junction with other features, preferably reproductive ones. Recognition of many taxa of Potamogeton, especially of the Pusilli, has been based upon slight morphological dif- ferences in the leaves (Hagstrom, 1916; Fernald, 1932). Dandy and Taylor (1940) suggested that, due to different ecological conditions, the vegetative variability of the spe- cies is too great to be useful as the only criterion for naming a new taxon. During the present study I have ob- served that several populations undergo a seasonal di- morphism, e.g., producing two morphologically dissimilar sets of leaves in one growing season. Often, plants of Potamogeton pusillus var. tenuissimus produce in the early summer a set of obtuse leaves that are near the limits of width for the species and which possess four to five rows of lacunae on each side of the midrib. Later in the growing season, these leaves die and a new set is formed. The leaves of this second set are usually acute, quite narrow, and pos- sess only one to three rows of lacunae. If one were to follow Fernald's (1932) taxonomy of this group, one could correctly identify the plant as one variety in the early sum- mer and another variety in the fall. The cause of this sea- sonal variation is unknown. Since the same genotype may produce two sets of leaves, each morphologically distinct, these vegetative characters should be considered with cau- tion when attempting to distinguish taxa. STIPULE: Associated with each leaf blade and sur- rounding the stem of the Pusilli is a tubular or convolute stipule. The stipules are free from the leaf blades and vary in color from white to brown to green. The number and coarseness of the veins varies from one species group to another. This vein character, along with the stipule color, can be useful for identification of species. In some taxa, e.g. Potamogeton friesii, the stipules are white with many coarse veins, With age the interveinal tissue near the apex decays leaving the coarse veins extending as fibers past the 1974] Potamogeton — Haynes 571 remaining interveinal tissue. In other species, e.g. P. pusil- lus, the stipules are usually brown to green with only a few veins, these being quite delicate. With age, the vein tissue decays along with the interveinal tissue. Hagstrom (1916) considered the tubular versus convolute condition of the stipules to be of great evolutionary signifi- cance. He, in fact, divided the Pusilli into two series — Pusilli Convoluti and Pusilli Connati — based on no char- acters other than the connation of the stipules. Later workers, Dandy and Taylor (1938), Ogden (1966), and Voss (1972) have accepted this character as important for determining species but have not accepted its use in sepa- rating the subsection into two series. For one of the species complexes in which the stipule character is considered most useful, according to Hagstrom and Fernald, — that being P. pusillus and P. panormitanus [P. pusillus var. tenuissi- mus and var. pusillus, respectively], I have compared the character with other vegetative and reproductive characters considered important by these workers. The data obtained from this comparison indicate only about a 70-80% corre- lation of this character with the other characters. In my opinion, when the situation is either the presence or ab- sence of a character, for that character to be given such high value in a classification system, there should be little variability of that character within a taxon. WINTER BUDS: The winter buds (turions) are veg- etative reproductive structures that function as agents for multiplication of the individuals and as perennating organs during the unfavorable season. Yeo (1966) planted one winter bud of Potamogeton crispus on 1 Apr. 1963. At the end of the growing season, 23,520 winter buds had been produced. This is obviously a very effective method of propagule production. In some species, the winter buds are produced early in the growing season and may, there- fore, germinate to produce new plants that year. In other species they are produced only at the end of the season, and do not germinate until the following year. 572 Rhodora [Vol. 76 The winter buds consist of a short stem apex with short- ened internodes. The leaves of the structures can be di- vided into two types: outer and inner. The outer leaves usually resemble the vegetative leaves, but the inner leaves may not. The inner leaves may be numerous, shortened, and oriented in a plane perpendicular to that of the outer leaves, thus giving a fan-shaped appearance, as in P. friesii. On the other hand, the winter buds may be few to several, compacted, and rolled into a hardened structure, thus ap- pearing fusiform, as in P. pusillus. Finally, they may be various in number and similar to the vegetative leaves, as in P. obtusifolius. As indicated above the winter buds may be useful as characters for the identification of the Pusilli. Hagstrom (1916) and Fernald (1932) have placed emphasis upon the location on the plant of the structures. I consider the loca- tion of the structure too variable to be of taxonomic value. Instead, the size of the structure and the modification of the inner leaves are of significant taxonomic value. INFLORESCENCE: The inflorescence of the Pusilli is either a capitate or cylindric spike consisting of from one to five whorls of four flowers each. Some authors have considered this structure to consist of a reduced compound spike (see comments under discussion of the flower). The inflorescence is a very good character for distinguishing species of the Pusilli. Important features are the length, shape, and number and separation of the whorls of the inflorescence. In some taxa, e.g. Potamogeton pusillus var. pusillus, with an inflorescence of more than three whorls, the whorls will usually be separated; however, in other taxa, e.g. P. foliosus, the whorls are rarely separated. PEDUNCLES: The peduncles of the Pusilli are either terminal on the stem or axillary to vegetative leaves. Often, axillary peduncles are reflexed, whereas the terminal ones usually are erect. In longitudinal section, the peduncles either have parallel sides or are clavate. In cross sections, 1974] Potamogeton — Haynes 573 they range from nearly terete to so compressed that they are three to four times as broad as high. The peduncle is often a very good character for dis- tinguishing taxa of the narrow-leaved pondweeds. In Potamogeton foliosus for example, the peduncle is short, clavate, usually reflexed, and axillary. In P. pusillus var. pusillus, on the other hand, the peduncle is elongate, paral- lel-sided, erect, and usually terminal. FLOWER: The flowering structures of Potamogeton have been interpreted variously. First, Eames (1961) in- terpreted each perianth segment as a sepaloid bract sub- tending and adnate to the corresponding stamen. He, therefore, regarded the flower of Potamogeton as an in- florescence comprising four staminate flowers, each of a single stamen and its adnate bract, and four apetalous carpellate flowers, each of a single carpel with one cam- pylotropus ovule. By this interpretation, the spike repre- sents a reduced compound inflorescence. Second, based upon vascular anatomy and floral morphology of seven species of Potamogeton, Singh (1965) chose to consider the flower of Potamogeton as a normal flower with four perianth seg- ments. Finally, Sattler (1965) considered it impossible to classify the structure as a true flower or as an inflorescence, because it displayed characters of both. He questioned the classical use of flower and inflorescence as two separate categories and emphasized the need for more developmental work in the “Helobiales” (an order that is now considered to be the Alismatidae). For the sake of convenience, Singh's concepts are followed in this revision. Controversy has also existed as to the real nature of the perianth-like structures that are adnate to the stamens. Ascherson (1889) and Rendle (1930) considered the struc- tures to be outgrowths of the stamens. They then called the structures stamen connectives. Sattler (1965) and Singh (1965) have reached a different conclusion. Sattler, for example, demonstrated in Potamogeton richardsonii that these structures are initiated on the floral apex prior 574 Rhodora [Vol. 76 to stamen initiation. Thus, they cannot be outgrowths of the stamens. He instead considered them to be perianth segments. After inception of the stamen primordia, growth occurs between the perianth primordia and that of the stamen. Thereby, the base of the developing stamen and perianth become united. The perianth is usually light green to greenish brown in color and consists of two parts — a claw and an expanded upper portion — each of approxi- mately equal length. The gynoecium consists of four separate, superior, uni- ovulate carpels with parietal placentation. The androecium consists of four bithecate stamens. Anthesis usually is approximately at the time of embryo sac maturity (Law- rence, 1951). Among the various taxa of the Pusilli, only slight mor- phological variation exists in the flowers. These structures, therefore, are of little value for distinguishing species. However, the size of the structures often is quite useful for separating various subsections of the genus. FRUITS: The fruits of the pondweeds have been vari- ously classified — e.g., as a drupe (Reichenbach, 1845), as a drupelet or drupaceous (Morong, 1893; Fryer and Ben- nett, 1915), as a drupe or an achene (Clapham et al., 1962) and as a druplet or nutlet (Lawrence, 1951). The use of such an array of terms suggests that no one term could adequately describe the fruits. Although the fruits resemble an achene in size and by having a thin, often leathery meso- carp, they do not conform with the definition of an achene in that the seed coat is not adnate to the pericarp, and under natural conditions the fruits are rarely ever dry. Muenscher (1936), in fact, has shown that when the fruits are subjected to drying, a very high rate of mortality oc- curs, The fact that the fruits rarely dry also rules out the possibility of their being a nut or nutlet. According to Aalto (1970) the fruits are histologically drupes since the peri- carp is differentiated into an exocarp, mesocarp, and stony endocarp. The fruit is, however, unlike a drupe because 1974] Potamogeton — Haynes 575 the dorsal area of the pericarp wall appears to open and then be covered by a lid (Aalto, 1970). At the junction of the lateral walls and the lid, a ridge is formed. This ridge, in many species, is pronounced into two lateral wings (one on each side of the fruit). The lid can be removed exposing the seed coat. Because of the histological similarity, Aalto chose to call the fruits of the Fennoscandian species drupes. Until a better term is proposed, I am considering the fruits drupe-like. The fruits are the single most important character for distinguishing the taxa of the Pusilli. Several fruit char- acters are useful for identification: 1) color — usually brown or green; 2) size; 3) shape — whether widest above the middle, at the middle, or below the middle; 4) presence of dorsal or lateral wings or ridges — one species, P. folio- sus, has a dorsal wing, three others two lateral and one dorsal ridge, others have no ridges; 5) amount of depres- sion on the sides; 6) length and position of the style (or beak). Aalto (1970) has demonstrated that for some species of Potamogeton the endocarp can be useful for identification. The majority of his work was done with subfossil endo- carps. He also exposed recent endocarps by boiling the fruits in 5% potassium hydroxide solution for 10 to 15 minutes. The exocarp and mesocarp disintegrate leaving the endocarp exposed. These were examined for shape and size of the cells and for shape, size, and appendages of the endocarp. VEGETATIVE DIMORPHISM Vegetative morphology has been used as a basis for sep- aration of certain taxa of Potamogeton. In Fernald’s study (1932), P. pusillus was divided into six varieties based on shape of the leaf apex and number of lacunae bordering the leaf midrib. The species was divided into two informal groups using the shape of the leaf apex. These groups then were further divided into the varieties with the num- 576 Rhodora [Vol. 76 ber of lacunae being the only character used for separa- tion. During the summer of 1878, E. S. Miller made two collections of P. pusillus from presumably the same locality of the Wading River, Suffolk County, New York: 1) sterile plants on 25 May 1878, and 2) fertile plants on 12 Jul. 1878. A sterile and fertile plant together were mounted on a herbarium sheet and these sheets were distributed. I have seen six such sheets and on every one the sterile plant had leaves with obtuse apices and two to three rows of lacunae each side of the midribs. Leaves of the fertile specimens, on the other hand, were acute and lacked lacunae. Fernald examined some of the specimens and consistently identified the sterile plants as P. pusillus var. mucronatus and the fertile plants as P. pusillus var. tenuissimus. This situation led me to consider the hypothesis that the taxonomic char- acter of the leaf apex and lacunae were environmentally controlled. I, therefore, began visiting populations several times throughout one year and, often, in subsequent years to see if any morphological changes occurred. Data gath- ered from these field studies indicate that P. pusillus and, also, P. friesii may undergo a dimorphism in which two morphologically dissimilar phenotypes may be observed in the same population. During the summer of 1969, I made a collection of Pota- mogeton pusillus (Haynes 3251) from Pelee Island, Ontario. These specimens had obtuse leaves with five to six rows of lacunae on each side of the midrib (Fig. 1). The popula- tion was visited separately by Dr. Ronald L. Stuckey and Mr. Marvin L. Roberts, both of The Ohio State University, several times during the next two years. In late August 1971, the population was found in fruit (Roberts 1562). The leaves from this collection were acute and had at most two rows of lacunae each side of the midrib (Fig. 1). Therefore, plants of this one population could be correctly identified, according to Fernald (1932), as var. mucronatus when sterile and var. tenuissimus when fertile. I also have observed this phenomenon in a population from New York (Haynes 3326). Here both types of leaves were found on 1974] Potamogeton — Haynes 577 the same plant. The obtuse, lacunate leaves were black and appeared dead at the time of collection and the acute, non- lacunate leaves were green and alive at the time of collec- tion. During the course of herbarium work, I have exam- ined many specimens in which both types of leaves were present on one plant, usually with the obtuse, lacunate ones appearing to have been dead at the time of collection. These data appear to indicate that plants with obtuse lacu- nate leaves are always sterile. This is not the case. Some are fertile; however, as compared with the number of fer- tile plants with non-lacunate, acute leaves, the former are few in number. A similar situation can be observed in Potamogeton fries. However, rather than two distinct sets of leaves being produced by one plant, apparently only one set of leaves is produced a year, but these leaves undergo a mor- phological change during the growing season. According to Voss (1972), P. friesii tends to have leaves without revolute margins, whereas P. strictifolius normally has ones with revolute margins. Data gathered during my field work indicate that in the early summer P. friesii does have leaves without revolute margins (Fig. 1). A popula- tion in Emmet County, Michigan, visited by me on 15 Jun. 1971 (Haynes 3694) had such leaves. However, on 30 Jul. 1971, I again visited the same population and found most plants with leaves having revolute margins (Haynes 3747). Thus, one population may produce both phenotypic expres- sions in one year. The two examples of vegetative dimorphism outlined above illustrate the vegetative plasticity of Potamogeton explained by Dandy and Taylor (1940). These data indicate that within the Pusillii and probably other groups of aquatic plants, the effects of differing environmental con- ditions upon the vegetative characters should be under- stood before one uses these characters in a classification. 578 Rhodora [Vol. 76 A B e D E K^ a» F Fig. 1l. Potamogeton pusillus leaf (A-D). A. Midsection, early summer. B. Apex, early summer. C. Midsection, late summer. D. Apex, late summer. Cross-section of Potamogeton friesii leaf (E-F). E. Early summer. F. Late summer. 1974] Potamogeton — Haynes 579 REPRODUCTIVE ECOLOGY REPRODUCTION: The reproductive ecology of Pota- mogeton is poorly known. According to Sculthorpe (1967) certain species of Potamogeton reproduce by rhizome frag- ments, tubers, and winter buds or turions. Also, Moore (1915) studied seed germination of Potamogeton and con- cluded that once a seed germinates and develops into a mature plant, several periods of vegetative reproduction must occur before the plant will produce flowers and fruits. In the Pusilli, Hagstrom (1916) and Fernald (1932) were of the opinion that the plants rarely reproduced by seeds, but instead, reproduced mostly by winter buds. For the species in which winter buds are rare or unknown, this idea of reproduction by winter buds may not be valid. However, for the species which prolifically produce winter buds, reproduction by these structures is probably the most common type, since fruiting plants often are found with winter buds attached to the underground parts. POLLINATION: According to Sculthorpe (1967) and to Cronquist (1968), the pollination trend in the Alismati- dae is from insect to wind, and ultimately, to water pollina- tion. Members of the genus Potamogeton including the Pusilli are, for the most part, wind pollinated (Sculthorpe, 1967). However, according to Voss (1972) and data based on personal field experience, many inflorescences of the Pusilli never become emergent. In such instances, if pol- lination does occur, the medium for the transport of the pollen must be the water. However, the mature, viable fruits might be formed as a result of apomixis. GERMINATION: As shown by Muenscher (1936), the germination rate of Potamogeton fruits is greatly reduced when the fruits are subjected to drying. Many species, according to Muenscher, require near-freezing tempera- tures for a period of one to three months. However, as shown by Sullivan (1967) for many of the broad-leaved species of Potamogeton, the fruits can be induced to germi- 580 Rhodora [Vol. 76 nate without a previous cold treatment if the exocarp and mesocarp are split, thus exposing the endocarp. As stated earlier, the fruits of the pondweeds are among the most common foods of waterfowl. Often only the exo- carp and mesocarp are digested, leaving the endocarp with the intact seed to be passed from the digestive system. Lohammar (1954) has shown that a high percentage of germination may occur after the fruits have passed through the digestive system of the waterfowl. CYTOLOGY Several workers, e.g. Palmgren (1939), Harada (1942; 1956), Lóve (1954), Lóve and Lóve (1961), and Stern (1961), have studied Potamogeton cytologically. However, according to Bolkhovskikh et al. (1969), these studies have produced chromosome counts for only 50 of the approxi- mately 100 species of the genus. Stern (personal commu- nication) indicates that the pondweeds are difficult to collect for meiotic material since the plants undergo meiosis quite rapidly in very early bud stage. The above-mentioned factor added to the fact that the aquatic environment is a difficult one for field work probably are the major reasons that few cytological studies have been undertaken on the pondweeds. During the summers of 1970, 1971, and 1972, I collected flower buds with the intention of obtaining chromosome counts from meiotic material. The buds were preserved in the usual manner with a 3:1 solution of ethyl alcohol and acetic acid. In some instances, the buds were examined within two or three days of collection; in others, the buds were examined after several months. In all in- stances I failed to have success in obtaining countable meiotic configurations. Therefore, all counts included in this paper are based upon previous reports in the literature. According to Stern (1961), the base number for Pota- mogeton is x — 13 or 14. The majority of the chromosome numbers reported for the floating-leaved species are multi- ples of these figures; e.g. P. amplifolius Tuckerm., n — 26; P. gramineus L., n= 26; P. illinoensis Morong, n = 52 1974] Potamogeton — Haynes 581 (Stern, 1961). On the other hand, most reports for the submersed-species indicate an n number of 13 or 14. A summary of the reported chromosome numbers for the Pusilli is included in Table 1. These reports of chromosome numbers in Potamogeton should be considered with caution, as the nomenclature has been considerably confused and most workers, excluding Stern (1961) and Taylor and Mulligan (1968), did not cite voucher specimens from which the counts were taken. TABLE 1 Published chromosome numbers for Potamogeton subsection Pusilli. Taxon Chromosome number P. berchtoldii Fieber n = 13 (Taylor & Mulligan, 1968) [P. pusillus var. tenuissimus] 2n — 26 (Taylor & Mulligan, 1968) P. foliosus Raf. 2n — 28 (Stern, 1961) P. groenlandicus Hagstrom 2n = 26 (Jørgensen, et al., 1958) P. mucronatus Schr. 2n — 26 (Palmgren, 1939) [P. friesii] P. obtusifolius M. & K. 2n = 26 (Palmgren, 1939) P. panormitanus Biv. 2n — 26 (Palmgren, 1939) [P. pusillus var. pusillus] P. strictifolius Ar. Benn. 2n — 52 (Lóve, 1954) PHYLETIC RELATIONSHIPS GENERIC RELATIONSHIPS: According to Cronquist (1968), Potamogeton is classified in the Class Liliatae (monocots), Subclass Alismatidae, Order Najadales, and family Potamogetonaceae. He considers the Alismatidae to be “a near-basal sidebranch, a relictual group which has retained a number of primitive characters." The group 582 Rhodora [Vol. 76 is, therefore, not considered to be in the main line of evolu- tion of the monocots. In studying the relationships of the Potamogetonaceae with other families of the “Helobiae,’ Chrysler (1907) came to the conclusion that Potamogeton was the most primitive genus of the group and that Potamogeton-like organisms gave rise to most of the other genera. He con- sidered the floating-leaved species, especially P. pulcher with its submersed leaves very similar to its floating leaves, to be more primitive than the submersed-leaved species. The submersed species were assumed to have been derived from the floating-leaved species as “a stage in the assump- tion of the aquatic life by the genus." One would conclude that Chrysler considered the Helobiae (Alismatidae) to have been derived from terrestrial species. Cronquist (1968), on the other hand, considered the primitive monocots to have been derived from a Nym- phaeales-like dicot, thus aquatic in origin. He considered the typical parallel-veined leaf of the monocot to be a modified, bladeless petiole. From this modified, flattened petiole, the expanded blade of, say Sagittaria, could have been derived by the spreading of the veins farther apart near the tip of the petiole. By applying this concept to Potamogeton, one could infer that the floating leaves might have been derived from the submersed leaves by an increase in the amount of tissue between the veins near the tip of the petiole. This would imply that possibly the floating- leaved species were derived from some ancestral stock of submersed-leaved plants. Published cytological data (cf. cytological section) sup- port this interpretation. The submerged-leaved species are, for the most part, diploid, whereas the floating-leaved spe- cies are, for the most part, tetraploid. Although Raven and Thompson (1964), DeWet (1965), and Anderson (1972) have demonstrated instances of polyhaploidy, based upon our present interpretation of cytological data as evidence for evolutionary relationships, the published chromosome 1974] Potamogeton — Haynes 583 numbers indicate that the primitive condition probably was that of total vegetative submergence. As stated earlier, pollination in the Alismatidae is of three types — insect, wind, and water. According to Cron- quist (1968), the reduction of the perianth in the Najadales reflects the abandonment of insect pollination, with water pollination, as in Najas and Zostera, being the most special- ized type. One would suspect, therefore, the primitive species of Potamogeton to be wind-pollinated. The water- pollinated species, then, would have been derived from some wind-pollinated ancestral stock. Again, this interpretation is supported by cytological data. Chromosome number reports for the P. filiformis subsection are the highest of all Potamogeton. This group is the only one thought to be totally water-pollinated. Palmgren (1939) reports numbers of 2n — ca. 66 for P. filiformis Pers. and 2n — ca. 88 for P. vaginatus Turcz. Based upon the scanty cytological, pollination, and mor- phological evidence, I here propose that the primitive spe- cies of Potamogeton were similar to P. zosterifolius in that they possessed narrow submersed leaves with many fine nerves and that they were wind pollinated. From the Pota- mogeton zosterifolius-like ancestor, several lines of differ- entiation probably occurred. One of these, featuring a reduction in vein number and retaining the wind pollina- tion, eventually gave rise to the Pusilli. As the scope of this paper is only the Pusilli, I will not postulate on the systematic relationships of the entire genus. Instead, my further comments on this subject will be restricted to the Pusilli. SUBSECTIONAL RELATIONSHIPS: Potamogeton subsection Pusilli is treated here to consist of 15 species. The species morphologically most similar to P. zosterifolius is P. friesii. The latter species, as does P. strictifolius, resembles P. zosterifolius in fruits, peduncles, inflorescence, leaves, and stipules. Potamogeton friesii, therefore, is prob- ably the most primitive species of the Pusilli. Potamogeton 584 Rhodora [Vol. 76 strictifolius, being a tetraploid, probably was derived from a P. friesii-like ancestor by polyploidy. One can postulate two main lines having developed from the ancestral Pusilli (Fig. 2): 1) dorsal and lateral keels absent; 2) dorsal and/or lateral keels present. The first line is represented by Potamogeton pusillus and P. groenlandicus. Potamogeton pusillus is widespread throughout the world, whereas P. groenlandicus is re- stricted to the coastal areas of Greenland. The latter spe- cies probably represents a population of P. pusillus that was isolated on Greenland and has differentiated from those of P. pusillus. The second line is represented by Potamogeton obtusi- folius, P. hillii, P. clystocarpus, and P. foliosus. Potamoge- ton foliosus, having a chromosome number of 2m — 28, probably represents a population which was separated from a P. obtusifolius-like ancestor by aneuploidy and has become widespread. In fact, it is so widely distributed in North America that it is difficult to draw any relationships as to its possible place of origin. However, other species of the line as well as P. strictifolius and P. friesii have very distinct distributions. These ranges will be discussed and used as supplemental evidence for the phyletic relation- ships. Potamogeton obtusifolius, P. strictifolius, and P. friesii, have distributions limited almost entirely to geo- graphical areas that were covered with ice during Pleisto- cene glaciation. These species probably survived glaciation elsewhere and migrated into the once glaciated areas fol- lowing the retreat of the ice. Apparently, species or vari- eties could have evolved in a matter of 10,000 years. One need only examine a few of the Great Lakes endemics, e.g. Tris lacustris (Guire & Voss, 1963), Cirsium pitcheri (John- son & Iltis, 1963), and Calamovilfa longifolia var. magna (Thieret, 1960). However, in comparison to these sand dunes endemics, most of the Pusilli are wide ranging cir- cumboreal species. If these Pusilli were to have evolved recently, then rapid circumpolar dispersal also would have had to occur. 1974] Potamogeton — Haynes 585 Hultén (1937) proposed that there were two main ref- ugia in North America — the Rocky Mountains and the continental shelf outside eastern America — where plants survived during continental glaciation. From these refugia, he stated, the plants probably spread in an easterly and westerly direction, respectively, toward the center of the continent. Some taxa probably survived in only one area, while other taxa possibly survived in both areas. Some of the taxa which survived in both refugia migrated until their ranges overlapped; thus, they now have a continuous range across North America. For others, however, migra- tion ceased before the ranges overlapped. Hultén conceded that species surviving in other areas south of the glacial boundary did migrate north to some extent into the once glaciated lands. However, he suggested that a much smaller proportion of the species now found in the glaciated areas survived in the vast areas south of maximum glaciation than in the two other refugia. Iltis (1965) and Stuckey (1972) have suggested that cer- tain species of Gentianopsis and Rorippa, respectively, sur- vived glaciation in the Rockies and migrated eastward following the retreat of the Wisconsin Ice sheet. The dis- tributions of the taxa discussed by these two workers are similar to that of Potamogeton obtusifolius, P. strictifolius, and P. friesii in that all have a more or less continuous range across northern North America and are mostly known from south of the glacial boundary only in the Rockies. Marie-Victorin (1938) proposed an explanation of the east- ward migration of such species by suggesting that ‘the unforested belt that must have existed along a receding icefront [served] as a kind of sidewalk extending from the Rockies to the Gulf of St. Lawrence." He cited no pond- weeds which demonstrated such distributions, but Stuckey (1972) indicated that P. friesii and P. strictifolius have ranges similar to a distribution pattern that Marie-Victorin probably had in mind when proposing this idea. At the time of Hultén's article (1937) Potamogeton ob- tusifolius from western America was represented by few 586 Rhodora [Vol. 76 collections, all from localities in glaciated areas. This exiguous number of specimens from western America led Hultén to regard the taxon as a cireumpolar species which survived in the eastern refugium. However, recent collec- tions indicate that the taxon is widespread, although uncom- mon, throughout this geographical area, including the Rocky Mountains. Therefore, I include P. obtusifolius as a taxon which survived continental glaciation in the western ref- ugium and then migrated eastward. Potamogeton hillii, on the other hand, survived conti- nental glaciation in eastern America just south of the glacial boundary, in the area of Pennsylvania, and migrated north and west with the retreat of the ice. In fact, the morpho- logical similarity of the species with P. obtusifolius would indicate that prior to glaciation, the two currently recog- nized taxa were perhaps represented by one large hetero- geneous population. Glaciation would have separated this large population into two smaller allopatric ones, one to the east and one to the west. During glaciation, the two populations differentiated, resulting in each population evolving into a different species. Once the period of glaci- ation was complete and the distributions overlapped, the reproductive barriers that had been established during glaciation, evidently now prevents gene flow from one popu- lation to the other. Iltis (1965) indicated that Gentianopsis procera and G. crinita possibly had a similar history. Sim- ilarly, P. clystocarpus is known from only one locality in Jeff Davis County, Texas. This species probably represents a population of P. obtusifolius in the southern mountains which was isolated and has differentiated from the parental population. TAXONOMY Of the more than 60 names that have been proposed for the North American Potamogeton subsection Pusilli, most of these are based on differences in venation, texture, color, lacunae number, and apex shape of the leaves. Dandy (1937), Dandy and Taylor (1940), and Clapham et al. 1974] Potamogeton — Haynes 587 groenlandicus var. pusillus P. pugtllus ar. tenuissimus ar. gemmiparus P. friesii-like ancestor friesii tricttfolius Lystocarpus obtueifolius hillii var. foliosus P. foltosu ar. fibrillosus Fig. 2. Hypothetical phylogenetic relationships of North Ameri- can Pusilli. (1962) have shown that the leaf form and anatomy of Potamogeton may vary widely with age, water depth, cur- rent speed, nutrient supply, light intensity, and perhaps other factors. Data from classical and experimental studies of other genera of aquatic vascular plants indicate that this 588 Rhodora [Vol. 76 heterophyllous condition is common among the submerged species. Fassett (1951) illustrated no fewer than ten quite distinct individuals of Callitriche heterophylla and drew attention to the close morphological similarities of ecological forms which actually belong to different species. Bostrack and Millington (1962) found in Ranunculus flabellaris that the shape of the leaf may be affected by temperatures, pho- toperiod, light intensity, and submergence acting directly upon the distribution and frequency of cell division in the primordium so that the inherent zones of growth initiated on the primordium are modified in their development. Cook (1966), in fact, after recognizing 20 taxa of Ranun- culus subgenus Batrachium listed 302 names which applied to these taxa. Williams (1970) found that by transplanting individuals of Nymphaea tuberosa to localities where N. odorata occurred, the individual would lose its “tuberosa” characters and obtain those of N. odorata. For reasons just outlined, these workers have found it necessary to compile a classification based primarily on reproductive characters. My approach to the systematies of Potamogeton subsec- tion Pusilli has been primarily that of combined herbarium and field studies. The data presented are based on three summers field work in northern United States and southern Canada and an examination of over 8000 herbarium speci- mens from the following 22 herbaria (Acronyms from Lan- jouw and Stafleu, 1964) : C, CAN, CGE, DAO, F, GH, ILL, K, LAF, MICH, MO,MU, NY, NYS, OS, PH, S, UC, UMBS, UPS, US, W. Evidence from the field work has indicated that the Pusilli are as morphologically plastic as Callitriche and Ranunculus. Just as workers in these other genera have resorted to fruiting characters as a basis of a classification, I think that a classification based for the most part upon the reproductive structures is the only one which would be workable. Therefore, one should always attempt to collect flowering or fruiting specimens (preferably fruiting) ! Just as identification of a sterile Aster or Solidago is nearly impossible, it is difficult at best to identify a sterile pond- weed (and then one can never be sure of his determination). 1974] Potamogeton — Haynes 589 I have thus refrained from mapping any specimen in which the determination was questionable. For one to completely adhere to the biological species concept, he should demonstrate, either experimentally or naturally, if hybridization between two taxa does occur. However, in genera such as Potamogeton, in which the flowers are so small as to render difficult the artificial cross- ing of individuals, one hardly can demonstrate whether two individuals are capable of crossing and producing fertile offspring. Ogden (1943) has suggested that the broad- leaved species of Potamogeton hybridize quite readily. He, however, did not demonstrate this phenomenon experi- mentally; rather he depended wholly upon the hybrid in- dividuals possessing characters intermediate between those of the two putative parents. With the Pusilli, however, it is not so easy. Vegetatively, the condition one would expect to find a hybrid, the species of the Pusilli are quite similar; thus, intermediates would be difficult to detect. Therefore, the techniques used by Ogden are not applicable to the Pusilli. Because of the difficulties outlined above with hybridiza- tion, I have chosen to stress a morphological species con- cept. I have followed the premise that species should be separated by discontinuities between the morphological ranges of the fruits, peduncles, inflorescence, and, to a lesser extent, the winter buds. Therefore, for the 60 names which have been applied to the North American Pusilli, I am accepting only eight species, with one of these, Potamogeton foliosus, being represented by two varieties and another, P. pusillus, being represented by three varieties. In an attempt for consistency, I have regarded varieties as mor- pho-geographic subdivisions of a species (Kapadia, 1964) that presumably reflect genetic differences. As will be noticed, I do not recognize subspecies or forms. It is my contention (as that of Raven, 1969) that only one infra- specific unit should be recognized. I have chosen to use variety over subspecies because of tradition in eastern North America. 590 Rhodora [Vol. 76 In the treatment that follows, keys are based upon fruit- ing material. Dimensions of leaves are taken from the fully expanded, longest leaves of a specimen. Measurements of the width were taken approximately at the widest point of the leaf. Dimensions of the spike and peduncles are based upon both flowering and fruiting specimens. Descriptions of the fruits are taken strictly from mature structures. POTAMOGETON Linnaeus, Species Plantarum 126. 1753 Hydrogeton Loureiro, Flora Cochinchinensis 244. 1790. Type Species: Hydrogeton heterophyllum — Potamogeton octandrus Poir. Patamogeton Honckeny, Syn. Plan. Germ. 2: 110. 1793. (orthographic variant). Potamogiton Rafinesque, Med. Repos. 5: 354. 1808. (or- thographic variant). Potamogetum Clairville, Man. d'Herb. Suisse & Valais 34. 1811. (orthographic variant). Peltopsis Rafinesque, J. Phys. Chim. Hist. Nat. Arts 89: 102. 1819. Type Species: Potamogeton perfoliatus. L. Spirillus J. Gay, Compt. Rend. Hebd. Seances Acad. Sci. 38: 703. 1854. (name without any listed species) ; emend. Nieuwland, Amer. Midl. Naturalist 3: 14. 1913. Type Species: (lectotype here designated) Potamogeton diversi- folius Raf. Plants herbaceous, aquatie, submerged in fresh or rarely brackish water, annual or perennial, propagated from seeds, winter buds, or rhizomes. Stems variable in length accord- ing to water depth, branched or unbranched, terete or com- pressed, rooting at the nodes. Leaves all submersed or both submersed and floating, alternate or subopposite; sub- mersed leaves pellucid, sessile or petiolate, linear to orbicu- lar, subulate to obtuse at the apex, the margins entire to serrate, rarely crimped, the nerves 1-35; floating leaves coriaceous, mostly petiolate, rarely subsessile, elliptic to 1974] Potamogeton — Haynes 591 ovate, acute to obtuse at apex, cuneate to rounded or cor- date at base, the margins entire, the nerves 3-51. Stipules tubular, sheathing the stem and young inflorescences, con- nate or convolute, either free or adnate to the base of sub- mersed leaves, free from base of floating leaves. Winter buds present or absent, with extremely shortened nodes, divided into inner and outer leaves; inner leaves few to numerous, either shortened and oriented at 90? angles with respect to outer leaves, rolled into a fusiform structure, or unmodified; outer leaves 1-5 per side, mostly similar to vegetative leaves, rarely corrugated near base. Inflorescence a capitate or cylindrie spike with 1 to 20 whorls of flowers, compact or moniliform, with 2 to 4 flowers each whorl, mostly buoyed above surface of water. Flowers bisporangi- ate, Perianth of 4, free, rounded, short-clawed, greenish segments. Androecium of 4 stamens: filaments adnate to the perianth claw; anthers bithecate, extrorse. Pollen spherical monoaperturate. Gynoecium of 4, free, unilocular, uniovulate carpels ; ovule campylotropus; placentation pari- etal. Fruit drupe-like; dorsally rounded or keeled ; embryo coiled; cotyledon one, endosperm absent. Chromosome base number: z — 13 or 14 (Stern, 1961). (Name from the Greek potamos, a river and geiton, a neighbor.) Type Spe- cies: Potamogeton natans L. (fide Taylor, 1909). POTAMOGETON Subsection PUSILLI Graebn. in Aschers. & Graebn. Das Pflanzenreich 4(11): 260. 1907. Plants submersed in fresh water, annual or perennial. Stems branched or unbranched, terete or compressed. Leaves all submersed, pellucid, sessile, linear, subulate to obtuse at apex, entire, 1-9-nerved. Stipules connate or con- volute, free from the base of the leaves. Winter buds pres- ent or absent. Inflorescence mostly emersed, a capitate or cylindric spike with 1-5 whorls of flowers, compact or moniliform, mostly with 4 flowers at each whorl. Fruit dorsally rounded or keeled, to 4.0 mm long. Chromosome base number x = 13 or 14. Type Species: Potamogeton pusillus L. 592 Rhodora [Vol. 76 KEY TO THE TAXA OF NORTH AMERICA 1. Leaves 7-9-nerved and inner-leaves of winter buds un- modified; plants of Greenland. .. 8. P. groenlandicus. 1. Leaves with up to 9 nerves, but if 7-9-nerved, then with inner-leaves of winter buds modified; plants from areas other than Greenland. ............. 0.000002 0 ee 2. 2. Fruits with a dorsal keel or ridge to 0.4 mm high, often with 2 lateral keels. .................... 3. 3. Inflorescence cylindric, 8 mm or more long; basal glands usually present, mostly 0.5 mm or larger in diam; leaves obtuse or apiculate, rufescent. Lecce eee ee ee ee ee eee eee 3. P. obtusifolius. 3. Inflorescence capitate or rarely cylindric, 7.5 mm or shorter; basal glands when present, mostly smaller than 0.5 mm diam; leaves acute, rarely obtuse, olive or green. .............0000005 4. 4. Peduncle cylindrie, mostly terminal, erect; basal glands present; fruit with basal tuber- cles, ....... 0.00. less 6. P. clystocarpus. 4. Peduncles usually clavate, axillary, often re- curved; basal glands uncommon; fruit with- out basal tubercles. ................00-. 5. 5. Fruits rounded on sides, 3-keeled, 3-4 mm long; dorsal keel ridge-like, to 0.2 mm high. ..............2000050. 4. P. hillii. 5. Fruits with concave sides, 1-keeled, to 2.7 mm long; dorsal keel wing-like and undu- late, to 0.4 mm high. ................ 6. 6. Spike rarely interrupted; fruit olive to green-brown, 1.5-2.7 mm long, 1.2-2.2 mm wide; keel mostly 0.2 mm high or higher; beak 0.2-0.6 mm long ; stipular veins decaying with age; basal glands rare, to 0.3 mm diam, .............. NEM 5a. P. foliosus var. foliosus. 1974] Potamogeton — Haynes 593 6. Spike mostly interrupted; fruit pale- green, 1.4-1.7 mm long, 1.1-1.2 mm wide; keel mostly less than 0.2 mm high; beak 0.2 mm long or less; stipu- lar veins with age remaining as long fibers; basal glands common, to 0.5 mm diam. .. 5b. P. foliosus var. fibrillosus. 2. Fruits with dorsal surface rounded; lateral keels IO ec ese ne eed ve ERI eee T. 7. Stipules more or less coarsely fibrous, whitish, the oldest tending to disintegrate into shreds; bases of winter-buds usually indurated and corru- gated; peduncles mostly clavate. ............ 8. 8. Leaf tips rounded to apiculate; leaves light green to rufescent, 5-7 (-9)-nerved, 1.2-3.2 mm wide; winter buds 1.5-4.0 mm wide, inner leaves modified into a fan-shaped structure, outer leaves corrugated at base; peduncles compressed; stem compressed. . 1. P. friesii. 8. Leaf tips acute, rarely obtuse, mostly deep green to olive, 3-5 (-7)-nerved, 0.6-2.0 mm wide; winter buds 0.8-2.2 mm wide; inner leaves modified into a fusiform structure, outer leaves rarely with corrugations at base; peduncles mostly terete; stem mostly terete. LOUER UERTASEEAITT RN FRA 2. P. strictifolius. 7. Stipules mostly delicate, whitish, green, or brown- ish, usually disintegrating with age; bases of winter-buds without corrugations; peduncles cy- Hndrie ees se es ce eg RE ee 9. 9. Fruits 2.5-3.6 mm long, 1.7-2.4 mm wide; winter buds 3.5-7.8 cm long, 2.3-5.1 mm wide, inner leaves unmodified. .. 3. P. obtusifolius. 9. Fruits 1.5-2.2 mm long, 1.2-1.6 mm wide; winter buds 0.9-3.2 cm long, 0.3-1.8 mm wide, inner leaves modified into a fusiform struc- INPUNE esc icem e iq EM RD 10. 594 Rhodora [Vol. 76 10. Leaves uninervate, subulate, 0.2-0.7 mm wide; plants of New England and south- ern Quebec. ............ 002 eee eee EN Tc. P. pusillus var. gemmiparus. 10. Leaves 1-5-nerved, acute to obtuse, 0.2- 2.5 mm wide; widespread throughout North America. ............. sess 11. 11. Mature fruit widest above the mid- dle, sides concave, beak central; pe- duncle filiform to cylindrical, usually 1-3 per plant; inflorescence usually of 2-4 distinct verticels; leaves with up to 2 rows of lacunae along mid- rib, apex acute, rarely apiculate; stipules mostly connate. .......... MEM (a. P. pusillus var. pusillus. 11. Mature fruit mostly widest at or be- low the middle, sides rounded, beak mostly forward, peduncles cylindri- eal, usually more than 3 per plant; inflorescence mostly of 1-2 adjacent verticels; leaves with 1-5 rows of lacunae along midrib, apex acute to obtuse; stipules mostly convolute. .. .. Tb. P. pusillus var. tenuissimus. 1. Potamogeton friesii Rupr. in Beitr. Pflanzenk, Russ. Reiches 4: 43. 1845. TYPE: (type not located), application of the name is from the illustration of Potamogeton com- pressus auct. non Linn.: In Reichenbach. Icones Florae Germanicae et Helveticae 7: pl. 24. 1845! Spirillus friesii (Rupr.) Nieuwl. Amer. Midl. Naturalist 3: 17. 1913. Potamogeton pusillus var. major Fries, Novitiae Florae Suecicae, ed. 2, 48. 1828. P. mucronatus Schr. ex Sonder, Flora Hamburgensis 99. 1851. P. major (Fries) Morong, Mem. Torrey Bot. Club 3: 41. 1893. TYPE: (type not lo- 1974] Potamogeton — Haynes 595 cated), application of the name is from the illustration of Potamogeton compressum auct. non L.: In Smith, English Bot. 2: pl. 418. 1797! Potamogeton pusillus var. latifolius Meyer, Chloris Han- overana, 525. 1836. P. oederi Meyer, Flora Hanoverana Excursoria 536. 1849. TYPE: (type not located), applica- tion of the name is from the illustration of Potamogeton compressum auct. non L.: In Oeder, Flora Danica 2: (4) t. 203. 1765! Stem pale green to pale brown, simple to profusely branched near the apex, compressed, slightly ridged, 10- 135 em long, 0.3-1.1 mm diam. Leaves usually light green, rarely olive-green to rufescent, delicate to rigid, 5-7 (-9)- nerved, 2.3-6.5 cm long, 1.2-3.2 mm wide; apex acute to apiculate, glands green, greenish-brown, or gold, to 0.7 mm diam; lacunae absent or 1 narrow row each side of the midrib; lateral nerves joining the midrib 0.2-0.9 mm below the apex. Stipules white, fibrous, shredding at apex, 5.5- 21.0 mm long, 0.7-2.5 mm diam. Winter buds common, terminal or lateral, 1.5-5.0 cm long, 1.5-4.0 mm wide; inner leaves reduced, arranged into a fan-shaped structure and oriented at right angles to the outer leaves; outer leaves 2-3 per side, apiculate to acute, corrugated at base. Pe- duncles usually slightly clavate, terminal or axillary, rarely recurved, 1.2-4.1 (-7.0) em long, (0.1-) 0.5-1.2 mm diam. Spike cylindrical, 7.0-16.0 mm long, 2.6-8.0 mm diam; ver- ticels 2-5, 1.5-5.0 mm apart. Perianth segment 1.2-1.5 mm long, 0.5-1.5 mm wide. Fruit olive-green to brown, without dorsal keel, 1.8-2.5 mm long, 1.2-2.0 mm wide; beak central, 0.3-0.7 mm long, 0.2-0.5 mm diam; sides rounded, rarely centrally depressed; wall texture smooth, rarely reticulate. Chromosome number, 2n — 26 (Palmgren, 1939). Distribution: Central Alaska to Newfoundland, south to northeastern Utah and southeastern Pennsylvania, Fig. 3. Fruiting from late June to September. Illustrations: Fernald (1932, pl. 6; 29, fig. 3; 33, fig. 1; and 39, fig. 8). 596 Rhodora [Vol. 76 Nomenclaturally, Potamogeton friesii has been poorly understood. The taxon was long referred to P. compressum L. (Smith, 1797; Oeder, 1765; Reichenbach, 1845). Fries (1828) regarded the taxon as a variety of P. pusillus and, therefore, proposed P. pusillus var. major. However, Mertens and Koch (1828) had made the same combination for a different taxon. Thus, Fries! name was a later homo- nym. The first combination at specific rank which appar- ently applied to this entity was P. mucronatus Schrader. However, to my knowledge, this name was never published by Sehrader, but first appeared in Romer and Schultes (1818) as a nomen nudum — “Qid P. mucronatus Schrader." I have been unable to locate the name again until used by Reichenbach (1845), definitely placing the name in synon- Fig. 3. Map of North America showing the documented distribu- tion of Potamogeton friesii. The barred line represents the southern limit of Wisconsin Glaciation in the United States east and west of the Rocky Mountains (after Flint, 1957; Flint, et al., 1959). 1974] Potamogeton — Haynes 597 omy under P. compressum — “P. mucronatus Schrader, R. S. III. p. 517!" Neither of these references to P. mucro- natus constitute publication, since a name is not considered validly published when it originally appears in synonomy. Apparently, the name was not validly published in specific rank until Sonder (1851) applied a description, giving credit to Schrader. However, Ruprecht (1845) had validly published Pota- mogeton friesii and accurately applied the name to the en- tity which we now consider to be this taxon. Although I have not seen any of his original specimens, he referred to the illustration cited above in Reichenbach (1845) and also to one in Oeder (1765), both of which are Potamogeton friesii, REPRESENTATIVE SPECIMENS CANADA: Alberta: Bow River Valley, Brown 692 (GH, NY, PH); Banff (Vermilion Lakes), Malte & Watson 961 (CAN, GH). British Columbia: Blackwater Lake, ca. 32 mi. N. of Golden, Hitchcock & Martin 7645 (NY, UC). Manitoba: Cross Lake, 45 mi N of Lake Winnipeg, Scoggan 3573 (CAN). New Brunswick: Tidehead, near Campbellton, Malte 784 (CAN). Newfoundland: St. Georg’s Pond, near Bay St. George, Fernald & Wiegand 2456 (CAN, GH, PH, NY). Northwest Territories: MacKenzie River delta, East branch, 68°40'N, Porsild 7243 (GH, US). Nova Scotia: GOLDCHESTER CO.: Salmon River, Truro, Bissell & Linden 19698 (GH). Ontario: HASTINGS CO.: no locality, Macoun s.n. (F, K, MO, NY, s); Charlton Island, James Bay, 52°N, 79°30’W, 4292 (CAN, US, photo at DAO). Prince Edward Island: KINGS CO.: Black Pond, Fernald & St. John 6772 (CAN, GH, K, NY, US); Cove Head, Macoun 3000 (CAN, GH, K, MO, NY). Quebec: Coffin Island, Magdalen Islands, Fernald et al. 6775 (CAN, GH, K, NY, PH, US); Ile de la Grand-Entree, Magdalen Islands, Marie-Victorin & Rolland-Germain 9921 (F, GH, NY, PH, US). Saskatchewan: Barrier River, ca. 4 mi. W, 3 mi. S. of McKague, Breitung 414 (CAN, DAO, NY, UC). Yukon: Craig Lake, vicinity of Carcross, Porsild 18418 (CAN). UNITED STATES: Alaska: vicinity of College, near Fair- banks, Argus 1097 (pao). Connecticut: NEW HAVEN CO.: Derby, Robbins (GH, NY). Idaho: JEFFERSON CO.: Mud Lake, Sperry & Martin 670 (us). Illinois: cook co.: South Chicago, Hill 79-1881 (ILL). Indiana: WHITLEY CO.: Goose Lake, ca. 7 mi. NW of Columbia City, Deam 48794 (GH). Iowa: DICKINSON CO.: East Lake Okoboji, Shimek (DAO, F, GH, MO). Maine: AROOSTOOK CO.: Washburn, Aroo- 598 Rhodora [Vol. 76 stook River, Ogden & Chamberlain 2692 (os). Massachusetts: MID- DLESEX CO.: Fresh Pond, Cambridge, Faxon (GH, NY, US). Michigan: CHEBOYGAN CO.: Black River between mouth and Alverno, Haynes 3377 (LAF, os); Haynes 3385 (Mo, 0S); Haynes 3698 (MO, NY, OS); Hook Point, Douglas Lake, Haynes 3361 (F, GH, MO, NY, OS, PH, UC, US); mouth of Bessey Creek, Douglas Lake, Haynes 3887 (os); Marl Bay, Douglas Lake, Haynes 3711 (ASU, GH, LAF, OS). EMMET CO.: small arm of Carp Lake, Haynes 3376 (os); Haynes 3694 (os); Haynes 8747 (os); Haynes 3889 (08S). MANISTEE CO.: Little Manistee River, Manistee, Hill (F, GH, ILL, MO, US). Minnesota: CASS CO.: Cullen Lake, Ballard (ILL, MICH, MO, UC, US). MARTIN CO.: Silver Lake, Cratty (GH, ILL, MO, NY, PH, UC, US). Montana: GLACIER CO.: marshes above Lower Two Medicine Lakes, Glacier National Park, Maguire 482 (GH). Nebraska: CHERRY CO.: Dewey Lake, near Valen- tine, Tolstead 477 (GH). New York: CORTLAND CO.: Upper York Lake, near Little York, Haynes 3347 (MICH, MO, OS, US); Haynes 3348 (F, GH, MO, NY, OS, PH, UC, US); Haynes 3349 (MICH, OS); Haynes 3350 (MICH, 08). ONTARIO CO.: Lake Seneca, Geneva, Morong (CAN, GH, NY, S). SARATOGA CO.: Saratoga Lake, Haynes 3311 (MICH, MO, OS). WARREN CO.: Dunham Creek near S. end of Lake George, Haynes 3330 (GH, MICH, MO, OS, UC, US); Haynes 3331 (MICH, MO, OS, US). North Dakota: cass co.: Fargo, Whitney (MU). Ohio: ERIE C0.: East Bay, Sandusky, Osburn & Williamson (GH). Pennsylvania: LEHIGH CO.: Bethlehem, Noble (GH, PH). South Dakota: FAULK CO.: ponds and rivers, near Jacques, Geyer (US). Utah: suMMIT co.: Lyman Lake, Blacksfork Creek, Hobson 52 (GH). Vermont: ADDISON CO.: Little Otter Creek, Ferrisburg, Faron (GH, US). Washington: OKANOGAN CO.: Bonaparte Lake, St. John, et al. 5298 (GH, MO, UC). Wisconsin: BROWN CO.: Point Sable, Green Bay, Schuette (F, GH, K, MICH, UC). MILWAUKEE CO.: Milwaukee, Lapham (GH, K, MO, PH). Wyoming: TETON CO.: shallow lake, 10 mi. s. of Moran, Porter & Porter 7869 (DAO, UPS, UC). 2. Potamoveton strictifolius Ar. Benn. J. Bot. 40: 148. 1902. Potamogeton pusillus var. pseudo-rutilus Ar. Benn. J. Bot. 39: 201. 1901. P. foliosus X P. rutilus? Hagstrom, Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. P. strictifolius var. typicus Fern., Mem. Amer. Acad. Arts 17: 56. 1932. Type: E. J. Hill [146], Wolf Lake, Ham- mond, [Lake Co.] Indiana, 3 Sept. 1900, (lectotype, fide Fernald, 1932, at K, but not located; photo of lectotype, CAN! NY!, isolectotype, ILL !). 1974] Potamogeton — Haynes 599 Potamogeton strictifolius var. rutiloides Fern. Mem. Amer. Acad. Arts 17: 57. 1932, P. pusillus var. rutiloides (Fern.) Boivin, Naturaliste Canad. 94: 527. 1967. TYPE: F. P. Metcalf 1453, Molly Lake, 10 mi. N of Brainerd, Crow Wing Co., Minnesota, 26 Aug. 1921, (holotype, GH!; iso- type, US!). Stem pale green to green-brown, branched or simple, more or less rounded, slighty ridged, 27-95 cm long, 0.4- 0.8 mm diam. Leaves green to olive, usually rigid, 3-5 (-7) - nerved, 1.2-6.3 cm long, 0.6-2.0 mm wide; apex acute to nearly bristle-tipped; glands white, green, greenish-brown, or gold, to 0.3 mm diam; lacunae usually absent; lateral nerves joining the midrib 0.8-1.5 mm below the apex. Stipules usually white, fibrous, shredding at tip, connate, 0.6-1.6 cm long, 0.6-2.0 mm diam. Winter buds common, terminal or lateral, 2.5-4.8 cm long, 0.8-2.2 mm wide; inner leaves undifferentiated; outer leaves 3-4 per side, acute, mostly without, rarely with, corrugations at base. Pedun- cles usually cylindric, rarely slightly clavate, terminal, mostly erect, rarely recurved, 1.0-4.5 em long, 0.3-0.9 mm diam. Spike cylindrical, 0.6-1.3 em long, 1.5-5.2 mm diam; verticels 3-4, 1.5-3.2 mm apart. Perianth segments 0.9-1.7 mm long, 0.8-1.5 mm wide. Fruit green-brown, without dorsal or lateral keels, 1.9-2.1 mm long, 1.3-1.8 mm wide; beak central, 0.3-0.5 mm long, 0.2-0.4 mm diam; sides rounded, often centrally depressed; wall texture smooth. Chromosome number, 2n == 52 (Lóve, 1954). Distribution: Northwestern Northwest Territories to eastern Quebec, south to northeastern Utah and north- western Connecticut. Fruiting from early July to late September. Fig. 4. Illustrations: Fernald (1932, pl. 7; 8; 29, fig. 4, 5; and 33, fig. 2, 3). Bennett, in the original description of Potamogeton pusil- lus var. pseudo-rutilus, cited two collections: *Lake Scugog, Ontario, Canada, 1897, W. Scott, ex Prof. Macoun;" and 600 Rhodora [Vol. 76 *Wolf Lake, Indiana, U.S.A., 1900, Rev. E. J. Hill." After receiving more material from Hill, Bennett later raised the taxon to specific rank. Hagstrom (1916) examined addi- tional material sent to him by Hill and ascertained this to be a hybrid between P. foliosus and P. rutilus. Hagstrom cited the original specimen as being from Lake George, East Chicago, rather than from Wolf Lake. I think Hag- strom made an error in citing this specimen, or possibly Lake George and Wolf Lake refer to the same lake. The original most certainly was from Wolf Lake. Hagstrom did not examine material from Lake Scugog, but based his decision on the 1902 description of P. strictifolius by Ben- nett and other Canadian material he had examined from Sea-Cow-Pond, Crane Lake, Assiniboia, Picanok River, Quebec, and Ottawa River, Quebec [which he said “seem excellently to answer to the description of the fruiting plant of Lake Scugog"]. He concluded that the Lake Scugog ma- terial was probably the same as P. panormitanous (P. pusillus var. pusillus). Fernald examined Bennett's collec- tion but overlooked the point of whether the Lake Scugog plant was the same taxon as the Wolf Lake plant. As he could not be certain that the two were the same, he did not deny or accept Hagstrom's decision. He did, however, desig- nate Hill's collection from Wolf Lake, Indiana, September 3, 1900, as the Type [lectotype] of P. strictifolius. Fernald did say, with which I concur, that Hagstrom had erred in calling P. strictifolius a hybrid of P. foliosus and P. rutilus. I also am unable to state with assurance that the speci- men from Lake Scugog is conspecific with the one from Wolf Lake. I have corresponded with curators of the herbaria (K, BM, & CGE) where the majority of Bennett's material is located. There is no sheet in Bennett's collection at these herbaria filed under Potamogeton pusillus, P. berch- toldii, or P. strictifolius. As I have photographs from CAN, NY, & US of a sheet with only the two specimens mounted on it, I am certain that it is not correctly filed under the name of another specimen on the same sheet. The photo- graph from CAN has a paragraph in Bennett's handwriting 1974] Potamogeton — Haynes 601 stating that the sheet was in his collection and that the photograph was taken by his daughter. I have, however, located a duplicate of Hill's collection from Wolf Lake, and also a second specimen collected by Seott at Lake Seugog on 3 August 1897 (the same day as the one in Bennett's collection). I can not ascertain if Scott's collection is a duplicate of the one sent to Bennett. With its convolute stipules, characteristic winter buds, and one row of lacunae along each side of the midrib, this specimen is rather characteristic of P. pusillus var. tenuis- simus. It is, however, unusual in that the stipules are white rather than the usual greenish-brown of P. pusillus var. tenuissimus. Until the specimen in Bennett’s collection can be located, I am following Fernald in that I cannot be sure to which taxon the Lake Scugog collection should be re- ferred. Fernald (1932) separated Potamogeton strictifolius into two varieties, var. strictifolius & var. rutiloids, based on the leaf apex shape, rigidity of the leaves, and the coarseness of the stipules. After examining many sheets and hundreds of plants in the field, I have concluded that this variability is both within and between populations. The leaf tips of the isotype of var. strictifolius at ILL, for example, vary from very gradually tapering acute to obtuse. Because of this variability within populations and of no apparent range distinctions, I consider P. strictifolius to be one taxon. The rigidity of the leaves, I believe to be of ecological rather than of genetic origin. Morphologically, Potamogeton strictifolius is most similar to P. friesii. The best way to distinguish the species is by the winter buds. In P. strictifolius, the inner leaves of the winter buds are only slightly modified, whereas in P. friesii the inner leaves are much shorter than the outer leaves and are oriented at 909 to the outer leaves. Differences in the leaves are difficult to assess. Fernald (1932) and Ogden (personal communication) stress the number of veins — three to five in P. strictifolius and five to seven in P. friesii. 602 Rhodora [Vol. 76 Street L0 bee. ~? vex = EN Fig. 4. Map of North America showing the documented distribu- tion of Potamogeton strictifolius. The barred line represents the southern limit of Wisconsin Glaciation in the United States east and west of the Rocky Mountains (after Flint, 1957; Flint et al., 1959). 1974] Potamogeton — Haynes 603 For both species, I find, as does Voss (1972), that the usual number is five. If specimens are sterile and lacking winter buds and I must use vegetative characters, I then rely upon the apex shape and whether the leaves are revolute. In P. strictifolius, the apex is usually acute and the leaves are revolute. In P. friesii, the apex is obtuse to apiculate and the leaves are usually flat. However, as I indicated earlier and pointed by Voss (1972) one must use these characters with caution. In the late summer the margins of P. friesii roll under, thus the leaves become revolute. REPRESENTATIVE SPECIMENS CANADA: Alberta: Large pond, Lacombe, Dixon 691 (DAO, NY). Manitoba: York Factory, Drummond (GH). Northwest Territories: MACKENZIE DIST.: E branch of Mackenzie River, 68°40’ to 68°55’, Reindeer Station, Porsild 7234 (CAN). Ontario: THUNDER BAY DIST.: Black Fox Lake, Haynes 3750 (CAN, GH, LAF, MO, 0S); Nipigon, Lake Superior, Macoun (CAN, NY, S). Quebec: GATINEAU CO.: Danford Lake, 45 mi. N of Ottawa, Dore 9032 (DAO, MO). Saskatchewan: Pike Lake, Saskatoon, Fraser 8 (DAO, GH). UNITED STATES: Connecticut: LITCHFIELD CO.: Indian Pond, Sharon, Eames 11868 (GH). Illinois: cook co.: Wolf Lake, Chicago, Chase 1709 (CGE, F, GH, ILL). Indiana: LAKE C0.: Lake George, East Chicago, Hill 144- 1903 (GH, ILL, NY). Michigan: CHARLEVOIX CO.; Cunningham Lake, ca 6 mi SW of Ironton, Voss & Haynes 13602 (MICH). CHEBOYGAN co.: Black River between its mouth and Alverno, Haynes 3379 (GH, MO, OS, US), Haynes 3384 (Mo, os, US), Haynes 3743 (GH, 0S); Black Lake, near mouth of Upper Black River, Haynes 3777 (os); Cheboy- gan River, Haynes 3798 (os); Douglas Lake near source of Maple River, Haynes 3739 (GH, MICH, MO, NY, OS, UMBS, US). WAYNE CO.: Detroit River near Belle Isle, Wheeler (GH, ILL, NY, US). Minnesota: BECKER CO.: DeSota Lake, Grant 3269 (GH, Ny, UC, US). Nebraska: CHERRY CO.: Willow Lake, Thomson 122 (Us). New York: MONROE co.: Clausen & Hinkey 4196 (GH, NY). North Dakota: MCHENRY co.: Red Willow Lake, Mabbott 372 (US). Ohio: ERIE CO.: Sandusky Bay, Pieters (US). Pennsylvania: ERIE CO.: Presque Isle, Garber (GH). Utah: BOX ELDER CO.: Bear River, Watson 1136 (GH, K, NY, US). Vermont: CALEDONIA CO.: Sarah Moor Pond, 2 mi NW of Barnet, Hotchkiss 7804 (Us). Wisconsin: WASHINGTON CO.: Big Cedar Lake, Hotchkiss & 'Hoehler 4264 (GH, US). Wyoming: ALBANY CO.: reservoir ca 6 mi. NE cf Lookout, Porter 7370 (DAO, NY, UC, USP). PARK CO.: Firehole River S of Madison Junction, Porter 6385 (DAO, MO, NY, UC, UPS). 604 Rhodora [Vol. 76 Potamogeton strictifolius X zosteriformis Fernald (1932) named Potamogeton longiligulatus based upon flowering specimens from Newfoundland. The entity was said to resemble P. hillii by its bristle-tipped leaves but differed from the latter species by having numerous- nerved leaves, Voss (1967) reported that, at least in north- ern Michigan, the entity appeared to be a result of hybridi- zation between P. strictifolius and P. zosteriformis, a spe- cies in the subsection Compressi of Hagstrom. Data gath- ered during my field studies in northern Michigan support Voss' conclusions. I have seen several instances where the putative hybrid was growing between populations of P. zosteriformis and P. strictifolius. The plants of the putative hybrid were morphologically intermediate between the two species. Therefore, until the putative hybrid can be studied in detail, I accept Voss' concept of the taxon. SPECIMENS EXAMINED: CANADA: Newfoundland: Straits of Belle Isle, pond in barrens S of Flower Cove, Fernald & Long 27330, (holotype, GH!; isotype, PH!, photo at DOA!). Ontario: BRUCE CO.: Bruce Peninsula, Stokes Bay, Krotkov 8627 (Us). LAMBTON C0.: Sarnia Bay, Dodge (GH, photo at DAO). UNITED STATES: Connecticut: LITCHFIELD CO.: Salisbury, Twin Lakes, Bissell (GH). Illinois: LAKE CO.: Grayslake, Dolbeare 1444 (MicH). Michigan: CHEBOYGAN CO.: Black River between its mouth and Alverno, Bedell & Prescott 55 (UMBS), Haynes 3380 (GH, Mo, os, us), Haynes 3381 (GH, MO, os, US), Haynes 3702 (os), Haynes 3744 (os), Majors (GH, NY, UMBS), Voss 11118 (GH), Voss 11359 (UMBS); Black Lake, bay at mouth of Black River, Stuckey 2411 (os), Stuckey 3275 (os, CAN), Voss 11745 (GH, NY), 11760 (UMBS); Cheboygan River, Haynes 3799 (OS). ST. CLAIR CO.: Algonac, Big Ditch, Dodge s.n. (US); Port Huron, Dodge s.n. (NY). SCHOOLCRAFT CO.: Lake McDonald, SE of Whitedale, Uhler 96 (Us). Minnesota: BECKER C0.: Cotton Lake, Hotchkiss 6341 (GH). HEN- NEPIN C0.: Lake Minnetonka, Keck & Stilwill 416 (GH). New York: COLUMBIA CO.: Beebe Pond near Queechy Lake, Haynes 3344 (os), Haynes 3801 (os), Smith & Ogden 45590 (os). 3. Potamogeton obtusifolius Mert. & Koch, in Róhling, Deutschland Flora 855. 1823. TYPE: (type not located), application of the name is from the illustration of Potamo- 1974] Potamogeton — Haynes 605 geton gramineus auct. non L.: In Smith, English Bot. 11: pl. 2253. 1811. Spirillus obtusifolius (Mert. & Kóch in Róhling) Nieuwl. Amer. Midl. Naturalist 3: 19. 1913. Potamogeton compressum. var. tenwius Wahl. Flora Up- saliensis 60. 1820. TYPE: G. Wahlenberg, Upsala Sraklen, Ekebysja near Quam, 22 Aug. 1818, (holotype, UPS!). Potamogeton obtusifolius var. angustifolius Fieber in Berchtold, Oekon.-tech. Flora Bohmens 275. 1838. TYPE: (not located). Potamogeton obtusifolius var. latifolius Fieber in Berch- told, Oekon.-tech. Flora Bohmens 275. 1838. TYPE: (not located). Stem green to green-brown, slightly compressed, usually without ridges, 35-90 cm long, 0.3-0.7 mm diam. Leaves light green to rufescent, usually flaccid, 3-nerved, 3.0-8.2 em long, 1.0-3.5 mm wide; apex round to round apiculate; glands yellow-green to gold, 0.2-1.0 mm diam; lacunae of 1-3 rows each side of midrib; lateral nerves joining the midrib 0.2-1.2 mm from apex. Stipules usually white, slightly fibrous, rarely shredding at tip, convolute, 0.6-1.8 em long, 0.5-1.2 mm diam. Winter buds abundant, terminal, 3.5-7.8 em long, 2.3-5.1 mm wide; inner leaves undifferen- tiated; outer leaves 3-4 per side, apiculate to obtuse, with- out corrugations at base. Peduncles cylindrical, axillary, rarely recurved, 0.8-1.9 (-4.2) em long, 0.5-1.0 mm wide. Spike cylindrical, 0.8-1.3 cm long, 4.6-7.0 mm wide; verti- cels 3, crowded or to 2.0 mm apart. Perianth segments 1.4- 1.5 mm long, 1.1-1.3 mm wide. Fruit olive-green to brown, 2.5-3.6 mm long, 1.7-2.4 mm wide; keels absent or present, when present, ridged, to 0.2 mm high; beak mostly central, rarely forward, (0.2-) 0.6-0.7 mm long, 0.5-0.7 mm diam; sides rounded; wall texture smooth or rough. Chromosome number 2n — 26 (Palmgren, 1939). Distribution: Eastern Yukon to eastern Quebec, south to Washington, Wyoming, and New Jersey. Fruiting from early July to late September. Fig. 5. 606 Rhodora [Vol. 76 Illustrations: Fernald (1932, pl. 14; 30, fig. 3; 34, fig. 4; 39, fig. 1). In its winter buds, fruit, and peduncles, Potamogeton obtusifolius most closely resembles P. hillii. When in fruit, however, it can easily be separated from the latter species by its spikes being cylindric, whereas those of P. hillii are capitate. Sterile specimens of P. obtusifolius can usually be distinguished from P. hillii by the former having obtuse leaves and the latter having acute leaves. However, plants of P. hillii occasionally possess obtuse leaves. o. uo CASH u T RRA; ` FAR AUTRES (CON, TA NES >; mw YIN AG = f 4d: Mac 2.* Jj Ys Y BANNA Fig. 5. Map of North America showing the documented distribu- tion of Potamogeton obtusifolius. The barred line represents the southern limit of Wisconsin Glaciation in the United States east and west of the Rocky Mountains (after Flint, 1957; Flint et al., 1959). h. REPRESENTATIVE SPECIMENS CANADA: Alberta: ca. 2 mi. W of Glenevis, Moss 12449 (CAN, DAO). British Columbia: VANCOUVER ISLAND: Cowichen, Glendenmire (CAN, GH). Manitoba: Fort Churchill, 58*67'N, 94^10'W, Schofield & Crum 6787 (CAN, F). Northwest Territories: MACKENZIE DIST.: small pool ca. 45.5 mi. WNW of Yellowknife, Thieret & Reich 7947 1974] Potamogeton — Haynes 607 (F, os, US). Nova Scotia: CAPE BRETON CO.: Mira Bay, Macoun 20755 (F, GH, CAN). Ontario: THUNDER BAY DIST.: Black Fox Lake, Haynes 3749 (CAN, GH, LAF, MO, OS, UC); Sibley Twp., Ravine Lake, Taylor et al. 232 (CAN, GH, uc). Quebec: Lake Memphremagog, Sargents Bay, Churchill (GH, K, MO). GASPE CO.: Riviere Bonaven- ture, Marie-Victorin et al. 33764 (F, GH, NY, PH). Saskatchewan: MACKENZIE DIST.: Meridian Creek near Amisk Lake, Hudson 1550 (DAO). Yukon: Sheldon Lake, Canol Road mi. 222, Porsild & Breitung 11511 (CAN). UNITED STATES: Connecticut: FAIRFIELD CO.: Norwalk River, Ridgefield, Eames 11717a (GH). Maine: PENOBSCOT co.: Pushaw Stream, Old Town, Ogden et al. 1617 (CAN, GH, MO, NY, US). PISCATAQUIS CO.: cold clear stream, Foxcroft, Fernald 478 (GH, MO, US). Massachusetts: ESSEX CO.: ditch in Wenham, Morong (F, GH, ILL, PH); Pleasant Pond, Wenham, Faxon (GH, NY, US), Ogden & Ogden 1763 (CAN, DAO, F, GH, ILL, MICH, MO, NY, PH, UC, UPS, US). Michigan: KEWEENAW CO.: Beaver Pond, ca. 4 mi, S of Central Mine, Robbins (GH, MO, NY, PH). LUCE CO.: Bodie Lake, Haynes 8787 (0S). SCHOOLCRAFT CO.: Canoe Lake, Haynes 3741la (os). Minnesota: sT. LOUIS CO.: West Two Rivers, Tower, Hill 218- 1889 (CAN, ILL). Montana: GLACIER CO.: Howe Lake, Glacier Na- tional Park, Hazzard 480 (GH). New Hampshire: GRAFTON CO.: pond, Enfield, Kennedy (GH). New Jersey: MORRIS CO.: Rockaway River near Milton, Williamson (GH, PH). New York: ALBANY CO.: Alcove, Shear (GH, NY, UC). Pennsylvania: WAYNE CO.: Howell’s Pond, Twining (GH, PH). Rhode Island: PROVIDENCE CO.: Fountain Spring Brook, Smithfield, Lovewell (GH). Vermont: ADDISON CO.: Little Otter Creek, Ferrisburg, Eggleston & Grout (F, GH, PH, US). CALEDONIA CO.: spring-fed pond, ca. 2 mi. N of Walden Village, Haynes 3829 (os); Coles Pond, Walden Twp., Haynes 3837 (os). Washington: SAN JUAN CO.: Summit Lake, Orcas Island, Sutherland & Kern 1182 (CAN, NY, UC). Wisconsin: ONEIDA CO.: stream, Three Lakes, Hoffman (GH, MO). Wyoming: TETON CO.: Beaver ponds near Moose, Porter & Porter 9405 (GH, UC). 4. Potamogeton hillii Morong, Bot. Gaz. (Crawfordsville) 6: 290. 1881. TYPE: E. J. Hill, Stagnant pools, Manistee, [Manistee Co.] Michigan, (holotype, NY!; isotypes, F[2 sheets] !, aH [2 sheets] !, ILL[4 sheets] !, K!, PH!). Potamogeton porteri Fern. Mem. Amer. Acad. Arts 17: 73. 1932. TYPE: T. C. Porter, Run in Dillerville Swamp, near Lancaster, [Lancaster Co.] Pennsylvania, (lectotype, PH!; isolectotypes, F[2 sheets]!, MO !, NY[2 sheets] !, PH[2 sheets] !). 608 Rhodora [Vol. 76 Stems green to olive, slightly compressed, heavily ridged, 30-60 em long, 0.5-1.0 mm diam. Leaves pale-green to olive- green, delicate, 3-nerved, 2.0-6.0 cm long, 0.6-2.5 (-4.0) mm wide; apex apiculate to bristle-tipped; glands present or absent, brown to green, 0.1-0.3 mm diam; lacunae of 1-2 rows each side of midrib; lateral nerves joining midrib (0.4-) 0.7-1.7 mm from apex. Stipules white to light brown, slightly fibrous, rarely shredding at tip, convolute, 7.0-16.0 mm long, 0.6-2.2 mm diam. Winter buds rare, terminal, 2.8-3.0 em long, 1.5-3.0 mm wide; inner leaves undifferenti- ated; outer leaves 3-4 per side, acute to apiculate, without corrugations at base. Peduncles slightly clavate, axillary or terminal, rarely recurved, 6.0-13.5 mm long, 0.3-1.0 mm diam. Spike globose, (2.0-) 4.0-7.0 mm long, 4.6-7.0 mm diam; verticels 1-2, when 2, these crowded, 0.5-1.0 mm apart. Perianth segments 1.3-1.5 mm long, 1.1-1.5 mm wide. Fruit brown to light greenish-brown, dorsally and laterally keeled, 2.3-4.0 mm long, 2.0-3.2 mm wide; keels forming ridges, without undulations, to 0.2 mm high; beak central, rarely forward, 0.3-0.7 mm long, 0.2-0.6 mm diam; sides rounded rarely centrally depressed; wall texture rough. Chromosome number unknown. Distribution: In cold stagnant or slow moving, often brown water, from northern Lower Peninsula Michigan to Vermont and south to northeastern Ohio and southeastern Pennsylvania. Fruiting from late June to late August. Fig. 6. Illustrations: Fernald (1932, pl. 13; 30, fig. 2; 34, fig. 2). In the original description, Morong cited only the follow- ing data: “excellent specimens obtained in August 1880, by Mr. Hill, at Manistee, Michigan . . ." He included no indi- cation as to the location of the specimen. Hill prepared, from Manistee in August 1880, no less than 14 herbarium specimens on at least two separate days. Fernald (1932, p. 169), also giving no specimen location, designated as the type collection the one gathered by Hill on 10 August 1880. 1974] Potamogeton — Haynes 609 However, in the collection at NY there is a specimen col- leeted on 5 August 1880 by Hill and labeled in Thomas Morong's handwriting as “Type Specimen." Of the many other specimens examined by Morong, none of these others were indicated as types by Morong. Therefore, the 5 Au- gust specimen, having been designated by the author, then is the holotype and the one cited by Fernald then is a topo- type. Fernald (1932) named Potamogeton porteri based on several sheets collected by Thomas Porter in mill ponds near Lancaster, Pennsylvania. In the body of the original description Fernald did not cite any one specimen as the type. The data he gave are “PENNSYLVANIA: Cold streams or rivulets in Dillerville Swamp, Lancaster Co., October 5 and 6, 1860, Thos. C. Porter, distributed as P. pusillus and P. obtusifolius (TYPE in Herb. Acad. Sci. Nat. Phila.; duplicate in Herb. Gray; fragmentary (sterile) du- plicates in Herb. Field Mus., Dudley, and Mo. Bot. Gard.).” In a footnote on the same page he states ‘“Porter’s labels, all of material collected on October 5, with the exception of one sheet marked October 6, indicate somewhat different habitats: ‘Cold rivulets near Lancaster,’ ‘Near Lancaster,’ ‘In a rivulet, Dillerville swamp, Lancaster,’ ‘Stream in Dil- lerville Swamp, near Lancaster,’ ‘Run in Dillerville Swamp, near Lancaster.’ ” I have seen the specimens at PH, GH, F, MO, and none of the sheets have the locality data given by Fernald in the body of the text. There are, in fact, three sheets at PH labeled by Fernald as P. porteri. He indicated “Type Col- lection” on two of these sheets, “Run in Dillerville Swamp, near Lancaster, Penn. October 5, 1860” and “Stream in Dil- lerville Swamp, near Lancaster, Penn. October 5, 1860.” On the other sheet, ‘Cold rivulets near Lancaster, Pa. October 5," Fernald did not indicate “Type Collection.” The two specimens labeled as “Type Collection” by Fernald are obviously the same taxon and both match the descrip- tion. However, the one with data of “Stream in Dillerville Swamp, near Lancaster, Penn. October 5, 1860” is fertile 610 Rhodora [Vol. 76 while the other is sterile. Hence, I am here designating the fertile one as the lectotype. Fernald states (p. 74), “In its large fruit with prominent keel and recurved beak and its essential lack of glands at the base of the stipules P. porteri is close to P. hillii, but the latter species is wholly different in its acute, attenuate and thinner stipules; and in its attenuate almost bristle- tipped leaves with a single row of lacunae each side of the midrib and with the lateral nerves confluent with the midrib well below the tip." The specimens annotated by Fernald as P. porteri vary from ones with narrow leaves with apiculate leaf tips to very broad ones with obtuse leaf tips. The broad leaved specimens are quite distinct from characteristic P. hillii. However, the ones with the nar- rower leaves closely resemble P. hillii in all characters men- tioned by Fernald. A locality of P. hillii in Cheboygan County, Michigan, first located by Dr. E. G. Voss in 1967, was visited by me during the summers of 1970, 1971, and 1972. I noticed a variability within the population from individuals with the characteristic narrow attenuate leaves with delicate stipules to ones with broad obtuse leaves with coarse stipules. Within this one population, a complete gradation exists from characteristic P. hillii to character- istic P. porteri. For this reason I believe P. porteri is a morphological extreme of P. hilli. With the capitate inflorescence, short axillary peduncles, and fruits with a small dorsal keel, Potamogeton hillii closely resembles P. foliosus. However, it can be distin- guished from the latter species by the large 2.3-4.0 X 2.0- 3.2 mm fruits (the largest of any Pusilli), by the nearly bristle-tipped leaves and by the stipules being convolute. By the nearly bristle-tipped leaves, some individuals re- semble P. longiligulatus, a proposed hybrid, but can be dis- tinguished from this putative hybrid by the leaves having only 3 veins, whereas those of P. longiligulatus have mostly 7-9 veins. From all other Pusilli, P. hillii can be separated by its large fruits, short peduncles, and capitate inflores- cences. 1974] Potamogeton — Haynes 611 Fernald (1932) states that the Pusilli reproduce, for the most part, not by seeds, but by winter buds. He also states that winter buds are unknown in Potamogeton hillii. I have seen one sheet with well developed winter buds and several with ones that were immature. The winter buds, when present, with the inner leaves being unmodified, resemble small ones of P. obtusifolius in shape and structure, even in having the leaves round apiculate. Potamogeton hillii has been considered to be extremely local throughout its range. This distribution is becoming more apparent with the destruction of the aquatic environ- ment by pollution and dredging. In fact, Voss (1967) re- ported that the type locality of P. hillii has been destroyed. UN 2 WARES Fig. 6. Map of eastern North America showing the documented distribution of Potamogeton hillii. The barred line represents the southern limit of Wisconsin Glaciation (after Flint et al, 1959). 612 Rhodora [Vol. 76 Around 1900, the species was present in northeastern Ohio and northwestern Pennsylvania. This area has become heavily populated, with few if any natural areas remaining. The two Ohio counties, Portage and Ashtabula, have been rather heavily collected over the past 20 years. Some col- lectors thought they had collected the species, but after examining the specimens, I found them all to be P. pusillus. On the other hand, intensive field work in northern Michi- gan has shown the species to occur in no fewer than four localities within a radius of 30 miles of Cheboygan County. The same situation may prove to be true in other less heav- ily populated areas of its range, especially northeast New York state, as botanists continue to search for the species. REPRESENTATIVE SPECIMENS CANADA: Ontario: ELGIN CO.: near St. Thomas, James (DAO). UNITED STATES: Connecticut: LITCHFIELD CO.: Indian Pond, Sharon, Eames 11867 (GH). Massachusetts: BERKSHIRE CO.; pond fed by Karner Brook, South Egremont, Weber 1626 (CAN, F, GH, MICH, MO, NY, UC, US). Michigan: CHEBOYGAN CO.: Elliot Creek, 4 mi. E of Cheboygan, Haynes 3389 (GH, MO, OS, UC, US), Haynes 3713 (os). EMMET CO.: Cecil Bay marsh, 4 mi. SW of Mackinaw City, Voss 14061 (CAN, GH, MICH, MSC, NY, OS, UMBS, WIS). MAC- KINAC CO.: small stream ca. 1 mi. S of Engadine, Haynes 4025 (ASC, DAO, F, FSU, GH, K, MICH, MO, MSC, NY, OS, PH, SMU, TENN, TEX, UC, US, VDB, WIS, WVA). MANISTEE CO.: Manistee, Morong (GH, MICH, NY, US). OTSEGO CO.: outlet of Grass Lake, Stuckey 1300 (GH, MICH, NY, OS, UMBS). PRESQUE ISLE CO.: mouth of Black Mallard River, Haynes 3390 (F, GH, MO, NY, OS, PH, UC, US), Haynes 3714 (08), Haynes 3740 (ASU, GH, LAF, MO, NY, OS, PH, QFA, UC, VDB, US), Haynes 3890 (os). New York: THOMPKINS CO.: Ithaca, Lake Cayuga, Dudley (CAN, F, GH, MICH, MO. US). WASHINGTON CO.: creek 2 mi. N of Dresden Station road, Haynes 3342 (GH, MICH, MO, OS, UC, US). Ohio: ASHTABULA CO.: pools, Ashtabula, Hill (F, ILL). Pennsylvania: BEDFORD CO.: millpond S of Woodbury, Hotchkiss 6003 (GH, US). Vermont: WINDSOR CO.: Evarts Pond, Windsor, Dudley (GH, NY, UC, US). 5. Potamogeton foliosus Raf., Med. Repos. Hexade 2, 5: 354. 1808. Stems green to olive, slightly compressed and ridged, 4-75 em long, 0.2-1.2 mm diam. Leaves pale green to olive, 1974] Potamogeton — Haynes 613 rarely rufescent, delicate, 1-3 (-5)-nerved, 1.3-8.2 cm long, 0.3-2.3 mm wide; apex acute to apiculate, rarely with a bristle; glands present or absent, black to gold, to 0.5 mm diam; lacunae absent or to 2 rows each side of midrib; lateral nerves joining the midrib 0.5-1.2 mm below the apex. Stipules greenish to brown, rarely white, delicate to fibrous, shredding or not at the tip, connate or convolute, 0.2-2.2 em long, 0.2-1.7 mm diam. Winter buds uncommon, lateral or terminal, 0.9-2.5 em long, 0.6-2.0 mm wide; inner leaves rolled into a hardened fusiform structure; outer leaves 1-3 per side, acute to apiculate, without corrugations at base. Peduncles clavate, mostly in axiles of lower, rarely upper, leaves, usually recurved, 0.3-1.1 (-3.7) cm long, 0.3- 1.4 mm diam. Spikes capitate to cylindric, 1.5-7.0 mm long, 1.0-6.0 mm diam; verticels 1-2, when 2, these usually crowded, 0.6-1.2 (-2.0) mm apart. Perianth segments 0.4- 1.4 mm long, 0.3-1.0 mm wide. Fruit pale green to olive or brown, dorsally keeled, 1.4-2.7 mm long, 1.1-2.2 mm wide; keel undulate winglike, to 0.4 mm high; beak central, rarely forward, 0.2-0.6 mm long, 0.1-0.4 mm diam; sides rounded to centrally depressed; wall texture smooth. Potamogeton foliosus is most similar to P. hillii in its short clavate peduncles and capitate inflorescence. How- ever, when in fruit, P. foliosus can be separated from the latter species by the presence of a dorsal undulate wing on the fruit. The fruit of P. hillii, on the other hand, is three-keeled — two lateral and one dorsal ridge which does not appear as a thin undulating wing. Vegetatively, P. foliosus is most similar to P. pusillus. From P. pusillus var. tenuissimus, P. foliosus often can be distinguished by the near lack of lacunae. From P. pusillus var. pusillus, how- ever, the separation is not that simple. Fernald (1932) suggests that P. foliosus may be separated from P. panor- mitanus [var. pusillus] by the absence of nodal glands. Voss (1972), on the other hand, has questioned this char- acter as reliable. In most fruiting specimens of P. pusillus [var. pusillus] from Michigan, Voss could not find evident nodal glands. Data gathered by me from examination of 614 Rhodora [Vol. 76 hundreds of herbarium specimens and many populations in the field support Voss's conclusions. Therefore, I give little value to the absence of glands as a character for separating these two taxa. Although intermediates occur, the only character which I have been able to consider with any re- liance is the coarseness of the stipular veins. Those of P. foliosus, for example, are usually evident, appearing as ridges extending the length of the stipules. The veins are not usually so evident in P. pusillus. 5a. Potamogeton foliosus var. foliosus P. foliorum Raf., Med. Repos. Hexade 3, 2: 409. 1811. (orthographic variant). P. pauciflorus Pursh, Flora Amer. Sept. 121. 1814. P. foliosus Raf. var. genuinus Fern., Mem. Amer. Acad. Arts 17: 43. 1932. TYPE: Michaux, in rivis affluente mari inundatis Carolinae inferioris [South Caro- lina] (holotype, P; photograph of holotype, aH !). Potamogeton niagarensis Tuckerm., Amer. J. Sci. Arts Series 2, 7: 354. 1849. P. pauciflorus Pursh var. niagaren- sis (Tuckerm.) Robbins in Gray, Man. Bot. North. U. S., ed. 2. 435. 1856. P. foliosus Raf. var. niagarensis (Tuck- erm.) Morong, Mem. Torrey Bot. Club 3: 39. 1895. Spiril- lus foliosus (Raf.) Nieuwl. var. niagarensis (Tuckerm.) Nieuwl., Amer. Midl. Naturalist 3: 18. 1913. P. foliosus Raf. f. niagarensis (Tuckerm.) Hagstrom. Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE: Tucker- man, Hogback, Niagara Falls, [Niagara Co.] New York, (holotype, NY!; isotypes, GH !, K!). Potamogeton pauciflorus Pursh var, californicus Morong, Bot. Gaz. (Crawfordsville) 10: 254. 1885. P. foliosus Raf. var. californicus (Morong) Morong, Mem. Torrey Bot. Club 3: 40. 1895. P. californicus (Morong) Piper, Contr. U.S. Natl. Herb. 11: 98. 1906. P. foliosus Raf. f. californicus (Morong) Hagstrom, Kongl, Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE: S. B. & W. F. Parish 940, submerged aquatic, streams, San Bernardino, San Ber- nardino Co., California, (holotype, NY!; isotypes GH !, MO!, PH!). 1974] Potamogeton — Haynes 615 Potamogeton foliosus Raf. var. macellus Fern., Mem. Amer. Acad. Arts 17: 46. 1932. TYPE: E. & C. E. Faxon Fresh Pond, Cambridge, [Suffolk Co.] Massachusetts, (holo- type, GH ; isotype, US!). Stems 4-72 cm long, 0.2-1.2 mm diam. Leaves 1-8 (-5)- nerved, 1.3-8.2 cm long, 0.3-2.3 mm wide; apex acute to apiculate, rarely with a bristle; glands rare, to 0.3 diam; lacunae absent or to 2 rows each side of midrib; lateral nerves joining the midrib 0.5-1.2 mm below the apex. Stipules greenish to brown, delicate to slightly fibrous, rarely shredding at the apex, connate, 0.2-2.2 cm long, 0.2- 1.7 mm diam. Winter buds lateral, 0.9-1.8 cm long, 0.6-1.0 mm wide. Peduncles usually clavate, recurved, 0.3-1.1 (-3.7) em long, 0.3-1.4 mm diam. Spikes usually capitate, rarely cylindric, 1.5-7.0 mm long, 1.0-6.0 mm diam; verticels 1-2, when 2, these crowded, 0.6-1.2 (-2.0) mm apart. Perianth segments 0.4-1.4 mm long, 0.5-1.0 mm wide. Fruit olive to green-brown, 1.5-2.7 mm long, 1.2-2.2 mm wide; keel 0.1- 0.4 mm high; beak 0.2-0.6 mm long, 0.2-0.4 mm diam. Chromosome number, 2n — 28 (Stern, 1961). Distribution: Widespread, in waters of lakes, springs, streams and rivers, from central Alaska to Nova Scotia and south to southern Guatemala and Jamaica. Fruiting from late May to late October. Fig. 8. Illustrations: Fernald (1932, pl. 3; 29, fig. 1; 32, fig. 3; 38, fig. 1). Potamogeton foliosus was first recognized by Michaux (1803) when he reported P. gramineus? L. for North America. Rafinesque (1808), noticing that many of the Western Hemisphere plants represented different species from those of Europe, listed several species of Michaux which he later intended to describe. Among these was “P. gramineum to be described later as P. folioswm." Indeed, he did later describe the taxon (Rafinesque, 1811), but it was spelled as P. foliorum rather than as P. foliosum. However, the name of P. foliosum [foliosus] was validly and effectively published in 1808 when he made direct ref- 616 Rhodora [Vol. 76 erence to the published description of P. gramineum of Michaux. Pursh (1814) applied a superfluous name — Potamogeton pauciflorus — to Michaux’s specimen. Other botanists either overlooked or neglected Rafinesque’s name and ac- cepted instead the one of Pursh. Tuckerman (1849) recog- nized that two forms — a large-leaved form and a small- leaved form — of P. pauciflorus Pursh [foliosus] existed. He divided the group into two species, applying P. niaga- rensis Tuckerm. to the larger-leaved individuals and retain- ing the name P. pauciflorus for the smaller-leaved ones. After noticing considerable morphological integration, Fer- nald (1932) chose to consider the two taxa at the varietal rank. After having examined Michaux's specimen at Paris, Fernald determined the specimen to be the large-leaved entity. Thus, P. foliosus and P. pauciflorus actually applied - Zz c EF] Lal m z Q P Dos e ^9 a e oœ z oe oo 9 © 0e L = 9 9 M ao 9 e C Oo O 9 o 9 9 99 c + + " 4 - Tt P LEAF LENGTH (CM) Fig. 7. Potamogeton foliosus var. foliosus. Scatter diagram com- paring leaf length, fruit length, and number of nerves per leaf. 1 nerve, open circle; 3 nerves, half-closed circle; 5 nerves, dot. 1974] Potamogeton — Haynes 617 to this growth form. Therefore, the smaller-leaved plants were unnamed. Fernald then named the smaller-leaved specimens P. foliosus var. macellus Fern. and called the larger river form var. genuinus [now correctly as var. foliosus]. I have examined thousands of specimens in the field and herbarium. Many of these have been plotted on scatter diagrams (see Fig. 7). Although only three char- Fig. 8. Map of North America showing documented distribution of Potamogeton foliosus var. foliosus. 618 Rhodora [Vol. 76 acters are used here, other characters were plotted and similar patterns developed. Data gathered from these studies indicate that a continuous gradient exists from ones with the largest leaves to those with the smallest leaves. The two groups do not appear to be separated into distinct geographical areas, with the intermediates in areas of overlap. Instead, these data indicate that the larger- leaved individuals occur in rapidly flowing waters, the inter- mediates occur in slow moving streams, and the smaller- leaved plants occur in lakes or ponds. Sculthorpe (1967) indicates that the size and shape of the leaves of many species of Potamogeton vary greatly with different rates of water flow. This is probably the situation with P. foliosus. I am, thus, considering var. macellus to be an ecological growth form of P. foliosus, and therefore, am not formally recognizing it. REPRESENTATIVE SPECIMENS CANADA: Alberta: Wood Buffalo Park, Raup 1562 (CAN, NY, uc). British Columbia: Revelstoke, Macoun 3037 (CAN, NY). Manitoba: Little Saskatchewan, Rapid City, Macoun 16482 (CAN, GH). Northwest Territories: small lake, 38.3 mi. WNW of Yellow- knife, Thieret & Reich 8356 (F, os). Nova Scotia: KINGS CO.: Canard River, Smith et al. 11446 (CAN, DAO). Ontario: FRONTENAC CO.: Kingston, Smith Falls, Fowler (CGE, MO, US). Prince Edward Island: QUEENS CO.: ponds, Southport, Fernald & St. John 6782 (CAN, K, NY, US). Quebec: NEW-RICHMOND CO.: Bonaventure River, Marie- Victorin et al. 33839 (CAN, F, MU, NY, PH). Saskatchewan: MOOSE JAW DIST.: Mortlach, Hudson 1143 (Dao). GUATEMALA: Dept. Esquintla: mountain stream, Finca San Filipe, Muenscher 12074 (cH, os). JAMAICA: Portland: Rio Grande, Maxon & Killip 265 GLL). MEXICO: Baja California: Rancho San Jacinto, 45 mi. S. of Eusenada, Wiggins & Demaree 4749 (F, MICH, MU, NY, US); stream, Santiago, Wiggins 5662 (MICH, NY, UC, US). Chihuahua: DEPT. DE GUERRERO: stream near Minaca, Shreve 7993 (F, GH). Guanajuato: DEPT. ATOTONILCO: just S of village of Santuario, between San Miguel Allenda and Dolores Hidalgo, Ogden 51149 (os). Michoacan: Rio Duero at Zamora, Ogden 51183 (os). Puebla: DIST. DE TEPEACA: San Hipalito, vicinity of Puebla, Arsene 2860 (ILL, NY, us). Sonora: Fronteras, Hartman 992 (GH, NY). Veracruz: Orizaba, Engenio, Rio Blenio, Muller 1330 (NY). UNITED STATES: Alabama: JACK- SON co.: Paint Rock River, Larkin Fork, Harper 2998 (GH, MO, NY, 1974] Potamogeton — Haynes 619 PH, US). Alaska: 2 mi. SW of College, small pond, Smith 1911 (CAN). Arizona: Colorado Havasupai Canyon, Clover 5218a (MICH, US). Arkansas: FULTON C0.: Mammoth Springs, Spring River, Thomas & Smith 17094 (os). California: Los ANGELES CO.: Lancaster, Elmer 3504 (GH, K, NY, UC, US). SISKIYOU CO.: near Mt. Shasta, Abrams Lake, Heller 13908 (F, ILL, MO, NY, PH, UC, US). Colorado: DENVER co.: Denver, Platte River, Jones 600 (F, NY, US). Connecticut: LITCHFIELD CO.: Litchfield-Morris Wildlife Sanctuary, Bantam Lake, Dwyer 2150 (NY). Delaware: NEW CASTLE CO.: near Wilmington, Brandywine Creek, Commons (Pn). District of Columbia: Wash- ington, Ward (GH, UPS). Georgia: CHATTOOGA CO.: 2.8 mi, E 30^N of Trion, small pond below Greeson Springs, Duncan & Harris 12842 (GH, LAF, MICH, US). Idaho: BOISE co.: 20 mi. S of Idaho City, marsh, Hitchcock & Muhlick 9947 (Ny, vc). Illinois: COOK CO.: Englewood, ditches, Hill (F, GH, ILL, US). ST. CLAIR CO.; East St. Louis, Eggert (CAN, F, ILL, K, MICH, NY, PH, OS, US). STARK CO.: near Wady Petra, prairie pond, Chase 1148 (ILL, MO, NY, PH). Indiana: LAKE CO.: Wolf Lake, West Bay, Chase 1461 (F, ILL, MO, PH, US). Iowa: CLAY CO.: Lake Twp, Dan Greene’s slough, N end, Hayden 820 (MO, NY, PH, US). EMMET CO.: Estherville, running water, Cratty (GH, ILL, MO, NY, US). PALO ALTO CO.: Highland Twp, 2 mi. S of Ruthven, Virgin Lake, Hayden 10125 (Mo, NY, PH, US). Kansas: RILEY CO.: ponds, Pond 1100 (MICH, MO, NY, US). Kentucky: FAYETTE CO.: near Lexington, brook, Peter (K, MICH, NY, PH). Maine: PISCATAQUIS CO.: Dover, river margin, F'ernald 479 (Mo, Us). Massachusetts: MIDDLESEX CO.: Cambridge, Fresh Pond, Morong (F, MO). Michigan: CHEBOYGAN CO.: Douglas Lake, Marl Bay. Haynes 3923 (os). EMMET CO.: small arm of Carp Lake, Haynes 3374 (GH, os, UC, US), Haynes 3705 (ILL, os). Haynes 3745 (os), Haynes 3888 (os). Minnesota: CLEARWATER CO.: Bohal Lake, Grant & Oosting 3228 (NY, UC, US). Missouri: BARRY CO: Eagle Rock, river, Bush 511 (CGE, GH, MO, US). BUTLER CO.: Poplar Bluff, Eggert (CAN, F, K, MO, OS, US). OREGON CO.: Thomasville, spring pond, Steyermark & Palmer 41701 (MO, NY. PH, US). TEXAS CO.: SW of Plato, Roubidoux Creek, Steyermark 25006 (DAO, F, NY. os). Montana: MISSOULA CO.: ca. 6 mi. E of Lolo Hot Springs, small slough, Hitchcock 23978 (DAO, ILL, NY, US). Nebraska: LINCOLN co.: North Platte, slough, Shear 4445 (GH, NY, US). New Hampshire: GRAFTON CO.: Hanover, Count River, Jesup (MO). New Jersey: MORRIS CO.: Oak Ridge, running water, Mackenzie 3736 (MO. Ny, US). New Mexico: LINCOLN CO.: Bonita River, ca. 616 mi. NW of Alto, Haynes 2923 (LAF). New York: COLUMBIA CO.: 1% mi. SSW of West Copake, Lower Rhoda Pond, Haynes 3802 (os). DUTCHESS CO.: 14% mi. SW of Pine Plains, Stissing Pond, Haynes 3804 (0s). ESSEX CO.: near Ticonderoga, outlet of Lake George, Haynes 3333 (os), Haynes 3336 (0S). NIAGARA CO.: 620 Rhodora [Vol. 76 Niagara Falls, mill race near Cataract House, Hill 26-1887 (ILL, MICH, MO). PUTNAM CO.: Carmel, Glendia Lake, Muenscher & Curtis 5447 (MO, NY, PH, UC, US). WARREN CO.: Lake Luzerne boat launch, Haynes 3814 (os). Nevada: ELKO CO.: Copper Mountains, 3 mi. S of Coon Creek Pass, Maguire & Holmgren 22411 (CAN, NY, US). North Carolina: CHEROKEE C0.: N of Marble, limestone quarry, Radford 4812 (NY). North Dakota: CASS CO.: Fargo, ditch, Stevens 1376 (CAN, UC, US). Ohio: AUGLAIZE CO.: Lake St. Marys Fish Hatchery, Haynes 3427 (GH, MICH, OS, us); 2 mi. SE of Wapakoneta, borrow pit, Haynes 3420 (0s, US). CHAMPAIGN CO.: Kaiser Lake State Park, Haynes 3435 (MICH, OS, UC, US). ERIE co.: 4 mi. NW of Castalia, Haynes 3247 (MICH). LOGAN CO.: 1.9 mi. due ESE of Russells Point, Haynes 3415 (MICH, OS, PH, us). OTTAWA CO.: Lake Erie, North Bass Island, Fox’s Marsh, Haynes 3145 (08). PORTAGE co.: 5% mi. NW of Ravenna, borrow pit, Haynes 3452 (GH, MICH, MO, OS, US). Oklahoma: OTTAWA CO.: Hattenville, overflow pond, Stevens 2481 (GH, ILL, K, MO, NY, US). WOODS CO.: Waynoka pond, Stevens 1768 (GH, ILL, K, MO). Oregon: pond on the Willamette River, Hall 492 (F, GH, ILL, MO, NY, us). Pennsylvania: BERKS CO.: 1 mi. SE of Pine Forge, Manatawny Creek, Wilkens 5619 (GH, PH). South Carolina: GEORGETOWN CO.: Georgetown, Hobeaw Plantation, Alexander 148 (Us). South Dakota: PENNINGTON CO.: Sheridan Lake, Porter 6593 (DAO, UPS). Tennessee: FRANKLIN CO.: Sher- wood, Eggert (GH, MO, US). Texas: JEFF DAVIS CO.: Limpia Canyon, Tracy 288 (GH, NY). Utah: CACHE CO.: Bear River Range, E side of Tony Grove Lake, Maguire & Snell 16048 (DAO, ILL, MICH, NY). GRAND CO.: Hill Creek near Weaver Reservoir, Holmgren et al, 2358 (DAO, ILL, NY, UPS). Vermont: ORLEANS CO.: Ca. 1 mi. W of Crafts- bury Common, backwater of Black River, Haynes 3839 (os); 1 mi. S of Craftsbury Common, backwater of Black River, Haynes 3840 (os). WINDSOR CO.: Plymouth, Eglleston (F, MO, PH, us). Virginia: BOTETOURT CO.: Cloverdale, Tinker Creek, Wood 6183 (GH). Wash- ington: GRANT CO.: Pond SW of Moses Lake, St. John et al. 4960 (Mo, UC). West Virginia: GREENBRIAR CO.: Dunlap Creek, Berkley 1194 (mo). Wisconsin: WAUSHARA CO.: Poy Sippi, Pine River, Hill (CAN, F, GH, ILL). Wyoming: LARAMIE CO.: Laramie Mountains, Middle Fork of Crow Creek, Porter 6320 (DAO, MO, NY, UPS). 5b. Potamogeton foliosus var. fibrillosus (Fern. Haynes & Reveal, Rhodora 75: 76. 1973. Potamogeton fibrillosus Fern. Mem. Amer. Acad. Arts 17: 51. 1932. TYPE: W. C. Cusick 2598, in warm spring, margin of Harney Valley, “P” Ranch, [Harney Co.] Ore- gon, (holotype, GH!; isotypes, F! K!, MO!, NY!, uc!, us!). 1974] Potamogeton — Haynes 621 Stems 20-75 cm long, 0.3-1.2 mm diam. Leaves (1-) 3- nerved, 2.1-5.5 mm long, 0.5-1.7 mm wide; apex acute apicu- late; glands usually present, black to gold, to 0.5 mm diam; lacunae absent; lateral nerves joining midrib 0.6-0.8 mm below apex. Stipules brown, rarely white, fibrous, shred- ding at tip, often convolute, 0.4-12.1 em long, 0.5-1.2 mm diam. Winter buds lateral or terminal, 1.2-2.5 cm long, 0.8.2.0 mm wide. Peduncles clavate, recurved, 6.5-9.2 (-22.0) mm long, 0.4-1.0 mm diam. Spike capitate to cylin- dric, 1.7-5.5 mm long, 1.0-4.0 mm diam; verticels 1-2, when 2, then 0.6-0.8 mm apart. Perianth segments 0.5-0.7 mm long, 1.1-1.2 mm wide. Fruit pale-green, 1.4-1.7 mm long, 1.1-1.2 mm wide; keel to 0.2 mm high; beak ca. 0.2 mm long, 0.1-0.2 mm wide. Chromosome number unknown. Distribution: In waters, often warm, of shallow lakes, springs, streams and rivers, from southwestern Washing- ton southward through southeastern Oregon, hence east- ward to southeastern Wyoming. Fruiting from mid-June to mid-September. Figure 9. Illustrations: Fernald (1932, pl. 5; 28. Fig. 5; 32, Fig. 5). In 1932, Fernald proposed Potamogeton fibrillosus based on specimens collected in Wyoming, Idaho, Oregon and Washington. He noted that the new species closely resem- bled P. foliosus Raf. in its foliage, peduncles and small dorsally keeled fruits. Fernald said that P. fibrillosus dif- fered from P. foliosus by its stipules which disintegrate into “rope-like” fibers and are thus open or convolute, and by having fruits with a less developed keel and more nearly median beak. Fernald (1932, p. 52) stated his reservations in assigning specific rank to this new entity: “The plant will doubtless be found to have a broader range, when it may prove to be a marked geographic variety of the wide- spread P. foliosus." Several collections, especially from Yellowstone National Park of Wyoming and from southeastern Oregon, are dis- tinct from Potamogeton foliosus. In general, these speci- 622 Rhodora [Vol. 76 mens differ from P. foliosus not only in having the stipules disintegrating into fibers and fruits with poorly developed keels, but also in having cylindric, interrupted inflorescences and leaves with basal glands — the last two characters not mentioned by Fernald, but are present even on the holotype. Other collections from the Yellowstone region are not so distinct. One such collection (Haynes 3849) has fruits, including the keel, which are identical with those of Pota- mogeton foliosus, but the inflorescence is capitate and the stipules only rarely disintegrate into fibers. Yet, basal glands are present. Flowering specimens from Albany County, Wyoming, taken by C. L. Porter (3473) are similar to P. fibrillosus in having basal glands and short, clavate peduncles. However, in this collection, the stipules only rarely disintegrate into fibers. Flowering specimens ob- tained by Maguire (21578) in northern Utah commonly produce fibers when the stipules decay, but basal glands are rarely found. Hitchcock (1969) regarded Potamogeton fibrillosus as a distinct species, but conceded its close relationship with P. foliosus. Porter (1963) reduced P. fibrillosus to synonymy, regarding it only as a local form of P. foliosus produced by the rather warm-water conditions of “geyser formations and hot springs" in the Yellowstone area. Porter stated (19683, p. 9) “The warm water hastens disintegrations of the sheaths by bacterial action." 'This is not always the case, however, in northern Utah or southeastern Oregon where the water is often cool. Here the stipules still form fibers. In the same general region, P. foliosus is known to occur in similar environmental conditions and at approxi- mately the same degree of development. Therefore, the possibility that mere bacterial activity (or some other type of mechanical breakdown) being the only cause of the fibrous stipules seems unlikely. For this reason and that the extremes can be distinguished quite easily, Haynes and Reveal (1973) thought that P. fibrillosus should be taxo- nomically recognized. However, since many intermediate 1974] Potamogeton — Haynes 623 forms can be found, they could not justify the specific rank and therefore ranked the taxon at the varietal level. From var. foliosus, the var. fibrillosus may be distin- guished by its poorly developed keel, cylindric and inter- rupted inflorescence, fibrous stipules and glands present at the base of most leaves. Fig. 9. Map of northwestern United States showing documented distribution of Potamogeton foliosus var. fibrillosus. 624 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS UNITED STATES: Idaho: BLAINE Co.: Russian John Ranger Station, Macbride & Payson 3790 (Uc). Oregon: HARNEY CO.: warm spring, near Burns, Henderson 8867 (GH); irrigating ditch at French- glen, Thompson 12068 (NY). KLAMATH CO.: N end of Klamath Marsh, Peck 21954 (UC). MALEUR CO.: Otis Creek, Leiberg 2340 (GH, US). Utah: CACHE CO.: stream, 3 mi. NE of Logan, Maguire 16685 (CAN, MO, NY); stream, 2 mi. NW of Logan, Maguire 21578 (DAO, F, GH, MICH, MO, NY, US). Washington: PIERCE CO.: outlet, Lake Spanaway, near Tacoma, Thompson 9657 (GH, NY); unknown locality, Wash- ington Territory, Brandegee 1127 (GH, UC). Wyoming: ALBANY CO.: N. fork of Pole Creek, Porter 3474 (Ny, US). PARK CO.: Firehole River, Jepson 2540 (GH); Firehole Canyon, Firehole River, Haynes 3848 (ILL, OS, PH, UC, US); Midway Geyser Basin, Firehole River, Haynes 3850 (GH, MICH, MO, NY, 0S); Upper Geyser Basin, Richard- son (GH); Upper Nez Perce Creek, Haynes 3849 (MICH, os). 6. Potamogeton clystocarpus Fern., Mem. Amer. Acad. Arts 17: 79. 1932. TYPE: J. A. Moore & J. A. Steyermark 3088, pool in rock, Little Aguja Canyon, Davis Mts., 1575 ft. Jeff Davis Co. Texas, (holotype, GH!; isotvpes, MICH!, MO!, NY!, PH!, ua, Stem light green to brown, terete to slightly compressed, rarely ridged, to 57 cm long, 0.5-0.7 mm diam. Leaves green, 3 (-5)-nerved, 3.2-7.8 cm long, 0.7-1.7 mm wide; apex acute; glands usually present, white to gold, 0.2-0.3 mm diam; lacunae to 4 rows each side of midrib, rarely absent; lateral nerves joining midrib 0.2-0.4 mm below apex. Sti- pules brown, delicate, not shredding at tip, usually convo- lute, to 6.2 mm long, 0.5-0.8 mm diam. Winter buds un- known. Peduncles cylindrical, axillary to terminal, erect, 3.2-4.8 cm long, 0.3-0.5 mm diam. Spike capitate to cylin- dric, 5.5-7.5 mm long, 3.0-5.7 mm diam; verticels 3, 1.5-1.7 mm apart. Perianth segments 1.7-2.0 mm long, 1.5-1.8 mm wide. Fruit brown to yellow-green, dorsally and laterally keeled, 2.0-2.2 mm long, 1.7-1.8 mm wide; keels ridged, without undulations, to 0.2 mm high; beak central, 0.5-0.6 mm long, 0.2-0.4 mm diam; sides depressed, with 1-3 tuber- cles near base; wall texture rough. Chromosome number unknown. 1974] Potamogeton — Haynes 625 Distribution: Known only from the type locality. Fruit- ing from early May to mid-June. Fig. 10. Illustrations: Fernald (1932, plate 15.30, fig. 5) ; Ogden (1966, plate 51). Potamogeton clystocarpus was named by Fernald (1932), based on the “gibbous-tuberculate-based fruits.” He sug- gested the fruits were “so similar to those of the western Eurasian and African P. trichoides C. & S. in having basal bosses that they might easily pass as fruits of that species. P. trichoides, however, as its name implied, has very bristle- tipped leaves . . . P. clystocarpus cannot be referred to it." No other North American pondweed, to my knowledge, has fruits with the basal “bosses.” In this character, P. clysto- carpus is quite distinct. But, as with other Pusilli, it is almost impossible to identify with sterile material! Fig. 10. Map of southwestern North America showing docu- mented distribution of Potamogeton clystocarpus. 626 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS UNITED STATES: Texas: JEFF DAVIS CO0.: Little Aguja Canyon, Davis Mts, Palmer 34526 (Mo, NY, PH), Correll & Ogden 25070 (NYS, UC). 7. Potamogeton pusillus L., Species Plantarum 1: 127. 1753. Stems pale green to olive, terete to slightly compressed, smooth to slightly ridged, 18-150 cm long, 0.1-0.7 mm diam. Leaves pale green to olive, rarely rufescent, delicate to coarse, 1-3 (-5)-nerved, 0.9-6.5 cm long, 0.2-2.5 mm wide; apex subulate to obtuse; glands present or absent, green, gold, brown, or rarely white, to 0.5 mm diam; lacunae absent or to 5 rows each side of midrib; lateral nerves, when present, joining midrib 0.1-1.0 (-2.0) mm below apex. Stipules brown to green or white, delicate, rarely appearing fibrous, not shredding at tip, connate or convolute, 3.1-9.2 mm long, 0.2-0.7 mm diam. Winter buds common, lateral or terminal, 0.9-3.2 em long, 0.3-1.8 mm wide; inner leaves rolled into a hardened fusiform structure; outer leaves 1-3 per side, subulate to obtuse, without corrugations at base. Peduncles filiform to slightly clavate, axillary or terminal, mostly erect, rarely recurved, 0.5-6.2 (-6.6) cm long, 0.2- 0.7 mm diam. Spikes capitate to cylindric, 1.5-10.1 mm long, 0.7-5.2 mm diam; verticels 1-3 (-4), 1.2-4.7 mm apart. Perianth segments 0.7-1.7 mm long, 0.5-1.2 mm wide. Fruit green to brown, without keels, 1.5-2.2 mm long, 1.2-1.6 mm wide; beak forward or central, 0.1-0.6 mm high, 0.2-0.5 mm diam; sides rounded; wall texture smooth or rough. Potamogeton pusillus has been interpreted variously by botanists. Bennett (1890-1894; 1900-1904) and Morong (1893), for example, considered the taxon to consist of two species, one being divided into several varieties. Hagstrom (1916), on the other hand, divided the taxon into several poorly defined species, with two of these — P. panormitanus [var. pusillus] and P. pusillus [var. tenuissimus] — being split into many infraspecific taxa. These two species were separated by the connate versus convolute character of the stipule. Fernald (1932) took the intermediate position and 1974] Potamogeton — Haynes 627 recognized three species — P. panormitanus [var. pusillus], P. gemmiparus [var. gemmiparus], and P. pusillus [var. tenuissimus]. He divided P. panormitanus into two vari- eties and P. pusillus into six varieties. These varieties were based on the size of the leaf, apex shape, and number of lacunae bordering the midrib. The latter concept of the taxon was accepted until Dandy and Taylor (1938), after studying the type specimen of P. pusillus in the Linnaean herbarium, decided that the name P. pusillus actually should be applied to the taxon which had been passing as P. panor- mitanus. For the taxon which had been passing as P. pusil- lus, the next available name in the specific rank was P. berchtoldii Fieber. Fernald (1940) finally made the com- binations of various varieties to conform with the revised nomenclature. This is the concept that appeared in the eighth edition of Gray’s Manual (Fernald, 1950). Gleason and Cronquist (1963), however, did not accept P. pusillus and P. berchtoldii as two distinct species. They, instead, suggested that although this character may be good for distinguishing certain species, there was no reason to as- sume that both states of the stipule and gradations be- tween could not oceur within one species. As I indicated earlier, à single plant may produce two sets of leaves during one growing season. The first set may have obtuse leaves with 4-5 rows of lacunae; the second, instead, would probably have acute leaves with 1-3 rows of lacunae. In the early summer, this plant would resemble var. mucro- natus [of Fernald, 1932] and in the late summer, it would resemble var. tenuissimus [of Fernald, 1932]. For this reason and because of the ecological variability as explained by Seulthorpe (1967), I have decided to recognize only three taxa. Fernald (1950) suggested that useful characters in dis- tinguishing Potamogeton pusillus from P. berchtoldii were length of inflorescence, branching of the plant, and whether the fruit is wider at, above, or below the middle. Voss (1972) had noticed that fruiting specimens with connate stipules [P. pusillus var. pusillus] from Michigan often 628 Rhodora [Vol. 76 lacked glands. Ogden (personal communication) suggested that the presence or absence of lacunae along the midrib often is a reliable character for distinguishing the species. Eighty specimens were examined for the characters men- tioned above and other features. Some correlation, although slight, was noticed between the stipule character, the diam- eter of the glands, and the number of lacunae bordering the midrib. However, for the majority of the characters measured, little or no correlation was observed. Regardless of the inconsistency of the stipule character, the shape of the inflorescence and of the fruit, and the length of peduncle seem to be associated. This is not to say that intermediates do not occur, because they do! In their extremes, the two entities can be separated by the length of the inflorescence, the shape of the fruit, the length of the peduncle, the number of laeunae bordering the midrib, the presence and size of nodal glands, and, finally, the connation of the stipules. No one character can definitely be depended upon, but by using a combination of as many of the above mentioned cnarac- ters as are present on the plant, one can place the specimen into the correct taxon 70-80% of the time. The remaining 20-30%, however, are almost impossible to identify without depending upon one character taxonomy. Also, var. tenuis- simus has a more northerly distribution, e.g. into Alaska, and var. pusillus has a more southerly distribution, e.g. into Mexico. Because of the morphological integradation and the slight range differences, I have decided to place the two taxa at the varietal level. When in fruit, Potamogeton pusillus should not be con- fused with any other pusilloid. Its fruit, lacking a keel, a wing, or basal warts, and its delicate stipules separate it from all other species. However, in the sterile state, identi- fieation is not so easy. Vegetatively, it closely resembles P. foliosus, P. clystocarpus, and some individuals of P. hillit. I can give no morphological characters which serve to sep- arate P. pusillus from P. clystocarpus if fruits are lacking. If the specimen is from areas other than western Texas, one can assume the specimen is not P. clystocarpus, although 1974] Potamogeton — Haynes 629 this may not be a valid assumption. The stipules of P. pusillus, usually lacking slightly fibrous veins, often serve to distinguish the species from P. hillii and P. foliosus. However, the stipules of the two latter species may, on occasions, appear to lack the fibrous veins. In this instance, the species are impossible to separate. Therefore, one should always make an effort to collect fertile material! Ta. Potamogeton pusillus var. pusillus Potamogeton pusillus L., Species Plantarum, 128, 1753. P. pusillus var. typicus Fern., Mem. Amer. Acad. Arts 17: 81. 1932. TYPE: Europe. (holotype, LINN; photograph of holotype, os!). Potamogeton panormitanus Ant. Biv. Bern. in And. Biv. Bern. Nuove piante inedite del barone Ant. Bivona Bernardi 6. 1838. (publication not seen). P. pusillus var. panormi- tanus (Ant. Biv. Bern. in And. Biv. Bern) Morong, Mem. Torrey Bot. Club 3: 46. 1893. TYPE: (location unknown). Potamogeton panormitanus var. major Ant. Biv. Bern. in And. Biv. Bern, Nuove piante inedite del barone Ant. Bivona Bernardi 6. 1838. (publication not seen). TYPE: (location unknown). Potamogeton panormitanus var. minor Ant, Biv. Bern. in And. Biv. Bern. Nuove piante inedite del barone Ant. Bivona Bernardi 6. 1838. (publication not seen). P. pusil- lus var. minor (Ant. Biv. Bern. in And. Biv. Bern.) Fern. & Schubert, Rhodora 50: 154. 1948. TYPE: (location un- known). Potamogeton pusillus var. vulgaris Fries, Novitiae Florae Suecicae 2nd ed. 49. 1828. TYPE: (type not located), ap- plication of the name is from the illustration of Potamoge- ton pusillus L. In Smith, English Bot. 2: pl. 215. 1794! Potamogeton pusillus var. vulgaris subvar. interruptus Robbins in Watson, Bot. U. S. Geol. Expl. Fortieth Parallel 338. 1871. TvPE: S. Watson, 1137, Parley's Park in the Wasatch, 6,000 ft. [Wasatch Co.], Utah, (holotype, NY!; isotype, US!). 630 Rhodora [Vol. 76 Stems green to brownish, mostly terete, slightly ridged, 18-150 em long, 0.2-0.7 mm diam. Leaves pale green, rarely olive, delicate, 1-3-nerved, 1.4-6.5 cm long, 0.5-1.9 mm wide; Fig. 11. Map of North America showing documented distribu- tion of Potamogeton pusillus var. pusillus. 1974] Potamogeton — Haynes 631 apex acute or rarely apiculate, rarely with a bristle; glands often present, green, gold, brown, or rarely white, to 0.3 mm diam; lacunae absent or to 2 rows each side of midrib; lateral nerves joining midrib 0.1-0.7 mm below the apex. Stipules greenish to brown, delicate, rarely fibrous, usually connate, 2.2-9.2 mm long, 0.2-0.7 mm diam. Winter buds uncommon, lateral or terminal, 0.9-1.5 em long, 0.3-1.8 mm wide; outer leaves 1-2 per side, acute to apiculate. Pedun- cles cylindric, axillary, to lower or upper leaves, rarely re- curved, 1.0-6.2 em long, 0.2-0.7 mm diam. Spikes cylindric, 2.5-10.1 mm long, 1.2-4.7 mm diam; verticels 2 or 3 (-4). usually interrupted, 1.4-4.5 mm apart. Perianth segments 1.0-1.5 mm long, 0.7-1.2 mm wide. Fruit green to brown, usually wider above the middle, 1.5-2.2 mm long, 1.2-1.5 mm wide; keel absent; beak forward, rarely central, 0.1-0.6 mm high, 0.2-0.5 mm diam; sides rounded, usually centrally depressed; wall texture smooth. Chromosome number 2n — 26 (Palmgren, 1939). Distribution: From central British Columbia to Nova Scotia, south to northern Guatemala and western Florida. Fruiting from early May to late September. Fig. 11. Illustrations: Fernald (1932, pl. 9; 10; 29, fig. 7, 8; 33, fig. 4, 5; 39, fig. 10). REPRESENTATIVE SPECIMENS CANADA: Alberta: Wood Buffalo Park, Raup 1568 (CAN, GH, NY, UC, US). British Columbia: pond near Similkameen River, Macoun 70317 (F, GH, NY). Manitoba: Desford, pothole, Love & Love 6092 (DAO, ILL, US). New Brunswick: KINGS CO.: Westfield, Ingleside, cove in St. John River, Fernald 1618 (GH, NY). Northwest Territories: MACKENZIE DIST.: Norman Wells, MacKenzie River, Cody & Gutte- ridge 1631 (F, NY, US). Nova Scotia: CAPE BRETON CO.: Fort Louis- bourg, Smith et al. 8281 (CAN, DAO). Ontario: OTTAWA CO.: Ottawa, Rideau Canal, Fletcher (MO, NY, PH). Prince Edward Island: Tig- nish, pools and ponds, Fernald et al. 6776 (CAN, K, NY). Quebec: Magdalen Islands, Grindstone Island, Fernald et al. 6780 (GH, K, NY, US), Fernald et al. 6781 (GH, NY, PH, US). Saskatchewan: MAPLE CREEK DIST.: 14.7 mi. N of Govenlock, middle Creek, Bird 1022 (Dao). GUATEMALA: Quiche Dept.: Quiche, pond, Muensclher 12085 (uc). MEXICO: Baia California: La Encantada, Sierra San Pedro Martir, Wiggins & Demaree 4906 (GH, MU, vs). Hidalgo: 632 Rhodora [Vol. 76 DIST. MOLANGO: Molango, Lake Atexca, Moore 3470 (uc). Morelos: Zempoala National Park, Lake Zempoala, Moore 3435 (GH, US). Puebla: vicinity of Puebla, Laguna de San Baltasar, Arsene 1328 (US). San Louis Potosi: San Louis Potosi, Schaffner 533 (K, NY). Tamaulipas: vicinity of San Jose, Sierre de San Carlos, Bartlett 10397 (F, MICH, US). Veracruz: Los Molinos, Perote, pond, Balls & Gourlay (K, MICH, UC, Us). UNITED STATES: Alabama: 7-8 mi. NE of Mobile, Chuckfey Bay, Hotchkiss & Ekvall 3902 (us). Arizona: GILA CO.: Young's Ranch, Lemmon (K, MO, NY, UC, US). Arkansas: MISSISSIPPI C0.: S. of Hornersville, Missouri, Metcalf 644 (GH, NY, US). California: HUMBOLDT CO.: Eel River, near Fern- bridge, Harris 808 (CAN, DAO, F, GH, ILL, MICH, MO, NY, PH, UPS, US). Colorado: ROUTT CO.: ponds between Hayden and Craig, Porter 6563 (CAN, DAO, NY, UC, UPS). TELLER CO.: base of Pike's Peak, Lake Osborne, Parry (GH, MO, NY). Connecticut: MIDDLESEX CO.: Grove Beach, creek, Enquist 366 (NY). Florida: JACKSON co.: N of Sneads, Lake Seminole, Adams 445 (us). Idaho: CUSTER CO.: 15 mi. N of Challis, ditch, Hitchcock et al. 23839 (DAO, ILL, NY, UC). Illinois: COOK CO.: South Chicago, ditches, Hill 89-1875 (F, ILL). Indiana: CASS CO.: Lake Cicott, Deam 49306 (GH). Iowa: CLAY CO.: Lake Twp, Round Lake, Hayden 10120 (GH, MO, NY, PH, UC, US). Kansas: DOUGLAS CO.: 14% mi. E and % mi. N of Lawrence, ponds, Richards 3079 (NY). Louisiana: CAMERON PAR.: Sabine National Wildlife Refuge, Valentine (os). Maine: AROOSTOOK CO.: Oxbow, Aroostook River, Oaden 2661 (os). Maryland: CECIL co.: Blairs Shore, Elk River, Long 57093 (pu). Massachusetts: BERKSHIRE C0.: New Marl- boro, Lake Buel, Churchill (Gn. MO). Michigan: EMMET CO.: small arm of Carp Lake, Haynes 3370 (CAN, FSU, GH, ILL, MICH, MO, MSU, NY. OS, PH, UC, US, WVA), Haynes 3706 (ILL. os). Minnesota: sT. LOUIS CO.: Armstrong, near Ely, Jones 18543 (ILL, NY. US). Missouri: LACLEDE CO.: Gasconade River, between Falcon and Nelio, Steyer- mark 13913 (GH, MO). Montana: GALLATIN CO.: ca. 12 mi. NW of West Yellowstone, Hegben Lake. Haynes 3841 (os). LAKE CO.: 0.2 mi. SE of Nine-Pipes Reservoir, pot-hole, Hitchcock et al. 23980 (CAN, F, ILL, NY, UC). Nebraska: CHERRY CO.: 30 mi. S of Valentine, small pond, Porter 6445 (DAO, NY, UPS). Nevada: STOREY CO.: vicinity of Reno, Sparks, irrigation ditch, Hitchcock 443 (GH, US). New Jersey: SUSSEX CO.: Catfish Pond, Griscom 9750 (GH). New Mexico: SAN MIGUEL CO.: Montezuma, Peterson Reservoir, Drouet & Richards 3308 (F, GH). New York: ST. LAWRENCE CO.: Lisbon, Sucker Brook, Phelps 269 (CAN, GH, NY, US). North Carolina: DARE CO.: Currituck Sound, inlet S of Duck, Radford 5758 (pAo, Uc). North Dakota: BURLEIGH CO.: 7 mi. E of Bismark, slough, Metcalf 356 (US). Ohio: PORTAGE CO.: ca. 54% mi. NW of Ravenna, small pond, Haynes 3450 (os). Oklahoma: COMANCHE co.: Wichita Mountains Wildlife 1974] Potamogeton — Haynes 633 Refuge, Lake Quanah Parker, McMurry 1219 (NY). Oregon: WHEELER co.: 15 mi. E of Mitchell, roadside pond, Hitchcock et al. 23779 (CAN, DAO, F, ILL, NY, UC). Pennsylvania: BERKS CO.: 1 mi. NW of Moselem Springs, Brwmbach 2841 (PH). South Dakota: CODINGTON co.: outlet of Lake Kampeska, Over 17149 (US). Tennessee: OBION co.: Walnut Log, Sharp 7933 (os). Texas: TRAVIS co.: Colorado River, ca. 5 mi. below Austin, Barkley 13313 (F, GH, Mo, PH, UC). WALLER CO.: Hempstead, Hall 620 (F, K, MO, NY, US). Utah: CACHE co.: 1 mi. W of Logan, marsh, Maguire 13883 (CAN, GH, DAO). Vermont: WINDSOR CO.: Windsor, mill pond, Hellquist 1818 (Boston State College). Virginia: FAIRFAX co.: Dyke, Metcalf & Sperry 1688 (GH). Washington: GRAYS HARBOR CO.: slough near ocean beach, Otis 2116 (GH, NY, PH, UC). Wisconsin: LA CROSSE CO.: Lake Onalaska, Hartley 1826 (DAO, F, ILL, Us). Wyoming: ALBANY CO.: Laramie River E of Lookout, Porter 6352 (DAO, MO, NY, UPS). 7b. Potamogeton pusillus var. tenuissimus Mert. & Koch in Rohling, 857. 1823. TYPE: (location unknown). Potamogeton tenuissimus (Mert. & Koch in Róhling) Reichen. Icones Florae Germanica Helveticae 7: 14. 1845. P. berchtoldii var. tenuissimus (Mert. & Koch in Rohling) Fern. & Schubert, Rhodora 42: 246. 1940. Potamogeton berchtoldii Fieber, in Berchtold, Oekon.- tech. Flora Bohmens, 277. 1838. TYPE: (location unknown). Potamogeton berchtoldii var. mucronatus Fieber in Berchtold, Oekon.-tech. Flora Bohmens 277. 1838. P. pusillus var. mucronatus (Fieber) Graebner in Ascherson & Graebner, Das Pflanzen. 4(11): 115. 1907. TYPE: (loca- tion unknown). Potamogeton berchtoldii var. acuminatus Fieber in Berchtold, Oekon.-tech. Flora Bohmens 278. 1838. TYPE: (location unknown). Potamogeton pusillus var. polyphyllus Morong, Bot. Gaz. (Crawfordsville) 5: 51. 1880. P. berchtoldii var. polyphyl- lus (Morong) Fern. Rhodora 42: 246. 1940. TYPE: T. Morong, oozy pool in South Natick, [Middlesex Co.], Mas- sachusetts, (holotype, NY!). Potamogeton pusillus var. elongatus Ar. Benn., in Macoun, Cat. Canadian Plants 4: 371. 1888. TYPE: J. Macoun 4140, Spullamasheen River at and above Enderby, British Colum- bia, Canada, (holotype, K[?]; isotype, GH!). P. pusillus 634 Rhodora [Vol. 76 var. capitatus Ar. Benn., nom. illeg., J. Bot. 39: 201. 1901. (Superfluous when published). Potamogeton pusillus var. cuspidatus G. Fisher, Ber. Bayer. Bot. Ges. 11: 116. 1907. TYPE: (location unknown). Potamogeton lacunatus Hagstrom, Kongl. Svenska Vet- enskapsakad. Handl. 55(5): 120. 1916. P. pusillus var. lacunatus (Hagstrom) Fern. Mem. Amer. Acad. Arts 17: 85. 1932. P. berchtoldii var. lacunatus (Hagstrom) Fern. Rhodora 42: 246. 1940. TYPE: T. Morong, Ashland, in lacu Wauhakune, [Middlesex Co.], Massachusetts, (lectotype here designated, UPS!). Potamogeton turionifera f. tenuis Hagstrom, Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE: J. Macoun, Algonquin Park, Cache Lake, Ontario, Canada, (lectotype here designated, c !; isolectotypes, C ! CAN I, GH!). Potamogeton turionifera f. mucronulatus Hagstrom, Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE: J. Macoun, Ottawa, Brigham’s Creek, Ontario, Canada, (holotype, C!: isotypes, C!, CAN], GH!). Potamogeton pusillus var. colpophilus Fern. Mem. Amer. Acad. Arts 17: 90. 1932. P. berchtoldii var. colpophilus (Fern.) Fern. Rhodora 42: 246. 1940. TYPE: J. F. Collins, M. L. Fernald, & A. S. Pease, brackish pools and dead waters near the mouth of Dartmouth River, Gaspe Co., Quebec, Canada, (holotype, GH!; isotypes, CAN!, NY!, us!). Stem light green to olive, terete to slightly compressed, usually smooth, 22-74 cm long, 0.1-0.5 mm diam. Leaves pale green to olive, rarely rufescent, delicate, 1-3 (-5)- nerved, 0.9-5.4 cm long, 0.2-2.5 mm wide; apex acute to obtuse; glands usually present, green to brown or white, to 0.4 mm diam; lacunae usually present, 1-5 rows each side of midrib; lateral nerves, when present, joining midrib 0.3-1.0 (-2.0) mm below apex. Stipules brown to green, rarely white, delicate, mostly convolute, 3.1-5.0 mm long, 0.2-0.5 mm diam. Winter buds common, lateral or terminal, 1.2-3.2 em long, 0.7-1.2 mm wide; outer leaves 1-3 per side, acute to obtuse. Peduncles cylindrical to slightly clavate, 1974] Potamogeton — Haynes 635 axillary or terminal, rarely recurved, 0.5-4.6 (-6.6) em long, 0.2-0.7 mm diam. Spikes capitate to cylindric, 1.5-7.2 mm long, 0.7-5.2 mm diam; verticels 1-8, when of 2 or 3, then these crowded, 1.2-2.5 mm apart. Perianth segments 0.7- 1.0 mm long, 0.5-0.7 mm wide. Fruit green to brown, usu- ally wider at or below the middle, 1.6-2.1 mm long, 1.2-1.5 mm wide; keels absent; beak usually central, rarely for- ward, 0.2-0.5 mm long, 0.2-0.4 mm diam; sides usually rounded, rarely centrally depressed; wall texture smooth. Chromosome number, n = 13; 2n = 26 (Taylor & Mulligan, 1968). Distribution: Central Alaska to Labrador, south to Cali- fornia and northern Florida. Fruiting from early June to late September. Fig. 12. Fig. 12 Map of North America showing documented distribu- of Potamogeton pusillus: var. tenuissimus. 636 Rhodora [Vol. 76 Illustrations: Fernald (1932, pl. 16-20; 30, fig. 6-9; 31, fig. 1; 35; 39, fig. 9). REPRESENTATIVE SPECIMENS CANADA: Alberta: CRAIGMYLE DIST.: Craigmyle, Brinkman 538 (us). British Columbia: QUEEN CHARLOTTE ISLANDS: Graham Island, 2 mi. NW of Tlell, n — 13, Calder & Taylor 35688 (DAO); Moresby Island, White Swan Lake, 2n — 26, Calder & Taylor 35297 (DAO). VANCOUVER ISLAND: Lost Lake near Victoria, Macoun (CAN, GH, NY, us). Labrador: Goose Bay, 53°20'N, 60°09'W, Gillett & Findlay 5910 (DAO, NY). Manitoba: CHURCHILL DIST.: Gillman Lake, Schofield 1524 (DAO, GH). New Brunswick: CHARLOTTE CO.: Grand Manan, Weatherby & Weatherby 7284 (CAN, GH). Newfoundland: Harry’s River, near Bay of St. George, Fernald & Wiegand 2453 (CAN, GH, NY). Northwest Territories: small pool 45.5 mi. WNW of Yellow- knife, Thieret & Reich 7944 (CAN, F, OS, us). Nova Scotia: Sable Island, St. John 1125 (CAN, GH, NY, PH, us). Ontario: THUNDER BAY DIST.: Black Fox Lake, 12 mi. E of Terrace Bay, Haynes 351a (os); small lake 0.2 mi. NE of Coldwell, Haynes 3773 (ASC, CAN, FSU, GH, ILL, MICH, MO, MSU, NY, OS, PH, UC, US, WVA). ESSEX CO.: Fox’s Pond, Haynes 3251 (0s), Roberts 1567 (oS). Prince Edward Island: QUEENS C0.: Watervale, Erskine (DAO). Quebec: MAGDALEN ISLANDS: ridges back of the Narrows, Alright Island, Fernald et al. 6778 (CAN, GH, K, NY, US). Saskatchewan: Cornwall Bay, Lake Atabaska, 59?27'N, 108°27'W, Raup 6633 (CAN, DAO, F, GH, NY). Yukon: Lewes River, Whitehorse, Porsild & Breitung 10644 (CAN). UNITED STATES: Alaska: Goldstream Creek, 51 mi. N of Fair- banks, 65°N, 147?30'W, Porsild & Porsild 115 (CAN, GH, US). Arkansas: DREW CO.: Taxodium swamp, Tillar, Demaree 17394 (F, MO, NY, UC). Arizona: COCONINO CO.: Crater Lake, N of Mt. Agassy, Lemmon 8244 (GH). California: BUTTE CO.: cold spring, Jonesville, Copeland 604 (F, GH, K, MICH, MO, MU, NY, uc, us). Colorado: JACKSON CO.: Big Creek Lake, Porter 6433 (DAO, ups). Connecticut: TOLLAND CO.: Stafford, Morris (F, GH, ILL, us). Florida: FRANKLIN co.: 6 mi. NE of Carrabelle, Adams 442 (GH, UC, US). Idaho: SHOSHONE CO.: St. Maries River, 7 mi. SE of Clarkia, Cronquist & Jones 6038 (CAN, DAO, GH, ILL, MICH, NY, UPS, UC, us). Illinois: COOK co.: South Chicago, Hill 88-1880 (ILL). Indiana: STEUBEN CO:. Crooked Lake, Williams (MO). Iowa: DICKINSON co.: Manhattan Pond, Lakeville Twp. Thorne 13015 (uc). Louisiana: CALDWELL PAR.: Horseshoe Lake, 8 mi. NNW of Columbia, Thomas 4224 (os). Maine: PENOBSCOT CO.: Stillwater River, Old Town, Ogden & Steinmetz 1600 (CAN, GH, NY, us). Maryland: HARFORD CO.: Spesutie Island, Moldenke 9398 (NY). Massachusetts: NORFOLK CO.: Charles River, Dedham, Fernald & Svenson 410 (CAN, DAO, F, GH, ILL, MICH, MO, 1974] Potamogeton — Haynes 637 NY, UC, UPS, US). Michigan: CHEBOYGAN CO.: Black River, between its mouth and Alverno, Haynes 3378 (GH, ILL, os, US), Haynes 3387 (os); Marl Bay NW corner of Douglas Lake, Haynes 3712 (GH, ILL, MO, 0S, UC), Haynes 3924 (os). LUCE CO.: Bodie Lake, Haynes 3790 (os); bog lake, 612 mi. NE of Newberry, Haynes 3718 (08). SCHOOL- CRAFT CO.: Seney Wildlife Refuge, Haynes 3741 (os). Minnesota: KOOCHICHING CO.: ditch 2 mi. E of Tilson Bay, Rainy Lake, Moore & Moore 11778 (DAO, GH, ILL, NY, UC, US). Montana: MISSOULA CO.: Ft. Missoula, Hitchcock 23968 (DAO, NY, UC). Nevada: ELKO CO.: Hot Creek Field, NE of Golliher Pasture, Holmgren 1401 (DAO, NY, uc). New Hampshire: CHESHIRE CO.: East Jaffrey, Deane (GH). New Jersey: CAMDEN CO.: Newton Creek, West Collingswood, Adams 370 (GH, MO, PH). New York: CORTLAND CO.: borrow pit just N of Little York, Haynes 3353 (0S). ESSEX CO.: outlet of Lake George, near Ticonderoga, Haynes 3336 (08). SARATOGA CO.: Saratoga Lake, Haynes 3301 (os), Haynes 3307 (0S). SUFFOLK CO.: Wading River, Riverhead, Miller (F, GH, US[4]). WARREN C0.: Dunham Bay, S end of Lake George, Haynes 3326 (GH, MICH, MO, NY, OS, US), Haynes 3329 (CAN, GH, ILL, MICH, MO, NY, OS, PH, UC, US); Lake Luzerne, Haynes 3812 (os). North Carolina: CRAVEN CO.: 2 mi. SW of Blades, Radford 5856 (DAO, GH, NY). North Dakota: STUTSMAN CO.: Jim Lake, Pingree, Mabbott 316 (NY). Ohio: JACKSON CO.: Sec. 10, Jefferson Twp., Roberts 795 (CAN, LAF, MICH, OS, PH, US). LOGAN co.: Indian Lake State Park, Haynes 3418 (08). OTTAWA CO.: Mid- dle Bass Island, Lake Erie, Haynes 3134 (os); Squaw Harbor, Put-in-Bay, Lake Erie, Haynes 3127 (MICH). PORTAGE CO.: Twin Lakes, Haynes 3448 (oS). VINTON CO.: Lake Alma State Park, Haynes 3468a (os). Oklahoma: COMANCHE C0.: Creek near Cache, Stevens 1369 (GH, ILL, K, MO, NY, US). Oregon: Porter Lake of Willamette River, 5 mi. S of Corvallis, Wentz 344 (MICH, MO, NY, os, vs). Pennsylvania: LUZERNE CO.: Lily Lake, Small (F, ILL, MU, NY). Rhode Island: NEWPORT CO.: Block Island, Fernald et al. 8448 (PH). South Carolina: GEORGETOWN CO.: Georgetown, Forster (NY). Texas: BOWIE COo.: Club Lake, 3 mi. W of New Boston, Correll & Ogden 25248 (LAF). Vermont: CALEDONIA CO.: beaver pond, Walden Twp., Haynes 3828 (MICH, OS). ESSEX CO.: Bruns- wick, Eggleston 1654 (GH, NY, US). LAMOILLE CO.: Wolcott Pond, Wolcott. Haynes 3826 (os). Virginia: SUSSEX CO.: Chappell’s Mill- pond, W of Lumberton, Fernald & Long 12236 (GH, PH). Wash- ington: KLICKITAT CO.: Columbia River, Bingen, Suksdorf 2570 (F, GH, Mo, US). Wisconsin: JUNEAU CO.: Glacial Lake Wisconsin, Cutler Twp. Hartley 8276 (GH, US). Wyoming: ALBANY CO.: Swas- tika Lake, Medicine Bow Mountains, Porter 6190 (DAO, GH, UC, UPS). CROOK CO.: pool 7 mi. N of Moorcroft, Porter & Miller 5974 (DAO, MO, UC, UPS). 638 Rhodora [Vol. 76 Potamogeton perfoliatus X pusillus var. tenuissimus Morong (1880) named Potamogeton mysticus based upon a collection from Mystic Pond, Massachusetts. The entity was said to have the habit of P. perfoliatus, a species of subsection Perfoliati of Hagstrom, but differed from this species by its delicate stature. Morong noted that the entity had never been known to fruit. Ogden (1943), however, based upon the extreme rarity of the entity and that it was not known to fruit, proposed it to be a hybrid with P. per- foliatus as one of its parents. He examined the stem anat- omy of the entity, as well as studying the plants in the field. The stem anatomy appeared to be intermediate between P. perfoliatus and the Pusilli. At the one locality in which Ogden found the putative hybrid, P. berchtoldii var. tenuis- simus [P. pusillus var. tenuissimus] was present. He thus suggested the entity to be a result of hybridization of P. perfoliatus and P. berchtoldii var. tenuissimus [P. pusillus var. tenuissimus]. Again, until the entity can be studied in depth, I am accepting the concept of Ogden. SPECIMENS EXAMINED: UNITED STATES: Maine: CUMBERLAND CO.: Scarboro, West Scarboro, Scarboro River, Ogden et al. 1731 (CAN, MO, NYS, US); Scarboro, West Scarboro, Stuart Brook, Steinmetz & Martson 539 (CAN, F, ILL, UC, US). Massachusetts: MIDDLESEX CO.: Medford, Mystic Pond, Booth (US), Faxon (ILL, US), Morong 11 Aug. 1879 (lectotype here designated, Ny!; isolectotypes, K!, MICH!, NY[3]!), Morong (K, NY). North Carolina: DARE CO.: Currituck Sound, inlet S of Duck, Radford 5757 (GH, NY, UC). 7c. Potamogeton pusillus var. gemmiparus Robbins in A. Gray Man. Bot. North. U. S. 5th ed. 489. 1867. TYPE: Robbins, Valley of the Blackstone, Uxbridge, Massachusetts, (lectotype here designated, NY!; isolectotypes, GH!, NY!, PH !, uS!). Potamogeton gemmiparus (Robbins in A. Gray) Morong, Bot. Gaz. (Crawfordsville) 5: 51. 1880. Stems pale green, terete, smooth, 30-140 em long, 0.1-0.3 mm diam. Leaves pale green, uninervate, 1.1-6.0 cm long, 0.2-0.7 mm wide; apex subulate; glands gold, to 0.8 mm 1974] Potamogeton — Haynes 639 diam; lacunae absent or to 2 rows each side of the midrib. Stipules white, convolute, 4.1-7.2 mm long, 0.2-0.4 mm diam. Winter buds abundant, terminal or lateral, 1.2-2.8 cm long, 0.7-1.5 mm wide; outer leaves 1-2 per side, subulate, much longer than inner. Peduncles cylindric, axillary or terminal, 1.1-2.8 em long, 0.2-0.5 mm diam. Spike cylindric, 2.5-5.5 mm long, 1.2-3.0 mm diam; verticels 2-3, to 3.5 mm apart. Perianth segments 1.0-1.7 mm long, 0.7-0.8 mm wide. Fruit rare, greenish-brown, 1.9-2.0 mm long, 1.5-1.6 mm wide; beak central, 0.1-0.2 mm long, 0.2-0.3 mm diam; sides cen- trally depressed; wall texture rough. Chromosome number unknown. Distribution: South-central Quebec south to central Mas- sachusetts and Rhode Island. Fruiting late August to late September. Fig. 15. Illustrations: Fernald (1932, pl. 12; 30, fig. 1; 34, fig. 1). o b \ y X 4 em Y ~y N P , x I: v om \ g Y \ "4 iu ws S \ " AD | 4 D 3 g Y e. . T , c ‘As P \ art o 9 At uU e a d D j i" à >Ê c - \ Q 4 [e] - A p ` E l t e q Pa i bo ; " © Le Eon \ [*] i ^ A A P OE c i n- 1 Fig. 13. Map of northeastern United States and southeastern Canada showing documented distribution of Potamogeton pusillus var. gemmiparus. 640 Rhodora [Vol. 76 REPRESENTATIVE SPECIMENS CANADA: Quebec: Saint-Tite, comte de Champlain, Lake Pierre- Paul Gauthier 2273 (GH, photo at DOA). UNITED STATES: Maine: CUMBERLAND CO.: Horseshoe Pond, North Fryeburg, Hellquist 4457 (MICH). KENNEBEC CO.: Belgrade Stream, Belgrade, Hellquist 4501 (MICH). PENOBSCOT CO.: Upper Stillwater, Fernald (uc, us), Fer- mald (GH). Massachusetts: HAMPSHIRE CO.: Amherst, Jesup (GH, NY). MIDDLESEX CO.: Charles River, South Natick, Morong (NY). NORFOLK CO.: Charles River, Dedham, Fernald & Svenson (GH). WORCESTER CO.: Uxbridge, Morong (CGE, F, ILL, MICH, MO, NY, US). New Hampshire: CARROLL CO.: Conway Lake, Eaton, Hellquist 4469 (MICH). HILLSBOROUGH CO.: Pine Island Pond, Manchester, Kroch- mal 265 (GH). Rhode Island: PROVIDENCE CO.: Central Pond, Ten Mile River, East Providence, Collins (GH). 8. Potamogeton groenlandicus Hagstrom, Kongl, Svenska Vetenskapsakad. Handl. 55(5): 127. 1916. TYPE: Hartz, Kingua Orpiksnit, 68°30’, Greenland, (lectotype here desig- nated, C!). Potamogeton pusillus ssp. groenlandicus (Hagstr.) Boch. 1952. Meddel. Groenland 147 (9) : 44. Stems light green, terete, smooth, 20-50 cm long, 0.2-0.3 mm diam. Leaves pale to deep green, 7-9 nerved, 2.6-8.8 mm long, 0.7-1.7 mm wide; apex acute to apiculate; glands usually present, white, 0.1-0.7 mm diam; lacunae absent or to three rows present each side of the midrib; lateral nerves joining the midrib 0.7-1.2 mm below the apex. Stipules brown, delicate, convolute, 4.5-19.1 mm long, 0.2-0.4 mm diam. Winter buds common, terminal or axillary, 4.2-8.4 em long, 1.4-2.5 mm wide; inner leaves unmodified; outer leaves 2 per side, acute to apiculate, without corrugations at base. Peduncles rare, cylindric, terminal, ca 18.2 mm long, ca 0.5 mm diam. Spikes cylindric, ca 5.0 mm long, ca 3.5 mm diam; verticels 3, crowded, ca 1.0 mm apart. Perianth segments ca 1.1 mm long, ca 0.7 mm wide. Fruit unknown. Chromosome number unknown. Distribution: Western Greenland. Flowering in late August. Fig. 14. Illustrations: (apparently none). 1974] Potamogeton — Haynes 641 In 1916, Hagstrom named Potamogeton groenlandicus from several sterile specimens. He distinguished this spe- cies from other Pusilli by its possessing 5-9 nerves per leaf. “In this respect,” he stated, “this species constitutes an interesting link between the Pusilli and the Oxyphylli and would be ranked with the latter as well as with the former." He added that the ligules and leaf-apices, however, unite it with the Pusilli. Fernald (1932), after examining as many specimens as available [but not the Type], concluded the taxon to be conspecific with P. pusillus var. mucronatus [var. tenuissimus]. He stated that he had been unable to convince himself that the leaves were more than three- nerved. He added that this character alone does not war- rant specific rank. I have examined the material at Copen- hagen and at all the large herbaria of the United States and Canada. In my opinion, the leaves are 5-9 nerved. I agree with Fernald, however, that one vegetative character is not enough to separate two Species, especially when one is Fig. 14. Map of Greenland showing documented distribution of Potamogeton groenlandicus. 642 Rhodora [Vol. 76 known only from non-fruiting material. However, both Fernald and Hagstrom overlooked the winter buds. In P. groenlandicus, the inner leaves of the winter buds are not modified into hardened fusiform structures, as in P. pusillus. Instead, the winter buds are similar to those of P. obtusi- folius, in that the inner leaves are unmodified. To my knowledge, only one population of fertile individuals has been found, this being in flower. The flowers are most similar to P. pusillus. Because the winter buds and number of nerves are so different from P. pusillus, I am currently accepting P. groenlandicus at the specific level. However, material should be gathered and cultivated. If fruiting does occur, then the correct affinities of this endemic taxon can be ascertained. REPRESENTATIVE SPECIMENS GREENLAND: Christianshaab Dist.: Christianshaab, Hartz (c). Egedesminde Dist.: Egedesminde, 67?44'N, Porsild (CAN, GH, MO, NY, US). Frederikshaab Dist.: Frederikshaab, 62^00'N, 49°40'W, Jorgensen & Larsson (CAN); Ivigtut, Berlin (c). Godhavn Dist.: Disko, near Arktisk Station, 69?15'N, Porsild (CAN, F, GH, US) ; Disko, Godhavn, Laegaard 350 (CAN, Us); Disko, near Godhavn, 69?14'N, Porsild 228 (GH, US); Disko, Sinigtik, 69°25'N, Porsild (CAN, GH); Disko, Skauseklippen, 69°25'N, Porsild (F). Godthaab Dist.: vicinity of Godthaab, 64°10'N, 51°43’W, Porsild 8273 (CAN) ; Godthaab Fjord, Itivnera, 64°22'N, Porsild (GH) ; Godthaab Fjord, Jersiutilik, Trap- nell 402 (GH, K). Jakobshavn Dist.: Jakobshavn, Sorensen (C). Julianehaab Dist.: Igaliko Fjord, Qagssaarssuk, Polunin 10779 (CAN); vicinity of Julianehaab, 60^43'N, 46°05'W, Porsild 8120 (CAN). Ritenbenk Dist.: Ritenbenk, 69°44'N, Hansen (C); Sarkak, pool on gmeiss, Seward & Holttum (cGE, K); 5 km E of Sarkak, Beschel 12163 (CAN, photo at DAO). Sukkertoppen Dist.: Ungarsivik Lake, 65?25'N, 52^50'W, Porsild 8870 (CAN). Upernavik Dist.: Laksefjord, Proven, Porsild 8837 (CAN). ACKNOWLEDGMENTS I am grateful to Dr. Ronald L. Stuckey, who supervised this study, for his advice and constant encouragement dur- ing the course of my work. Appreciation is also owed to Dr. Edward G. Voss for help with nomenclature; to Miss Judy Canne, Mr. Robert C. Gardner, and Mr. David Keil 1974] Potamogeton — Haynes 643 for their helpful suggestions; to the curators of the herbaria from which specimens were borrowed; to the staffs at Kew Botanical Gardens and the British Museum for their pains- taking search for the holotype of Potamogeton strictifolius; to the Linnean Society of London for the photograph of the holotype of P. pusillus; to my colleagues at the many librar- ies who supplied xerox copies of many original descriptions; to Mr. Robert C. Gardner, Dr. Pete Hostetter, and Mr. Robert Kalinsky for their help with the photographs; to Dr. Eugene C. Ogden, Mr. Stanley Smith, Dr. Edward G. Voss, and Dr. Steven Young for help in locating populations of Potamogeton in the field; and to Mr. Marvin L. Roberts, Dr. John A. 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DEPARTMENT OF BIOLOGICAL SCIENCES LOUISIANA STATE UNIVERSITY IN SHREVEPORT SHREVEPORT, LOUISIANA 71105 SARRACENIA ALABAMENSIS, A NEWLY RECOGNIZED SPECIES FROM CENTRAL ALABAMA' FREDERICK W. CASE, JR. AND ROBERTA B. CASE In February, 1953, while collecting pitcher plants at Elmore, Alabama, for taxonomic studies, we located a colony of Sarracenia with frozen leaves. As we were seek- ing plants of S. oreophila (Kearney) Wherry at Wherry's published range extension for that species at Elmore, and since the frozen leaves were mostly large and widely ex- panded at the orifice, much resembling the published photo- graphs of the yellow flowered S. oreophila, which we had not at that time seen, we assumed our discovery to repre- sent that species. When the plants flowered in our green- house, however, the flowers were small and red. This seemed to place the plant in S. jonesii Wherry, which was widely accepted at that time. After observing and growing the plant for two seasons, we became convinced that either the published description of S. jonesii was too narrow, or that the plant was not S. jonesii at all. Our next step was to visit the type locality of S. jonesii in North Carolina to observe and obtain plants for comparison. They proved not to be the same, so we began the studies leading to this paper. With the help of the late Dr. R. M. Harper, who made his field notes available to us, and who, in 1955, guided us to his colonies of pitcher plants in Central Alabama, we were able to obtain a wide sampling of the population and to locate other colonies of our own. Because there has long been controversy and confusion among Sarracenia taxon- omists over the treatment of the taxa related to S. rubra Walter, we visited the Elmore and Chilton County colonies eight times at various seasons in the period from 1953 to 1973 to better understand this related species. 'Manuscript received January, 1973. 650 1974] Sarracenia — Case & Case 651 Since 1953, we have grown this plant, which we call the Alabama Canebrake Pitcher Plant, continuously, both in our greenhouses, and, since 1966, outdoors as well, at Sag- inaw, Michigan. Although its small red flowers with obo- vate petals indicate a close relationship to S. rubra, it shows a number of differences. In this connection, those interested in significant taxonomic characteristics in Sarracenia may wish to refer to the remarks of MacFarlane (1908), Wherry (1929, 1935), Bell (1949), and MeDaniel (1966). Unlike Sarracenia rubra (MacFarlane, 1908) our plant produces ftvo types of pitchers and occasional phyllodia each season. Its spring pitchers, similar in size to the leaves of S. rubra, differ markedly in developing a curved form and in lacking the external dark maroon coloring of the veins, pitcher rim and hoods found in that species. The summer pitchers, straight and usually much larger and more tapered than those of S. rubra (excepting S. rubra ssp. jonesii), differ from those of both of those taxa in possessing a softer texture, a visibly fine pubescence, and a paler yellow-green color. The major veins are maroon col- ored mainly only on the inside of the pitcher tube. The hoods of the summer pitchers are from 2-8 times larger than those of summer leaves produced in S. rubra, The greater expansion of the hood, the stronger reflexion of the hood neck margins, the strongly rolled orifice rim, the leaf size, and overall yellowish green coloration give the summer leaves a profile more suggestive of the pitcher of S. flava or S. oreophila than of S. rubra. The flowers, while less useful than the leaves for species delineation within this genus (c.f. Bell, 1949, p. 140), differ from Sarrecenia rubra in their larger size, lighter red coloring, petal shape, in their erose petal margins, and in their fainter fragrance (Fig. 1). In addition, the plant produces flower scapes abundantly; often two or three scapes develop from one terminal growth in vigorous plants, making large flowering clumps spectacular. The regular production of multiple blooms from one terminal bud is an unusual feature in Sarracenia. 652 Rhodora [Vol. 76 Figure 1. Comparison of flowers of Sarracenia alabamensis, Chil- ton Co. Ala. (left) and S. rubra, Crestview, Florida (right). Note differences in color tone, shape of petal lobe, and detail of petal margins. Since, in addition to its structural differences, its range is disjunct from Sarracenia rubra and probably from that of all other species except S. oreophila, we consider it a dis- tinct species. MacFarlane (1908) has pointed out that all immature Sarracenia leaves tend to resemble closely those of Sar- racenia minor Walter and do not show good specific dif- ferences. While he spoke mostly of very young pitcher 1974] Sarracenia — Case & Case 653 leaves, we believe that all specific differences are best dis- played by the largest leaves of the season. In our descrip- tion we give the usual size ranges, but we urge readers to standardize their comparisons, when possible, by examining the largest seasonal leaves of related taxa. Because we believe that vague size references without standardized points of measurement have contributed to the problems in understanding the taxa related to S. rubra, we have tried to be quite exact in our measurements. Pitcher length was measured from the point of attachment to the rhizome to the rim of the pitcher orifice; it does not include the hood. The length of hood refers to the distance from the narrow- est part of the hood-constriction (neck) to the tip of the hood, and hood width refers to measurement across its widest point. To facilitate measurement comparisons of the tubular pitchers in fresh or herbarium material, we give width figures for pressed pitchers rather than diam- eters for inflated pitchers. The panduriform petals present problems for accurate measurement comparisons, being divided into a cuneate basal portion, an isthmus, and an obovate distal portion. For purposes of comparison, the length of the basal cuneate portion of the petal was measured as the distance from the petal attachment along the median line to the widest point of the segment; the length of the distal petal lobe we measured from the widest point of the basal portion to the tip of the petal. We have based the following description upon both culti- vated specimens and those collected in the wild. Sarracenia alabamensis Case & Case, sp. nov. Rhizoma valde ramosum, ramis confertis; folia trimor- pha; folia vernalia ascidiformia, maxima plerumque plus minusve sigmoidea, tubo basi angusto, sursum gradatim ampliato, 17.7-49.5 cm. longo, 0.7-3 cm. lato; ala ventralis [adaxialis] ad medium expansa, 0.4-1.5 em. lata, versus orem et basin attenuata; tubi venae extus plerumque in- coloratae, intus atropurpureae; operculum cordatum, acu- 654 Rhodora [Vol. 76 tum, subundulatum vel convexum, 1.9-7 cm. longum, 1.7-6 cm. latum, venis intus prope basin coloratis; collum 0.8-3 em. latum. Folia maxima ascidiformia aestivalia quam vernalia multo majora, foliis S. flavae vel S. oreophilae similia, suberecta, recta vel subsigmoidea, extus dense pubescentia, 24.5-48.5 em. longa, ad basin longe angustata, in quarta parte superiore parum ampliata, 3.1-6.7 em. lata; peristom- ium valde revolutum, adaxiale incisum; ascidii textura tenuis, mollis, ex salmoneo flavo-viridis, demum pallide flavo-viridis; tubus prope orem extus obscure areolatus, areolis albidis, venis intus valde purpureis, extus plerum- que (maximis interdum exceptis) incoloratis; operculum magnum, expansum, cordatum, fere tam latum quam longum, 3.7-9.5 cm. longum, 3.4-8.6 cm. latum; collum 1.3- 3.4 cm. latum; textura inter venulas convexa; operculum salmoneum, demum pallide flavo-viride, intus (necnon praecipue collum peristomiumque) iridescens. Phyllodia laminaria pauca, interdum e gemmis terminali- bus majoribus oriunda, ascidia vernalia sequentia, aestivalia praecedentia, prope basin valde recurvata, ultra medium latiora, apicem versus denuo recurvata, 4-6.5 cm. longa, 0.3-0.8 em. lata. Folia squamiformia pauca, triangularia, amplitudine admodum variabilis. Scapi numerosi, interdum aliquot per ramulum termina- lem, 34.5-57.6 em. longi. Calyx applanatus 4.5-6 cm. latus. Sepala ovata, rotundata vel subacuta, 1.7-2.6 cm. longa, 1.1-2.2 cm. lata, sub anthesi faleato-incurvata, ovarium mox cingentia, demum patentia vel reflexa, marginibus vetustate valde revolutis. Petala panduriformia, 3.4-4.6 cm. longa, extus sanguinea (anglice “Oxblood Red"); pars basalis cuneata 0.7-1 cm. longa, basi 0.3-0.4 em. lata, maxime 0.9-1.1 cm. lata; con- strictio mediana marginibus reflexis; lamina ultra constric- tionem in isthmum brevem parallelilateralem tum in lobum 1974] Sarracenia — Case & Case 655 obovatum erose marginatum 2.7-3.6 cm. longum, 1.8-2.5 cm. latum expansa. Ovarium tuberculatum, paullo compressum, 0.8 cm. longum, 0.9 cm. latum. Stylus umbraculiformis, umbraculo valde convexo, 3.5-4.2 em. lato. TYPE: Alabama: ELMORE CO.: very rare and local in wet gravelly or peaty soil and around springheads and rills in open grassy swales, and in thickets of Rhododendron, Alnus, Magnolia virginiana, Rhus vernix and Arundinaria tecta; along the railroad between Elmore and Speigner. Type plant collected in June 1971, Case & Case S-500 (us), but all specimens prepared from clonal divisions grown in our greenhouse and taken May 1, 1972 (flowering) and July 27, 1972 (large summer leaves). REPRESENTATIVE SPECIMENS EXAMINED Alabama: CHILTON CO.: Clanton, Pollard & Mason (us); sloping gravelly bog near Jasmine, 28 Apr. 1921, Harper (NY, Us); Jasmine, 18 Apr. 1921, Harper (NY); gravelly bog, near Adams, 17 June 1955, F. & R. Case, C-100 (MicH), C-101, C-102 (NY); boggy rill feeding Mulberry Creek, Pletcher, 17 June 1955, F. & R. Case et al., C-108 (us); cultivated material, Canebrake-Magnolia glauca bog, Adams, grown at Saginaw, Michigan, F. & R. Case 8-520, S-522 (us), S-524, S-527, S-528 (Ncvu), S-521, S-525, S-526 (MICH); boggy soil around springhead, with Arundinaria tecta, Magnolia virginiana, Jas- mine, 19 Apr. 1973, F. & R. Case C-104 (NcU), C-105 (US). ELMORE co.: Slightly damp sandy soil 112 miles S. of Speigner, 14 Apr. 1934, Harper (PH); boggy depression along railroad near station, Elmore, 13 July, 1935, Wherry (PENN); approximately 114 miles north of Elmore along railroad, damp boggy ground, specimen from a culti- vated plant (Oct. 1955), collected 16 Feb. 1953, F. & R. Case, C-106 (us). DISTRIBUTION AND ECOLOGY This extremely rare plant is known to occur only in small, very localized colonies among the Fall Line Sand Hills of Chilton, Autauga, and Elmore Counties, Alabama, north of the Alabama River in an area roughly bounded by the Coosa River to the east, and the Mulberry River to the west. The soils there derive from rocks generally regarded as Upper Cretaceous in origin (Harper, 1922). Except for Wherry’s report of Sarracenia oreophila at Elmore, Ala- 656 Rhodora [Vol. 76 bama, no other Sarracenia is reported to grow within the range of S. alabamensis. Sarracenia alabamensis occurs in somewhat sandy-grav- elly bogs, usually on sloping ground, or in damp peaty or mucky soil around small spring-heads and tiny rills. It is not usually a plant of sphagnum beds although species of Sphagnwm may occur in isolated patches nearby. The colonies we have observed are extremely small compared to those of most of the other southern species, and range from single clumps to one group of perhaps 100 plants in a 20 by 50 foot area. According to Harper's (1922) ob- servations and our own, the following plants commonly grow with the Canebrake Pitcherplant: Magnolia virgin- tana, Alnus sp. Arundinaria tecta, Acer rubrum, Rhus vernix, Smilax laurifolia, Osmunda cinnamomea, Pteridium aquilinum var., Juncus sp., Drosera capillaris, Pinguicula sp., Aletris aurea, Rhexia sp., Pogonia ophioglossoides, Rhododendron sp., and Calopogon tuberosum. The pitcher plant grows most often in open stands of Magnolia virginiana, Arundinaria tecta, and Alnus, in the “Canebrakes” of the region, or in open boggy swales. Tol- erant of more shade than most of the trumpet-leaf sarra- cenias, the species nevertheless reaches its most vigorous and distinctive development when growing in full sunlight. Although few of the large, distinctive summer leaves de- velop on heavily shaded plants, the species is able to flower sparingly even when so shaded that not even the curved spring pitchers can develop without severe deformity. Hardest to detect of the sarracenias, this species probably has always been very limited in distribution in modern times, for its habitat is very restricted and by nature it occurs in small stands among other plants. But recent de- velopments within its range have caused further restriction. In 1921, Harper (1922) reports, the bogs in which Sar- racenia alabamensis grew were open, with woody plants rather small and scattered, the bulk of the vegetation sun- loving. It was still generally thus at both Elmore and in Chilton County when we first saw this species in 1953 and 1974] Sarracenia — Case & Case 657 1955. At that time the swamps were shrubby but open, and small rills lined with pitcher plants were frequent. The entire area was sparsely inhabited, remote, and the roads were limited, temporary, and unditched. In contrast, on visits in the past few years, we found much change. The bogs have not burned in some years. No longer are they open ; dense stands of magnolia and maple, solid canebrakes, tangles of Smilax, and alder, rob the bogs of light. The sarracenia is much rarer today, and seedlings scarcely can be found. Good roads reach the area, and farms, cattle raising operations, use of herbicides, ditching and draining, all have taken considerable toll of the habitat. Perhaps the greatest change in the pitcher plant habitat here has been the invasion of the bogs and openings by the Japanese honeysuckle, Lonicera japonica Thunb., which destroys vir- tually all native species in the bogs it invades. DISCUSSION The Alabama Canebrake Pitcher Plant has many features of interest. It is the only trumpet leaved Sarracenia to produce pitchers which differ significantly in size and form with the season (Fig. 2). The usually recurved, shorter, spring pitchers are less voluminous than the often very much larger, erect, summer pitchers. In these spring leaves, the laminar wings expand at or just below the middle, pushing the pitcher into a loosely sigmoid curve. The upper, sharply expanded part of the tube and hood is erect. In effect, these early leaves seem almost deformed, as in a plant growing under dras- tically shaded conditions or in one which shows the effects of an application of an herbicide. This leaf form, produced in other species only under conditions of ecological duress, is entirely normal and comprises the usual spring pitcher in this species, even on plants growing in full sun (Fig. 2). But occasionally, on extremely vigorous, uncrowded plants in very wet, sunny situations, the plant produces the large summer type pitchers without first producing the recurved leaves. 658 Rhodora [Vol. 76 j2 328 is 15 ie i z 4 4 Figure 2. Sarracenia alabamensis. Three spring type, curved leaves (left) and three of the larger, straight summer leaves (right). Note the obscure white, areole-like markings near summit of pitcher tube on the right-hand specimen. 1974] Sarracenia — Case & Case 659 A single clone may produce either type of pitcher in the first wave of growth in different years, but the trend is clearly toward recurved spring pitchers. In a group of about 50 study clones grown in our greenhouses, roughly 90% of the plants will produce the recurved pitchers in a given season. In the wild, in more competitive (and per- haps darker) situations all except two of several hundred clones we have observed have produced the recurved spring pitchers. These two plants had produced only the large summer-type pitchers that season. The variable form of the early leaves of the season, some- times as recurved, reduced smallish pitchers with an un- usual development of wing tissue, sometimes as fully de- veloped phyllodia, and occasionally as erect, summer-type pitchers, suggests to us that these variations occur at least partly in response to a combination of degree of light in- tensity and increasing daylength. Phyllodia and the re- curved, reduced pitchers do not reappear in the autumn; at that time only smaller, fully erect, summer-type pitchers develop. MacFarlane (1908) has stated that phyllodia represent an advanced leaf type, developed by reduction of the su- perior pitchered portion of the wing while the inferior lamina remains full sized or enlarged. He notes that oc- casional leaves intermediate between normal pitchers and phyllodia appear in all the phyllodium-forming species. We suggest that the variable structure of the spring leaves in Sarracenia alabamensis represents a stage in the evolution of a laminar phyllodium from a pitcher, one in which the exact degree of pitcher reduction in response to daylength and light intensity has not yet become stabilized. The fact that these variable leaves occur only in spring and that in other species which regularly produce phyllodia, such leaves are strictly seasonal, seems to strengthen this interpreta- tion. The summer pitchers of the Canebrake Pitcherplant are fully erect, remarkable for their relatively large size, and great volume. The tissue of these large summer pitchers 660 Rhodora [Vol. 76 possesses a soft substance lacking the waxy-thickened qual- ity of the other trumpet leaf species. This, together with the fine, dense pubescence, gives to the pitcher a unique "feel" prompting Dr. Wherry to remark that this is the only “rubra type" Sarracenia one can readily identify in the dark. Other features of the summer pitcher include its very light green color when grown in bright light, and the pres- ence in many leaves of a few rather obscure, whitish areole- like areas scattered on the outer pitcher-tube surface near its orifice. These cannot be seen from within. These faint areoles seem to be the consequence of the extreme tissue expansion characteristic of the summer pitchers. They are not structurally quite the same as those in the truly areolate species, but they may offer interesting evidence as to the mode of origin of areoles within this genus. Because all taxa related to Sarracenia rubra have become controversial and because other taxa in the rubra alliance do not show areoles at all, some botanists may seize upon this feature as evidence that S. alabamensis represents a hybrid popu- lation of some sort, not a species. We cannot regard these areole-like tissues as evidence of any recent hybridization with an areolate species for sev- eral reasons, The areolae do not appear on the spring leaves in this taxon; in the known hybrids between other areolate and non-areolate species, the areolae occur in all leaves of mature plants and even on the larger seedling leaves. In known areolate hybrids, the white transparent areoles extend from outer to inner tube tissues. Also, no areolate species occurs in contact with the known range of this species. If these areole-like formations be of hybrid origin, it is ancient hybridization and the whitish markings have become a fixed although obscure characteristic of the larger summer leaves. Although we have already stated our interpretation of the nature of the recurved spring pitchers above, the inter- mittent production of full phyllodia in Sarracenia ala- bamensis, coupled with Wherry's report of S. oreophila 1974] Sarracenia — Case & Case 661 within its range at Elmore, may lead some to say that S. alabamensis represents a hybrid swarm of S. oreophila with S. rubra. Several points ought to be made in this connec- tion. Since S. oreophila is a large yellow-flowered species, and S. rubra a small red-flowered one, any hybrid popula- tion ought to produce orange, or yellow and red mottled flowers of intermediate size as are reported for all other hybrids between S. rubra and yellow flowered species. Not one plant of our more than 50 study clones produces any- thing but medium to dark oxblood red small flowers. S. rubra Walter does not occur in the region of S. alabamensis. The population of S. alabamensis is an extremely uniform one in structure and behavior: even the variability of the spring leaf structure and the variability in occurrence of the areole-like markings on the larger summer leaves are uniform features of the species. There is none of the type of variation in leaf structure or coloration so characteristic of Sarracenia populations where hybridization can be dem- onstrated. In an effort to ascertain whether this plant might repre- sent a population of direct hybrid origin between Sarracenia oreophila and S. rubra, we hand pollinated a plant of S. rubra from Fruitdale, Ala., with pollen of S. oreophila from De Soto Falls, Ala. The resulting hybrid plants, grown in our greenhouse, were intermediate between the parents in leaf structure and size and produced the orange-red flowers typical of such hybrids. The hybrids did produce laminar phyllodia, but, surprisingly, these were of an erect type as in S. leucophylla and did not resemble closely those of either S. oreophila or S. alabamensis. Leaf coloration patterns and pitcher hood conformation were unlike S. alabamensis. No recurved pitcher leaves of the S. alabamensis spring pitcher type have appeared. We could not find justification in the structure of these hybrids for suspecting a recent hybrid origin for S. alabamensis involving these two possi- ble parents. The tendency toward phyllodia, the presence of very large summer leaves resembling more in profile those of Sar- 662 Rhodora [Vol. 76 racenia flava, S. oreophila, or S. leucophylla than those of S. rubra which it resembles in flower and spring leaf, and the tendency to produce tissue areas resembling faint, ran- dom areolae, could be interpreted as placing this species in a central primitive position from which differing selective factors have led in one direction toward the S. rubra types and in another toward the phyllodium-producing groups. Its presence as a relict in an area of ancient soil derivation (Harper, 1922) in an area largely devoid of other pitcher plants suggests to us that this is a possibility. HISTORY In the earliest published reference to this plant, R. M. Harper (1918) referred the plant to Sarracenia sledgeii MacF. [S. alata (Wood) Wood], although he was certainly skeptical, for in his discussion of the distribution of S. sledgeti he says: “East of Mobile Bay, I have seen a few specimens [of true sledgeii] in the southern part of Bald- win County, Alabama, and what appears to be the same thing (though it may be a variety of the next [S. flava]) in boggy places among the long-leaf pine hills in Chilton and Autauga Counties, near the center of the State.” At that time Harper had not seen the plant in flower. In April, 1921, Harper found the flowers to be small and dark red, so he identified the plants as S. rubra Walt., although his locality was farther inland than the known distribution of that species (Harper, 1922). In 1932, Wherry, guided by Harper, visited some of these colonies but stated: “We located a colony of a red flowered pitcher plant but it was not in good enough condition to establish its identity" (Wherry, 1934) [Possibly Wherry's uncertainty was because of the presence of the peculiar, recurved spring pitchers which simulate pitchers deformed by shading]. In the September, 1973 issue of the Carnivo- rous Plant Newsletter, Dr. Wherry refers to this plant which he saw in our greenhouse in September 1955, after having seen it with Harper in the wild, as an *undescribed species from Alabama." 1974] Sarracenia — Case & Case 663 Wherry's discovery of Sarracenia oreophila at Elmore, Ala. represents the only known contact of the range of S. alabamensis with another species. At this station both species apparently grew in proximity, for both have been collected in summer leaf there and determined by various taxonomists as S. oreophila. The difficulty of determining certain summer plants of S. alabamensis from S. oreophila is emphasized by the fact that Wherry's July 13, 1935, Elmore, Ala. specimen (PENN), determined by both Wherry and McDaniel as S. oreophila, is in our opinion S. ala- bamensis. This is indicated by the small size of the flowers, dark sepals, the shape and size of the phyllodia, and in this particular specimen, smaller pitcher size than is typical of flowering-sized S. oreophila (see Fig. 3). The difficulty of separating all specimens of the two species without the presence of petals to indicate flower color, is further pointed up by McDaniel's comments (1966, pp. 45, 46) on the plants of the Elmore area: “... the Harper collection from Elmore County, Ala. differs in certain details of which the much reduced phyllodia seem to be the most significant. Addi- tional collecting both in Elmore County and in the lower Coosa River Valley is needed to determine the status of this central Alabama population." Indeed, the dissimilar spring and summer pitchers on well developed plants of Sarracenia alabamensis have led to much historical confusion and to determinations of or references to it in the literature in the past as S. sledgeii, S. flava, S. jonesii, S. rubra, and S. oreophila. Yet those who have seen vigorous living specimens agree it is distinct. ACKNOWLEDGEMENTS We wish to thank the curators of the herbaria whose specimens we have studied. We are grateful to Dr. W. H. Wagner, Jr., University of Michigan, and to Dr. James Wells, Cranbrook Institute of Science, for their valuable counsel and for securing the specimen loans for us at vari- ous times in this study. 7 Rhodora [Vol. 76 ka 1 Scan NUS E i oo SN mm a. A@NVOOW A3NQN$ 430 s *$cót T PONE c i QU J ÉXisQ (aeuiwe,) WTTITTTIS VYITSTUTIVI "wT W ‘pret zweu PRT Tt yoye woperaidəp Cog è facea (ÉmAnwey) vipudoeio wrusdwicer poc uA sigh T o iu oin Figure 3. A specimen of Sarracenia alabamensis from Elmore, Ala. showing the fully reduced laminar phyllodia occasionally pro- duced in this species which, in combination with the large summer leaves, give to this specimen a strong resemblance to S. oreophila. 1974] Sarracenia — Case & Case 665 To Dr. Rogers McVaugh, University of Michigan Herbar- ium, we are especially grateful for examining our manu- seript, for help with several difficult points, and for pre- paring the Latin description. To G. L. Burrows, IV, Mr. and Mrs. Charles F. Moore, Mr. and Mrs. Bernard Klein, and to David Case go our thanks for help in the field search for pitcher plant stations. Over the years we have received special help from the late Dr. R. M. Harper, Tuscaloosa, Ala., who furnished us with transcripts of his field notes and who guided us in 1955, and Dr. Edgar T. Wherry, Philadelphia, Pa. No words could express our gratitude to Dr. Wherry, nor acknowledge adequately all of the assistance and kindness he has extended to us. LITERATURE CITED BELL, C. R. 1949. A cytotaxonomic study of the Sarraceniaceae of North America. Jour. Elisha Mitchell Sci. Soc. 65: 137-166. pls. 8-14. Harper, R. M. 1918. The American pitcher plants. Jour. Elisha Mitchell Sci. Soc. 34: 110-125. . 1922. Some pine-barren bogs in Central Alabama. Torreya 22: 57-59. MACFARLANE, J. M. 1908. Sarraceniaceae. In: A. ENGLER, Das Pflanzenreich IV. 110(34): 1-38. McDANIEL, S. T. 1966. A taxonomic revision of Sarracenia (Sar- raceniaceae). Unpubl. Ph.D. Diss., Florida State Univ. (Avail- able from Univ. Microfilms, Ann Arbor, Mich.) Ripeway, R. 1912. Color standards and color nomenclature. A. Hoen & Co., Baltimore. Wuerry, E. T. 1929. Acidity relations of the sarracenias. Jour. Wash. Acad. Sci. 19: 379-390. . 1934. Exploring for plants in the southeastern states. Sci. Monthly 38: 80-85. 1935. Distribution of the North American pitcher plants. Im: M. V. WarcoTT, Illustrations of North American pitcher plants. Smithsonian Inst., Washington, D. C. 1973. Reminiscences on carnivorous plants. Carnivo- rous Plant Newsl. 2: 35, 36. 7275 THORNAPPLE LANE, SAGINAW, MICHIGAN 48603 666 Rhodora [Vol. 76 INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA Manuscripts should be submitted in duplicate and should be double-spaced or preferably triple-spaced (not on cor- rasable bond), and a list of legends for figures and maps provided on a separate page. Footnotes should be used sparingly, as they are usually not necessary. Do not indi- cate the style of type through the use of capitals or under- scoring, particularly in the citations of specimens, except that the names of species and genera may be underlined to indicate italics in discussions. Specimen citations should be selected critically especially for common species of broad distribution. Systematic revisions and similar papers should be prepared in the format of “The Systematics and Ecology of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particularly with reference to the indentation of keys and synonyms, Papers of a floristic nature should follow, as far as possible, the format of “Contribution to the Fungus Flora of Northeastern North America. V.," H. E. Bigelow & M. E. Barr, Rhodora 71: 177- 203. 1969. For bibliographic citations, a recommended list of standard journal abbreviations is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300. 1958. Volume 76, No. 808, including pages 497-680, was issued Feb. 28, 1975. Dodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALFRED LINN BOGLE, Editor-in-Chief ROLLA MILTON TRYON ) STEPHEN ALAN SPONGBERG Associate Editors GERALD JOSEPH GASTONY RICHARD EDWIN WEAVER J VOLUME 76 1974 The Neu Lngland Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02138 INDEX TO VOLUME 76 Entries include: authors; title words such as geographic area; and new records, systematic revisions, or ecological studies by taxa. Lists or tables of taxa and maps are also classified separately under “Lists” and “Maps”. Note that only one page number per article is given for each taxon entered. New scientific names and combinations are printed in bold face type. Adoxa moschatellina 43 Alabama, Sarracenia alabamensis, A Newly Recognized Species From Central 650-665 Algal Ecology, The Role Of Some Haloragaceae In 446-459 Alternanthera brasiliana 476 Anaphalis margaritacea 44 Andrus, Richard E., Significant New Distributional Records For The Genus Sphagnum In The Northeastern United States 511- 518 Antirrhinum orontium 467 Arkansas, Lindera melissaefolium In 525 Ascophyllum mackaii 408; nodo- sum 59-63 Asiatic Species Of Desmodium: A Review 142-144 Asparagopsis hamifera 410 Asperococcus echinatus 407 Authority For Cardamine clema- tis (Cruciferae), The Correct 53-57 Averett, John E., Typification of Chamaesaracha coronopus 311 Bahama Flora, Name Changes For The Seed Plants In The 67-138 Bahama Flora, Range Extensions And New Records For The 471- ATT Bald Cypress (Taxodium disti- chum (L.) Richard) In Suffolk County, Long Island, New York, The Occurrence Of 25-26 Baldwin, Henry I., The Flora of Mount Monadnock, New Hamp- shire 205-228 Barkley, T. M. and Arthur Cron- quist, Cacalia rugelia: A New Combination For A North American Senecionoid 48-50 Bauhinia variegata 476 Bidens beckii 44 Biosystematic Revision Of Lupi- nus lyallii 422-445 Black Spruce, Environmental Con- trol Of Needle Characteristics In Subalpine 519-524 Blanchard, Orland J., Chromosome Numbers In Kosteletzkya Presl (Malvaceae) 64-66 Bogle, A. Linn (see Lane, David M.) Bollard, Peter M. (see Hehre, Edward J.) Bryophytes of Southern Manitoba, Studies On The III. Collections From Grand Beach Provincial Park 27-38, list 29-34 Cacalia rugelia: A New Combina- tion For A North American Senecionoid 48-50 Calamagrostis lapponica 40 669 670 Caltha natans 42 Campanula rotundifolia var. velu- tina 468 Cape Cod, Massachusetts, Investi- gations Of New England Ma- rine Algae VI: Distribution Of Marine Algae Near 537-563 Capsosiphon fulvescens 401 Cardamine clematis (Cruciferae), The Correct Authority For 53- 57 Carex adusta 40; exilis, Geocaulon lividum And Other Plants Of Interest In Pittsburg, New Hampshire 307-309; rufina 41 Carya pallida 462 Case, Frederick W. and Roberta B. Case, Sarracenia alabamen- sis, A Newly Recognized Spe- cies From Central Alabama 650-666 Case, Roberta B. (see Case, Frederick W.) Catling, Paul M. and James E. Cruise, Spiranthes casei, A New Species From Northeastern North America 526-536 Central Plains States, New and Interesting Plants From The 489-490 Chabot, Brian F., Environmental Control Of Needle Character- istics In Subalpine Black Spruce 519-524 Chamaesaracha coronopus, Typi- fication Of 311 Chenopodium 462; bushianum 463; pallescens 463; strictum var. glaucophyllum 463-464 Chromosome Numbers In Koste- letzkya Presl. (Malvaceae) 64- 66 Cladophora laetevirens 403 Coleman, Douglas C. and Arthur C. Mathieson, Investigations Of Rhodora [Vol. 76 New England Marine Algae VI: Distribution of Marine Algae Near Cape Cod, Massachusetts 587-563 Color-Forms From Maine, Two New 470 Colt, L. C., Jr. and C. Barre Hell- quist, The Role Of Some Halo- ragaceae In Algal Ecology 446-459 Combination For North Ameri- can Senecionoid, Cacalia ru- gelia: A New 48-50 Conrad, Melvin L., Panicum ensi- folium Baldw. — New For Mexico 509-510 Corallorhiza wisteriana 489 Cox, B. J., A Biosystematic Revi- sion Of Lupinus lyallii 422-445 Crataegus marshallii 465 Crinum asiaticum 476 Cronquist, Arthur (see T. M. Barkley) Crotalarias Of North America (Leguminosae), A Systematic Treatment Of The Native Uni- foliate 151-204; Crotalaria bre- vipedunculata 164-166, fig. 3, map 166; bupleurifolia 194-198, map 197; var. bupleurifolia 198, fig. 11; var. robusta 198, fig. 12, map 197; mexicana 184- 186, fig. 9, map 166; nayari- tensis 198-200, fig. 13, map 197; nitens 161-164, map 159; var. nitens 164, fig, 2; var. gracilis 164, fig. 2, map 159; pilosa 157-161, fig. 1; map 159; poly- phylla 166-169, fig. 4, map 166; purshii 177-181, fig. 7, map 179; quercetorum 181-184, fig. 8, map 166; rotundifolia 186-194; X purshii 202; var. rotundi- folia 189-193, fig. 10, map 191; var. vulgaris 193-194, fig. 10, Southern 1974] map 192; sagittalis 172-177, fig. 6, map 174; XX quercetorum 202; X rotundifolia var. vul- garis 202; X stipularia 202; stipularia 169-172, fig. 5, map 159 Cruise, James E. (see Catling, Paul M.) Cucurbita pepo var. ovifera 468 Datura inoxia 476 Desmodium: A Review, The Asiatic Species Of 142-144 Digitaria sanguinalis, A New Form; f. illinoensis 57-58 Distributional Records For The Genus Sphagnum In The North- eastern United States, Signifi- cant New 511-518 Draba reptans var. micrantha 464; verna var. boerhaavii 464 Drosera From Venezuela, A New; felix 491-493, fig. 1 Dudley, T. R., The Correct Au- thority For Cardamine clematis (Cruciferae) 53-57 Ebinger, John E., A New Form Of Digitaria sanguinalis 57-58; A Systematic Study Of The Genus Kalmia (Ericaceae) 315- 398 Enteromorpha ahlneriana 401; groenlandica 402 Environmental Control Of Needle Characteristics In Sub- alpine Black Spruce 519-524 Eryngium prostratum var. pros- tratum 490 Euphrasia arctica var. dolosa 43 Evans, Dan K. (see Mohlenbrock, Robert H.) Fissidens adianthoides 32 Index to Volume 76 671 Flora Of Mount Monadnock, New Hampshire, The 205-228, list 211-221 Florida, A. New Species Of Lechea (Cistaceae) From Peninsular 478-483 Fogg, John M., Jr., Stone’s Plants Of Southern New Jersey 139- 141 Form Of Digitaria sanguinalis, A New 57-58 Fucus vesiculosus 59-63 Fuirena scirpoidea 460 Geocaulon lividum And Other Plants Of Interest In Pittsburg, New Hampshire, Carex exilis 307-309 Gillis, William T., Name Changes For The Seed Plants In The Bahama Flora 67-138 Grindelia lanceolata f. latifolia 469 Growth Of Some New England Perennial Seaweeds 59-63 Habenaria orbiculata 41 Haloragaceae In Algal Ecology, The Role Of Some 446-459 Harman, Willard N., Phenology and Physiognomy Of The Hy- drophyte Community In Otsego Lake, N.Y. 497-508 Harms, Vernon L. and John H. Hudson, Some New Or Note- worthy Vascular Plant Records From Northwestern Saskatche- wan 39-44 Harriman, Neil A., Pimpinella saxifraga L. (Umbelliferae) In Wisconsin 483 Haynes, Robert R., A Revision Of North American Potamogeton Subsection Pusilli (Potamoge- tonaceae) 564-649 672 Rhodora Hehre, Edward J. and Peter M. Bollard, The Occurrence Of Bald Cypress (Taxodium dis- tichum (L.) Richard) In Suf- folk County, Long Island, New York 25-26 Hellquist, C. Barre, A White- Flowered Form Of Utricularia purpurea In New Hampshire 19; (see Colt, L. C., Jr.) Hepper, F. N., Parasitic Witch- weed: Striga asiatica Versus S. lutea (Scrophulariaceae) 45-47 Hill, Steven R., Range Extensions And New Records For The Ba- hama Flora 471-477 Hodgdon, A. R., Carex exilis, Geocaulon lividum And Other Plants Of Interest In Pittsburg, New Hampshire 307-309 Holosteum umbellatum 489 Hudson, John H. (see Harms, Vernon L.) Hydrophyte Community In Ot- sego Lake, N.Y. Phenology And Physiognomy Of The 497- 508 Illinois Field and Herbarium Studies 460-470 Jacquemontia tamnifolia 466 Kalmia (Ericaceae), A Syste- matic Study Of The Genus 315-398; angustifolia 367-381; var. angustifolia 368-378; f. candida 377-378; var. carolini- ana 378-381; cuneata 381-383; ericoides 386-390; var. ericoides 388-389; var. aggregata 389- 390; hirsuta 383-386; latifolia 352-366; f. angustata 363; f. fuscata 363-364; f. myrtifolia 364; f. obtusata 365; f. poly- [Vol. 76 petala 365-366; microphylla 333-342; var. microphylla 334- 340; var. occidentalis 340-342; f. alba 342; polifolia 343-351; f. leucantha 351 Kitfield, Aminta, Range Exten- sion Of Woodwardia areolata 312 Kosteletzkya presl (Malvaceae), Chromosome Numbers In 64-66, table 66 Lactuca saligna f. ruppiana 490 Lagenaria siceraria 468 Laminaria digitata, 59-63; sac- charina 59-63 Lane, David M. and A. Linn Bogle, A Second Report Of The Prothallia Of Lycopodium in- undatum In North America 484-488 Lechea (Cistaceae) From Penin- sular Florida, A New Species Of; lakelae 478-483 Lindera melissaefolium In Ar- kansas 525 Lists (Tables): Bryophtes From Grand Beach Provincial Park 29-34; Chromosome Counts In Kosteletzkya 66; Chromosome Numbers In Perityle Section Perityle 236; Distribution Of Marine Algae Near Cape Cod 553-558; Flora Of Mount Mo- nadnock 211-221; Marine Algae Of Rhode Island 400-416; Name Changes In The Bahama Flora 10-124; Range Extensions And New Records For The Bahama Flora 473-477 Lithophyllum lenormandi 411 Lobelia dortmanna 43 Longleaf Pine, On The Scientific Name Of 20-24 Lophocolea heterophylla 29 1974] Lóve, Áskell, A Magnificent Swiss Flora Manual: A Review Lupinus lyallii, A. Biosystematic Revision Of 422-445, map 444; subsp. lyallii 426-435; var. lyallii 426-429; var. danaus 429-431; var. fruticulosus 431- 432; var. macroflorus 432-4535; var. roguensis 433-435; var. villosus 435-436; subsp. alcis- temporis 436-438; subsp. mi- nutifolius 438-439; subsp. sub- pandens 439-442; subsp. washo- ensis 442-443 Lycopodium inundatum In North America, A Second Report Of The Prothallia Of 484-488, fig. 3, 4, map 488 Magrath, Lawrence K. and Ronald R. Weedon, New And Interest- ing Plants From The Central Plains States 489-490 Maine, Two New Color-Forms From Southern 470 Malphigia glabra 476 Manitoba, Studies On The Bry- ophytes Of Southern III. Col- lections From Grand Beach Provincial Park 27-38 Maps: Crotalaria spp. 159, 166, 174, 179, 191, 192, 197; Lupinus lyallii 444; Lycopodium inun- datum In New England 488; Marine Algae Stations On Cape Cod 539; Perityle Section Peri- tyle spp. 251, 257, 267, 281, 285, 300; Potamogeton Section Pu- silli spp. 596, 602, 606, 611, 617, 623, 625, 630, 635, 639, 641; Sphagnum spp. In North- eastern United States 512; Spiranthes casei 534; Xanthis- ma texanum 3, 5 Index to Volume 76 613 Marine Algae Near Cape Cod, Massachusetts, Distribution Of, Investigations Of New England Marine Algae VI: 537-563, list 553-558 Marine Algae Of Rhode Island 399-421, list 400-416 Matelea decipiens 467 Mathieson, Arthur C. (see Cole- man, Douglas C.) Mentzelia decapetala 465-466 Mexico, Panicum ensifolium Balw. — New For 509-510 Mohlenbrock, Robert H. and Dan K. Evans, Mlinois Field And Herbarium Studies 460-470 Monostroma leptodermum 402 Mount Monadnock, New Hamp- shire, The Flora Of 205-228 Myriophyllum alterniflorum 43 Name Changes For The Seed Plants In The Bahama Flora 67-138, list 70-124 New And Interesting Plants From The Central Plains States 489- 490 New England Perennial Sea- weeds, The Growth Of Some 59-63 New Hampshire, A White-Flow- ered Form Of Utricularia pur- purea From 19 New Hampshire, Carex exilis, Geocaulon lividum And Other Plants Of Interest In Pittsburg 307-309 New Hampshire, New Station For Nuphar microphyllum In Durham 310 New Hampshire, The Flora Of Mount Monadnock 205-228 New Jersey: A Reprint, Stone’s Plants Of 139-141 674 New York, The Occurrence Of Bald Cypress (Taxodium dis- tichum (L.) Richard) In Suffolk County, Long Island 25-26 Newly Recognized Species From Central Alabama, Sarracenia alabamensis, A 650-665 North America, A Second Report Of The Prothallia Of Lycopo- dium inundatum In 484-488 North America (Leguminosae), A Systematic Treatment Of The Native Unifoliolate Crotalarias Of 151-204 North American Potamogeton Subsection Pusilli, A Revision Of 564-649 Northeastern North America, Spiranthes casei, A New Species From 526-536 Northeastern United States, Significant New Distributional Records For The Genus Sphag- num In The 511-518 Nuphar microphyllum In Dur- ham, New Hampshire, New Station For 310 Opuntia rubescens 476 Otsego Lake, N.Y., Phenology And Physiognomy Of The Hydrophyte Community In 497- 508 Panicum ensifolium Baldw. — New For Mexico 509-510 Penstemon cobaea 467-468 Percursaria percursa 402 Perityle Section Perityle (Com- positae-Peritylinae), Taxonomy Of 229-309, table 236; aglossa 289-291, map 285; aurea 262- 264, map 251; californica 298- 302, map 300; canescens 275- 276; ciliata 276-277, map 267; Rhodora [Vol. 76 cordifolia 291-292, map 285; coronopifolia 277-279, map 267; crassifolia 248-253; var. crassi- folia 248-250, map 251; var. robusta 250-253, map 251; cu- neata 302-305; var. cuneata 302- 305, map 300; var, marginata 304-305, map 300; emoryi 255- 262, map 257; feddemae 283- 284, map 281; hofmeisteria 279- 280, map 267; incana 264-266; jaliscana 280-282, map 281; leptoglossa 292-294, map 285; lineariloba 268-269, map 267; lloydii 294-295, map 285; lobata 295-296, map 285; microcephala 269-270, map 267; microglossa 270-275; var. microglossa 270- 273, map 267; var. saxosa 273- 275, map 267; palmeri 296-298, map 285; parryi 284-288, map 285; rosei 282-283, map 281; socorrensis 253-255; trichodonta 283, map 281; turneri 266-268, map 267; vaseyi 288-289, map 285 Phenology And Physiognomy Of The Hydrophyte Community In Otsego Lake, N.Y. 497-508 Phleum pratense f. viviparum 460 Physalis macrophysa 467 Phytogeography And Systematies Of Xanthisma texanum DC. (Asteraceae): Proper Usage Of Interspecific Categories 1-19 Picea mariana 519-524 Pimpinella saxifraga L. (Umbelli- ferae) In Wisconsin 483 Pinus palustris 20-24 Poa bulbosa 460 Polygonum ramosissimum f. alba 470 Polypremum procumbens 466 Populus X smithii 461-462 1974] Porphyra miniata 409 Potamogeton Subsection Pusilli (Potamogetonaceae), A Revi- sion Of North American 564- 649; clystocarpus 624-626, map 625; foliosus 612-624; var. foliosus 614-620, map 617; var. fibrillosus 620-624, map 623; friesii 594-598, fig. 1, map 596; groenlandicus 640-642, map 641; hillii 607-612, map 611; obtusi- folius 604-607, map 606; pusil- lus 626-640, fig. 1; var. pusillus 629-633, map 630; var. gemmi- parus 638-640, map 639; var. tenuissimus 633-637, map 635; X perfoliatus 638; strictifolius 598-603, map 602; X zosteri- formis 604 Powell, A. Michael, Taxonomy Of Perityle Section Perityle (Com- positae-Peritylinae) 229-306 Prothallia Of Lycopodium inun- datum In North America, A Second Report Of The 484-488 Pyrus pyrifolia 464-465 Quercus velutina f. missouriensis 462 Range Extension Of Woodwardia areolata 312 Range Extensions And New Rec- ords For The Bahama Flora 471-477, list 473-477 Ranunculus hyperboreus 42; tes- ticulatus 490 Records From Northwestern Sas- katchewan, Some New Or Note- worthy Vascular Plant 39-44 Reviews: Asiatic Species Of Des- modium 142-144; Stone’s Plants Of Southern New Jersey 139- 141; Swiss Flora Manual 145- 149 Index to Volume 76 675 Revision Of North American Potamogeton Subsection Pusilli (Potamogetonaceae) 564-649 Reynolds, Norman B., The Growth Of Some New England Peren- nial Seaweeds 59-63 Rhizoclonium tortuosum 404 Rhode Island, Marine Algae Of 399-421 Rhodymenia palmata 59-63 Salix X myricoides 461 Sargassum natans 409 Sarracenia alabamensis, A Newly Recognized Species From Cen- tral Alabama 650-665 Saskatchewan, Some New Or Noteworthy Vascular Plant Records From Northwestern 39-44 Schubert, Bernice G., The Asiatic Species Of Desmodium: A Re- view 142-144 Scirpus americanus 51-52; olneyi 51-52; pungens 51-52 Scleranthus annuus 489 Seed Plants In The Bahama Flora, Name Changes For The 67-138 Semple, John C., The Phytogeog- raphy And Systematics Of Xan- thisma texanum DC. (Astera- ceae) 1-19 Senecio jacobaea 469 Sherardia arvensis 468 Smith, L. B. (see Steyermark, J, A2) Sparganium chlorocarpum 39 Spergula arvensis 42; media 464 Sphagnum In The Northeastern United States, Significant New Distributional Records For The Genus 511-518, map 512; henry- ense 511-513; jensenii 516; por- toricense 513-514; pulchrum 676 515-516; riparium 515; strictum 514; subfulvum 516-517; tenel- lum 516; torreyanum 514 Spiraea latifolia f. rosea 470 Spiranthes caseii, A New Species From Northeastern North America 526-536, fig. 1, 2, map 534; gracilis 41-42 Spongomorpha lanosa 404; spines- cens 404 Steyermark, J. A. and L. B. Smith, A New Drosera From Venezuela 491-493 Stone's Plants Of Southern New Jersey: A Reprint 139-141 Striga asiatica Versus S. lutea (Scrophulariaceae), Parasitic Witchweed 45-47 Stringer, Muriel H. L. (see Stringer, Paul W.) Stringer, Paul W. and Muriel H. L. Stringer, Studies On The Bryophytes Of Southern Mani- toba III. Collections From Grand Beach Provincial Park 277-38 Suaeda depressa 464 Swiss Flora Manual, A Review, A Magnificent 145-149 Systematies Of Xanthisma texa- num DC. (Asteraceae): Proper Usage Of Intraspecific Cate- gories. The Phytogeography And 1-19 Systematic Study Of The Genus Kalmia (Ericaceae), A 315-398 Taxodium distichum 25-26 Taxonomy Of Perityle Section Perityle (Compositae-Perityli- nae) 229-306 Thelesperma gracile 469 Thlaspi perfoliatum 490 Trailliella intricata 415 Triadenum virginicum 465 Rhodora [Vol. 76 Tucker, G. E., Lindera melissae- folium In Arkansas 525 Typification And Application Of The Names Scirpus americanus Pers., S. olneyi Gray, And S. pungens Vahl 51-52 Typification Of Chamaesaracha coronopus 311 Utricularia purpurea From New Hampshire, A White-Flowered Form Of; f. alba 19 Venezuela, A New Drosera From 491-493 Villalard-Bohnsack, Martine (see Wood, R. D.) Vinca major 466 Ward, Daniel B., On The Scientific Name Of The Longleaf Pine 20-24 Weedon, Ronald R. (see Magrath, Lawrence K.) Wells, Theodore W., Two New Color-Forms From Southern Maine 470 Wicks, Johonet C., New Station For Nuphar microphyllum In Durham, New Hampshire 310 Wilbur, Robert L., A New Species Of Lechea (Cistaceae) From Peninsular Florida 478-483 Windler, Donald R., A Systematic Treatment Of The Native Uni- foliolate Crotalarias Of North America (Leguminosae) 151- 204 Wisconsin, Pimpinella saxifraga L. (Umbelliferae) In 483 Witchweed: Striga asiatica Ver- sus S. lutea (Scrophulariaceae) Parasitic 45-47 1974] Index to Volume 76 677 Wood, R. D. and Martine Villa- geography And Systematics Of lard-Bohnsack, Marine Algae 1-19, fig. 3, map 3; subsp. texa- Of Rhode Island 399-421 num 14; var. texanum 14; var. Woodwardia areolata, Range Ex- orientale 14-16; f. orientale 15; tension Of 312 f. rubrum 15-16; subsp. drum- mondii 16 Xanthisma texanum DC. (Astera- ceae): Proper Usage Of Infra- Zephyranthes citrina 476; puerto- specific Categories, The Phyto- ricensis 476 STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION Title of Publication: RHODORA Date of Filing: September 27, 1974 Frequency of Issue: Quarterly Location of Known Office of Publication: Botanical Museum, Harvard University, Oxford Street, Cambridge, Mass. 02138 Location of the Headquarters of the Publishers: Botanical Museum, Harvard University, Oxford Street, Cambridge, Mass. 02138 Publisher: New England Botanical Club, Inc., Botanical Museum, Harvard University, Cambridge, Mass. 02138 Editor: Dr. Alfred L. Bogle, Dept. Botany, University of New Hampshire, N.H. 03824 Managing Editor: Dr. Herman R. Sweet, Botanical Museum, Harvard University, Cambridge, Mass. 02138 Owner: New England Botanical Club, Inc., Botanical Museum, Oxford Street, Cambridge, Mass. 02138 No stockholders Known Bondholders, Mortgagees, and other Security Holders: None Average No. Copies each issue during preceding 12 months Total No. copies printed 850 Paid Circulation Sales through dealers, etc. none Mail subscriptions 778 Total paid circulation 778 Free distribution 12 Total distribution 790 Office use, left-over, etc. 60 Total 850 Single issue nearest to filing date 850 none 785 785 12 797 53 850 I certify that the statements made by me above are correct and complete. Herman R. Sweet, Business Manager and Treasurer. ui 175 CONTENTS: — continued Lindera Melissaefolium in Arkansas CORE MEME I ears acta a a ecg I IU 525 Spiranthes casei, a New Species from Northeastern North America Paul M. Catling and James E. Cruise .............. eee 526 Investigations of New England Marine Algae VI: Distribu- tion of Marine Algae near Cape Cod, Massachusetts Douglas C. Coleman and Arthur C. Mathieson ................ 537 A Revision of North American Potamogeton Subsection Pusilli (Potamogetonaceae) BE SEET a AEE EI A ett reno nnt entente pn e N I A 564 Sarracenia alabamensis, a Newly Recognized Species from Central Alabama Frederick W. Case, Jr., and Roberta B. Case .................. 650 Instructions for Contributors ................. eene 666