Hovdova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON Associate Editors RADCLIFFE BARNES PIKE ELIZABETH ANNE SHAW STEPHEN ALAN SPONGBERG Vol. 74 March, 1972 No. 797 CONTENTS: Revision of the Genus Melampodium (Compositae: Heliantheae). Tod F. Stues8y nouses 1 In Memoriam Charles Schweinfurth, 1890-1970. MEM rc ueniens siet tec qu eia s HERE 71 Nomenclatural Changes in Lesquerella. Reed C. Rollins and Elizabeth A. Shaw .............. ne 76 The Parker Cleaveland Herbarium of Bowdoin College. Edward Hehre, A. R. Hodgdon and R. B. Pike ............. 80 Studies on the Audouinella microscopica (Naeg.) Woelk. Complex (Rhodophyta). William J. Woelkerling ............ 85 Nomenclatural Clarification of Two Species of Verbesina (Compositae) Endemic to Florida. James R. Coleman ... 97 (Continued on Inside Back Cover) The Nem England Botanical Club, Inc. 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 double-spaced throughout. Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Albion R. Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire 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 TRbooora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUD Vol. 74 March, 1972 No. 797 REVISION OF THE GENUS MELAMPODIUM (COMPOSITAE: HELIANTHEAE)' Top F. STUESSY Probably every plant collector in Mexico at some time during his travels has come upon large fields of weedy yellow-headed Compositae. More often than not, these herbs belong to the genus Melampodium. Such abundance of num- bers, however, is not matched at all by our knowledge of the group, and in fact, the only comprehensive treatment of the genus is B. L. Robinson's synopsis published in 1901. Although this study brings together partial synonymy and provides a key for the group, all taxa are not adequately described nor does the key allow for easy and accurate iden- tification. Many of the recent specimens being collected in Mexico and Central America have been extremely difficult if not impossible to identify satisfactorily. This incomplete understanding of relationships within Melampodium sug- gested that a thorough revision should be undertaken. The present investigation is based on five years of study, including four months of field work and an examination of over 8000 herbarium specimens. The treatment presented here is essentially a straight-forward revision of Melampo- dium. Evolutionary considerations regarding the whole genus have been kept to a minimum as these evaluations have been the subject of a separate paper (Stuessy, 1971a). *Publication No. 795 from the Department of Botany, The Ohio State University, Columbus. 2 Rhodora [Vol. 74 Comments on probable phyletic relationships among se- lected species, however, are interspersed throughout the text, and data from experimental studies either in progress or already published are brought to bear on the relation- ships when appropriate. It is realized that many system- atic problems still persist within Melampodium, but it is hoped that the present study dispels the confusion that has prevailed. TAXONOMIC HISTORY The first description of Melampodium appeared in Lin- naeus’ Hortus Cliffortianus (1738) which was the refer- ence cited in the Species Plantarum of 1753. Linnaeus in his Critica Botanica of 1737 (p. 76) clearly indicated that the name he gave to the genus was derived from “Melam- pus medicus graecus." Apparently overlooking this ex- planation, many later workers (e.g., Gray, 1884; Cockerell, 1905) erroneously have believed the name to come from the Greek words meaning ''black-foot." From the time of Linnaeus' original description of M. americanum in 1738, the number of the species described in Melampodium has increased to over 88. Difficulties of specific cireumscription and synonymy, resulting from the many described taxa, have been surpassed by the more perplexing problems dealing with generic and sectional de- limitation. Because discussions have already been pub- lished on the history of generic problems associated with Melampodium (Stuessy, 1969b, 1970a), the following com- ments will emphasize the development of attempts to sub- divide the genus. Humboldt, Bonpland, and Kunth (1820) established the first subdivisions of the genus. Based primarily on the genera that were included in synonymy, they designated three groups (here recognized as subgenera) : Melampodia, Dysodia (Rich. in Pers.) H.B.K., and Alciniae (Cav.) H.B.K. No descriptions or comments were added to explain the rationale for the groupings beyond the inclusion of species. 1972] Melampodium — Stuessy 3 Morphological criteria for the sections later recognized by Candolle (1836) were first stated by Cassini (1829). Cassini recognized the separate genera, Melampodium, Al- cina, and a new genus, Zarabellia, all of which had inner phyllaries tightly enclosing and fused with a single ray achene, but each of which possessed distinctive characters as follows: Melampodium L. (1) Inner phyllaries each crowned with a large hooded appendage (2) Sterile disc ovaries very large. Zarabellia Cass. (1) Inner phyllaries with no hood but with two small valve-like processes (2) sterile disc ovaries nearly or completely absent. Alcina Cav. (1) Inner phyllaries with no hood or valve- like processes and smooth at the sides (2) Sterile disc ovaries nearly or completely absent. Candolle (1836) put Zarabellia and Alcina into Melam- podium but maintained the taxa as three sections based on Cassini’s bract differences: Humelampodium DC., Zara- bellia (Cass.) DC., and Alcina (Cav.) DC. Later, Bentham and Hooker (1873) and Hoffmann (1890) did not recognize sections in Melampodium, perhaps to stress the morpholog- ical unity of the genus (Turner and King, 1962). Baker, treating Brazilian species (1884), suggested two subgen- era: Dysodium (Rich. in Pers.) H.B.K., and Unzia (L.f.) Baker, which were characterized by pedunculate and sub- sessile heads, respectively. Baillon (1882) departed signif- icantly from previous authors by lumping Acanthospermum Schrank and Lecocarpus Dene. into Melampodium, and these included genera formed the basis of his two recog- nized sections: Acanthospermum (Schrank) Baillon and Lecocarpus (Dene.) Baillon. Robinson in his synopsis of Melampodium (1901) sub- merged section Alcina (containing the single species, M. perfoliatum) into section Zarabellia. The two other sec- tions of Candolle were retained on the basis of the hood versus non-hood bract character. Robinson acknowledged the variability of this feature but wrote that (p. 455) 4 Rhodora [Vol. 74 * ., the presence or absence of a hood can usually be determined readily, and the two sections Eumelampodium and Zarabellia may conveniently be retained.” Using cytological data, Turner and King (1962) re- opened considerations of sectional subdivision. Based on chromosome counts from 89 populations of 21 species’, the taxa of section Melampodium (Humelampodium DC.) were shown to be on a base of « — 10, and the species within section Zarabellia to be multibasic with x = 9, 11, 12, 16° and 23+. In addition to bract and chromosome base num- ber criteria, “small plants" and “densely pubescent or to- mentose (rarely merely hispid) foliage" were suggested as characterizing section Melampodium, with “heterogeneous” morphology and “hispid pubescence” more indicative of section Zarabellia. No one has ever proposed series within the two recog- nized sections of Melampodium. In fact, Robinson experi- enced frustration in attempting to create such order (1901, p. 455): “Much difficulty has been experienced in giving the species a natural sequence, and after many efforts the hope of securing such an arrangement has been abandoned.” Turner and King (1962) suggested that within the more morphologically heterogeneous section Zarabellia some for- mal subdivisions might be desirable after further study. They mentioned several informal groups such as: (1) “the small-headed, annual species having a chromosome base of x = 9"; (2) “those rhizomatous species centering about M. montanum with chromosome numbers on a base of x — 11"; and (3) "those annual species with large involucral bracts and smooth achenes having chromosome numbers of n = 11 and 12." *See Stuessy (1971b) for a reassignment of these chromosome counts to the presently recognized taxa. ‘Due to the generic transfer of M. camphoratum (n = 16) back into Umzia (Stuessy, 1969b), this number no longer belongs in Melampodium. ‘Because of recent counts of n — 12 for M. dicoelocarpum (Stuessy, 1971b), the n — 23 count for this species seems best interpreted as an aneuploid at the tetraploid level on a base of x — 12. 1972] Melampodium — Stuessy D Figs. 1-4. Disc florets of representative species of Melampodium showing the two different types of abortive ovaries. Figs. 1 & 2, M. leucanthum, Stuessy 752 (OS); Figs. 3 & 4, M. divaricatum, Stuessy 547 (OS). Drawn from fresh material. The corollas and stamens in Figs. 2 and 4 have been removed, but they are the same florets as in Figs. 1 and 3, respectively. The arrow in Fig. 1 points to a marked annular constriction at the point of ovary and corolla tube juncture, which is characteristic of species in sect. Melampodium. That same marked constriction is absent from the florets of species in the other five sections of the genus (Fig. 3). Both types of ovaries are capped by ovarian discs (Figs. 2 & 4). In the present treatment I depart significantly from pre- vious workers by recognizing six sections in the genus. Based on coherence of morphology and chromosome num- bers, section Melampodium seems a phyletic unit and there- fore is retained, although here for the first time it is di- vided into five series. Section Zarabellia s.l. of previous authors, however, has been circumscribed in the past by 6 Rhodora [Vol. 74 the absence of hoods of the inner phyllaries (to be dis- cussed in detail later) and by the possession of chromo- some numbers other than on a base of x = 10 (i.e, xz = 9, 11, and 12). The only positive feature shared by all taxa of the section is the rudimentary ovary of the disc florets which contrasts markedly with the more conspicuous ovary of those taxa in section Melampodium (Figs. 1-4; these ovary differences mentioned earlier by Cassini, 1829, as having generic significance). The diversity of other mor- phological features and chromosome numbers in taxa of section Zarabellia s.l., however, suggests that the one uni- fying ovary feature may have been derived by parallel evolution (Stuessy, 1971a). From this phyletic viewpoint section Zarabellia s.l. seems an artificial unit and is there- fore divided into five separate sections based on chromo- somal and morphological differences. Two of the sections, Zarabellia, s.str. and Rhizomaria, correspond respectively to the first two informal groups mentioned by Turner and King (1962). A list of the recognized sections, series, and included taxa follows: Synopsis of Classification of Melampodium L. I. Section Melampodium 1. Series Melampodium 1. M. americanum L. 2. M. diffusum Cass. 3. M. pilosum Stuessy 4. M. longipes (A. Gray) Robins. 5. M. linearilobum DC. 2. Series Leucantha Stuessy 6. M. leucanthum Torr. & A. Gray 7. M. cine- reum DC. Ta. M. cinereum DC. var. cinereum Tb. M. cinereum DC. var. hirtellum Stuessy Tc. M. cinereum DC. var. ramosissimum (DC.) A. Gray 8. M. argophyllum (A. Gray ex Robins.) Blake 3. Series Sericea Stuessy 9. M. sericeum Lag. 10. M. pringlei Robins. 11. M. strigosum Stuessy 12. M. longicorne A. Gray 13. M. nayaritense Stuessy 1972] Melampodium — Stuessy 7 4. Series Cupulata Stuessy 14. M. cupulatum A. Gray 15. M. appendiculatum Robins. 16. M. sinuatum Brandg. 17. M. rosei Robins. 18. M. tenellum Hook. & Arn. 19. M. glabribracteatum Stuessy 5. Series Longipila Stuessy 20. M. longipilum Robins. II. Section Zarabellia (Cass.) DC. 21. M. longifolium Cerv. ex Cav. 22. M. mimuli- folium Robins. 23. M. gracile Less. 24. M. microcephalum Less. 25. M. paniculatum Gardn. III. Section Serratura Stuessy 26. M. divaricatum (Rich. in Pers.) DC. 27. M. costaricense Stuessy 28. M. dicoelocarpum Robins. 29. M. tepicense Robins. 30. M. sina- loense Stuessy IV. Section Bibractiaria Stuessy 31. M. bibracteatum S. Wats. 32. M. repens Sessé & Moc. V. Section Rhizomaria Stuessy 33. M. montanum Benth. 33a. M. montanum Benth. var. montanum 33b. M. montanum Benth. var. viridulum Stuessy 34. M. aureum Brandg. VI. Section Alcina (Cav.) DC. 35. M. perfoliatum (Cav.) H.B.K. 36. M. glabrum S. Wats. 37. M. nutans Stuessy DISTRIBUTIONAL SUMMARY Melampodium is a tropical and subtropical genus mostly restricted to Mexico and Central America (Fig. 5) with four species located in the southwestern United States, and three species scattered in Colombia and Brazil. Although the Brazilian distributions could indicate South American origins for the two species, M. paniculatum and M. divari- catum, the fact that these species are found abundantly also in Central America and Mexico suggests that their presence in Brazil is due to recent introductions by man. 8 Rhodora [Vol. 74 Fig. 5. Map of North and South America showing generalized distribution of Melampodium. Collections from Burma and the Philip- pine Islands not shown. The exceedingly weedy nature of M. divaricatum and M. perfoliatum has allowed successful introductions of the genus to Burma, Cuba, Puerto Rico and the Virgin Islands. Another Mexican species, M. diffuswm, has become estab- lished near Manila in the Philippine Islands. 1972] Melampodium — Stuessy 9 Melampodium occupies primarily moist habitats from mountain pine-oak to tropical deciduous forests. Only two species, M. leucanthum and M. cinereum, are clearly xero- morphic, found in the drier regions of northern Mexico and adjacent United States. SPECIFIC AND VARIETAL CATEGORIES It would be a formidable task to treat 37 species of a genus exhaustively with many experimental approaches to discover all the effective isolating mechanisms. As a re- sult, in the present study a morphological species concept has been stressed. With the exception of chromosome num- bers for many taxa, only in a few cases have data from ex- perimental work been used to define more accurately the reproductive limits of each taxon. It is assumed, however, that the qualitative and quantitative morphological discon- tinuities formally recognized in this study do represent genetic differences that in some fashion are responsible for maintaining the integrity of each specific and varietal unit. An attempt has been made to treat specific and varietal categories consistently with regard to the data available in each case. Varieties are regarded as morpho-geograph- ical subdivisions of a species (Kapadia, 1964) that pre- sumably reflect genetic differences. MORPHOLOGY AND TAXONOMIC CRITERIA As an aid to understanding the specific and varietal cate- gories used in the present study, the taxonomic value of various morphological features is discussed below: Habit. — Both herbaceous annuals and suffruticose or rhizomatous perennials are found in the genus. The peren- nial habit is the less common condition found only in the following seven species of sections Melampodium and Rhi- zomaria: M. americanum, M. leucanthum, M. cinereum, M. argophyllum, M. sinuatum, M. montanum, and M. aureum. Roots. — All taxa of the genus are tap-rooted except for the two rhizomatous perennial species of section Rhizo- maria: M. montanum and M. aureum. 10 Rhodora [Vol. 74 Leaves. — The shape, size, vesture, and type of margin of leaves are often useful as discriminating taxonomic characters at various levels in the hierarchy from section to variety. However, in some taxa leaf shape and size can be quite variable, especially in the widespread species, and this plasticity has been the cause of recognition of forms as species or varieties by previous workers (e.g., in M. divaricatum). Vesture. — The surface of various organs ranges from glabrous to copiously sericeous, and the vesture often can be used as a reliable taxonomic character. Heads. — Many features of the outer and inner involu- cral bracts and the ray florets are of diagnostic value. Within limits, the number of disc and ray florets is some- times useful as a specific indicator, and as mentioned pre- viously, the shape of the abortive ovaries of the disc florets is useful as a criterion for delimiting section Melampodium. “Fruit” in this treatment refers to the mature ray achene and enveloping inner involucral bract (Robinson, 1901). The fruits are often capped by extensions of the inner in- velucral bracts of varying shapes and sizes upward from the achene apexes (Figs. 6-9). The presence of one partic- ular type of extension called a “hood” (the term apparently first used by Cassini, 1829, and followed by most subsequent authors) correlates well with other features of section Melampodium. Within most species, however, the hood size, shape, and appendage vary considerably. In fact, the amount of variation prevalent among fruits in different plants of a single population (Fig. 10) and sometimes even among bracts in a single head (cf. Fig. 9E in Stuessy, 1970b) is astonishing. Such permutations, especially in the sculpturing of the lateral outer surfaces of the inner phyl- laries, have been judged by some students of the group (e.g., Robinson, 1901) as indicative of varieties or even as distinct species. However, with few exceptions (such as in M. divaricatum and M. dicoelocarpum) the lateral bract surface features usually are not reliable as taxonomic char- acters. 1972] Melampodium — Stuessy Bi Figs. 6-9. Representative fruits of Melampodium illustrating variation in achenal apexes. Fig. 6, M. glabrum, apex moderately sculptured, Stuessy 709 (OS); Fig. 7, M. longifolium, apex with abaxial protuberance, Sprengel s.n. (P); Fig. 8, M. longipilum, apex with adaxial awn, Stuessy 634 (OS); Fig. 9, M. leucanthum, apex with hood, Stuessy 752 (OS). Fig. 10. Representative fruits of Melampodium linearilobum from a single population in Michoacán, Mexico (Stuessy 698, OS) illus- trating variations in structure of the inner involucral bracts. 12 Rhodora [Vol. 74 TAXONOMIC TREATMENT MELAMPODIUM L. Melampodium L. [Hort. Cliff. 425. 1738.] Sp. Pl. 921. 1753. Gen. Pl. ed. 5. 392. 1754. Type species: Melampo- dium americanum L. Cargilla Adans. Fam. 2:130. 1763. nom. superfl. in- cluding Chrysogonum L. and Melampodium L. Alcina Cav. Ic. 1:10. t. 75. 1791. Type species: Alcina perfoliata Cav. = Melampodium perfoliatum (Cav.) H.B.K. Dysodium Rich. in Pers. Syn. 2:489. 1807. non Dys- sodia Cav. 1802. Type species: Dysodium divaricatum Rich. in Pers. = Melampodium divaricatum (Rich. in Pers.) DC. Melampodium L. subg. Alcina (Cav.) H.B.K. [as subg. Alciniae]. Nov. Gen. Sp. 4:274. 1820. Melampodium L. subg. Dysodium (Rich. in Pers.) H.B.K. [as subg. Dysodia]. Nov. Gen. Sp. 4:273. 1820. Melampodium L. subg. Melampodium [as subg. Melam- podia]. H.B.K. Nov. Gen. Sp. 4:272. 1820. Camutia Bonat. ex Steud. Nom. Bot. 146. 1821. pro syn., nom. nud. Based on Camutia perfoliata Bonat. ex Steud. Carnutia [attributed to Bonat. ex Steud. by] Baker in Martius, Fl. Bras. 6(3) :159. 1884. Orthogr. var. Zarabellia Cass. Dict. Se. Nat. 59:240. 1829. [non Neck. 1790. invalid fide Art. 20, Intern. Code Bot. Nomen.] Type species: Zarabellia rhomboidea Cass. = Melampo- dium longifolium Cerv. ex Cav. Annual herbs to perennial subshrubs. Tap roots in all but 2 species (M. aureum and M. montanum with fibrous roots and rhizomes). Stems decumbent to erect, terete to finely striate, dichotomously branched with peduncles aris- ing from the middle and upper dichotomies. Peduncles very short and stout to long and filiform. Leaves opposite, de- cussate, linear to ovate-rhombic, at apex acuminate to ob- tuse, at base attenuate to auriculate-connate, with upper surfaces glabrous to pilose, with lower surfaces glabrous to sericeous, at margin entire to toothed. Heads solitary. Receptacle paleaceous, convex, sometimes elevated on a short cylindrical stalk 2-3 mm above the outer involucre. 1972] Melampodium — Stuessy 13 Involucre biseriate. Outer involucre spreading to cupulate; bracts 2-5, more or less equal, with margins entire, separate to connate more than 2/3 their length, with adaxial surface glabrous. Innermost phyllaries each enclosing a single ray achene, completely covering or sometimes partially open at apex, often extending upward into a hood or other apical appendage (Figs. 6-9). Ray florets 3-13; ligules varying shades of yellow or cream-white (2 species), at the apex bi- or tri-dentate, attached on adaxial side of achene apex; tube nonexistent or very short; style branches filiform, flattened, obtuse at the apex, with stigmatic surfaces marginal and enlarged and running the length of the branches, recurved when fruit maturing; achenes asymmetrically obovoid and laterally compressed. Disc florets 3-110; corollas yellow- green to yellow-orange, regular, 5-lobed (rarely 4-lobed), with lobes partially reflexed or extended at right angles, throat salverform-funnelform; anthers brown, exserted from corolla 1/3-1/2 their length; style linear-capillaceous, unbranched, exserted from within anthers; ovary sterile, capped by a disc; pappus absent. Paleae scarious, con- duplicate around disc corollas. Chromosome numbers, n = 9, 10, 11, 12, 18, 20, 23, 25 + 1, 27, 30, and 33. ARTIFICIAL KEY TO TAXA a. Ligules white. (b) b. Outer phyllaries connate more than half their length. (c) c. Stems and leaves strigillose; leaves with margins usually entire-sinuate (less often pinnately lobed) ; heads 20-37 mm diam‘; ligules 7-13 mm long, 2.5-8 mm wide; plants 15-50 em tall MEE cs qx onda se sees TATEM 6. M. leucanthum c. Stems and leaves canescent-tomentose; leaves with margins usually pinnately lobed-parted (less often entire) ; heads 10-23 mm diam; lig- ules 2.5-9 mm long, 1.5-4.5 mm wide; plants 12- 22 em tall ................ 8. M. argophyllum 'In the keys and descriptions head diameter excludes the outer involueral bracts. 14 Rhodora [Vol. 74 b. Outer phyllaries connate a third or less their length. (d) d. Leaves with basal margins hispid with hairs 0.6- d. 1.5 mm long; stems strigose-hispid with hairs 0.2-1.5 mm long . 7b. M. cinereum var. hirtellum Leaves with basal margins strigose with hairs 0.3-0.6 mm long; stems strigose with hairs 0.1- 0.8 mm long. (e) e. Leaves linear, 7-32 mm long, 2-5 mm broad; heads 3-5 mm tall; outer involucre 4.5-9 mm diam; outer phyllaries 2.7-4 mm long, 1.5- 2.8 mm wide; hood 0.8-1.5 mm tall ........ ZEN 7c. M. cinereum var. ramosissimum e. Leaves linear-oblong, 12-55 mm long, 1-14 mm broad; heads 5-8 mm tall; outer involucre 7-14.38 mm diam; outer phyllaries 3.5-7.5 mm long, 2.3-5 mm wide; hood 1.3-2.9 mm tall .. es Ta. M. cinereum var. cinereum a. Ligules yellow. (f) f. Outer phyllaries 2. (g) g. g. Leaves oblong to obovate-oblanceolate; ray flor- ets 5-6 (rarely 3); stems erect to decumbent; plants subaquatic ........ 31. M. bibracteatum Leaves obovate; ray florets 2-3; stems prostrate ; plants terrestrial .............. 32. M. repens f. Outer phyllaries 3-5. (h) h. Outer phyllaries 3. (i) i. Heads 3-5.5 mm diam; ligules usually less than 2 mm long; peduncles copiously stipi- tate-glandular ........ 25. M. paniculatum i. Heads 5-15 mm diam; ligules usually more than 2 mm long; peduncles strigillose to his- pid to weakly stipitate-glandular. (j) j. Leaves subauriculate at base .......... ee ee ee 23. M. gracile j. Leaves attenuate to obtuse at base. (k) k. Leaves with tapering petioles 1-3 cm long; inner phyllaries open in a small 1972] Melampodium — Stuessy 15 hood at achene apex, often extended Into-awn ... 0 . 9T. M. nutans k. Leaves sessile or with petioles 0.1-0.4 cm long; inner phyllaries completely closed over achene apex, with no ex- tended appenddgsg 8541 eas Po ote alg s E 24. M. microcephalum h. Outer phyllaries 5. (1) l. Leaves sericeous beneath. (m) m. Outer phyllaries with the margin scari- OUS ................. 5. M. linearilobum m. Outer phyllaries with the margin herba- ceous. (n) n. Leaves narrowly ovate to lanceo- late. (0) o. Plants perennial; heads 11-17 mm diam; ligules 4-6 mm long ...... i las ate RE l. M. americanum o. Plants annual; heads 6-11 mm diam; ligules 2.5-4.5 mm long. (p) p. Plants 8-28 cm tall; stems pil- ose; outer phyllaries ovate to narrowly ovate; leaves copious- ly sericeous beneath ......... PNIS 3. M. pilosum p. Plants 25-40 cm tall; stems stri- gillose-hispidulous; outer phyl- laries ovate; leaves moderately sericeous beneath ..... ..... DENN ee 2. M. diffusum n. Leaves linear, oblanceolate, or deeply pinnately divided. (q) q. Ligules 2 mm long or less. (r) r. Leaves linear and entire to deeply pinnately divided ; ligules ca 2 mm long; heads 8-11 mm diam; disc florets 15-25 ...... E esuna 13. M. nayaritense 16 Rhodora [Vol. 74 r. Leaves linear-oblong to oblance- olate, often lobed or parted; ligules ca 1 mm long; heads 4- 8 mm diam; disc florets 2- 12. (s) s. Peduncles 5-22 mm long; dise florets 5-12; ligules on undersurface yellow at apex; outer involucre 6-11 mm diam .. 9. M. sericeum s. Peduncles 1-2 mm long; disc florets 2-3; ligules on under- surface purple at apex; outer involucre 3-4 mm diam ENDE 10. M. pringlet q. Ligules more than 2 mm long. (t) t. Plants annual; stems yellow to green ......... 4. M. longipes t. Plants perennial; stems purple to green .... 1. M. americanum l. Leaves glabrous or strigose beneath. (u) u. Outer phyllaries with the margin narrow- ly scarious. (v) V. Peduncles 1.3-1.7 cm long; outer phyl- laries abaxially glabrous ..... DEM 19. M. glabribracteatum Peduncles usually more than 2 cm long; outer phyllaries abaxially sub- glabrous, strigose or pilose. (w) w. Plants perennial with fibrous roots and rhizomes. (x) x. Heads 19-38 mm diam; ligules yellow-orange to yellow, 5-12 mm long; disc corollas yellow; fruit 2-2.3 mm long*; chromo- *Fruit length in the key and descriptions excludes hoods and ap- pendages. 1972] Melampodium — Stuessy 17 yellow, 4-7.5 mm long; disc corollas yellow-green; fruit 1.6- 1.8 mm long; chromosome num- ber, n = 11. (y) y. Ligules on undersurface light green at apex and on veins .. 33b. M. montanum var. viridulum y. Ligules on undersurface dark purple at apex and on veins .. 33a. M. montanum var. montanum w. Plants annual or perennial with tap roots. (z) Z. Plants perennial; leaves on un- dersurface copiously tomentose, at margin markedly sinuate .. DER op oe Pune 16. M. sinuatum Plants annual; leaves on under- surface strigose, at margin en- tire, lobed or divided. (aa) aa.Leaves at base attenuate. (bb) bb. Outer phyllaries ovate to narrowly ovate, 4-6 mm long; heads 7-8 mm tall .. ........ 14. M. cupulatum bb. Outer phyllaries ovate to orbieulate, 2.5-3.5 mm long; heads 3.5-5. mm tall .... 18. M. tenellum aa. Leaves at base obtuse to sub- auriculate. (cc) cc. Outer phyllaries connate 2/3 their length ........ 18 Rhodora [Vol. 74 .. 15. M. appendiculatum cc. Outer phyllaries connate less than 1/4 their length ZEE 17. M. rose u. Outer phyllaries with the margin herba- ceous. (dd) dd. Outer involucre 15-32 mm diam ; outer phyllaries 6-20 mm long ; fruits 4-7 mm long; lower leaves conspicuously per- foliate at base .... 35. M. perfoliatum dd. Outer involucre 3-10 (—-14) mm diam; outer phyllaries 2-6(—6.8) mm long; fruits 1.6-3 (—4) mm long; lower leaves attenuate to subauriculate at base. (ee) ee. Leaves markedly petiolate. (ff) ff. Ligules 3.5-7 mm long. (gg) gg. Fruit at apex with a flattened adaxial appendage extending upward into a cirrhous awn (Fig. 8) .... 20. M. longipilum gg. Fruit at apex variously ridged and sculptured but not as above EE 26. M. divaricatum ff. Ligules less than 3 mm long. (hh) hh. Fruit on lateral surfaces with 2 deep oval cavities .......... . 28. M. dicoelocarpum hh. Fruit on lateral surfaces vari- ously ribbed, tubercled, and sculptured but not as above. (ii) ii. Heads 6-10 mm diam; stems usually 2-4 mm diam ...... ZEN 27. M. costaricense ii. Heads 3-5 mm diam; stems usually less than 2 mm diam. (jj) jj. Peduncles 0.8-8 mm long; 1972] Melampodium — Stuessy 19 disc florets 3-7(-12) ..... hee 29. M. tepicense jj. Peduncles 10-29 mm long; disc florets ca 14 ........ NS 30. M. sinaloense ee. Leaves sessile or subsessile. (kk) kk. Plants subaquatic; stems thick and yellowish .... 36. M. glabrum kk. Plants terrestrial; stems greenish to greenish-purple. (ll) ll. Ligules less than 2 mm long; outer phyllaries lanceolate. (mm) mm. Fruits with an abaxial pro- tuberance (Fig. 7) ...... `... 21. M. longifolium mm. Fruits with hoods (Fig. 9) or minutely sculptured at apex. (nn) nn. Plants usually 15-20 cm tall; stems ascending; peduncles usually 0-3 mm long; leaves at margin entire-pinnately lobed ; heads 3-4 mm tall, 4-8 mm diam; outer phyl- laries lanceolate-elliptic . e i 11. M. strigosum nn. Plants usually 20-40 cm tall; stems erect; pedun- cles usually 10-20 mm long; leaves at margin en- tire to dentate; heads 4-7 mm tall, 7-13 mm diam; outer phyllaries elliptic to obovate-rhombic ....... T" 12. M. longicorne ll Ligules more than 2 mm long; 20 Rhodora [Vol. 74 outer phyllaries ovate or nar- rowly ovate. (00) oo. Leaves attenuate at base .... DENEN 24. M. microcephalum oo. Leaves subauriculate to ob- tuse at base. (pp) pp. Stems copiously pilose near apex; outer phyllaries ovate-lanceolate, 5-6 mm long, at apex acute ...... 22. M. mimulifoliwm pp. Stems moderately hispid- pilose and weakly stipitate- glandular near apex; outer phyllaries ovate, 3.5-5 mm long, at apex acuminate INE 23. M. gracile I. MELAMPODIUM section Melampodium Melampodium L. sect. Eumelampodium DC. Prodr. 5:518. 1836. Type species: Melampodium americanum L. Tap-rooted annuals or perennials; leaves linear to ovate, sessile to markedly petiolate, with margins entire or lobed or obscurely serrate; outer involucre cupulate, with bracts 5, at margins herbaceous or scarious; ovaries of the disc florets linear, 1.2-2.3 mm long; tube of disc florets with a marked basal annular constriction at point of ovary junc- ture (Fig. 1, arrow) ; fruits usually capped (Fig. 9) with a hood (absent in M. pringlei, and M. sinwatum; sometimes absent in M. americanum, M. hispidum, M. longicorne, and M. sericeum; rudimentary in M. cupulatum) or with a flat- tended adaxial awn (M. longipilum, Fig. 8) ; chromosome base number, x — 10. Species 1-20. 1. Series MELAMPODIUM Annual herbs or suffruticose perennials (in M. ameri- canum) ; leaves narrowly ovate to linear, often pinnately divided, sericeous beneath, ligules orange-yellow, more than 1972] Melampodium — Stuessy 21 or rarely about 2 mm long; margins of outer phyllaries her- baceous or scarious (in M. linearilobwm) ; fruits usually hooded. Species 1-5. Type species: Melampodium ameri- canum L. 1. Melampodium americanum L. [Hort. Cliff. 425. 1738.] Sp. Pl. 921. 1753. TYPE: MEXICO: Veracruz, near Vera- cruz, Mar 1731, W. Houstoun s.n. (Holotype, BM; photo- graph of holotype, Ny! US!; photograph of BM isotype, US!). Calendula decumbens Mill. Gard. Dict. ed. 8. n. 9. 1768. nom. superfl., based on type of Melampodium americanum. Melampolium heterophyllum Lag. Gen. et Sp. Nov. 33. 1816. TYPE: “NOVA HISPANIA": grown in Roy. Bot. Gard. Madrid, source and collector of seeds uncertain (Holotype, MA?; tracing of DC isotype [?], GH!). Melampodium sericeum H.B.K. Nov. Gen. Sp. 4: 272, t. 398. 1820. non Lag. 1816. nom. illegit. TYPE: MEXICO: Guerrero, between Tehuilotepec and Taxco, 5520 ft, Apr 1803, A. J. Bonpland 3968 (Holotype, P; isotype, P!; frag- ment of holotype, P! US!; photograph of P isotype, OS! TEX!; photograph of probable B isotype, TEX!). Melampodium angustifolium DC. Prodr. 5:519. 1836. TYPE: PERU [?]: “in Peruviae montanis, ad Oronocum," 1790-91, T. Haenke s.n. (Holotype, PR; isotypes, P [2]!; probable isotype, F!; photograph of G-DC isotype, F! NY! us! Ipc! 800. 927: IIL1!; photograph of P isotype, os! TEX!). If the collection actually is from Peru, then the range of M. americanum would be extended much further south than its present southern limit in Guatemala. How- ever, apparent mixing of label data of Haenke's specimens has been reported in at least three instances by other "The method of citing specimens on Interdocumentation (IDC) microfiches described by Hepper (1968) is used here throughout the text. Taking the above citation as an example, the first number (800) refers to the specific herbarium in the IDC collection (here G-DC) followed by the numbers of: the individual microfiche (927), the particular line on the card (III), and the specific frame (—speci- men) on the line (1). 22 Rhodora [Vol. 74 workers (Hitchcock, 1909; Merrill, 1925; Tryon, 1955). Because Haenke also collected in Mexico in the region from Acapulco to Mexico City (Safford, 1905; Alston, 1934) where M. americanum is common, it seems likely that mix- ing of label data also has occurred here. Melampodium kunthianum DC. Prodr. 5:519. 1836. nom. nov. Based on type of M. sericeum H.B.K. Meiampodium nelsonii Greenm. Proc. Amer. Acad. Arts & Sci. 41:260. 1905. TYPE: MEXICO: Michoacán, Volcano of Jorullo, 28 Mar 1903, E. W. Nelson 6939 (Lectotype chosen, GH!; isotypes, NY! US!; photograph of US isotype, TEX !). Perennial subshrubs, 15-60 cm tall. Stems ascending, 1-2 mm diam, strigose to hispid-pilose with hairs 0.2-1 mm long. Peduncles 2.5-7.5 em long. Leaves sessile, linear to ovate, 9-6 em long, 0.2-2.7 cm wide, at apex acute-obtuse, at base attenuate to obtuse, with upper surface strigose with hairs 0.2-0.5 mm long, with lower surface sericeous; margin entire or lobed to pinnately divided into linear segments, at base usually pilose-hirsute with hairs up to 1 mm long. Heads 7-8 mm tall, 11-17 mm diam. Outer involucre cupu- late, 7-10 mm diam; bracts 5, slightly connate at base, im- bricate 2/3 their length (rarely separate), ovate to rhombic, 5-7 mm long, 3-4 mm wide, at apex acuminate, with abaxial surface strigose with hairs 0.5 mm long near apex, near base pilose to sericeous, at margin herbaceous. Fruits 2-3 mm long, with lateral surfaces smooth with ribs and few tubercles to verrucate-aculeate; hood apex mucronate (rare- ly muticous) to cirrhous, with tapering appendage up to 2 mm long. Ray florets 8-13; ligules yellow-orange, oblong- elliptic, 4-6 mm long, 1.5-3 mm wide. Disc florets 50-100; corollas yellow-orange, 2.1 mm diam, with throat and tube each 1 mm long. Paleae oblanceolate, 4 mm long, 1 mm wide; apex yellow, with margin entire to undulate becom- ing dentate-erose laterally; midrib prominent, pubescent with hairs 0.4 mm long. Chromosome number, n = 10. 1972] Melampodium — Stuessy 23 Fig. 11. Map of Mexico and adjacent Guatemala showing distri- bution of Melampodium americanum (dots), M. glabribracteatum (square), and M. pringlei (triangle). Three collections of M. americanum near Tepic, Nayarit, not shown. Habitats ranging from savannas to pine-oak forests in Mexico on the eastern slope of the Sierra Madre Oriental and on the western slopes of the Sierra Madres Occidental and Sur, with extensions into Guatemala (Fig. 11), 210- 2330 m. Flowering dates, Jan-Aug. This species is very widespread, ranging from north in Tamaulipas, Mexico, to south into Guatemala and is found in diverse habitats from mountain pine forests to lowland savannas. As might be expected from such geographical 24 Rhodora [Vol. 74 and ecological diversity, much morphological variability also is present. Although no formal varieties are proposed at this time, morpho-geographic trends do exist that need further study: (1) plants with short leaves with few and broad lobes and large heads found in the coastal lowlands near Alvarado, Veracruz; (2) plants with long leaves, pin- nately divided into narrow lobes, found in the mountains of Chiapas and Guatemala; (3) short plants with small heads and a tendency toward the annual habit found in the mountains of the western Sierra Madre in Jalisco, Micho- acan and Guerrero. Three collections (Feddema 832; King 3699; Rose, Stand- ley and Russell 14307) found in the northwestern range of this species (north of Tepic, Nayarit) also may deserve formal taxonomic status. The outer involucral bracts of these collections are more rhombic with narrower atten- uate apexes as compared with the more typical ovate bracts of the rest of the species, and the ligules are longer and narrower than usual. A collecticn from Guerrero, how- ever, Hinton 9134, approaches these Nayarit collections and seems to vitiate clear varietal or specific recognition. Ob- viously additional work, especially in the Tepic area, is needed. 9 REPRESENTATIVE SPECIMENS: Guatemala. BAJA VERAPAZ: ca 3 mi S of Salamá, 10 Jul 1960, King 3260 (DS, NY, TEX, UC, Us); ca 14 mi W of Salama, 14 Jul 1960, King 3358 (DS, NY, TEX, UC, US); ca 15 mi S of Rabinal, 14 Jul 1960, King 3364 (DS, NY, TEX, UC, US). EL PROGRESO: 35 mi NE of Guatemala, 30 Jul 1966, Stuessy 602 (TEX). HUEHUETENANGO: Uaxackanal, 1 Aug 1896, Volkem 2973 (F [photograph of US specimen], GH, Ny, US). Mexico. CHIAPAS: ca 10 mi E of the Oaxaca-Chiapas border on rte 190, 23 Jun 1960, King 2980 (DS, NY, TEX, UC, US); ca 12 mi E of Cintalapa, 23 Jun 1960, King 2987 (DS, NY, TEX, UC, US); ca 33 mi S of Tuxtla Gutiérrez, 27 Jun 1960, King 3105 (Ds, NY, TEX, UC, US); 30 mi SE of Comitán, 26 Jul 1966, Stuessy 571 (TEX); 1 mi N of Entronque Santa Isabel, 10 Aug 1966, Stuessy 632 (TEX). COLIMA: Colima, 24 Oct 1910, Orcutt 4516 (ps, F, GH); Colima, 9 Jan-6 Feb 1891, Palmer 1172 (G, GH, NY, UC, US); 5 mi NW of rte 110 on rd to Alzada, 25 Aug 1966, Stuessy 724 (TEX); Alzada, 25 Aug 1966, Stuessy 727 (TEX); Man- zanillo, 1863-64, Xantus s.n. (F, GH [2], Ny [2], vs [2]). GUERRERO: 1972] Melampodium — Stuessy 25 Placeres, Puerta, 22 Jul 1936, Hinton 9134 (ARIZ, TEX, US); ca 9 mi NW of Taxco, 14 Mar 1961, King 4168 (F, NY, TEX, UC, US); ca 25 mi NE of Acapulco, 14 Mar 1961, King 4178 (F, NY, TEX, UC, US). JALISCO: Río Cuvianes, 13 Jun 1892, Jones 274 (pom [2], US); Río Cuvianes, 13 Jun 1892, Jones 369 (POM, US); % km S of Puente San Pedro on rd from Colima to Ciudad Guzmán, 31 Jul 1960, Koeppen & Iltis 618 (TEX, UC); 10 mi S of Autlán toward La Resolana, 28 Jun 1949, R. & C. Wilbur 1895 (us). MÉXICO: 8 km SW of Luvianos, 2 Sep 1965, Rzedowski 20748 (08). MICHOACÁN: Coalcomán, 5 Jan 1939, Hinton et al. 12871 (Ny, US); Uruapán, Tancitaro, 26 Oct 1940, Hinton et al. 15587 (ARIZ, GH, TEX, US); ca 32 km N of Playa Azul (near Los Encinos), 25-31 Oct 1961, King & Soderstrom 4992 (NY, SMU, TEX, UC, US); Apatzingán, canyon below Acahuato, 15 Aug 1941, Leavenworth & Hoogstraal 1589 (F, GH); 7 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 688 (TEX). NAYARIT: Mirador del Águila, ca 14 mi N of Tepic, 21 Aug 1959, Feddema 832 (MICH [2]); ca 25 mi N of Tepic, 12 Aug 1960, King 3699 (DS, NY, TEX, UC, US); vicinity of Acaponeta, Tepic, 10 Apr 1910, Rose, Standley & Russell 14307 (vs); Ixtlan del Río — San Marcos, 5 Aug 1921, Thompson s.n. (NY). OAXACA: Estación San Marcos, Istmo de Tehuantepec, 21 Jan 1907, Conzatti 1106 (F, GH, US) ; vicinity of Yalalag [Hidalgo Yalalag], Jul 1894, Nelson 947 (US); rd between Llano Grande & Pinotepa, 19 Feb 1895, Nelson 2339 (US); between Guichocovi & Lagunas, 27 Jun 1895, Nelson 2740 (GH, US). SAN LUIS POTOSÍ: San Dieguito, 13-16 Jun 1904, Palmer 459 (US). TAMAULIPAS: 10 km NW of El Progreso, 22 Aug 1941, Stanford, Retherford & Northeraft 1077 (ARIZ, DS, GH, NY, UC). VERACRUZ: La Purga, 27 Jan 1906, Greenman 279 (F, GH, NY); ca 26 mi E of Cuitláhuac, 6 Jun 1960, King 2682 (DS, NY, TEX, UC, US); ca 14 mi SE of Alvarado, 7 Jun 1960, King 2718 (Ds, NY, TEX, UC, US); vicinity of Pueblo Viejo, 2 km S of Tampico, 1-2 Jun 1910, Palmer 535 (GH, NY, US); 63 mi S of jet rtes 110 & 105, 6 Jul 1966, Stuessy 471 (TEX). 2. Melampodium diffusum Cass. Dict. Sci. Nat. 59 :238. 1829. TYPE: PHILIPPINE ISLANDS: Manila, summer 1825, F. L. Busseuil s.n. (Holotype, P?). Melampodium manillense Less. Linnaea 6:155. t. 2, f.G (fruit). 1831. TYPE: PHILIPPINE ISLANDS: “in Luconia" [Luzon], 1831, A. Chamisso s.n. (Holotype, B?; photograph of G-DC isotype (?), Ipc 800. 927:I. 6!). Melampodium lanceolatum DC. Prodr. 5:519. 1836. TYPE: PHILIPPINE ISLANDS or MEXICO: locality and date unknown, L. Née s.n. (Holotype, G-DC; isotype, MA!; photo- 26 Rhodora [Vol. 74 graph of holotype, Us!, pc 800. 927:1I. 6!; photograph of MA isotype, OS! TEX!). Melampodium diffusum Cass. var. lancelatum (DC.) Robins. Proc. Amer. Acad. Arts & Sci. 36:460. 1901. Annual herbs, 25-40 cm tall. Stems erect, 1-5 mm diam, subglabrous at base to strigillose-hispidulous above with hairs up to 0.8 mm long. Peduncles 0.8-6.5 em long. Leaves usually sessile (sometimes with petioles up to 3 mm long), lanceolate to narrowly ovate, 2.5-5.5 cm long, 0.5-1.5 (—3) cm wide, at apex acute, at base attenuate to somewhat obtuse, with upper surface strigose with hairs 0.3 mm long, with lower surface sericeous; margin entire to 1-3 cleft, hispid with hairs up to 1.5 mm long. Heads 5-7 mm tall, 6-11 mm diam. Outer involucre cupulate, 5-9 mm diam; bracts 5, slightly connate at base, imbricate 1/2 their length, ovate, 2.5-6 mm long, 1.5-4 mm wide, at apex acuminate, with abaxial surface strigillose with hairs 0.3 mm long, at margin herbaceous. Fruits 1.8-2.3 mm long, with lateral surfaces smooth and longitudinally ribbed to tuberculate- aculeate; hood apex mucronate to cirrhous (rarely muti- cous)^, with appendage up to 2 mm long. Ray florets 8-13; ligules yellow-orange, oblong-elliptic, 2.5-4 mm long, 1-3 mm wide. Disc florets 40-60; corollas yellow-orange, 1.3 mm diam, with throat 0.6 and tube 0.4 mm long. Paleae ob- lanceolate, 3 mm long, 0.9 mm wide; apex yellow, with margin entire to moderately erose; midrib prominent, most- ly glabrous. Chromosome number, n — 10. Tropical deciduous forests in Colima and near Acapulco, Guerrero, Mexico, and Manila in the Philippine Islands (Fig. 12), 30-300 m. Flowering dates, Jun-Jan. One collection, Winbery & Rowell 2492, approaches the perennial habit in this characteristically annual species. Be- cause this collection also possesses heads and leaves larger than usual, the suggestion of hybridization with nearby M. americanum, especially at higher altitudes in Guerrero, can- not be excluded. *One collection, Palmer 3 (F), has fruits with hoods and no hoods in one head of the single plant. 1972] Melampodium — Stuessy 27 Fig. 12. Map of Mexico, adjacent Guatemala, and the Philippine Islands (inset), showing distribution of Melampodium diffusum (triangles), M. longifoliwm (closed squares), M. longipilum (dots), M. mimulifolium (open square), and M. pilosum (circles). Robinson (1901, p. 456) previously noticed the unusual bicentric distribution of this species and offered the follow- ing plausible explanation: *As the genus is otherwise American, the occurrence of this species in a region so remote has always been prob- lematie, and it has been a matter of no small interest to find the Philippine plant closely matched by specimens recently collected by Dr. Edward Palmer, about Acapulco, Mexico. There can therefore be scarcely a doubt that the genus is in reality of New World origin, and that a single Mexican species was accidentally introduced into the Philippines, where it attracted scientific attention before it was recognized in America. This seems the more likely from the circumstance that Mexico and the Philippines were under the same national control, and early con- 28 Rhodora [Vol. 74 nected by a certain amount of oceanic traffic [demon- strated by Merrill (1954)]. This being the case, the transference of seed from Acapulco, the most important Pacific port of Mexico, to the neighborhood of Manila, presents no inherent improbability." REPRESENTATIVE SPECIMENS. Mexico. COLIMA: 5-10 mi N of Teco- man, 25 Dec 1958, Thompson & Fields 332( TEX). GUERRERO: above Playa Hornos ca 1 mi E of Acapulco, 9 Jan 1944, Barkley 14142 (F, GH, POM, TEX, UC, US); near Pie de la Cuesta NW of Acapulco, 5 Jan 1944, Barkley 14163 (TEx, us); 20 mi NE of Acapulco, 20 Aug 1947, Barkley, Webster & Paxson 17M724 (TEX); between Acapulco & Pueblo Nuevo, 13 Nov 1882, Hancock 25 (F, K); 8 mi E of Acapulco, 22 Jun 1952, Mockford & Rowell 2769 (SMU); near Acapulco, 25-31 Oct 1895, Palmer 3 (F, GH, Ny, UC, US); near Acapulco, Dec 1895, Palmer 281 (GH, US); 26 mi W of Acapulco, 19 Aug 1961, Powell & Edmondson 758 (F, TEX); 5 mi SW of Tierra Colorada, 17 Jun 1952, Winbery & Rowell 2492 (sMU); 1 mi N of Acapulco, 27 Aug 1965, Stuessy 366 (TEX); near S shore of Acapulco bay, 28 Aug 1965, Stuessy 369 (TEX). Philippine Islands. LUZON: Cavite, Aug 1905, Foxworthy 167 (NY, US); Cavite, Binacayan, 21 Jan 1917, Merrill 10641 (NY, US); Cavite, Bacoor, Jul 1910, Robinson 11835 (F); Laguna, 17 Apr 1953, Canicosa 1103 (US); Nueva Vizcaya, Bam- bang, Jan 1924, Clemens s.n. (UC); Nueva Vizcaya, Benquet, Dec 1908, Curran & Merritt 15837 (US); Nueva Vizcaya, Jan 1913, MeGregor 20181 (vs). 9. Melampodium pilosum Stuessy, Brittonia 22:115. f. 6. 1970. TYPE: MEXICO: Guerrero, Montes de Oca, Vallecitos, 28 Sep 1937, G. B. Hinton 11428 (Holotype, NY!; isotypes, G! GH! uc! us!). Annual herbs, 8-28 cm tall. Stems erect to decumbent, 1-2 mm diam, pilose with hairs up to 1.5 mm long. Pedun- cles 3-3.7 cm long. Leaves with petioles 2 mm long, lance- olate to narrowly ovate, 3-6 cm long, 1-1.3 cm wide, at apex acute, at base obtuse, with upper surface near apex strigose with hairs 0.3 mm long, near base pilose with hairs 1.5 mm long, with undersurface markedly sericeous; margin en- tire to rarely 1-2 lobed, irregularly ciliate with hairs 0.3 mm long, toward base pilose with hairs up to 2 mm long. Heads 6-7 mm tall, 7.5-11 mm diam. Outer involucre cupu- late, 6.5-9 mm diam; bracts 5, slightly connate at base, im- 1972] Melampodium — Stuessy 29 bricate 1/2 their length, ovate to narrowly ovate, 4-5 mm long, 2-2.5 ram wide, at apex acuminate, with abaxial sur- face pilose with hairs 1 mm long, at margin herbaceous. Fruits 1.6-2.5 mm long, with lateral surfaces 3-ribbed and smooth to very weakly tuberculate; hood apex mucronate to cirrhous?, with tapering appendage up to 2 mm long. Ray florets 7-8; ligules yellow-orange, ovate-oblong, 3-4.5 mm long, 2-3 mm wide. Disc florets 55-80; corollas yellow, 1.2 mm diam, with throat and tube each 0.7 mm long. Paleae oblanceolate, 4 mm long, 1 mm wide; apex yellow, with margin erose becoming laciniate laterally; midrib promi- nent, hirsutulous with hairs 0.3 mm long. Chromosome number, » — 10. Tropical deciduous forests in Michoacán and Guerrero, Mexico (Fig. 12), 910-1220 m. Flowering dates, Aug-Sep. REPRESENTATIVE SPECIMENS. Mexico. GUERRERO: Coyuca, Chaca- merito, 14 Aug 1934, Hinton 6436 (F, G, GH, NY [2], US). MICHOACÁN: ca 1 mi N of Huacana, 11 Aug 1965, Melchert & Sorensen 6085 (ENCB); 25 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 695 (TEX). 4. Melampodium longipes (A. Gray) Robins. Proc. Amer. Acad. Arts & Sci. 36:459. 1901. Melampodium sericeum Lag. var. longipes A. Gray. Proc. Amer. Acad. Arts & Sci. 22:423. 1887. TYPE: MEXICO: Jalisco, Tequila, “base of mountains," Aug 1886, E. Palmer 391 (Holotype, GH ! ; isotypes, G! K! Mo! Ny 12]! US!; photograph of Us isotype, TEX!). Annual herbs, 10-46 cm tall. Stems erect, 1-2 mm diam, hispid-pilose with hairs up to 1 mm long. Peduncles 5-8 cm long. Leaves sessile, linear and entire to markedly pin- nately parted, 3-7 cm long, 0.3-3.5 cm wide, at apex acute- obtuse, at base mostly attenuate (less often obtuse), with upper surface strigose with hairs 0.3-1 mm long, with lower surface sericeous; margin entire, at base hispid-pilose with hairs up to 2 mm long. Heads 6.5-8.5 mm tall, 10-12 mm *The holotype has fruits possessing hoods with both mucronate and cirrhous apexes. 30 Rhodora [Vol. 74 Fig. 18. Map of central Mexico showing distribution of Melam- podium dicoelocarpum (triangles), M. glabrum (dots), M. longipes (squares), and M. nutans (circles). One collection of M. nutans from Oaxaca not shown. diam. Outer involucre cupulate, 7-8 mm diam; bracts 5, slightly connate at base, imbricate 2/3 their length, obovate, 4-5 mm long, 2.5-3 mm wide, at apex acuminate, with abaxial surface pilose with hairs 1 mm long (more dense near base) ; margin herbaceous. Fruits 2-3 mm long, with lateral surfaces tuberculate with 3-4 longitudinal ribs; hood apex mucronate to cirrhous, with tapering appendage up to 3 mm long. Ray florets 8-10; ligules yellow-orange, ovate- oblong, 3-5 mm long, 1-3 mm wide. Disc florets 50-80; corollas yellow-orange, 2 mm diam, with throat 0.8 mm and tube 0.7 mm long. Paleae oblanceolate, 3.5 mm long, 0.8 mm wide; apex yellow, with margin entire to undulate; midrib distinct, mostly glabrous. Chromosome number, n = 10. 1972] Melampodium — Stuessy 31 Dry hillsides in Jalisco, Nayarit and Sinaloa, Mexico (Fig. 13), ca 1220 m. Flowering dates, Aug-Sep. This species is very similar to both M. americanum and M. linearilobum. M. longipes usually can be distinguished from either of these two taxa by its annual habit (peren- nial in M. americanum) and herbaceous outer involucral bracts (conspicuously scarious in M. linearilobum). The leaves divided markedly into pinnatifid segments and the yellowish hue of the stems also serve as additional features to help recognize M. longipes from among its close relatives. One collection, Rose 3183, from Sinaloa tends more to- ward a perennial habit, and the leaves are more distinctly pinnatifid. Nevertheless, these specimens appear to fall within the limits of M. longipes. REPRESENTATIVE SPECIMENS. Mexico. JALISCO: ca 2 mi NW of Tequila, 8 Aug 1960, King 3662 (DS, MICH, NY, TEX, UC, US); Tequila, 22 Sep 1893, Pringle 4598 (F, G [2], GH, MICH, MSC, NY, P, UC, US); 2 mi NW of Tequila, 3 Sep 1965, Stuessy 397 (TEX), 26 Aug 1966, Stuessy 738 (TEX). NAYARIT: El Maguey, Sierra Madre, Tepic, 26 Aug 1905, Goldsmith 132 (Ds, F, GH, NY, UC, US). SINALOA: be- tween Rosario & Colomas, 13 Jul 1897, Rose 3183 (GH, US). 5. Melampodium linearilobum DC. Prodr. 5:518. 1836. TYPE: MEXICO: “ad Guilotepec et Las Bacas," Sep 1831, L. Alamán s.n. (Holotype, G-DC; isotype, G!; photograph of holotype, inc 800. 927: II. 2!; photograph of G isotype, F! os! TEX! us!). Melampolium canescens Brandg. Zoe 5:222. 1905. TYPE: MEXICO: Sinaloa, Cofradia [ca 30 mi E of Culiacán near Durango border], 22 Oct 1904, T. S. Brandegee s.n. (Lecto- type chosen, UC!; isotypes, GH! POM! US!). Annual herbs, 6-50 cm tall. Stems erect, 1-3 mm diam, strigillose-strigose with hairs 0.8 mm long. Peduncles 3.5- 5.5 em long. Leaves sessile, linear and entire to markedly pinnately parted, 3-5.5 cm long, 0.3-3 em wide, at apex acute-obtuse, at base attenuate to obtuse, with upper sur- face strigose with hairs 0.2-1 mm long, with lower surface sericeous; margin entire, at base hispid-pilose with hairs up to 2 mm long. Heads 5-7 mm tall, 7-11 mm diam. Outer 32 Rhodora [Vol. 74 involucre cupulate, 6-10 mm diam; bracts 5, slightly con- nate at base, imbricate 1/2 their length, narrowly ovate to somewhat obovate, 3.5-5.5 mm long, 2.5-3 mm wide, at apex acute, with abaxial surface pilose-strigose with hairs 1-1.8 mm long; margin conspicuously scarious. Fruits 2-3 mm long, with lateral surfaces smooth and striate to tubercu- late; hood apex muticous to cirrhous, with appendage up to 2.5 mm long. Ray florets 7-8; ligules yellow-orange, ovate-orbicular (often with imbricate margins), 2-4 mm long, 2-5 mm wide. Disc florets 45-75; corollas yellow- orange, 2.4 mm diam, with throat 0.9 mm and tube 0.5 mm long. Paleae oblanceolate, 4 mm long, 1 mm wide; apex yellow, with margin entire (rarely erose) ; midrib promi- nent, mostly glabrous. Chromosome number, » — 10. Tropical deciduous and pine forests of the western Sierra Madres of Mexico and into Central America to Costa Rica (Fig. 14), 20-1620 m. Flowering dates, Jul-Oct. The conspicuous scarious margins (colored orange-yel- low when fresh) of the outer involucral bracts of M. linear- ilobum. The type of M. canescens from Sinaloa has linear, members of series Melampodium. Were it not for this char- acteristic bract feature, M. linearilobum would be very dif- fieult to distinguish from M. longipes. In spite of considerable vegetative variability, no intra- specific categories have been recognized within M. linear- ilobum. The type of M. canescens from Sinaloa has linear, entire leaves, as do other collections from nearby areas, but plants with similar leaves also appear in Central America and in scattered localities in Mexico. As a further indica- tion of the plasticity of leaf shape, in many plants the upper leaves are linear and entire, while those lower on the stem are once or twice pinnatifid. Due to this vegetative plastic- ity, M. canescens is considered herein to be synonymous with M. linearilobwm. REPRESENTATIVE SPECIMENS. Costa Rica. GUANACASTE: Bolson River, 16 Jul 1918, W. & H. Rowlee 179 (Ny, US). El Salvador. AHUACHAPÁN: 1921, Padilla 1 (US), 1921, Padilla 242 (us), 1923, Q2 e» 1972] Melampodium — Stuessy Fig. 14. Map of Mexico and Central America showing distribu- tion of Melampodium linearilobum. Padilla 586 (US). MORAZAN: ca 8 km SW of Montecristo, 3 Dec 1941, Tucker 455 (F, GH, NY, SMU, UC, US). SANTA ANA: near Chalchuapa, 1922, Calderén 979 (GH, NY, US). USULUTAN: 24 mi E of turnoff to San Vicente on rte 1, 3 Aug 1966, Stwessy 613 (TEX). Guatemala. CHIQUIMULA: near Ipala, 23 Oct 1939, Steyermark 30309 (F). HUEHUETENANGO: between San Ildefonso Ixtahuacán & Cuilco, 16 Aug 1942, Steyermark 50724 (F, NY). JUTIAPA: between Suchitán & Sta Catarina, Jul 1870, Bernoulli 709 (NY); near Jutiapa, 20 Dec 1938, Standley 60493 (F); between Jutiapa & Plan de Urrutia, 28 Oct 1940, Standley 75620 (F, US); 8 mi NE of Jutiapa, 2 Aug 1966, Stuessy 607 (TEX). QUICHE: 1942, Ignacio 1495 (F). ZACAPA: near divide on rd between Zacapa & Chiquimula, 9 Oct 1940, Standley 73774 (F); between Río Hondo & Santa Cruz, 11 Oct 1940, Standley 74071 (F); between Río Hondo & Santa Cruz, 11 Oct 1940, Standley 74125 (F, US); near La Fragua, 14 Oct 1940, Standley 74813 (F); near Sta Rosalia, 4 Oct 1939, Steyermark 29041 (F). Honduras. CHOLUTECA: vicinity of Pespire, 18-27 Oct 1950, Standley 27175 (F). FRANCISCO MORAZÁN: El Chile, Tegucigalpa, 18 Sep 1950, 34 Rhodora [Vol. 74 Standley 26708 (F). OLANCHO: near Rio Telica, 20 Nov 1963, Molina 13346 (F). VALLE: 21 Jan 1956, Molina 5899 (F). Mexico. CHIAPAS: Chicomucelo, 14 Jul 1941, Matuda 4444 (4504) (F); valley of Jiquipilas, 16-18 Aug 1895, Nelson 2949 (GH, US). DURANGO?: Lodiego [25°N, 106*45'W; McVaugh, 1956], 9-15 Oct 1891, Palmer 1609 in part (vs [2]). GUERRERO: Coyuca de Catalán, Pungarabato, 10 Jul 1934, Hinton et al. 6270 (F, GH, NY, US); Mina, Placeres, 18 Aug 1936, Hinton et al. 9118 (ARIZ, GH, TEX, US); near Iguala, 26 Apr 1900, Pringle 9162 (F, GH, Ny, US); Los Amates Station, 28 Sep 1905, Pringle 10065 (ARIZ, F, G, NY, SMU, UC, US); ca 50 mi S of Cuernavaca, 26 Aug 1965, Stuessy 361 (TEX). MÉXICO: Tema- scaltepec, Limones, 7 Nov 1932, Hinton 2513 (GH, NY [2]); Tema- scaltepec, Palmar, 9 Aug 1934, Hinton et al. 6420 (Ny, US); San Antonio, Tlatlaya, 20 Jul 1954, Matuda et al. 31141 (vs); Palmar Chico, Amatepec, 24-25 Aug 1954, Matuda et al. 31314 (Us). MICHOACAN: Zitacuaro, Zitacuaro-Tiamaro, 9 Sep 1938, Hinton et al. 13214 (ARIZ, GH, TEX, US); Tancitaro, above Apatzingan, 15 Aug 1941, Leavenworth & Hoogstraal 1480 (F, GH, NY); Apatzingán, below Acahuato, 15 Aug 1941, Leavenworth & Hoogstraal 1588 (F); 1 mi N of Apatzingán, 21 Aug 1966, Stuessy 697 (TEX); 4 mi N of Nueva Italia, 21 Aug 1966, Stuessy 698 (OS, TEX). OAXACA: Salina Cruz, 23 Aug 1935, Fisher 35321 (ARIZ, F, NY, SMU, US); ca 37 mi W of Tehuantepec, 23 Jul 1960, King 3454 (DS, Ny, TEX, UC, US) ; ca 3 mi NE of Huajuapán de León, 28 Jul 1960, King 5539 (TEX); Playa de Puerto Ángel, Oct 1917, Reko 3812 (vs); 24 mi S of Sola de Vega, 18 Aug 1966, Stuessy 650 (TEX). SINALOA: Cerro Colorado, near Cofradia, 2 Nov 1904, Brandegee s.n. (GH, UC); Cofradia, 25 Nov 1939, Gentry 5039 (ARIZ, GH, NY); Las Mesas, Sierra Surotato, 25 Aug 1941, Gentry 6153 (ARIZ, GH, NY); La Noria, 10 Oct 1925, Mexia 240 (uc), 14 Oct 1925, Mexia 349 (POM, UC, US). Nicaragua. GRANADA: 1 mi S of Granada, 6 Aug 1966, Stuessy 619 (TEX). 2. Series Leucantha Stuessy, ser. nov. Plantae perennes suffruticosae; folia anguste ovata vel lineata, saepe lobata, subtus strigosa; ligulae eburneae, plus quam vel raro circa 2 mm longae; involucrum extimum marginibus herbaceis; fructi cucullati. Species 6-8. Typus: Melampodium leucanthum Torr. & A. Gray. 6. Melampodium leucanthum Torr. & A. Gray, Fl. N. Amer. 2(2) :271. 1842. TYPE: TEXAS: without locality or date, J. L. Riddell s.n. (Holotype, Nv?). Type not located, but the following collection cited by Gray (1852) clearly is repre- 1972] Melampodium — Stuessy 35 sentative: “Hills near El Paso," Apr 1849, C. Wright 311 (GH! UC! UST): Perennial subshrubs, 15-60 cm tall. Stems ascending, 1.1- 2.5 mm diam, strigillose (rarely hispid) with hairs 0.1-0.2 mm long. Peduncles 3-7 cm long. Leaves sessile, linear- oblong, 2.1-4.5 em long, 0.13-1 cm wide, at apex and base obtuse, with both surfaces strigillose with hairs 1-2 mm long; margin entire to pinnately 6-lobed, at base strigillose with hairs 0.1-0.2 mm long. Heads 6-8 mm tall, 20-37 mm diam. Outer involucre cupulate, 10-13 mm diam; bracts 5, connate 1/2 to 3/4 their length, ovate, 5-7 mm long, 4-5.1 mm wide, at apex acute, with abaxial surface strigillose- strigose with hairs 0.1-0.7 mm long; margin herbaceous. Fruits 1.5-2.6 mm long, with lateral surfaces aculeate-ver- rucate; hood apex muticous (very rarely mucronate). Ray florets 8-13; ligules cream-white, oblong-elliptic, 7-13 mm long, 2.5-8 mm wide. Disc florets 25-50; corollas yellow, 1 mm diam, with throat 1.2 mm and tube 0.8 mm long. Paleae oblong-elliptic, 2.9 mm long, 1.1 mm wide; apex yellow, with margin laciniate; midrib prominent, often strigillose with hairs 0.1 mm long. Chromosome numbers, » — 10 and 20. Caleareous soils throughout western and central Texas (stopping at the Edward's Plateau), Arizona, New Mexico, and portions of Oklahoma, Kansas, Colorado, and northern Mexico (Fig. 15), 490-2590 m. Flowering dates, Apr-Sep. This species is the familiar ‘‘blackfoot daisy” of the southwestern United States that has been gathered exten- sively by many different collectors. Despite its abundance, however, the relationships to M. cinereum and M. argophyl- lum have been unclear for many years. Recent studies have shown (Stuessy, 1971c; present treatment) that M. leucan- thum has a separate distribution that comes only within ten miles of M. cinereum at the edge of the Edward's Plateau. Both 2n and 4n cytological races have been reported for this species (Turner and King, 1962; Stuessy, 1971b & c), 36 Rhodora [Vol. 74 Fig. 15. Map of the southwestern United States and adjacent Mexico showing distribution of Melampodium lewcanthum. and they are morphologically indistinguishable. Evidence for treating these cytotypes as informal races having been derived by spontaneous autopolyploidy has been reviewed by Stuessy (1971c). REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: 7 mi N of Mestenas, 25 Sep 1938, Johnston 7959 (GH, US); Organos Mts, 8 Sep 1937, LeSueur 1456 (F, TEX); Colonia Diaz, 20-21 Sep 1899, Nelson 6447 (GH, US); hills near Chihuahua, 5 Oct 1886, Pringle 1019 (F, NY, US); 20 mi S of Ciudad Jüarez, 21 Jul 1937, Shreve 7918 (ARIZ, GH, US); 59 mi N of Villa Ahumada, 23 Aug 1967, Stuessy 1122 (TEX). COAHUILA: Del Carmen Mts, 6 Sep 1936, Marsh 855 (F, GH, TEX); Cañon del Indio Felipe, close to the Chihuahuan boundary, 27-29 Sep 1940, Stewart 84 (GH). SONORA: Colonia Morelos, 15 Sep-4 Oct 1941, White 4587 (ARIZ, GH); 5 mi E of Esqueda on rd to Río de la Tierra, 10 May 1948, Wiggins 11765 1972] Melampodium — Stuessy 37 (Ds, US). UNITED STATES. Arizona: COCHISE CO., near Douglas, 16 Aug 1907, Goodding 2399a (DS, GH, US). GILA CO., ca 3 mi N of Dripping Springs turnoff, 11 Apr 1965, Niles 572 (ARIZ, TEX); GREENLEE CO., 7 mi N of Metcalf, 6 Jun 1935, Maguire, Richards & Moeller 11849 (ARIZ, GH, NY); MARICOPA CO., 3 mi N of Sunflower, 9 Apr 1960, Russell 11511 (smu [2], UC); MOHAVE co., Kingman, 18 Apr 1935, Kearney & Peebles 11133 (ARIZ, GH, US); PIMA CO., 3 mi N of Greaterville, 17 Aug 1967, Turner 5735 (TEX); YAVAPAI CO., 10 mi SW of Congress, 21 Apr 1962, Turner 4790 (DS, FSU, SMU, TEX); YUMA CO. Squaw Canyon, Harquahala Mts, 9 Sep 1952, Wright 42-54 (ARIZ). Colorado: BACA co. 25 mi S of Pritchett, 11 Jul 1947, Porter 4262 (GH, RSA, SMU, TEX); EL PASO CO., below Colorado Springs, 29 May 1878, Jones 117 (F, POM [2]); FREMONT co., Canon City, 2 Jul 1920, Clokey 3946 (DS, F, GH, NY, POM, UC, US) ; PROWERS CO. 31 mi S of Lamar, 6 Jun 1967, Irving 825 (TEX); PUEBLO CO., 12 mi S of Pueblo, 7 Jun 1922, Wiegand & Upton 4439 (NY). Kansas: FORD CO., 3 mi S of Dodge City, 9 Jul 1950, Horr 3444 (GH, US); HAMILTON CO., vicinity of Syracuse, 15 Sep 1912, Rose & Fitch 17043 (NY, US); MEADE CO., 16 mi SE of Meade, 16 Aug 1939, Horr & Franklin E278 (F, GH, SMU, TEX, UC, US) ; SEWARD CO., Jul 1891, Carleton 331 (ARIZ, F, US). New Mexico: BERNALILLO CO. ca 2 mi E of Albuquerque, 1915, Kammerer 48 (DS, TEX); CHAVES CO., 20 mi S of Roswell, Aug 1900, F. & E. Earle 295 (NY, POM, US); DE BACA CO., 6 mi E of Yeso, 8 Aug 1967, Turner 5673 (TEX); DONA ANA CO., Organ Mts, 9 Jul 1897, Wooton 117 (ps, GH, NY, POM, UC, US) ; GRANT CO., Mangas Springs, 18 mi NE of Silver City, 15 May 1903, Metcalfe 66 (ARIZ, DS, NY, POM, UC [2], vs); LINCOLN CO., 15 mi E of Capitan, 25 Jul 1938, Hitchcock, Rethke & van Raadshooven 4244 (DS, GH, UC); ROOSEVELT CO., 5 mi NE of Portales, 14 Jul 1930, Goodman & Hitchcock 1130 (ps, GH, NY, UC) ; SAN MIGUEL co., 10 mi SE of Las Vegas, 27 Jul 1924, Bacigalupi 607 (Ds, GH, UC) ; SANTA FE CO., 20 mi S of Golden, 24 Jul 1938, Hitchcock, Rethke & van Raadshooven 4214 (DS, GH, UC); SIERRA CO., Kingston, 29 Aug 1904, Metcalfe 1271 (F, GH, NY, POM, UC, US). Oklahoma: BEAVER co., near Knowles, 5 May 1913, Stevens 325 (ps, NY, SMU, US); CIMARRON CO., Kenton, low sides of Black Mesa, 28 Jul 1936, Demaree 13344 (GH, NY, POM, SMU); JACKSON CO. El Dorado, 18 Apr 1936, Demaree 12210 (SMU, US). Texas: ANDREWS CO, 5 mi WSW of Andrews, 15 Sep 1966, Shinners 31559 (SMU); BELL CO. 2% mi S of Nolanville, 7 Mar 1954, C. & G. York 54037 (SMU, TEX); BEXAR co., 5 mi NW of San Antonio, 4 May 1948, Burr 135 (ps, Ny [2]); BLANCO CO. 10 mi N of Johnson City, 18 Apr 1958, Thompson & Graham 18 (SMU, TEX); BREWSTER CO. Marathon, 15 Jul 1965, Stuessy 230 (TEX); BROWN CO., 7 mi S of Brownwood, 24 Apr 1966, Guthrie 43 (SMU, TEX); CALLAHAN CO., ca 2 mi W of Baird, 27 Mar 38 Rhodora Vol. 74 1968, Henderson 63-88 (FSU, SMU, TEX); COLEMAN CO., 1 mi SE of Santa Ana, 9 Jul 1957, Shinners 26390 (SMU); COMAL CO., Comanche Spring, Mar 1849, Lindheimer 949 (ARIZ, F, GH, NY, TEX, UC, US); COMANCHE CO. 7 mi W of Comanche, 6 Apr 1966, Teeters 18 (LL, SMU, TEX); CORYELL CO., 4 mi S of Gatesville, 2 Oct 1965, Baize 5 (LL, SMU, TEX); CROCKETT CO., 31 mi W of Ozona, 8 Jul 1965, Stuessy 147 (TEX); CULBERSON CO. 1 mi S of Texas-New Mexico line E of Guadalupe Mts, 14 Aug 1942, Waterfall 3781 (ARIZ, GH, NY) ; DAWSON co., plains between Lamesa & Tahoka, 29 Apr 1925, Small & Wherry 12134 (NY, TEX); ECTOR CO., 5.8 mi S of Odessa, 8 May 1966, Shinners 31236 (SMU); EL PASO CO., 28 mi SE of El Paso, 24 Aug 1967, Stuessy 1127 (TEX); ERATH CO., Stephenville Pk, 9 mi N of Stephen- ville, 2 Apr 1950, Shinners 12190 (SMU); FLOYD CO., intersec- tion of hwys 97 & 2009, 22 Apr 1962, Melchert 180 (TEX); GIL- LESPIE C0., 4.8 mi NE of Lawrence Jung Ranch, 31 Mar 1967, Mears 1383 (TEX); HAMILTON CO. 17 mi S of Hico, 16 Apr 1945, Shinners 7183 (GH, LL, SMU, UC); HARDEMAN CO., 11 mi S of Quanah, 4 Apr 1966, Turner 7 (LL, TEX); HAYS CO., 1.5 mi E of Dripping Springs, 30 May 1966, Stuessy 414 (TEX); HEMPHILL Co., 18 mi SW of Canadian, 15 Sep 1950, Tharp & Miller 51-312 (RSA, TEX); HUDSPETH CO., ca 20 mi N of Allamore, 29 Jul 1943, Waterfall 4818 (GH, NY, SMU); JEFF DAVIS CO. gravel flat between Big & Little Aguja Canyons, Davis Mts, 16 Jun 1931, Moore & Steyermark 3115 . (DS, GH, NY, UC, US); KENDALL CO. 5 mi W of Sisterdale, 24 May 1965, Flyr 489 (ns, SMU); KERR CO., Kerrville, 19-25 Apr 1894, Heller 1632 (ARIZ, F, GH, NY [2], POM, SMU, UC [2], US); KNOX CO., 5 mi E of Benjamin, 22 Apr 1962, Melchert 169-C (TEX); LLANO CO., Marble, 31 Mar 1967, Mears 1413 (TEX); LOVING CO, 1 mi W of Mentone, 10 Jul 1965, Stuessy 180 (TEX); LUBBOCK CO., Lubbock, 4 May 1930, Demaree 7594 (DS, GH, US); NOLAN CO., Sweetwater, 5 May 1927, Stanfield s.n. (NY, TEX); OCHILTREE CO., 7.8 mi SE of Perryton, 13 Jul 1957, Wallis 4807 (ARIZ, SMU); OLDHAM CO., 16 mi N of Vega, 17 May 1967, Turner 5632 (TEX); PECOS CO., 10 mi E of Ft. Stockton, 19 Apr 1946, Warnock 46217 (FSU, TEX); POTTER co., 1 mi N of Canadian River Bridge, 19 May 1945, Jespersen 2678 (DS, F, NY, SMU, UC, US); PRESIDIO CO., 7.5 mi N of Candelaria, 17 Apr 1947, MceVaugh 7986 (ps, F, LL [2], SMU, TEX [2]) ; RANDALL co., Palo Duro State Pk, 20 Oct 1945, Cory 50422 (GH, NY, SMU); SHACKELFORD CO., 2 mi NE of Jones Co. line, 27 Mar 1963, Henderson 63-98 (SMU, TEX); STEPHENS CO. 3 mi N of Ranger, 29 Apr 1939, Culwell & Timmons 3091 (SMU, TEX); TAYLOR CO, 5 mi SW of Abilene, 30 Jun 1962, Turner & Melchert 4840 (TEX); TRAVIS CO., 7 mi SW of Zilker Pk, Austin, 13 Oct 1966, Stuessy 752 (0s, TEX) ; UVALDE CO., Garner State Pk, 21 Jun 1958, Sullivan & Turner 34 (FSU, TEX); VAL VERDE CO, 13 mi W of Langtry, 23 Mar 1941, 1972] Melampodium — Stuessy 39 Innes & Warnock 589 (DS, GH, NY, TEX); WARD CO., Barstow, 15 Apr 1902, Tracy & Earle 23 (F, GH, NY [2], TEX, US), WILBARGER CO., 16.9 mi S of Electra-Waggoner pastures, 12 May 1945, Whitehouse 9841 (SMU, TEX); WINKLER CO., 1 mi W of Kermit, 10 Jul 1965, Stuessy 167 (TEX). 7. Melampodium cinereum DC. Prodr. 5:518. 1836. Perennial subshrubs, 14-20 cm tall. Stems suffruticose, ascending. Leaves sessile, at apex and base obtuse, at mar- gin revolute when dry. Outer involucre cupulate; bracts 5, connate 1/6 to 1/3 their length, ovate, at apex acute, with abaxial surface strigose with hairs 0.2-1 mm long, at mar- gin herbaceous. Fruits with lateral surfaces aculeate-ver- rucate. Ligules cream-white, oblong-elliptic. Disc corollas yellow. Paleae oblong-oblanceolate; apex yellow, with mar- gin laciniate; midrib prominent, glabrous or puberulous with hairs 0.1 mm long. Ta. Melampodium cinereum DC. var. cinereum Melampodium cinereum DC. Prodr. 5:518. 1836. TYPE: MEXICO[?]: San Fernando de las Presas, Oct 1830, J. L. Berlandier 2243 [—823] (Holotype, G-DC; isotypes, G! GH! K!; photograph of holotype, Ipc 800. 927: II. 4!; photo- graph of K isotype, B! US!; photograph of G isotype, F! Us!). San Fernando is a town about 83 miles SW of Matamoros, Tamaulipas, Mexico, an area visited by Berlandier (Geiser, 1948). However, this variety is known to occur only near Laredo and the surrounding territory in southcentral Texas, indicating that the locality may have been interchanged with M. ramosissimum as suggested by Gray (1884) and Robinson (1901). Stems 0.8-1.3 mm diam, strigose with hairs 0.1-0.8 mm long. Peduncles 1.5-7 cm long. Leaves linear-oblong, 1.2- 5.0 em long, 0.12-1.4 cm wide, with both leaf surfaces strigose with hairs 0.2-0.6 mm long; margin entire to pin- nately 10-lobed or parted, near base strigose with hairs up to 0.6 mm long. Heads 5-8 mm tall, 9-23 mm diam. Outer involucre 7-13 mm diam; bracts 3.5-7.3 mm long, 2.3-5 mm wide. Fruits 1.4-2.2 mm long; hood apex muti- 40 Rhodora [Vol. 74 Fig. 16. Map of southern Texas and adjacent Mexico showing distribution of Melampodium cinereum var. cinereum, small form (circles), large form (dots); M. cinereum var. hirtellum (open squares); M. cinereum var. ramosissimum (closed squares); and M. argophyllum (triangles). Shaded area indicates generalized south- eastern distribution of M. leucanthum. cous to cirrhous, with appendage up to 2.2 mm long. Ray florets 7-13; ligules 3-8.2 mm long, 1-3.6 mm wide. Disc florets 30-50; corollas 1.1 mm diam, with throat 0.8 mm and tube 0.7 mm long. Paleae 2.5 mm long, 1.1 mm wide. Chromosome numbers, » — 10 and 20. 1972] Melampodium — Stuessy 41 Mesquite-grasslands of Rio Grande Plains of Texas south to Cameron Co., east to Jim Wells Co., north to Bexar Co. and west to Maverick Co., extending into Tamaulipas and Nuevo Leon, Mexico, 30-240 m (Fig. 16). Flowering dates, Mar-Nov. All the varieties of M. cinereum can be distinguished most easily from M. leucanthum and M. argophyllum by the outer phyllaries that are connate only 1/4 their length at the base. The outer involucral bracts of the latter two species always are fused at least 1/2 or even 2/3 their length. The usefulness of this feature has not been pre- viously noted, but it is the best diagnostic morphological character that I have discovered for separating M. cinereum from the other taxa of the white-rayed complex. Scattered within the range of var. cinereum are two morphological forms (small and large) that are weakly differentiated by quantitative features of habit and head size. These informal units presumably correspond to the 2n and 4n autopolyploid chromosomal races reported within this variety by Turner and King (1962) and Stuessy (1971b & c). Additional information concerning the presumptive autopolyploid origin of these races can be found in Stuessy (1971c). REPRESENTATIVE SPECIMENS (SMALL FORM). Mexico: NUEVO LEÓN : S of Nuevo Laredo, 8 Jun 1935, Clark 6635 (NY); 15 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 854 (TEX). TAMAULIPAS: 5 km S of Nuevo Laredo, 11 Nov 1961, Domínguez 63 (TEX); 24 km S of Nuevo Laredo, 24 Mar 1962, Dominguez & McCart 8217 (SMU, TEX); 59 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 857 (TEX). UNITED STATES. Texas: CAMERON CO., Santa Ana National Wild- life Refuge, 26 Sep 1961, Fleetwood 3840 (TEX); DIMMIT CO., 3 mi W of Bigwells, 21 Apr 1945, Shinners 7395 (GH, LL, SMU, UC) ; DUVAL co., 4 mi NE of Freer, 8 Oct 1954, Tharp & Johnston 541 [802A] (TEX); FRIO CO., 5.5 mi S of Pearsall, 1 May 1954, Johnston, Tharp & Turner 3472 (SMU, TEX); HIDALGO CO., 6 mi E of Sullivan City, 8 Mar 1959, Turner 4490 (TEX); JIM HOGG CO., 8 mi E of Hebbron- ville, 12 Jun 1952, Jones 719 (SMU); JIM WELLS CO., near Alice, 21 Jun 1935, Drushel 10441 (NY); LA SALLE CO., 1 mi W of Encinal, 21 Apr 1963, Sánchez 105 (SMU, TEX); MAVERICK CO. 1 mi E of Eagle Pass, 22 May 1898, Bray s.n. (TEX); STARR CO., 5.8 mi E of 42 Rhodora [Vol. 74 Rio Grande City, 9 Oct 1954, Tharp & Johnston 541922 (TEX); WEBB CO., 22 mi NW of jct rtes 83 & 35, 5 Jul 1967, Stuessy 868 (TEX); ZAPATA CO., 13 mi N of San Ygnacio, 31 Jan 1954, Shinners 17659 (SMU); ZAVALA CO., 6 mi S of Batesville, 6 May 1964, Turner 5006 (SMU, TEX). REPRESENTATIVE SPECIMENS (LARGE FORM). Mexico: TAMAU- LIPAS: 3 mi S of Nuevo Laredo, 11 Nov 1961, Escalante 21 (TEX); 50 mi SE of Nuevo Laredo, 28 Mar 1964, A. & R. Garcia 47 (ARIZ, SMU, TEX) ; 12 mi SE of Nuevo Laredo, 17 Mar 1962, C. & L. de la Garza 46 (SMU, TEX). UNITED STATES. Texas: BEXAR CO., High- land Hills, San Antonio, 1 Apr 1963, Martinez & García 13 (SMU); BROOKS CO., 15 mi E of Hebbronville, 17 Mar 1968, Ramirez 44 (UL, TEX); CAMERON CO., 12 mi N of Brownsville, 18 Apr 1965, Ríos & Cavazos 264 (LL, SMU); DUVAL CO., 25 mi N of Hebbronville, 20 Jun 1966, Stuessy 429 (TEX); FRIO CO. 9 mi S of Moore, 20 Nov 1967, Stuessy & Renold 1284 (TEX); HIDALGO CO., 1 mi E of Sullivan City, 1 Apr 1941, C. & A. Lundell 9866 (LL [3], RSA); JIM HOGG CO., 5 mi E of Hebbronville, 20 Jun 1966, Stuessy 425 (TEX); JIM. WELLS CO., 2 mi E of San Diego, 25 Nov 1962, Dohnke 3 (SMU, TEX); KINNEY co., [?], Bracket [Brackettville?], 21 Mar 1900, Canby 133 (vs); LA SALLE CO., 1 mi N of Encinal, 16 Mar 1963, Solis 124 (SMU, TEX) ; LIVE OAK CO., ca 32 mi S of Whitsett, 3 Jun 1967, Stuessy 771 (TEX); MCMULLEN CO., 22% mi SW of George West, 17 Apr 1965, Ríos & Cavazos 231 (LL, SMU); MEDINA CO., ca 3 mi S of Devine, 28 Oct 1952, Correll 15204 (LL, US); NUECES CO., ca 4 mi W of Mathis, 8 May 1957, Jones 1359 (SMU); STARR CO., 3 mi N of Roma, 31 Jan 1954, Shinners 17708 (SMU); UVALDE CO., Leona River, 7 mi SE of Uvalde, 23 Jun 1935, Munz 13303 (POM); WEBB CO., 14 mi NE of Laredo, 9 Mar 1963, Cisneros 33 (LL); ZAPATA CO., 10 mi N of San Ygnacio, 7 Apr 1963, Arzola 178 (LL, SMU). 7b. Melampodium cinereum DC. var. hirtellum Stuessy, Sida 3:348. 1969. TYPE: MEXICO: Nuevo León, 5 km S of Sabinas Hidalgo on rd to Monterrey, 21 Apr 1939, T. C. & E. M. Frye 2415 (Holotype, US!; isotypes, DS! GH! NY! RSA! UC!). Stems 0.8-1.3 diam, strigose-hispid with hairs 0.2-1.5 mm long. Peduncles 0.7-8 cm long. Leaves linear-oblong, 1.1-3.4 em long, 0.2-1 cm wide, with both surfaces strigose with hairs 0.3-0.8 mm long, with midrib on undersurface hispid with hairs up to 1.5 mm long; margin pinnately 8- lobed or parted, near base hispid with hairs 0.4-1.5 mm long. Heads 4.5-5 mm tall, 9-18 mm diam. Outer involucre 7-10 mm diam; bracts 3.3-6 mm long, 2.2-3.6 mm wide. 1972] Melampodium — Stuessy 43 Fruits 1.3-2 mm long; hood apex muticous to mucronate, with appendage up to 0.3 mm long. Ray florets 8-13; ligules 3-6 mm long, 1.3-2.8 mm wide. Disc florets 25-50; corollas 1.1 mm diam, with throat 0.7 mm and tube 0.5 mm long. Paleae 3.2 mm long, 0.9 mm wide. Chromosome number, n — 10. Mesquite-grasslands in eastern Coahuila, northwestern Nuevo León and Tamaulipas, Mexico, and into Val Verde, Kinney, Uvalde and Maverick Cos. of Texas (Fig. 16), 210-520 m. Flowering dates, Feb-Oct. REPRESENTATIVE SPECIMENS. Mexico: COAHUILA: Saltillo, Aug 1913, Adole 29 (F, GH, NY, US); Sabinas, 16 Sep 19957, Kenoyer 5 (F); 30 mi S of Monclova, 14 Aug 1948, Kenoyer & Crum 2594 (A); Muzquiz, Apr 1938, Marsh 1176 (F, GH, TEX); Soledad, 25 mi SW from Monclova, 9-19 Sep 1880, Palmer 556 (GH, NY, US); Rio Grande Valley near Diaz, 17 Apr 1900, Pringle 9008 (F, GH, US); 4 mi W of Nueva Rosita, 14 Aug 1967, Stuessy 902 (TEX); 21 mi S of Monclova on rte 57, 14 Aug 1967, Stuessy 912 (TEX); 23 mi N of Sabinas, 23 Aug 1959, Waterfall 15805 (F, SMU); Monclova, 25 mi SW of Sabinas, 19 Jun 1936, Wynd & Muller 214 (ARIZ, FSU, GH, NY, US). NUEVO LEÓN: Monterrey, Obispado, Jul 1911, Abloon 187 (us); 36 mi NE of Sabinas Hidalgo, 24 Mar 1944, Barkley 14575C (GH, NY); between Laredo & Monterrey, 8 Feb 1945, Bonner 55 (F); 12 mi N of Sabinas Hidalgo, 26 Mar 1944, Heard & Barkley 14542B (TEX); Cerro del Obispado, 20 Oct 1946, Lacás 58 (F); 8 mi S of Sabinas Hidalgo, 8 Feb 1964, May 13 (sMU); 12 mi W of Monterrey, 27 Feb 1944, Painter, Lucas & Barkley 14297 (TEX); 47 mi S of Nuevo Laredo, 28 Apr 1962, Reséndez 52 (SMU, TEX) ; 17 mi NE of Sabinas [Hidalgo], 20 Mar 1963, Rodriguez 72 (SMU); 26 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 855 (TEX); 38 mi N of Sabinas Hidalgo, 12 Jun 1967, Stuessy 856 (TEX). UNITED STATES. Texas: KINNEY C0., 9 mi W of Brackettville, 28 Mar 1947, McVaugh 7694 (DS, F, LL [2], SMU, TEX); MAVERICK co., 30 mi SE of Eagle Pass, 14 Mar 1964, Bruni 15 (LL, SMU, TEX) ; UVALDE CO. 2 mi N of Cline, 8 Jul 1936, Hedrick 156 (UC); VAL VERDE C0., Del Rio, 20 Apr 1930, Jones 25916 (DS, POM). 7c. Melampodium cinereum DC. var. ramosissimum (DC.) A. Gray, Smithson. Contrib. Knowledge 3, Art. 5: 103. 1850. Melampodium ramosissimum DC. Prodr. 5:518. 1836. TYPE: TEXAS[?]: between the Nueces River & Laredo, 44 Rhodora [Vol. 74 Jul 1829, J. L. Berlandier 2017 [—607] (Holotype, G-DC; isotypes, F! G! GH[2]! HAL! K! Mo[2]! w[2]! wis!; photo- graph of holotype, Ipc 800. 927: II. 1!; photograph of K isotype,US!; photograph of HAL isotype, OS! TEX!). Melampodium cinerascens S. F. Blake, Contrib. U. S. Nat. Herb. 22:605. 1924. TYPE: MEXICO: Tamaulipas, Hacienda Buena Vista, ca 20 mi E of Abasolo, 18 Jun 1919, E. O. Wooton s.n. (Holotype, US!; photograph of holotype, TEX!). Stems 0.7-0.8 mm diam, strigose with hairs 0.1-0.8 mm long. Peduncles 0.7-3.8 cm long. Leaves linear, 0.7-3.2 cm long, 0.2-0.5 em wide, with both leaf surfaces strigose with hairs 0.2-1 mm long; margin entire to pinnately 10-lobed or cleft, near base strigose with hairs up to 1 mm long. Heads 3-5 mm tall, 5-14 mm diam. Outer involucre 4.5-9 mm diam; bracts 2.7-4 mm long, 1.3-2.8 mm wide. Fruits 1.5-1.7 mm long; hood apex mucronate to cirrhous (rarely muticous), with appendage up to 1.5 mm long. Ray florets 7-8; ligules 2-3.8 mm long, 1-2.3 mm wide. Disc florets 25-35; corollas 1.8 mm diam, with throat and tube each 0.4 mm long. Paleae 2.5 mm long, 0.9 mm wide. Chromo- some number, n —10. Mesquite-grasslands of northern Tamaulipas, Mexico, and adjacent Hidalgo, Cameron and Webb Cos. of Texas (Fig. 16), 15-90 m. Flowering dates, Jun-Dec. There is some character intergradation where the three varieties of M. cinereum overlap near the Rio Grande Val- ley of southern Texas (Fig. 16). Although no detailed studies have been carried out to clarify the nature of this intergradation, hybridization is likely to be occurring. REPRESENTATIVE SPECIMENS. Mexico. TAMAULIPAS: 20 mi E of San Fernando-Santander Jiménez hwy on rd to Loreto (11 mi W of Loreto), 15 Sep 1960, Crutchfield & Johnston 5527 (TEX); 13 mi E of Abasolo turnoff on the Santander-Jiménez-Pesca rd, 15 Dec 1960, Crutchfield & Johnston 6141b (TEX); 48 mi S of Reynosa, 19 Oct 1959, Graham & Johnston 4878 (TEX); 1 mi N of San Fernando, 4 Jul 1966, Stuessy 450 (TEX); ca 2 mi SE of Reynosa, 4 Jun 1967, Stuessy 778 (TEX); 27 mi S of Reynosa, 4 Jun 1967, 1972] Melampodium — Stuessy 45 Stuessy 787 (TEX); 3 mi S of San Fernando, 18 Nov 1967, Stuessy & Renold 1261 (TEX). UNITED STATES. Texas: HIDALGO CO., Mission, Lomita Alta, 13 Jul 1937, Cameron 248 in part (TEX) ; McAllen, 10 Aug 1937, Cameron 248 in part (TEX); La Joya gravel pit, 27 Aug 1942, Walker 93 (LL, TEX); WEBB CO., 11 mi W of Encinal, 5 Jul 1960, H. & F. Iltis & Koeppen 35 (wIs). 8. Melampodium argophyllum (A. Gray ex Robins.) S. F. Blake, Contrib. U. S. Nat. Herb. 22:606. 1924. Melampodium cinereum DC. var. argophyllum A. Gray ex Robins. Proc. Amer. Acad. Arts & Sci. 36:458. 1901. (A. Gray in S. Wats. Proc. Amer. Acad. Arts & Sci. 18: 104. 1883. nom nud., but based on the type collection) TYPE: MEXICO: Coahuila Sierra Madre S of Saltillo [ca 60 km SE of Saltillo; McVaugh, 1956], Feb-Oct 1880, FE. Palmer 2068 (Holotype, GH!). Melampodium leucanthum Torr. & A. Gray var. argo- phyllum (A. Gray ex Robins.) Stuessy, Sida 3:348. 1969. Perennial subshrubs, 12-22 cm tall. Stems ascending, 0.8- 1.2 mm diam, canescent-tomentose with hairs 0.2-1 mm long. Peduncles 3.5-7.7 cm long. Leaves sessile, linear-ob- long, 1-2.8 cm long, 0.2-1.5 cm wide, at apex and base obtuse, with both surfaces tomentose with hairs 0.2-1 mm long; margin pinnately 8-lobed or parted (rarely entire), near base strigose-hispid with hairs 0.5-0.7 mm long. Heads 4-7 mm tall, 10-23 mm diam. Outer involucre cupulate, 6.2-12 mm diam; bracts 5, connate 1/2 to 3/5 their length, ovate, 3.5-6 mm long, 2.5-5 mm wide, at apex acute, with abaxial surface tomentose with hairs 0.1-1 mm long; mar- gin herbaceous. Fruits 0.8-1.9 mm long, with lateral sur- faces aculeate-verrucate; hood apex muticous to cirrhous (usually mucronate). Ray florets 8-9; ligules cream-white, oblong-elliptic, 2.5-9 mm long, 1.5-4.5 mm wide. Disc florets 25-50; corollas yellow, 1 mm diam, throat and tube each 0.8 mm long. Paleae oblong-elliptic, 2.6 mm long, 1.2 mm wide; apex yellow, with margin laciniate; midrib promi- nent, often strigillose with hairs 0.1 mm long. Chromosome number unknown. 46 Rhodora [Vol. 74 Arid mountains in western Nuevo León and adjacent Coahuila, Mexico (Fig. 16), 1830-2440 m. Flowering dates, Feb-Oct. Although M. argophyllum was first described as a variety of M. cinereum, in subsequent years the status of the former taxon has been changed twice. Morphologically M. argophyllum shows similarity to both M. leucanthum and M. cinereum but seems to have a greater total resemblance to the fcrmer species. Due to recent chromatographic stud- ies on M. argophyllum and other taxa of the white-rayed complex (Stuessy, 1971c), however, I have reversed my earlier judgment (1969a) based solely on morphological criteria which treated M. argophyllum as a variety of M. leucanthwm. I now concur with Blake's (1924) recognition of M. argophyllum as being a distinct species. Refer to my recent paper (1971c) for additional discussions on the sta- tus of M. argophyllum and on hypothetical evolutionary relationships within the entire white-rayed complex. REPRESENTATIVE SPECIMENS. Mexico. COAHUILA: G. Cepeda, Sierra de la Paila (V. Seco), 6 Jul 1944, Hinton et al. 16563 (GH, NY, US); 45 km SW of Monterrey, 28 Feb 1946, Johnson & Barkley 16250M (TEX); Castanos, “Puerto de San Lázaro," Sierra de San Lazar, 30 Aug 1939, Muller 3040 (F, GH, UC); Sierra de la Pata Solana, Mar 1905, Purpus 1003 (F, GH, NY, UC), Feb 1905, Purpus 1003a (uc); Sierra de la Paila, Oct 1910, Purpus 4730 (uc); Sierra de Parras, Mar 1905, Purpus s.n. (UC); Cañon Espantosa, W slope of Sierra de Saa Vicente, ca 20 km ESE of Cuatro Cienegas, 25 Mar 1941, Schroeder 120 (GH); 9 km S of Parras on Sierras Negras, 3 Jul 1941, Stanford, Retherford & Northcraft 186 (ARIZ, DS, F, GH, NY [2], UC); Castanos, San Lázaro, “rocky slopes of El Puerto de San Lázaro," 16 Jun 1936, Wynd & Muller 132 (ARIZ, US). NUEVO LEÓN: Las Salinas, 1924, Orcutt 1395 (us); 24 km W of Icamole, 3 Feb 1907, Safford 1264 (us). 3. Series Sericea Stuessy, ser. nov. Herbae annuae; folia lanceolata vel pinnatim divisa, sub- tus sericea vel strigosa; ligulae luteae, minus quam vel raro circa 2 mm longae; involucrum extimum marginibus her- baceis; fructi plerumque cucullati. Species 9-13. Typus: Melampodium sericeum Lag. 1972] Melampodium — Stuessy 47 9. Melampodium sericeum Lag. Gen. et Sp. Nov. 32. 1816. non H.B.K. 1820. TYPE: “NOVA HISPANIA": 1787-1804, seeds collected by M. Sessé & J. M. Mocino s.n. (Holotype, MA?; isotype, G!; photograph of G isotype, F! os! TEX! US!; photograph of G-DC isotype, IDC 800. 937: I. 8!). Melampodium hispidum H.B.K. Nov. Gen. Sp. 4:273. t. 399. 1820. TYPE: MEXICO: Michoacán, between Pazcuaro & Valladolid [Morelia], 6000 ft, Sep 1803, F. H. A. von Humboldt & A. J. Bonpland. s.n. (Holotype, P; fragments of holotype, P!). Melampodium sericeum Lag. var. brevipes A. Gray, Proc. Amer. Acad. Arts & Sci. 22: 423. 1887. TYPE: MEXICO: Guanajuato, León, W of Guanajuato, 1829, “Menzies” [probably J. Méndez] s.n. (Holotype, GH!; photograph of G-DC isotype, TEX!; tracing of probable isotype [G?], GH!). Melampodium sericeum Lag. var. exappendiculatum Robins. Proc. Amer. Acad. Arts & Sci. 36:459. 1901. TYPE: MEXICO: San Luis Potosí, “in mountains near Morales," Aug 1876, J. G. Schaffner 271 (Lectotype chosen, GH!). Annual herbs, 3-40 cm tall. Stems ascending to erect, 0.4-2.5 mm diam, hispid-pilose with hairs up to 1 mm long. Peduncles 5-22 mm long. Leaves sessile, linear-oblong to oblanceolate, 2-6.5 em long, 0.2-3 em wide, at apex acute- obtuse, at base attenuate to obtuse, with upper surface strigose with hairs 0.5 mm long, with lower surface serice- ous; margin entire to pinnately parted with 1-4 lobes, ir- regularly ciliate with hairs 0.2 mm long, at base hispid with hairs 1-1.5 mm long. Heads 4-5 mm tall, 4-8 mm diam. Outer involucre cupulate, 6-11 mm diam; bracts 5, slightly connate at base, separate to imbricate, narrowly rhombic to obovate, 3-7 mm long, 2-4 mm wide, at apex acute, with abaxial surface pilose to hispid with hairs 1-1.5 mm long; margin herbaceous. Fruits 2.5-3.2 mm long, with lateral surfaces smooth and striate to strongly verrucate; hood apex muticous to cirrhous, with tapering appendage up to 2 mm long. Ray florets 5-7; ligules yellow, narrowly ovate, 0.8-1.2(2.2) mm long, 0.6-0.8(—1.2) mm wide. Disc florets Cruz, Sep 1851, G. Thurber 937 (Lectotype, GH! K! Mo! 48 Rhodora [Vol. 74 Fig. 17. Map of Mexico and adjacent Central America showing distribution of Melampodium sericeum. (—5) 8-12; corollas yellow, 1.1 mm diam, with throat 0.6 mm and tube 0.5 mm long. Paleae oblanceolate, 3.5 mm long, 0.3 mm wide; apex yellow, with margin dentate-erose; mid- rib prominent, glabrous. Chromosome number, n = 30. Pine-oak forests of Mexico, Guatemala and El Salvador (Fig. 17), 1100-2590 m. Flowering dates, Jul-Sep. This is a widespread, variable species. Although no varieties are recognized here in M. sericeum, there are sev- eral morphologically distinct collections that should be noted. One specimen from Guatemala, Steyermark 50530, is unusually robust and has brown-purple disc florets. Also from Guatemala are several collections, Morales 1300, Rojas 86, Standley 59769 and 82849, which possess fruits with 1972] Melampodium — Stuessy 49 broad, wide-spreading hoods in contrast to the much nar- rower hoods typical of the species. REPRESENTATIVE SPECIMENS. El Salvador. SANTA ANA: near Chal- chuapa, 1922, Calderón 1019 (GH, US). Guatemala. BAJA VERAPAZ: Santa Rosa, Oct 1912, Tuerckheim 3955 (US). ESCUINTLA: 1942, Ignacio 1567 (F). GUATEMALA: Laguna, Amatitlan, 1929, Morales 1300 (F); Chilloni, 21 Jun 1921, Rojas 86 (GH, NY, US); Finca Bretafia, rd between Guatemala & Riscal, 12 Dec 1938, Standley 59769 (F, NY, SMU). HUEHUETENANGO: ca 3 mi S of Huehuetenango, 18 Jul 1960, King 3416 (NY, TEX, UC); Huehuetenango, 9 Nov 1934, Skutch 1592 (GH); between San Sebastian H. & Rio San Juan, 9 Jan 1941, Standley 82849 (F, NY, US); between Huehuetenango & San Sebastian H., 12 Aug 1942, Steyermark 50392 (F, GH, US); between San Sebastián H. & San Rafael Pétzal, 14 Aug 1942, Steyermark 50530 (F). QUICHE: 1942, Ignacio 1404 (F). Mexico. CHIHUAHUA: La Mesa Colorada, 14 Oct 1939, Gentry 550 [550M] (ARIZ, DS, F); Loreto, Río Mayo, 1 Sep 1936, Gentry 2553 (ARIZ, GH, F, UC, US); Temosachie, Cañón Huahuatán, 10 mi SE of Madera, 23 Sep 1939, Muller 3484 (GH, UC). COLIMA: Alzada, 25 Aug 1966, Stuessy 730 (TEX). DISTRITO FEDERAL: Olivar, 15 Aug 1910, Orcutt 3605 (F); above Santa Fe, 4 Sep 1901, Pringle 8609 in part (F, GH, NY, POM, UC, US [2]). DURANGO: vicinity of Durango, Iron Mt, Oct 1896, Palmer 926 (GH, US). GUANAJUATO: 20 mi NE of Irapuato, 28 Aug 1947, Barkley, Rowell & Paxson 735 (TEX); near Guanajuato, Sep [year?], Dugés 44 (GH [2]); Guanajuato, 1909, Furness s.n. (F); near Acambaro, 6 Oct 1892, Pringle 5309 (GH); 14 mi SE of León, 16 Aug 1957, Waterfall & Wallis 13886 (F, SMU). GUERRERO: Petaquillas, 27 Aug 1965, Stuessy 364 (TEX). JALISCO: 56 mi SE of Guadalajara, 21 Aug 1953, W. & M. Manning 531212 (GH, TEX); ca 11 mi SE of Lagos de Moreno, 7 Sep 1952, McVaugh 12822 (SMU, US); Guadalajara, Jul 1886, Palmer 260 (GH, US); near Guadalajara, 24 Aug 1901, Rose & Hay 6275 (GH, US); 2 mi NW of Tequila, 26 Aug 1966, Stuessy 740 (TEX). MEXICO: Temascaltepec, Plaza de Gallos, 22 Jul 1934, Hinton et al. 6324 (GH, NY, US). MICHOACÁN: ca 10 mi W of Morelia, 5 Aug 1960, King 3626 (Ds, NY, TEX, UC, US) ; Cerro Potrerillos, ca 5 mi N of Cotija, 5-9 Oct 1961, King & Soder- strom 4584 (NY, SMU, TEX, UC, US) ; 0.8 mi NW of Zitacuaro, 1 Sep 1965, Stuessy 377 (TEX); 7 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 690 (TEX); 11 mi NW of Zamora, 23 Aug 1966, Stuessy 703 (TEX). MORELOS: Cuernavaca, 31 Aug 1910, Orcutt 3887 (F). NAYARIT: 5 mi SE of Ixtlán del Río, 16 Aug 1961, Waterfall 16342 (SMU, UC). OAXACA: 53 mi S of Tehuacán, 13 Aug 1961, Powell & Edmondson 660 (F, TEX); Valley of Oaxaca, 5 Jul 1897, Pringle 6728 (F, GH, NY, UC, US [2]) ; 38 mi S of Zimatlán, 12 Aug 1966, 50 Rhodora [Vol. 74 Stuessy 642 (TEX); 4 mi S of Sola de Vega, 13 Aug 1966, Stuessy 647 (TEX); % mi NW of Las Sedas, 15 Aug 1966, Stuessy 660 (TEX). PUEBLA: Los Molinos between Puebla & Atlixco, 18 Sep 1944, Sharp 44961 (NY). QUERÉTARO: Jul 1904, Kuntze 27465 (Ny [2]); 6 mi W of Querétaro, 6 Aug 1961, Powell & Edmondson 579 (F, TEX); 4.7 mi N of Querétaro, 8 Aug 1959, Rock M-442 (TEX). SAN LUIS PoTOsÍ: 15-16 mi E of Ciudad del Maiz, 20 Sep 1960, Crutchfield & Johnston 5664 (TEX); 3 mi E of Ciudad Maiz, 2 Sep 1948, Kenoyer & Crum 4093 (GH); near San Luis Potosí, 1877, Schaffner 270 in part (NY). SONORA: ca 2 mi W of Tres Ríos (ca 88 mi W of Vieja Casa Grandes, Chihuahua), 29 Aug 1952, Tucker 2565 (ARIZ, US); Cañón de Huépari, N of Aribabi, 2-3 Sep 1939, White 2642 in part (Gi). ZACATECAS: between Colotlán & Plateado, 31 Aug 1897, Rose 3617 (F, US). 10. Melampodium pringlei Robins. Proc. Amer. Acad. Arts & Sci. 36:461. 1901. TYPE: MEXICO: Oaxaca, Las Sedas, 1850 m, 15 Sep 1894, C. G. Pringle 5722 (Holotype, GH!). Annual herbs, 30 em tall. Stems ascending, 1-2 mm diam, hirsute-hispid with hairs up to 1 mm long. Peduncles 1-2 mm long. Leaves sessile, linear-oblong to oblanceolate, 2- 2.8 em long, 0.7-1.6 em wide, at apex acute-obtuse, at base attenuate to obtuse, with upper surface strigose with hairs 1 mm long, with lower surface copiously sericeous; margin entire to 2-lobed, irregularly ciliate with hairs 0.3 mm long, near base hirsute with hairs 1-2 mm long. Heads 3-4 mm tall, 3.5-4 mm diam. Outer involucre cupulate, 3-4 mm diam; bracts 5, slightly connate at base, separate, lanceo- late, 3-3.5 mm long, 1 mm wide, at apex acute-acuminate, with abaxial surface hirsute with hairs 1 mm long; margin herbaceous. Fruits 3 mm long, with lateral surfaces 3- ribbed with large tubercles. Ray florets 3-4; ligules yellow, with undersurface near apex light purple, ovate, 1 mm long, 0.5 mm wide. Disc florets 2-3; corollas yellow, 1.1 mm diam, with throat 0.6 mm and tube 0.5 mm long. Paleae oblanceolate, 2.5 mm long, 0.8 mm wide; apex purple, with margin erose-dentate; midrib prominent, weakly pilose with hairs 0.3 mm long. Chromosome number unknown. Known only from the type collection from pine-oak for- ests near Las Sedas, Oaxaca, Mexico (Fig. 11), 1860 m. Flowering date, Sep. 1972] Melampodium — Stuessy 51 Although this taxon is very similar to M. sericeum, it seems sufficiently distinct to be maintained as a separate species at the present time. Additional collections, however, may indicate that varietal status is preferable. I have searched the countryside around Las Sedas, the type lo- cality, but have been unsuccessful in rediscovering the taxon. Characters (in addition to those listed in the key) which distinguish M. pringlei from its closeset relative are: M. sericeum M. pringlei 1. Heads 4-5 mm tall, 4-8 1. Heads 3-4 mm tall, mm diam. 3.5-4 mm diam. 2. Outer involucral bracts 2. Outer involucral bracts narrowly rhombic to lanceolate, 3-3.5 mm obovate, 3-7 mm long, long, 1 mm wide. 2-4 mm wide. 3. Ray florets 5-7. 3. Ray florets 3-4. 4. Paleae yellow-tipped. 4. Paleae purple-tipped. 11. Melampodium strigosum Stuessy, sp. nov. Melampodium coronopifolium Sch. Bip. ex Hemsl. Biol. Centr. Am. Bot. 2:145. 1881. nom. nud. Based on several specimens including C. C. Parry & E. Palmer 444 1/2 (GH! K! MO! NY! US!). Herbae annuae, 3-35 cm altae. Caules ascendentes, 0.6-2 mm diametro, moderate hispido-strigillosi, pilis 0.3-1 mm longis. Pedunculi 0-3(-11) mm longi. Folia sessilia, ob- longo-linearia vel oblanceolata, 0.6-5 cm longa, 0.2-1.5 cm lata, apice acuto-obtusa, basi obtuso-subauriculata, utrinque strigosa pilis 0.2-0.6 mm longis, marginibus integris vel pinnatim 2-4 lobatis, basalibus strigoso-hispidis, pilis 0.3- 1.1 mm longis. Capitula 3-4 mm alta, 4-8 mm diametro. Involucrum extimum cupulatum, 3-8 mm diametro; bracteae 9, basi leviter connatae, separatae, lanceolato-ellipticae, 2.7- 6 mm longae, 1-2.9 mm latae, apice acutae, extus strigoso- hispidae in nervis principalibus, pilis 0.3-0.8 mm longis, marginibus herbaceis. Fructus 2.2-3 mm longi, lateribus striatis vel verrucatis, cucullis apice muticis vel cirrhosis, 52 Rhodora [Vol. 74 Fig. 18. Map of Mexico and adjacent United States showing dis- tribution of Melampodium strigosum (dots), M. sinaloense (square), and M. tepicense (triangles). One collection of M. strigosum from Colorado not shown. 1972] Melampodium — Stuessy 53 mucrones 2-2.5 mm longis. Flosculi radii 5-8; ligulae flavae, oblongo-ovatae, 0.6-1.1 mm longae, 0.5-0.8 mm latae. Flos- culi disci 4-6; corollae flavae, 1-2 mm diametro, faucibus 0.6 et tubis 0.5 mm longis. Paleae oblongo-ellipticae, 2.3 mm longae, 0.8 mm latae, apice flavae, marginibus dentatis, cos- tis conspicuis, glabris vel puberulis pilis 0.1 mm longis. Chromosomatum numerus, n = 20. TYPUS: MEXICO: Chihuahua, ca 5 mi E of Ciudad Guer- rero on rte 16, 21 Aug 1967, T. F. Stuessy 1054 (Holotype, US!; isotypes, F! GH! MICH! NY! os! SMU! TEX! UC!). Pine-oak forests of central and northwestern Mexico, reaching Cochise, Pima and Santa Cruz Cos. of southeast- ern Arizona, Jeff Davis Co. of Texas, and Chaffee Co. of Colorado (Fig. 18), 1310-2590 m. Flowering dates, Aug- Oct. This new species is a common taxon that has masquer- aded under the name of M. hispidum H.B.K. for many years. Dr. Rogers McVaugh first mentioned to me that the holotype of M. hispidum in the herbarium at Paris seemed conspecific with the even more common species, M. sericeum Lag. Having obtained additional information from Dr. Alicia Lourteig at Paris regarding the holotype of M. his- pidum, I believe it is definitely conspecific with M. sericeum. The placement of M. hispidum in synonymy with M. sericeum leaves M. coronopifolium Sch. Bip. ex Hemsl. as the only other name applicable to this taxon, but this latter epithet is a nomen nudum and therefore not validly pub- lished. By reference to three specimens cited in the proto- logue as being representative, however, M. coronopifolium is clearly referable to this species. I could now validate this available epithet, but I prefer to name this undescribed Species with an epithet that emphasizes the distinctive strigose pubescence of the leaves in contrast to the mark- edly sericeous leaves of the morphologically similar M. sericeum. REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: Parral, 19 Sep 1898, Goldman 110 (NY, Us); San Diego Canyon, Sierra Madre Mts, 16 Sep 1903, Jones s.n. (POM); Sta Clara Mts, 10-19 Oct 1935, 54 Rhodora [Vol]. 74 LeSueur 341 (ARIZ, F, GH, TEX); near Chihuahua, Sep-Oct 188», Pringle 297 (F, GH, NY, US [2]), 1 Jun 1885, Pringle 10 (F, GH, K, NY [2], US); mts near Chihuahua, Sep 1886, Pringle 754 (F, NY [2], uc [2], us [2]); near Chihuahua, 12 Sep 1886, Pringle 1045 (ps, NY, UC); ca 2-3 mi W of Parral, 18 Aug 1967, Stuessy 1016 (TEX); 2 mi S of Cuauhtémoc, 20 Aug 1967, Stuessy 1038 (TEX) ; near Colonia García in Sierra Madre Mts, 26 Sep 1899, Townsend & Barber 351 (GH, NY, US). DISTRITO FEDERAL: Pyramid of Cuicuilco, Tlalpan, 1 Sep 1936, MacDaniels 747 (F); Valley of Mexico, 9 Sep 1896, Pringle 6491 (F, GH, NY, UC, US); above Santa Fe, Valley of Mexico, 4 Sep 1901, Pringle 8609 in part (GH); near Club Golf de Chapultepec, 4 Sep 1946, Zamora, Paxon & Barkley 16M905 (TEX). DURANGO: vicinity of Durango, Apr-Nov 1896, Palmer 486 (F, GH, NY, UC, US). GUANAJUATO: Guanajuato, Aug 1899, Dugès s.n. (US) ; Guanajuato, 1909, Furness s.n. (F). HIDALGO: near El Salto, 17 Sep 1901, Pringle 9331 (GH, NY, US); near Landa Station, 3 Sep 1903, Pringle 11548 (ARIZ, F, GH, SMU, US). QUERÉTARO: 22 mi NE of San Juan del Río, 10 Sep 1962, Turner & Powell 1116 (TEX) ; 14 mi SE of San Juan del Río, 17 Aug 1957, Waterfall & Wallis 13960 (SMU, US). SONORA: Cañón de Huépari N of Aribabi, 2-3 Sep 1939, White 2642 in part (ARIZ), 7 Sep 1939, White 2788 (ps). UNITED STATES. Arizona: COCHISE CO. Chiricahua Mts, 18 Sep 1907, Blumer 1665 (ARIZ, F, GH, NY, US); Portal, Chiricahua Natl Forest, 17-19 Sep 1914, Eggleston 10729 (us); Lanner Canyon, Huachuca Mts, 24 Aug 1910, Goodding 814 (ARIZ, GH, US); Huachuca Mts, 20 Aug 1893, Holzner 1962 (vs); Chiricahua Mts, 22 Sep 1931, Jones s.n. (POM); Huachuca Mts, near Ft Huachuca, Sep 1882, Lemmon 2777 (F, GH, K, NY, UC); PIMA CO., Rincon Mts, Dec 1907, Goodding 18 (ARIZ); SANTA CRUZ CO., Nogales, 25 Oct 1926, Jones 22681 (pom); Patagonia Mts, 18 Aug 1928, Kearney 5587A (us); Nogales, 7 Aug 1927, Peebles, Harrison & Kearney 4614 (us). Colorado: CHAFFEE CO., Buena Vista, Jones s.n. (POM). Texas: JEFF DAVIS co., Davis Mts, Madera Canyon, Aug 1936, Hinckley 620 (NY), 8 Oct 1936, Hinckley 881 (ARIZ, F, GH, LL, NY [2], SMU); Davis Mts, Madera Creek, old Fisher Ranch House, 16 Sep 1944, Hinckley 3293 (LL, US); Madera Canyon, Mt Livermore, Aug 1936, Hinckley sm. (TEX); Madera Canyon, Davis Mts, 25 Aug 1967, Stuessy 1128 (TEX); 11 Sep 1959, Turner 4620 (SMU, TEX) ; N of Merrill ranch on rd to Mt Locke, 13 Aug 1950, Warnock 9229 (SMU) ; upper Madera Canyon, Davis Mts, 5 Oct 1955, Warnock 13507 (LL) ; Davis Mts, 18 Sep 1918, Young s.n. (TEX). 12. Melampodium longicorne A. Gray, Mem. Amer. Acad. ser. 2. 5:321. 1855. TYPE: MEXICO: Sonora, near Santa 1972] Melampodium — Stuessy 55 Cruz, Sep 1851, G. Thurber 937 (Lectotype, GH!; isotypes, GH! K! MO! NY!; photograph of K isotype, USE Annual herbs, 12-60 cm tall. Stems erect, 0.9-2.8 mm diam, subglabrous to hispid-strigose with hairs 0.5-1 mm long. Peduncles 4-30 mm long. Leaves sessile, linear-ob- long to oblanceolate, 3-5 em long, 0.3-1.2 cm wide, at apex obtuse, at base truncate-subauriculate, with both surfaces strigose with hairs 0.3-0.8 mm long; margin entire to ob- scurely dentate, at base hispid-hispidulous with hairs 0.5-1 mm long. Heads 4-7 mm tall, 7-13 mm diam. Outer in- volucre cupulate, 6-8(—10) mm diam, bracts 5, slightly con- nate at base, usually imbricate 2/3 their length (less often separate), elliptic to obovate-rhombic, 4.3-6.8 mm long, 2-4 mm wide, at apex acute, with abaxial surface moderately strigose with hairs 0.5-1 mm long; margin herbaceous. Fruits 5-3.6 mm long, with lateral surfaces granulate to aculeate-verrucate; hood apex cirrhous, with appendage over 4 mm long. Ray florets 7-12; ligules yellow, oblong- elliptic, 1.2-1.5 mm long, 0.7-0.9 mm wide. Disc florets 8-10; corollas yellow, 1 mm diam, with throat 0.5 mm and tube 0.4 mm long. Paleae oblong-elliptic, 3.4 mm long, 1.1 mm wide; apex yellow, with margin bidentate; midrib promi- nent, strigillose with hairs 0.1 mm long. Chromosome num- ber, n = 30. Mountains of southeastern Arizona in Cochise, Pima and Santa Cruz Cos., and in adjacent Sonora, Mexico (Fig. 19), 1130-1830 m. Flowering date, Sep. Found exclusively in southeastern Arizona and neighbor- ing Sonora, this species is often conspicuous with its long coiled awns on the hoods of the ray achenes. Just as often, however, the awns are absent, making separation from the morphologically similar M. strigosum difficult. In the area of sympatry in Arizona, there is a trend in M. strigosum toward longer peduncles, more entire leaves, and more erect stems, suggesting that hybridization may be occurring be- tween the two taxa. Asa Gray seems to have experienced some difficulty in delimiting M. longicorne, even though he himself described 56 Rhodora [Vol. 74 Fig. 19. Map of northwestern Mexico and adjacent United States showing distribution of Melampodium appendiculatum (squares), M. cupulatum (triangles), M. longicorne (circles), and M. sinuatum (dots). it as new. He first (1855) confused a Wright 1205 collec- tion (distinct M. longicorne) with M. strigosum and later (1859) emended his original description to include a Schott 72 collection, the latter being treated subsequently as be- longing to M. appendiculatum (Robinson, 1901; present treatment). 1972] Melampodium — Stuessy 57 Confusion has existed regarding the correct spelling of the specific epithet, and some workers (Robinson, 1901; Tidestrom and Kittell, 1941; Gentry, 1942) have elected to use the form longicornu rather than longicorne as originally published. Even Gray himself ceased to use the original adjectival spelling and reverted to the substantive form longicornu in his subsequent publications of 1859 and 1884. However, as longicorne is the original correct spelling, I have used this form in the present treatment, as have other workers in recent times (Kearney and Peebles, 1942, 1964; Shreve and Wiggins, 1964). REPRESENTATIVE SPECIMENS. Mexico. SONORA: 2 mi E of Agua Prieta-Nacozari rd to Oputo & Angostura, 7 Sep 1961, Turner, Dodge & Mason 2068 (ARIZ, DS); near Santa Cruz, 1851, Wright 1205 (G, GH [3], K, NY, US). UNITED STATES. Arizona: COCHISE CO., 10 mi N of Chiricahua Station, 8 Sep 1942, Barneby 5135 (NY); Chiri- cahua Mts, N of Wilgus Ranch, 2 Sep 1907, Blumer 2128 (F, GH, NY); Ft Huachuca Military Reservation, 29 Sep 1962, Goodding 267-62 (ARIZ); Huachuca Game Preserve, Garden Canyon, 21 Sep 1949, Goodding 580-49 (ARIZ [2]) ; Montezuma Canyon, Huachuca Mts, 17 Oct 1958, Goodding 699-58 (ARIZ, UC); Ramsey Canyon, Huachuca Mts, 29 Sep 1929, Jones 25045 (Ny [2], POM, TEx [2], vc [2]); Huachuca Mts, 3 Sep 1903, Jones s.n. (NY, POM); near Apache Pass, Chiricahua Mts, Sep 1881, Lemmon 331 (F, Uc [2]); Huachuca Mts, Aug 1882, Lemmon 2795 (F, G, NY, US); 4% mi S of Sonoita & Parker Canyon rds, 10 Oct 1965, Reese 94 (ARIZ); Bear Canyon, Huachuca Mts, 28 Aug 1936, Shreve 7709 (ARIZ); near Ft Huachuca, Aug 1894, Wilcox 335 (us); Ft. Huachuca, Aug 1893, Wilcox s.n. (NY); PIMA CO, Oak Tree Canyon, Thurber Ranch, Santa Rita Mts, 14 Oct 1940, Benson 10598 (ARIZ); Madera Canyon, Santa Rita Mts, 6 Sep 1944, Clark 12377 (ARIZ); 17 mi S of Tucson- Benson hwy on rte 83, 21 Oct 1944, Gould 2883 (ARIZ); Thurber Ranch, Santa Rita Mts, 14 Sep 1945, Gould & Haskell 3268 (ARIZ, DS, GH, NY, UC, US); Santa Rita Mts, 24 Aug 1903, Jomes s.n. (DS, POM); Madera Canyon, Santa Rita Mts, 24 Aug 1926, Loomis & King 2890 (ARIZ); Santa Rita Mts, 16 Sep 1884, Pringle 55 (GH, SMU); Greaterville, 16 Sep 1916, Shreve 4965 (ARIZ) ; Empire Ranch, 26 Aug 1936, Shreve 7685 (ARIZ, SMU); Santa Rita Mts, Rozemont, 20 Sep 1915, Thornber 7404 (ARIZ), 10 Sep 1915, Thornber 8121 (ARIZ), 20 Sep 1915, Thornber 9050 (ARIZ); SANTA CRUZ CO., Syca- more Creek, ca 28 mi NW of Nogales, 20 Sep 1964, Cronquist 10003 (NY); Sycamore Canyon, near Ruby, 30 Sep 1944, Darrow & Haskell 2012 (ARIZ, NY, UC, US); near Canoa Tank, Summit Motorway, on 58 Rhodora [Vol. 74 Nogales-Ruby rd, 14 Oct 1944, Darrow & Haskell 2256 (ARIZ); Patagonia, 14 Sep 1959, Goodding 248-59 (ARIZ); Sonoita Creek, 30-31 Aug [year?], Harrison 8181 (F); Nogales, 30 Aug 1931, Harrison & Fulton 8153 (ARIZ, Us); Sonoita, 2 Sep 1928, Harrison & Kearney 5702 (us); Patagonia Mts, 15 Sep 1934, Kearney & Peebles 10052 (uc); Sycamore Canyon, SW of Atascosa Mts & SE of Ruby, 3 Oct 1951, Parker 7707 (ARIZ, F, RSA, UC, US); Patagonia Mts (Nogales), 28 Aug 1927, Peebles & Harrison 4680 (ARIZ, F); Harshaw, 22 Aug 1932, Shreve 6003 (ARIZ, F). 13. Melampodium nayaritense Stuessy, Brittonia 22:113. f. 5. 1970. TYPE: MEXICO: Nayarit, Canon de Jesús María, 1000 ft, 29 Aug 1905, P. Goldsmith 133 (Holotype, NY!; isotypes, F! GH! MO! Uc! US!). Annual herbs, 10-30 cm tall. Stems ascending to erect, 1-2 mm diam, hirtellous with hairs 0.5 mm long. Peduncles 1.5-3.6 em long. Leaves sessile, linear and entire to deeply pinnately divided, 3-4.5 cm long, 0.2-3.7 em wide, at apex acute, at base attenuate to obtuse, with upper surface finely strigose with hairs 1 mm long, with lower surface sericeous ; margin at base pilose-hispid with hairs up to 1.5 mm long. Heads 5-6 mm tall, 8-11 mm diam. Outer involucre cupu- late, 7-10 mm diam; bracts 5, slightly connate at base, sep- arate, obovate-rhombie, 1.5-5.5 mm long, 3-3.5 mm wide, at apex acute-acuminate, with abaxial surface pilose with hairs 1 mm long; margin herbaceous. Fruits 3 mm long, with lateral surfaces ribbed and smooth to markedly tuber- culate; hood apex cirrhous, with tapering appendage up to 2 mm long. Ray florets 5-6; ligules yellow, ovate-orbicu- late, 2 mm long, 2 mm wide. Disc florets 15-25; corollas yellow, 1.2 mm diam, with throat 0.8 mm and tube 0.6 mm long. Paleae broadly oblanceolate, 3.5 mm long, 1.6 mm wide; apex yellow, with margin entire and undulate; mid- rib prominent, weakly pubescent with hairs 0.3 mm long. Chromosome number unknown. Known from woodland hills in northeastern Nayarit, and adjacent Durango, Mexico (Fig. 20), 180-300 m. Flowering dates, Aug-Sep. *In one head of an isotype collection (UC) are found both smooth and roughly tuberculate fruits. 1972] Melampodium — Stuessy 59 Fig. 20. Map of Mexico showing distribution of Melampodium bibracteatum (dots), M. nayaritense (squares), M. repens (circles), M. rosei (closed triangles), and M. tenellum (open triangles). One collection of M. bibracteatum from Guatemala not shown. REPRESENTATIVE SPECIMENS. Mexico. DURANGO: Huasemota, 14 Aug 1897, Rose 3476 (US). NAYARIT: Valley of Río Jesús María, near village of Jesus María, 17 Sep 1960, Feddema 1209 (MICH). 4. Series Cupulata Stuessy, ser. nov. Herbae annuae vel perennae (M. sinuatwm); folia an- guste ovata (saepe lobata), subtus strigosa; ligulae luteae vel flavo-aurantiacae, plus quam vel raro circa 2 mm longae; involucrum extimum marginibus scariosis; fructi plerum- que eucullati. Species 14-19. Typus: Melampodium cupula- tum A. Gray. 60 Rhodora [Vol. 74 14. Melampodium cupulatum A. Gray, Proc. Amer. Acad. Arts & Sci. 8:291. 1870. TYPE: MEXICO: Sonora, Rio Yaqui [27937' N, 110°38’ W; McVaugh, 1956], Nov-Dec 1869, FE. Palmer 20 (Holotype, GH!; isotypes, NY! US[2]!; photo- graph of US isotype, TEX!). Eclipta pusilla M. E. Jones, Contrib. West. Bot. 18:70. 1933. TYPE: MEXICO: Baja California, Loreto, Arroyo Undo Ranch, 26 Oct 1930, M. E. Jones 27739 (Holotype, POM!). Annual herbs, 10-35 cm tall. Stems erect, 1-4 mm diam, moderately puberulent to hispidulous with hairs 0.1-0.3 mm long. Peduncles 3.5-8 em long. Leaves sessile or with short petioles 2-3 mm long, ovate-lanceolate, 3.3-7.5 cm long, 0.8- 2.1 em wide, at apex acute, at base attenuate, with both surfaces strigillose with hairs 0.2-0.8 mm long; margin en- tire to broadly crenate or sinuate on upper 3/4 of blade. Heads 7-8 mm tall, 9-17 mm diam. Outer involucre cupu- late, 6.5-10 mm diam; bracts 5, connate 1/2 their length, ovate to narrowly ovate, 4-6 mm long, 2.7-3.3 mm wide, at apex acute-acuminate, with abaxial surface strigillose with hairs 0.2-0.8 mm long; margin scarious. Fruits 3-3.4 mm long, with lateral surfaces with small tubercles and reticula- tions." Ray florets 8-12; ligules yellow-orange, oblong-el- liptie, 3-6 mm long, 1-3 mm wide. Disc florets 40-60; corollas yellow-orange, 2 mm diam, with throat 0.7 mm and tube 0.3 mm long. Paleae oblanceolate, 3.5 mm long, 1 mm wide; apex yellow, with margin erose; midrib prominent, puberulous with hairs 0.1 mm long. Chromosome number, n — 10. Arid regions of Baja California, Sinaloa, Sonora, and Nayarit, Mexico (Fig. 19), 30-790 m. Flowering dates, Sep- Mar. REPRESENTATIVE SPECIMENS. Mexico. BAJA CALIFORNIA: vicinity of La Cumbre de Alta Gracia, 1 Nov 1964, Carter 4880 (Ds, UC); Mesa del Potrero de San Javier, 19 Sep 1965, Carter 4964 (Ds, UC); Cerro Gabilán, S of Portezuelo de Gabilán, 30 Sep 1965, Carter 5055 (UC); "Although hoods are usually rudimentary in this species, a few coiled awns on small hoods are found in Feddema 1219 from Nayarit. 1972] Melampodium — Stuessy 61 NE end of Valle de Los Encinos (S side of Cerro Giganta), 28 Sep 1967, Carter & Moran 5314 (uc); 2 mi E of Comondü on rd to Loreto, 28 Aug 1955, Chambers 965 (pns, UC); N of Comondú, 3 Oct 1941, Hammerly 182 (ns, POM); 8 mi S of Querétaro, 29 Nov 1946, Wiggins 11507 (Ds, GH, UC, US). NAYARIT: near Jesús Maria, 18 Sep 1960, Feddema 1219 (MICH). SINALOA: 5 mi W of Culiacán on rd to Altata, 28 Jan 1964, Flyr 112 (TEX); Cerro Tecomate, W of Pericos, 28 Feb 1940, Gentry 5750 (ARIZ, DS, GH, NY, UC); Maratón, 12 mi W of Culiacán, 9 Mar 1944, Gentry 7005 (DS, F, GH, NY, UC, US); near Culiacán, Oct [year?], Schaffner s.n. (GH). SONORA: near rr station, Masiaca, 13 Sep 1930, Abrams 12811 (Ds); Ciudad Obregón, 29 Sep 1933, Gentry 290 (Ds); Alamos, 26 Mar-8 Apr 1890, Palmer 726 (GH); 4 mi S of La Puerca, SW of Hermosillo toward Tastiota, 4 Sep 1941, Wiggins & Rollins 281 (DS, GH, NY, UC, US). 15. Melampodium appendiculatum Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE: MEXICO: Chihua- hua, near Frailes [ca 12 km NE of Guachochic; McVaugh, 1956], ca 7400 ft, Oct 1885, E. Palmer 245 (Holotype, 3H !; isotype, US!; photograph of US isotype, TEX!). Melampodium appendiculatum Robins. var. leiocarpum Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE: MEXICO: Sonora, Alamos, 16-30 Sep 1890, E. Palmer '126 (not 726, 26 Mar-8 Apr 1890) (Holotype, GH!; isotypes, NY! US!). Melampodium appendiculatum Robins. var. sonorense Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE: MEXICO: Sonora, Cochuto, 5100 ft, 2 Oct 1890, C. V. Hart- man 71 (Holotype, GH!). Annual herbs, 10-40 em tall. Stems erect, 1-3 mm diam, subglabrous at base to hispid or pilose above with hairs 0.5-1.5 mm long. Peduncles 1-8 cm long. Leaves sessile, linear to lanceolate, 2.5-8.3 cm long, 0.4-1.8 em wide, at apex acute, at base subauriculate, with both surfaces stri- gose-hispid with hairs 1-1.5 mm long; margin entire to obsoletely crenate, at base hispid with hairs up to 2 mm long. Heads 5-8 mm tall, 7-17 mm diam. Outer involucre strongly cupulate, 4.5-8 mm diam; bracts 5, connate 2/3 their length, ovate, 3.5-4.5 mm long, 2-3 mm wide, at apex acute, with abaxial surface pubescent-pilose with hairs 0.5 62 Rhodora [Vol. 74 mm long (longer near base) ; margin scarious. Fruits 2.3- 2.8 mm long, with lateral surfaces smooth and glabrous to very tuberculate and striate; hood apex cirrhous (rarely muticous or mucronate) ?, with appendage up to 4 mm long. Ray florets 8-13; ligules yellow-orange, oblong-elliptic, 2-8 mm long, 1-2.9 mm wide. Disc florets 25-40; corollas yellow, 1.3 mm diam, with throat 0.7 mm and tube 0.4 mm long. Paleae oblancolate, 3.2 mm long, 1 mm wide; apex yellow, with margin dentate-laciniate; midrib prominent, puberu- lent with hairs 0.1 mm long. Chromosome number un- known. Pine-oak forests in mountains of Sonora, Chihuahua and northwestern Durango, Mexico (Fig. 19), 790-1390 m. Flowering dates, Jul-Nov. Robinson (1901) described two new varieties within this species based on inner involucral bract differences. From an examination of specimens that indicate intra- and inter- populational variability, however, it seems clear that these described varieties represent simply forms within the mor- phological range of one species. REPRESENTATIVE SPECIMENS. Mexico. CHIHUAHUA: Guasaremos, Río Mayo, 5 Aug 1935, Gentry 1570 (ARIZ, F, GH, NY, UC, US), 3 Aug 1936, Gentry 2353 (ARIZ, F, GH); Rio Aros, 23 Jul 1937, LeSueur 1466 (TEX), 28 Jul 1937, LeSueur 1472 (F); Sierra Madre Mts near Seven Star Mine, 27 Aug 1899, Townsend & Barber 376 (F, GH, NY, POM, UC, US [2]). DURANGO?: Lodiego [ca 25° N, 106°45' W; Mc- Vaugh, 1956], 9-15 Oct 1891, Palmer 1609 in part (F, GH, NY, US). SONORA: 20 mi NE of Ures, 16 Nov 1939, Drouet, Richards & Lock- hart 3609 (F); Baviácora, W of Rio de Sonora, 17 Nov 1989, Drouet, Richards & Lockhart 3624 (F); Alamos, Quirocoba, 13 Nov 1933, Gentry 790M [790] (ARIZ, DS); San Bernardo, Río Mayo, 14 Jul 1935, Gentry 1492 (F, Gu) ; 6 mi S of Nogales, 7 Aug 1965, R. & J. Matthews 480 (TEX); San Bernardino Ranch, 22 Aug 1893, Mearns 1997 (US); ca 12 mi from Cananea at km 20 on rd to Bacanuchi, 25 Aug 1963, Moreno MS-198 (ARIZ); Nácori Chico, Pie de la Cuesta, 6 Oct 1939, Muller 3659 (F, UC) ; Aduana, 10 Aug 1930, Russell & Souviron 2 (Us); Sta Magdalena, Nov 1855, Schott 72 (F, GH, NY), "In the collection Shreve 6758 (US), most flowering heads have fruits with long cirrhous appendages, but in one head all the achenes are muticous or only very slightly mucronate, 1972] Melampodium — Stuessy 63 Schott s.n. (F); 21 mi SE of Moctezuma, 25 Sep 1934, Shreve 6758 (ARIZ, US); Arispe, 18 Aug 1958, Turner 192 & Lowe 2074 (ARIZ); 91 mi S of Nogales along Rio de los Alisos, 8 Sep 1934, Wiggins 7021 (Ds, TEX [2], US); ca 5 mi S of Babiácora along Sonora River, 21 Sep 1934, Wiggins 7388 (ARIZ, ps, TEX [2], US); 21 mi S of Moctezuma, 25 Sep 1934, Wiggins 7444 (ARIZ, DS, TEX, US); 8 mi S of Nogales & 2 mi W of rr, 3 Oct 1934, Wiggins 7533 (Ds, Us); 8 mi from Matape toward Batuc, 9 Sep 1941, Wiggins & Rollins 433 (ARIZ, DS, GH, NY); Puerto de Huépari, NW of Aribabi, 7 Sep 1939, White 2784 (ARIZ, GH, US); Cañón de la Gallina, 25 Aug 1940, White 3516 (ARIZ, GH); El Rancho de la Nacha, 25 mi W of La Angostura, 14-20 Aug 1941, White 3908 (GH); Cañón de la Palomita, N of El Tigre, 29 Aug 1941, White 4153 (GH). 16. Melampodium sinuatum Brandg. Proc. Calif. Acad. Sci. ser. 2. 3:144. 1891. TYPE: MEXICO: Baja California, San José del Cabo, 16 Sep 1890, T. S. Brandegee 302 (not 302, Sep 1891; not 302, 10 Mar 1892) (Lectotype chosen, UC!). Suffruticose perennials, 8-35 cm tall. Stems erect, 1-2 mm diam, puberulent-canescent with hairs 0.2-0.5 mm leng. Peduncles 3-9 em long. Leaves sessile, linear to elliptic, 2-4.5 cm long, 0.5-1.6 cm wide, at apex obtuse, at base subauriculate, with both surfaces densely strigillose with hairs 0.2-0.3 mm long; margin sinuate-undulate (rarely repand) to irregularly lobed. Heads 6-7 mm tall, 10-20 mm diam. Outer involucre cupulate, 6-9.5 mm diam; bracts 5, connate 3/4 their length, orbiculate, 3-4.5 mm long, 2.5-4.5 mm wide, at apex obtuse, with abaxial surface strigillose- tomentose with hairs 0.4-0.7 mm long; margin slightly scarious. Fruits 2.9-3.2 mm long, with lateral surfaces tuberculate-striate or reticulate. Ray florets 9-13; ligules yellow-orange, elliptic-linear, 4-9 mm long, 1.5-3 mm wide. Disc florets 50-80; corollas yellow-orange, 1.3 mm diam, with throat 1 mm and tube 0.5 mm long. Paleae broadly oblanceolate, 3.5 mm long, 0.8 mm wide; apex yellow, with margin erose-laciniate; midrib prominent, puberulent with hairs 0.1 mm long. Chromosome number unknown. Low granitic hills near San José del Cabo, Baja Califor- nia, Mexico (Fig. 19), ca 60 m. Flowering dates, Sep-Mar. 64 Rhodora [Vol. 74 M. sinuatum is one of the three species in the genus, along with M. cupulatum and M. divaricatum that are found in Baja California. The distributional and morpho- logical similarity of the former two taxa, and the restricted distribution of M. sinuatum at the tip of Baja California suggest that this species may have been derived from a mainland ancestral population of M. cupulatum. REPRESENTATIVE SPECIMENS. Mexico. BAJA CALIFORNIA: San José del Cabo, Sep 1891, Brandegee 302 (GH, US); Sierra de la Trinidad, Nov 1902, Brandegee s.n. (vc [2], US); 12.2 km SW José del Cabo, 16 Dec 1947, Carter, Alexander & Kellogg 2216 (DS, UC, us); San José del Cabo, Mar 1901, Purpus 416 (uc, US) ; b mi W of San José del Cabo, 17 Dec 1958, Wiggins 14377 (DS, GH, TEX, UC). 17. Melampodium rosei Robins. Proc. Amer. Acad. Arts & Sci. 36:461. 1901. TYPE: MEXICO: Sinaloa, between Rosa- rio & Concepcion, 28 Jul 1897, J. N. Rose 3271 (Holotype, us!; photograph of holotype, TEX!). Melampodium arenicola Robins. Proc. Amer. Acad. Arts & Sci. 36:457. 1901. TYPE: MEXICO: Sinaloa, Mazatlan, Isla Piedra, 31 Dec 1894, F. H. Lamb 361a (Holotype, GH!; iso- types, G[2] ! Mo! NY! U$!; photograph of US isotype, TEX !). Melampodium rosei Robins. var. subintegrum Robins. Proc. Amer. Acad. Arts & Sci. 36:462. 1901. TYPE: MEXICO: Sinaloa, Rosario, 7 Jul 1897, J. N. Rose 1568 (Holotype, US!; isotypes, F! GH!; photograph of holotype, TEX!; photograph of K isotype, US!). Annual herbs, 10-50 cm tall. Stems erect to decumbent, 1-3 mm diam, glabrous near base to hispidulous above with hairs up to 0.5 mm long. Peduncles 2-5 em long. Leaves sessile, lanceolate to oblong-ovate, 3-5.5 em long, 0.5-2 cm wide, at apex obtuse to acute, at base subauriculate (rarely obtuse), with both surfaces strigose with hairs 0.3-1 mm long; margin subentire to strongly lobed. Heads 4-6 mm tall, 7-12 mm diam. Outer involucre cupulate, 5-8 mm diam; bracts 5, connate 1/4 or less their length at base, separate to imbricate 3/4 their length, orbiculate to ovate, 2.5.3.8 mm long, 1.5-3.5 mm wide, at apex broadly rounded to acute, with abaxial surface subglabrous to strigillose 1972] Melampodium — Stuessy 65 with hairs 0.3 mm long (longer near base) ; margin scari- ous. Fruits 1.1-2.3 mm long, with lateral surfaces smooth to striate (sometimes very tuberculate); hood apex mu- ticous or less often mucronate to cirrhous, with appendage up to 3 mm long. Ray florets 8-9; ligules yellow-orange, ob- long-elliptic, 3-6 mm long, 1-2.3 mm wide. Disc florets 40- 60; corollas yellow-orange, 1.6 mm diam, with throat 0.7 mm and tube 0.5 mm long. Paleae broadly oblanceolate, 2.7 mm long, 0.8 mm wide; apex yellow, with margin lacini- ate; midrib prominent, weakly puberulent with hairs 0.2 mm long. Chromosome number, n = 10. Tropical deciduous and thorn forests in Sinaloa and ad- jacent Durango, Mexico (Fig. 20), 0-400 m. Flowering dates, Aug-Jan. Some of the specimens on Isla Piedra off the coast of Mazatlán (type locality of M. arenicola) possess more lobed leaves, a tendency toward decumbent stems, larger heads, and narrower and longer ligules than is typical for the species. Because these forms exhibit considerable char- acter intergradations and are found also intermixed in at least one population on the mainland (personal observa- tions), they are not here formally recognized. With M. arenicola and M. rosei regarded as being conspecific and because both names were published at the same time, one of the epithets must be selected for use. M. rosei has been chosen because this name has been applied almost exclu- sively in identifications of previous collections of this taxon. Collections from Ymala (Imala), Sinaloa, at the north- west portion of the range of this species morphologically approach M. cupulatum, a species found nearby (Fig. 19). Hybridization between the two taxa cannot be exluded. REPRESENTATIVE SPECIMENS. Mexico. DURANGO: La Bajada, Tama- zula, Nov 1921, Ortega 4446 (US). SINALOA: Mazatlán, 3 Nov 1893, Brandegee s.n. (US) ; 2 mi SE of Mazatlán, 4 Aug 1938, Eyerdam & Beetle 8687 (ARIZ, UC); Mazatlán, 15 Sep 1927, Ferris 5020 (ps); vicinity of Labradas, 18 Sep 1925, Ferris & Mexia 5078 (ps, GH); near Mazatlán, 29 Jan 1964, Flyr 138 (TEX); ca 2 mi N of Escuinapa, 13 Aug 1960, King 3707 (Ds, NY, TEX, UC, US); ca 12 mi N of Escui- napa, 13 Aug 1960, King 3709 (DS, NY, RSA, TEX, UC); ca 13 mi N 66 Rhodora [Vol. 74 of Rosario, 13 Aug 1960, King 8710 (Ds, NY, TEX, UC, US); ca 21 mi N of Rosario, 13 Aug 1960, King 8712 (DS, NY, TEX, UC, US); ca 3 mi S of Mazatlan, 13 Aug 1960, King 3715 (DS, NY, TEX, UC, US) ; ca 10 mi NE of jet rtes 15 & 40, 14 Aug 1960, King 3716 (DS, NY, TEX, UC); Mazatlán, on Isla Piedra, 31 Dec 1894, Lamb s.n. (DS, UC, US); Mazatlán, 2 Jan 1895, Lamb s.n. (DS, GH, NY, UC, US); San Agustín, 1921, Ortega 4008 (Us) ; Escuinapa, 1926, Ortega 6125 (us), 1933, Ortega 7004 (F); Ymala [Imala], [25°42’ N, 107^15' W; Mc- Vaugh, 1956], 16-25 Aug 1891, Palmer 1457 (F, GH, US), 25 Sep-8 Oct 1891, Palmer 1757a (ARIZ, GH, US); 10 mi E of jet rtes 15 & 40, 26 Aug 1961, Powell & Edmondson 908 (TEX); near Colomas, 19 Jul 1897, Rose s.n. (US); between Mazatlán & Villa Unión, 14 Dec 1936, Shreve 7821 (ARIZ, F); 4.5 mi N of Escuinapa, 4 Sep 1965, Stuessy 404 (TEX); Isla Piedra, Mazatlán, 28 Aug 1966, Stuessy 748 (TEX); 4 mi S of Mazatlán, 28 Aug 1966, Stuessy 750 (TEX); 12-15 km SE of Mazatlán, 4 Aug 1938, Worth & Morrison 8807 (vc, US) ; Mazatlán & vicinity, Dec 1888, Wright 1213 (ps, F, UC, US). 18. Melampodium tenellum Hook. & Arn. Bot. Beech. Voy. 299. 1838. TYPE: MEXICO: Nayarit, Tepic, 1825-28, G. T. Lay & A. Collie et al. s.n. (Holotype, K!; photograph of holotype, F! MICH! OS! TEX! US!; photograph and fragment of holotype, US!). Annual herbs, 6-30 cm tall. Stems erect (rarely decum- bent), 1-2 mm diam, subglabrous to hispidulous with hairs 0.3 mm long. Peduncles 3-7 cm long. Leaves sessile or with petioles 2-10 mm long, lanceolate to narrowly ovate, 2-5.5 cm long, 0.5-1.4 em wide, at apex obtuse (less often acute), at base attenuate (rarely obtuse), with both surfaces scat- teredly strigose with hairs 0.3 mm long; margin subentire to shallowly lobed. Heads 3.5-5.5 mm tall, 6-14 mm diam. Outer involucre cupulate, 3.5-7 mm diam; bracts 5, connate 1/4-1/2 their length, separate to imbricate, orbiculate to ovate, 2.5-3.5 mm long, 2.5-3 mm wide, at apex acute to rounded, with abaxial surface strigillose-hispidulous with hairs 0.8 mm long; margin scarious. Fruits 1.8-2.6 mm long, with lateral surfaces usually with 4 longitudinal rows of tubercles (rarely smooth); hood apex muticous. Ray florets 8-9; ligules yellow-orange, oblong-elliptic, 2-5 mm long, 1-2.8 mm wide. Disc florets 40-60; corollas yellow- orange, 1.3 mm diam, with throat 0.8 mm and tube 0.5 mm 1972] Melampodium — Stuessy 67 long. Paleae oblanceolate, 2.5 mm long, 0.7 mm wide; apex yellow, with margin laciniate; midrib prominent, glabrous. Chromosome number, n — 10. Tropical deciduous forests and savannas in Nayarit, Jalis- co, and Michoacán, Mexico (Fig. 20), 30-920 m. Flowering dates, Jul-Nov. M. tenellum is morphologically similar to M. rosei, but each species does possess unique morphological and geo- graphic features. As in M. rosei, however, M. tenellum also has considerable variation in vegetative and outer phyl- lary merphology. Much additional work will be needed before this intraspecific variation can be understood satis- factorily. REPRESENTATIVE SPECIMENS. Mexico. JALISCO: Puerto Vallarta, 20 Jul 1932, Howell 10303 (US). MiCHOACÁN: 24 km S of Arteaga, 29 Nov 1968, Rzedowski 26618 (os). NAYARIT: E of Tepic along Río Tepic, 22-24 Aug 1935, Pennell s.n. (GH); Santiago Ixc., Mezcal- titan, Jan 1926, González 6125 (ps, US); ca 38 mi S of Nayarit- Sinaloa border on rte 15, 12 Aug 1960, King 3703 (Ds, NY, TEX, UC, US); ca 28 mi S of Nayarit-Sinaloa border on rte 15, 12 Aug 1960, King 3704 (DS, NY, RSA, TEX, UC); ca 21 mi S of Nayarit-Sinaloa border on rte 15, 13 Aug 1960, King 3705 (DS, NY, TEX, UC, US); ca 1 mi S of Nayarit-Sinaloa border on rte 15, 13 Aug 1960, King 9706 (DS, NY, RSA, TEX, UC); 27.9 mi SE of Nayarit-Sinaloa border on rte 15, 4 Sep 1965, Stuessy 401 (TEX); 10 mi NW of jct rte 15 & rd to Tuxpán, 27 Aug 1966, Stuessy 745 (TEX). 19. Melampodium glabribracteatum Stuessy, Brittonia 22: 112. f. 3. 1970. TYPE: MEXICO: Oaxaca, Cerro de San Antonio, Ocotlán, 1600 m, 29 Aug 1936, C. Conzatti 5169 (Holotype, K!; photograph of holotype, os! TEX!). Annual herbs, 15-22 cm tall. Stems erect to decumbent, 1-1.5 mm diam, subglabrous at base to pilose above with hairs up to 1 mm long. Peduncles 1.3-1.7 cm long. Leaves sessile, blades lanceolate, 2-3.3 em long, 0.4-0.7 cm wide, at apex obtuse, at base obtuse-subauriculate, with both sur- faces sparingly strigose with hairs up to 1 mm long; margin entire or rarely 2-lobed, near base pilose with hairs up to 1 mm long. Heads 5-6 mm tall, 6-9 mm diam. Outer in- volucre cupulate, 7-8 mm diam; bracts 5, slightly connate 58 Rhodora [Vol. 74 at base, imbricate 3/4 their length, elliptic-orbiculate, 3-4 mm long, 2.4-3.2 mm wide, at apex rounded, with abaxial surface glabrous; margin narrowly scarious, ciliate with hairs up to 0.6 mm long. Fruits 2.1-2.5 mm long, with laterai surfaces ribbed and tuberculate-aculeate; hood apex cirrhous, with tapering appendage up to 2 mm long. Ray florets 6-8; ligules yellow, ovate-oblong, 3.5 mm long, 2-2.5 mm wide. Disc florets ca 30; corollas yellow, 1 mm diam, with throat 1 mm and tube 0.7 mm long. Paleae oblance- olate, 3.3 mm long, 0.8 mm wide; apex yellow, with margin smooth-erose; midrib prominent, glabrous. Chromosome number unknown. Known only from the type collection from pine-oak for- ests near Ocotlán, Oaxaca (Fig. 11), 1600 m. Flowering date, Aug. 5. Series Longipila Stuessy, ser. nov. Herbae annuae; folia late ovata, subtus sericea; ligulae flavae-aurantiacae, plus quam 2 mm longae; involucrum extimum marginibus herbaceis; fructi apice arista adaxiali complanato cirrhoso. Species 20. Typus: Melampodium longipilum Robins. 20. Melampodium longipilum Robins. Proc. Amer. Acad. Arts & Sci. 27:173. 1892. TYPE: MEXICO: San Luis Potosi, San José Pass, 11 Jul 1890, C. G. Pringle 3639 (Holotype, GH!). Melampodium villicaule Greenm. Field Col. Mus. Pub. Bot. 2:345. 1912. TYPE: MEXICO: Guerrero, Rio Baisas, 26 Aug 1910, C. R. Orcutt 4386 (Holotype, F!). Annual herbs, 7-30 cm tall. Stems erect, 0.7-4 mm diam, villous with hairs 2 mm long. Peduncles 2-11.5 cm long. Leaves with short and broad petioles (rarely sessile) 3-6 mm long, with blades ovate to elliptic, 2-7.5 cm long, 0.5-3 em wide, at apex acute, at base attenuate to partially sub- auriculate, with both surfaces pilose with hairs 1 mm long; margin entire. Heads 5-7.5 mm tall, 7-12 mm diam. Outer involucre markedly cupulate, 6.5-9.5 mm diam; bracts 5, 1972] Melampodium — Stuessy 69 slightly connate at base, imbricate 1/2 their length, ovate, 3.5-5 mm long, 2-3.5 mm wide, at apex acute, with abaxial surface pilose with hairs 1 mm long; margin herbaceous. Fruits 2.2-2.8 mm long, with lateral surfaces tuberculate or striate with tapering cirrhous appendages up to 8 mm long. Ray florets 7-8; ligules yellow-orange, ovate-elliptic, 3.5-5.5 mm long, 1.8-4 mm wide. Disc florets 40-70; corollas yel- low-orange, 1.2 mm diam, with throat 0.8 mm and tube 0.5 mm long. Paleae oblanceolate, 3.5 mm long, 0.8 mm wide; apex yellow, with margin laciniate-dentate; midrib prominent, pubescent with hairs 0.5 mm long. Chromosome number, n = 10." Scattered in tropical deciduous and pine-oak forests of western Guatemala and the Mexican states of San Luis Potosi, Hidalgo, Puebla, Guerrero, and Oaxaca (Fig. 12), 1870-3050 m. Flowering dates, Jul-Nov. In my opinion this species is morphologically closest to M. tenellum, although Turner and King (1962) have sug- gested that its relationship is with M. divaricatum (in sec- tion Serratura). 'The unusual adaxially flattened appendage (Fig. 8) makes M. longipilum unique within the genus, and this feature along with other characters such as large ovate leaves and markedly cupulate outer involucres suggests separation of the species as a monotypic series. Considered somewhat rare before the present treatment, M. longipilum is quite common especially throughout Oaxaca. REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Cumbre Papal between Cuilco & Ixmoquí, 19 Aug 1942, Steyermark 50916 (F, NY). Mexico. GUERRERO: 19 mi N of Chilpancingo, 29 Aug 1965, Stuessy 374 (TEX). HIDALGO: Metztitlán, SE of Metztitlán, 24 Nov 1942, Moore 2283 (GH). OAXACA: ca 64 mi SE of Oaxaca, 23 Jul 1960, King 3461 (DS, NY, TEX, UC, US); 13 mi NW of Tehuantepec, 23 Aug 1965, Stuessy 328 (TEX); 3.8 mi NW of Huajuapán de León, 25 Aug 1965, Stuessy 344 (TEX) ; 10 mi NW of Tehuantepec, 11 Aug 1966, Stuessy 634 (os, TEX); 51 mi NW of Tehuantepec, 11 Aug 1966, Stuessy 636 (TEX); 87 mi NW of Tehuantepec, 11 Aug 1966, Stuessy 637 (TEX); 28 mi S of Zimatlán, 12 Aug 1966, Stuessy 639 (TEX). PUEBLA: Tehuacán, Sep 1911, Purpus 5619 (GH, NY, UC); near "See Stuessy (1971b) for a discussion of the n — 11 count for this species. 70 Rhodora [Vol. 74 Tehuacán, 36 Aug-8 Sep 1905, J. N. & J. S. Rose & Painter 10157 (US); Asunción de Chila, 25 Aug 1965, Stuessy 346 (TEX); 32 mi NW of Huajuapán de León, 25 Aug 1965, Stuessy 347 (TEX); 2 mi NW of Tehuitzingo, 16 Aug 1966, Stwessy 667 (TEX). SAN LUIS POTOSÍ: Villar, 14 Sep 1893, Pringle 4537 (F, G, NY, UC, US [2]). (To be continued) IN MEMORIAM CHARLES SCHWEINFURTH, 1890-1970 November 16, 1970 marks the end of an outstanding era in the annals of American orchidology. Charles Schweinfurth was born on April 13, 1890 in Brookline, Massachusetts, in one of the oldest and most 71 72 Rhodora [Vol. 74 historic regions of the United States. During his formative years he attended public schools in Brookline where he quickly developed an intense devotion to various aspects of natural history. While his interest in bird watching and plant collecting started as an avocation, they became the driving force for his whole life soon after graduation from college. He entered Harvard University in 1909 and majored in chemistry, graduating in 1913 with a degree of A.B., cum laude. 'This was a remarkable achievement, for in his sopho- more year he succumbed to polio which paralyzed his right arm. To regain the use of his arm, with a self discipline that eharacterized his whole life, he soon started daily ex- ercises, aided by his devoted parents. He maintained this discipline almost to his last days. When he was informed by his doctors that he must abandon chemistry, he found a life-long career in his avocation. Charles’ commitment to botany became officially regis- tered on April 5, 1912 when he was elected a member of the New England Botanical Club. He faithfully supported, as well as promoted, its interest, often beyond the call of duty. It was, indeed, a rare occurrence when Charles would succumb to some more compelling business than the attend- ance of the club’s monthly meetings. It was in this same spirit that he accepted the burdensome responsibility of corresponding secretary in 1949, a post he held in the club until 1965, when he had to resign from it due to his doctor’s order. His memberships included, among others, also the American Association for the Advancement of Science, the International Association for Plant Taxonomy, the National Audubon Society, the Appalachian Mountain Club, and the Boston Museum of Science. Charles had many avocations in addition to his ardent bird watching. During the summer months he enjoyed traveling, since he was greatly interested in the classical aspects of our cultural past. Consequently photography became a second nature to him in recording precious im- pressions, especially of paintings, sculptures and architec- 1972] Schweinfurth — Garay 73 tural designs for his leisure viewing at home or for sharing them with others at some later time. During winter months he spent much of his extra-curricular time on writing for local newspapers, as well as in pursuit of winter sports, notably ice skating. The professional life for Charles Schweinfurth started in 1914 when he accepted a position offered to him by Pro- fessor Oakes Ames to tend the living orchid collection at North Easton, Massachusetts. Within less than a year Pro- fessor Ames, recognizing Charles’ great ability for remem- bering scientific names and fine details, took him on as a personal assistant to work in his private “Ames Botanical Laboratory", also in North Easton. It was then that his orchidaceous career started. From January, 1915, Charles Schweinfurth was an orchidologist. He worked intensively on the orchids of the Philippines, those from Mt. Kinabalu, British North Borneo and various Pacifie islands, followed by studies of the orchids of Hon- duras, Costa Rica and Panama. In 1936, he co-authored with Ames and Hubbard an extensive monographic study, “The Genus Epidendrum in the United States and Middle America". These works alone would suffice to record him among out- standing orchidologists. Charles, however, has built an even more imposing monument for posterity. True to his good nature and encouraged by Professor Ames, in 1922 he ac- cepted an offer made by J. Francis Macbride to write up the orchid family for his proposed “Flora of Peru". He labored year after year and on April 9, 1958, the first of the four parts of his monumental work appeared in pub- lication. This work was the first descriptive treatise on orchids of any Andean country. Charles, like many of the outstanding botanists, while working on a given project, concurrently undertook others. He accepted the very onerous task of identifying the nu- merous colored plates prepared under the supervision of Celestino Mutis during La Real Expedición Botániea del Nuevo Reino de Granada, between 1760 and 1817. The first 74 Rhodora Vol. 74 volume of this labor of love was published in 1963, while the second volume appeared in 1969. In 1958, Charles attended the 3rd South American Bo- tanieal Congress in Lima, Peru, where he was awarded Catedratico Honorario (Honorary Professor of Botany) by Universidad Mayor de San Marcos, the oldest university of the Americas. It was during that year, when he also visitel Colombia, that the picture reproduced here was taken by his colleague, Dr. Hernando Garcia-Barriga of Universidad Nacional, Bogota, during a visit to the Paramo de Chipaque. Charles received another award of Catedratico Honorario in 1962, from Universidad de Cuzco. In 1964, he was hon- ored by the American Orchid Society with a Gold Medal inscribed “For outstanding contributions to orchidology”, and on July 12, 1966, he was elected Miembro Honorario by Sociedad Colombiana de Orquideologia. On November 29, 1965, the staff of the Botanical Museum of Harvard University honored him on the occasion of his Golden Jubilee in Orchidology with a scroll, inscribed as follows: “Fifty years have passed since our colleague Charles Schweinfurth initiated his productive career in orchidology. Dean of the world’s orchidologists, like his predecessors Lindley, Rolfe, Reichenbach, Schlechter, Kraenzlin, Smith and Ames, he has made invaluable con- tributions to our knowledge of the systematics of the orchids of both the Old and the New World. His great number of publications, his elucidation of the intricate structure of orchid flowers, his augmentation to our knowledge of the phytogeography of orchids, his acute powers of observation for minute details, his loyalty and his dedication to his chosen field stand as an inspiration to both present and future students of orchidology. In grateful recognition of these fifty years of service to orchidology, we of the Botan- ical Museum of Harvard University present this scroll.” The most meaningful event in Charles’ life, however, oc- curred in December 1963, when, after a visit to the Holy Land, he married the lifelong friend of the Schweinfurth 1972] Schweinfurth — Garay 75 family, Miss Maria Elizabeth Westergren of Stockholm, Sweden. The blessings of this mature companionship he openly cherished throughout the rest of his life. The orchid world is vastly richer today for there was a Charles Schweinfurth intimately associated with it. And for those few who had the privilege of sharing his meaning- ful friendship, not only the orchid world, but the whole world was just a little better to live in. LESLIE A. GARAY ORCHID HERBARIUM OF OAKES AMES BOTANICAL MUSEUM HARVARD UNIVERSITY NOMENCLATURAL CHANGES IN LESQUERELLA REED C. ROLLINS AND ELIZABETH A. SHAW During the last twenty years there has been a gradual introduction into the International Code of Botanical No- menclature of provisions for the automatic recognition of tautonymous names and epithets when a taxon in a cate- gory of infrafamilial rank (other than genus or species) is segregated from one of the next higher rank. In the Code adopted at Stockholm (Reg. Veg. 3: Arts. 34, 35. 1952) it was clearly stated that valid publication of the name of a taxon of infraspecific rank which does not include the type of the name of the taxon of the next higher rank automaticaliy establishes in parallel the name of a second taxon of the same subordinate rank, this having the same type as does the name of the taxon of higher rank and the same epithet. At Paris similar provisions were made for categories of infrageneric rank (Reg. Veg. 8: Art. 22. 1956) and at Montreal for categories of infrafamilial rank (Reg. Veg. 23: Art. 19. 1961). However it soon became apparent that the very broad application of this useful principle to the name of any taxon which includes the nomenclatural type of the name of the taxon next higher in rank can give rise to complications. Although the International Code of Botanical Nomencla- ture (Reg. Veg. 46: Arts. 2, 3. 1966) makes it clear that every Recent plant belongs to a species, a genus, and a fam- ily (as well as to an order, a class, and a division) it nowhere provides that a plant must belong to a subspecies or to a section, a series, or to a tribe. The use of such cate- gories of intermediate, and optional, rank reflects only a recognition of the possibility of using more hierarchical steps within the framework of the required classification. The sweeping extension of the principle of automatic rec- ognition of tautonyms to names of taxa in such optional ranks creates the possibility that a taxon with a particular circumscription, position and rank might have, according 76 r 1972] Lesquerrella — Rollins and Shaw Ti to the classification used, two correct names. Classification and nomenclature are thus entangled in a way contrary to Principle IV of the Code. Such distressing situations were discussed by Wood & Webster (1968) and a series of care- fully worded proposals intended to clarify matters was sub- mitted to, and accepted by, the 11th International Botanical Congress in 1969. In essence, these modifications restrict the automatic recognition of tautonyms in the optional ranks to names of only those taxa which do include the type of the name of a species, a genus, or of a family. In the other optional ranks, strict priority of publication ap- plies. Difficulties arising from the entanglement of classification and nomenclature have come to light more often in dealing with categories ranked between family and genus, or be- tween genus and species, than with those below the rank of species, and Wood & Webster were not entirely satisfied with the example used with their proposal for modification of Article 26, which deals with names of taxa of infra- specific rank, in the then current edition of the Code. In the course of our revision of the North American species of Lesquerella (in press), we made certain decisions result- ing in our recognition of two subspecies of Lesquerella lasiocarpa (Hook. ex Gray) Watson, and, in that subspecies not including the type of the name of the species, of two varieties. We realized that one taxonomic decision in par- ticular had produced precisely the situation Wood & Web- ster had in mind when proposing the change in Article 26. Since this is being used as an example in the forthcoming edition of the Code, we wish to make the necessary nomen- clatural changes here and to provide some explanation of them. Our classification of L. lasiocarpa is summarized as follows: (1) Lesquerella lasiocarpa subsp. lasiocarpa Vesicaria lasiocarpa Hook. ex Gray, Smithson. Con- trib. Knowl. 5: 13. 1853. Type: Texas. between Bexar & Trinity River, Berlandier in 1828 (K) 78 Rhodora [Vol. 74 (2) Lesquerella lasiocarpa subsp. berlandieri (Gray) Rol- lins & Shaw, comb. nov. Synthlipsis berlandieri Gray, Bot. Mex. Bound. Sur- vey 34. 1859. Lectotype (here designated): Mexico. Tamaulipas, Matamoros, Berlandier 3017 (GH), iso- types (MO, US) (22) L. lasiocarpa [subsp. berlandieri] var. hispida (Wats.) Rollins & Shaw, comb. nov. Synthlipsis berlandieri var. hispida Wats., Proc. Amer. Acad. 17: 321. 1882. Holotype: locality uncertain, Palmer 26 (GH) S. berlandieri Gray (see above) L. lasiocarpa var. berlandieri (Gray) Payson, Ann. Missouri Bot. Gard. 8: 189. 1922 L. lasiocarpa var. ampla, Rollins, Rhodora 57: 245. 1955. Holotype: Mexico. Tamaulipas, vicinity of Victoria, Palmer 41 (GH), isotype (NY) (2b) L. lasiocarpa [subsp. berlandieri] var. hetero- chroma (Wats.) Rollins, Rhodora 57: 245. 1955 Synthlipsis heterochroma Wats., Proc. Amer. Acad. 17: 321. 1882 According to the provisions of the 1966 edition of the Code, it would have been necessary to use the name L. lasio- carpa, [subsp. berlandieri] var. berlandieri [without an au- thor— not *(Gray) Payson"] for that variety (2a) of subsp. berlandieri which includes the type of the name of the subspecies, no matter how it might be circumscribed, when recognizing taxonomically, as we do, this optional category. Still following the 1966 edition of the Code, if one chose nct to adopt the category “subspecies” in classify- ing the infraspecific taxa of Lesquerella lasiocarpa, but used only “varietas,” the correct name for this taxon, still with the same circumscription, position, and rank, would be L. lasiocarpa var. hispida (Wats.) Rollins & Shaw. This is the earliest available epithet in the rank of variety for a taxon which does not include the type of the name of the taxon of next higher rank. Thus, depending upon the classi- 1972] Lesquerrella — Rollins and Shaw 79 fication used (whether subspecies or not in the present in- stance), this one variety could have two correct names, a situation contrary to Principle IV. The crux of the matter is our decision to circumscribe the variety which includes the type of the name L. lasiocarpa subsp. berlandieri so as to include Synthlipsis berlandieri var. hispida Watson. Article 26 as modified at Seattle now provides that use of a tautonymous epithet without citation of an author’s name applies only to the names of taxa of infraspecific rank which do include the type of the name of the species to which they are assigned. Since variety 2a does not include the type of the name L. lasiocarpa, the earliest available epithet in varietal rank, if one such exists, must then be used for it, and in this case it is “hispida”. In Payson’s classification, based upon different criteria, S. ber- landieri var. hispida was not placed in the synonymy of S. berlandieri, and he was thus quite correct in his use of the name L. lasiocarpa var. berlandieri (Gray) Payson. How- ever, according to the Code as modified at Seattle, the taxon, as circumscribed by us, is to be called Lesquerella lasio- carpa var. hispida (Wats.) Rollins & Shaw. LITERATURE CITED Woop, C. E. & G. L. WEBSTER. 1968. Tautonyms and Confusion in tbe International Code. Taxon 17: 645-651. GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138 THE PARKER CLEAVELAND HERBARIUM OF BOWDOIN COLLEGE EDWARD HEHRE, A. R. HODGDON, AND R. B. PIKE! The Herbarium of Bowdoin College, Brunswick, Maine, which was assembled in the 19th century under the direc- tion of Parker Cleaveland (1780-1858) was stored away in the attic of Searle’s Science Building at Bowdoin College for over half a century. Professor Cleaveland was a dis- tinguished naturalist in the nineteenth century tradition. He is best known for the collection of minerals housed for many years in the *Cleaveland Cabinet" in Massachusetts Hall at Bowdoin College. Cleaveland had studied under Goethe who recommended him highly when he began his career in Brunswick in 1805. That he had much knowledge of botany and of botanists is suggested by the important collections that came his way. However, the Cleaveland Collection, like many herbaria in small colleges, ceased to grow and became isolated and inaccessible after its aus- picious beginning, and because of neglect, failed to fulfill the purpose for which it had been assembled. Through the kindness of the President and overseers of Bowdoin College, the Cleaveland Herbarium is now on a ten-year loan to the University of New Hampshire. By means of support provided by Mr. Sumner Pike of Lubec, Maine, the collection of approximately 9,400 speci- mens was given a thorough and painstaking rehabilitation by the senior author. Most specimens were in good condi- tion with a minimum of damage by mold and insects but nearly all required remounting and, for a great many, some nomenclatural updating. The entire collection now is incor- porated in the University of New Hampshire Herbarium (NHA), each sheet being stamped with the Bowdoin Seal for ready recognition. "Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as scientific contribution number 610. 80 1972] Herbarium — Hehre, Hodgdon & Pike 81 For a relatively small collection, the Cleaveland Her- barium is notable for its representation of important collectors. One can speculate that Professor Cleaveland intended at the beginning to create an important botanical center at Bowdoin. The first two successful collectors who worked for Gray in the 1840s are represented with 124 specimens by Ferdi- nand Lindheimer (1801-1879) and 207 from New Mexico by Augustus Fendler (1813-1883) of which 39 are isotypes of species described by Gray and others. There are even duplicate isotypes for 13 of the taxa. Dupree (1959) briefly and McKelvey (1955) at much greater length discuss the activities of these two botanical collectors of the American Southwest. There follows a partial and by no means complete list of other botanists whose collections are to be found in the Cleaveland Herbarium: Michael S. Bebb (1833-1895) — 157 specimens, preponder- antly from Illinois. Joseph Blake (1814-1888) — nearly 1500, chiefly from Cumberland County in Maine and some in Cóos, Carroll, Grafton and Belknap Counties in New Hampshire. William Boott (1805-1887) — 16 from the White Mountains of New Hampshire. William N. Canby (1831-1904) — 256 from Delaware and adjoining areas. Parker Cleaveland (1780-1858) — about 500, mostly from Brunswick, Maine. Chester Dewey (1784-1867) — 4 of Carex. Asa Gray (1810-1888) — 16 (ex herbario W. S. Poor) John S. Henslow (1796-1861) — father-in-law of Joseph Dalton Hooker and whose recommendation led to Dar- win's accepting the post of naturalist on the “Beagle” — 271 from Great Britain. Marcus E. Jones (1852-1934) — 66 collected about 1880 in Rocky Mountain area. Peter D. Knieskern, M.D. (1798-1871) —119 from New Jersey. 82 Rhodora [Vol. 74 William Oakes (1799-1848) —52 from Massachusetts and New Hampshire, without date. Thomas C. Porter (1822-1901) — 129 from Pennsylvania. Henry P. Sartwell (1792-1867) — 114 from western New York State, 87 of these being Carex. Frank Lamson Seribner (1851-1938) — the first U.S.D.A. Plant Pathologist — 140 from Manchester, Maine. Edward Tuckerman, Jr. (1817-1886) — 250 mostly from New England, a few from central Europe. George Vasey (1822-1893) — 160 from Illinois. Oliver R. Willis (1815-1902) — 335 from New Jersey. Nearly all the specimens of any particular collector are of different species and since many of the collectors were specialists on certain groups and worked in different parts of the world, the collection contains a very great number of species. The Herbarium of the University of New Hampsnire (NHA) is now being computerized and it will soon be pos- sible to provide interested persons with considerable infor- mation as to the collectors represented as well as informa- tion on the labels of the plants they collected. For the Parker Cleaveland collection, in the case of each important collector, we have made lists of all species represented. Thus we are in a position now to supply this information. Wooton & Standley (1915) give type localities for all taxa and often specify the collectors. For Fendler, usually the collection number is cited making it comparatively easy to locate his isotypes. A large percentage of Fendler’s isotypes of New Mexican plants is represented in the Cleaveland collection. List of Fendler isotypes in the Parker Cleaveland Her- barium: 1. Selaginella Underwoodii Hieron., (in duplicate), Fend- ler 1024. Cheilanthes Fendleri Hook., Fendler 1015. Poa Fendleriana (Steud.) Vasey, (in duplicate), Fend- ler 932. oo po Herbarium — Hehre, Hodgdon & Pike 83 P. Bigelovii Vasey & Scribn., Fendler 931. Aristida Fendleriana Steud., Fendler 973. A. longiseta Steud., (in duplicate), Fendler 978. Cyperus Fendlerianus Boeckel, Fendler 865. Salix Fendleriana Anderss., Fendler 816. S. irrorata Anderss., Fendler 812. Quercus novemexicana (A.DC) Rydb., Fendler 809. . Arceuthobium cryptopodium Engelm., Fendler 283. Phoradendron juniperinum Engelm., Fendler 281. Abronia Fendleri Standley, Fendler 739. Lesquerella intermedia (S. Wats.) Heller, (in dupli- cate), Fendler 38. . Draba neomexicana Greene, (in duplicate), Fendler 43, . Arabis Fendleri (S. Wats). Greene, Fendler 27. Potentilla propinqua Rydb., Fendler 198. Astragalus ceramicus Sheld., (in duplicate), Fendler 161. A. lonchocarpus 'Torr., Fendler 160. Euphorbia Fendleri Torr. & Gray, (in duplicate), Fendler 800. Sphaeralcea Fendleri A. Gray, Fendler 78. Galpinsia Fendleri (A. Gray) Heller, Fendler 230. Cymopterus Fendleri A. Gray, Fendler 274. Daphnidostylis Fendleriana Klotzsch, Fendler. Dodecatheon radicatum Greene, Fendler 549. Mertensia Fendleri A. Gray, Fendler 625. Oreocarya fulvocanescens (S. Wats.) Greene, (in du- plicate), Fendler 632. Oreocarya multicaulis (Torr.) Greene, Fendler 636. Hydrophyllum Fendleri (A. Gray) Heller, Fendler 642. Lycium pallidum Miers., Fendler 670. . Physalis Fendleri A. Gray, (in triplicate), Fendler. . P. similas A. Nels., Fendler 575. Eupatorium Fendleri A. Gray, Fendler 347. Erigeron cinereus A. Gray, (in duplicate), Fendler 914. 84 Rhodora [Vol. 74 35. Actinella argentea A. Gray, (in duplicate), Fendler 457. 36. Actinella Richardsonii Nutt. var. floribunda A. Gray (in duplicate), Fendler 460. 37. Diplopappus ericoides Torr. & Gray var. hirtella A. Gray, (in duplicate), Fendler 348. 38. Bidens tenuisecta A. Gray, Fendler 449. 39. Agoseris purpurea (A. Gray) Greene, Fendler 487. For many years there has been a diminishing support for small herbaria, some of which, like the Cleaveland Col- lection at Bowdoin College, are of historic importance and have scientific value. The space they occupy has been en- croached upon by classrooms and by modern research setups. This does not mean that the collections are less important than they ever were. It does suggest that the custodians of such herbaria should take steps, as Bowdoin College has done, to make their specimens available to the scientific community. LITERATURE CITED DUPREE, A. HUNTER. 1959. Asa Gray, Harvard University Press, 505 p. MCKELVEY, SUSAN DELANO. 1955. Botanical Exploration of the Trans-Mississippi West 1790-1850. The Arnold Arboretum of Harvard University, 1144 p. Wooton, E. O. and PAUL STANDLEY. 1915. Flora of New Mexico, Contrib. U.S. Nat. Herb., Vol. 19. DEPARTMENT OF BIOLOGY SOUTHAMPTON COLLEGE OF LONG ISLAND UNIVERSITY SOUTHAMPTON, NEW YORK 11968 DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 DEPARTMENT OF PLANT SCIENCE UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 STUDIES ON THE AUDOUINELLA MICROSCOPICA (NAEG.) WOELK. COMPLEX (RHODOPHYTA) WILLIAM J. WOELKERLING Several recent studies (Abbott 1962, West 1968, Woelker- ling 1970, 1971) have indicated that the taxonomic status of numerous species in the Audouinella complex (Acrochae- tium-Rhodochorton complex) of the red algae is in need of review. Many of these taxa have been described from meagre material and (or) without regard for possible in- traspecific variation, and recent work on several species (Abbott 1968, West 1969, Woelkerling 1970) has resulted in a reduction of some taxa to synonomy. Preliminary work of the author on the audouinelloid algae of the New England coast of North America has led to a detailed consideration of the relationships of Audowi- nella microscopica (Naegeli) Woelkerling to six closely re- lated taxa: Acrochaetium crassipes Boergesen (1909, p. 1, Fig. 1; 1915, p. 20, Figs. 11-13), A. catenulatum Howe (1914, p. 84, pl. 31, Figs. 12-18), A. microfilum Jao (1936, p. 240, pl. 10, Figs. 1-5), Kylinia collopoda (Rosenvinge) Kylin (see Rosenvinge 1898, p. 41, Figs. 10-11; 1909, p. 81), K. compacta (Jao) Papenfuss (see Jao 1936, p. 241, pl. 10, Figs. 6-14), and K. moniliformis (Rosenvinge) Kylin (see Rosenvinge 1909, p. 98, Figs. 28-29). (It should be noted here that although the taxonomic proposals of Woelkerling (1971) have been adopted in this study, older generic names have been employed in cases of probable synonomy in order to avoid making new and unnecessary nomencla- tural combinations). A. crassipes and A. catenulatum were described respectively from material collected in the Vir- gin Islands (Boergeson 1909) and Peru (Howe 1914) while the other four taxa have been described from or are re- ported to occur along the New England and adjacent coasts. These seven species have been distinguished from one another on slight differences in habit, branching, cell size, 85 86 Rhodora [Vol. 74 chromoplast shape, and spore size. These apparent differ- ences, however, may not be as taxonomically reliable as formerly thought, and Woelkerling (1971) has suggested that several or all of the taxa under discussion may be con- specific. The aims of the present investigation have been: 1) to critically examine and compare morphologically the type collections and other populations of these seven taxa, and 2) to clarify taxonomic limits within this species com- plex, particularly with reference to the New England flora. MATERIALS AND METHODS The morphological techniques employed in these studies have been detailed elsewhere (Woelkerling 1970). Line drawings have been made with the aid of a Leitz drawing head microscope attachment; herbarium abbreviations fol- low Lanjouw and Stafleu (1964). Wherever possible, results have been based on the study of populations (Table 1) rather than isolated individuals. This approach has been facilitated by the nature of the material; i.e., audouinelloid algae sometimes occur in large numbers on various substrates and by virtue of their small size, dozens or hundreds of individuals may be present in a single collection. The data presented here in most cases represents the results of study on numerous individuals within each population; an exception is the type collection of Acrochaetium crassipes Boergesen which is represented in C only by several drawings. Data on this taxon has been taken from the accounts of Boergesen (1909, 1915). MORPHOLOGY Audouinella microscopica, the earliest described member of this complex, was first characterized by Kuetzing (1849), and later Naegeli (1861) discussed and illustrated it in somewhat greater detail. Although Kuetzing (1849, p. 640) cited the Bay of Naples as the type locality, Hamel (1927; 1928) has indicated that the type collection came from Torquay, England and is represented by specimen 454 in Hauck and Richter's ^Phykotheca Universalis" (as Chan- 1972] Audouinella — Woelkerling 87 | M Figs. 1-14. Audouinella microscopica (Naegeli) Woelkerling. Figs. 1-6. Monosporangial plants from type collection. Note variation in habit and in development of basal cell wall. Figs. 7-8. Spermatangial plants from type collection. Figs. 9-11. Cystocarpic plants from type collection. Note remains of trichogyne and an apparent transversely divided carpogonium (Fig. 9). Figs. 12-14. Sexual plants from Den- mark removed from the host, Chordaria. Note variation in develop- ment of basal cell wall and carpogonium with attached spermatium (Fig. 13). 88 Rhodora [Vol. 74 transia secundata (Lyngbye) Thuret). The type specimen in the Kuetzing collections at L contains only plants col- lected by Naegeli in England (identical to those distributed in Hauck and Richter) and none from the Bay of Naples. The location given by Kuetzing (1849) is, therefore, ap- parently in error. Plants from the type collection of A. microscopica in L as well as isotypes in FH and NY have been examined during this study. The plants form a dense population on Enter- omorpha (Chlorophyta), are 40-100 (-200) p tall, and are attached to the host by unicellular bases with or without enlarged lower cell walls (Figs. 1-11). One or occasionally several erect filaments arise from the basal cell (Figs. 3, 4), and these may remain unbranched (Fig. 1-2) or bear several secundly or irregularly arranged laterals (Figs. 3- 11). Cells are doliiform to cylindrical in shape, 6-8 (-12) & wide and 6-12, (1-2 diameters) long; terminal hairs up to 40u long occur occasionally. Both monosporangial and sexual plants are present in the material examined. The monosporangia occur singly or in pairs, are 7-10, long and 5-7, wide, are sessile or stalked, and are scattered over the erect filaments (Figs. 1- 6). Spermatangia are ovoid, up to 4u long, occur singly or in pairs, are sessile or stalked and are scattered over the erect filaments (Figs. 7-8). Unfertilized carpogonia have not been observed definitely in the type collection material, but the remains of at least one apparently transversely divided fertilized carpogonium has been seen (Fig. 9). Gonimoblasts are several celled and give rise to terminal or lateral carposporangia 7-10, long and 6-8, wide (Figs. 10-11). Only the clustered arrangement of the carpospor- angia distinguishes the gonimoblast from a monosporangial bearing branch. Tetrasporangial individuals have not been observed but are reported by Schiffner (19531). Hamel (1927, 1928) previously reported sexual individ- uals, and Lund (1942) noted possible antheridia in Danish plants. One Danish collection on loan from Dr. Lund has been examined and found to contain numerous sexual in- 1972] Audouinella — Woelkerling 89 dividuals including specimens with carpogonia bearing at- tached spermatia (Figs. 12-14). Lund (1942) referred the Danish material to Chantransia (= Kylinia) collopoda, a taxon here considered conspecific with Audouinella micro- scopica. Monosporangial plants more or less agreeing with the above description of A. microscopica occur along the New England and adjacent coasts, but up to the present time, they have been referred (Edelstein & McLachlan 1966; Edelstein et. al. 1967; Jao 1936; Taylor 1937, 1957) to four other species (see above) including two (A. microfilum and K. compacta) with type localities in the Cape Cod region. An analysis (Table 1) of a number of New England populations including those cited by the above authors, strongly indicates that they agree in all essential features with A. microscopica and are therefore justifiably referred to that species. The analysis further indicates that while the range in basal attachment, height, branching, number of erect axes, chromoplast shape, cell size, and spore size may vary some- what from one population to the next (probably attribu- table to variation in environmental factors and age), con- siderable overlap in these characters exists between various collections and in no cases can distinct specific limits be drawn. Consequently, it appears that all these plants are best regarded as members of a single, variable species — A. microscopica. Woelkerling (1971) has found similar vari- ation in southern Australian populations of this species. Ecological data on A. microscopica in New England re- mains scant. Specimens have been collected from July through February, but the species probably occurs through- out the year and has thus far escaped detection by virtue of its small size. Sexual plants have not been reported to date in New England, and indeed have been recorded only twice from European waters. A. microscopica has been found growing on a number of algae in New England waters (Chaetomorpha, Chondria, Chondrus, Chordaria, Clado- phora, Entromorpha, Polysiphonia, Porphyra) all of which 90 Rhodora [Vol. 74 occur in the sublittoral or in the drift. The species no doubt enjoys a much wider host distribution and is to be sought particularly on old and heavily epiphytized algae. SYSTEMATIC IMPLICATIONS The results of this study again (see Woelkerling 1971) raise the question as to whether a number of taxa closely related to Audowinella microscopica are really distinct species. The type collections of six of these taxa have been available for study, and a detailed analysis (Table 1) strongly indicates that taxonomic distinctions cannot be made among them on the bases of height, number of erect axes, cell size, or spore size. As is the case for the various New England collections, considerable overlap in the above characters is evident in the type collection pop- ulations, and specific limits cannot be clearly drawn. More- over, a comparison of the type collection illustrations (see Boergesen 1909; Howe 1914; Jao 1936; and Rosenvinge 1909) also indicates the great similarity of these taxa. In addition to the above characteristics, apparent differ- ences in habit, development of the basal cell wall, degree of branching, origin of laterals, chromoplast shape, pres- ence or absence of hairs, and position of sporangia have been used in making specific distinctions. As Woelkerling (1971) has shown in a lengthy review, the degree of branching, presence or absence of hairs, and sporangial position are not taxonomically reliable in general for making specific distinctions in the Awdouinella complex, and the present study supports this view. Thus, for example, the type col- lection of A. microscopica contains plants which vary in the degree of branching, may or may not have unicellular hairs, and possess both terminal and lateral sporangia (Figs. 1- 11). Hamel (1927, 1928) attempted to distinguish A. micro- scopica from other members of this complex on the basis of lateral branch origin. Thus, according to Hamel, the first lateral branch in A. microscopica always arises from the first cell above the base. The type collection of this species, 1972] Audouinella — Woelkerling 91 however, also contains unbranched plants (Figs. 1-2) and plants in which the lateral arises from the basal cell (Fig 4) or from two or more cells above the base (Figs. 9-11). Woelkerling (1971) found similar variation in Australian populations of A. microscopica, and this variation also oc- curs in New England populations. Thus it appears that specific distinction based on origin of lateral branches is not taxonomically reliable. The presence of a much enlarged basal cell wall has been used (Rosenvinge 1898, 1909; Lund 1942) to distinguish Kylinia collopoda from Audouinella microscopica. However, considerable variation in basal cell wall development occurs in the type collection of A. microscopica (Figs. 1-11) as well as collections made by Lund (Figs. 12-14), and Woel- kerling (1971) reported a similar situation in southern Australian populations of this species. This variation sug- gests that this character also is not taxonomically reliable for delimiting species in this complex. Chromoplast shape (parietal vs. stellate) has also been used (Taylor 1957) to distinguish these species. However, recent work (West 1968, p. 92, 95; Woelkerling 1971) has indicated that plastid shape shows considerable intraspe- cific variation and therefore is not a generally trustworthy taxonomic criterion. Some variation has already been re- corded in plastids of A. microscopica (Woelkerling 1971), and studies (e.g., Abbott 1962, p. 100; Boergesen 1937, p. 39, 41; Drew 1928, p. 156, 176, 177, 182; Feldmann 1962, p. 220; Levring 1937, p. 94) have noted similar variation in other species. It appears, therefore, that the taxa under discussion cannot be distinguished from one another solely on apparent differences in chromoplast shape. This study has not revealed any other criteria by which these taxa may be reliably separated into distinct species; consequently all are regarded here as conspecific with Au- douinella microscopica. The relationships of A. microscopica to four other taxa remain uncertain, primarily because the type collections have not been available for examination. The taxa include 92 Rhodora [Vol. 74 Chantransia meditierranea Levring (1942, p. 30, Figs. 1a- g), C. minutissima Reinsch (1874-5, p. 33, tab. V, Fig. 2a, tab. XI, Fig. 3a; not of other authors), C. trifila Buffham (1892, p. 24, pl. 3, Figs. 1-4), and Kylinia scapae Lyle (1929, p. 245, Figs. 6-7). A comparative examination of the type collections of these taxa will almost certainly show them to be conspecific with A. microscopica. The andro- phores described and illustrated by Lyle (1929) probably represent young, unelongated unicellular hairs. COLLECTIONS EXAMINED Types and Isotypes: DENMARK: Kattegat Channel, 17. vii. 1890, Rosenvinge (C, Rosenvinge 863, Algae marinae Danicae, type of Chantransia moniliformis Rosenvinge). ENGLAND: Torquay, 1845, Naegeli (L 940285 .. . 306, type of Audouinella microscopica (Naegeli) Woelkerling). Tor- quay, 1845, Naegeli (FH, No. 454, *Phykotheka Univer- salis", isotype of A. miscroscopica). Torquay 1845, Naegeli (NY, No. 454, *Phykotheka Universalis", isotype of A. mi- croscopica). GREENLAND: Holstenborg, 9. vii. 1895, Hanson (c, type of Chantransia collopoda (Rosenvinge) Rosen- vinge). PERU: La Punta, region of Callao, 25. i. 1907, Coker (NY, type of Acrochaetium catenulatum Howe). UNITED STATES: Black Rock, Sconticut Neck, New Bedford, Massa- chusetts, 25. vii. 1934, Jao (MICH, Woods Hole, No. 275, type of Acrochaetium compactum Jao). Norton Point, Martha’s Vineyard, Massachusetts, 3. viii. 1934, Jao (MICH, Woods Hole, No. 280 [not 274 as reported by Jao 1936, p. 240], type of Acrochaetium microfilum Jao). VIRGIN ISLANDS: St. Thomas (The Harbour), i. 1906, Boergesen (c, type of Acrochaetium crassipes (Boergesen) Boerge- sen). Other collections: DENMARK: Vorupor, NW coast of Jutland, 30. vii. 1929, Lund (C). MASSACHUSETTS: Cape Codder Point (Falmouth), 19. xi. 1969, Woelkerling (2292, personal collection). West Falmouth Harbor, 17. x. 1970, Woelkering (2826, personal collection). Woods Hole (Nobska Point), 4. ii. 1970, Woelkerling (2320, personal 1972] Audouinella — Woelkerling 93 collection). NOVA SCOTIA: Cranberry Cove, 12. ix. 1965, Edelstein (Nat. Res. Council. Herb., Halifax, No. 1867, as Acrochaetium microfilum Jao). Herring Cove, 18. i. 1966, Edelstein (Nat. Res. Council. Herb., Halifax, No. 2217, as Kylinia compacta (Jao) Papenfuss). Ketch Harbour, 7. ii. 1966, McLachlan & Edelstein (Nat. Res. Council. Herb., Halifax, No. 2244, as Kylinia collopoda (Rosenvinge) Pap- enfuss). Peggy’s Cove, 26. viii. 1965, Edelstein (Nat. Res. Council. Herb., Halifax, No. 2105, as K. collopoda). SUMMARY The relationship of Awudouinella microscopica to six close- ly related taxa has been investigated with particular ref- erence to the New England and adjacent coasts. The mor- phology of monosporangial and sexual plants in the type collection of A. microscopica is discussed and illustrated. Collections from the New England region previously re- ferred to Acrochaetium microfilum Jao, Kylinia collopoda (Rosenvinge) Kylin, K. compacta (Jao) Papenfuss, and K. moniliformis (Rosenvinge) Kylin have been found to rep- resent specimens of Audouinella microscopica. A compara- tive study of the type collections of these five taxa as well as Acrochaetium crassipes Boergesen and A. catenulatum Howe indicates that all taxa are conspecific with Audouinella microscopica. Chantransia meditierranea Levring, C. min- utissima Reinsch non. al., C. trifila Buffham, and Kylinia scapae Lyle are regarded as probable synonyms. ACKNOWLEDGMENTS Sincere thanks are due Dr. T. Edelstein (Atlantic Re- gional Laboratory, National Research Council, Halifax, Nova Scotia), Dr. I. Mackenzie Lamb (FH), Dr. Soren Lund (c), Dr. W. F. Prud’homme van Reine (L), Dr. Clark Rogerson (NY), and Dr. W. R. Taylor (MICH) for the loan of herbarium material. Miss Susan Heller kindly aided in the preparation of the drawings. This work was supported in part by National Science Foundation Grant GB-13250 to the Systematics-Ecology Program. 94 Rhodora [Vol. 74 Contribution No. 242 from the Systematics-Ecology Pro- gram, Marine Biological Laboratory. REFERENCES ABBOTT, I. A. 1962. Some Liagora-inhabitating species of Acrochae- tium. Occ. Pap. Bernice P. Bishop Mus. 23: 77-120. ABBOTT, I. A. 1968. An examination of the type specimens of some species of Acrochaetium (Rhodophyceae) Taxon 17: 518-520. BOERGESEN, F. 1909. Some new or little known West Indian Flor- ideae. Bot. Tidsskr. 30: 1-19. BOERGESEN, F. 1915. The marine algae of the Danish West Indies. Vol. II. Rhodophyceae. Dansk bot. Ark. 3: 1-80. BOERGESEN, F. 1937. Contributions to a south Indian marine algal flora. J. Indian bot. Soc. 16: 1-56, pl. 1. BUFFHAM, T. H. 1892. Chantransia trifila, a new marine alga. J. Quekett microsc. Club (Ser. 2) 5: 24-26. Drew, K. M. 1928. A revision of the genera Chantransia, Rhodo- chorton, and Acrochaetium, etc. Univ. Calif. Publs. Bot. 14(5) : 139-224, pl. 37-48. EDELSTEIN, T. and MCLACHLAN, J. 1966. Species of Acrochaetiwm and Kylinia new to North America. Br. phycol. Bull. 3: 37-41. EDELSTEIN, T., J. MCLACHLAN, and J. S. CRAIGIE. 1967. Investiga- tions of the marine algae of Nova Scotia. II. Species of Rho- dophyceae new or rare to Nova Scotia. Can. J. Bot. 45: 202, pl. 1-8. FELDMANN, J. 1962. The Rhodophyta order Acrochaetiales and its classification. Proc. IX Pacif. Sci. Congr. 4: 219-221. HAMEL, G. 1927. “Recherches sur les genres Acrochaetium Naeg. et Rhodochorton Naeg.” Dissertation. (Paris). HAMEL, G. 1928. Sur les generes Acrochaetium Naeg. et Rhodo- chorton Naeg. Revue algol. 3: 159-210. Hauck, F., and P. RICHTER. 1894. *Phykotheka Universalis”. X. Leipzig. HowE, M. A. 1914. The marine algae of Peru. Mem. Torrey bot. Club 15: 1-185, pl. 1-66. Jao, C. C. 1936. New Rhodophyceae from Woods Hole. Bull. Torrey Bot. Club 63: 237-257. KUETZING, F. T. 1849. “Species Algarum" (Leipzing). LANJOUW, J., and F. A. STAFLEU. 1964. Index herbariorum. Part 1. The herbaria of the world. Ed. 5. Regnum Vegetabile 31: 1-251. LEVRING, T. 1937. Zur kenntnis der algenflora der Norwegischen Westkueste. Acta Univ. Lund (N.F. Avd. 2) 33(8): 1-147. LEVRING, T. 1942. Meeresalgen aus dem Adriatischen meer, Sizilien, und dem golf von Nepal. K. fysiogr. Sallsk. Lund Forh. 12(3): 25-41. 1972] Audouinella — Woelkerling 95 LUND, S. 1942. Chantransia collopoda K. Rosenv., a new rhodo- phyceous alga of the Danish waters. Bot. Tidsskr. 46: 53-57. LvLE, L. 1929. Marine algae of some German warships in Scapa Flow and of the neighboring shores. J. Linn. Soc. (Bot). 48: 231-257. NAEGELI, C. 1861. Beitraege zur morphologie und systematik der Ceramiaceae. Sher. bayer. Akad. Wiss. 2: 297-415. REINSCH, P. F. 1874-5. "Contributions ad algologiam et Fungolo- giam". 1. Nuernbert. RosENVINGE, L. K. 1898. Deuxieme memorie sur les algues du Groenland. Meddr. Groenland 20: 1-125. RosENVINGE, L. K. 1909. The marine algae of Denmark. I. Rho- dophyceae 1. K. danske Vidensk. Selsk. Skr. Afd 7 Raekke 7(1): 1-151. ScHIFFNER, V. 1931. Neue und bemerkenswerte meeresalgen. Hed- wigia 71: 139-205. TAYLOR, W. R. 1937. “Marine Algae of the Northeastern Coast of North America”. (Ann Arbor). TAYLOR, W. R. 1957. “Marine Algae of the Northeastern Coast of North America", Ed. 2. (Ann Arbor). WEST, J. A. 1968. Morphology and reproduction of the red alga Achrochaetium pectinatum in culture. J. Phycol. 4: 89-99. WEST, J. A. 1969. The life histories of Rhodochorton purpureum and R. tenue in culture. J. Phycol. 5: 12-21. WOELKERLING, W. J. 1970. Acrochaetium botryocarpum (Harv.) J. Ag. (Rhodophyta) in Southern Australia. Br. phycol. J. 5(2): 159-171. WOELKERLING, W. J. 1971. Morphology and Taxonomy of the Au- douinella complex (Rhodophyta) in Southern Australia. Aust. J. Bot., Suppl. Ser., Suppl. 1(1971): 1-91. DEPARTMENT OF BOTANY UNIVERSITY OF WISCONSIN MADISON, WISCONSIN 53706 [Vol. 74 Rhodora 96 ‘ON 'extj 9SOY} JO suorjoe[[oo edAj Əy} Jo uoryeuruexo UB uo peseq sr suonv[ndod UdAVS jsig ƏY} 10g BA, '"(IL6I) Sumay -POM Wolf €jep Sjuoso1do4 (j'ON :peururexo suonoo[[po pueru MƏN JE Worl v3ep Jo ejrsoduioo e sQuoseudoi g "i L- "OT-G ZCL "OT-g ZI- "0pI-0c (ț-)g-1 suone[ndoq uvl[eijsny 6 "-p "GI-9 eI "TI-g "or-eg "00T-0c p- suone[ndoq pue[gun[^'N $8 "1-9 IGI- ZI TPI-L "II-L 0GT-OF 8-3 «SINUOAITINON,, 4 1i L-G VOT-L cr UeI-9 "1-9 "002-0p (2) I .NANOIMOOSOUDIN,, 9 ""g-p ig -9 I US Ig -g "0p -0c reg «INQYILAOWOIN,. € "L-G "S -9 cr 16 -G Wy, -Q "09 -OF (c) I «4SHdISSVHO. P "-G "S -9 cr "or-e IQ -G "0G -0c (p-) &(-Z) «IWüLOVdWOO. $8 "g-), IGT-8 yc "ep-0T "6 -L "00z-0c g «WAdOdJOTTIOD, z "1-9 "TI-6 I VOT-L TI-A VOPL-OF I «NOLVIOANALVO, I UjplA ujsueT Oey ujsue] YPM 4uST9H SoxXy 9UIeN ‘ON o1odg a10ds M/A ee) 1199 pər Jo ‘ON X H'Id WOO VOIdOOSOOIW V AO SNOLLV'I1dOd NO VLVd 'IVOIDO'IOHdHONW LI W'ISV.L NOMENCLATURAL CLARIFICATION OF TWO SPECIES OF VERBESINA (COMPOSITAE) ENDEMIC TO FLORIDA: JAMES R. COLEMAN Two species of Verbesina are evidently endemic to Flor- ida. One of these is restricted to a few coastal counties of the Apalachicola region of western Florida whereas the second has been collected from several counties in north- eastern Florida. The species are disjunct by approximately 150 miles and are readily distinguished from each other by several morphological features. Most notably, the eastern species has decurrent leaves which form winged stems and has rayed heads whereas the western species has wingless stems and rayless heads. I have examined approximately 50 collections of these species and have observed no excep- tions to these differences. In 1822 Nuttall was the first to describe the eastern spe- cies, naming it Actinomeris pauciflora and in 1841 he elab- orated on his original description. Actinomeris pauciflora was described as having decurrent leaves and 3-4 rays. The description was based on a collection made by Ware. I have been unable to locate this specimen and do not know whether it still exists. However, there is no doubt as to the species being described. In 1872 Chapman applied the name A. pauciflora Nutt. to a coastal species from western Florida. The species was described as having wingless stems and the florets as hav- ing “the marginal ones abortive.” Chapman’s lucid descrip- tion leaves no doubt that he was erroneously applying the name A. pauciflora Nutt. to the species from the Apalach- icola region. In 1878 Chapman, evidently still unaware of his error, redescribed the eastern species as new, giving it the name Actinomeris heterophylla. He described A. heterophylla as having winged stems, 5-10 rays and as oc- "This investigation was supported by National Science Foundation Grant GB-13923. 97 98 Rhodora [Vol. 74 curring in eastern Florida. I have examined the type and it clearly belongs to the eastern species. The next event in the nomenclatural history of these spe- cies occurred in 1883 when Gray transferred the eastern species to Verbesina forming the combination V. hetero- phylla (Chapm.) A. Gray. Although no description was provided, Gray did list Actinomeris heterophylla Chapm. as asynonym. The description Gray provided in his Synoptical Flora (1884) clearly reveals that he was referring to the eastern species. In his 1883 publication Gray also trans- ferred A. pauciflora Nutt. to Verbesina. However, since the specific epithet was preoccupied by a Mexican Verbesina described by Hemsley in 1881, he formed the new name Verbesina warei A. Gray and gave Actinomeris pauciflora Nutt. as a synonym. Gray’s description of Verbesina waret in his Synoptical Flora is confusing in that he described the stems as wingless but described the heads, in quotes, as being 3-4 rayed. He stated that the species occurs near the coast but did not specify the eastern or western coast. Col- lections by Ware and by Chapman were cited; however, Chapman collected both species and, as far as I am aware, Ware collected only the eastern. There is, therefore, serious doubt as to whether Gray was applying the name Verbesina warei to the eastern or western species. These species had been very infrequently collected prior to 1884 and it is pos- sible that Gray never saw specimens of one or the other or either species. It appears probable that Gray compiled his description of V. warei largely or entirely from Nuttall’s and Chapman’s descriptions of Actinomeris pauciflora Nutt. It should be recalled that Chapman’s description was ac- tually of the eastern species. Were this assumption true, it could account for the mixing of characteristics of both spe- cies in Gray's description of Verbesina warei, for his use of quotes in referring to the rayed condition and to his fail- ure to indicate whether the species occurred on the east or west coast as the two descriptions conflict on this point. In citing the collections of Ware and Chapman, Gray was probably again simply following Nuttall and Chapman. 1972] Verbesina — Coleman 99 In their revision of the genus Verbesina, Robinson and Greenman (1899) applied the name Verbesina heterophylla A. Gray to the eastern species and listed Actinomeris heter- ophylla Chapm. in synonymy. The western species was called Verbesina warei A. Gray and Actinomeris pauciflora Nutt. was given as a synonym. Small (1903), in his Flora of the Southeastern United States, also called the eastern species Verbesina hetero- phylla (Chapm.) A. Gray, but formed a new combination, Verbesina pauciflora (Nutt.) Small, for the western species and listed both Actinomeris pauciflora Nutt. and Verbesina warei A. Gray in synonymy. Small also was evidently un- aware that Nuttall’s name was originally applied to the eastern species and was either unaware of, or ignored, the fact that the combination Verbesina pauciflora was pre- occupied. Alexander, in treating these species in Small’s Manual of the Southeastern United States (1933), transferred both to the genus Pterophyton, forming the new combination P. pauciflorum (Nutt.) Alex. for the western species and P. heterophyllum (Chapm.) Alex. for the eastern species. The genus Pterophyton Cass. had been reduced to a section of Verbesina by Torrey and Gray in 1842. This reduction appears justified when the genus Verbesina is considered in its entirety rather than on a regional basis. The question arises as to the correct names for the two species under consideration. Although the earliest binom- ial applied to the eastern species is Actinomeris pauciflora Nutt. (1822), the epithet pauciflora cannot be used in com- bination with Verbesina since the combination V. pauci- flora is preoccupied by Hemsley’s Mexican species. The second oldest binomial given to this species is Actinomeris heterophylla Chapm. (1878), the specific epithet of which was transferred by Gray to form the correct name for the eastern species, Verbesina heterophylla (Chapm.) A. Gray. Although Verbesina warei A. Gray has been used fairly commonly for the western species, Gray did give Actinom- eris pauciflora Nutt. as a synonym in making the transfer. 100 Rhodora [Vol. 74 The name Verbesina warei A. Gray is, therefore, associated with the type of Actinomeris pauciflora which belongs to the eastern species. Verbesina warei must then be con- sidered a synonym of V. heterophylla (Nutt.) A. Gray. Verbesina heterophylla (Chapm.) A. Gray, Proc. Amer. Acad. 19: 12, 1883. Actinomeris pauciflora Nutt., Sill. Journ. 5: 301, 1822; Trans. Amer. Phil. Soc. 7: 364, 1841. Type: East Florida, Ware s.n. (location unknown). Actinomeris heterophylla Chapm. in Coult., Bot. Gaz. 3: 6, 1878. Type: East Florida, Chapman s.n., 1871. (NY!) Verbesina warei A. Gray, Proc. Amer. Acad. 19: 12, 1883. nom. nov. for A. pauciflora Nutt., not Verbesina pauciflora Hemsl. 1881. Pterophyton heterophyllum (Chapm.) Alex. in Small, Man. S. E. Fl. 1444, 1933. The earliest description of the western species was given by Chapman in 1872; however, he failed to recognize it as a new species and misidentified it as Actinomeris pauci- flora Nutt. Probably the most commonly used name for this species is Verbesina warei A. Gray. However, this name belongs to the eastern species. Since the western species has never been given a valid name, it is described as a new species. Verbesina chapmanii J. R. Coleman, sp. nov. Perennis ca 0.5-1.1 m alta; caulis simplex non alatus; folia opposita vel alterna, sessilia vel subsessilia, lanceolata, elliptica vel lata, scabra; capitula 1-5 terminalia pedunculata, plerum- que 1-2 cm lata, discoidea; involucra 2-seriata, ca 8-12 mm elliptico-oblonga, plerumque 5-9 cm longa, usque ad 3.2 cm alta; flores lutei; achaenia alata, oblongo-obovata, plerum- que 6-8 mm longa, 4-5 mm lata, glabra vel subglabra, epap- posa vel pappi setis usque ad ca 1 mm longis praedita. Perennial herbs mostly 0.5-1.1 m tall with heavy rhi- zomes; stems numerous, unbranched below the inflores- cence, wingless, hispidulous near the heads, otherwise glabrous or subglabrous; leaves alternate or opposite, be- 1972] Verbesina — Coleman 101 coming bractiform above, rigid, sessile or subsessile, lance- olate, elliptical or elliptic-oblong, serrulate to subentire, obtuse or less frequently acute, to ca 12 cm long, commonly 5-9 cm long, to ca 3.2 cm wide, scabrous; inflorescence solitary or to 5-headed; heads mostly 1-2 cm wide, discoid; phyllaries 2-seriate, ca 8-12 cm tall; florets yellow, mostly 12-14 mm tall; paleae ca 8-10 mm tall, mostly red-purple apically; achenes mostly red-purple, oblong-obovate, mostly 6-8 mm long, 4-5 mm wide, glabrous or sparingly hirtel- lous apically, lateral wings to ca 1 mm wide, continuous about the crown to form a shallow cup, pappus of 2 short horns or lacking. Type: open prairie-like pine-cypress savanna, 3 mi N of Orange, Liberty Co, Florida, McDaniel 4468 (GH!) DEPARTMENT OF BOTANY THE UNIVERSITY OF GEORGIA ATHENS, GEORGIA 30601 LITERATURE CITED CHAPMAN, A. W. 1872. Flora of the southern United States. Ivi- son, Blakeman, Taylor and Co., N. Y. 621 pp. GRAY, A. 1884. Synoptical Flora of North America. Vol. I. Part II. Smithsonian Institution, Washington, D. C. HEMSLEY, W. B. 1881. Biologia centrali-americana. Bot. Vol. II. London. SMALL, J. K. 1903. Flora of the southeastern United States. Pub- lished by the author, N. Y. 1370 pp. . 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel Hill. 1554 pp. ROBINSON, B. L. and J. M. GREENMAN. 1899. A synopsis of the genus Verbesina, with an analytical key to the species. Proc. Amer. Acad. 34: 534-5064. TORREY, J. and A. GRAY. 1842. A Flora of North America Vol. 2 pt. 2. Wiley and Putnam, N. Y. SOMATIC CHROMOSOME NUMBERS FOR SOME ASTERACEAE MILOSLAV KOVANDA Few plant groups can claim the attention of botanists that the American Asteraceae have been enjoying in recent years. In addition to thorough monographic studies an un- precedented series of chromosome number reports has ap- peared, and it may seem trivial to publish additional counts for a handful of miscellaneous taxa. Yet the need for re- peated chromosome sampling cannot be stressed enough. Previous studies have revealed that intraspecific polyploidy is widespread, and chromosome counts from as many popu- lations as possible throughout the range of each species are required to gain an idea of the nature and significance of this diversity. Many of the taxa discussed below are wide- ranging, polymorphous entities that have been counted but once or twice, making the "cytofloristic" approach impera- tive. The counts for Artemisia saxicola and Haplopappus lyalli are believed to be the first to be published for these species. A new ploidy level is established for Chrysopsis fulcrata. MATERIALS AND METHODS All chromosome counts were obtained from squashes of root-tips of young seedlings grown in the laboratory. Achenes were collected in the field in the late summer and fall of 1970. All germinated without special treatment in 2 to 9 days when placed on moist filter paper in Petri dishes and maintained at room temperature. Excised root-tips were pretreated in a saturated solution of 8-hydroxyquinoline for 3 hours, fixed in 1 : 3 acetic alcohol, and stained in lacto- propionic orceine (for details, see Kovanda 1970). The counts were verified in five or more plantlings. Voucher specimens were collected for all plants sampled and will be deposited at the Herbarium of the National Museum in Prague (PR). Duplicates, when available, will be deposited 102 1972] Kovanda — Asteraceae 103 in the United States National Herbarium (US). A portion of the material examined has been planted in the experi- mental plot of the Botanical Institute, Czechoslovak Acad- emy of Sciences, Prague, for future observations. All col- lections were made by Kovanda or by Kovanda and S. G. Shetler. OBSERVATIONS TRIBE VERNONIEAE 1. Elephantopus carolinianus Willd. Maryland, Montgomery County: thicket along Chesapeake & Ohio Canal near Glen Echo, vicinity of Washington, D.C., Kovanda 3558. 2n = 22. This species recently has been examined cytologically from two widely separated areas: Virginia (Baldwin & Speese, 1955) and Texas (Lewis et al., 1962). In both cases, the authors report the same chromosome numbers, 2n — 22 and n = 11, respectively. The entire genus seems to have the base number x — 11, and all species counted heretofore have proved to be diploid. The count 2n = 44 for E. tomen- tosus L., ascribed by Fedorov et al. (1969) to S. B. Jones (1966), is in error, Jones having counted n — 11 and listed the earlier count of 2n = 22 made by Baldwin & Speese. TRIBE EUPATORIEAE 2. Eupatorium coelestinum L. Maryland, Montgomery County: dry woodland near Cabin John Bridge, vicinity of Washington, D.C., Kovanda 3529. 2n — 20. This number was reported previously by Grant (1953) for material from Pennsylvania, Kentucky, and Tennessee. By its chromosome morphology this species differs from all other species of Hupatorium having z = 10 and would be best referred perhaps to the genus Conocliniwm DC. (see Grant 1953). 9. Liatris punctata Hook. Colorado, Larimer County: Rist Canyon, grassy roadside about 8 mi. W. of Bellevue, 6800 ft., Kovanda & Shetler 3169. 2n = 40. 104 Rhodora [Vol. 74 In her study of the genus Liatris, Gaiser (1950) has dem- onstrated that two chromosome races exist in this species, which she was able to identify with previously described infraspecific taxa. Diploids (2n — 20) have been referred to var. nebraskana Gaiser, and the tetraploid level (2n = 40) is represented in var. punctata (var. typica Gaiser) and var. mexicana Gaiser. The two cytotypes are not al- ways easily recognizable on a morphological basis but show, in addition to marked differences in the beginning of anthesis, also a clear geographical separation. Tetraploids seem to occur throughout most of the extensive range of the species (with var. mexicana, being limited to its south- ern part) while the diploid variety is obviously localized. The Rist Canyon population certainly falls within the typ- ical variety but approaches somewhat var. nebraskana by its leaves which are almost entirely devoid of cilia. TRIBE ASTEREAE 4. Aster occidentalis (Nutt.) T. & G. Colorado, Larimer County: brushy hillside north of Stove Prairie School, S. of Poudre River, 6500 ft, Kovanda & Shetler 3183. 2n — 10. The count reported here is consistent with those pub- lished by Huziwara (1958, 1959) and Solbrig et al. (1969). The Western Aster is one of the few aster species not in the x = 9 series. The number x = 8, likewise x = 5 in A. exilis Ell. and A. foliaceus Lindl., and x — 13 in A. chil- ensis Nees, is clearly secondary (Solbrig et al., 1969). Other species with « — 8 include, for instance, A. adscendens Lindl., A. ericoides L., A. hirtifolius Blake, A. lateriflorus (L.) Britt., and A. puniceus L.; most of them are tetra- ploid, with 2n = 32. It is interesting to note that base num- bers other than x — 9 are not known to occur in species native to the Old World, which supports the contention that the center of origin of this huge genus is America. 5. Chrysopsis fulcrata Greene Colorado, Clear Creek County: abandoned campground in Clear Creek valley, about 12 mi. south of Georgetown, 11,440 1972] Kovanda — Asteraceae 105 ft., Kovanda & Shetler 3331. 2n — 18. Colorado, Larimer County: Rist Canyon, grassy roadside about 8 mi. west of Bellevue, 6800 ft. Kovanda & Shetler 3172. 2n = 36. Counts by Raven et al. (1960), Turner et al. (1962), Sol- brig et al. (1964), Chuksanova et al. (1968), and Solbrig et al. (1969) indicate that both diploids and tetraploids are present in the Chrysopsis villosa species aggregate. In C. fulcrata, a segregate defined mainly by leaf-like bracts sub- tending the heads, glandular phyllaries and sessile leaves, only the diploid level (» = 9) has been sampled (Solbrig et al. 1964, 1969). An examination of material from Rist Canyon clearly showed 36 somatic chromosomes. The col- lection was compared with Greene’s type material of C. ful- crata in the U. S. National Herbarium, and no specific dif- ferences could be seen except for the leaves. These are sometimes slightly tapering at the base in my plants but are distinctly semi-amplexicaul in the type specimen, col- lected in the Organ Mountains, New Mexico. In the Clear Creek diploid this tapering is even more conspicuous, mark- ing an intergradation with C. hispida (Hook.) DC. and C. viscida (A. Gray) Greene. The taxonomy of the entire com- plex is not very well understood, and there appears to be little correlation between the cytological findings and mor- phological segregation of the several species. 6. Chrysothamnus nauseosus (Pallas) Britt. subsp. nau- seosus Wyoming, Albany County: stony roadside just east of Cen- tennial, 8000 ft., Kovanda & Shetler 3205. 2n = 18. The Rabbitbrush was the subject of a detailed cytotax- onomic study by Anderson (1966) who demonstrated that polyploidy is rare in this genus. He found only the diploid number 2n — 18 in many plants of C. nauseosus divided among 12 subspecies. The same number has been counted by Raven et al. (1960) and Solbrig et al. (1964). The count reported here is only a further proof (if one is needed) that the astonishing morphological variation shown by this spe- cies cannot be credited to polyploidy. 106 Rhodora [Vol. 74 7. Erigeron speciosus (Lindl.) DC. Wyoming, Teton County: Teton Pass, 9000 ft., Kovanda & Shetler 3252. 2n = 18. Previous counts seem to indicate that this species com- prises diploid and tetraploid cytotypes. Ferri (1961), Zhu- kova (1964, 1967) and Huziwara (1965) give, alternatively, ^ — 9 or 2n — 18, contrasting with 2» — 36 counted by Vilmorin and Chopinet (1954). Unfortunately, all these counts were made on cultivated material of uncertain origin, and it remains to be seen whether both the races occur in natural habitats. The Teton Pass population proved to be diploid and appears referable to var. macranthus (Nutt.) Cronquist, which has been counted as E. macranthus Nutt. by Bergman (1942) and Zhukova (1964). Both counts were 2n — 18, too, but the sources of the material were not given. 8. Grindelia aphanactis Rydb. New Mexico, Rio Arriba County: Rio Grande Canyon, about 5 mi. NE. of San Juan Pueblo, 5800 ft., Kovanda 3404. 2n — 24. The only previous count for this species was n — 12 by Raven et al. (1960) on material from Arizona. Grindelia Willd. is a natural genus, and, chromosomally, a very homo- geneous one, with the base number only x — 6. The majority of species seem to be diploid, with 2n = 12 (see also Whit- aker and Steyermark 1935, Dunford 1964, 1970). 9. Haplopappus lyallii A. Gray Colorado, Clear Creek County: Loveland Pass, 11,900 ft., Kovanda & Shetler 3382. 2n — 18. No previous cytological work seems to have been done on this tiny goldenweed. Two members of the section Tones- tus, H. eximius Hall and H. peirsonii (Keck) Howell, have been shown to have 2n — 18 and ca. 90, respectively (Steb- bins in Howell 1950), which would indicate that the base number for that group is 9. But considering the variety of chromosome numbers encountered in other species (see e.g., Raven et al. 1960, Solbrig et al. 1964, 1969), any such statement would be premature. In the genus Haplopappus, 1972] Kovanda — Asteraceae 107 as broadly conceived by Hall (1925), « = 9 as a basic chromosome number is not uncommon; however, it is puz- zling to discover that it is largely associated with the shrubby groups of the southwestern deserts, such as sec- tions Ericameria and Stenotopsis. The only other herba- ceous species hitherto known to have the base number « = 9 is H. clementis (Rydb.) S. F. Blake of the section Pyr- rocoma. The genus Tonestus was proposed by Nelson (1904) to accommodate low perennial herbs from the Rocky Moun- tains area and “to bring together under one name these species [H. laceratus Henderson, H. lyallii A. Gray, and H. pygmaeus T. & G.] which are allied by habit and mor- phological characters to each other and are aberrant in any recognized genus or genera in which they can be placed." When more cytological information is available, it may be- come necessary to reconsider the generic and sectional limits. 10. Solidago flexicaulis L. Maryland, Harford County: woodland along Susquehanna River below Deer Creek, between Schweers Landing and Lapidum, Kovanda & Shetler 3638. 2n — 18. The same diploid number has been found in plants from several Canadian localities (see Beaudry and Chabot 1959, Kapoor and Beaudry 1966, Kapoor 1970). A tetraploid cytotype was reported from Michigan (Solbrig et al. 1964). 11. Solidago multiradiata Ait. Colorado, Clear Creek County: Clear Creek valley, about 12 mi. west of Georgetown, 11,400 ft., Kovanda & Shetler 3872. 2n — 18. This arctic-alpine species was once thought to be diploid (Beaudry and Chabot 1959, Beaudry 1963) but has recently been shown to have tetraploid races, 2n — 36, in western Canada and Alaska (Taylor 1967, Johnson & Packer 1968, Packer 1968). The count recorded here is the first made on material from the Southern Rockies where the species reaches its southernmost limit. The collection is var. scopu- lorum A. Gray. 108 Rhodora [Vol. 74 12. Solidago sempervirens L. Delaware, Sussex County: salt marsh about 2 mi. S. of Rehoboth Beach, Kovanda 3589. 2n — 18. Several counts have been made on this halophyte, reveal- ing the presence of diploid and tetraploid levels. However, these races do not correspond with the two major segments recognized taxonomically within S. sempervirens. The southern entity, var. mexicana (L.) Fern., was first re- ported to be diploid but was later found to have also 2n = 36, and the same holds for the more northern var. semper- virens (Goodwin 1937, Beaudry and Chabot 1959, Beaudry 1963, Kapoor 1970). Likewise, the geographical distribu- tion of the cytotypes does not seem to form a definite pattern, but there are too few counts to allow any generali- zation. Their morphological differences and ecological pref- erences, if any, also await future research. TRIBE HELIANTHEAE 13. Rudbeckia hirta L. Maryland, Frederick County: Catoctin Mountains, old field near cabin of David Scott, west of Thurmont, Kovanda 2896. 2n = 38. Colorado, El Paso County: Pikes Peak, 9000-foot level, streamside, Kovanda & Shetler 3398. 2n — 38. Rudbeckia hirta is an extremely variable species, and many local variants have been given varietal or specific rank. Perdue (1957) greatly reduced their number by recognizing five geographical varieties: var. hirta, var. pulcherrima Farwell, var. corymbifera Fernald, var. angus- tifolia (T. V. Moore) Perdue, and var. floridana (T. V. Moore) Perdue. Most controversial perhaps is the treat- ment of var. pulcherrima, which was first described as R. serotina Nutt. and to which both the populations sampled by the present author belong. Fernald (1948) considered it a western entity which had spread eastwards, becoming thoroughly naturalized throughout most of the United States, and advocated its specific status. His ideas, however, received little support from later workers (Perdue 1957, Core 1962). In contrast to its intricate morphological struc- 1972] Kovanda — Asteraceae 109 ture, the species complex appears remarkably uniform in cytology, 2n = 38 being the only chromosome number so far obtained (Battaglia 1946, 1947, Perdue 1959). The same chromosome number has also been reported for R. serotina (Battaglia 1947, Mulligan 1959). 14. Viguiera multiflora (Nutt.) S. F. Blake Utah, Daggett County: Uinta Mountains, between Cub Creek and Y Creek, ca. 6 mi. W. of Red Canyon Road, 8500 ft., Kovanda & Shetler 3275. 2n = 16. Two earlier counts for this species give the gametic chromosome number as n = 8 (Heiser and Smith 1955, Heiser 1963). TRIBE HELENIEAE 15. Hymenoxys richardsonii (Hook.) Cockerell New Mexico, Santa Fe County: 2 mi. NE. of Santa Fe, 7200 ft., Kovanda 3428. 2n — 30. The chromosome number of this species has been deter- mined repeatedly as either n — 15 or 2n — 30 (Speese and Baldwin 1952, Strother 1966). The count 2n — 28, made by Taylor and Brockman (1966) on a collection from Saskatch- ewan, suggest that there are two chromosome races in this species whose morphology and geographical distribution re- quire further study. Material examined by the present au- thor belongs to var. floribunda (A. Gray) Parker and was collected in or near the type locality (“rocky hills, as well as plains and creek bottoms, around Sante Fe", A. Gray in Mem. Amer. Acad., n.s. 4: 101, 1849). 16. Pericome caudata A. Gray New Mexico, Santa Fe County: Sangre de Cristo Moun- tains, along the Chamisa Trail, 7600 ft, Kovanda 3426. 2n = 36. Pericome caudata has recently been counted by Turner and Flyr (1966) and Powell (1968). Both obtained a game- tic chromosome number of » — 18. Raven and Kyhos (1961) have found the same number in the closely related P. glandulosa Goodm. It would appear that the small genus Pericome has the base number x = 9. 110 Rhodora [Vol. 74 TRIBE ANTHEMIDEAE 17. Artemisia borealis Pallas Colorado, Clear Creek County: Loveland Pass, 11,990 ft., Kovanda & Shetler 3376. 2n = 18. Artemisia borealis is taxonomically complex in the Old and New World, and attempts to subdivide it into more natural entities have not been very successful. Diploids and tetraploids, based on x — 9, are known to exist in this group. Both these races have been shown to occur in North Amer- ica, but their disposition is not at all clear. Diploids (2n = 18) have been discovered in Greenland (Jörgensen et al. 1958), northern Quebec (Hedberg 1967), and Alberta (Packer in Johnson and Packer 1968), but tetraploids (2n = 36) have so far been found only in Alaska (Johnson and Packer 1968). The Loveland Pass record is important because the count was made on material from the most southern portion of the species range. The plants examined have basal leaves three times pinnatifid, densely hairy on both surfaces; stems almost totally glabrous; heads erect, arranged in a narrow, spike-like inflorescence, somewhat interrupted in the lower half; phyllaries villous; achenes silky-hairy. 18. Artemisia frigida Willd. Colorado, Park County: 2 mi. N. of Fairplay on Route 9, 9900 ft., Kovanda & Shetler 3379. 2n = 18. This number was obtained previously by Lóve and Lóve (1964) on plants from Manitoba and by Knaben (1968) on plants from Alaska. Kawatani and Ohno (1964) give 2n — 18 for garden material of unspecified origin. 19. Artemisia saxicola Rydb. Colorado, Grand County: alpine tundra above Berthoud Pass, 12,400 ft., Kovanda & Shetler 3324. 2n — 18. Colorado, Summit County: near Pass Lake, just S. of Loveland Pass, 11,000 ft., Kovanda & Shetler 3367. 2» = 18. These are probably the first counts for the Rocky Moun- tains segregate of A. arctica Less. This species, ranging in northeastern Asia and northwestern North America and 1972] Kovanda — Asteraceae 111 itself a segregate of the European A. norvegica Fries, has been counted many times on Asiatic material and is known to have two chromosome numbers, 2n — 18 and 36. (Kita- mura 1957, Sokolovskaya 1963, Kawatani and Ohno 1964, Zhukova 1964, 1965, 1967). Only two records are available from North America (Johnson and Packer 1968, Taylor and Mulligan 1968). Both counts are tetraploid and were made on plants from Alaska and the Queen Charlotte Islands, respectively. It is extremely interesting to discover that the Rocky Mountain populations are diploid and, there- fore, more primitive than the northern A. arctica. This, combined with the geographical isolation and a degree of morphological differentiation, would justify the specific rank assigned to this race by Rydberg but disregarded by most later students. TRIBE SENECIONEAE 20. Senecio longilobus Benth. New Mexico, Santa Fe County: near Bishop's Lodge, N. of Santa Fe, 6900 ft., Kovanda 3425. 2n — 40. This number conforms to the findings of Jackson (1959) and Ornduff et al. (1967) who determined n = 20 on plants from New Mexico and Texas. Powell and Turner (1963) have reported the same chromosome number for Mexican material under the name of S. filifolius Nutt. The section Suffruticosi, confined to the western United States and northern Mexico, appears chromosomally uniform. All spe- cies hitherto examined have proved to have » — 20 (Stout- amire and Beaman 1960; Turner, Ellison and King 1961; Ornduff et al. 1963, 1967), and Ornduff et al. (1963) have proposed that the base number is either x = 10 or x = 20. ACKNOWLEDGMENTS This work was carried out during my appointment as a Visiting Research Associate in the Department of Botany at the Smithsonian Institution, Washington, D.C., 1970-71. It is my pleasant duty to express my appreciation to S. G. Shetler, Associate Curator, Department of Botany, for ar- ranging the Rocky Mountain trip, making my stay in the 112 Rhodora [Vol. 74 United States an eventful and interesting one, and for many stimulating ideas. My sincere thanks are due to W. A. Weber, University of Colorado, Boulder, who kindly checked or made identifications of my collections from the Colorado Rockies; however, I accept full responsibility for the tax- onomic opinions expressed here. The fieldwork was sup- ported in part by the Smithsonian Research Foundation (Sg-0653061/C2). LITERATURE CITED ANDERSON, L. C. 1966. Cytotaxonomic studies in Chrysothamnus (Astereae, Compositae). Amer. J. Bot. 53(2): 204-212. BALDWIN, J. T. and B. M. SPEESE. 1955. Chromosomes of Elephan- topus and Pluchea in the range of Gray's Manual of Botany. Amer, J. Bot. 4(2): 123-125. BATTAGLIA, E. 1946. Ricerche cariologiche ed embriologiche sul genere Rudbeckia (Asteraceae). 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Senecioneae. Amcr. J. Bot. 54(2): 205-213. PACKER, J. G. in A. LóvE. 1968. IOPB Chromosome number reports XVII. Taxon 17(3): 285-288. PERDUE, R. E. 1957. Synopsis of Rudbeckia subgenus Rudbeckia. Rhodora 59: 293-299. 1959. The somatic chromosomes of Rudbeckia and related genera of the Compositae. Contr. Gray Herb. 185: 129- 162, PowELL, A. M. 1968. Chromosome numbers in Perityle and related genera (Peritylanae-Compositae). Amer. J. Bot. 55(7): 820-828. and B. L. TURNER. 1963. Chromosome numbers in the Compositae. VII. Additional species from the southwestern United States and Mexico. Madroño 17(4): 128-140. 1972] Kovanda — Asteraceae 115 RAVEN, P. H. and D. W. Kynos. 1961. Chromosome numbers in Compositae. II. Heleneae. Amer. J. Bot. 48(9): 842-850. , O. T. SoLBRIG, D. W. KvHos, and R. Snow. 1960. Chromosome numbers in Compositae. I. Astereae. Amer. J. Bot. 47 (2): 124-132. SOKOLOVSKAYA, A. P. 1963. Geograficheskoe rasprostraneniye poli- ploidnykh vidov rasteniy. 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Queen’s Printer, Ottawa. TURNER, B. L. and D. FLYR. 1966. Chromosome numbers in the Compositae. X. North American species. Amer. J. Bot. 53(1): 24-33. , W. L. ELLISON and R. M. KiNG. 1961. Chromosome numbers in the Compositae. IV. North American species, with phyletic interpretations. Amer. J. Bot. 48(3): 216-223. , M. POWELL, and R. M. Kine. 1962. Chromosome numbers in the Compositae. VI. Additional Mexican and Guate- malan species. Rhodora 64: 251-271. VILMORIN, R., DE and R. CHOPINET. 1954. Contribution à l'étude des nombres chromosomiques des races et variétés cultivées chez nos plantes ornamentales. Caryologia 6, Suppl.: 1006-1015. WHITAKER, T. W. and J. A. STEYERMARK. 1935. Cytological aspects of Grindelia species. Bull. Torrey Bot. Club 62(2): 69-73. 116 Rhodora [Vol. 74 Zuukova, P. G. 1964. Kariologia nekotorykh vidov Compositae v Polyarno-Alpijskom botanicheskom sadu. Bot. Zhurn. 49(11): 1656-1659. 1965. Kariologischeskaya kharakteristika neko- torykh rastenyi Chukotskogo poluostrova. Bot. Zhurm. 50(7): 1001-1004. 1967. Kariologia nekotorykh rastenyi, kultivirue- mykh v Polyarno-Alpijskom botanicheskom sadu. In Pereselenie rastenyi na polyarnyj sever II, A. AVRORIN, ed., Leningrad. p. 139-149. INSTITUTE OF BOTANY CZECHOSLOVAK ACADEMY OF SCIENCES PRAGUE INTERACTIONS BETWEEN APIS MELLIFERA (HYMENOPTERA: APIDAE) AND TRIBULUS CISTOIDES (ZYGOPHYLLACEAE) DANIEL F. AUSTIN In southern Florida a common weed of disturbed areas is Tribulus cistoides L. (Burr Nut or Puncture Vine). A frequent visitor to this plant is the Honey Bee, Apis melli- fera L. Little is known about the reproductive biology of this weedy plant (Porter, 1971; pers. comm.), but observa- tions in Florida suggest some interesting interactions be- tween Apis and Tribulus. According to Grant (1950) there is a division of labor among field bees of Apis mellifera. Some of the workers carry water; others gather pollen, nectar or propolis, while others search for new food sources. The constancy of an individual to her job and to a particular plant species has been amply verified by numerous authors and summarized by Grant (op. cit.). While I have observed Hemiptera, Lepidoptera, Diptera, and other Hymenoptera on Tribulus in Florida, the most frequent visitors observed on the plants near Boca Raton, Palm Beach County, are Apis mellifera. There is a marked behavioral division among the Honey Bee individuals visit- ing the plants. Most of the Honey Bees visiting the flowers approach in the “normal” bee fashion described by Meeuse (1961) and Fagri & Pijl (1966). Instead of landing on the stamens and stigmas as they should, however, they circle around the flower and land on the outside of the calyx and corolla (Figs. 1-4). After landing they separate two petals with their front legs, insert their tongues, and sip nectar (Fig. 3). They continue this procedure around the flower until they have collected nectar between all the petals. Once a flower has been utilized, the bees fly to another and repeat the process. Several flowers are usually visited before the bee disappears. 117 118 Rhodora [Vol. 74 Figure 1-4. Apis mellifera visiting the flowers of Tribulus cis- toides from the back. Note the tongue of the bee in Fig. 3. Other Honey Bees approach and land within the corolla in the *normal" manner. In so doing they position their bodies over the stigma and stamens (Figs. 5-7). Normally they insert their tongue between the petals and the nectary (Fig. 7), sip nectar, and turn around the flower to drink nectar from other nectaries. They dust much of their body with pollen as they turn (Figs. 5-6). After the nectar has been collected, they often brush the stamens with their front legs to gather pollen. After nectar has been taken and pollen gathered with the front legs, the bees may fly to another flower to repeat the process. It is common, however, for an individual to brush pollen, fly off and hover in front of the flower, re- turn to brush more pollen, and hover again. This may be done three or four times before the process is repeated at another flower. 1972] Tribulus cistoides — Austin 119 | " t LÀ Tn ; Figure 5-8. Bees visiting the flowers of T'ribulus cistoides from the front. Fig. 5-7. Apis mellifera. Fig. 8. Chloralictus. Those individuals which exhibit this hovering behavior usually have their pollen baskets full. The bees with pollen in the pollen baskets have little or no pollen adhering to the hairs of their legs and body. Bees which have no pollen in their baskets rarely hover, and their bodies and legs usually are dusted with pollen. The individuals illustrated here (Figs. 5-7) were photographed between 3:40 and 4:40 p.m. in December. Although their bodies are dusted with pollen, the pollen baskets are empty. There has been a seasonal shift in the labor of the in- dividuals visiting the plants. From September until Decem- ber about 18 out of 20 Honey Bees gathered nectar. None of the nectar gathering bees were ever seen collecting pollen. In February the visitation had dropped from about 20 bees to about 4 bees per hour. All of those bees seen in Febru- ary were gathering pollen. 120 Rhodora [Vol. 74 Associated with the reduction of visits by Honey Bees was an increase of visits by other insects. During Febru- ary the flowers were visited by five bee species, three Dip- tera, one Hemiptera, and one Lepidoptera in addition to the Honey Bees. Equivalent observations in December re- vealed visits by Honey Bees with one other bee species (Chloralictus — Fig. 8), two Diptera, and one Lepidoptera. Honey Bees were most common in December; other bees (Agapostemon, Chloralictus (2 species), Halictus, and Lasi- oglossum) in February. The reason for the apparent preference for Tribulus nec- tar over pollen during the period from September to De- cember is not clear. Nor is it clear that these observations are the result of a rigid division of labor. There must be some strong force (attractant?) which guides the Honey Bees around the visual guides usually followed in flowers. Since much of the underside of some Honey Bees is dusted with pollen, self- or cross-pollination could easily be accomplished. The plants have not been tested for autog- amy or self-compatibility, but the ample fruits present suggest that Honey Bees or other visitors may successfully complete pollination. Porter (pers. comm.) indicates that T. cistoides is protandrous and thus probably outcrossing. Self-compatibility, however, should not be ruled out until demonstrated. Tribulus cistoides is native to tropical and subtropical Africa (north to Cape Verde on the west and Mozambique on the east). Since it is now widely dispersed throughout the drier tropics, it often becomes a bothersome weed. Apis mellifera, probably native to the Mediterranean area, is also naturalized in the New World. The interactions between Apis and Tribulus in Florida probably did not evolve as the result of foraging activities of the Honey Bee in the Old World since 1) Apis is polytropic by nature, and 2) their distributions do not permit a long history of co- evolution. It must have been only recently that Apis learned to utilize Tribulus as a nectar and pollen source, especially since Apis is capable of rapid learning (Meeuse, 1961; 1972] Tribulus cistoides — Austin 121 Adams, pers. comm., 1971). Regardless of where and when Apis learned, pollen is frequently transported from one Tribulus flower to the other by the visits of Apis. An important factor in the reproduction of the plants is the approach used by the Apis individuals. Those bees ap- proaching from the back of the flowers merely “steal” nec- tar without transferring pollen, while those workers enter- ing the flowers from the front effect pollination. The majority of species other than Apis visit the flowers of Tribulus in the “normal” front approach. Deviations from this are the one Lepidoptera (Melachroia) which took nectar from the back of the flower, and occasional individ- uals of the Dance Fly (Diptera: Empididae) species. Most of the Dance Flies utilized the flowers in the normally ex- pected front approach. The flies are too small, however, to accomplish much pollination. The largest visitor observed on the flowers was a Diptera, probably a Syrphid fly, but no collection was made. Sev- eral flowers were visited by the fly before attempts at photography frightened it away. Another fly, a Tachinid Fly, has been seen occasionally throughout the period of observation. This species is commonly prey to yellow preda- tory spiders in the flowers. Hymenoptera other than Apis have been seen on the flowers. One, a small Chloralictus bee, crawled under the stamens and stigma (Fig. 8). Although the individual in the photograph was not captured, others with the same behavior have been collected. These small insects moved among the stamens as they went from nectary to nectary. Due to the visits among the stamen their bodies often be- came heavily dusted with pollen (Fig. 8). In spite of this none of them has been seen in contact with the stigma; pollination by these small bees must be rare. An Ash-Gray Leaf Bug (Hemiptera: Piesmatidae) exhibited much the same behavior as the small Chloralictus bee. Four bee species other than Apis are probably effective in the pollination of Tribulus. Agapostemon, Chloralictus nymphaerum, Halictus, and Lasioglossum all collect pollen 122 Rhodora [Vol. 74 and nectar in the “normal” manner, and the undersides of their bodies are heavily dusted with pollen. A fifth species was observed on the plants, but not captured. This large wasp-like Hymenoptera visited the plants near Boynton Beach and gathered only nectar on the single visit observed. PLANT COLLECTIONS: FLORIDA: PALM BEACH COUNTY: hammock end north of public beach in Boynton Beach, Austin 4895 (FAU); campus of Florida Atlantic University, Austin 4422 (FAU, MO). INSECT COLLECTIONS: HEMIPTERA: PIESMATIDAE: Austin 4422- 15, 4422-16 (FAU). LEPIDOPTERA: GEOMETRIDAE: Melanchroia cephise (Cramer), Austin 4422-24 (FAU). DIPTERA: EMPIDIDAE: Austin 4422-18, 4422-14 (FAU); TACHINIDAE: Austin 4422-27 (FAU). HYMENOPTERA: APIDAE: Apis mellifera L., Austin 4422-1, 4422-2, 4422-3, 4422-17, 4422-18, 4422-19, 4422-20 (FAU); HALICTIDAE: Agapos- temon splendens Lef.. Austin 4422-21 (FAU) ; Chloralictus nymphaerum Robertson, Austin 4422-5, 4422-23 (FAU); Chloralictus aff. marinis Crawf., Austin 4422-26 (FAU); Halictus sp., Austin 4422-4, 4422-28 (FAU); Lasioglossum sp., Austin 4422-6, 4422-23 (FAU). ACKNOWLEDGEMENTS Thanks are due Ralph M. Adams (Florida Atlantic Uni- versity, Boca Raton, Florida) and Duncan M. Porter (Mis- souri Botanical Garden, St. Louis, Missouri) who read the manuscript and offered suggestions. The Lepidoptera were determined by Thomas E. Pliske (University of Miami, Miami, Florida); the Hymenoptera were determined with a reference collection donated by Calaway H. Dodson (Uni- versity of Miami, Miami, Florida). The assistance given by these men is gratefully acknowledged. The remaining insects were determined with Borror and DeLong (1954) and Borror and White (1970). This study was supported by a grant from the Florida Atlantic University Division of Sponsored Research. LITERATURE CITED Borror, D. J. and D. M. DELoNG. 1964. An Introduction to the Study of Insects, Holt, Rinehart & Winston, New York. and R. E. WHITE. 1970. A Field Guide to the Insects of America North of Mexico. Peterson Field Guide Series, Houghton Mifflin Co., Boston. FAGRI, K. and L. VAN DER PIJL. 1966. The Principles of Pollination Ecology. Pergamon Press, New York. 1972] Tribulus cistoides — Austin 123 GRANT, V. 1950. The Flower Constancy of Bees, Bot. Rev. 16: 379- 398. MEEUSE, B. J. D. 1961. The Story of Pollination. Ronald Press Co., New York. PORTER, D. M. 1971. Notes on the Floral Glands in Tribulus (Zygo- phyllaceae). Ann. Missouri Bot. Gard. (in press). HERBARIUM DEPARTMENT OF BIOLOGICAL SCIENCES FLORIDA ATLANTIC UNIVERSITY BOCA RATON, FLORIDA 33432 REDEFINITION OF CEDRELA OAXACENSIS C. DC. & ROSE C. EARLE SMITH, JR. During the course of preparation of “A Revision of Ced- rela (Meliaceae)” (Fieldiana: Bot. 29: 295-341. 1960) dif- ficulties in defining inadequate collections of Spanish cedar from Mexico and Central America led me to place many species in synonymy. Among these, I recognized a grouping of specimens with (for the genus) exceptionally large cap- sules under the name Cedrela oaxacensis C. DC. & Rose. Subsequently, Dr. Faustino Miranda kindly showed me his well-collected specimens with very good notes which indi- cated that the ecological disparity among the large-capsuled cedrelas was, indeed, solidly based on the confusion of at least two species. A large-fruited species in higher eleva- tions from Chiapas to Panama bears several capsules in a hanging inflorescence. Another species, bearing single erect capsules, grows in less well watered habitats primarily in the semi-deciduous Pacific slope forests from Mexico southward. In Flora of Panama, VI. Family 92, Meliaceae (Ann. Mo. Bot. Gard. 52: 60 seq. 1965), I partially untangled this skein of my own making by recognizing C. tonduzii C. DC. as the upland, moist-forest species with large cap- sules in hanging inflorescences. I included in this C. sal- vadorensis Standl. and I have no evidence at the present time which leads me to change this opinion. At the time that I visited the type locality of C. oaxacensis (Monte Alban, Oaxaca) in 1957 in search of material identi- fiable with Pringle’s collection 4802, the arborescent veg- etation was only beginning to recover from a severe clearing which had accompanied archaeological work at this impor- tant pre-Conquest site. Unable to find Cedrela in the area to corroborate the original collection, I mistakenly concluded that the capsules accompanying the Pringle collection were anomalous. I have since made three collections of Cedrela oaxacensis in the Oaxaca Valley (one at Monte Alban in 124 1972] Cedrela oaxacensis 125 sprout clumps from original stumps on the site) which prove that the foliage and capsules of Pringle 4802 are indeed related. I am now redefining C. oaxacensis to exclude all of the large-capsuled collections which I had previously placed there. Cedrela oaxacensis C. DC. & Rose. Contrib. U. S. Nat. Herb. 5: 190. 1899. Lectotype collection: C. G. Pringle 4802. Monte Alban, Oaxaca. Aug., 1894. (US, B, F, MO, NY, PH). C. discolor Blake. Proc. Biol. Soc. Wash. 33: 108. 1920. Type collection: Palmer 184. San Ramon, Durango. Tree to 10 m tall. Branchlets thick, lenticels small, scattered. Leaves with, usually, 7 pairs of leaflets to 19 cm long, 7 cm wide, ovate, base rounded to obtuse to abruptly trun- cate, rarely slightly acute, apex shortly acuminate, acute, blade markedly pubescent beneath; rachis and petiolules frequently pubescent, petiolules 4 mm long or less. In- florescence moderately open, to ca. 25 cm long in flower, to ca. 30 cm long in fruit, shorter than the leaves, puberu- lent; bracts early deciduous; flowers pinkish, calyx to 2.5 mm deep, 5-lobed, lobes to 1.75 mm wide at base, puberu- lent, petals 5.0 to 6.0 mm long, lanate without, puberulent within, pinkish, but fading toward the thinner margins; filaments fleshy, broad at point of attachment and tapering gradually upward to the anther, connective broad, apiculum short, obtuse; anthers ca. 1.0 mm long; ovary ovoid, gla- brous, style to 2.0 mm long, thick, tapering very gradually to the thick, capitate stigma. Fruit 3.5 to 4.0 em long, valves thin, less than 1.0 mm thick, outer surface lenticellate, outer layer peeling away from the inner as the capsule opens; central column with wings extending only to the margin of a distinct rostrum 6.0 to 8.0 mm in diameter, the scars from the seed attachments 5.0 to 8.0 mm long imme- diately beneath the rostrum between the wings. Trees of hillslopes in the Oaxaca Valley, north and west as far as Durango and probably to the southeast along the Pacific slopes. 126 Rhodora [Vol. 74 Isotype collections of Pringle 4802 and Palmer 184 were recently reexamined in the U. S. National Herbarium, con- firming their conspecificity. Unfortunately, other material was not then available and I was unable to place other col- lections previously cited as C. oaxacensis. However, I have made the following collections in Oaxaca: Rd. to Guelatao ca. 5 km above intersection with Pan American Highway. Stream margin. Smith & Schoenwetter 4346. Aug. 6, 1966.; Monte Alban, on side of mound. Smith & Kitchen 4823. July 2, 1968.; Barranca above molino. San Gabriel Etla. Smith & S. Kitchen 4825. July 4, 1968. I leave unresolved the disposition of C. saxatilis Rose and C. poblensis Miranda. I have already indicated my previous placement of C. salvadorensis Standl. with C. tonduzii. I believe that Miranda felt that C. poblensis is a synonym of C. salvadorensis (annotations on specimens in U. S. Na- tional Herbarium). Neither the specimens nor the localities clearly indicate the proper place of this species. Collections made on the Pacific slope are more apt to be the short tree with the single capsules. However, with increase in eleva- tion, many areas support humid forests in which the larger, multi-capsuled species might be found. Additional field col- lecting with careful documentation is needed to delimit the ranges of the large-capsuled species. I was unable to see a specimen of Pringle 11806 (type collection of C. saxatilis) and I make no disposition, although I suspect that it belongs with C. oaxacensis. I thank the staff of the U. S. National Herbarium for their generous cooperation and my colleague, Joab Thomas, for his patience and helpful comments. PROFESSOR OF ANTHROPOLOGY AND BIOLOGY UNIVERSITY OF ALABAMA TUSCALOOSA, ALABAMA 35486 AN ABNORMAL SPECIMEN OF LAMINARIA DIGITATA (L.) LAMOUROUX ARTHUR C. MATHIESON, EMERY F. SWAN AND RICHARD A. FRALICK! The present report describes the occurrence and mor- phology of an abnormal specimen of Laminaria digitata (L.) Lamouroux. The plant was found (by Emery Swan) de- tached in a deep tide pool near mean low water at Odiornes Point, Rye, New Hampshire, U.S.A. (70°42’42” W longi- tude, 43°03’33” N latitude) on May 16, 1971. Figures 1 and 2 illustrate the general morphology of the plant; a single stipe and two attached blades are evident. The overall length of the plant was 208 cm. The maximum dimensions of the fronds were 120X 98 and 76 X 74 cm respectively, exclusive of the stipe. The larger blade was darker in color, healthier in appearance, and it seemed to be more actively growing, as evidenced by its segmentation (Fig. 2), than the smaller frond. Laminaria digitata is a common plant in the vicinity of Odiornes Point. It grows abundantly on solid rocks from 0 to 14 m below mean low water, with maximum biomass occurring from -2 to 10 m (Mathieson, Hehre and Rey- nolds, in press). It is speculated that the plant broke off its holdfast and then regenerated a second frond. Recent studies by Markham (1968) demonstrated exten- sive regenerative capacities in Laminaria sinclairii (Har- vey) Farlow, Anderson e£ Eaton. Detached portions of its haptera, as short as 2.5 mm in length, were capable of de- veloping into entire plants. Laminaria digitata also exhibits extensive regenerative capacities (unpublished data, A. Mathieson), for new blades can be produced from intact stipes if they are injured or destroyed. Similar regenera- tive capacities are probably evident in healthy, pelagic specimens. Figure 3 shows a conspicuous swelling of the 'Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution Number 127 128 Rhodora [Vol. 74 t $ RINI LLUTETELECLEETERLETULELTELTLELTIEE] 1972] Laminaria — Mathieson, Swan and Fralick 129 stipe in the abnormal specimen; it probably represents the site of regeneration of the second frond. The swelling is several cm below the transition zone —the area of “ac- tive” cell division. Thus, regeneration occurred in an area presumably restricted to elongation and differentiation of cells. The older appearance of the smaller frond suggests that it is the original one. However, in order to preserve the intact specimen a critical examination (section) of the stipe was not made. Burrows (1956, 1958) reported extensive populations of loose-lying algae (particularly Chorda filum, Desmarestia aculeata, Laminaria saccharina and Saccorhiza polyschides) from the subtidal zones at Port Erin Bay, Isle of Man — particularly after periods of calm weather and high insola- tion. Most of the plants were initially attached. Extensive SCUBA observations near Odiornes Point have never shown quantities of loose-lying L. digitata or any other algae — perhaps because of the exposed nature of the site. The accidental deposition of the specimen in a deep tide pool may have provided a relatively stable habitat and allowed regeneration to occur. According to Burrows (1958) mem- bers of the loose-lying populations may serve as a means of restocking habitats devoid of reproductive plants. No re- productive sori were found on the specimen of L. digitata. The vegetative stae of the plant may have been due to the time of its collection, for L. digitata exhibits limited re- production during the spring in New England (Mathieson, Hehre and Reynolds, in press). It should also be noted that several pelagic algae (e.g. Sargassum, Focus and Gracil- aria) are reported (Fritsch, 1959) to have limited repro- ductive capacities. Figure 1. Intact specimen of Laminaria digitata, with a single stipe and two attached blades. Figure 2. Close up of stipe and fronds, showing darker and healthier appearance of larger (left) frond. Active segmentation of the larger frond is also visible. Figure 3. Conspicuous swelling of the stipe, that may represent the site of regeneration of the second frond. 130 Rhodora [Vol. 74 According to van Overbeek (1940 a, b) auxins may be found in Macrocystis pyrifera and other known brown algae. If this were true for L. digitata, then the loss of polarity, due to a pelagic state, may explain the regenera- tion of a second frond rather than a holdfast. Extensive regeneration and the loss of polarity must also occur in other unattached algae that produce spherical morphologies, such as Cladophora holsatica Kiitzing described by Smith (1950) and the “beach form” of Ascophyllum nodosum (L.) Le Jolis ecad Mackaii (Turner) Cotton reported by Gibb (1957). In conclusion the occurrence of an abnormal specimen of L. digitata poses a variety of questions concerning the growth and ecology of the species. Additional studies should be carried out to determine the growth, reproduction and ecological significance of pelagic specimens of L. digitata. LITERATURE CITED Burrows, E. M. 1956. A preliminary survey of the sublittoral algae of Port Erin Bay, Isle of Man. Phycol. Bull. 4: 14-15. , 1958. Sublittoral algae populattions in Port Erin Bay, Isle of Man. J. Mar. Biol. Assoc. U.K. 37: 687-703. FRITSCH, F. E. 1959. The structure and reproduction of the algae. Vol. II. Cambridge University Press, 939 pp. GIBB, D. C. 1957. The free-living forms of Ascophyllum nodosum (L.) Le Jol. J. Ecol. 45: 49-83. MARKHAM, J. W. 1968. Studies on the haptera of Laminaria sin- clairii (Harvey) Farlow, Anderson et Eaton. Syesis 1: 125-131. MATHIESON, A. C., HEHRE, E. J. and ReyNoups, N. B. Investigations of New England marine algae I: A floristic and descriptive ecological study of the marine algae at Jaffrey Point, New Hampshire, Nova Hedwigia (in press). OVERBEEK VAN, F. 1940a. Auxin in marine algae. Plant Physiol. 15: 291-299. SMITH, G. M. 1950. The fresh-water algae of the United States, McGraw-Hill Book Co., N. Y., 719 pp. JACKSON ESTUARINE LABORATORY OF THE UNIVERSITY OF NEW HAMPSHIRE DURHAM, N.H., U. S. A. 03824 RANGE EXTENSIONS OF VASCULAR AQUATIC PLANTS IN NEW ENGLAND C. BARRE HELLQUIST During 1970 and 1971 extensive collecting of vascular aquatic plants was conducted in New England. New records for some states as well as interesting extensions of range were noted. In New England the aquatic flora has been neglected during the past twenty years, so these ranges may have expanded or may have been overlooked. Such botanists as M. L. Fernald and A. S. Pease did exten- sive collecting in the time span of 1920 to 1955. Since then no single person has devoted much time to the aquatics in New England except William Countryman who has done extensive collecting in Vermont. The plants collected and observed are discussed below. Representative specimens have been deposited in the author’s personal herbarium as well as those of Boston State College, the University of New Hampshire and the New England Botanical Club. Potamogeton filiformis Pers. var. borealis (Raf.) St. John Both Potamogeton filiformis and its variety borealis are found in New England only in the extreme north. A plant of calcareous waters, (St. John, 1916), this plant has been collected from several areas in Aroostook County, Maine, and in northern Vermont (Seymour, 1969). A specimen in the University of New Hampshire herbarium (S B. Krochmal, 142a) from Piermont, New Hampshire, Grafton County was examined by this author and keyed out as P. filiformis Pers. var. Macounii Morong, which has never been reported from New England. This identification is questionable and more specimens of this plant would be desirable for absolute identification. In October 1970, several specimens of the variety borealis were collected for the first time in New Hampshire in Lombard Pond, Cole- brook, Cóos County (Hellquist 93 and 94). This pond has 191 132 Rhodora [Vol. 74 the highest alkalinity of any in the state with a Methyl Orange ppm. reading of 90 (Newell, 1960). Of all the ponds in New Hampshire this is one of the few where this Potamogeton would find the requirements of an alkaline body of water in the northern portion of the state. Lime Pond in Colebrook and Ladd Pond in West Stewartstown might also harbor this species. Potamogeton crispus L. This has increased its range extensively in the past fifty years. Records in the Gray Herbarium and the New Eng- land Botanical Club Herbarium include very few reports of the species. Seymour (1969) noted this plant from Middlesex County in Massachusetts, Chittenden County in Vermont, Providence County in Rhode Island and a number of areas in Connecticut. Collections of this plant were made from a number of ponds and lakes in Massachusetts and a couple from northwestern Connecticut. Ogden (1943) noted it from Spy Pond in Arlington, Massachusetts, Fresh Pond in Cambridge, Massachusetts and the Sudbury River in Concord, Massachusetts. All of these are in Middlesex County. In Connecticut, the Housatonic River in Litchfield County was the only area noted. From the recent collections by the present author it is evident that Potamogeton crispus L. has increased its range extensively, especially in the hard water regions of Berkshire County, Massachusetts. This plant was ex- tremely common in the Stockbridge Bowl, Stockbridge, Massachusetts where it was the most abundant pondweed and along with Myriophyllum spicatum L. var. exalbescens (Fernald) Jepson made up about 80% of the vascular vege- tation. This plant was also extremely common at Lake Buel in Monterey and New Marlboro, Massachusetts, Mill Pond, South Egremont, Massachusetts and the Mill Pond in Shef- field, Massachusetts. A new record in Middlesex County for this species is Fisk Pond, Framingham, Massachusetts. From Connecticut it was found in Mudge Pond and Indian Lake, Sharon, Litchfield County. In all the above mentioned 96 1972] Aquatic Plants — Hellquist 133 areas except Stockbridge Bowl it was not the predominant species but was nevertheless common. This species has been known to be more tolerant of polluted waters (Fassett, 1966). After observing the water where collections were made this statement would seem to hold true. The ponds from western Massachusetts and Connecticut were in alka- line regions where this plant does well (Moyle, 1945). This plant should be looked for in almost all hard water regions of New England especially where pollution may occur. Potamogeton strictifolius Ar. Benn. Var. rutiloides Fern. This is one of the narrow-leaved pondweeds of the section Pusilli and is characteristic of basic or alkaline waters of the northern United States and Canada (Fernald, 1932). Fernald at that time noted it from Lake Champlain, Ver- mont. Seymour (1963) recorded the variety rutiloides from three other areas of western Vermont not far from Lake Champlain as well as from Berkshire County, Massa- chusetts. A new eastern record for Potamogeton strictifolius Ar. Benn. Var. rutiloides Fern. in New England is the Mill Pond in Windsor, Vermont, Windsor County. It was abundant in some of the shallow areas on the eastern side of the pond. This plant, upon first observation, was mis- taken by the writer for Potamogeton Friesii Rupr. with which it may be confused easily. Potamogeton strictifolius and its variety generally have three veins, but this speci- men had three to five. Fernald (1932) noted that P. stricti- folius may rarely have five veins. The Mill Pond has the typical vegetation of the more alkaline lakes including Najas flexilis (Willd.) Rostk. & Schmidt., Vallisneria americana Michx. and Potamogeton pectinatus L. (Moyle, 1945). Potamogeton longiligulatus Fern. One of the rarer pondweeds in the United States as well as New England is P. longiligulatus. Only two specimens have been reported and both were from northwestern Con- necticut. Fernald (1932) noted it from Twin Lakes, Salis- 134 Rhodora [Vol. 74 bury, Connecticut, where it was collected by C. H. Bissell in 1906. A second report was from Indian Lake in Sharon, Connecticut, (Seymour, 1969). This specimen was de- posited in the New England Botanical Club Herbarium but after careful study by the writer it was found to be a very narrow-leaved form of Potamogeton zosteriformis Fernald. The second actual record for this specimen comes from Evarts’ Pond (Lake Runnymede) in Windsor, Vermont, Windsor County. Evarts’ Pond is a privately owned pond which is of interest in that it was one of the ponds where Potamogeton Hillii Morong. had been reported in 1875 (Fernald, 1932). More recent collections of P. Hillii from Evarts’ Pond in the late thirties are in the New England Botanical Club Herbarium. In an attempt to find Pota- mogeton Hillii, permission was received from the owners to collect in the pond. A narrow-leaved pondweed believed to be Potamogeton Hillii was collected by the author at the northern portion of the northeast bay and on the southwestern end of the pond. In both areas it was found in water approximately three feet deep. Upon examining these collections care- fully, it was determined that the two rare Potamogetons, Hillii and longiligulatus were both found. Fernald (1932) commented on much confusion in identifying these two species. Both have sharply acute, cuspidate leaves. Pota- mogeton Hillii has three veins while P. longiligulatus has five to nine. Both specimens were in flower at this time so they had the same initial appearance. In talking with Mr. Evarts, one of the owners, it was learned that the state of the pond had deteriorated over the past two summers. The water previously had been very clear, but now becomes cloudy as the water warms up in the summer. He stated that corn has been planted along the north side of the pond, and is fertilized extensively with chemicals. This most likely has led to the turbid water with its poor light penetration so it was rocommended that the corn crop be discontinued in 1972. Hopefully this would clear up the problem. Since this pond has no inlets, but 1972] Aquatic Plants — Hellquist 135 is spring-fed, it shouldn’t take long for the water to become clear again. It would be a shame for the ecology of this pond to be so disturbed that these two valuable pondweeds would be destroyed. Nymphaea tuberosa Paine was also discovered here and will be discussed later in this paper. Potamogeton gemmiparus Robbins. This uncommon narrow-leaved pondweed has been re- ported from only four New England states: Maine, Massa- chusetts, Connecticut and Rhode Island (Fernald, 1932). Since it is from eastern New England it is found in more acid waters. This plant had never been reported in New Hampshire, where it should occur. Seymour (1969) records it as near as Fryeburg, Maine, where it is found in the Saco River. Two stations of this plant were found by the author in Carroll County, New Hampshire, both in the Saco River drainage. One station where it was somewhat rare was the southern end of Conway Lake in Eaton, New Hamp- shire. Here it was found with other acid water plants such as Potamogeton Berchtoldii Fieber, P. Berchtoldii Fieber, var. tenuissimus (Mert. & Koch) Fern., P. epihydrus Raf. var. ramosus (Peck) House, P. gramineus L. var. grami- neus and P. capillaceus. Bearcamp Pond in Sandwich, New Hampshire was the other site of collection. Here the speci- men was collected at the inlet of the Bearcamp River. Again, it was not plentiful. Potamogeton gemmiparus was growing along with P. natans L., P. gramineus L. var. myriophyllus Robbins. P. gramineus L. var. gramineus, P. confervoides, Reichenb., P. capillaceus Poir. and P. Spirillus Tuckerm. Egeria densa Planchon Egeria has been reported from three different towns in New England — two in Massachusetts and one in Vermont (Seymour, 1969). The reports from Massachusetts were from Abington in Plymouth County and Quincy in Norfolk County. In Vermont it was collected from Townshend, Windham County. It was observed by the writer to be very 136 Rhodora [Vol. 74 common in Hemenway Pond, Milton, Massachusetts, Nor- folk County, near the Boston city limits where it was growing with Elodea Nuttallii (Planch.) St. John, Pota- mogeton epihydrus Raf. var. ramosus (Peck) House, Myriophyllum humile (raf.) Morong and Utricularia geminiscapa Benj. This plant was introduced and natu- ralized from South America and is one to be looked for around cities because it is being sold in pet shops. Wolffia columbiana Karst. This plant is continuing to spread its range to the north in New Hampshire. Previously reported from Hillsborough County, (Countryman, 1968) and Rockingham County (Colt, et. al., 1971) it was discovered by the author in Durham, New Hampshire, Strafford County. In Durham, it was reported from two new locations — the Mill Pond and Beard’s Creek Pond. This plant was not observed in either place in 1970, but was very abundant at Beard’s Creek Pond and common at the Mill Pond in 1971. Wolfia punctata Griseb., which usually occurs with the former, was not found but should be looked for in the future. In talking with Dr. Albion Hodgdon at the University of New Hampshire in Durham, it was agreed that this plant probably is rapidly extending its range. One explana- tion for the rapid spread may be the passage of ducks in the region moving between ponds. Wolffia would stick to their bodies and be carried around easily. Also anyone transferring a boat from ponds could easily introduce the plants into new bodies of water. Nymphaea tuberosa Paine Seymour (1969) recorded this plant from six stations in western Vermont, Winchester, Massachusetts, and three townships in Connecticut. This species has been found by the writer to be quite abundant. In Vermont, a new eastern record for the state is herein reported from Evarts’ Pond, Windsor County. Here it was the only member of the Nymphaeaceae and was found extensively around the pond. The first two records for this plant in New Hampshire are 1972] Aquatic Plants — Hellquist 137 Lake Kanasatka, Moultonboro, Carroll County and Beard’s Creek Pond, Durham, Strafford County. This pond lily was found at the southern end of Lake Kanasatka near the gift shop “The Horse of Another Color" along Route 25. It appears that this plant was introduced here some time ago, due to its localized range at the southern end of the lake. It was found with other members of the same family in- cluding Nymphaea odorata Ait., Nuphar variegatum Engelm., and Brasenia Schreberi Gmel. This is a lake with a higher alkalinity than most other lakes in Carroll County as shown by the presence of Potamogeton zosteri- formis Fernald and P. praelongus Wulfen. At Beard’s Creek in Durham Nymphaea tuberosa Paine was found near Route 4 where it has spread throughout the southern end of the pond. The water is probably in much the same condition as Taylor River in Hampton, New Hampshire (Colt, et al., 1971) since this also is a tidal estuary which has been blocked off by a dam. Much salt in the bottom might tend to raise the alkalinity of the water. Other plants of interest growing along with N. tuberosa here are Myrio- phyllum verticillatum L. var. pectinatum Wallr. and Elodea Nuttallii (Planch.). In Massachusetts this plant was discovered by the writer in three locations, all within Middlesex County. These were the Sudbury and Concord Rivers in Concord, Massachusetts and a small pond by Upper Mystic Lake in Winchester, Massachusetts which is near the previously reported site. Mr. Richard Eaton of Lincoln, Massachusetts told the author in 1971 that Nymphaea odorata used to be very com- mon in these rivers and was almost completely destroyed due to the pollution. Nymphaea tuberosa is found in the highly productive lakes of the midwest and seems to be more tolerant of polluted waters. In the Sudbury River Nymphaea odorata was found in limited numbers while N. tuberosa was more common. The area observed in the Concord River had Nymphaea tuberosa as the only water lily and it was found at only one location. The location at Winchester, Massachusetts was a small 138 Rhodora [Vol. 74 pond at the recreation area on the east side of Upper Mystic Lake. It appears that this lily was planted here as it does not extend into Upper Mystic Lake which connects with this pond. This plant may be more common than previously ob- served. Many of the specimens reported as Nymphaea odorata Ait. var. gigantea Tricker may actually be N. tu- berosa. The leaves of N. tuberosa are generally larger and green underneath, or with only a slight purplish tinge. Probably the best identifying characteristic of this plant is the large green petiole with brown stripes as opposed to the purple petiole of Nymphaea odorata. Fernald (1950) noted that the flowers are odorless or barely fragrant which may be misleading. The flowers have a noticeable odor, but when compared with N. odorata they are less fragrant. Another interesting feature found on a number of speci- mens was the presence of pubescence on the petiole near the blade. In some this was so marked that the striping in the petiole was obscured. Other characteristics which aid in identification are the more rounded petals and the ten- dency of the leaves to become elevated above the water by as much as one foot. Nymphaea alba L. forma rosea Hartm. A European species, this pond lily has not been reported previously from New England. It was found by the writer to be relatively abundant in Winkley Pond, Barrington, New Hampshire, Strafford County. It occurs around the perimeter of the Pond where it grows with Nymphaea odorata and a possible hybrid between N. odorata and N. alba f. rosea which has the pointed more numerous petals and is a light pink. Winkley Pond is an acid body of water in a semi-bog condition with many ericaceous plants. Of particular interest as an associated plant is Potamogeton confervoides Reichenb. which is extremely common there. Nymphaea alba f. rosea is characterized by 12-24 rounded deep pink petals (Fernald, 1950). The leaves are similar to N. odorata with deep purple undersides. The following 1972] Aquatic Plants — Hellquist 139 observations were made on N. alba f. rosea: many of the petioles were pubescent as seen in N. tuberosa and the leaves were elevated above the surface of the water along the edge of the pond. Because of the attractive blossom, this water lily may have been planted more extensively than observed. Many specimens labeled Nymphaea odorata Ait. forma rubra Guillon may actually be N. alba f. rosea. Myriophyllum alterniflorum D. C. Except for the present report M. alterniflorum has been found in New England only from numerous areas in Vermont and Maine and from Cóos County in northern New Hampshire (Seymour, 1969). Last year the first record of the species south of Cóos County in New Hamp- shire was noted in Upper Danforth Pond, town of Freedom, Carroll County, (Hellquist, 1971). Two other locations for this plant have also been found in Carroll County by the writer. One is from Cooks Pond in Madison, New Hamp- shire where it was scarce. Isolated plants were found in this acid-water pond which also included Potamogeton confervoides Reichenb. (Hodgdon, et. al, 1946). Lake Kanasatka, Moultonboro, New Hampshire was the other station. The Myriophyllum was found at the south end of the lake near the previously reported Nymphaea tuberosa Paine. As more extensive studies are carried out in central New Hampshire, this plant should prove to be more abundant. Myriophyllum heterophyllum Michx. This plant has been quite uncommon in New Engiand until recently if the few herbarium specimens are a fair indication. It had been recorded from Bridgeport, Con- necticut, Uxbridge and Sutton, Massachusetts (Seymour, 1969). A specimen from Sandy Pond, Ayer, Massachu- setts, collected by Edward Richardson in 1940 is on deposit in the University of Massachusetts herbarium. In southern Worcester and Middlesex Counties, Massa- chusetts, the writer found M. heterophyllum to be extremely common in a number of ponds and lakes, where: it forms 140 Rhodora [Vol. 74 large beds around the shore. It has been found in White- hall Reservoir and North Pond in Hopkinton, Middlesex County; Pratt Pond, Upton; Silver Lake, Grafton; and Hopedale Pond, Hopedale, Worcester County. Houghton’s Pond (Hoosiwhisick Pond), Canton, Norfolk County, Mill Pond, Wareham, and Agawam River, Wareham, Plymouth County are other areas where this plant was found to be abundant. Fassett (1966) noted this plant to range from Virginia to Florida, west to Ontario and Minnesota, so it appears that this specimen is spreading rapidly in southern New England. One cause of this recent and rapid spread may be the fact that the plant is sold extensively in pet shops for aquarium use. It may be discarded into ponds and streams as has Egeria densa Planchon. LITERATURE CITED Corr, L. C., JR., C. BARRE HELLQUIST and W. J. L. ZUBRIN. 1971. An Interesting Association of Rare Aquatic Plants from New Hamp- shire. Rhodora 73: 296-297. CoUNTRYMAN, W. D. 1968. Wolfia in New Hampshire. Rhodora 70: 491. Fassett, NoRMAN C. 1966. A Manual of Aquatic Plants. ( Appen- dix by E. C. Ogden). University of Wisconsin Press. Madison, Wisconsin. 405 p. FERNALD, M. L. 1932. The Linear-leaved North American Species of Potamogeton, section Axillares. Memoirs of the American Academy of Arts and Sciences Vol. 17, Part I. 183 p. 1950. Gray's Manual of Botany, eighth edition. American Book Company, New York. 1632 p. HELLQUIST, C. BARRE, 1971. Vascular Flora of Ossipee Lake, New Hampshire and its Shoreline. Rhodora 73: 249-261. Hopepon, A. R., P. GIGUERE, S. B. KROCKMAL and A. RIEL. 1954. New Potamogeton Records in New Hampshire. Rhodora 54: 237-246. MovLE, Jonn B. 1946. Some Chemical factors Influencing the Dis- tribution of Aquatic Plants in Minnesota. Am. Midl. Nat. 34: 402-420. NEWELL, ARTHUR E. 1960. Biological Survey of the Lakes and Ponds in Coos, Grafton, and Carroll Counties. Survey Report no. 8a.New Hampshire Fish and Game Department. 297 p. OGDEN, E. C. 1943. The Broad Leaved Species of Potamogeton of North America North of Mexico. Rhodora 45: 57-105, 119-163, 171-214. 1972] Triphora trianthophora — Eastman 141 SEYMOUR, FRANK C. 1969. The Flora of New England. The Charles Tuttle Co., Rutland, Vermont. 596 p. St. JoHN, HaROLD. A Revision of the North American Species of Potamogeton of the Section Coleophylli. Rhodora 18: 121-138. BIOLOGY DEPARTMENT BOSTON STATE COLLEGE 625 HUNTINGTON AVENUE BOSTON, MASSACHUSETTS 02115 A SECOND OCCURRENCE FOR TRIPHORA TRIAN- THOPHORA (SW.) RYDB. IN MAINE. Another station of the orchid, Triphora trianthophora, the second to be found in the last few years, was discovered on August 31, 1971 at Evans Notch, Batchelders Grant, Oxford County by C. Paul Wight and me. This location is about three miles in a direct line from the Stow station (Rhodora 71: 509, 1969). This new station consists of many individuals grow- ing on a wooded hillside in depressions which are filled with thickly matted beech leaves. It is difficult to estimate how many of the orchids occur at this station, as they tend to grow singly, spreading out over a large area, unlike the Stow station, where the orchids were growing in large groupings. We were fortunate in finding many of the plants still in flower for the majority had already lost their blos- soms. Other interesting plants were found among the Triphoras, namely, Corallorhiza maculata, Conopholis amer- icana, Epifagus virginiana, and members of the Botrychium group. Specimens have been deposited at the Herbaria of the University of Maine and the New England Botanical Club. LESLEY M. EASTMAN OLD ORCHARD BEACH, MAINE 04064 ILLINOIS FIELD AND HERBARIUM STUDIES ROBERT H. MOHLENBROCK AND DAN K. EVANS Continued field and herbarium studies of Illinois plants have resulted in the discovery of several unreported or pre- viously undocumented taxa for the state. These new addi- tions to the flora are reported in this paper, in addition to locality records for other rare taxa in southern Illinois. All specimens cited are in the herbarium of Southern Ili- nois University (SIU), unless otherwise indicated. TAXA NEW TO ILLINOIS Cyperus iria L., an introduced southern sedge, was known to range in North America from Virginia east to southeast- ern Missouri, southwest to Texas, and east to Florida. Now Illinois may be added to the range. In Illinois, this species may be found sparingly in the wet meadows at Horseshoe Lake, Alexander County. Of the twenty-one taxa of Cyperus found in Illinois, this species may be most likely confused with those in the fol- lowing key. a. Scales 1.0-1.5 mm long; achenes 0.8-1.0 mm long. b. b. Achenes white; spikelets about 1 mm broad; scales closely arranged ................ C. erythrorhizos b. Achenes brown or black; spikelets about 1.5 mm broad; scales somewhat remote from each other . a C. iria a. Scales 1.5-4.5 mm long; achenes 1.0-2.8 mm long. c. c. Scales remote, the tip of one just reaching the base of the one above; base of the achene enclosed in a corky sheath ............ sees C. engelmannit c. Scales approximate and overlapping ; no corky sheath present at base of achene. d. d. Some or all of the mature spikelets reflexed; spikelets subterete .......... C. lancastriensis d. None of the spikelets (except sometimes the low- est pair) reflexed; spikelets flattened. e. 142 1972] Illinois Studies —- Mohlenbrock & Evans 143 e. Rhizomes scaly and usually ending in a tuber; scaletips slightly spreading .... C. esculentus e. Rhizomes absent or merely hard and corm- like; scaletips appressed. f. f. Plants: annual without rhizomes; scales ferrugineous or golden-brown, 1.7-3.0 mm long; achenes obovoid-oblongoid . ...... M ar ee: C. ferruginescens f. Plants perennial with hard, corm-like bases; scales stramineous, 3.5-5.0 mm long; achenes linear ..... .. C. strigosus COLLECTION DATA: Alexander County: Horseshoe Lake, wet meadows, July 30, 1968, E. Estes s.n. Ulmus procera Salisb. Several specimens of the English elm oecur in the vicinity of an abandoned farm along the “Giant City blacktop” about 1 1/4 miles north of Giant City State Park. The colony is apparently spreading by suckers. This elm is distinguished by its scabrous upper leaf surface and the presence of corky wings on the branches. COLLEC- TION DATA: Jackson County: along blacktop road between Carbondale and Giant City State Park, June 26, 1970, R. H. Mohlenbrock 19256. Amaranthus caudatus L. This often cultivated Tassel Flower was collected from a spontaneous colony in waste ground. Although it has undoubtedly escaped elsewhere in Illinois, this is the first report of such an occurrence. COL- LECTION DATA: Jackson County: waste ground, Oakland Street, Carbondale, July 29, 1970, D. Evans 1112. Ranunculus repens L. var. pleniflorus Fern. The garden form of the creeping buttercup, with most of its stamens transformed into petals, was collected in a grassy area at the edge of the campus of Southern Illinois University, Carbondale. COLLECTION DATA: Jackson County: Carbon- dale, August 14, 1967, R. H. Hohlenbrock s.n. Nigella damascena L. The Love-in-a-mist, a member of the Ranunculaceae, is occasionally grown as a flower garden ornamental and is rarely escaped from cultivation. How- ever, a few specimens were observed at the edge of an 144 Rhodora [Vol. 74 abandoned lot at the rear of 411 South 20 Street, Murphys- boro, during the summer of 1968. The deeply divided in- volucre which subtends the blue flowers accounts for the common name. COLLECTION DATA: Jackson County: lot, rear of 411 S. 20 Street, Murphysboro, July 27, 1968, R. H. Mohlenbrock 16121. Rorippa islandica Oeder var. islandica. This variety of Yellow Cress, new to the Illinois flora, was collected from mud and sand flats along the Mississippi River. In that habitat it occurs with the more abundant variety fernald- iana and a similar second species, R. sessiliflora. Although normally found in the northeastern United States, Quebec, and Greenland, Steyermark (1960) reports a station in Harrison County, Missouri. COLLECTION DATA: Jackson County: Mississippi River mud and sand flats, Grand Tower, November 1, 1969, D. Evans 656. Rubus procerus P. J. Muell., a native of Europe, is re- ported by Fernald (1950) as naturalized in the United States from Delaware to Virginia. Steyermark (19623) re- ports a Palmer collection from Newton County, Missouri. The authors collected this Illinois record from a roadside habitat in Randolph County, where it grows in abundance. The large drupelets are most edible. Among the many tax- onomically difficult species of Rubus one encounters in Illi- nois, only three have leaflets with the underside whitened. These may be identified in the field by the following key: a. Stems glaucous . .. ............... R. occidentalis a. Stems not glaucous. b. b. Mature fruit red; prickles small, bristle-like or ab- sent; leaves ovate-lanceolate ... ................ NENNEN R. idaeus var. strigosus b. Mature fruit black; prickles abundant, flat with a broad base; leaves broadly ovate ...... R. procerus COLLECTION DATA: Randolph County: roadside, 10 miles south of Chester, Illinois, July 21, 1970, R. Mohlenbrock & D. Evans 1113. Croton texensis (Klotzsch) Muell. Jones and Fuller (1955) report that although Ferguson (1901) cites a Hall 1972] Illinois Studies — Mohlenbrock & Evans 145 collection of this species from Athens, Illinois, it is excluded from the Illinois flora since no specimen could be found to verify this report. However, a specimen of this species, collected by Hall from Athens, is on deposit in the her- barium of the Missouri Botanical Garden. This poisonous species normally ranges from South Dakota, south to Mex- ico, with introductions known from western Missouri, Ala- bama, and New England. COLLECTION DATA: Menard County Athens, Hall 514. Hibiscus syriacus L. The Rose-of-Sharon is a small tree often planted, particularly in the past, as an ornamental. Occasionally specimens may be found around abandoned homesteads, although they generally do not appear to be spreading. A small grouping of specimens occurs along an abandoned road on the Little Grassy camp facilities of Southern Illinois University. At this site, there is indication that the plants have multiplied since their original planting some forty years ago. COLLECTION DATA: Jackson County: along dirt road, Little Grassy camp, Southern Illinois Uni- versity, August 14, 1969, R. H. Mohlenbrock s.n. Ascyrum hypericoides L. var. hypericoides. The range of the typical variety of A. hypericoides generally lies to the south and east of Illinois. On the other hand, var. multi- caule (Michx.) Fern., is a relatively common taxa in dry woods, on slopes, and along ridges across the southern tip of Illinois. Variety hypericoides has never been reported from Illinois although there is a specimen of it, housed in the herbarium of the Missouri Botanical Garden, annotated by Dr. Preston Adams, the most recent monographer of the group. It is interesting to note, however, that Adams fails to attribute this variety to Illinois in his monograph (1957). Variety hypericoides differs from var. multicaule by its more ascending habit and by its narrower leaves which attain widths of only 4(-5) mm. COLLECTION DATA: Hancock County: Augusta, July, 1842, S. B. Mead s.n. Hypericum punctatum Lam. var. pseudomaculatum (Bush) Fern. When a typical specimen of the Large Spotted St. Johns-wort is encountered, the first reaction is to con- 146 Rhodora [Vol. 74 sider it a distinct species since the large flowers are so strikingly different from the similar but smaller-flowered var. punctatum. However, no other characters seem fo ex- ist which can reliably distinguish var. pseudomaculatum from var. punctatum. A frequently used character to sep- arate these two entities is the uppermost leaves obtuse in var. punctatum and acute in var. pseudomaculatum. Al- though all specimens which we have observed of var. pseud- omaculatum do have acute upper leaves, a few specimens of var. punctatum also have acute upper leaves. Jones (1963) and Jones, et al. (1955) attribute Hypericum pseud- omaculatum to Illinois, but do not indicate localities. Since we have never before observed a specimen of this plant from Illinois, we are documenting its occurrence in Illinois by the following collection. COLLECTION DATA: Jackson County: dry woods, July 26, 1964, J. Ozment & DeFilipps 1726. Hypericum densiflorum Pursh. This shrubby St. Johns- wort previously was known from southern Missouri, but the collection cited below is apparently the first from Illi- nois. This species closely resembles H. lobocarpum from which it differs only by the presence of three styles and a 3-celled capsule. Since these seem to be fundamental dif- ferences to us, we are treating the two as separate species. Other botanists prefer to treat H. lobocarpum as a variety of H. densiflorum. Although Fernald (1950) indicates that the tip of the leaf may be used to separate H. densiflorum from H. lobocarpum, we do not find this character to be reliable. The other shrubby St. Johns-wort in Illinois with three styles is H. spathulatum, a species usually larger in all respects. COLLECTION DATA: Alexander County: swampy woods near Miller City, August 28, 1964, J. Ozment & R. DeFilipps 2712. Hypericum sphaerocarpum Michx. var. turgidum (Small) Svenson. This southern variant of H. sphaerocarpum is recognized by its narrow, revolute leaves which have no apparent lateral nerves. In some cases, it looks more like H. denticulatum Walt. or H. dolabriforme Vent. than H. 1972] Illinois Studies — Mohlenbrock & Evans 147 sphaerocarpum var. sphaerocarpum. It is distinguished from H. denticulatum by its united styles and from H. dolabriforme (which does not occur in Illinois) by its shorter sepals and somewhat smaller flowers. COLLECTION DATA: Jackson County: railroad prairie, 6 miles north of Murphysboro, June 13, 1953, R. H. Mohlenbrock s.n.; Ma- coupin County: railroad prairie southwest of Carlinville, August 15, 1968, J. White 335; Monroe County: limestone bluffs, one mile south of Fults, August 7, 1962, J. Ozment s.n. Hypericum sphacrocarpum Michx. var. turgidum (Small) (1945) indicated that the Northern St. Johns-wort was known from Illinois, we are unaware of any Illinois collec- tions of this species prior to the 1964 collection cited here. Since the overall range of H. boreale reaches from New- foundland across to eastern Iowa, its occurrence in north- ern Illinois is not unexpected. 'This species is similar in appearance to the other small-flowered species of Hyperi- cum in Illinois, but differs in the bracteal leaves which are similar to the cauline leaves in shape and size. COLLECTION DATA: Iroquois County: Iroquois County Conservation Area, August 17, 1964, J. Ozment & R. DeFilipps 2298. Aesculus glabra Willd. var. leucodermis Sarg. This variety differs from typical A. glabra in its distinctly whitened lower leaf surfaces and its whitish bark. At its only Illinois location, it grows in a mesophytic woods. COL- LECTION DATA: Jackson County: along Kinkaid Creek, three miles south of Ava, July 14, 1970, R. H. Mohlenbrock s.n. Acer rubrum L. f. tomentosum (Desf.) Dansereau. This form is enigmatic in that the densely tomentose lower leaf surface of this upland taxon is identical with that of the swamp-inhabiting A. rubrum L. var. drummondii (H. & A.) Sarg. Only the fruits can be used reliably to separate these two taxa. Forma tomentosum has samaras up to 2.5 cm long, while var. drummondii has samaras over 3.0 cm long. COLLECTION DATA: Union County: upland woods, Pine Hills, July 3, 1970, R. H. Mohlenbrock s.n. Euonymus fortunei (Turcz. Hand.-Maz. This climbing 148 Rhodora [Vol. 74 evergreen ornamental has become rampant in a low woods in Giant City State Park. COLLECTION DATA: Jackson County: low woods, Giant City State Park, February 6, 1971, R. H. Mohlenbrock 21626. Elaeagnus umbellatus Thunb. is previously unreported from Illinois. Collections from this small tree were made from an island in Lake-of-Egypt, Williamson County. At this site a large number of plants are well established and are spreading spontaneously. No evidence of a homesite exists on the island. Elaeagnus umbellatus differs from E. angustifolia, also a rare adventive in Illinois, by displaying both brown and silver scales on the leaves and twigs, while the latter has only silver scales present. COLLECTION DATA: Williamson County: island in Lake-of-Egypt, August, 1970, J. Swayne s.n. Cynosciadium digitatum DC. represents a genus new to the Illinois flora. Collections of this species were made from the Pin Oak flats in the Greentree Reservoir, Jackson County. This area, long set aside as a waterfowl preserve, has seldom been botanized during the growing season since it is mostly innundated and infested with mosquitos. Col- lections made in 1969 and again in 1970 from widely sep- arated areas indicate this species is well established at this southern Illinois site. Associated species include Glyceria striata, Carex squarrosa, Quercus palustris, and Ptilim- nium costatum. The overall range of this species previously reached from Mississippi and Texas north to Oklahoma, and east to southeastern Missouri. COLLECTION DATA: Jack- son County: Greentree Reservoir, Pin Oak flats, June 6, 1969, R. Anderson & D. Evans 1001. Scutellaria ovata Hill var. rugosa (Wood) Fern. This tax- on is very different in appearance from the other varieties of S. ovata (var. ovata and var. versicolor) in Illinois. Its small, rather sprawling habit is contrasted sharply with the large, erect stature of the other varieties. Perhaps the most striking difference exhibited by var. rugosa is found in the leaves, which are strongly wrinkled and seldom reach a length greater than 4 cm. In general, var. rugosa 1972] Illinois Studies — Mohlenbrock & Evans 149 occupies dry, rocky slopes, whereas var. ovata and var. versicolor occur more frequently in more mesic woodlands. Fernald (1950) indicates that var. rugosa is further dis- tinguished by its smaller corolla (about 1 cm long). Ilinois as well as Missouri material assignable to var. rugosa fre- quently has the corolla up to 1.5 cm long, while some corollas in var. ovata and var. versicolor may be less than 1.5 em long. COLLECTION DATA: Monroe County: south of Fults, Illinois, August 14, 1952, W. Bailey & J. Swayne 2818. Hypochoeris glabra L. This introduced species is not re- corded in Gray’s Manual of Botany, 8th edition (Fernald, 1950). The station closest to Illinois seems to be eastern South Carolina as reported by Radford, et al. (1964). In Illinois, this species was collected from waste ground where it grew with other pioneer-type weeds. The beakless outer achenes of H. glabra distinguish it from H. radicata, also a rare species in Illinois, in which all the nutlets are beaked. COLLECTION DATA: Jackson County: rear of 1401 Tripoli, Carbondale, July, 1969, P. Thomson s.n. ADDITIONAL LOCALITIES FOR RARE SOUTHERN ILLINOIS TAXA Carex hystricina Muhl. This species is relatively common in the northern two-thirds of the state, but is known else- where in Illinois only from Pulaski and now Union County. Union County: LaRue Swamp, June 28, 1969, D. Tindall & S. Wunderle s.n. Lychnis alba Mill. This Union County collection is from the farthest south locality for this species in Illinois. Union County: along Route 51, near Cobden, May 15, 1969, R. H. Mohlenbrock & D. Evans s.n. Ranunculus carolinianus DC. The achenes of this species are larger (over 3.5 mm long) than those of R. septen- trionalis and have a conspicuous broad, high keel near the margin of the achene. Previously this species was known from collections in Champaign and Hancock counties in the central part of the state. The collection reported here is 150 Rhodora [Vol. 74 the first for southern Illinois. Union County: McCann Springs, May, 1968, C. Ott s.n. Lotus corniculatus L. This adventive is becoming increas- ingly more common in Illinois. There are now three south- ern Illinois counties for this species, as well as several northern counties. Johnson County: roadside, near Gore- ville, July, 1969, D. Evans & R. H. Mohlenbrock 1046. Trifolium pratense L. f. leucochraceum Aschers. & Prantl. Only a specimen from McLean County was previously known from Illinois. Randolph County: roadside, north of Prairie de Rocher, July 21, 1970, D. Evans 1108. Hibiscus trionum L. The collection reported here is the farthest south for this species in Illinois. Jackson County: mud and sand flats of the Mississippi River, Grand Tower, October 9, 1968, D. Evans 441. Myriopsyllum exalbescens Fern. This species was pre- viously known only from the northern half of Illinois. Johnson County: Lake-of-Egypt, Beaver Neck, September 6, 1968, J. Swayne s.n. Lysimachia terrestris (L.) BSP. This species was not known from the southern half of the state until it was collected at the edge of a swamp. Union County: LaRue Swamp, June 25, 1968, M. Sadler s.n. Verbena hastata X urticifolia Pepoon. This hybrid is scattered in Illinois, but its only southern Illinois localities previously had been from Hardin and Wabash Counties. Jackson County: roadside, Southern Illinois University campus, August 22, 1969, D. Evans & R. H. Mohlenbrock 1010. Physalis pendula Rydb. The only previously known col- lection of this species is from Union County. Jackson County: mud flats of the Mississippi River, near Fountain Bluff, October 26, 1968, D. Evans 543. Centaurea solstitialis L. Jackson County can now be added to Massac County for a second southern Illinois locality for this adventive. Jackson County: along Illinois Central Railroad, 1 mile S of Elkville, August 11, 1970, D. Evans 1091. 1972] Illinois Studies — Mohlenbrock & Evans 151 LITERATURE CITED ADAMS, P. 1957. A revision of the genus Ascyrum (Hypericaceae). Rhodora 59: 73-95. FERGUSON, A. M. 1901. Crotons of the United States. Annual Re- port of the Missouri Botanical Garden 12: 33-73. FERNALD, M. L. 1945. An incomplete flora of Illinois. Rhodora 47: 204-219. 1950. Gray's Manual of Botany, ed. 8. The Amer- ican Book Co., New York. 1632 pp. JONES, G. N. 1963. Flora of Illinois, ed. 3. The University of Notre Dame Press, Notre Dame, Indiana. 401 pp. , G. D. FULLER, G. S. WINTERRINGER, H. E. AHLES, & A. FLYNN. 1955. Vascular Plants of Illinois. The University of Illinois Press, Urbana, and the Illinois State Museum, Springfield. 593 pp. RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1964. Guide to the Vascular Flora of the Carolinas. The Book Exchange, Univer- sity of North Carolina, Chapel Hill. 383 pp. STEYERMARK, J. A. 1963. Flora of Missouri. The Iowa State Uni- versity Press. Ames. 1725 pp. DEPARTMENT OF BOTANY SOUTHERN ILLINOIS UNIVERSITY CARBONDALE, ILLINOIS 62901 CHRYSOPSIS MARIANA IN NEW ENGLAND: Until very recently, the genus Chrysopsis was believed to be rep- resented in New England by the single species C. falcata (Pursh) Ell. It is a plant of the coastal plain ranging from southeastern Massachusetts (notably Cape Cod) to New Jersey, according to Gray’s Manual, 8th edition. Another species, the wider ranging C. mariana (L.) Ell. is reported from Block Island, Washington Co., Rhode Island, by Mr. Robert Marks of Attleboro, Massachusetts, and authenticated by specimens collected by him in 1969 and 1971. In a recent letter to me (November, 1971), he states that three hundred or more plants occur in an area of approximately one quarter mile on the Lewis farm in the south-west corner of the island. There appears to be a gradual decrease in abundance from near the ocean north- ward and eastward. Vouchers are being processed for in- clusion in the herbarium of the New England Botanical Club. It is not surprising that this species of Golden Aster should turn up on Block Island. It has been known from near-by Long Island, New York, for more than one hun- dred years, e.g., Hempstead, D.C. Eaton, 1860, (NY) Judg- ing from specimens in the herbarium of Brooklyn Garden dated between 1886 and 1912, it seems to have been a fairly common plant in all sections of the island including the eastern end. I learn from correspondence that there is a recently collected specimen at the New York Botanical Gar- den from Montauk at the eastern tip (Andrews 3-84, 1962). What is surprising is that such a conspicuous plant should have escaped the attention of previous visitors to Block Island, as early as ninety years ago, and especially M. L. Fernald and companions who collected extensively there in 'W. W. Bailey in Notes on the Flora of Block Island (Bull. Torr. Bot. Club 20: 231-239, 1893) published a list of 294 vascular plants found by himself and his *youthful companion J. F. Collins" between July 19 and August 31, 1892. If seen by them, C. mariana should have been recognizable at the end of August. 152 1972] Chrysopsis — Eaton 153 August and again in September 1913.' It may have been present at that time, mostly in a vegetative state, having been suppressed by over-grazing by cattle, and only re- cently is recovering or re-capturing its former habitat among thickets of bayberry and other shrubs and stunted trees now replacing the closely-cropped turf of former pas- ture-land. Alternatively, Block Island may constitute a re- cent extension of range. I think this is unlikely. Block Island is downwind from nearby Montauk and presumably has been exposed to invasion by wind-blown seeds of Chry- sopsis for several millenia. However, the colony should be watched. Mr. Marks knows the island intimately and is confident that the Chrysopsis is localized in its southwestern corner. If it gradually spreads throughout, then recent colonization may be an attractive supposition. RICHARD J. EATON 22 DIVINITY AVENUE CAMBRIDGE, MASSACHUSETTS 02138 ERIOGONUM HARPERI GOODMAN IN TENNES- SEE: Eriogonum harperi was described by George J. Good- man in 1947 (Bull. Torr. Bot. Club 74(4) : 329-331). The description was based on material collected on a “marly glade” in Colbert County, Alabama (Roland M. Harper 3944 in 1943). Two other locations for the species, based on Harper's notes, were reported by Goodman. One was “a calcareous prairie area", also in Colbert County, and the other was “a small cedar glade” in the vicinity of Rus- sellviile in adjacent Franklin County. These three locations are within fifteen miles of each other. Our specimens (Rogers and Watson 4073, August, 1970, MISSA, TENN and University of Southern Mississippi) were collected on a limestone bluff overlooking the Caney Fork River where it intersects 1-40, about 7 to 8 miles east of the Carthage exit, Wilson County, Tennessee. Perhaps a half dozen plants were growing along the bluff near the high- way. This Tennessee locality is apparently the first new site to be discovered since the description of the species and the first site outside of Alabama. It extends the known range of the species approximately 150 miles to the Northeast. J. RAY WATSON, JR. DEPARTMENT OF BOTANY MISSISSIPPI STATE UNIVERSITY STATE COLLEGE, MISSISSIPPI KEN E. ROGERS DEPARTMENT OF BIOLOGY UNIVERSITY OF SOUTHERN MISSISSIPPI HATTIESBURG, MISSISSIPPI 39401 LATHYRUS APHACA L. NEW TO TENNESSEE AND THE SOUTHEAST: Lathyrus aphaca L. has been collected in Madison County, Tennessee, as a wild plant by the senior author. This species is a new record for Tennessee and, apparently, for the Southeast as well. Its identification was sought in all the manuals of the Eastern United States in addition to Mahler’s recent monograph (1970) to no avail. The bright yellow flowers always lead to Crotalaria in these keys which it surely is not. Dr. Robert L. Wilbur of Duke University kindly agreed to examine a specimen, and he reports the identification to be L. aphaca. According to Dr. Wilbur this is the first record of the species in the Southeastern United States. Specimen cited: "Tennessee. Madison County. 2.2 mi. S of Jackson on side of highway 45S. North bank. 15 June 1970. Richard L. Beardsley, 591. A specimen is deposited in the Memphis State Univer- sity Herbarium, and others are to be distributed elsewhere. Observations in 1971 indicate that this species is persistent and may be spreading some from the original collection site. The origin of the species in Tennessee can be only speculative. LITERATURE CITED MAHLER, W. F. 1970. Manual of the legumes of Tennessee. J. Tenn. Acad. Sci. 45(3): 65-96. RICHARD L. BEARDSLEY EDWARD T. BROWNE, JR. DEPARTMENT OF BIOLOGY MEMPHIS STATE UNIVERSITY MEMPHIS, TENNESSEE 38111 RECORDS ON THE FLORA OF WISCONSIN: Leonurus Marrubiastrum L. (Labiatae), a weedy adventive from Eu- rope, is reported to range from Pennsylvania and Delaware to Florida and Illinois. This is to announce an extension of its range into southern Wisconsin: Dane County, waste places in Kegonsa State Park, section 18, R11E, T6N, Pleasant Springs Township, Hansen 164, 27 September 1970. One sheet is deposited in the herbarium, Wisconsin State University-Oshkosh ; no duplicates were collected. Dr. Hugh Iltis has kindly confirmed the identification; there are no Wisconsin records for the plant at WIS, MIL, or UWM. R. C. Koeppen (1957. Preliminary Reports on the Flora of Wisconsin. No. 41. Labiatae — Mint Family. Trans. Wis. Acad. Sci. Arts and Letters, 46: 115-140) does not report the plant for Wisconsin. Aethusa Cynapium L. (Umbelliferae), another weedy ad- ventive from Europe, is reported by Fernald (Gray's Man- ual, 8th Edition, 1950) to range from Nova Scotia to south- ern Ontario and Minnesota, south to Delaware, Pennsyl- vania, and Ohio. Gleason & Cronquist (Manual of Vascu- lar Plants, 1963) assign it a more restricted range, from Nova Scotia and Maine to Pennsylvania and Ohio. Dr. Gerald Ownbey at MIN informs me that there are indeed two Minnescta records for the plant in their collections, the latest from 1902. However, the curators at WIS, MIL, and UWM have no Wisconsin records for the plant, which can now be tentatively added to the state's flora: Winnebago County, along fence in back yard at 212 Oxford Avenue, in the city of Oshkosh; growing as a weed in a fenceline separating two vegetable gardens. The residents on either side testify that it was certainly not planted there; Mis- terek 037, 23 September 1970. Dr. Iltis has confirmed my identification; the single specimen, deposited in the her- barium here, has abundant fruits and perhaps may become more than a mere casual element in our flora. Linaria dalmatica (L.) Mill. (Scrophulariaceae) is re- ported by Fernald (loc. cit.) to range from Cape Breton 156 1972] Wisconsin — Harriman 157 Island to Pennsylvania and Ohio. This is to report its rather extensive occurrence in Wisconsin: Bayfield County, abundant and well established on a sandy roadside on county trunk J, ca. 1.5 miles N of its junction with county trunk I, section 2, R4W, T50N, Bayfield Township; Harriman 7124, 17 July 1971. One sheet is deposited in the herbarium here, with duplicates at UW and MIL. Iltis at UW informs me (personal communication) that there are three rec- ords for the plant from Bayfield County deposited there, plus one each from Vilas, Sawyer, Sauk, Dane, and Green counties. It is certainly to be included now in the Wiscon- sin flora; it was not reported by P. J. Salamun (1951. Pre- liminary Reports on the Flora of Wisconsin. No. 36. Scro- phulariaceae. Trans. Wis. Acad. Sci, Arts and Letters. 40: 111-138). NEIL A. HARRIMAN BIOLOGY DEPARTMENT UNIVERSITY OF WISCONSIN OSHKOSH 54901 LOMENTARIA CLAVELLOSA (TURNER) GAILLON: AN ADDITION TO THE MARINE ALGAL FLORA OF NEW HAMPSHIRE. On 13 August 1971, several plants of Lomentaria clavellosa were collected at Dover Point, Dover, New Hampshire. Attached plants were found on rocks about 3 feet below mean low water near the concrete sup- port of the northbound section of the General Sullivan Bridge. It was reported in North America for the first time in 1963, from three stations in Boston Harbor, Massa- chusetts (Wilce and Lee, 1964). Since 1963, it has also been reported from Marthas Vineyard and Woods Hole in Massachusetts. The occurrence of L. clavellosa at Dover Point is not only a northward extension of its previously known North American range but it is the first record of this species from an estuarine environment in North Amer- ica. However, although Dover Point is located several miles from the open coast (it is the confluence of the Piscataqua River and Little Bay), its algal flora is unique, with definite coastal as well as estuarine species present (Mathieson, Reynolds & Hehre, in press). Voucher specimens are de- posited in the Algal Herbaria at Southampton College, the University of New Hampshire and the University of Puerto Rico at Mayaguez. I would like to thank the following people for their assistance in obtaining specimens of Lomentaria clavellosa: Messrs. R. Shepp, R. Szita, D. Fallon, Miss Dorothy Meyers and Miss Debbie Near. LITERATURE CITED MATHIESON, A. C., REYNOLDS, N. B., and HEHRE, E. J. Investiga- tions of New England Marine Algae II. Species Composition, Distribution and Zonation of Seaweeds in the Great Bay Estuary System and adjacent Coastal Waters. Nova Hedwigia (in press). WILCE, R. T. and LEE, R. W. 1963. Lomentaria clavellosa in North America. Botanica Marina 6(3/4). EDWARD J. HEHRE SOUTHAMPTON COLLEGE SOUTHAMPTON, NEW YORK 11968 158 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, particularly 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- atics 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: 177-208. 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 74. No. 797. including pages 1-159, was issued April 7. 1972. CONTENTS: — continued Somatic Chromosome Numbers for Some Asteraceae. Miloslav Kovanda eieaeoeaii a a oee a 102 Interactions between Apis mellifera (Hymenoptera: Apidae) and Tribulus cistoides (Zygophyllaceae) PO o eL mmerterénni blctlaodoneanadeteussiiuuliiaaseatene TI Redefinition of Cedrela oaxacensis C. DC. & Rose. C. Earle Smith, Jr. AEN A EE E E 124 An Abnormal Specimen of Laminaria digitata (L). Lamouroux. Arthur C. Mathieson, Emery F. Swan and Richard A. Fraliel ennea eene eaae o inten etta tatto sette aeta P eods 127 Range Extensions of Vascular Aquatic Plants in New Eng- land. C. Barre Hellquist........... esee 131 A Second Occurrence for Triphora trianthophora (SW.) Rydb. in Maine. Lesley M. Eastman ........... ee 141 Illinois Field and Herbarium Studies. Robert H. Mohlenbrock and Dam K. Evang ............ ee 142 Chrysopsis mariana in New England. Richard J. Eaton .... 152 Eriogonum harperi Goodman in Tennessee: J. Ray Watson, Jr. and Ken E. Rogers ...... eee 154 Lathyrus aphaca L. New to Tennessee and the Southeast. Richard L. Beardsley and Edward T. Browne, Jr. ........ 155 Records on the Flora of Wisconsin. Neal A. Harriman .... 156 Lomentaria clavellosa (Turner) Gaillon: An Addition to the Marine Algal Flora of New Hampshire. Edward J. Hehe ...cccorccssocsscccsssssscsssssccessssssssesoncgasovanconcstennseasere 158 Hovdova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON »Associate Editors RADCLIFFE BARNES PIKE ELIZABETH ANNE SHAW STEPHEN ALAN SPONGBERG Vol. 74 June, 1972 No. 798 CONTENTS: Revision of the Genus Melampodium (Compositae: Heli- antheae). Tod F. Stwessy essent Spores, Chromosomes and Relations of the Fern Pellaea atropurpurea. Alice F. Twryom viccccccccsssssssssssssscsssescsssesscscescece The Genus Coriaria (Coriariaceae) in the Western Hemi- Sphere. Lawrence E. Skog essees. List of the Cladoniae of Southeastern Massachusetts, with Special Reference to the Collection of Charles Albert Robbins. Lawrence B. Mish ciccccccccccssscsssssssssssssssssssssscessaene On the Marine Algae of Kent Island, Bay of Fundy. Kenneth L. Koetzner and R. D. Wood aese ( Continued on Inside Back Cover) 242 Che New England Botanical Club, Bue. 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 double-spaced throughout. Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Albion R. Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire 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 Rbodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 74 June, 1972 No. 798 REVISION OF THE GENUS MELAMPODIUM (COMPOSITAE: HELIANTHEAE)' Top F. STUESSY (Continued from page 70) II. Melampodium section Zarabellia (Cass.) DC. Melampodium L. sect. Zarabellia (Cass.) DC. Prodr. 5: 519. 1836. Zarabellia Cass. Dict. Sc. Nat. 59:240. 1829. Type species: Melampodium longifolium Cerv. ex Cav. Tap-rooted annuals; leaves rhombic to deltoid (rarely narrowly ovate), sessile or shortly petiolate, with margins entire to obscurely serrate; outer involucre usually cupu- late or less often spreading, with bracts 3-5, at margins herbaceous; ovaries of the disc florets ovoid, less than 0.7 mm long, rudimentary (Figs. 3 & 4) ; fruits at apex nearly smooth or moderately sculptured, or sometimes with an abaxial protuberance (in M. longifolium, Fig. 7) ; chromo- some base number, x — 9. Species 21-25. 21. Melampodium longifolium Cerv. ex Cav. Anal. Cien. Nat. 6:333. 1803. non Brouss. ex Willd. 1809. TYPE: "NUEVA HISPANIA": grown from seed in the Royal Bot. Gard. Madrid, flowering Jun-Sep, V. Cervantes s.n. ( Holo- type, MA?; isotype, G!; photograph of G-DC isotype, IDC 800. 927: III. 6!; photograph of G isotype, F! us!). Melampodium longifolium Brouss. ex Willd. Enum. Plant. Hort. Berol. 934. 1809. nom. illegit. non Cerv. ex Cav. 1803. 161 162 Rhodora [Vol. 74 TYPE: MEXICO: locality, date, and collector unknown, culti- vated in Bot. Gard. Berlin and harvested in 1808 (Holotype, B; photograph of holotype, TEX!). Zarabellia rhomboidea Cass. Dict. Sc. Nat. 59:241. 1829. TYPE: cultivated in Bot. Gard. Paris, source of seeds un- known (Holotype, P?; photograph of possible G-DC isotype, Ipc 800. 927: III. 7!). Dysodium divaricatum Hort. ex DC. Prodr. 5:520. 1836. non Rich. in Pers. 1807. nom. illegit., pro syn. Melampodium rhomboideum (Cass.) DC. Prodr. 5:520. 1836. Melampodium parvulum Brandg. Zoe 5:262. 1908. TYPE: MEXICO: Puebla, Cerro Santa Lucia, Jul 1907, C. A. Purpus 2813 (not 2813’) (Holotype, UC!; isotypes, F! GH! NY! US!; photograph of US isotype, TEX!). Annual herbs, 8-45 cm long. Stems erect, 1.5-4 mm diam, moderately hispid near base to copiously hispid above with hairs 0.8 mm long. Peduncles 0-21 mm long. Leaves sessile, rhombic (rarely elliptic, younger leaves often lanceolate- ovate), 2.5-6 cm long, 0.5-2.7 cm wide, at apex acute, at base subauriculate, with both surfaces moderately pubescent with hairs 0.3 mm long; margin obscurely serrate. Heads 3-4 mm tall, 5-7 mm diam. Outer involucre spreading, 7-10 mm diam; bracts 5 (rarely 4), separate, lanceolate, 3-5 mm long, 1.2-2 mm wide, at apex acute-acuminate, with abaxial surface glabrous to moderately pubescent (very pubescent at base) with hairs 0.7 mm long; margin herbaceous. Fruits 2.2-3 mm long, with lateral surfaces with 2 large irregular cavities. Ray florets 5-8; ligules yellow, ovate- elliptic, 1.3-1.5 mm long, 0.8-0.9 mm wide. Disc florets 10- 15; corollas yellow, 0.8 mm diam, with throat 0.7 mm and tube 0.6 mm long. Paleae oblong, 1.4 mm long, 0.7 mm wide; apex colorless, with margin erose; midrib distinct, glabrous. Chromosome number, » — 9. Scattered throughout pine-oak forests in the Mexican states of Nuevo Léon, San Luis Postosi, México, Distrito 1972] Melampodium — Stuessy 163 Federal, Puebla, and Oaxaca (Fig. 12), 1830-2740 m. Flow- ering dates, Aug-Sep. REPRESENTATIVE SPECIMENS. Mexico. DISTRITO FEDERAL: Tlalpan, Pedregal, Valley of Mexico, 20 Aug 1896, Harshberger 176 (GH); Tlalpan, Pyramid of Cuicuilco, 15 Aug 1935, MacDaniels 53 (F); Valley of Mexico near Tlalpan, 20 Aug 1896, Pringle 6455 (F, GH, NY, UC, US); Pena Pobre near Tlalpan, 8 Aug 1944, Sharp 44208 (NY). MEXICO: near El Oro, 14 Sep 1901, Pringle 9325 (F, GH, US). NUEVO LEON: Hacienda Pablillo, Galeana, 8 Aug 1936, Taylor 115 (DS, F, NY, RSA, TEX). OAXACA: N limits of Las Sedas, 15 Aug 1966, Stuessy 659 (TEX). PUEBLA: vicinity of Puebla, 9 Sep 1906, Arséne 76 (vs), 27 Oct 1907, Arsène 1199 (US), 10 Sep 1910, Nicolas 5447 (GH, US), s.n. (NY); Teocalli de Cholula, near Puebla, 25 Sep 1909, Nicolas s.n. (NY); Rancho Posadas, near Puebla, 25 Sep 1909, Nico- las sn. (NY); vicinity of San Luis Tultitlanapa, Aug 1908, Purpus 2813’ (F, GH, NY, UC, US). SAN LUIS POTOSÍ: vicinity of San Luis Potosi, 1878, Parry & Palmer 444 (F, GH, NY, US); 22 mi E of San Luis Potosí, 5 Aug 1961, Powell & Edmondson 551 (F, TEX). 22. Melampodium mimulifolium Robins. Proc. Amer. Acad. Arts & Sci. 36:462. 1901. TYPE: MEXICO: Oaxaca, Totonto- pec, 1700-2150 m, 15 Jul 1894, E. W. Nelson 740 (Holotype, US!; photograph of holotype, TEX !). Annual herbs, over 30 cm tall. Stems erect, 2-3 mm diam, subglabrous below to pilose-villous above with hairs 1 mm long. Peduncles 3-4.2 cm long. Leaves sessile, lanceolate- elliptic, 4.5-6.8 cm long, 1.5-2.2 cm wide, at apex acuminate, at base subauriculate to auriculate, with both surfaces stri- gose with hairs 0.4-1 mm long; margin entire to serrulate. Heads 6-7 mm tall, 8-10 mm diam. Outer involucre spread- ing to shallowly cupulate, 8-9 mm diam; bracts 5, slightly connate at base, ovate-lanceolate, 5-6 mm long, 2-2.5 mm wide, at apex acute, with abaxial surface moderately pilose with hairs 0.5-1 mm long; margin herbaceous. Fruits 2-2.1 mm long, with lateral surfaces smooth-striate with enlarged tuberculate margins. Ray florets ca 8; ligules yellow-orange, ovate, 3-4 mm long, 1.5-3 mm wide. Disc florets ca 50; corollas yellow, 1 mm diam, with throat 0.8 mm and tube 1.4 mm long. Paleae oblanceolate-obovate, 2.2 mm long, 0.8 mm wide; apex yellow, with margin erose- 164 Rhodora [Vol. 74 laciniate; midrib weak, glabrous. Chromosome number un- known. Known only from the type collection from pine-oak for- ests at Totontopec, Oaxaca, Mexico (Fig. 12), 1700-2150 m. Flowering date, Jul. M. mimulifolium is very similar to M. gracile, and in fact, additional collecting may disclose transitional specimens that might suggest merger of the two species. At present, however, M. mimulifolium seems treated best as a distinct species that can be distinguished from M. gracile by the former's shorter peduncles, narrower leaves, longer and narrower outer phyllaries, and absence of stipitate-glandu- lar hairs anywhere on the plant. 23. Melampodium gracile Less. Linnaea 6:407. 1831. TYPE: MEXICO: Veracruz, Papantla, Jan 1829, C. J. W. Schiede & F. Deppe 1254 (Lectotype, HAL!; isotypes, HAL! P! w! Mo! photograph of lectotype, 0S! TEX!; photograph of HAL iso- type, OS! TEX!; photograph of P isotype, oS! TEX!). Melampodium oblongifolium DC. Prodr. 5:519. 1836. TYPE: MEXICO: Veracruz, “rancho de los huevos” [from label] near Tantoyuca, Jan 1832, J. L. Berlandier 2153 [= 733] (Lectotype chosen, G-DC; isotypes, F! G [2]! GH! K [2]! MO! P!; photograph of lectotype, IDC 800. 927: III. 2!; trac- ing of lectotype, GH!; photograph of G-DC isotype, IDC 800. 927: IIL 3!; photograph of K isotypes, vus [2]!). Melampodium gracile Less. var. oblongifolium (DC.) A. Gray, Proc. Amer. Acad. Arts & Sci. 5:182. 1861. Melampodium microcarpum S. F. Blake, Contrib. U. S. Nat. Herb. 22:606. 1924. TYPE: MEXICO: Tamaulipas, vi- cinity of Gómez Farias, ca 350 m, 13-21 Apr 1907, E. Palmer 319 (Holotype, US!; isotype, GH! MO!; photograph of holotype, TEX! UC!). Annual herbs, 17-50 cm tall. Stems erect, 1.2-4 mm diam, hispid-pilose and weakly stipitate-glandular with hairs 0.1-2 mm long. Peduncles 5.5-11.5 cm long. Leaves sessile, 1972] Melampodium — Stuessy 165 eri WA ] n » k bs. / Va Fig. 21. Map of Mexico and adjacent Guatemala and British Honduras showing distribution of Melampodium gracile (triangles), and M. microcephalum (dots). usually deltoid (younger leaves often lanceolate), 5-10 em long, 2-4 cm wide, at apex acute to acuminate, at base nar- rowed and dilated near stem to subauriculate (rarely ob- tuse), with both surfaces strigillose with hairs 0.2 mm long; margin obscurely crenate to irregularly cleft. Heads 6-7 mm tall, 8-15 mm diam. Outer involucre spreading to shal- lowly cupulate, 7-10 mm diam; bracts 3 (sometimes 4 or 5), slightly connate at base, imbricate 1/3 their length, ovate, 3.5-5 mm long, 2.5-3.5 mm wide, at apex acuminate, with abaxial surface strigose with hairs 0.3-1 mm long near apex, near base stipitate-glandular with hairs 0.1-0.2 mm long; margin herbaceous. Fruits 2.3-2.5 mm long, with lat- eral surfaces reticulate and striate to tuberculate. Ray florets 5-8; ligules yellow-orange, ovate, 2-5 mm long, 3-4 mm wide. Disc florets 30-45; corollas yellow-orange, 1.5 166 Rhodora [Vol. 74 mm diam, with throat and tube each 0.9 mm long. Paleae oblanceolate-obovate, 2.2 mm long, 0.8 mm wide; apex yellow-orange, with margin entire to dentate-laciniate lat- erally; midrib absent. Chromosome number, n — 9. Pine-oak, tropical deciduous, tropical evergreen, and rain forests on the Gulf side of the Sierra Madre Oriental and the Yucatan Peninsula, and on the Pacific sides of the Sierra Madres Occidental and Sur, with extensions into British Honduras and northwestern Guatemala (Fig. 21), 30-1830 m. Flowering dates, Jul-Oct. Melampodium gracile is very similar morphologically and geographically to M. microcephalum, but the two differ in several features such as habit, and the shapes of leaf bases, outer involucral bracts, and ligules. Single popula- tions of both species have been crossed reciprocally in the greenhouse, and although robust F, hybrids are obtained, they are almost completely sterile. Detailed analyses of hybridization among all three species in the genus that have stipitate-glandular peduncles (M. gracile, M. micro- cephalum and M. paniculatum) currently are in progress and the results will be reported in a separate paper. REPRESENTATIVE SPECIMENS. British Honduras. EL CAYO: San Agus- tin, Jul-Aug 1936, Lundell 6759 (ARIZ, GH, NY, TEX [2], Us). Guate- mala. HUEHUETENANGO: between San Sebastián H. & San Rafael Pétzal, 14 Aug 1942, Steyermark 50539 (F); between Nentón & Miramar, 29 Aug 1942, Steyermark 51452 (F, UC). PETÉN: vicinity of La Libertad, Aug-Nov 1933, Aguilar 25 (F [photograph of US specimen], NY, US). Mexico. CAMPECHE: Champotón, 7-15 Jul 1932, Steere 1805 (us); 1 mi W of Champotón, 18 Jul 1966, Stuessy 531 (TEX); 16 mi N of Champotón, 18 Jul 1966, Stuessy 533 (TEX); 12 mi S of Campeche, 25 Jul 1966, Stuessy 546 (TEX). CHIAPAS: ca 28 mi SE of Comitán, 25 Jun 1960, King 3042 (DS, NY, TEX, UC, US); ca 17 mi S of Tuxtla Gutiérrez, 27 Jun 1960, King 3096 (DS, NY, TEX, uc, Us); 10 mi S of Bochil, 25 Jul 1966, Stuessy 562 (TEX); 32 mi SE of Comitán, 26 Jul 1966, Stuessy 574 (TEX). COLIMA: 7 mi S of Colima, 25 Aug 1966, Stuessy 722 (TEX). GUERRERO: Acuitlapan, 24 Sep 1937, Abbott 439 (GH); Rio Balsas, Aug 1910, Orcutt 4365 (F); near Iguala, 14 Sep 1900, Pringle 9177 (F, GH, SMU, UC, US), 10 & 12 Aug 1905, J. N. & J. S. Rose, & Painter 9308 (GH, US). HIDALGO: Jacala, 20 Oct 1937, Kenoyer 427 (F). MÉXICO: Temas- 1972] Melampodium — Stuessy 167 caltepee, Anonas, 24 Jul 1934, Hinton 6338 (US). MICHOACÁN: ca 4 mi E of Jiquilpán, 6 Aug 1960, King 3636 (DS, NY, TEX, UC, US) ; ca 5 mi E of Zamora, 6 Aug 1960, King 3646 (DS, NY, TEX, UC, US); 30 mi NW of Zamora, 2 Sep 1965, Stuessy 393 (TEX); 25 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 694 (TEX) ; 2 mi S of jct rte 15 & rd to Cotija, 24 Aug 1966, Stuessy 713 (TEX). MORELOS: ca 11 mi 8 of Cuernavaca, 14 Mar 1961, King 4165 (TEX) ; 6 mi W of Yautepec, 14 Aug 1950, Pipes 81 (SMU); valley near Jojutla, 17 Jun 1901, Pringle 8466 (F, GH, NY, POM, UC, US); near Yautepec, 27 Oct 1902, Pringle 9937 (F, GH, Ny, US); 7 mi NW of Cuautla, 25 Aug 1965, Slessy 356 (TEX). SAN LUIS POTOSÍ: El Pujal, Río Tampaón valley, 21 Jul 1939, Chase 7533 (ARIZ, F, GH, NY); El Salto, 15 Jun 1951, Holman & Dittman 791 (SMU) ; El Salto, 14 Jun 1951, Kay & Higgins 202 (SMU); El Salto, 20 Feb 1961, King 3887 (F, NY, TEX, US); 48 mi N of Tamazunchale, 23 Aug 1957, Waterfall & Wallis 14288 (smu). TAMAULIPAS: 7 mi N of Padilla, 9 Jun 1951, Paynet & Hulan 1213 (SMU, TEX); 10 km NW of El Progreso, 22 Aug 1941, Stanford, Retherford & Northeraft 1078 (ARIZ, DS, GH, NY) ; 15 mi SW of Ciudad Mante, 9 Aug 1965, Stuessy 274 (TEx [2]); ca 3-4 mi E of Nuevo Morelos, 5 Jul 1966, Stuessy 463 (TEX); 3 mi E of Gómez Farias, 8 Jun 1967, Stuessy 826 (TEX). VERACRUZ: Huasteca, Wartenberg, near Tantcyuca, 1858, Ervendberg 92 (GH); 13 mi W of Orizaba, 18 Nov 1959, Graham & Johnston 4777 (TEX); 7 mi SW [SE] of Morelos, 12 Aug 1961, Powell & Edmondson 646 (F, TEX): Jalapa, 1894, Smith 1605 (au, UC); 18 mi S of ject rtes 110 & 105, 6 Jul 1966, Stuessy 466 (TEX). YUCATAN: Progreso, Gawmer 2349 (F); Izamal, 1895, Gaumer & sons 789 (Ds, F, GH [2], Ny [2], pom, vc [3], US); at km 13 on Merida-Progreso rd, May-Aug 1938; C. & A. Lundell 7972 (ps, F, TEX, UC, US); 13 km N of Merida, 20 Jul 1966, Stuessy 537 (TEX); 86 mi W of Valladolid, 21 Jul 1966, Stuessy 544 (TEX [2]). 24. Melampodium microcephalum Less. Linnaea 9:268. 1854. TYPE: MEXICO: in cultivation near Hacienda de la Laguna, Jul 1829, C. J. W. Schiede 217 (Holotype, HAL!; isotype, P!; photograph of holotype, os! TEX!; photographs of P isotype, F! os! TEX! US!). Melampodium lanceolatum Sessé & Moc. Fl. Mex. ed. 2. 196. 1894. nom. illegit. non DC. 1836. TYPE: “calidoribus Novae Hispaniae regionibus,” Jul 1795-1804, M. Sessé et al. “3978” (Lectotype chosen, MA! [drawing of stem with two pairs of leaves on reverse side of label] ; isotypes, MA [3]!). 168 Rhodora [Vol. 74 Annual herbs, 15-75 cm tall. Stems erect to decumbent and rooting at nodes, 1-2 mm diam, villous and weakly stipitate-glandular with hairs 0.1-1 mm long. Peduncles 2.5.6.5 cm long. Leaves sessile or with petioles 1-4 mm long, ovate-lanceolate, 3-6.5 cm long, 0.9-3.5 cm wide, at apex acute to obtuse, at base attenuate to obtuse, with both surfaces strigose with hairs 0.2-1 mm long; margin entire to obscurely crenate. Heads 3-4 mm tall, 5-10 mm diam. Outer involucre spreading to shallowly cupulate, 5-8 (-10) mm diam; bracts 3 (sometimes 4 or 5), slightly connate at base, separate, narrowly ovate, 3-4.5 mm long, 1.5-3 mm wide, at apex acuminate, with abaxial surface pilose-villous and stipitate-glandular with hairs 0.1-1 mm long; margin herbaceous. Fruits 1.7-2 mm long, with lateral surfaces rugose-reticulate with enlarged tuberculate margins. Ray florets 5-8; ligules yellow-orange, elliptic, 2.5-4 mm long, 2.9 mm wide. Disc florets 35-50; corollas yellow-orange, 1.6 mm diam, with throat 1 mm and tube 0.7 mm long. Paleae oblanceolate-obovate, 2 mm long, 0.8 mm wide; apex yellow, with margin serrulate-laciniate; midrib absent. Chromo- some number, n = 9. Mostly pine-oak, tropical deciduous and tropical ever- green forests in many states of Mexico in the Mesa Central and the Sierra Madres Occidental and Sur, and in adjacent Guatemala (Fig. 21), 50-1740 m. Flowering dates, Jun- Nov. M. microcephalum is often confused with M. gracile and M. paniculatum. Although all these taxa possess three outer involueral bracts, stipitate-glandular stems and peduncles, and chromosome numbers on a base of x — 9, the size of the heads and the shapes of the leaf bases can be used to distinguish easily one from another (see key). In most taxa of Melampodium the number of outer in- volucral bracts is very constant, but in both M. microcepha- lum and M. gracile, as seen from field and greenhouse studies, plants occasionally are found with four or five bracts instead of the usual three. 1972] Melampodium — Stuessy 169 REPRESENTATIVE SPECIMENS. (Guatemala. HUEHUETENANGO: ca 6 mi S of Huehuetenango, 18 Jul 1960, King 3425 (DS, NY, TEX, UC, US). Mexico. CHIAPAS: 10 mi SE of Tonalá, 10 Aug 1966, Stuessy 627 (TEX). GUANAJUATO: near Acambaro, 18 Oct 1904, Pringle 13069 (F, GH, SMU, UC). GUERRERO: Taxco, 28 July 1937, Abbott 304 (GH); 20 mi NE of Acapulco, 20 Aug 1947, Barkley, Webster & Paxson 17M732 (TEX); Mina, Manchón, 7 Aug 1936, Hinton 9205 (ARIZ); Mina, Río Frio, 25 Sep 1937, Hinton et al. 10719 (ARIZ, GH, POM, TEX, US). JALISCO: Cerro Viejo, above Zapotitan de Hidalgo, ca 45 mi S of Guadalajara, 27 Jun 1956, Gregory & Eiten 205 (SMU). MÉXICO: Temascaltepec, Volcán, 10 Sep 1932, Hinton 1661 (F, GH, NY [2], US); Temascaltepec, Bejucos, 6 Oct 1932, Hinton 2008 (Ds, F, NY [2], US) ; Temascaltepec, Ixtapán, 22 Jun 1934, Hinton et al. 6207 (F, GH, US). MICHOACÁN: 7 mi W of La Piedad, 6 Oct 1962, Cronquist 9613 (NY, TEX, US) ; ca 3 mi S of Ciudad Hidalgo, 4 Aug 1960, King 3607 (Ds, NY, TEX, UC, US); ca 22 mi S of Uruapan, Cerro de Carboneras, 16-22 Oct 1961, King & Soderstrom 4804 (NY, SMU, TEX, UC, US); Lake Cuitzeo, 26 Oct 1892, Pringle 4322 (F, GH [2, one a tracing], Ny, vc [2], US); 6 mi NW of Tuxpán, 1 Sep 1965, Stuessy 383 (TEX). MORELOS: El Teycal, 9 km SW of Tepoztlan, 30 Aug 1955, Clausen s.n. (NY); Cuantla, Aug 1904, Kuntze 23527 (NY [3]) ; Yautepec, 30 May 1939, Nagel 8030 (GH); Barranca, near Cuernavaca, 2 Nov 1896, Pringle 7321 (GH, US); Moyotepec, 8 Nov 1964, Ripley & Barneby 13708 (NY). OAXACA: Cerro de San Felipe, 26 Sep 1897, Conzatti & González 550 (GH, US); 10-12 km E of Niltepec, 18 Jul 1959, King 1833 (NY, SMU, TEX, UC, US) ; vicinity of Choapám, 28-29 Jul 1894. Nelson 870 (US); 1 mi E of Monte Alban, 12 Aug 1966, Stuessy 638 (TEX); 29 mi S of Zimatlán, 12 Aug 1966, Stuessy 640 (TEX). PUEBLA: Mt Orizaba, Botteri 809 (F, GH); near Puebla, Jul 1888, Hoffmann 832 (GH); Orizaba, 27 Mar 1887, Mills- paugh s.n. (F); Mt Orizaba, 24 Aug 1891, Seaton 461 (F, GH, NY, US). VERACRUZ: Tantoyuca, Dec 1830, Berlandier s.n. (GH); valley of Córdoba, 20 Jan 1866, Bourgeau 1628 (GH, US) ; valley of Córdoba, 23 Apr 1865-66, Bourgeau 1629 (US); Pacho, near Jalapa, Jan 1894, Smith 1655 (F, NY, RSA). 25. Melampodium paniculatum Gardn. Hook. Lond. Jour. Bot. 7:287. 1848. TYPE: BRAZIL: Goias, near Arraias, Apr [Feb?] 1840, G. Gardner 3844 (Holotype, K!; isotypes, F [2]! c [2]! Ny [2]! P! w!; photograph of holotype, us!; photograph of G isotype, F! US!). Melampodium laxum Sch. Bip. Ber. Bot. Zeit. 3:155. 1845. nom. nud. Based on G. Gardner 3844. 170 Rhodora [Vol. 74 Melampodium brachyglossum J. D. Smith, Bot. Gaz. 13: 74. 1888. TYPE: GUATEMALA: Alta Verapaz, Coban, 4300 ft, May 1886, H. von Tuerckheim 114 (Lectotype chosen, US!; isotype, US!; photograph of holotype, TEX!). K, NY & P specimens that bear exactly the same printed label as the lectotype are clearly Jaegeria hirta (Lag.) Less. (see Torres, 1968, for agreement). It is probably a labeling error. Annual herbs, 13-70 em tall. Stems erect, 0.7-5 mm diam, hispid and stipitate-glandular with hairs 0.1-1 mm long. Peduncles 1.1-5 cm long. Leaves sessile or with petioles up to 5 mm long, ovate-rhombic (younger leaves approaching lanceolate), 3-9 cm long, 1-5 cm wide, at apex acute to acuminate, at base attenuate, often dilated almost to sub- auriculate, with both surfaces strigose with hairs 0.3-1 mm long; margin entire to irregularly serrate-crenate. Heads 2.2-4.5 mm tall, 3-5.5 mm diam. Outer involucre cupulate, 5-7 mm diam; bracts 3, separate, ovate, 1.8-4 mm long, 1.5-2.8 mm wide, at apex acute-acuminate, with abaxial surface strigose and stipitate-glandular with hairs 0.1-0.8 mm long; margin herbaceous. Fruits 2-2.8 mm long, with lateral surfaces with longitudinal ribs and striations. Ray florets 3-5; ligules yellow, ovate, 1-2 mm long, 1-2 mm wide. Disc florets 10-15; corollas yellow, 1.1 mm diam, with throat and tube each 0.8 mm long. Paleae elliptic, 1.3 mm long, 0.7 mm wide; apex yellow-orange, with margin laciniate; midrib weak, glabrous. Chromosome numbers, n = 8 and 27. Low tropical mountains of Chiapas, Mexico, Central America (excluding British Honduras, Nicaragua, and Pan- ama), and isolated points of Colombia and Brazil in South America (Figs. 22 & 23), 370-2010 m. Flowering dates, Jan-Dec. Adding to the chromosomal diversity already existing in Melampodium are the new counts in M. paniculatum of n = 27 reported from two different populations in Brazil 1972] Melampodium — Stuessy 171 Fig. 22. Map of Central America showing distribution of Me- lampodium paniculatum. See Fig. 23 for South American localities. 172 Rhodora [Vol. 74 Fig. 23. Map of South America showing distribution of Melampo- dium divaricatum (dot), and M. paniculatum (circles). by Coleman (1970)'*. I have examined the cited voucher specimens, and due to an absence of significant morpholog- ical differences between the » — 18 and n — 27 cytotypes, it seems best at this time to consider these chromosomally distinct populations as simply polyploid races of a single species. It is interesting to note, however, that the tetra- ploid counts of n = 18 prevail in Central America while the chromosomally derived hexaploid populations of m — 27 are found only in Brazil where M. paniculatum prob- ably has been recently introduced by man. M. paniculatum thus joins M. cinereum, M. dicoelocarpum and M. leucan- thum as the fourth species in the genus known to have intraspecific euploidy. "These new counts actually were listed as » — 26-27 and n — 26. In recent correspondence regarding these counts, however, Dr. Cole- man states that “On restudying the slide, I have decided that the correct number of the material I counted is n — 27." 1972] Melampodium — Stuessy 173 REPRESENTATIVE SPECIMENS. Brazil. FEDERAL DISTRICT: Córrego Landim, ca 20 km N of Brasilia, 15 Dee 1965, Irwin, Souza & Santos 11303 (TEX); Corallino, 1817-21, Pohl 1368 [1276] (NY). GOIAS: near Arraias, Apr 1840, Gardner 287 (NY). MATO GROSSO: Sta Terezinha, 12 Feb 1948, Macedo 1408 (US). MINAS GERAIS: 1838, Clausen s.n. (F, NY [2], Us); Belo Horizonte, Pampulha, Feb 1945, Williams 5643 (vc, US). RÍO DE JANEIRO: Río de Janeiro, Feb 1882, Glaziou 12794 (K). SAO PAULO: São Joao de Boa Vista, 6 Sep 1893, Lofgren & Edwall 16695 (Us). Colombia. CUNDINAMARCA: Bogotá, Triana 1331 (NY, US). Costa Rica. CARTAGO: 24 Aug 1924, Torres 50 (Us). GUANACASTE: coffee plantations of San Francisco de Guadalupe, Jul 1892, Pittier 6963 (GH); Hatillo, 4 Jan 1935, Solís 52 (F). HEREDIA: Río Virilla on hwy to Heredia, 1 Dec 1937-1 Jan 1938, Allen 585 (F); Río Virrilla near La Uruca, Aug 1888, Pittier 429 (F, GH); Río Virilla near San Juan, Jun 1896, Tonduz 10144 (vs [2]). SAN JOSE: San José, 4 Sep 1938, Orozco 260 (F); vicinity of San José, 6 Nov 1938, Orozco 418 (F); vicinity of Las Pavas, 29 Feb 1924, Standley 36058 (US); vicinity of San José, Feb 1924, Standley 38971 (vs); Desamparados, 27 Jun 1923, Stevens 136 (vs). EI Salvador. AHUA- CHAPÁN: vicinity of Ahuachapán, 9-27 Jan 1922, Standley 20260 (GH), 16-25 Jan 1947, Standley & Padilla 2755 (F). LA LIBERTAD: Finca Paraíso, 4 mi S of Sta Tecla, 14 Jan 1946, Carlson 106 (F, UC, US) ; Sta Tecla, 20 Jul 1941, García 51 (UC). SAN SALVADOR: Volcán de San Salvador, Apr 1922, Calderón 459 (GH, NY, US) ; San Salvador, 1922, Calderón 779 (GH, NY, US); vicinity of San Salvador, 30 Mar-24 Apr 1922, Standley 22687 (GH, US); Volcan de San Salva- dor, 7 Apr 1922, Standley 22864 (GH, NY, US); San Salvador, Nov 1906, Velasco 8946 (US). SANTA ANA: Cerro Miramundo, above Hacienda Los Planos, NE of Metapán, 25 Feb 1946, Carlson 873 (F); Finca of Los Naranjos, E of Cerro de los Naranjos, 27 Apr 1942, Tucker 1313 (F, NY, UC, US). SAN VICENTE: vicinity of San Vicente, 7-14 Feb 1947, Standley & Padilla 3630 (F). Guatemala: ALTA VERA- PAZ: ca 4 mi NE cf San Pedro Carcha, 12 Jul 1960, King 3329 (ps, NY, TEX, UC, US); 28 mi E of San Miguel Uspantán, 28 Jul 1966, Stuessy 589 (TEX); 1 mi NE of San Pedro Carchá, 29 Jul 1966, Stuessy 594 (TEX): Coban, Jul 1885, Tuerckheim 761 (us), Mar 19083, 8417 (F. GH, NY, US). CHIMALTENANGO: near Río Pixcavó between Chimaltenango & San Martin Jilotepeque, 3 Feb 1939, Standley 64303 (F, NY); Finca La Alameda near Chimaltenango, 11-22 Dec 1940, Standley 79869 (F). EL QUICHE: Nebaj, 19 Nov 1934, Skutch 1709 (F, GH); 4 mi E of Cunén, 28 Jul 1966, Stuessy 585 (TEX); 11 mi E of San Miguel Uspantan, 28 Jul 1966, Stuessy 587 (TEX). ESCUINTLA: Finca Monterrey, Volcán de Fuego, 5 Feb 1939, Standley 64595 (F, NY); Los Diamantes, 9 Mar 1896, Volkem 2564 (GH, NY). GUATE- MALA: near Finca La Aurora, 1938-39, Aguilar 322 (F). HUEHUE- 174 Rhodora [Vol. 74 TENANGO: ca 3 mi S of Huehuetenango, 18 Jul 1960, King 3417 (DS, NY, TEX, UC, US); vicinity of Cuilco near Rio Cuileo, 17 Aug 1942, Steyermark 50773 (F, GH); 2 mi S of Huehuetenango, 26 Jul 1966, Stuessy 578 (TEX); 1 mi W of Aguacatán, 27 Jul 1966, Stuessy 582 (TEX); Chaculá, 20 Aug 1896, Volkem 2774 (GH, NY, US). JALAPA: vicinity of Jalapa, 7-18 Nov 1940, Standley 76429 (F); between Jalapa & San Pedro Pinula, 12 Nov 1940, Standley 77073 (F); mts near Chahuite, NW of Jalapa, 16 Nov 1940, Standley 77440 (F); between Jalapa & Montana Miramundo, 7 Dec 1939, Steyermark 32847 (F). QUEZALTENANGO: vicinity of San Francisco de Miramar, Costa Cuca, Apr 1905, Pittier 57 (F, US), Pittier 60 (F, US); between Colomba & Las Mercedes, 20 Feb 1941, Standley 87933 (F); San Francisco Miramar, 15 Mar 1921, Tonduz & Rojas 203 (Us). RETALHULEU: Finca Helvetia, 6 May 1937, Muenscher 12467 (F). SACATEPÉQUEZ: Antigua, 12 Sep 1959, O. & I. Degener 26504 (US); Antigua, 13 Feb 1905, Kellerman 4501 (F, US); Antigua, Apr 1890, Smith 2334 (us); near Antigua, Nov 1938-Feb 1939, Standley 60326 (F, NY); Barranca above Duefias, 21 Jan 1939, Standley 63265 (F, NY). SAN MARCOS: Río Cabüs near Malacatán, 15 Mar 1939, Standley 68866 (F). SANTA ROSA: near Cerro Redondo, 20 Dec 1938, Standley 60413 (F); La Joya de Limón, E of Cuilapa, 25 Nov 1940, Standley 78268 (F). soLotA: Panajachel, near Lake Atitlán, 11 Sep 1959, O. & I. Degener 26559 (US) ; behind Lago de Atitlán near Panajachel, 5 Jul 1960, King 3242 (ps, NY, TEX, UC, US). SUCHITEPÉQUEZ: Finca Moca, 27 Oct 1934, Skutch 1550 (GH). Honduras. CHOLUTECA: vi- cinity of San Marcos de Colón, 12-22 Jan 1949, Standley 15756 (F). COMAYAGUA: vicinity of Siguatepeque, 14-27 Feb 1928, Standley 56290 (vs), 56522 (F, US), 25 Mar-5 Apr 1947, Standley & Chacón 6625 (F), 6660 (r); Rittenhouse's hacienda, 30 Jun 1936, Yuncker, Dawson & Youse 5519 (F, GH). EL PARAÍSO: Güinope, Dec 1943, Valerio 1662 (F). INTIBUCA: vicinity of La Esperanza & Intibucá, 31 Jan-12 Feb 1950, Standley 25149 (F). MORAZÁN: Río Yeguare, 22 Jul 1948, S. F. Glassman 2003 (r, Nv), 26 Aug 1951, Williams 18257 (F); near Río Yeguare, below El Zamorano, 10 Aug 1947, Standley 12105 (F); Cerro de Uyuca between Las Flores & Tatumbla, 17 Aug 1949, Standley 22718 (F); La Leona, Tegucigalpa, 6 Sep 1951, Standley 28678 (F); Zamorano, 7 Jan 1946, Valerio 3735 (F, GH). OLANCHO: vicinity of Juticalpa, 5-16 Mar 1949, Standley 17710 (F). Mexico. CHIAPAS: Tenejapa, Pokolum, 10 Jul 1964, Breedlove 6085 (ps, F); Chamula, Toh Tik on rd to Chenalho, 12 Dec 1964, Breedlove 7861 (ps, F); Los Lagos, 3 mi NW of Rancho San José, 18 Jan 1952, Carlson 2254 (F). 1972] Melampodium — Stuessy 175 III. Melampodium section Serratura Stuessy, sect. nov. Herbae annuae radicibus palaribus; folia rhombica vel deltoidea (raro anguste ovata), petiolata, marginibus ser- ratis (raro integris); involucrum extimum cupulatum vel interdum patens, bracteis 5, marginibus herbaceis; ovaria flosculi disci ovoidea, minus quam 0.7 mm longa, rudimen- tales (Figs. 3 & 4) ; fructus apice fere laeves vel moderate sculpti et sine cucullis, vel complanata abaxiali arista (M. tepicense) ; chromosomatum numerus basicus, x = 12. Species 26-30. Typus: Melampodium divaricatum (Rich. in Pers.) DC. 26. Melampodium divaricatum (Rich. in Pers.) DC. Prodr. 5:520. 1836. Dysodium divaricatum Rich. in Pers. Syn. 2:489. 1807. non Hort. ex DC. 1836. TYPE: COLOMBIA: “Gairam, prope St. Martham," 1785-89, L. C. M. Richard s.n. (Lectotype chosen, P!; isotype, P!; photograph of holotype, os! TEX!; photograph of P isotype, os!). Wedelia ovatifolia Willd. Enum. Suppl. 61. 1814. TYPE: source of seed unknown, apparently grown in Bot. Gard. Berlin, “2421” (Holotype, B?; isotype, w!; photograph of w isotype, OS! TEX!; photograph of probable G-DC isotype, IDC 800. 928: I. 4!). Alcina ovatifolia (Willd.) Jacq. f. Eclog. P1. 1:115. t. 78. 1815. Wedelia minor Hort. ex Hornem. Enum. Hort. Hafn. 2:855. 1815. TYPE: introduced into the Bot. Gard. Copen- hagen in 1812 from Berlin Bot. Gard., source of seed unknown (Lectotype, c!; isotypes, C [2]!; photograph of lectotype, oS! TEX!). Alcina ovalifolia Lag. Gen. et Sp. Nov. 32. 1816. TYPE: "NOVA HISPANIA" [MEXICO] : seeds cultivated and grown in Bot. Gard. Madrid, 1805, M. Sessé s.n. (Holotype, MA!; isotype, G!; photograph of holotype, os! TEX!; photograph of G isotype, OS! TEX!). 176 Rhodora [Vol. 74 Melampodium paludosum H.B.K. Nov. Gen. Sp. 4:287. 1820. TYPE: PANAMA: “in humidis prope ostia fluminis sinu, juxta litora Dariensis," Mar 1801, F. H. A. von Humboldt & A. J. Bonpland 1421 (Holotype, P; isotype, P!; photo- graph of P isotype, OS! TEX!). Melampodium ovatifolium Rchb. Icon. Bot. Exot. 30. t. 42. 1824. nom. superfl. Based on type of Dysodium divaricatum Rich. in Pers. Melampodium berterianum Spreng. Syst. Veg. 3:619. 1826. TYPE: “IND. occ.” [COLOMBIA]: near Sta Martha, C. G. Bertero s.n. (Holotype, P!; probable isotype, GH!; photo- graph of holotype, os! TEX [2]! US!; photograph of G-DC isotype, IDC 800. 928: I. 3!). Alcina minor Cass. Dict. Sc. Nat. 59:243. 1829. TYPE: MEXICO: cultivated in Bot. Gard. Paris (Holotype, P?). Dysodium radiatum Hort. ex Desf. Cat. Hort. Paris. ed. 3. 182. 1829. pro syn. Melampodium pumilum Benth. Pl. Hartw. 64. 1840. TYPE: MEXICO: exact locality unknown, 1830, G. J. Graham 5 (Holotype, K!; isotype, GH!; photograph of holotype, os! TEX! US!). Melampodium tenellum Hook. & Arn. var. flaccidum Benth. Bot. Voy. Sulphur 115. 1845. TYPE: MEXICO: Nay- arit, Tepic, 1844, A. Sinclair s.n. (Lectotype chosen, K!; isotype, K !; photograph of lectotype, US [3] !). Melampodium divaricatum (Rich. in Pers.) DC. var. macranthum Schlecht. Linnaea 24:198. 1851. TYPE: GUATE- MALA: locality and date unknown, seeds collected and grown in Bot. Gard. Turin, J. von Warscewicz s.n. (Holo- type, HAL; isotype, W !; photograph of w isotype, 0S! TEX!). Melampodium flaccidum (Benth.) Benth. in Oerst. Kjoeb. Vidensk. Meddel. 5-7:86. 1852. Melampodium copiosum Klatt, Engl. Bot. Jahrb. 8:41. 1887. TYPE: GUATEMALA: Alta Verapaz, near Cobán, 1300 1972] Melampodium — Stuessy 177 m, Feb [May?] 1882, F. C. Lehmann 1434 (Holotype, GH!; isotypes, F! G [2]! K! US!; photograph of K isotype, US!; photograph and fragment of K isotype, US!). Melampodium panamense Klatt, Engl. Bot. Jahrb. 8:42. 1887. TYPE: PANAMA: “ad margines silvarum," 20 Aug 1880, F. C. Lehmann 69 (Holotype, GH!; isotypes, G! K! US!; photograph of K isotype, US!). Spilanthes guatemalensis Vatk. ex J. D. Smith, Enum. PI. Guat. 1:23. 1889. nom. nud. Based on H. von Tuerckheim 124 (F! GH! NY! US [2]!). Melampodium rhombifolium Sessé & Moc. Fl. Mexic. ed. 2. 195 [first on page]. 1894. nom. illegit. non Sessé & Moc. 1890. nec Sessé & Moc. [second on page]. 1894. TYPE: MEXICO: Michoacan, Temascaltepec, Jul 1790-1791, M. Sessé et al. “3983” (Holotype, MA!; isotype, F!; photograph of holotype, OS! TEX!). Eleutheranthera divaricata (Rich. in Pers.) Millsp. Field Col. Mus. Pub. Bot. 1:53. 1895. Annual herbs, 15-100 cm tall. Stems erect, with lateral stems sometimes decumbent and rooting at the nodes, 1.3-6 mm diam, glabrous to pubescent with hairs up to 0.7 mm long. Peduncles 1.5-13.5 cm long. Leaves with petioles 2-20 mm long, ovate to rhombic (younger leaves approaching lanceolate), 1.5-15 cm long, 0.5-9.5 cm wide, at apex acumi- nate to obtuse, at base attenuate (rarely obtuse to cordate), with both surfaces infrequently strigillose with hairs 0.2 mm long; margin entire to coarsely dentate-crenate. Heads 5-10 mm tall, 10-21 mm diam. Outer involucre cupulate, 6-9 mm diam; bracts 5, connate 1/3-1/4 their length, imbricate, ovate-orbiculate, 3.5-6 mm long, 3-5 mm wide, at apex obtuse, with abaxial surface glabrous to tomentose near peduncle; margin herbaceous, near base tomentose. Fruits 2.8-4 mm long, with lateral surfaces with diagonal stria- tions and enlarged margins. Ray florets 8-13; ligules yel- low-orange, oblong-elliptic, 3.5-7 mm long, 1.6-3 mm wide. Disc florets 40-70; yellow-orange, 2 mm diam, with throat Fig. 24. Map of Mexico, Central America, northern South Amer- ica, and the Caribbean region showing distribution of Melampodium divaricatum. See Fig. 23 for the one Brazilian locality. Collections from Burma and the Virgin Islands not shown. 1 mm and tube 0.5 mm long. Paleae obovate, 2.5 mm long, 0.9 mm wide; apex yellow-orange, with margin dentate- erose; midrib weak, glabrous. Chromosome number, n = 12. Very widely spread in many subtropical habitats through- out Mexico, Central America, and northwestern Colombia, also in eastern Brazil, and introduced into Cuba, Burma, Puerto Rico and the Virgin Islands (Figs. 23 & 24), 15- 2990 m. Flowering dates, Jan-Dec. This is the most widespread and weedy species of the genus. Although field and garden studies have shown leaf shape to be extremely variable, which partially accounts for the many synonyms accorded this taxon, the morphology of the outer involucral bracts is very constant and allows the species to be recognized without difficulty. One plant from Nicaragua, Stwessy 615, has fruits with an anomalous hooded condition; other plants from the same population are hoodless. REPRESENTATIVE SPECIMENS. Brazil. BAHÍA: 1842, Glocker 19 (US); Marau, 22 Jan 1965, Pereira 9616 & Pabst 8505 [34972] (F). British 1972] Melampodium — Stuessy 179 Honduras. EL CAYO: El Cayo, near Camp 6, 11 Mar 1938, Gentle 2325 (ARIZ, F, GH, NY). Burma. MANDALA: Maymyo, Nov 1950, White 100 (US). Colombia. ANTIOQUÍA: near Dabeiba, 20 Dec 1947, Barkley & Gutiérrez 1777 (F). BOLIVAR: vicinity of Turbaco, 6-22 Nov 1926, Killip & Smith 14169 (F, Ny, US). CORDOBA: W of Monteria, near Río Sinu, 6 Jan 1949, Cervantes, Molina & Barkley 19Bo061 (vus). MAGDALENA: Sta Marta, Bonda, 9 Nov 1898-1899, Smith 514 in part (NY [3], Uc); Sta Marta, Jul 1898-1901, Smith 514 in part (F, RSA, SMU, TEX, UC, US). Costa Rica. ALAJUELA: between San Josecito & La Garita, 18 Aug 1963, Jiménez 1073 (F, US). GUANACASTE: Nicoya, Dec 1899, Tonduz 13615 (NY, US). PUNTARENAS: 21 km SE of Puntarenas, near bridge over Río Barranca, 20 Aug 1938, Stork & Morrison 8920 (GH, UC). Cuba. HAVANA: Vedado, shore of Arigu- anabo River, San Antonio de los Baños, 3 Sep 1939, Léon, Victorin & Alain 17344 (NY). El Salvador. AHUACHAPÁN: vicinity of Ahua- chapán, 9-27 Jan 1922, Standley 19722 (GH, US). LA LIBERTAD: Finca Paraíso, 4 mi S of Sta Tecla, 14 Jan 1946, Carlson 79 (F, UC, US). MORAZÁN: Río San Miguel opposite Barrios Mine, ca 5 km W of Montecristo, 19 Dec 1941, T'ucker 585 (F, NY, UC, US). SAN MIGUEL: 39 mi W of La Unión, 7 Aug 1966, Stwessy 623 (TEX). SAN SALVADOR: 18 mi E of San Salvador, 2 Aug 1966, Stuessy 611 (TEX). SANTA ANA: 4 mi SE of Sta Ana, 2 Aug 1966, Stuessy 609 (TEX). SAN VICENTE: vicinity of San Vicente, 2-11 Mar 1922, Standley 21648 (GH, NY, US). SONSONATE: vicinity of Sonsonate, 18-27 Mar 1922, Standley 21789 (GH, NY, US). Guatemala. ALTA VERAPAZ: ca 2 mi E of Coban, 12 Jul 1960, King 3322 (ps, NY, TEX, UC, US); 28 mi E of San Miguel Uspantán, 28 Jul 1966, Stuessy 588 (TEX); 1 mi NE cf San Pedro Carchá, 29 Jul 1966, Stuessy 592 (TEX). BAJA VERAPAZ: ca 1 mi S of Rabinal, 14 Jul 1960, King 3359 (DS, NY, TEX, UC, US). CHIMAL- TENANGO: San Martín J., 1928, Morales 1221 (F). CHIQUIMULA: Montana Nonojá, 3-5 mi E of Camotán, 11 Nov 1939. Steyermark 31670 (F). EL PROGRESO: between Finca Piamonte & Finca San Miguel, 9 Feb 1942, Steyermark 43745 (F, UC). EL QUICHE: 11 mi E of San Miguel Usnantaén, 28 Jul 1966, Stuessu 586 (TEX). ESCU- INTLA: 9 mi NE of Escuintla, 8 Aug 1966, Stuessy 624 (TEX). GUATEMALA: ca 4 mi E of Guatemala, 9 Jul 1960, King 3249 (ps, NY, TEX, UC, US). HUEHUETENANGO: Chacula, 29 Jul 1896, Volkem 3013 (GH, NY, US). IZABAL: Cristina, 23 May 1919, Blake 7632 (US). JALAPA: vicinity of Jalapa, 7-18 Nov 1940, Standley 77149 (F). JUTIAPA: 25 mi E of Cuilapa, 2 Aug 1966, Stuessy 605 (TEX); 8 mi NE of Jutiapa, 2 Aug 1966, Stuessy 608 (TEX). QUEZALTENANGO: Colomba, 31 Dec 1934, Skutch 2041 (F, GH). RETALHULEU: 1 mi W of jet rtes 2 & 9S, N of Retalhuleu, 10 Aug 1966, Stuessy 625 (TEX). SANTA ROSA: Volcán Jumaytepeque, Dec 1892, Heyde et Lux 4213 (F, GH, NY, US [2]). SUCHITEPEQUEZ: Mazatenango, 28 Feb 1905, 180 Rhodora [Vol. 74 Kellerman 4958 (SMU, US). ZACAPA: Gualán, 18 Jan 1905, Deam 247 (F, GH, NY, US). Honduras. EL PARAÍSO: Llano de Lizapa, 24 Aug 1948, Molina 1082 (F). FRANCISCO MORAZÁN: near San Francisco along Yeguare River, 16 Nov 1947, Williams & Molina 13398 (F, GH). VALLE: Amapala, 11 Sep 1945, Valerio 3341 (F [2]). Mexico. BAJA CALIFORNIA: San José del Cabo, 12 Sep 1890, Brandegee 301 (UC). CAMPECHE: 1 mi W of Champotón, 18 Jul 1966, Stuessy 532 (TEX) ; 14 mi N of Champotón, 18 Jul 1966, Stuessy 534 (TEX); 14 mi E of Campeche, 18 Jul 1966, Stuessy 535 (TEX); 12 mi S of Campeche, 22 Jul 1966, Stuessy 545 (TEX). CHIAPAS: ca 15 mi S of Suchiate, 20 Jul 1960, King 3434 (DS, NY, RSA, TEX, UC); 19 mi S of Pichucalco, 23 Jul 1966, Stuessy 550 (TEX). COLIMA: 4 mi E of Colima, 25 Aug 1966, Stuessy 723 (TEX); 7 mi NW of rte 110 on rd to Alzada, 25 Aug 1966, Stuessy 725 (TEX); Alzada, 25 Aug 1966, Stuessy 728 (TEX). DISTRITO FEDERAL: Sta Rosa, 4 Oct 1952, Matuda 26621 (US). GUANA- JUATO: valley near Irapuato, 20 Sep 1889, Pringle 2819 (F, GH). GUERRERO: vicinity of Acapulco, Oct 1894-Mar 1895, Palmer 186 (F, GH, NY, US) ; 4 mi S of Taxco, 17 Aug 1961, Powell & Edmondson 756 (F, TEX); S limits of Petaquillas, 27 Aug 1965, Stuessy 365 (TEX). HIDALGO: Jacala, 25 Jun 1939, Chase 7122 (F). JALISCO: Río Cuvianes, 13 Jun 1892, Jones 273 (PoM [2], US). MEXICO: Tema- scaltepec, 2 Sep 1932, Hinton 1520 (NY). MICHOACÁN: ca 45 mi W of Morelia, 5 Aug 1960, King 3634 (DS, NY, RSA, TEX, UC); 16 mi NW of Zitacuaro, 19 Aug 1966, Stuessy 677 (TEX); S city limits of Uruapán, 21 Aug 1966, Stuessy 702 (TEX). MORELOS: ca 11 mi S of Cuernavaca, 14 Mar 1961, King 4164 (F, NY, TEX, UC, US) ; 35.5 mi NW of Izácar de Matamoros, 25 Aug 1965, Stuessy 348 (TEX) ; 6 mi NW of Cuautla, 25 Aug 1965, Stuessy 351 (TEX); 10 mi S of Cuernavaca, 26 Aug 1965, Stuessy 359 (TEX). NAYARIT: ca 16 mi NW of Ixtlán del Río, 10 Aug 1960, King 3678 (Ds, NY, TEX, UC, US) ; 1 mi N of Tepie, 27 Aug 1966, Stuessy 743 (TEX); 10 mi NW of jet rte 16 & rd to Tuxpán, 27 Aug 1966, Stuessy 746 (TEX). NUEVO LEÓN: Villa de Santiago, Las Adjuntas, 21 Jun 1940, Leavenworth 157a (F). OAXACA: ca 1 mi W of the Oaxaca-Chiapas border on rte 190, 9 Jun 1960, King 2743 (DS, NY, TEX, UC, US); 0.7 mi N of Huajuapán de León, 24 Aug 1965, Stuessy 342 (TEX); 40 mi S of Zimatlán, 13 Avg 1966, Stuessy 645 (TEX). PUEBLA: just SE of the Morelos-Puebla border on rte 115, 19 Jun 1960, King 2917 (DS, NY, TEX, UC, US); Asunción de Chila, 25 Aug 1965, Stuessy 345 (TEX). QUERÉTARO: 2 mi W of Landa, 13 Dec 1960, Crutchfield & Johnston 6120b (TEX). QUINTANA ROO: Coba, Jun-Jul 1938, C. & A. Lundell 7879 (US). SAN LUIS POTOSÍ: ca 10 mi S of Tamazunchale, 23 Mar 1961, King 4239 (F, NY, TEX, UC, US); Cardenas, 3 Nov 1891, Pringle 3923 (DS, F, GH, NY, vc [2], Us); 2 mi off rte 80 on rd to Quadal- cazar, 10 Aug 1965, Stuessy 282 (TEX). SINALOA: ca 15 mi N of 1972] Melampodium — Stuessy 181 Rosario, 13 Aug 1960, King 3711 (DS, NY, RSA, TEX, uc); 4.5 mi N of Escuinapa, 4 Sep 1965, Stuessy 403 (TEX). TABASCO: 57 mi SE of Coatzacoalcos, 17 Jul 1966, Stuessy 527 (TEX); 84 mi E of Coat- zacoalcos, 17 Jul 1966, Stuessy 528 (TEX); 5 mi W of Villa Hermosa, 23 Jul 1966, Stuessy 547 (0S, TEX); 37 mi S of Villa Hermosa, 23 Jul 1966, Stuessy 548 (TEX). TAMAULIPAS: ca 4-5 mi W of Ciudad Mante, 18 Feb 1961, King 3839 (F, NY, TEX, US); 4 mi S of Juamave, 6 Jul 1949, Stanford, Taylor & Lauber 2318 (DS, GH, NY, RSA, SMU, TEX, UC, US [2]) ; 1 mi E of Gómez Farias, 8 Jul 1967, Stuessy 829 (TEX). VERACRUZ: ca 25 mi SE of Poza Rica, 10 Mar 1961, King 4138 (F, NY, TEX, US); 23.2 mi SE of Alvarado, 21 Aug 1965, Stuessy 318 (TEX); 12 mi S of Tantoyuca, 6 Jul 1966, Stuessy 473 (TEX); 49 mi SE of Catemaco, 16 Jul 1966, Stuessy 526 (TEX). YUCATÁN: Izamal, 1895, Gawmer 563 (F, GH [2], NY [2], us); Progreso, 20 Jul 1966, Stuessy 543 (TEX). Nicaragua, CHINANDEGA: vicinity of Chichigalpa, 12-18 Jul 1947, Standley 11226 (F). GRANADA: Volcan Mombacho, 20 Feb 1903, Baker 2482 (GH, POM, UC, us); NE limits of Granada, 6 Aug 1966, Stuessy 620 (TEX). MANAGUA: 1 mi SE of Managua, 5 Aug 1966, Stwessy 616 (TEX); 5 mi E of Managua, 6 Aug 1966, Stuessy 621 (TEX). MASAYA: near Masaya, SW slopes of Santiago Voleano, 5 Jul 1923, Maxon 7694 (US). MATAGALPA: Sebaco, 5 Aug 1966, Stuessy 614 & 615 (TEX). PUNTARENAS: 21 km SE cf Puntarenas, 20 Aug 1938, Stork & Morrison 8920 (GH, UC). Panama. CANAL ZONE: vicinity of Frijoles, 9 Jul 1962, King 5226 (UC, US). DARIÉN: trail between Pinogana & Yavisa, 17 Mar 1937, Allen 241 (cH [3], NY, US). PANAMA: near Arraiján, 21 Jul 1938, Woodson, Allen & Seibert 1870 (GH, NY, US). PEARL ARCHIPELAGO: San José Is, Naval Station, 22 Jul 1945, Erlanson 500 (DS, GH, TEX, US). Puerto Rico: Mayaguez, 14-15 Feb 1914, Britton & Cowell 1561 (NY). Virgin Islands. ST. CROIX: near Mt Pleasant, 19 Aug 1896, Ricksecker "S" (F, NY). 27. Melampodium costaricense Stuessy, Brittonia 22:118. f.7. 1970. TYPE: COSTA RICA: Alajuela, Naranjo, Cerro del Espíritu Santo, 1200 m, 11 Jul 1941, A. Smith 2922 (Holo- type, F!). Annual herbs, 15-35 em tall. Stems erect, with lateral stems often decumbent, 0.8-4 mm diam, glabrous to tomen- tose with hairs 0.8 mm long. Peduncles 0.5-7 cm long. Leaves with petioles 4-25 mm long, ovate-rhombic (younger leaves approaching lanceolate), 2.5-7 cm long, 1.5-5.5 cm wide, at apex acuminate to obtuse, at base obtuse-attenuate, with upper surface infrequently strigose with hairs 0.6 mm 182 Rhodora [Vol. 74 long, with lower surface glabrous; margin obscurely to coarsely serrate. Heads 4-5 mm tall, 6-7 (10) mm diam. Outer involucre cupulate, 5-10 mm diam; bracts 5, connate 1/3-1/4 their length, imbricate, ovate-orbiculate, 2.2-5 mm long, 2-4 mm wide, at apex obtuse, with abaxial surface glabrous; margin herbaceous, ciliate with hairs 0.3 mm long. Fruits 2.9-3.1 mm long, with lateral surfaces with diagonal ridges and enlarged margins. Ray florets 5-8; ligules yellow-orange, elliptic, 1-1.5 mm long, 0.5-1 mm wide. Disc florets 15-25; corollas yellow, 1.1 mm diam, with throat 0.8 mm and tube 0.5 mm long. Paleae oblong-elliptic, 2 mm long, 0.7 mm wide; apex yellow, with margin erose; midrib weak, glabrous. Chromosome number, n = 25 + 1. Tropical habitats in British Honduras, Nicaragua, Costa Rica and Panama of Central America, and Colombia, South America (Fig. 25), 0-1980 m. Flowering dates, Jan-Dec. REPRESENTATIVE SPECIMENS, British Honduras. EL CAYO: El Cayo, Jun-Aug 1936, Lundell 6107 (F, GH, NY, US). Colombia. CUNDINA- MARCA: Caqueza to Rio Sananie, 24 Aug 1917, Pennell 1335 (Ny). Costa Rica. ALAJUELA: banks of the Río Rosales y Carret, 8 Dec 1933, Brenes 17471 (NY); San Pedro de San Ramón, 29 Jul 1935, Brenes 20572 (F). CARTAGO: above Tres Ríos, 19 Mar 1965, Godfrey 67156 (FsU); Turrialba, Instituto Interamericano de Agrícolas, 20 Aug 1962, King 5348 (TEX, UC, US); Finca Las Concavas, 7-8 Dec 1925, Standley 41531 (US); rr below Turrialba, 9 Jun 1928, Stork 2486 (F, US). GUANACASTE: Hatillo, 4 Jan 1935, Solís 56 (F). LÍMON: marshes of Suerre & Dos Bocas, 3 Oct 1951, Shank & Molina 4236 (F, GH); Hamburg Finca, on the Río Reventazon below Cairo, 19 Feb 1926, Standley & Valerio 48814 (US). SAN JosÉ: Alajuelita, 8 Sep 1946, Echeverría 628 (F); San Isidro, 26 Feb 1965, Godfrey 6664 (FSU); San Pedro de Montes de Oca, 2 Aug 1938, Orozco 294 (F); El General [San Isidro del], Jan 1939, Skutch 3968 (GH, K, NY, US); between San Pedro de Montes de Oca & Curridabat, 2 Feb 1924, Standley 32757 (us); San José, Feb 1924, Standley 39021 (US). Nicaragua. GRANADA: near Granada, 1845-48, Oersted 9010 (photograph of K specimen, US). Panama, BOCAS DEL TORO: Changuinola Valley, 15 Mar 1924, Dunlap 557 (F, US). CHIRIQUT: 25 mi N of Concepción, 29 Jul 1962, King 5290 (TEX, UC, US). 28. Melampodium dicoelocarpum Robins. Proc. Amer. Acad. Arts & Sci. 44:619. 1909. TYPE: MEXICO: Guerrero, El Cala- 1972] Melampodium — Stuessy 183 Fig. 25. Map of Central America and neighboring countries show- ing distribution of Melampodium costaricense. bazal, 300 m, 20 Oct 1898, E. Langlassé 482 (Holotype, GH!; isotypes, G! K! P! US!; photograph of K isotype, US!; photo- graph of US isotype, TEX!). Annual herbs, 10-50 cm tall. Stems erect, 0.6-1.6 mm diam, glabrous to moderately pilose with hairs 0.3-1.3 mm long. Peduncles deflected, 0.4-7.5 cm long. Leaves with petioles 2-10 mm long, ovate to somewhat deltoid (younger leaves often ovate-lanceolate), 2-6 cm long, 0.7-3 cm wide, at apex acuminate-attenuate, at base obtuse-attenuate, with both surfaces moderately pilose with hairs 1 mm long; margin obscurely to coarsely serrate. Heads 2-4 mm tall, 2.5-8 mm diam. Outer involucre spreading, 2-7 mm diam; bracts 5 (rarely 3 or 4), slightly connate at base, separate, ovate-lanceolate, 1.5-4.5 mm long, 1-2.5 mm wide, at apex acuminate, with abaxial surface moderately strigose with 184 Rhodora [Vol. 74 hairs 0.6-1 mm long; margin herbaceous. Fruits 1.7-3 mm long, with lateral surfaces with 2 deep oval depressions. Ray florets 3-5; ligules yellow-orange, oblong-elliptic, 2.2- 2.5 mm long, 1.2-1.8 mm wide. Disc florets 25-35; corollas yellow-orange, 0.8 mm diam, with throat 0.7 mm and tube 0.5 mm long. Paleae oblong-elliptic, 2 mm long, 0.7 mm wide; apex yellow, with margin erose to laciniate; midrib weak, glabrous. Chromosome numbers, n — 12 and 23. Tropical deciduous forests in the Mexican states of Jalis- co, Michoacán, Guerrero and México (Fig. 13), 200-1890 m. Flowering dates, Jun-Oct. The cytological vouchers for the two chromosome levels are morphologically distinguishable, the aneuploid at the tetraploid level being larger in most characters. However, without intensive field and garden studies, it seems prema- ture to propose formal recognition of these forms. Among all the taxa of Melampodium, M. dicoelocarpum can be distinguished easily by the two deep oval cavities on the lateral sides of the fruit. REPRESENTATIVE SPECIMENS. Mexico, GUERRERO: Mina, Zihuagio, 17 Oct 1936, Hinton 9714 (G, GH, MICH, NY, P, UC, US); Galeana, Atoyac-San Juan, 12 Oct 1939, Hinton et al. 14614 (ARIZ, F, GH, NY, US). JALISCO: Tecalitlán, Barranca de San Juan de Dios, ca 15 km E of Pihuamo, 24 Oct 1963, Feddema 2204 (MICH). MÉXICO: Temas- caltepee, Anonas, 24 Jun 1934, Hinton et al. 6334 (GH, MICH, NY). MICHOACÁN: ca 5 mi N of Cotija, 5-9 Oct 1961, King & Soderstrom 4646 (MICH, NY, SMU, TEX, UC, US); 25 mi S of Ario de Rosales, 20 Aug 1966, Stuessy 693 (TEX); 15 mi S of jet rte 15 on rd to Cotija, Stwessy 715 (TEX). 29. Melampodium tepicense Robins. Proc. Amer. Acad. Arts & Sci. 44:620. 1909. TYPE: MEXICO: Nayarit, Tepic, " 5 Jan-6 Feb 1892, E. Palmer 1814 (Holotype, GH!; iso- types, F! NY! Us!). Melampodium cornutum S. F. Blake, Jour. Wash. Acad. Sei. 16:421. 1926. TYPE: MEXICO: Colima, Alzada,4 Nov 1910, C. R. Orcutt 6601 (Holotype, Us!; photograph of holo- type, TEX!). 1972] Melampodium — Stuessy 185 Annual herbs, 7-35 cm tall. Stems decumbent to erect, 0.5-1.3 mm diam, mostly glabrous to puberulent with hairs 0.1 mm long. Peduncles 0.8-8 mm long. Leaves with nar- rowly marginate petioles 1-18 mm long, ovate to rhombic, 0.8-5.7 cm long, 0.7-3.5 cm wide, at apex acute, at base at- tenuate to obtuse, with both surfaces sparsely strigose with hairs 0.3-1 mm long; margin crenate to serrate, near base entire. Heads 2-3 mm tall, 3-5 mm diam. Outer involucre cupulate, 3.5-7 mm diam; bracts 5, slightly connate at base, separate, elliptic, 2-4 mm long, 1.2-2.2 mm wide, at apex obtuse, with abaxial surface glabrous; margin herbaceous. Fruits 1.6-2 mm long, with lateral surfaces tuberculate to aculeate-reticulate; apex with a flattened abaxial append- age’, with tapering cirrhous awn up to 4 mm long. Ray florets 3-7; ligules pale-yellow, ovate-oblong, 0.8-1.5 mm long, 0.4-0.8 mm wide. Disc florets 3-7(12) ; corollas yel- low-green, 0.8 mm diam, with throat 0.4 mm and tube 0.7 mm long. Paleae oblong-oblanceolate, 1 mm long, 0.5 mm wide; apex pale-yellow, with margin irregularly dentate; midrib weak, glabrous. Chromosome number unknown. Pine-oak and tropical deciduous forests in Nayarit, Jal- isco, Colima, and Michoacan, Mexico (Fig. 18), 100-1740 m. Flowering dates, Aug-Feb. Recent collections frem Nayarit, Mexico, show much var- iation and intergradation in the leaf and inner involucral bract characters used by Blake (1926) to separate M. tepi- cense from M. cornutum. In my opinion they are conspe- cific. REPRESENTATIVE SPECIMENS. Mexico. COLIMA: SW foothills of Nevado de Colima, 1-1.5 mi S of Hacienda San Antonio, 11 Aug 1957, McVaugh 16112 (MiCH). JALISCO: ea 10 km NE of Puerto Vallarta near Milagro on rd to Mascota, 15 Nov 1963, Feddema 2577 (MICH); 10 mi S of Autlán, 19 Aug 1949, R. & C. Wilbur 2416 (MICH). MICHOACÁN: Coalcoman, Parotas, 28 Sep 1938, Hinton et al. 12276 (MICH, NY, UC, US). NAYARIT: 10 mi SE of Ahuacatlán on rd to Bar- "Similar to the appendage in M. longipilum, except here in M. tepicense it is on the abaxial rather than the adaxial side of the achene apex. 186 Rhodora [Vol. 74 ranca del Oro, 11-12 Aug 1959, Feddema 279 (MICH); 1 km N of El Cuatante, ca 40 km NNE of Puerto Vallarta, 18 Nov 1963, Feddema 2651 (MICH); 8 mi W of Tepic, 10 Sep 1960, McVaugh 18906 (MICH) ; 9 mi N of Compostela, 12 Nov 1959, McVaugh & Koelz 542 (MICH). 30. Melampodium sinaloense Stuessy, Brittonia 22:122. f. 10. 1970. TYPE: MEXICO: Sinaloa, Quebrado de Mansana, Sierra Surotato, 4000-4500 ft, 10-14 Sep 1941, H. S. Gentry 6507-A (Holotype, GH ! ; isotypes, ARIZ! DS! MICH! MO! NY!). Annual herbs, 25-35 em tall. Stems decumbent, 1-2 mm diam, sparsely tomentose to moderately hispidulous with hairs 0.1-0.3 mm long. Peduncles 1-2.9 cm long. Leaves with petioles 10-15 mm long, ovate, 3.5-5 cm long, 2-2.5 em wide, at apex acute, at base attenuate, with both surfaces sparingly strigose with hairs 0.3-1.2 mm long; margin ser- rate. Heads 3-4 mm tall, 3-4 mm diam. Outer involucre cupulate, 5-10 mm diam; bracts 5, slightly connate at base, separate, narrowly elliptic to elliptic, 2.5-6 mm long, 1.2- 1.5 mm wide, at apex obtuse, with abaxial surface with 2-5 hairs 0.3 mm long on midrib and major lateral veins; margin herbaceous. Fruits 1.8-2 mm long, with lateral surfaces ridged. Ray florets 6; ligules pale-yellow, elliptic, 1.5-Z mm long, 1-1.2 mm wide. Disc florets ca 14; corollas pale-yellow, 0.8 mm diam, with throat 0.7 mm and tube 0.6 mm long. Paleae elliptic, 1.3 mm long, 0.6 mm wide; apex pale-yellow, with margin irregularly and weakly den- tate; midrib weak, glabrous. Chromosome number un- known. Known only from the type collection from oak forests in Sinaloa, Mexico (Fig. 18), 1220-1370 m. Flowering date, Sep. IV. Melampodium section Bibractiaria Stuessy, sect. nov. Herbae annuae radicibus palaribus; folia ovata vel ob- ovata, sessilia vel subsessilia, marginibus integris; invol- ucrum extimum cupulatum, bracteis 2, marginibus herba- ceis vel leniter scariosis; ovaria flosculi disci ovoidea, minus quam 0.7 mm longa, rudimentales (Figs. 3 & 4) ; fructus apice fere laeves vel moderate sculpti et sine cucullis ; 1972] Melampodium — Stuessy 187 chromosomatum numerus basicus ignotus. Species 31 & 32. Typus: Melampodium bibracteatum S. Wats. 31. Melampodium bibracteatum S. Wats. Proc. Amer. Acad. Arts & Sci. 26:140. 1891. TYPE: MEXICO: México, Del Rio [ca 13 mi N of Toluca], 30 Aug 1890, C. G. Pringle 3230 (Holotype, US!; isotypes, DS! F! G[2]! GH! MA! MO! MSC! NY [2]! P! uc! US! W!; photograph of holotype, TEX!; photograph of K isotype, US!). Annual herbs, 2-32 cm tall. Stems ascending to decum- bent, 1.8-3.5 mm diam, glabrous below to pubescent above with hairs 0.2 mm long. Peduncles 0-3.5 mm long (heads often sessile). Leaves sessile, with blades oblong to ob- ovate or oblanceolate, 0.9-4 cm long, 0.3-1.8 cm wide, at apex obtuse-acute, at base obtuse-subauriculate to some- what connate, with both surfaces with a conspicuous midrib (very large basally) extending 1/3-1/2 of blade, with upper surface glabrous, with lower surface subglabrous; margin entire to 6-lobed, moderately ciliate with hairs 0.1 mm long. Heads 3.5-4.5 mm tall, 3.5-4.8 mm diam. Outer involucre cupulate, 4-7 mm diam; bracts 2, separate, ovate, 3-4.2 mm long, 1.5-2.5 mm wide, at apex acute, with abaxial surface glabrous; margin herbaceous to weakly scarious, rarely with several hairs 0.2-0.5 mm long. Fruits 2-2.7 mm long, with lateral surfaces smooth with several conspicuous nerves. Ray florets (3-) 5-6; ligules yellow, elliptic-ovate, 1 mm long, 0.4 mm wide. Disc florets 4; corollas yellow, 0.8 mm diam, with throat 0.7 mm and tube 0.5 mm long. Paleae oblanceolate, 1.5 mm long, 0.6 mm wide; apex color- less, with margin irregularly laciniate; midrib very weak, glabrous. Chromosome number unknown. Mainly in low, moist areas in pine-oak forests of central Mexico (Hidalgo, México and Puebla) but populations oc- eur also in southwestern Durango and in Guatemala (Fig. 20), 2140-3360 m. Flowering dates, Aug-Oct. REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Sierra de los Cuchumatanes, 1.5 mi W of Llano de San Miguel on trail to Todos Santos, 2 Aug 1960, Beaman 3977 (GH, TEX, UC). Mexico. 188 Rhodora [Vol. 74 DURANGO: 10 mi W of El Salto, 2 Oct 1962, Cronquist 9585 (NY); 3 mi SW of El Salto, 12 Aug 1961, Waterfall 16198 (SMU); 57 mi SW of Durango, 13 Aug 1957, Waterfall & Wallis 13702 (smu, Us). HIDALGO: near Buena Vista Station, 10 Aug 1904, Pringle 13068 (ARIZ, F, GH, SMU, US). MEXICO: Tultenango Station, 14 Sep 1901, Pringle 9301 (F, GH, K, NY, US); !? mi W of Station Del Rio, 18 Aug 1966, Stuessy 675 (TEX); 7 mi W of Toluca, 19 Aug 1966, Stuessy 676 (TEX). PUEBLA: near Cerro El Pinar, 28 Aug 1945, Alexander & Hernández 2229 [XA 218] (Nv); Laguna de San Baltasar, vicinity of Puebla, 20 Sep 1966, Arsène 294 (US). 32. Melampodium repens Sessé & Moc. Fl. Mex. ed. 2. 195. 1894. TYPE: MEXICO: Veracruz, *Habitat in medietate Vul- cani altissimi de Orizava," Jun 1795-1804, M. Sessé et al. “3981” (Holotype, MA!; photograph of holotype, 0S! TEX!). Melampodium arvense Robins. Proc. Amer. Acad. Arts & Sci. 36:464. 1901. TYPE: MEXICO: Distrito Federal, Val- ley of Mexico, 7500 ft, 19 Oct 1896, C. G. Pringle 7327 (Holotype, GH!; isotypes, F! POM! US!; photograph of US isotype, TEX!). Annual herbs, 2-30 cm tall. Stems prostrate (often form- ing a mat), 0.8-1.8 mm diam, puberulent with hairs 0.2 mm long. Peduncles 0-3 mm long (heads often sessile). Leaves sessile or with petioles 2 mm long, obovate, 0.5-2.5 cm long, 0.3-1.6 cm wide, at apex acute-cbtuse, at base at- tenuate-obtuse to somewhat connate, with both surfaces with a conspicuous midrib (very large basally) extending less than 1/3 of blade, with upper surface glabrous, with lower surface subglabrous; margin obscurely crenate, sometimes moderately ciliate with hairs 0.1 mm long. Heads 2-3 mm tall, 3.5-4 mm diam. Outer involucre cupulate, 4-5 mm diam; bracts 2, separate, cvate, 3.5 mm long, 1.8-2.2 mm wide, at apex acute, abaxial surface glabrous; margin herbaceous. Fruits 2-2.5 mm long, with lateral surfaces smooth with several conspicuous nerves. Ray florets 2-3; ligules yellow, elliptic-oblong, 0.8-1.2 mm long, 0.3 mm wide. Disc florets 6; corollas yellow, 0.5 mm diam, with throat 0.4 mm and tube 1.0 mm long. Paleae oblanceolate, 1.2 mm long, 0.4 wide; apex colorless, with margin laciniate; mid- rib very weak, glabrous. Chromosome number unknown. 1972] Melampodium — Stuessy 189 Pine-oak forests in the Mexican states of Distrito Fed- eral, México, Michoacan, and Veracruz (Fig. 20), 1380- 3150 m. Flowering dates, Aug-Sep. The two species of section Bibractiaria are perhaps the most unusual taxa of the whole genus, particularly in their over-all diminutive appearance. Chromosome counts have not been obtained yet from these two species, although collections have been made and counts attempted. In view of the wide diversity of chromosome numbers already found in Melampodium, it will be interesting to learn if new num- bers may be obtained from these two taxa. REPRESENTATIVE SPECIMENS. Mexico, DISTRITO FEDERAL: Cima, 24 Aug 1910, Orcutt 3780 (F); Eslava, Valley of Mexico, 7 Sep 1901, Pringle 8610 (ARIZ, F, G, GH, K, MSC, NY, POM, SMU, TEX, UC, US). DURANGO [?]: “in mountains near Santa Angela,” Sep 1855, Schaff- ner s.n. (GH). MÉXICO: Parque Nacional de Laguna Zempoala, 26 Jul 1947, Barkley, Webster & Rowell 7438 (TEX); Temascaltepec, Cucha, 4 Sep 1933, Hinton 4628 (GH, NY, us); Temascaltepec, Teju- pilco, 27 Sep 1933, Hinton 4826 (GH, K, NY, US); Toluca, 26 Sep 1892, Pringle 5257 (GH); Salto de Agua, Oct 1905, Purpus 1809 (F, UC); Popocatépetl, 22 Aug 1901, Rose & Hay 6244 (NY, US); Llano Grande, near Rio Frio, 28 Jul 1944, Sharp 44116 (NY); 2 mi E cf Cuajimalpa, 18 Aug 1966, Stuessy 670 (TEX). MICHOACAN: SW side of Cerro San Andres, ca 12 km N of Ciudad Hidalgo, 6 Sep 1960, Beaman 4352 (GH). VERACRUZ: Loma Grande, 27 Aug 1938, Balls & Gourlay B5372 (x). V. Melampodium section Rhizomaria Stuessy, sect. nov. Plantae perennae rhizomatosae; folia anguste ovata vel oblonga, sessilia vel interdum sessilia, marginibus integris vel subintegris; involucrum extimum patens vel aliquan- tum cupulatum, bracteis 5, marginibus scariosis; ovaria flosculi disci ovoidea, minus quam 0.7 mm longa, rudimen- tales (Figs. 3 & 4); fructus apice fere laeves vel moderate sculpti et sine cucullis; chromosomatum numerus basicus, x = 11. Species 33 & 34. Typus: Melampodium montanum Benth. 33. Melampodium montanum Benth. Pl. Hartw. 64. 1840. Suffrutescent perennials. Stems erect to ascending. Leaves usually sessile, ovate to elliptic, at apex obtuse 190 Rhodora [Vol. 74 (rarely acute), at base obtuse to subauriculate. Outer in- volucre spreading or somewhat cupulate; bracts 5, connate 1/5 their length, imbricate, ovate, 4-5 mm long, 4-5 mm wide, at apex acute, with margin scarious. Fruits 1.6-1.8 mm long, with lateral surfaces smooth to nervose or mar- ginally tuberculate. Ray florets 9-13; ligules yellow, nar- rowly elliptic-oblong. Disc florets 80-110; corollas yellow- green, with tube 0.8 mm long. Paleae oblanceolate, 2 mm long, 0.6 mm wide; apex yellow, with margin entire; mid- rib prominent, glabrous. 33a. Melampodium montanum Benth. var. montanum Melampodium montanum Benth. Pl. Hartw. 64. 1840. TYPE: MEXICO: Oaxaca, “In summo jugo (Cumbre) inter Oaxaca et la Sierra,” 1839, C. T. Hartweg 475 (Holotype, K!; photograph of holotype, os! TEX! US!). Melampodium liebmanii Sch. Bip. ex Klatt, Leopoldina 23:89. 1887. TYPE: MEXICO: Oaxaca, “Cumbre de Estepa et Yavesía," Jun 1842, F. M. Liebmann 232 (Lectotype, C; isotype, P!; photograph of lectotype, US!; photograph of C isotype, F! NY! us [2] !; photograph of P isotype, 0S! TEX!; drawing and presumed fragment of lectotype, GH !). Plants 6-28 em tall. Stems 0.8-1.2 mm diam, copiously to sparsely villous with hairs 0.8-1.5 mm long. Peduncles 3- 7.5 cm long. Leaves sessile or with petioles 1 mm long, 1.6-4 cm long, 0.8-1.7 em wide, with both surfaces strigose with hairs 0.8 mm long; margin entire to obscurely denticulate. Heads 5-6 mm tall, 11-21 mm wide. Outer involucre 8-13 mm diam; bracts at apex purple-tinged, with abaxial sur- face villous with hairs 1 mm long. Ligules at apex and on veins on undersurface dark purple, 4-7.5 mm long, 1.5-2.5 mm wide. Disc florets 1 mm diam, with throat 1.4 mm long. Chromosome number, n = ca 11. Pine-oak forests in north-central Oaxaca, Mexico (Fig. 26), 1740-3360 m. Flowering dates, Jul-Aug. 1972] Melampodium — Stuessy 191 Fig. 26. Map of Mexico and adjoining Guatemala showing dis- tribution of Melampodium aureum (squares), M. montanum var. montanum (dots), and M. montanum var. viridulum (triangles). REPRESENTATIVE SPECIMENS. Mexico, OAXACA: Cerro San Felipe, 18 Aug 1921, Conzatti 4169 (US) ; Jul-Aug 1900, Conzatti & González 998 (GH); vicinity of Cerro Zempoaltepet| [Zempoaltepec], 10 Aug 1950, Hallberg 900 (Us); ca 10 mi N of jet rtes 190 & 175, 26 Jul 1960, King 3492 (ps, NY, TEX, UC, US); NW side of summit of Mt Zempoaltepec, 9 Jul 1894, Nelson 655 (US); Sierra de San Felipe, 28 May 1894, Pringle 4666 (GH, Us). 33b. Melampodium montanum Benth. var. viridulum Stues- Sy, var. nov. Plantae 10-35 cm altae. Caules 0.8-1.4 mm diametro, vil- losi pilis 1-1.5 mm longis (raro glabri). Pedunculi 1-6.5 cm longi. Folia subsessilia (vel petiolis 2-3 mm longis), laminae 1.6-3.3 longae, 0.7-1.4 em latae, utrinque strigosae vel moderate vilosae pilis 1-1.5 mm longis (subtus raro glabrae); margines integri. Capitula 4.5-6 mm alta, 12-17 mm diametro. Involucrum extimum 8-14 mm diametro; bracteae apice interdum purpurascentes, extus villosae pilis 192 Rhodora [Vol. 74 1 mm longis (interdum subglabrae). Ligulae subtus apice venaque viridulae, 4-5 mm longae, 1.2-1.9 mm latae. Flos- culi disci 0.8 mm diametro, faucibus 1.2 mm longis. Chro- mosomatum numerus, n = 11. TYPUS: MEXICO: Chiapas, ca 5 mi E of San Cristóbal de Las Casas on rte 190, 11 Jun 1960, R. M. King 2801 (Holotype, US!; isotypes, Ds! NY! TEX! UC!). Scattered in pine-oak forests in several Mexican states in the Sierra Madres Oriental, Occidental and Sur, and in- cluding Chiapas and northwestern Guatemala (Fig. 26), 610-2740 m. Flowering dates, Jun-Oct. REPRESENTATIVE SPECIMENS. Guatemala. HUEHUETENANGO: Sierra de los Cuchumatanes, between kms 324 & 325 on rte 9N (between Chemal & San Juan Ixcoy), 4 Aug 1959, Beaman 3043 (GH, NY, UC); San Pedro Coloma, 20 Aug 1934, Skutch 1036 (F, GH, US); SW slopes of Sierra de los Cuchumatanes, between Chiantla & Patio de Bolas, 6 Jul 1942, Steyermark 48234 (F); along Río Selegua, oppo- site San Sebastián H., !3 Aug 1942, Steyermark 50490 (F, GH). Mexico. CHIAPAS: Jitotol, 3 mi S of Jitotol, 12 Feb 1965, Breedlove 8906 (ns); Tenejapa, Tuk, Matsab, 30 Sep 1965, Breedlove 12522 (ps); Comitán, 30 Apr 1904, Goldman 914 (US) ; ca 17 mi W of San Cristóbal de Las Casas, 9 Jun 1960, King 2796 (DS, NY, TEX, UC, US) ; ca 1 mi E of Teopisca, 12 Jun 1960, King 2843 (TEX); San Cristóbal de Las Casas, Cerro San Cristóbal, 11 Jun 1966, Laughlin 1050 (DS) ; Amatenango del Valle, 12 Jun 1945, Matuda 6002 (F); 34 mi S of Ishuatán on gravel rd to Tuxtla Gutiérrez, 24 Jul 1966, Stuessy 559 (TEX); 20 mi S of San Cristóbal, 25 Jul 1966, Stuessy 566 (TEX); 8 mi S of San Cristóbal, 25 Jul 1966, Stuessy 567 (TEX); 2 mi SE of Teopisca, 25 Jul 1966, Stuessy 568 (TEX); 9 mi SE of Tecpisca, 25 Jul 1966, Stuessy 570 (TEX). GUERRERO: Taxco, 23 Jul 1936, Abbott 162 (GH). JALISCO: Sierra del Tigre, 3 mi S of Mazamitla, 18 Sep 1952, MeVaugh 13033 (SMU, US). MÉXICO: Temascaltepec, Bejucos, 7 Oct 1932, Hinton 2017 (ARIZ, F, MA, NY [2], vs). MICHOACÁN: Morelia, Cerro Azul, 1910, Arséne 6575 (US). OAXACA: 15 km S of Sola de Vega, on rd to Puerto Escondido, 30 Sep 1965, Rzedowski 21322 (TEX). PUEBLA: vicinity of Puebla, woody hill farther than Cerro Tepoxuchitl, 16 Sep 1907, Arséne 1896 (GH, NY, us); hills near Amozoc, 10 Sep 1901, Pringle 9361 (F, GH, US). QUERÉTARO: 8 km W of El Lobo, municipio de Landa, 31 Aug 1957, Rzedowsk: 9320 (ENCB). SAN LUIS POTOSÍ: “barranca of Las Canoas,” 18 Aug 1891, Pringle 3818 (ps, F, GH [2], Ny [2], vc [2], US). TAMAULIPAS: Gómez Farías Region, between Rancho del Cielo & Charco de los Perros, spring 1965, Webster 140 (TEX). 1972] Melampodium — Stuessy 193 34. Melampodium aureum Brandg. Univ. Calif. Pub. Bot. 4:94. 1910. TYPE: MEXICO: Puebla, Cerro de Gavilan, 8000- 9000 ft, Aug 1909, C. A. Purpus 3822 (Holotype, UC!; iso- types, F! G! GH! MO! NY! US!; photograph of US isotype, TEX!). Suffrutescent perennials, 10-35 cm tall. Stems erect to ascending (occasionally decumbent), 0.8-2 mm diam, mod- erately villous with hairs 0.5-1 mm long. Peduncles 4.5-11.5 cm long. Leaves usually sessile or with petioles 2-3 mm long, ovate to weakly spatulate, 2.7-6 em long, 1-2.7 em wids, at apex obtuse to acute, at base obtuse to subauricu- late, with both surfaces strigose with hairs 0.5-0.8 mm long; margin entire to serrulate, ciliate with hairs 0.3 mm long. Heads 6-7 mm tall, 19-38 mm diam. Outer involucre spread- ing or somewhat cupulate, 10-14 mm diam; bracts 5, con- nate 1/5 their length, imbricate, ovate, 4-6 mm long, 3.5- 6 mm wide, at apex acute and sometimes purple-tinged, with abaxial surface villous with hairs 0.5 mm long (some- times mostly glabrous) ; margin scarious. Fruits 2-2.3 mm leng, with lateral surfaces smooth to rugose-aculeate. Ray florets 11-14; ligules yellow-orange to yellow, narrowly el- liptic-oblong to oblanceolate, 5-13 mm long, 2-3 mm wide. Disc florets 80-110; corollas yellow, 1 mm diam, with throat 1.8 mm and tube 0.7 mm long. Paleae oblanceolate, 3.2 mm long, 0.6 mm wide; apex yellow, with margin entire; midrib prominent, glabrous. Chromosome number, n = Scattered in pine-oak forests in Nuevo León, Michoacán, Oaxaea, and Pueb!a, Mexico (Fig. 26), 1800-2500 m. Flowering dates, Jul-Sep. The morphology of the hexaploid M. aureum suggests that it may have been derived from the diploid M. mon- tanum by autopolyploidy, perhaps through the union of un- reduced gametes from diploid and tetraploid plants. The two taxa are so similar, in fact, that experimental studies may indicate that M. aureum should be treated as simply a polyploid race of M. montanum. The extant chromosomal voucher specimens, however, do show a considerable amount 194 Rhodora [Vol. 74 of morphological divergence albeit of a quantitative na- ture. In view of these morphological differences and in the absence of intensive field and laboratory studies, I prefer to maintain the two taxa as distinct species at the present time. REPRESENTATIVE SPECIMENS. Mexico. MICHOACAN: vicinity of Morelia, Cerro San Miguel, 15 Sep 1910, Arsène 5219 (GH, NY, US); ca 20 mi W of Ciudad Hidalgo, 4 Aug 1960, King 3617 (DS, NY, TEX, uc, Us); 23 mi NW of Ciudad Hidalgo, 19 Aug 1966, Stuessy 682 (TEX); 21 mi NW of Ciudad Hidalgo, 19 Aug 1966, Stuessy 684 (TEX). NUEVO LEÓN: Dulces Nombres, 20 Jul 1948, Meyer & Rogers 2828 (vs), 10 Aug 1948, 2911 (US). OAXACA: Nochistlán, Cuesta Blanca, rd to Montelobos, 22 Jun 1907, Conzatti 1865 (F); ca 8 mi SE of Nochistlan, 27 Jul 1960, King 3529 (DS, NY, TEX, UC, US); 7 mi SE of Nochistlán, 15 Aug 1966, Stuessy 663 (TEX). PUEBLA: Cerro del Oro, Aug 1909, Purpus 3822a (F, GH, NY, UC, US). VI. Melampodium section Alcina (Cav.) DC. Melampodium L. sect. Alcina (Cav.) DC. Prodr. 5:520. 1836. Alcina Cav. Ic. 1:10. t. 15. 1791. Type species: Melampodium perfoliatum (Cav.) H.B.K. Tap-rooted annuals; leaves ovate or oblong, sessile or petiolate, with margins entire or weakly serrate; outer in- volucre spreading, with bracts 3-5, at margins herbaceous, ovaries of the disc florets ovoid, less than 0.7 mm long, rudi- mentary (Figs. 3 & 4); fruits at apex nearly smooth or moderately sculptured and without hoods; chromosome base number, x — 11. Species 35-37. 35. Melampodium perfoliatum (Cav.) H.B.K. Nov. Gen. Sp. 4:274. 1820. Alcina perfoliata Cav. Cav. le. 1:11. f. 15. 1791. TYPE: “IMPERIO MEXICANO": cultivated in Bot. Gard. Madrid, flowering 29 Nov (Lectotype chosen, MA!; isotypes, MA [2] !; photograph of lectotype, OS! TEX!). Wedelia perfoliata (Cav.) Willd. Sp. Plant. 3(3) :2335. 1803. Polymnia perfoliata (Cav.) Poiret, Encycl. Méthod. Botan. 5:506. 1804. 1972] Melampodium — Stuessy 195 Camutia perfoliata Bonato ex Steud. Nom. Bot. 146. 1821. pro syn. Melampodium connatum Sessé & Moc. ex DC. Prodr. 5:521. 1836. pro syn. Polymnia revoluta Autor. ex Steud. Nom. Bot. ed. 2. 2: 377. 1841. pro syn. Annual herbs, 20-150 cm tall. Stems erect, 1-11 mm diam, glabrous below to very minutely puberulent above. Pe- duncles 0.6-11 em long. Leaves sessile, rhombic-deltoid (rarely ovate-obovate) and tapering to an auriculate-con- nate base (sometimes obtuse on upper leaves), 2-21 em long, 0.9-15 em wide, at apex acute, with both surfaces strigose with hairs 0.2-0.3 mm long; margin obscurely to strongly irregularly serrate. Heads 5-7 mm tall, 8-11 mm diam. Outer involucre spreading, 15-32 mm diam; bracts 5, slightly imbricate and connate at base, oblong-elliptic, 6-20 mm long, 4-11 mm wide, at apex obtuse, with abaxial surface occasionally strigose with hairs 0.2-0.5 mm long ; margin herbaceous. Fruits 4-7 mm long, with lateral sur- faces smooth to moderately striate or nervose. Ray florets 8-13; ligules yellow-orange, oblong-elliptic, 2.5-4 mm long, 0.9-2.3 mm wide. Disc florets 30-45; corollas yellow, 1.5 mm diam, with throat 1 mm and tube 0.8 mm long. Paleae oblong-elliptic, 2.1 mm long, 0.7 mm wide; apex light yel- low, with margin dentate-laciniate; midrib distinct, gla- brous. Chromosome numbers, n = 11 & 12. Tropical deciduous and pine-oak forests in Mexico, Guate- mala, and Costa Rica, introduced to Cuba and Los Angeles, California (Fig. 21), 610-2440 m. Flowering dates, Jan- Dec. M. perfoliatum is the second most widely spread species in the genus (after M. divaricatum), but apparently it has been introduced only at one point in the United States (near Los Angeles). Because the city of Los Angeles and surrounding suburbs have grown tremendously in the last 196 Rhodora [Vol. 74 Fig. 27. Map of Mexico, Central America and neighboring coun- tries showing distribution of Melampodium perfoliatum. 60 years, it is likely that M. pei foliatum, however weedy and aggressive, has not managed to survive. Specimens ex- amined from Los Angeles Co., California, support this con- tention, as there are no records later than 1902. At maturity M. perfoliatum is very easy to recognize because the large fruits remain temporarily in a conspicu- ous ring within the spreading foliaceous outer phyllaries. In the field the bases of the lower leaves are markedly per- foliate and also help distinguish the species. On herbarium specimens, however, this latter characteristic is less help- ful because the bases of the upper leaves are sometimes obtuse, and such leaves, due to their more convenient size, most frequently are selected and pressed by collectors. 1972] Melampodium — Stuessy 197 An interesting nomenclatural problem is associated with the earliest date of publication for the combination Melam- podium perfoliatum. It has generally been attributed to H.B.K. in 1820. An anonymous author (perhaps M. A. Valdes, fide Langman, 1964) in the Gazeta de México of 1792-93 (vol. 5, p. 243) reported the events taking place at a public botanical examination in La Real y Pontifica Uni- versidad in Mexico City on December 7, 1792. At this gathering a student, Sebastian Gomez Moron, read a paper in which he “. . . determinó y describió con la mayor prop- iedad el Melampodio reunido, o Melampodium perfoliatum. ... A footnote elaborates in some detail the circumstances surrounding this new combination which precedes that of H.B.K. by at least 27 years. According to the anonymous author, it was the “opinion” of the “Expedición” (i.e., Sessé, Castillo, and Mocino) traveling through Mexico, that Alcina perfoliata of Cavanilles should not be generically separated from the well-known Melampodium. The stu- dent, Gómez Moron, apparently looked into the matter more closely and agreed with this contention. The author adds another sentence in the footnote, however, that changes the nomenclatural complexion of the situation: ‘‘Los Botanicos que tuvieren ocasion de cotejar la Planta, 6 la exacta de- scripcion de este autor [Cavanilles] con qualquiera de los Melampodios conocidos, pesarán los fundamentos que me- dian entre una y otra parte, y fixarán el nombre con que debe conocerse en lo sucesivo esta nueva especie." The au- thor states that botanists who might compare the species with the existing taxa of Melampodium will have to weigh the facts and decide what the correct name should be. I interpret this hesitation to mean that the author did not accept the new combination at this time, which therefore precludes its validation (Article 34, Provisions 1 and 2, Intern. Code Bot. Nomen.; Lanjouw et al. 1966). REPRESENTATIVE SPECIMENS. Costa Rica. ALAJUELA: between San Josecito & La Garita, 18 Aug 1963, Jiménez 1074 (F). CARTAGO: vicinity of Turrialba, 5 Feb 1965, Godfrey 66184 (FSU); Turrialba, 198 Rhodora [Vol. 74 Instituto Interamericano de Agricolas, 20 Aug 1962, King 5350 (TEX, uc, US); 4 km E of Cartago, 28 Aug 1962, King 5407 (TEX, UC, US). SAN JOSE: Rio Torres, San José, 31 Jan 1965, Godfrey 66064 (FSU). Cuba. LA HABANA: Santiago de las Vegas, 30 Jun 1904, Baker & Wilson 516 (F, NY, UC, US); Laguna de Ariguanabo at Cayo La Rosa, 17 Jul 1923, Ekman 16932 (NY); Santiago de las Vegas, 10 Oct 1904, Hermann 148 (F, NY, US); Laguna de Castellano, 27 Dec 1910, Wilson 9553 (NY [2]). PINAR DEL RIO: near Potrerito, Sumidero, 19 May 1951, Alain 1819 (NY). Guatemala. GUATEMALA: city limits of Guatemala, 9 Jul 1960, King 3248 (DS, NY, TEX, UC, US); Guate- mala City, 1 Aug 1966, Stuessy 603 (TEX). HUEHUETENANGO: ca 1 mi S of Huehuetenango, 18 Jul 1960, King 3410 (DS, NY, TEX, UC, US) ; 2 mi S of Huehuetenango, 26 Jul 1966, Stuessy 580 (TEX). QUICHÉ: 1942, Aguilar 1942 (F). Mexico, AGUASCALIENTES: below Presa Calles, 31 Aug 1939, Shreve 9278 (ARIZ, GH). CHIHUAHUA: Arroyo Hondo, Sierra Charuco, 11 Sep 1985, Gentry 1752 (ARIZ, F, GH, NY, uc, US); Municipio de Guadalupe & Calvo, Nabogame, 11 Oct 1959, Gentry, Correll & Arguelles 17970 (US). COLIMA: Colima, 9 Jan- 6 Feb 1891, Palmer 1163 (GH, NY, US); Alzada, 25 Aug 1966, Stuessy 729 (TEX). DISTRITO FEDERAL: Mixcoac, Olivar, 7 Sep 1913, Arsène s.n. (US); San Andrés, Aug 1930, Lyonnet 569 (US); Pyramid of Cuicuileo, Tlalpan, 15 Aug 1935, MacDaniels 41 (F); near Tlalpan, 1 Sep 1936, MacDaniels 723 (F); San Ángel, 12 Aug 1910, Orcutt 3548 (F). DURANGO: vicinity of Durango, Jun 1896, Palmer 286 (F, GH, NY, UC, US) ; ca 30 mi N of Durango, 12 Nov 1959, Gentry & Arguelles 18152 (US). GUANAJUATO: Guanajuato, Dugés 461 (GH), 462 (GH); Obregón, 13 Oct 1895, Volkens 1134 (GH, NY). GUERRERO: Taxco, 28 Sep 1937, Abbott 445 (GH), 5 Sep 1959, Degener 26220 (NY); Mina, Zihuaqueo-Filo Mayor, 20 Aug 1936, Hinton et al. 9293 (GH, US). JALISCO: La Barranca, 24 Nov 1930, Jones 27773 (ns [2], F, NY, POM, UC); NE slopes of Nevado de Colima, below Canoa de Leoncito, 17 Oct 1952, McVaugh 13576 (SMU, Us); near Santa Monia, 12-13 Nov 1952, MeVaugh 14107 (smu, Us); near Guadalajara, 29 Oct 1888, Pringle 1767 (F, GH, NY [2], UC, US); 2 mi NW of La Barca, 23 Aug 1966, Stuessy 705 (TEX). MEXICO: Valley of México, Tizapán, 12 Feb 1865-66, Bourgeau 841 (GH); Temascaltepec, Temas- caltepec, 2 Sep 1932, Hinton 1521 (NY); Temascaltepec, Chorrera, 25 Sep 1932, Hinton 1838 (GH, NY, US); Temascaltepec, Ixtapán, 2 Nov 1932, Hinton 2480 (Ny, UC, US); % mi N of Tepetlixpa, 16 Aug 1966, Stuessy 668 (TEX). MICHOACÁN: 13 mi S of Uruapán, 28 Oct 1962, Cronquist 9745 (NY, SMU, TEX, US); ca 11 mi W of the Michoacán-Mexico state border on rte 15, 3 Aug 1960, King 3600 (DS, NY, SMU, TEX, UC, US); E of San Juan Nuevo (ca 8 km S of Uruapán), 11-15 Oct 1961, King & Soderstrom 4707 (NY, SMU, TEX, UC, US); 2 mi W of Ciudad Hidalgo, 21 Aug 1961, Powell & Edmondson. 1972] Melampodium — Stuessy 199 816 (DS, F, TEX); 0.8 mi NW of Zitacuaro, 1 Sep 1965, Stuessy 379 (TEX). MORELOS: vicinity of Cuernavaca, 28 Jan 1932, Asplund 517 (NY), 31 Aug 1910, Orcutt 3892 (F, US). NAYARIT: vicinity of Jalisco, 10 Nov 1925, Ferris 5890 (DS, GH, US); Tepic, 14 Feb 1927, Jones 23398 (pom), 10 Feb 1927, Jones 23402 (POM). PUEBLA: vicinity of Puebla, Manzanilla, Nov 1908, Arsène 3523 (US); ca 5 mi W of Puebla, 29 Jul 1960, King 3560 (NY, TEX, UC, US) ; vicinity of Puebla, Hacienda Batan, 20 Dec 1911, Nicolas 6150 (GH, US); Teteles, 9 Jul 1966, Stuessy 497 (TEX); 3 mi W of Puebla, 14 Jul 1966, Stuessy 503 (TEX). OAXACA: Cerro San Felipe, 20 Sep 1908, Conzatti 2291 (F); vicinity of Totontepec, 15-20 Jul 1894, Nelson 715 (US); Sierra de San Felipe, Oct 1894, Smith 351 (F); N city limits of Zimatlán, 14 Aug 1966, Stuessy 654 (TEX). SAN LUIS POTOSÍ: 32 mi E of San Luis Potosí, 19 Sep 1960, Crutchfield & Johnston 5643a (TEX); near San Luis Potosí, Aug 1876, Schaffner 270 in part (GH, NY). SINALOA: Ocurahui, Sierra Surotato, 1-10 Sep 1941, Gentry 6350 (ARIZ, DS, GH, NY). VERACRUZ: Zacualpán, Dec 1907, Purpus 2849 (F, GH, NY, UC, US) ; Mt Orizaba, 22 Aug 1891, Seaton 54 (F, GH, NY, us); 8 mi NW of Jalapa, 9 Jul 1966, Stuessy 494 (TEX) ; 34 mi NW of Tehuacán, 14 Jul 1966, Stuessy 505 (TEX); 4% mi W of San Juan, near Pico de Orizaba, 15 Jul 1966, Stuessy 511 (TEX). UNITED STATES. Cali- fornia: LOS ANGELES CO.: Cienega, Aug 1902, Braunton 625 (DS, UC [2], us); Los Angeles, Aug 1902 [037], Grant 763 (ARIZ, F, UC), Sep 1879, James s.n. (NY, US); between Los Angeles & Compton, Nov 1895, McClatchie s.n. (DS, NY, POM); Los Angeles, Oct 1882, S. & W. Parish 1171 (ps, F, GH, NY, US [2]). 36. Melampodium glabrum S. Wats. Proc. Amer. Acad. Arts & Sci. 26:139. 1891. TYPE: MEXICO: Guanajuato, val- ley near Irapuato, 20 Sep 1889, C. G. Pringle 2821 (Holo- type, GH!). Annual herbs, 25-60 cm tall. Stems erect, 1-10 mm diam, somewhat soft and watery when fresh, glabrous near base and covered with bladdery hairs toward apex. Peduncles 2.5-5.5 em long. Leaves sessile, oblong-oblanceolate, 2.5-6 cm long, 0.7-2.1 em wide, at apex obtuse, at base subauricu- late, with both surfaces waxy and glabrous; margin sinu- ate-dentate. Heads 4.3-5 mm tall, 7-11 mm wide. Outer involucre spreading, 8-10(14) mm diam; bracts 5, fleshy, slightly connate at base, imbricate, ovate-orbiculate, 4.2-6 mm long, 3.8-4.5 mm wide, at apex obtuse with abaxial sur- face glabrous; margin herbaceous. Fruits 2.5-2.9 mm long, 200 Rhodora [Vol. 74 with lateral surfaces smooth to moderately striate or ner- vose. Ray florets 6-8; ligules yellow, oblong-ovate, 1.3-3.4 mm long, 0.7-1.4 mm wide. Disc florets 30-50; corollas yellow, 1 mm diam, with throat 0.9 mm and tube 0.8 mm long. Paleae oblanceolate-elliptic, 1.6 mm long, 0.5 mm wide; apex yellow, with margin entire; midrib prominent, glabrous. Chromosome number, n = 11. Very moist areas in pine-oak forests or high altitude grasslands in the Mexican states of Guanajuato, Jalisco, and Michoacan (Fig. 13), ca 1890 m. Flowering dates, Aug- Sep. This species is an attractive, fleshy, little herb found often in moist depressions on roadsides. The large folia- ceous outer involucral bracts and conspicuous blackish fruits as well as the n — 11 chromosome number indicate that M. glabrum is allied closely to M. perfoliatum. REPRESENTATIVE SPECIMENS. Mexico. GUANAJUATO: near San Miguel Allende, Aug 1947, Kenoyer 2421 (GH). JALISCO: valley of the Lerma near La Barca, 16 Sep 1891, Pringle 3863 (DS, G, GH [2], MA, Msc, NY [2], P, vc [2], US, w); 3 mi N of La Barca, 24 Aug 1966, Stuessy 708, 709 (TEX). MICHOACÁN: vicinity of Morelia, 13 Sep 1910, Arsène s.n. (US); 9 mi S of jet rte 15 on rd to Cotija, 24 Aug 1966, Stuessy 714 (TEX). 37. Melampodium nutans Stuessy, Brittonia 22:122. f. 9. 1970. TYPE: MEXICO: Colima, 12 mi S of Colima on rte 110, 25 Aug 1966, T. F. Stuessy 721 (Holotype, US!; isotypes, DS!F!GH! MICH! NY! TEX! UC!). Annual herbs, 16-40 em tall. Stems erect, 1.2-2 mm diam, hispidulous-strigillose with hairs 0.2-0.5 mm long. Pedun- cles 5.5-6.5 cm long. Leaves ovate to rhombic, with taper- ing petioles 1-3 cm long, blades 4-5 cm long, 2-4 cm wide, at apex acuminate, at base attenuate, with both surfaces strigose with hairs 0.5 mm long; margin subentire to ser- rate. Heads 4-6 mm tall, 7-8(10) mm diam. Outer in- voluere spreading, 8-14 mm diam; bracts 3 (rarely 4), slightly connate at base, separate, ovate to narrowly ovate, 4-7 mm long, 2.5-4 mm wide, at apex acute, with abaxial 1972] Melampodium — Stuessy 201 surface strigose with hairs 0.3-0.5 mm long; margin her- baceous. Fruits 1.8-2.1 mm long, with lateral surfaces ribbed and reticulated; hood at apex muticous to cirrhous'*, with tapering appendage up to 6 mm long. Ray florets 5-7; ligules yellow-orange, orbiculate, margins imbricate, 2-3 mm long, 2-4 mm wide. Disc florets 40-80; corollas yellow- orange, 1.7 mm diam, with throat 0.7 mm and tube 1 mm long. Paleae narrowly obovate, 2 mm long, 1 mm wide; apex yellow, with margin laciniate; midrib distinct, pube- rulent with hairs 0.1 mm long. Chromosome number, n — 11. Tropical deciduous forests of Colima, Jalisco, Michoacán and Oaxaca, Mexico (Fig. 13), 50-500 m. Flowering dates, Aug-Nov. The hoods on fruits of M. nutans are shorter and thicker than are the hoods of species of section Melampo- dium. Such a difference is not unexpected, however, be- cause M. nutans differs in many other respects from the taxa in that section. Due to the spreading outer involucral bracts, rudimentary disc ovaries, herbaceous habit, and chromosome number of » — 11 of this species, it clearly belongs in section Alcina near M. perfoliatum and M. gla- brum. REPRESENTATIVE SPECIMENS. Mexico. COLIMA: ca 11 mi SSW of Cd Colima, 21 Sep 1958, McVaugh 18070 (MICH), 25 Nov 1959, McVaugh & Koelz 1054 (MICH); 12 mi S of Cd Colima, 25 Aug 1966, Stuessy 720 (TEX). JALISCO: between Sierra de los Corales & Tepal- catepec, Michoacán, 26 Oct 1963, Feddema 2238 (MICH); Tecalitlan, 26 Oct 1963, Rzedowski 17514 (MICH, OS). MICHOACÁN: ca 5 km NW of Aquila, 23 Nov 1963, Feddema 2700 (MICH); 15 mi SSW of Apatzingán, 2 Aug 1940, Leavenworth 431 (F, GH, NY), 432 (F); 8 km NW of Aquila, 23 Nov 1963, Rzedowski 17921 (MICH, 0s). OAXACA: 3 km SW of Puerto Ángel, 24 Sep 1965, Rzedowski 21155 (os). “Isotype material in my preserved personal collection has fruits with both muticous hoods and hoods with long cirrhous appendages in the same head (cf. Fig. 9E in Stuessy, 1970b). 202 Rhodora [Vol. 74 DOUBTFUL SPECIES Melampodium angulatum Sessé & Moc. Pl. Nov. Hispan. 149. 1890. TYPE: MEXICO: Guerrero, Ayahualtempa, Aug 1789, M. Sessé et al. s.n. (Holotype, MA?). From the de- scription and locality data this taxon seems most similar to M. perfoliatum, although the leaf undersurface in the latter species is never “tomentose.” Melampodium dichothomum Sessé & Moc. Pl. Nov. His- pan. 149. 1890. TYPE: MEXICO: Guerrero, Mazatlán [ca 15 km S of Chilpancingo; Sprague (1926), and Rickett (1947) ], Jun 1789, M. Sessé et al. “3977.” MA has a mixed sheet with both M. montanum and M. divaricatum and an- other sheet with M. divaricatum. A sheet at F has a frag- ment of only M. divaricatum. The description more closely resembles M. montanum, but both M. divaricatum and M. montanum are found in Guerrero, and neither seems ex- plicitly excluded by the description. Melampodium diversifolium Sessé & Moc. Fl. Mex. ed. 2. 197. 1894. TYPE: locality unknown, 1787-1804, M. Sessé et al. s.n. (Holotype, MA?). The description seems to refer to M. linearilobum or M. longipes, but not mentioned are sev- eral diagnostic characters such as the leaf undersurface and margins of the outer involucral bracts. Melampodium pinnatum Sessé & Moc. Pl. Nov. Hispan. 149. 1890. TYPE: MEXICO: Michoacan, Apatzingán, Oct 1790-91, M. Sessé, J. M. Mocino & J. Castillo s.n. (Holo- type, MA?). The description is too general to allow any specific placement. Melampodium rhombifolium Sessé & Moc. Pl. Nov. His- pan. 149. 1890. TYPE: MEXICO: México, San Augustín, Sep 1787-1804, M. Sessé et al. s.n. (Holotype, MA?). The character “Receptaculum nudum,” if correct, would exclude this taxon from Melampodium. Melampodium vhombifolium Sessé & Moc. Fl. Mexic. ed. 2. 195 [second on page]. 1894. nom. illegit. non Sessé & Moc. 1890. nec Sessé & Moc. [first on page]. 1894. TYPE: 1972] Melampodium — Stuessy 203 MEXICO: “Hortis Sancti Augustini, prope Mexicum," Aug- Sep 1787-1804, M. Sessé et al. s.n. (Holotype, MA?). Spe- cific affinity cannot be determined from the description. EXCLUDED NAMES Melampodium L. sect. Acanthospermum (Schrank) Baillon, Hist. Plant. 8:231. 1882. =Acanthospermum Schrank, Pl. Rar. Hort. Acad. Monac. 53. 1820. Melampodium achillaeoides (Less.) Hemsley, Biol. Centr. Am. Bot. 2:145. 1881. —Villanova achillaeoides (Less.) Less. Syn. Gen. Comp. 256. 1832. TYPE: MEXICO: “los Llanos de Perote," Sep 1828, C. J. W. Schiede 354 (Holotype, HAL!; photograph of holotype, os! TEX !). Melampodium anomalum M. E. Jones, Contr. West. Bot. 18:72. 1933. TYPE: MEXICO: Jalisco, near Guadalajara, La Barranca, 17 Nov 1930, M. E. Jones 27727 (Lectotype, POM; isotype, MO! NY!; photograph and fragment of lecto- type, US!). =Tragoceras schiedeanum Less. Linnaea 9:269. 1834. fide Blake (1945) and Torres (1963). Melampodium australe Loefl. Iter Hisp. 268. 1758. TYPE: VENEZUELA: vicinity of Barcelona, 15 Feb 1755, P. Loefling "151" (Holotype, LINN). —Acanthospermum australe (Loefl.) Kuntze, Rev. Gen. Pl. 1:303. t. 22, h-m. 1891. Melampodium baranguillae Spreng. Syst. 3:619. 1826. TYPE: COLOMBIA: “Ad fl. Magdalenae," date unknown, C. G. Bertero s.n. (Holotype, P-co). Blake (1930) saw the type of M. baranguillae in the Schultz-Bipontinus Her- barium and found “it to be identical with Sclerocarpus co- lumbianus Rusby & Blake." Melampodium bonairense Boldingh, Fl. Neder. West-Ind. Bil 393. 1913. Fl. Dutch West Ind. Is. 2:107. t. 9. 1914. TYPE: DUTCH WEST INDIAN ISLANDS: Island Curacao, Mal- pais, 28-30 Oct 1909, I. Boldingh M.14 (Holotype, U; para- type [Boldingh 7401], P!; fragment of P paratype, Us!; photograph of K paratype [Boldingh 7401], us!). —Balti- mora sp. 204 Rhodora [Vol. 74 Melampodium camphoratum (L.f.) Baker in Martius, Fl. Bras. 6(3) :161. 1884. —Unzia camphorata L.f. Suppl. 368. 1781. TYPE: SURINAM: “locis arenosis," date unknown, C. G. Dalberg s.n. (HOLOTYPE, LINN; photograph of holotype, os!). Melampodium digynum (Steetz in Seem.) Jackson, Index Kew. 2:188. 1895. Unxia digyna Steetz in Seem. Bot. Voy. Herald. t. 30. 1853. text 154. 1854. TYPE: PANAMA: “about Panama [City]," Nov 1846, B. C. Seemann 46 (HOLOTYPE, BM; photograph and fragment of K isotype, us!; photo- graph of K isotype, F! us [2]!). =Unxia camphorata L.f. Suppl. 368. 1781. Melampodium? dombeyanum DC. Prodr. 5:520. 1836. TYPE: PERU: locality and date unknown, J. Dombey s.n. (Holotype, G-DC; isotype, P!; photograph of holotype, IDC 800. 923: III. 6!; photograph of P isotype, OS! TEX!). S. F. Blake (1930) saw the type in the Prodromus Herbarium and notes that it “has shown its identity with Sigesbeckia flosculosa” L’Her. Stirp. Nov. 37. t. 19. 1785-86. Melampodium durandi (A. Gray) M. E. Jones, Contrib. West. Bot. 15:156. 1929. Hemizonia durandi A. Gray, Proc. Amer. Acad. Arts & Sci. 6:549. 1865. TYPE: CALIFORNIA: Nevada Co., “.. . in the vicinity of Nevada [City]" [Du- rand, 1855], ca. 3000 ft., 1851, H. Pratten s.n. (Holotype, PH?). —Madia minima (A. Gray) Keck, Madroño 10:22. 1949. Melampodium geminatum Brandg. Zoe 5:223. 1905. TYPE: MEXICO: Sinaloa, Cofradia [ca 30 mi E of Culiacán near Durango border], 24 Oct 1904, T. S. Brandegee s.n. (Lecto- type chosen, UC!). = Baltimora sp. Melampodium? hidalgoa DC. Prodr. 5:521. 1836. nom. superfl. of Melampodium ternatum (Llave & Lex.) DC. —Hidalgoa ternata Llave & Lex. Nov. Veg. Descr. 1:15. 1824. 1972] Melampodium — Stuessy 205 Melampodium hirsutum (Rich.) Jackson, Index Kew. 2: 188. 1895. Unwia hirsuta Rich. Act. Soc. Hist. Nat. Par. 1:105. 1792. TYPE: FRENCH GUIANA: locality unknown, 1792, J. B. Leblond 346 (Holotype, P; isotype, G!; photo- graph of G isotype, OS! TEX!). =Unxia camphorata L.f. Suppl. 368. 1781. Melampodium humile Sw. Prodr. Veg. Ind. Occ. 114. 1788. TYPE: JAMAICA: “Domingo,” 1783-1787, O. Swartz s.n. (Holotype, S; photograph of B isotype, TEX!). —Acan- thospermum humile (Sw.) DC. Prodr. 5:222. 1836. Melampodium L. sect. Lecocarpus (Dcne.) Baillon, Hist. Plant. 8:231. 1882. —Lecocarpus Dene. Bot. Voy. Venus, t. 14. 1846. Melampodium minimum (A. Gray) M. E. Jones, Con- trib. West. Bot. 15:156. 1929. Hemizonia minima A. Gray, Proc. Amer. Acad. Arts & Sci. 6:549. 1865. TYPE: IDAHO: "Dry soil, near Soda Springs," 8600 ft., 1860-1862, W. H. Brewer s.n. (Holotype, GH?). —Madia minima (A. Gray) Keck, Madrofio 10:22. 1949. Melampodium minutiflorum M. E. Jones, Contrib. West. Bot. 18:72. 1933. TYPE: MEXICO: Guadalajara, La Bar- ranca, 17 Nov 1930, M. E. Jones 27738 (Lectotype, POM; isotype, MO! US!; photograph and fragment of lectotype, US!). —Galeana pratensis (H.B.K.) Rydberg, Fl. N. Amer. 94(1) :42. 1914. Melampodium paludicola Taub. Engl. Bot. Jahrb. 21:455. 1896. TYPE: BRAZIL: Goias, “in paludibus ad flumen Parana- hyba [Paranaiba]," Feb 1893, E. Ule 2978 [—397] (Holo- type, P!; isotype, F!; fragment of holotype, US!; photo- graph of holotype, F! os! TEX! Us!). Definitely not a Mel- ampodium; “the description of which suggests a Sclerocar- pus" (Robinson, 1901). Melampodium? ruderale Sw. Fl. Ind. Occ. 3:1372. 1806. TYPE: JAMAICA: “in ruderatis Jamaicae australioris," date 206 Rhodora [Vol. 74 unknown, O. Swartz s.n. (Holotype, 8?). — Eleutheran- thera ruderalis (Sw.) Sch. Bip. Bot. Zeit. 24:259. 1866. Melampodium suffruticosum Baker in Martius, Fl. Bras. 6(3):162. 1884. TYPE: VENEZUELA: “prope Esmeralda in ditione fluminis Orinoco," Dec 1853, R. Spruce 3225 (Holo- type, K!; isotypes, G! NY! P!; photograph of holotype, os! TEX! US!). =Unxia suffruticosa (Baker) Stuessy, Brit- tonia 21:319. 1970. Melampodium? ternatum (Llave & Lex.) DC. Prodr. 5: 521. 1836. —Hidalgoa ternata Llave & Lex. Nov. Veg. Descr. 1:15. 1824. TYPE: MEXICO: Veracruz, “. .. in mar- gine densissimarum silvarum, prope ultimam cataractam fluminis Blanco, non longe à S. Jose del Corral [near Ori- zaba]," Apr [year?], P. La Llave s.n. (Holotype, MA?). Melampodium L. subg. Unxia (L.f.) Baker in Martius, Fl. Bras. 6(3) :162. 1884. —Unzia L.f. Suppl. 56. 1781. ACKNOWLEDGEMENTS This paper represents a modification of a dissertation submitted to the Graduate School of the University of Texas at Austin in partial fulfillment of the requirements for the degree of Doctor of Philosophy. Sincerest thanks go to Dr. B. L. Turner for the initial suggestion of the problem and for his guidance throughout the development of the study. Appreciation is also owed to Drs. Marshall C. Johnston, Rogers MeVaugh, Lloyd H. Shinners, and Mr. Imre J. Eifert for their help with literature and nomen- clature; Mr. Geza Knipfer for preparation of most of the base maps used for the species distributions; Messrs. Ar- nold B. Birdsong, Imre J. Eifert, John B. Schoenhofen and Drs. Robert S. Irving and Walter Renold for their help and companionship on field trips; Dr. C. E. B. Bonner for cam- era lucida drawings of diagnostic features of the type of Melampodium angustifolium; Dr. Alicia Lourteig for in- formation concerning type-material at Paris; Dr. J. F. M. Cannon for information regarding Calendula decumbens; 1972] Melampodium — Stuessy 207 Dr. Harold Robinson for help in interpreting ovarian disc structures; Dr. Rogers MeVaugh for special help with typi- fication of the Sessé & Mocifio names; Miss Suzanne Moor- head for drawing Figs. 6-9 and 10; and Drs. Rogers Mc- Vaugh, Ronald L. Stuckey, Rolla Tryon, and B. L. Turner for critically reading the entire manuscript. Finally, many thanks are due my wife, Carol, who courageously typed cards for representative specimens and spent countless hours typing the drafts of the manuscript. Field work was partially supported by NsF Traineeship 4128 (to W. F. Blair), NSF grant GB-1458 (to H. J. Arnott), and a Sigma Xi Grant-in-Aid. Collective appreciation is expressed to the various cura- tors and directors of herbaria from which loans of speci- mens were made as indicated in the following list (abbre- viations after Lanjouw and Stafleu, 1964) : A, ARIZ, B, C, DS, ENCB, F, FSU, G, GH, HAL, K, LL, MA, MICH, MO, MSC, NY, P, POM, RSA, S, SMU, TEX, UC, UPS, US, W, WIS. LITERATURE CITED ALSTON, A. H. G. 1934. Notes on Selaginella. VI. The Selaginellae collected by Thaddeus Haenke and described by Karel Borewog Presl. J. Bot. (Lond.) 72: 223-226. BAILLON, H. E. 1882. Histoire des Plantes, vol. 8, p. 231. Paris. BAKER, J. G. 1884. Melampodiwm, p. 159-162. In C. F. P. von Martius, Flora Brasiliensis, vol. 6(3). Munich. BENTHAM, G., and J. D. HOOKER. 1873. Genera Plantarum, vol. 2, p. 348-349. Reeve & Co., London. BLAKE, S. F. 1924. New American Asteraceae. Contrib. U. S. Nat. Herb. 22: 587-661. 1926. Five new American Melampodinae. J. Wash. Acad. Sci. 16: 418-422. 1930. Notes on certain type specimens of American Asteraceae in European herbaria. Contrib. U. S. Nat. Herb. 26: 227-268. 1945. Asteraceae described from Mexico and the southwestern United States by M. E. Jones, 1908-1935. Contrib. U. S. Nat. Herb. 29: 117-137. CANDOLLE, A. P. DE. 1836. Prodromus, vol. 5, p. 517-521. Paris. Cassini, A. H. G. 1829. Melampodium, Zarabellia, and Alcina, p. 237-244. In G. Cuvier (ed.), Dict. Sci. Natur., ed. 2, vol. 59. Paris. 208 Rhodora [Vol. 74 COCKERELL, T. D. A. 1905. The name Melampodium. Torreya 5: 70. COLEMAN, J. R. 1970. Additional chromosome numbers in Brazilian Compositae. Rhodora 72: 94-99. DURAND, E. 1855. Plantae Prattenianae. J. Acad. Nat. Sci. Philad. n.s. 3: 79-104. GEISER, S. W. 1948. Naturalists of the Frontier, ed. 2, p. 30-54. Southern Methodist Univ, Press, Dallas. GENTRY, H. S. 1942. Rio Mayo Plants. Carnegie Inst. Wash. Publ. 527: 262. GRAY, A. 1852. Plantae Wrightianae Texano-neo-Mexicanae. Smith- son. Contrib. Knowl. 3(5): 103. 1855. Plantae Novae Thurberianae. Mem. Amer. Acad. ser. 2, 5: 321. 1859. Melampodium, p. 85. In J. Torrey, Botany of the Boundary, United States and Mexican Boundary Survey, vol. 2(1). Washington, D. C. 1884. Synoptical Flora of North America, vol. 1(2), p. 238-239. Ivison, Blakeman, Taylor, and Co., N. Y. HEPPER, F. N. 1968. A standard citation of microfiches. Taxon 17: 604. HITCHCOCK, A. S. 1909. Catalogue of the grasses of Cuba. Contrib. U. S. Nat. Herb. 12: 183-256. (cf. p. 210) HOFFMANN, O. 1890. Melampodium, p. 218. In A. Engler and K. Prantl, Die natürlichen Pflanzenfamilien, vol. 4(5). Leipzig. HUMBOLDT, F. H. A., A. J. BONPLAND, and C. S. KUNTH. 1820. Nova Genera et Species Plantarum, vol. 4, p. 272-274. Paris. KAPADIA, Z. J. 1964. Varietas and subspecies: a suggestion to- wards greater uniformity. Taxon 12: 257-259. KEARNEY, T. H., and R. H. PEEBLES. 1942. Flowering Plants and Ferns of Arizona. U. S. D. A. Misc. Publ. No. 423: 941. , and . 1964. Arizona Flora, ed. 2, p. 810. Univ. Calif. Press, Berkeley. LANGMAN, I. K. 1964. A Selected Guide to the Literature on the Flowering Plants of Mexico, p. 756. Univ. Penn. Press, Phila- delphia. LANJOUW, J., et al. (eds.). 1966. International Code of Botanical Nomenclature. Regnum Veg. 46: 37-38. , and F. A. STAFLEU. 1964. The herbaria of the world. Index Herbariorum, part 1, ed. 5. Regnum Veg. 31: 205-228. LINNAEUS, C. 1737. Critica Botanica. Leiden. 1738. Hortus Cliffortianus, p. 425. Amsterdam. 1753. Species Plantarum, vol. 2, p. 921. Stockholm. McVauaGH, R. 1956. Edward Palmer: Plant Explorer of the American West. Univ. of Oklahoma Press, Norman. MERRILL, E. D. 1923. An Enumeration of Philippine Flowering 1972] Melampodium — Stuessy 209 Plants, pp. 1-128. Philip. Dept. Agr. and Nat. Sci., Publ. No. 18, vol. 1(1). Manila. (cf. p. 49) 1954. The botany of Cook's voyages. Chron. Bot. 14: 161-384. RICKETT, H. W. 1947. The royal botanical expedition to New Spain. Chron. Bot. 11: 1-86. ROBINSON, B. L. 1901. Synopsis of the genus Melampodium. Proc. Amer. Acad. Arts & Sci. 36: 455-466. SAFFORD, W. E. 1905. The useful plants of the island of Guam. Contrib. U. S. Nat. Herb. 9: 1-416. (cf. p. 25) SHREVE, F., and I. L. WiagGINS. 1964. Vegetation and Flora of the Sonoran Desert, p. 1522. Stanford Univ. Press, Calif. SPRAGUE, T. A. 1926. Sessé & Mocifio’s Plantae Novae Hispaniae and Flora Mexicana. Kew Bull. Misc. Inform. 9: 417-425. STUESSY, T. F. 1969a. A new variety and new combination in Melampodiwm (Compositae-Heliantheae). Sida 3: 348-349. 1969b. Re-establishment of the genus Unxia (Com- positae-Heliantheae). Brittonia 21: 314-321. 1970a. The genus Acanthospermum (Compositae- Heliantheae-Melampodiinae): taxonomic changes and generic affinities. Rhodora 72: 106-109. 1970b. Six new species of Melampodium (Com- positae: Heliantheae) from Mexico and Central America. Brit- tonia 22: 112-124. 1971a. Chromosome numbers and phylogeny in Melampodium (Compositae). Amer. J. Bot. 58: 732-736. 1971b. Chromosome studies in Melampodium (Compositae-Heliantheae). Madroño 20: 365-372. 1971c. Systematic relationships in the white-rayed species of Melampodium (Compositae). Brittonia 23: 177-190. TIDESTROM, I., and T. KrrTELL. 1941. A Flora of Arizona and New Mexico, p. 436. Catholic Univ. of Amer. Press, Washington, DC.: TORRES, A. M. 1963. Revision of Tragoceras (Compositae). Brit- tonia 15: 290-302. 1968. Revision of Jaegeria (Compositae-Helian- theae). Brittonia 20: 52-73. TRYON, R. 1955. Selaginella rupestris and its allies. Ann. Mo. Bot. Gard. 42: 1-99. (cf. p. 82) TURNER, B. L., and R. M. KiNG. 1962. A cytotaxonomic survey of Melampodium (Compositae-Heliantheae). Amer. J. Bot. 49: 263-269. DEPARTMENT OF BOTANY THE OHIO STATE UNIVERSITY COLUMBUS, OHIO 43210 210 TAP ON E SERERERESEEEEREEESEEEEE Rhodora LIST OF TAXA OF MELAMPODIUM americanum L. diffusum Cass. pilosum Stuessy longipes (A. Gray) Robins. linearilobum DC. leucanthum Torr. & A. Gray cinereum DC. M. cinereum DC. var. cinereum M. cinereum DC. var. hirtellum Stuessy M. cinereum DC. var. ramosissimum (DC.) A. Gray . argophyllum (A. Gray ex Robins.) Blake . sericeum Lag. . pringlei Robins. . strigosum Stuessy . longicorne A. Gray nayaritense Stuessy . cupulatum A. Gray appendiculatum Robins. . sinuatum Brandg. rosei Robins. . tenellum Hook. & Arn. glabribracteatum Stuessy . longipilum Robins. longifolium Cerv. ex Cav. mimulifolium Robins. . gracile Less. . microcephalum Less. . paniculatum Gardn. . divaricatum (Rich. in Pers.) DC. . costaricense Stuessy . dicoelocarpum Robins. [Vol. 74 1972] 29. M. tepicense Robins. 30. M. sinaloense Stuessy 31. M. bibracteatum S. Wats. 32. M. repens Sessé & Moc. 33. M. montanum Benth. Melampodium — Stuessy 211 33a. M. montanum Benth. var. montanum 33b. M. montanum Benth. var. viridulum Stuessy 34. M. aureum Brandg. 35. M. perfoliatum (Cav.) H.B.K. 36. M. glabrum S. Wats. 37. M. nutans Stuessy D. Doubtful species E. Excluded taxon INDEX TO EXSICCATAE Numbers in parentheses represent the taxa recognized in this study (see preceding list). Abbott, R. Q. 162 (33b); 304 (24); 439 (23); 445 (35). Abloon, Bro. 137 (7b). Abrams, L. R. 12811 (14). Adole, Fr. L. 29 (7b). Aguilar H., M. 25 (23); 322 (25); 1404 (9); 1495 (5); 1567 (9); 1942 (35). Alain, Bro. 1819 (35). Alamán, L. s.n. (5). Alexander, E. L. & E. Hernández X. 2229 [XA218] (31). Allen, P. H. 241 (26); 585 (25). Arséne, Bro. G. 76 (21); 294 (31); 1199 (21); 1896 (33b); 3523 (35); 5219 (34); 6575 (33b); s.n. (35), (36). Arzola, O. 178 (7a). Asplund, C. 517 (35). Bacigalupi, R. C. 607 (6). Baize, R. M. 5 (6). Baker, C. F. 2482 (26). Baker & Wilson 516 (35). Balls, E. K. & W. B. Gourlay B5372 (32). Barkley, F. A. 14142, 14163 (2); 14575C (7b). Barkley, F. A. & G. Gutierrez V. 1777 (26). Barkley, F. A., C. M. Rowell, Jr. & J. B. Paxson 735 (9). Barkley, F. A., G. L. Webster & J. B. Paxson 17M724 (2); 17M732 (24). Barkley, F. A., G. L. Webster & C. M. Rowell, Jr. 7488 (32). Barneby, R. C. 5135 (12). Beaman, J. H. 3043 (33b); 3977 (31); 4852 (32): Benson, L. 10598 (12). Berlandier, J. L. 2017 [607] (Tc); 2153 [733] (23); 2243 [8238]. (Ta) "sa. :(24): Bernoulli, C. G. 709 (5). Bertero, C. G. s.n. (26) ; s.n. (E). 212 Blake, S. F. 7632 (26). Blumer, J. C. 1665 (11); 2128 (12). Boldingh, I. M.14 (E). Bonner, Mrs. J. 55 (Tb). Bonpland, A. J. 3968 (1). Botteri, M. 809 (24). Bourgeau, E. 841 (35); 1629 (24). Brandegee, T. S. 301 (26); 302 (16); sn. (5), (16), (17). Braunton, E. 625 (35). Bray s.n. (Ta). Breedlove, D. E. 6085, 7861 (25); 8906, 12522 (33b). Brenes, A. M. 17471, 20572 (27). Brewer, W. H. s.n. (E). Britton, N. L. & J. F. Cowell 1561 (26). Bruni, T. 15 (7b). Burr, R. D. 135 (6). Busseuil, F. L. s.n. (2). 1628, Calderón, S. 459, 779 (25); 979 (5); 1019 (9). Cameron, E. R. 248 in (7c). Canby, W. M. 133 (7a). Canicosa, E. 1103 (2). Carleton, M. A. 331 (6). Carlson, M. C. 79 (26); 873, 2254 (25). Carter, A. 4880, 4964, 5055 (14). Carter, A., A. M. Alexander & L. Kellogg 2216 (16). Carter, A. & R. Moran (14). Cervantes, J., J. A. Molina & F. A. Barkley 19Bo061 (26). Cervantes, V. s.n. (21). Chambers, K. L. 965 (14). Chamisso, A. s.n. (2). Chase, V. H. 7122 (26); 7533 (23). Cisneros, F. 33 (7a). part 106, 5314 Rhodora [Vol. 74 Clark, O. M. 6635 (Ta); 12377 (12). Clausen, P. s.n. (25). Clausen, R. T. s.n. (24). Clemens, M. S. s.n. (2). Clokey, I. W. 3946 (6). Conzatti, C. 1706 (1); 1865 (34); 2291 (35); 4169 (33a); 5169 (19). Conzatti, C. & V. González 550 (24); 998 (33a). Correll, D. S. 15204 (Ta). Cory, V. L. 50422 (6). Cronquist, A. 9585 (31); 9613 (24); 9745 (35); 10003 (12). Crutchfield, J. & M. C. Johnston 5527 (Te); 5643a (35); 5664 (9); 6120b (26); 6141b (Te). Culwell & Timmons 3091 (6). Curran, H. M. & M. L. Merritt 15837 (2). Dalberg, C. G. s.n. (E). Daniel, Bro. 2290 (26). Darrow, R. A. & H. S. Haskell 2012, 2256 (12). Deam, C. C. 247 (26). Degener, O. 26220 (35). Degener, O. & I. 26504, 26559 (25). Demaree, D. 7594, 12210, 13344 (6). Dohnke, W. C., Jr. 3 (7a). Dombey, J. s.n. (E). Domínguez M., M. 63 (7a). Domínguez M., M. & W. McCart 8217 (Ta). Drouet, F., D. Richards & W. A. Lockhart 3609, 3624 (15). Drushel, J. A. 10441 (7a). Duges, A. s.n. (11); 44 (9); 461 (35). Dunlap, V. C. 557 (27). Earle, F. S, & E. S. 295 (6). Echeverría, J. A. 628 (27). 1972] Eggleston, W. W. 10729 (11). Ekman, E. L. 16932 (35). Erlanson, C. O. 500 (26). Ervendberg, L. C. 92 (23). Escalante, L. 21 (7a). Eyerdam, W. J. & A. A. Beetle 8687 (17). d); 2204 2651 Feddema, C. 279 (29); 832 1209 (18); 1219 (14); (28); 2238 (37); 2577, (29); 2700 (37). Ferris, R. S. 5020 (17); (35). Ferris, R. S. & Y. Mexia 5078 (17). Fisher, G. L. 35321 (5). Fleetwood, R. J. 3840 (7a). Flyr, D. 112 (14); 138 (17); 489 (6). Foxworthy, F. W. 167 (2). Frye, T. C. & E. M. 2415 (7b). DE xn): En. 5890 Furness, (11). García, A., Jr. & R. 47 (7a). García, C. 51 (25). Gardner, G. 287, 3844 (25). Garza, C. & L. de la 46 (7a). Gaumer, G. F. 563 (26); 2349 (23). Gaumer, G. F. & sons 789 (23). Gentle, P. H. 2325 (26). Gentry, H. S. 290 (14); 550 [550M] (9); 790 [790M], 1492, 1570 (15); 1752 (35); 2353 (15); 2553 (9); 5039 (5); 5750 (14); 6153 (5); 6350 (35); 6507-A (30); 7005 (14). Gentry, H. S. & J. Arguelles 18152 (35). Gentry, H. S., D. S. Correll & J. Arguelles 17970 (35). Glassman, S. F. 2003 (25). Melampodium — Stuessy 213 Glaziou, A. 12794 (25). Glocker, E. F. von 19 (26). Godfrey, R. K. 6664 (27) ; 66064, 66184 (35); 67156 (27). Goldman, E. A. 110 (11); 914 (33b). Goldsmith, P. 132 (4); 133 (13). González O., J. 6125 (18). Goodding, L. N. 13 (11); 248-59, 267-62, 580-49, 699-58 (12); 814 (11); 2399a (6). Goodman, G. J. & C. L. Hitchcock 1130 (6). ` Gould, F. W. 2883 (12). Gould, F. W. & H. S. Haskell 3268 (12). Graham, G. J. 5 (26). Graham, J. & M. C. Johnston 4378 (7c); 4777 (23). Grant, G. B. 763 (35). Greenman, J. M. 279 (1). Gregory, D. P. & G. Eiten 205 (24). Guthrie, N. 43 (6). Haenke, T. s.n. (1). Hallberg, B. 900 (33a). Hammerly, B. J. 182 (14). Hancock, W. 25 (2). Harrison, G. J. 8181 (12). Harrison, G. J. & H. J. Fulton 8153 (12). Harrison, G. J. & T. H. Kearney 5702 (12). Harshberger, J. W. 176 (21). Hartman, C. V. 71 (15). Hartweg, C. T. 475 (33a). Heard, N. T. & F. A. Barkley 14542B (7b). Hedrick, D. C. 156 (7b). Heller, A. A. 1632 (6). Henderson, N. C. 63-88, 63-98 (6). Hermann, H. A. van 148 (35). Heyde, E. T. & E. Lux 4213 (26). 214 Hinckley, L. C. 620, 881, 3293, s.n. (11). Hinton, G. B. 1520 (26); 1521 (35); 1661 (24); 1838 (35); 2008 (24); 2017 (33b); 2480 (35); 2513 (5); 4628, 4826 (32); 6207 (24); 6270 (5); 6324 (9); 6334 (28); 60338 (23); 6420 (5); 6436 (3); 9113 (5); 9134 (1); 9205 (24); 9293 (35); 9714 (28); 10719 (24); 11428 (3); 12276 (29); 12871 (1); 13214 (5); 14614 (28); 15587 (1); 16563 (8). Hitchcock, C. L., R. V. Rethke & R. van Raadshooven 4214, 4244 (6). Hoffmann, O. 832 (24). Holman, R. S. & A. F. Dittman 791 (23). Holzner, F. X. 1962 (11). Horr, W. H. 3444 (6). Horr, W. H. & L. H. Franklin E278 (6). Houstoun, W. s.n. (1). Howell, J. T. 10303 (18). Humboldt, F. H. A. von & A. J. Bonpland 142 (26); s.n. (9). Iltis, H. & F., & R. Koeppen 35 (7c). Innes, R. R. & B. H. Warnock 589 (6). Irving, R. 825 (6). Irwin, H. S., R. Souza & R. Reis dos Santos 11303 (25). James, J. F. s.n. (35). Jespersen, B. & H. 2678 (6). Jiménez M., A. 1073 (26); 1074 (35). Johnson, J. C. & F. A. Barkley 16250M (8). Johnston, I. M. 7959 (6). Rhodora [Vol. 74 Johnston, M. C., B. C. Tharp & B. L. Turner 3472 (7a). Jones, F. B. 719, 1359 (7a). Jones, M. E. 117 (6); 273 (26); 274, 369 (1); 22681 (11); 23398, 23402 (35); 25045 (12); 25916 (Tb); 27727, 27738 (E); 27739 (14); 27773 (35); s.n. (11); s.n. (12). Kammerer, A. L. 48 (6). Kay, V. W. & J. E. Higgins 202 (23). Kearney, T. H. 5587A (11). Kearney, T. H. & R. H. Peebles 10052 (12); 11133 (6). Kellerman, W. A. 4501 4958 (26). Kenoyer, L. A. 5 (Tb); 427 (23) ; 2421 (36). Kenoyer, L. A. & Crum 2594 (1b); 4093 (9). Killip, E. P. & A. C. Smith 14169 (26). King, R. M. 1833 (24); 2718 (1); 2743 (26); (25) ; 2682, 2796, 2801, 2843 (33b); 2980, 2987 (1); (23); 3105 (1); 3948 (35); 3249 (1); 3322 (26); 3358 (1); 3359 (1); 3410 (35); 3417 (25); 3425 (26); 3454 (5); 3492 (33a); 3529 (5); 8560, 3600 (24); 3617 (34); 3634 (26); 3636, 3662 (4); 8678 (1); 3703, 3704, (18); 3707, 3709, 3711 (26); 3712, (17); 3833 (26); 2917 (26); 3042, 3096 3242 (25); (26); 3260 3329 (25); (26); 3364 3416 (9); (24); 3434 3461 (20); (34); 3539 (35); 3607 3626 (9); 3646 (23); (26); 3699 3705, 3706 3710 (17); 3715, 3716 3887 (23); 1972] 4138, 4164 (26); 4165 (23); 4168, 4178 (1); 4239 (26); 5090 (27); 5226 (26); 5348 (21); 5350, 5407 (35). King, R. M. & T. R. Soderstrom 4584 (9); 4646 (28); 4707 (35); 4804 (24); 4992 (1). Koeppen, R. & F. Iltis 618 (1). Kuntze, O. 23527 (24); 27465 (9). Lacás, M. M. 58 (7b). Lamb, F H. 361a, s.n. (17). Langlassé, E. 482 (28). Laughlin, R. M. 1050 (33b). Lay, G. T. & A. Collie s.n. (18). Leavenworth, W. C. 157a (26); 431 (37). Leavenworth, W. C. & H. Hoog- straal 1480, 1588 (5); 1589 t Leblond, J. B. 346 (E). Lehmann, F. C. 69, 1434 (26). Lemmon, J. G. 331 (12); 2777 (11); 2795 (12). Léon, Bro. M. Victorin & Bro. Alain 17344 (26). LeSueur, H. 341 (11); 1456 (6) ; 1466, 1472 (15). Liebmann, F. M. 232 (33a). Lindheimer, F. J. 949 (6). Llave, P. La s.n. (E). Loefling, P. “151” (E). Löfgren, A. & G. Edwall 16695 (25). Loomis, H. F. & C. J. King 2890 (12). Lundell, C. L. 6107 (27); 6759 (23). Lundell, C. L. & A. A. 7879 (26) ; 7972 (23); 9866 (7a). Lyonnet, E. 569 (35). MacDaniels, L. H. 41 (35); 53 (21); 723 (35); 747 (11). Melampodium — Stuessy 215 Macedo, A. 1408 (25). Maguire, B., B. L. Richards & T. Moeller 11849 (6). Manning, W. E. & M. S. 531212 (9). Marsh, E. G., Jr. 855 (6); 1176 (7b). Martinez, P. & A. Garcia 13 (7a). Matthews, R. W. & J. R. 480 (15). Matuda, E. 4444 [4504] (5); 6002 (33b); 26621 (26); 31141, 31314 (5). Maxon, W. R. 7694, 7696 (26). May, J. 13 (7b). McClatchie, A. J. s.n. (35). McGregor, C. 20181 (2). McVaugh, R. 7694 (7b); 7986 (6); 12822 (9); 13033 (33b); 13576, 14107 (35): 16112 (29); 18070 (37); 18906 (29). McVaugh, R. & W. N. Koelz 542 (29); 1054 (37). Mearns, E. A. 1997 (15). Mears, J. A. 1383, 1413 (6). Melchert, T. E. 169-C, 180 (6). Melchert, T. E. & P. Sorensen 6085 (3). Mendez, J. [^Menzies"] s.n. (9). Merrill, E. D. 10641 (2). Metcalfe, O. B. 66, 1271 (6). Mexia, Y. 240, 349 (5). Meyer, F. G. & D. J. Rogers 2828, 2911 (34). Millspaugh, C. F. s.n. (24). Mockford, J. P. & C. M. Rowell, Jr. 2769 (2). Molina, J. A. & F. A. Barkley 19Bo150 (26). Molina R., A. 1082 (26); 5899, 13346 (5). Moore, H. E., Jr. 2283 (20). Moore, J. A. & J. A. Steyermark 3115 (6). 216 Morales, R., H. 1221 (26); 1300 (9). Moreno, E. M. MS-198 (15). Muenscher, W. C. 12467 (25). Muller, C. H. 3040 (8); 3434 (9); 3659 (15). Munz, P. A. 13303 (7a). Nagel, O. 8030 (24). Née, L. s.n. (2). Nelson, E. W. 655 (33a); 715 (35); 740 (22); 870 (24); 947, 2339, 2740 (1); 2949 (5); 6447 (6); 6939 (1). Nicolas, Bro. 5447 (21); €150 (35); s.n. (21). Niles, W. E. 572 (6). Oersted, A. S. 9010 (27). Orcutt, C. R. 1395 (8); 3548 (35); 3605 (9); 3780 (32); 3887 (9); 3892 (35); 4365 (23); 4386 (20); 4516 (1); 6601 (29). Orozco, J. M. 260 (25); (27); 418 (25). Ortega, J. G. 4008, 4446, 6125, 7004 (17). 294 Padilla, S. A. 1, 242, 586 (5). Painter, J. T., W. D. Lucas & F. A. Barkley 14297 (7b). Palmer, E. 3 (2); 20 (14); 186 (26); 245 (15); 260 (9); 281 (2); 286 (35); 319 (23); 391 (4); 459 (1); 486 (11); 535 (1); 556 (7b); 726 in part (14), (15); 926 (9); 1163 (35); 1172 (1); 1457 (17); 1609 in part (5), (15) ; 1757a (17); 1814 (29): 2068 (8). Parish, S. B. & W. F. 1171 (35). Parker, K. F. 7707 (12). Parry, C. C. & E. Palmer 444 (21); 444% (11). Rhodora [Vol. 74 Paynet, A. M. & H. D. Hulan 1213 (23). Peebles, R. H. & G. J. Harrison 4680 (12). Peebles, R. H., G. J. Harrison & T. H. Kearney 4614 (11). Pennell, F. W. 1335 (27); sn. (18). Pereira, E. & G. Pabst 8505 [34972] (26). Pipes, D. 81 (23). Pittier, H. 57, 60, 429, 6963 (25). Pohl, J. E. 1368 [1276] (25). Porter, C. L. 4262 (6). Powell, A. M. & J. Edmondson 551 (21); 579 (9); 646 (23); 660 (9); 756 (26); 758 (2); 816 (35); 908 (17). Pratten, H. s.n. (E). Pringle, C. G. 10 (11); 55 (12); 297, 754 (11); 1019 (6); 1045 (11); 1767 (35); 2819 (26); 2821 (36); 3230 (31); 3639 (20); 3818 (33b); 3863 (36); 3923 (26); 4322 (24); 4537 (20); 4598 (4); 4666 (33a); 5257 (32); 5309 (9); 5722 (10); 6455 (21); 6491 (11); 6728 (9); 7321 (24); 7327 (32); 8466 (23); 8609 in part (9), (11); 8610 (32); 9008 (7b); 9162 (5); 9177 (23); 9301 (31); 9325 (21); 9331 (11); 9361 (33b); 9937 (23); 10065 (5); 11548 (11); 13068 (31); 13069 (24). Purpus, C. A. 416 (16); 1003, 1003a (8); 1809 (32); 2813, 2813’ (21); 2849 (35); 3822, 3822a (34); 4730 (8); 5619 (20); s.n. (8). Ramírez, J. 44 (Ta). Reese, J. A. 94 (12). Reko, B. P. 3812 (5). 1972] Reséndez, O. I. 52 (7b). Richard, L. C. M. s.n. (26). Ricksecker, A. E. *S" (26). Riddell, J. L. s.n. (6). Ríos, A. & H. Cavazos 231, 264 (Ta). Ripley, H. D. & R. C. Barneby 13708 (24). Robinson, C. B., Jr. 11835 (2). Rock, H. F. L. M-442 (9). Rodríguez, J. J. 72 (7b). Rodriguez, J. V. 1662 (25); 3341 (26); 3735 (25). Rojas 86 (9). Rose, J. N. 1568 (17); 3183 (4); 3271 (17); 3476 (18); 3617 (9); s.n. (17). Rose, J. N. & W. R. Fitch 17045 (6). Rose, J. N. & R. Hay 6244 (32); 6275 (9). Rose, J. N., J. H. Painter & J. S. Rose 9308 (23); 10137 (20). Rose, J. N., P. C. Standley & P. G. Russell 14307 (1). Rowlee, W. W. & H. E. 179 (5). Russell, N. H. 11511 (6). Russell, P. G. & M. J. Souviron 2 (15). Rzedowski, J. 9320 (33a) ; 17514, 17921, 21155 (37); 20748 (1); 21322 (33b); 26618 (18). Safford, W. E. 1264 (8). Sánchez, A. 105 (7a). Schaffner, J. G. 270 in part (9), (35); 271 (9) ; s.n. (14), (32). Schiede, C. J. W. 217 (24); 354 (E). Schiede, C. J. W. & F. Deppe 1254 (23). Schott, A. 72, s.n. (15). Schroeder, A. H. 120 (8). Seaton, H. E. 54 (35); 461 (24). Seemann, B. C. 46 (E). Melampodium — Stuessy 217 Sessé, M. “3977” (D); “3978” (24)3.5 489817 1982) 5: 39831, s8m(26)5 s.n: Gb): Sessé, M. & J. M. Mocino s.n. (9). Shank, P. J. & A. Molina R. 4236 (27). Sharp, A. J. 44116 (32); 44208 (21); 44961 (9). Shinners, L. H. 7183 (6); 7395 (7a); 12190 (6); 17659, 17708 (7a); 26390, 31286, 31559 (6). Shreve, F. 4965, 6003 (12) ; 6758 (15); 7685, 7709 (12); 7821 (17); 7918 (6); 9278 (35). Sinclair, A. s.n. (26). Skutch, A. F. 1036 (33b); 1550 (25); 1592 (9); 1709 (25); 2041 (26); 3968 (27). Small, J. K. & E. T. Wherry 12134 (6). Smith, A. 2922 (27). Smith, C. L. 351 (35); (23); 1655 (24). Smith, H. H. 514 in part (26). Smith, J. D. 2334 (25); 2382 (26). Solís, F. 52 (25); 56 (27). Solís, M. J. 124 (7a). Spruce, R. 3225 (E). 1605 Standley, P. C. 11226 (26); 12105, 15756, 17710 (25); 19722 (26); 20260 (25); 21648, 21789 (26); 22687, 22718, 22864, 25149 (25); 26708, 27175 (5); 28678 (25); 32757 (27); 36058, 38971 (25); 89021, 41581 (27); 56290, 56522 (25); 59769 (9); 60326, 60413 (25); 60493 (5); 63265, 643808, 64595, 68866 (25); 73774, 74071, 74125, 74813, 75620 (5); 76429, 77073 (25); 77149 (26); 77449, 218 78268, 79869 (25); 82849 (9); 87933 (25). Standley, P. C. & J. Chacón P. 6625, 6660 (25). Standley, P. C. & E. Padilla V. 2755, 3630 (25). Standley, P. C. & J. Valerio 48814 (27). Stanfield, Mrs. V. E. s.n. (6). Stanford, L. R., K. L. Rether- ford & R. D. Northcraft 186 (8); 1077 (1); 1078 (23). Stanford, L. R., L. A. Taylor & S. M. Lauber 2313 (26). Steere, W. C. 1805 (23). Stevens, F. L. 136 (25). Stevens, G. W. 325 (6). Stewart, R. M. 84 (6). Steyermark, J. A. 29041, 30509 (5); 31670 (26); 32847 (25); 43745 (26); 48234 (33b); 50392 (9); 50490 (33b); 50530 (9); 50539 (23); 50724 (5); 50773 (25); 50916 (20); 51452 (23). Stork, H. E. 2486 (27). Stork, H. E. & J. L. Morrison 8920 (26). Stuessy, T. F. 147, 167, 180, 230 (6); 274 (23); 282, 318 (26); 328 (20); 342 (26); 344 (20); 345 (26); 346, 347 (20); 348, 351 (26); 356 (23); 359 (26); 361 (5); 364 (9); 365 (26); 366, 369 (2); 374 (20); 377 (9); 379 (35); 383 (24); 393 (23); 397 (4); 401 (18); 4053 (26); 404 (17); 414 (6); 425, 429 (Ta); 450 (Tc); 463, 466 (23); 471 (1); 473 (26); 494, 497, 503, 505, 511 (35); 526, 527, 528 (26); 531 (23); 582 (26) ; 533 (23) ; 534, 535 (26); 537 (23); 543 (26); 544 (23); 545 (26); 546 (23); 547, 548, Rhodora [Vol. 74 550 (26); 559 (33b); 562 (23) ; 566, 567, 568, 570 (33b) ; 571 (1); 574 (23); 578 (25); 580 (35); 582, 585 (25); 586 (26); 587 (25); 588 (26) ; 589 (25); 592 (26); 594 (25); 602 (1); 603 (35); 605 (26); 607 (5); 608, 609, 611 (26); 613 (5); 614, 615, 616 (26); 619 (5); 620, 621, 623, 624, 625 (26); 627 (24); 632 (1); 634, 636, 637 (20); 638 (24); 639 (20); 640 (24); 642 (9); 645 (26); 647 (9); 650 (5); 654 (35); 659 (21); 660 (9); 663 (34); 667 (20); 668 (35); 670 (32) ; 675, 676 (31) ; 677 (26); 682, 684 (34); 688 (1); 690 (9); 693 (28); 694 (23); 695 (3); 697, 698 (5); 702 (26); 703 (9); 705 (35); 708, 709 (36); 713 (23); 714 (36); 715 (28) ; 720, 721 (37); 722 (23); 723 (26); 724 (1); 725 (26); 727 (1); 728 (26); 729 (35); 730 (9); 738 (4); 740 (9); 743 (26); 745 (18); 746 (26); 748, 750 (17); 751, 752 (6); 771 (7a); 778, 787 (Te); 826 (23); 829 (26); 854 (Ta); 855, 856 (Tb); 857, 868 (Ta); 902, 912 (7b): 1016, 1038, 1054 (11); 1122, 1127 (6); 1128 (11). Stuessy, T. F. & W. Renold 1261 (7c); 1284 (7a). Sullivan, W. & B. L. Turner 34 (6). Swartz, O. s.n. (E). Taylor, M. 115 (21). Tetters, S. 18 (6). Tharp, B. C. & M. C. Johnston 541 [802A], 541922 (7a). Tharp, B. C. & Miller 51-312 (6). 1972] Thompson, B. & B. Fields 332 (2). Thompson, B. & A. Graham 18 (6). Thompson, E. s.n. (1). Thornber, J. J. 7404, 8121, 9050 (12). Thurber, G. 937 (12). Tonduz, A. 10144 (25); 13615 (26). Tonduz, A. & Rojas 203 (25). Torres R., R. 50 (25). Townsend, C. H. T. & C. M. Barber 351 (11); 376 (15). Tracy, S. M. & F. S. Earle 23 (6). Triana, J. J. 1331 (25). Tucker. J. M. 455 (5); 585 (26); 1313 (25); 2565 (9). Tuerckheim, H. von 114 (25); 124 (26); 761 (25); 3955 (9); 8417 (25). Turner, B. L. 4490 (7a); 4620 (11); 4790 (6); 5006 (Ta); 5632, 5673, 5735 (6). Turner, B. L. & T. E. Melchert 4840 (6). Turner, B. L. & A. M. Powell 1116 (11). Turner, C. 7 (6). Turner, R. M., R. A. Dodge & C. T. Mason, Jr. 2068 (12). Turner, R. M. & C. H. Lowe, Jr. 2074 (15). Ule, E. 2978 [397] (E). Velasco, L. V. 8946 (25). Volkem 2564, 2774 (25); 2973 (1); 3013 (26). Volkens, G. L. A. 1134 (35). Walker, M. L. 93 (Tc). Wallis, C. 4807 (6). Warnock, B. H. 9229, (11); 46217 (6). Warscewicz, J. von s.n. (26). Waterfall, U. T. 3781, 4818 (6); 13507 Melampodium — Stuessy 219 15805 (7b); 16198 (31); 16342 (9). Waterfall, U. T. & C. S. Wallis 13702 (31); 13886 (9); 13960 (11); 14288 (23). Webster, M. B. 140 (33b). White, O. E. 100 (26). White, S. S. 2642 in part (9), (11); 2784 (15); 2788 (11); 3516, 3908, 4153 (15); 4587 (6). Whitehouse, E. 9841 (6). Wiegand, M. C. & G. B. Upton 4439 (6). Wiggins, I. L. 7021, 7388, 7444, 7533 (15); 11507 (14); 11765 (6); 14377 (16). Wiggins, I. L. & R. C. Rollins 281 (14); 433 (15). Wilbur, R. L. & C. R. 1395 (1); 2416 (29). Wilcox, T. E. 335, s.n. (12). Williams, L. O. 5643, 18257 (25). Williams, L. O. & A. Molina R. 13398 (26). Wilson, P. 9553 (35). Winbery, J. E. & C. M. Rowell, Jr. 2492 (2). Woodson, R. E., Jr., P. H. Allen & R. J. Seibert 1370 (26). Wooton, E. O. 117 (6) ; s.n. (7c). Worth, C. R. & J. L. Morrison 8807 (17). Wright, C. 311 (6); 1205 (12). Wright, J. T. 42-54 (6). Wright, W. G. 1213 (17). Wynd, F. L. & C. H. Muller 132 (8); 214 (7b). Xantus, L. J. s.n. (1). York, C. L. & G. 54037 (6). Young, M. S. s.n. (11). Yuncker, T. G., R. F. Dawson & H. R. Youse 5519 (25). Zamora, F., J. B. Paxon & F. A. Barkley 16M905 (11). SPORES, CHROMOSOMES AND RELATIONS OF THE FERN PELLAEA ATROPURPUREA ALICE F. TRYON The Purple Cliff-Brake, Pellaea atropurpurea, is the only American species of the large complex of cheilanthoid ferns that has a range extending from the tropics into northern regions, including New England. Only triploid plants are known, which evidently are hybrids, and they are apog- amous with 87 chromosomes in the sporophyte and spore mother cells. The species is a tool of some biological interest as there appears to be no evidence of sexual reproduction, although the specimens are highly variable throughout the geographic range. Northern Mexico and the adjacent southwestern United States is regarded as a center of dis- persal of Pellaea section Pellaea in America, for eleven of the fourteen species have distributions that converge there or have close relatives that do. Studies of the cytology and spores of P. atropurpurea and two other species, P. notabilis and P. ternifolia, that occur in this central area, provide new data for establishing relationships among these species, and insights on the possible origin of P. atropurpurea. MATERIAL AND METHODS The general geographic distributions of the taxa are outlined on the maps; localities more than 600 miles from the main range are indicated by separate dots. Ranges are based on data from my revision of Pellaea section Pellaea (A. Tryon, 1957) and from recent reports of range exten- sions. The wide cytological sampling of populations is shown on the maps. The new cytological] records are com- bined with earlier ones in Table 1. Meiosis was studied in sporangia fixed in 3:1 absolute ethyl alcohol: glacial acetic acid. Mitosis was examined in root tips fixed in the same solution following pretreatment for three hours with saturated aqueous solution of paradichlorobenzine. Scanning electron microscope photographs were obtained from spores undercoated with carbon prior to coating with an alloy of palladium-gold while rotating and tilting the 220 221 Pellaea atropurpurea — Tryon 1972] « AM) SƏNYM ‘WN :vsa (on) ‘us upeqoQ Xp 8c + ‘OD IJepuexo[y “D'N :vsa (CHOIN) 900F9 42946044 Xp 86 « “SPU €eumeje) "es “ZY :vsa (OSI) FZ9T 420]qgowu Xg »norur,g DUMGY DA M “ABA « “SPU €umejep "es “ZUY :vsa (OSW) PZ9T |20]qg0ouqy — X& »noru-/g myjojutto? DINIT ugeg eH “qong :oorxo]py (AXIN) GIT DRY XF 8€ (8T PT "si oj[ongry ues “d "['S :09IX9]N (115) 838S Uot, P SUMOY XF 86 x OAUIg[ 9p AJLA “Xa :02IXO]N (OW) GOTS Uou, Y uoa, X? 8G ugeg H “qond :091Xo]y (u5) SOF DQ XG 66 (LT ‘ST "erp oPpONSI ues *q "T'S :ootxo]q (11D) 61c8€ Uou, P swo XZ 62 D10jftw42] “ABA x BOBABILIONY “IOW :02IX9]A (OW) COTS Uot, 2p wofi.t XZ 62 pio[rua429] DINYA T OdsTOUBIY UBS 7D) “I'N OION (HD) eegp wmbossnpyy XZ 62 (ST ‘SE "sup OOSIOUBLY UBS ‘O “I'N :09IX9TA (HD) E961 20]qouxM XZ 62 811100301 020]]24d vsopy onburdryD “TI'N :091X9]q (HD) 098€ wofis] P suyo Xg onorour-jg (TT SIA) esə w onburdrq;) “TI'N :001X9]A (H5) 098S uou, 3p sumoy Xg —onoru-/g (9T ‘6 'sSup) « Avg s,[[eqdum;) “ond :epeuv;) (vod) OG6T Ul 2332uo]y Xg onorourjg x SduridsjoH 3unouLi(e, “o'g :eprueg (011) TOGT ut *4o]fin I, Xg DJOTIWU- Lg x Huung Aui “OW :vsn (OW) &ZGG woh, 3» UohaT XE 2O9nomu-/g mnoandando4jn vanyaq Aq] Boor] 10129[[02) Apold Ioquin N sənədg (1x03 eos “Grodad snoraəad,) vavTisd AO SAAIWAN AWOSOWONHOD '[ ATA 222 Rhodora [Vol. 74 sample. The original magnifications of the photographs of whole spores were taken at 1000 X and portions in greater detail at 5000 X. Spore measurements are of the greatest diameter of spores in lactic acid, including the outer perine layers. Range and mean measurements are based on samples of 100 spores — 25 from each of four collections — except for Pellaea notabilis, in which 50 spores were measured from each of two collections. Measurements of the pinnae are of the longest fertile pinnae in the central portion of the lamina. Data are drawn from the revisionary study cited and additional specimens in collections of the Gray Herbarium, the Field Museum of Natural History, and the Smithsonian Institution. ACKNOWLEDGMENTS I am most appreciative of the help of Drs. Donald M. Britton and Rolla Tryon for reading and helpful comments on the manuscript. I am also grateful to Dr. Robert H. Mohlenbrock for the use of the Cambridge Stereoscan scanning electron microscope, at the University of Southern Illinois, Carbondale and the assistance there of Judith Murphy with Figs. 31-33. Other spores were examined with an AMR instrument model 900, Harvard University; the help of G. R. Pierce in the McKay Laboratory is much appreciated. Drawings of the habit and leaf detail were done mostly by Sarah Babb Landry but also by Mary Robbins. GEOGRAPHY AND ECOLOGY The leaves of Pellaea atropurpurea (L.) Link are di- morphic, and several specimens from Mexico have leaves with mostly simple pinnae as in Fig. 1. The species ranges from Guatemala northward along the Sierra Oriental of Mexico, northeast across the United States to Vermont, southern Quebec and Ontario, and there are disjunct records from areas in southern British Columbia and Alberta and also north of Lake Athabaska, Saskatchewan (Fig. 3). It occurs on exposed or shaded ledges and in creviees of limestone cliffs, on talus slopes, masonry walls, 1972] Pellaea atropurpurea — Tryon 223 Figs. 1, 2. Habit of Pellaea: Fig. 1. P. atropurpurea with leaves of most simple form, Puebla, Arséne 3548 (GH); Fig. 2. P. notabilis with dimorphic leaves, Nuevo Leon, Knobloch 1963, (GH), a. Young plant, b, Fertile leaves on mature plant; both from spores. All X !4. 224 Rhodora [Vol. 74 or in rocky loam in woods, at 300-5200 meters. At its southernmost station in Guatemala it grows in crevices of ancient Indian pyramids. At Campbell’s Bay, in Quebec, near Ottawa, it occurs on precambrian rocks in which lime- stone is incorporated. In Ontario it is reported by Rigby (1968) to occur in sheltered locations on talus or limestone paving, rather than exposed cliffs. Leaves of mature plants of Pellaea notabilis Maxon are usually dimorphic and the pinnae simple. The first leaves are entire and cordate, and the older ones rarely have a compound lamina as in Fig. 2. Collections are known from the vicinity of Victoria, Tamaulipas, at 320 m. and from Nuevo Leon (Fig. 3). In the latter state it grows on cal- careous rocks at 1100 m. in San Francisco canyon, south of Monterrey, and P. atropurpurea has also been collected there. Plants of P. notabilis, grown from spores of two collections from this canyon, produced fertile leaves in six months. The rapid and vigorous growth of these plants in culture suggests that scme external factors are responsible for the limited distribution of the species. Peliaea ternifolia (Cav.) Link car. ternifolia is the widest ranging of all the American pellaeas occurring from south- ern Texas to Argentina and Hawaii, at 200-4000 m. Both diploids and tetraploids (Figs. 5, 6) are known in the variety, and beth occur at the same localities in the states of Durango 2nd Michoacán as well as the stations reported here. Ecological data from these collections as well as field observations show that the tetraploids grow in mcre shaded, wetter sites than the diploids. The localities for the three cytologically documented plants are shown on the map of the general range of the variety in continental North America (Fig. 7). In addition to these tetraploids other specimens which are morphologically similar, including those reported by Pray (1967) from Mexico and Texas, are included on the map. The leaves of Pellaea ternifolia vav. Wrightiana (Hook.) A. Tryon (Fig. 4) most closely resemble the form of the diploids in var. ternifolia, and variation in these is noted Fig. 3. Pellaea atropurpurea and P. notalilis: P. atropurpurea outlined dots-disjunct stations, circles-cytological records; P. notabilis stars-the upper a cytological record. 226 Rhodora [Vol. 74 Figs. 4-6. Habit of Pellaea ternifolia: Fig. 4. var. Wrightiana, Burro mts, New Mexico, Rusby in 1880 (GH); Figs. 5, 6. var. terni- folia: Fig. 5. Diploid, Puebla, Riba 408 (GH). Fig. 6. Tetraploid from culture, San Luis Potosí, Rollins & Tryon 58218 (GH). All x lé. 1972] Pellaea atropurpurea — Tryon 227 7 Figs. 7, 8. Range of Pellaea ternifolia: Fig. 7. var. ternifolia outlined, dot-disjunct station, stars and circles-tetraploids, circles- cytological records, half circles-localities of both diploids and tetra- ploids; Fig. 8. var. Wrightiana, circles-tetraploids, star-triploid, 228 Rhodcra [Vol. 74 in the section under morphology. The geographic range of var. Wrightiana extends from southern Oklahoma south- west to Arizona and across the Mexican border in northern Chihuahua, Coahuila and Baja California, at 1400-2300 m. (Fig. 8). A disjunct station is also reported about 1000 miles eastward in Alexander Co. in western North Caro- lina. This latter record may indicate a formerly more continuous distribution of these plants across the southern United States similar to the present range of P. atropur- purea, or possibly long distance dispersal from populations farther west. Specimens are mostly from exposed situa- tions, among sandstone or granitic rocks or rarely on lime- stone. A few collections of somewhat larger specimens are from shade, in moist forested sites. CYTOLOGY Mitosis and meiosis were examined in plants of Pellaea atropurpurea from Chipinque Mesa, south of Monterrey, Mexico. In the mitotic cells many chromosomes have sub- terminal centromeres as shown in Fig. 11. Differences in the position of centromeres, in chromosome size, and the occurrence of satellites, are diagnostic aspects useful in karyotype analyses. In these apogamous plants most spor- angia have eight spore-mother-cells, and some sixteen-celled sporangia are also formed. In meiotic nuclei, the nucleoli are conspieuous and persist through diakensis. Their struc- ture can be distinguished from the bivalents in Fig. 9 by the circular form and lighter stain. In the eight-celled sporangia the chromosomes are usually associated in bi- valents (Figs. 9, 16), and this is also reported by Manton (1950) and Rigby (1968). In apogamous plants the chromosome number in sporangia with eight spore-mother- cells, according to Manton, undergoes premeiotic doubling to accommodate reduction. In sixteen-celled sporangia, at meiosis, Manton (1950) and Britton (in Tryon & Britton, 1958) report approximately 29 pairs and 29 single chromo- somes. The pairing associations in several] cells from both types of sporangia were studied by Rigby. In these, two cells from sixteen-celled sporangia (a specimen from Que- a Figs. 9-14. Meiotic and mitotic nuclei: Figs. 9-11. Pellaea atropurpurea: Fig. 9. Nucleus from 8 celled sporangium with 87 bivalents, the nucleolus at upper right, X 660; Fig. 10. Early stage in sporogenesis, micronucleus and 4 nuclei at center, X 600; Fig. 11. Mitotic chromosomes from young sporangium. All triploid, n=87, Rollins & Tryon 5860 (GH). Fig. 12. Meiosis, P. notabilis, diploid, n—29 Marroquin 1855 (GH) 660. Figs. 13, 14. Meiosis, P. ternifolia var. ternifolia: Fig. 13. Diploid P. ternifolia var. ternifolia; Fig. 13. Diploid, »—29, Riba 408 (GH) X 1000; Fig. 14. Tetraploid, »—58, Riba 412 (MEXU) X 1000. 230 Rhodora [Vol. 74 bec) had equal numbers of pairs and univalents, but one had nine and the other six trivalents. A second collection (from British Columbia) had 31 bivalents, 22 univalents and one trivalent. Affinities between the parental genomes are expressed in these cells by the associations of more than 29 bivalents and also the trivalent formations. The varia- tions in chromosome associations in these cells and also the lack of multivalents in the eight-celled sporangia, which have doubled the chromosome number, raise problems in the interpretation of chromosome pairing in these plants. There are frequent irregularities as shown in the early stage of sporogenesis in Fig. 10, in the sixteen celled sporangia, with an extra micronucleus. Two collections of Pellaea notabilis from Nuevo Leon uniformly had 29 bivalents at meiosis as in Figs. 12, and 15. This establishes the species as a diploid with the basic chromosome number for Pel/laea. The number is consistent with the species being proposed as one of the elements involved in the origin of the triploid P. atropurpurea. Both diploid and tetraploid plants of Pellaea ternifolia var. ternifolia were studied from the same localities near San Luis Potosi and also farther south near Puebla. There are earlier records of these two polyploid levels, reported by Britton, from localities near Mexico City but not at the same site. The diploid with 29 bivalents and the tetraploid with 58, in Figs. 17, 18 are collections from Puebla. Con- trast in the number of bivalents at late metaphase in these two polyploid levels is readily apparent in the figures. The terminal portions of the chromosomes in the centromeric regions in Figs. 12, 14 are stretched thinner than the re- mainder of the chromosomes forming the bivalents. These attenuated portions show tension on the late metaphase chromosomes as they prepare to move to the poles. The tetraploid plants generally resemble the diploids, but the pinnae tend to be entire and the rachises more pubescent (Figs. 5, 6) as compared to the diploids, which have ternate pinnae and nearly glabrous leaves. The drawing of the tetraploid in Fig. 6 represents an especially robust plant 1972] Pellaea atropurpurea — Tryon 231 "AE Sr te at? 9v ot de : ; = 9 ade an Ge = \ n - D. i 4 ag} Y E wt daa Kg &1? ally oie & “ipo IS ( l6 $ tom E J» AS » a "2s at T opets’ ‘os ie * oe 94 sty "v, > etw 2 17 e v Ig Figs. 15-18. Explanatory diagrams of meiotic nuclei (Figs. 9-14) showing bivalents in focus in black, others in outline, nucleolus stip- pled, from the same cells except P. atropurpurea: Fig. 15. P. nota- bilis, Fig. 16. P. atropurpurea, n=87, Rollins & Tryon 5860 (GH); Figs. 17, 18. P. ternifolia var. ternifolia with thin portions omitted: Fig. 17. Diploid, Fig. 18. Tetraploid. 232 Rhodora [Vol. 74 that was grown in culture for several months. At Hacienda Batán, near Puebla, the tetraploids are rare, occurring in moist, shaded places while the diploids grow in exposed, rocky crevices. Other specimens, which appear to be tetra- ploids on the basis of the characters noted above, are quite widely distributed as is shown by the stars on the map in Fig. 7. The cytological records of Pellaea ternifolia var. Wright- iana are based on three earlier reports from localities in the United States. Tetraploids with » — 58 were noted from the western part of the range in New Mexico by Lloyd (1968) and from the disjunct station in North Carolina by Wagner (1965). These plants were considered to be amphiploid hybrids of the diploids P. ternifolia var. terni- folia and P. truncata Goodd. (P. longimucronata Hook. ). A triploid plant of var. Wrightiana with 29 bivalents and 29 univalents from Arizona was reported by Knobloch & Britton (1963). This plant was considered to have origi- nated as a hybrid between the diploid P. truncata and a tetraploid plant of var. ternifolia. Knobloch & Britton report the same chromosome number in both mitotic and meiotic cells and also 64 irrgular spores in the sporangia. A specimen (Pringle, May 6) from the Santa Catalina mountains, which also appears to be a variant as noted in the survey of the spores, indicates the need for further cytological sampling of var. Wrightiana. SPORES In Pellaea atropurpurea only spores of the 32-spored sporangia mature, and this reduced number reflects the apogamous type of reproduction. The spores are globose or somewhat ellipsoidal (Fig. 19) and unusually large, ranging from 54-92 „ in diameter. The triradiate scar scarcely projects, and is apparent only in cleared material, with arms extending 34, or more of the radius. The perine ridges are similar to those in P. notabilis but are sharper and more prominent. At higher magnification the perine surface consists of compacted sporopollenin particles (Fig. 20). 1972] Pellaea atropurpurea — Tryon 233 Figs. 19-22. Spores cf Pellaea: Figs. 19, 20. P. atropurpurea: Fig. 19. Nuevo Leon, Rollins & Tryon 5860 (GH), X 660; Fig. 20. Perine surface, Hac. Batán, Arsène 3548 (GH), X 3300. Figs. 21:22. P. notabilis: Fig. 21. Knobloch 1963 (GH), X 660; Fig. 22. Perine surface, X 3300. 234 Rhodora [Vol. 74 In Pellaea notabilis sporangia have 64 well formed spores that are globose or somewhat ellipsoidal (Fig. 21) and range from 40-62 , in diameter. The slightly larger spore size in this diploid, relative to diploid spores in P. terni- folia var. ternifolia, may be partly due to the inclusion of the prominent ridges in the measurements. These dense ridges obscure the triradiate scar, but in cleared material it projects slightly and the arms extend ?4 of the spore radius. At higher magnification the perine deposition ap- pears uniformly verrucate on both floor and ridge surface (Fig. 22). In both diploid and tetraploid forms of Pellaea ternifolia the sporangia uniformly have 64 spores. Those of the tetraploid are consistently larger, ranging from 46-62 , as compared to the diploids which range from 38-58 ,. The size differences are especially apparent in Figs. 23 and 25 and have been reported also in other collections by Pray (1968). Spore shape is generally spherical or tetrahedral- globose especially in the tetraploid which has broader sur- faces between the arms of the triradiate scar. These arms extend 34, or more of the spore radius. The perine is generally similar in spores of both polyploid levels, but variation occurs in density and prominence of the ridges. These may be especially prominent in the diploid spores (Fig. 24), and on the smoother surface of the tetraploid spores (Fig. 26) there are often numerous sporopollenin particles. Spores of Pellaea ternifolia var. Wrightiana were ex- amined from populations throughout its range. Sporangia usually have 64 spores of relatively uniform size. A speci- men (Pringle, May 6, 1883, GH) collected in the Santa Catalina mountains in Arizona had many irregular spores and some small ones as shown in Fig. 34. This specimen may represent another triploid, similar to that reported from the same area by Knobloch and Britton (1963). The spores of var. Wrightiana are usually globose with a promi- nent triradiate scar extending 9/4 or more of the spore radius. The form of the perine is relatively uniform as Figs. 23-28. Spores of Pellaea ternifolia: Figs. 23-26. var. terni- folia: Fig. 23. Diploid, X 660; Fig. 24. Perine surface, X 3300, both San Luis Potosí, Rollins & Tryon 58219 (Gu); Fig. 25. Tetraploid, X 660; Fig. 26. Perine surface with portion of triradiate scar, X 3300, both Rollins & Tryon 58218 (GH); Figs. 27, 28. var. Wright- iana: Fig. 27. Gillispie Co., Texas, Correll & Correll 12762 (Gu), X 660; Fig. 28. Santa Catalina mts. Arizona, Phillips & Reynolds 2943 (GH), X 660. 236 Rhodora [Vol. 74 Figs. 29-34. Spores of Pellaea ternifolia var. Wrightiana: Fig. 29. Lower spore with portions of perine eroded, smooth exine surface at right, X 660; Fig. 30. Perine surface with portion of triradiate scar, X 3300, both, Coahuila, Johnston & Miller 1509 (GH); Figs. 31- 33. Alexander Co., North Carolina, Bozeman et al. 45152 (stu): 9 Fig. 31. X 660, Fig. 32. Perine surface, X 3300, Fig. 33. Spore with partly eroded perine and subtending scabrous layer, X 660, Fig. 34. Santa Catalina mts. Arizona Pringle, May 6, 1883 (GH), X 660. 1972] Pellaea atropurpurea — Tryon 237 shown in the series of specimens from Mexico, Arizona, Texas, and North Carolina (Figs. 27-34), although there is some variation in size and prominence of the perine. The specimens from North Carolina have an especially rugose perine with stratification as shown in Fig. 33, showing a coarse outer rugose layer subtended by a strongly scabrous one. Similar stratification is also evident in the specimen from Mexico in Fig. 29, which has some scabrous deposition on the smooth surface of the spore exine where the perine has been abraded. The differences in prominence of the triradiate scars in these species may imply distinctions in spore development. The prominent triradiate scars with scant perine deposition characteristic of the spores of P. ternifolia indicate that these may have stronger tetrad associations. In contrast to these, the obscure triradiate scars in P. notabilis and P. atropurpurea suggest the tetrads may be disassociated before completion of the perine deposition. MORPHOLOGY Comparisons of the morphology of the species and varieties are made in Table 2. Similarities between Pellaea notabilis and P. atropurpurea (Figs. 1, 2) are especially evident in the dimorphic leaves, terete shape of the petiole, longer pinnae, and concolorous rhizome scales. These two species are readily distinguished from P. ternifolia by these characters. There are also marked similarities among the three kinds of P. ternifolia (Figs. 4-6) in the petiole shape, monomorphic leaves, and bicolorous rhizome scales. The form of the perine, in addition to the characters shown in the chart, also establishes a close relationship among them. These similarities express relationships that, by compari- son to other species of Pellaea, can appropriately be recog- nized by including them in a single species. Morphological variation in collections of Pellaea ternifolia var. Wrightiana was surveyed over the whole geographic range. Most of the specimens have leaves most closely resembling the form of the diploid var. ternifolia (Figs. 4,5). Some variation is shown in the division of the leaves, { Vol. 74 Rhodcra 238 edrrjs orjoadoe[os [?-13u929 peo.q ‘ugg 'sno.o[oorq (ve) " G9-9y xod? je Ajer -9odso oqvuv[ 929]J.Ins [etxepe uo ouv[d snaand.indo.3e &I "un (PLT ayeuuld T 9je[ooou?| Ájmoa -IU 10 .I&9UI[ orgdaourouour edrrjs orjodo9[os [?.13u99 AAO.LLEU ‘ugg 'sno.to[ooIq (97) " 86-86 juaosoqnd Á[os -ieds 10 sno.qe[3 ooej.Ins [eixepe uo oue[d sno1ind.indo.t3e 10 SNIUVJSLI T 0} ‘g Asou “Wo op-c0 ayeuuld T 9j?[ooou?[ ATMO -IVU 10 TRaUT] on[d.ourououtr odrijs orj30.19[os [€.11u99 peo.q ‘AISI ‘SNOLO]OIIG (6F) 19G-P§ j1uoosoqnd Á[os -ieds 10 snoiqey.s 9oPj.ins ]?rxepe uo ouv[d snou?jseo ITE "up C po ayeuuld g qem -3UVLII} ÁjmoIreu orn[d.rourouour Ajsni 10 uj *sno.to[0ouoo (9) n G6-Y6 prds *qjuooseqnd reond 10 939193 SNIULJSLI ÁJƏILI *sno1ndando.t3e SrT "uo &rT o9jeuutd e-T xen -3UV11}-947¥ 3U0JƏ rydrowp ÁAjsni 10 uey ‘Sn0.10[00U09 (0€) 7 69-0F j1uoosoqnd AVY STS 949.19} aseq 3? sneind -Indore ‘ueg "Wd L-g əyeuurd T 378 A0-97e.3 -U0ĮƏ 10 proq[op ərydrowrp XP vuofiuao) ‘apa VAVTIJd AO SHIOddS AAUHL AO SNOSIdIVdWOO 'IVOIDO'IOHd:IOWN XZ DYOfimsaz ‘una D0 fruta} DUDIZY HUA ‘Da paindindosyD $1110020u s3]D28s IWOZIYY (uvour) -ƏZIS ILOS juoumpur ədeys 10109 3101324 Jaquinu juourdos [?.13u92 Y4sus] vud UOISIAID adeys nwuiwmn' soano' '6 M'IdV.L 1972] Pellaea atropurpurea — Tryon 239 especially on the same plant, and this appears to be corre- lated with leaf size. In these specimens the leaves with shorter pinnae rachises and nearly ternate pinnae, re- sembling those of var. ternifolia, are usually smaller and seem to be produced before the more highly divided leaves resembling those of P. truncata. A similar correlation between leaf size and division is also evident in the plant from North Carolina figured by Wagner (1965). Thus, variation in leaf form in plants of var. Wrightiana shows types intermediate between var. ternifolia and P. truncata and also differences in developmental sequence. DISCUSSION The report of 29 bivalents in Pellaea notabilis establishes this as one of the basic species among the American pellaeas. Similarities of the spores and other morphological characters indicate that it is probably one of the elements involved in the origin of the widely distributed triploid, P. atropurpurea. On the basis of this new information several possibilities may be considered for the origin of P. atropurpurea. The strong morphological resemblances between the two species suggest that Pellaea atropurpurea may be an auto- ploid of P. notabilis. The triploid could have originated from two gametes of P. notabilis, one of them unreduced. The studies of P. atropurpurea by Rigby show that affinities do exist in the two parental genomes. However, several unique characters of Pellaea atropur- purea must also be considered. The broad leaves up to three times pinnate with densely pubescent rachises sug- gest that a second distinctive parent is involved in addition to P. notabilis. The cytological and morphological survey of P. ternifolia, the most likely parent in northern Mexico, shows that none of its variants completely fills the morpho- logical requirements of the second parent. The spores of P. ternifolia are especially diagnostic, and the perine form is quite different from that of P. atropurpurea. The unique morphological features of Pellaea atropurpurea are in- consistent with an autoploid origin from P. notabilis, and 240 Rhodora [Vol. 74 the characters of its closest relatives near the present range of that species are inconsistent with an amphiploid origin involving one of them. The most reasonable conclusion at this time is that either the second parental element of P. atropurpurea belongs to another species group possibly not within the geographic center, or it may be extinct. The rare occurrence of P. notabilis, its closest relative in north- ern Mexico, suggests that the other parent may no longer be extant. The extensive distribution of P. atropurpurea is un- doubtedly partly based on the absence of syngamy. Elimi- nation of the conditions required for fertilization results in more broadly adapted apogamous gametophytes. The large spores may also provide additional reserves, increased survival capacity, and a greater energy source for more rapid development of apogamous gametophytes and spore- lings. In Pellaea ovata (Desv.) Weath., which has both a sexual (2x) and an apogamous (32) race, the range of the latter is over 4000 miles greater than that of the sexual race. The effectiveness of the apogamous reproductive system is shown by the wider geographic distribution of the apogamous race in that species as well as in P. atro- purpurea. GRAY HERBARIUM, HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138. 1972] Pellaea atropurpurea — Tryon 241 LITERATURE CITED KNOBLOCH, I. W. & D. M. BRITTON. 1963. The chromosome number and possible ancestry of Pellaea Wrightiana. Amer. Journ. Bot. 50: 52-55. LLovp, R. M. 1966. I.O. P. B. Chromosome number reports. Taxon 15: 283. MANTON, I. 1950. Problems of Cytology and Evolution in the Pteridophyta. Cambridge University Press. Pray, T. R. 1967. Notes on the distribution of some American cheilanthoid ferns. Amer. Fern Jour. 57: 52-58. RIGBY, S. J. 1968. An investigation of Pellaea glabella and Pellaea atropurpurea and their relationships. University of Guelph, Guelph, Ontario. M. S. thesis, 120 pp. Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Missouri Bot. Gard. 44: 125-193. TRYON, A. F. & D. B. BRITTON. 1968. Cytotaxonomic studies on the fern genus Pellaea. Evolution 12: 137-145. WAGNER, W. H. 1965. Pellaea Wrightiana in North Carolina and the question of its origin. Jour. Elisha Mitchell Sci. Soc. 81: 95-103. THE GENUS CORIARIA (CORIARIACEAE) IN THE WESTERN HEMISPHERE’ LAURENCE E. SKOG With the recent awakening of research in hallucinogenic plants, some attention has been given to the genus Coriaria. Plants of this genus are known to be toxic to animals when taken internally, as well as being hallucinatory in small quantities. Studies of its toxicity and the application of the toxic elements in treatments of barbituate poisoning have made necessary a basic taxonomic study of Coriaria. This paper deals with taxonomic studies of the plants which occur in the Western Hemisphere. Coriaria inhabits warm temperate regions of southern South America, and, in the tropical areas, high elevations of the northern Andes, ranging north through Central America to the Sierra Madre Occidental in Mexico. Coriaria is known also from southern Europe, North Africa, the Himalayas, islands of the Pacific coast of Asia and the western Pacific Ocean, in- cluding New Zealand. The genus makes an interesting study of geographical distribution and dispersal which has resulted in highly disjunct populations. The plants of the Western Hemisphere have close relatives on the other side of the Pacific Ocean, but these are excluded from considera- tion here. PHYLOGENY OF THE GENUS As the only genus in the family Coriariaceae, Coriaria has no close relatives and no close morphologic affiliation with any other family. In 1816 De Candolle placed Coriaria in the Rhamnaceae. Since that time the position of Coriaria in a phylogenetic system has changed often with various authors, ranging from alliances with the Urticaceae (Max- imowicz, 1881) to a much more advanced position near the Araliaceae (Russow, 1884). Most often the genus has been ‘Based on a thesis submitted to the Graduate School of the Uni- versity of Connecticut in partial fulfillment of the requirements for the degree of Master of Science. 242 1972] Coriaria — Skog 243 placed near the Anacardiaceae as was done by Bentham and Hooker in 1862. Hutchinson in 1964 placed Coriaria in its own order, Coriariales, between the Dilleniales and the Rosales, and derived from the former. Cronquist (1968), with a note of doubt, has placed the Coriariaceae in the Ranunculales. The phylogenetic relationships of the Coriariaceae can- not be determined accurately without, perhaps, the dis- covery of some “pre-Coriaria” which would link the family to another group. It is apparently not closely related to any single other family; it has long been distinguished and kept separate from other families and probably should not be closely associated with others in the same order. There is, however, no single character of the family that sets it apart, but a combination of traits. If the dicots are viewed as polyphyletic, Coriaria could be one of the distinct lines which branched from the main line of evolution some- time in the Cretaceous, retaining many primitive characters while evolving other more advanced or specialized traits. If one lists the characteristics of dicotyledonous families and then attempts to place a family among them, correla- tions become evident which can sometimes be interpreted as showing evolutionary relationships; with Coriaria this has not been successful. Various characters of dicots are considered as being either primitive or advanced. These often determine the position of the family in a phylogenetic order. Maximowicz (1881) attempted to draw conclusions about relationships by using morphological characters of various families, then weighting them as to their import- ance in Coriaria. After totaling the values, Maximowicz concluded the genus was most closely allied to the Sima- roubaceae, and also had affinities with the Phytolaccaceae. If one had to select another family in which Coriaria could be placed, one might choose the Rosaceae. The characters of Coriaria fall within the broad limits of this family, as defined or delimited by Lawrence (1951) and Hutchinson (1964). Within the Rosaceae, though, there is no single genus to which Coriaria is closely related. The genus fits 244 Rhodora [Vol. 74 less well in the Simaroubaceae or Erythroxylaceae, and only superficially in the Brunelliaceae. Paralleling this viewpoint, Hutchinson has placed Coriaria in a distinct order between the Brunelliaceae and Rosaceae, and Hallier (1905) placed the Coriariaceae in the Rosales, but allied to the Simaroubaceae. TAXONOMY Coriaria L., Genera Plantarum, ed. 5. 459. 1754. Type: Coriaria myrtifolia L. Heterocladus Turez., Bull. Soc. Imp. Naturalistes Moscou 20: 152. 1847. Type: H. caracasanus Turcz. Non Hetero- cladia Dnd. as Hererocladia Dnd. ex Turcz., sphalm. ex Jacks. Heterophylleia Turez., Bull. Soc. Imp. Naturalistes Moscou 21: 591. 1848. Type: H. caracasana (Turcz.) Turcz. Subshrubs to small trees. Roots fibrous, with prominent nitrogen-fixing nodules. Stems rhizomatous, the branches sharply quadrangular, becoming terete, eventually arching to pendulous, the lenticels corky. Clusters of axillary buds produced in the axils of the leaves of the main axes. Leaves simple, whorled to opposite, appearing distichous through alternate twisting of the branches, often approximate above, short-petiolate to subsessile, the minute stipules withering and persistent or caducous, the blades palmately veined, lanceolate to ovate, entire, cordate, truncate to rounded at the base, the apex rostrate to mucronate. Racemes erect to pendent, terminating main or lateral axes. Flowers small, basically pentamerous, actinomorphic, or the calyx slightly zygomorphic, perfect, polygamo- monoecious, andromonoecious or appearing so through marked proterogyny, hypogynous, green, red or dark pur- ple, the pedicels ebracteolate to several-bracteolate, the sepals at fertilization, but accrescent and prominently fleshy subulate, persistent, the petals distinct, shorter than the sepals at fertization, but accrescent and prominently fleshy in fruit, ultimately black, red or yellow, stamens 10 in two 1972] Coriaria — Skog 245 whorls, with filiform filaments 2 mm. long, each anther sagittate, smooth to verruculose, 4-celled, dehiscing by longitudinal slits, the pollen spheric, 3-colporate, pistils 5 to 10 (-12), distinct, simple, the styles fasciculate and strict or divergent, papillose and stigmatic over their entire surfaces. Ovule solitary, pendant. Fruits dorsally keeled achenes, each bilaterally compressed, all enclosed by the fleshy accrescent petals. Seeds with a single mem- branous testa and scanty endosperm. Key to the species and subspecies of Coriaria. (All known species included here for completeness. ) a. Bases of lateral branches nearly always without bud scales; inflorescences produced terminally, either on the main stem or on branches, but on the current flush of growth; racemes sparsely to very densely pubescent; bracts subtending the flowers lanceolate to elliptic, en- tire; flowers strictly perfect; inner whorl of stamens free; achenes 1 to 2 mm. long. (b) b. Stems herbaceous, dying to the ground following maturation of fruits; leaves of the flowering axes ovate to suborbiculate, with 7 to 9 main basal veins; fruit pedicels more than 1.5 cm. long. Himalayas of Tibet to China. .... 4478 Coriaria terminalis b. Stems woody, perennial; leaves of the flowering axes lanceolate to elliptic, with 3 to 5 main basal veins; fruit pedicels less than 1 cm. long. .. V. Coriaria ruscifolia b e oo o o D e $4.» o o we. 68) Q ^. 9 a ee bearing numerous short branches bearing inflor- escences terminally; leaves 0.5 to 3.1 cm. long. Western slopes of the Andes from Mexico to Peru, and cool mountains of New Guinea and New Zealand. ose nun PO aa ea "e Coriaria ruscifolia subsp. microphylla c. Large shrubs to small trees; main axes bearing inflorescences in the axils of the leaves or rarely on short branches; leaves 1.0 to 7.5 cm. long. Temperate coasts of Chile, New Zealand and 246 Rhodora [Vol. 74 other Pacific islands. ....................00. ZEE Coriaria ruscifolia subsp. ruscifolia a. Bases of lateral branches surrounded by persistent bud scales; inflorescences produced from the axils of the leaves of the previous flush of growth; bracts subtend- ing the flowers ovate to orbiculate; flowers usually andromonoecious; inner whorl of stamens epipetalous. Shrubs of the Northern Hemisphere. (d) d. Racemes finely puberulent to densely pubescent; achenes 1.5 to 2.5 mm. long; bracts subtending the flowers subentire to erose. Northern India, Nepal and Western China. ......... Coriaria nepalensis d. Racemes glabrous; achenes more than 3 mm. long: bracts subtending the flowers entire. (e) e. Accrescent petals far exceeding the mature achenes and enclosing them; leaves ovate-lanceo- late; styles linear to fusiform. Japan, Formosa and the Philippines. ....... Coriaria japonica e. Accrescent petals not exceeding the exposed mature achenes; leaves subelliptic; styles fili- form. West-Mediterranean region. ............ 2*2 25 2 2-222929» Coriaria myrtifolia Coriaria ruscifolia L., Species Plantarum, ed. 1. 1037. 1753. Plants fruticose; new branches sprouting and developing rapidly, fleshy, at first terete, green to purple to gray, secondary growth producing a quadrangular stem which is brown or gray at maturity, cork developing early, lenticels present only on the previous flush of growth, scaly terminal buds not formed. Leaves subsessile, glabrous and dark green above, puberu- lent on the emersed veins and paler beneath ; leaves of the main stem orbiculate, with 5 to 9 main basal veins, those of the lateral branches elliptic to lanceolate, with 3 to 5 main basal veins, acute to cordate at the base, the apex acute to mucronate. Racemes few to numerous, pendent, finely to densely pubescent, the pedicels ebracteolate, 3 to 6 mm. long, at 1972] Coriaria — Skog 247 maturity less than 1 cm. long, with entire, lanceolate to subulate bracts. Flowers 2 to 3 mm, in diameter, usually pentamerous, regular, perfect, proterogynous, sepals ovate, green to red, 1.5 to 2 mm. long, 1 to 1.5 mm. wide, petals smaller, ovate, anthers yellow to reddish, styles linear, fasciculate, green to purple, matured achenes enclosed by the accrescent fleshy, black petals, 1 to 2.5 mm. long. The numerous populations of Coriaria found in the West- ern Hemisphere are here regarded as two subspecies of a single species, polymorphic in leaf size and shape and habit in response to geographic location and altitude. The char- acters which unite all these plants into one species are listed below: 1. The inflorescences are produced on wood of the cur- rent growing season. These racemes are either borne terminally on the lateral leafy branches or from the axils of the large orbiculate leaves on the main axes. 2. The leaves on the lateral branches have 3 to 5 veins. 3. The flowers have the same morphology throughout the distributional range and are always perfect, a trait shared only with Coriaria terminalis of the Himalayas. The two subspecies have heretofore been maintained as separate species since they were first described. Reducing these taxa from species to subspecies reflects their simi- larity and at the same time illustrates the differences in morphology which are ecological adaptations to altitudinal changes. At higher elevations, 600 m. to 4000 m. in the Andes, small-leaved forms occur while larger-leaved forms grow at elevations from sea level to 300 m. in southern South America. Another character which varies with alti- tude and which is noticeable in dried specimens as well as in the living plants is the increased numbers of lateral branches occurring with increased elevation. The correla- tion with altitude of these variations, independent of genetic constitution, has been demonstrated by comparisons of seedlings of South American forms grown under identical greenhouse conditions at the University of Connecticut. Seeds were collected in Colombia at 3200 m. (Skog 1000) 248 Rhodora [Vol. 74 and Chile at 5 m. (Skog 1075), germinated and grown to- gether in the greenhouse. The resulting plants were similar in leaf size and shape, and bore few lateral branches. (Herbarium vouchers are on file at CONN of the repre- sentative plants, Skog 1375, 1376.) Studies of herbarium specimens and field observations indicate that when plants otherwise assignable to the small-leaved subspecies, C. ruscifolia subsp. microphylla, are found at lower eleva- tion (less than 1000 m.) the leaves are larger and the individuals have fewer lateral brances. Coriaria ruscifolia L. subsp. ruscifolia Coriaria ruscifolia L., Species Plantarum, ed. 1. 1037. 1755. Type: Feuillée, Journal des observations physiques, mathématiques et botaniques 3 [Histoire des plantes méde- cinales . . . du Perou & du Chily]: 17, t. 12. 1725? Large shrubs to small trees. Main axes to 7 m. long. bearing whorled or opposite leaves 1.0 to 7.5 cm. long, 0.8 to 3.2 em. wide. Racemes arising from the axils of the leaves of the main axes, or at the apices of the few lateral branches, to 25 cm. long. Vernacular names: CHILE: deu, dehue-lahuen, deó, hiuque, mataratones, veu. Distribution and habitat in the Western Hemisphere: River banks or lake shores in western Argentina in the province of Neuquén, and in central and southern Chile in the provinces of Arauco, Biobio, Cautin, Chiloé, Concepción, Llanquihue, Malleco, Maule, Nuble, Osorno, Santiago and Valdivia. Representative specimens examined: ARGENTINA. PROVINCE OF NEUQUÉN: Lago Nahuel Huapi, Isla Victoria, Cordini 8 (F, US). CHILE. PROVINCE OF ARAUCO: Contulmo, Gunckel 21784 (CSG) ; Santa Cruz s. n., 1.835 (BH). PROVINCE OF BIOBIO: Fundo ‘Los Prados,’ Riegel s. n., Jan. 1, 1955 (CSG). PROVINCE OF CAUTIN: Rio Quepa, Temuco, D. Bullock s. n., 31 Jan. 1906 (BM); Quebrada del Río Pal- quin, Gunckel 15413 (csc); Puerto Saavedra, Hollenmayer 8 (CSG, M). PROVINCE OF CHILOÉ: Chiloé, J. Anderson 61 (BM); Piruquina, Junge s. n., Mar. 25, 1932 (csG) ; Junge 386 (M, MO); near Puerto Varas, 50 m., Morrison 17537 (BH, GH, MO) ; Castro, 1-10 m., Pennell 1972] Coriaria — Skog 249 12600 (F, GH, SGO, US); cerros cerca de Chonchi, Ricardi 5308 (CONC, CONN). PROVINCE OF CONCEPCION: Laraquete, Castillo s. n., Feb. 1946 (csG); Conception, Cuming 1461 (BM); Tomé, Germain s. n., Nov. 1855 (GH, sco); Talcahuano, Póppig 131 (BM, MO); near Tumbes, 5 m., Skog 1040 (CONN). PROVINCE OF LLANQUIHUE: Lago Llanquihue, Calvert s. n. in 1912 (BM); Petrohue, Gunckel 9227 (csG); Piedra Azul, Ricardi 5286 (CONC, CONN); Casa Panque, Shan- non & Shannon 24 (US); near Los Riscos, 200 m., Skog 1073, 1074, 1075 (CONN); near Ensenada, 100 m., West 4678 (GH, MO). PROVINCE OF MALLECO: Puren, Claude-Joseph 3029 (US). PROVINCE OF MAULE: Constitución: Burnier s. n. Sept. 1958 (csG), Gunckel 21785 (csa), Volekmann s. n. (SGO). PROVINCE OF NUBLE: Valle Rio Renegado: Alvares Ramirez s. n., Mar. 1954 (csG), Cuming 146 (E, GH); Valle de Alico, Jarpa s. n., Feb. 10, 1935 (csG); Reed s. n., 1871 (BM, GH); Sargent s. n., Jun. 29, 1906 (A). PROVINCE OF OSORNO: Fundo Río Blanco, southwest of Purranque, Eyerdam 10758 (F, US); Lago Llanquihue, F. Phillipi & Barchers s. n., 15/1/85 (BM); north shore of Lago Llanquihue, Senn 4635 (MO, US); southwest of Puerto Klocker, near Lago Llanquihue at base of Volcán Osorno, Skog 1062, 1064 (CONN). PROVINCE OF SANTIAGO: Claude-Joseph 957 (GH, US). PROVINCE OF VALDIVIA: Valdivia, Bridges 612 (BM); Uferbüsch des Calle-calle, Buchtien s. n., Sept. 17, 1896 (E, GH, M, US) ; ad margines rivulorum, Gay 1019 (sao); Isla del Ray, Carbonares, Gunckel 1097, 1149 (cse); Valdivia, Buchtien s. n., 12.X1.1904 (LD); Valdivia, Lechler s. n. (M). Coriaria ruscifolia L. subsp. microphylla (Poir.) L. Skog stat. nov. Coriaria microphylla Poir., Encyclopédie méthodique. Botanique 4: 87. 1804. Type: J. Jussieu s. n., P-JU. Coriaria thymifolia Humb. & Bonpl. ex Willd., Species Plantarum, ed. 4. 4: 819. 1805. Type: Humboldt & Bon- pland. 3018, B, P. Coriaria phylicifolia Humb. & Bonpl. ex Willd., Species Plantarum, ed. 4. 4: 819. 1805, ex char. Coriaria atropurpurea DC., Prodromus systematis na- turalis regni vegetabilis 1: 740. 1824. Type: De Candol!e, Calques des dessins de la flore du Mexique de Mocino et Sessé, t. 1167. Heterocladus caracasanus Turcz. Bull. Soc. Imp. Natu- ralistes Moscou 20: 152. 1847. Type: Galeotti 293, LE. Heterophylleia caracasana (Turez.) Turez., Bull. Soc. Imp. Naturalistes Moscou 21: 591. 250 Rhodora [Vol. 74 Coriaria cuneifolia Sessé & Moc., Plantae Novae His- paniae, ed. 1. 173. 1890, ex char. Suffruticose herbs to large shrubs. Main axes to 4 m. long bearing numerous short lateral branches. Leaves op- posite to approximate, 0.5 to 3.1 cm. long, 0.2 to 1.6 cm. wide. Racemes terminating the lateral branches, to 15 cm. long. Vernacular names: COLOMBIA: rebentadera. ECUADOR: pinan, shanshi, shanchi or zhanzhi, tinta. GUATEMALA: moco tinto, moco de chompipe. MEXICO: helecho de tierra, tlalocopetate, tlalocopatlatl. PERU: mio-mio, saca-saca, mio-venenosa, raqui-raqui. VENEZUELA: tisis, helecho de playa, helecho-uvite. Distribution and habitat in the Western Hemisphere: Moist montane forest openings and hillsides in Mexico, Guatemala and Panama and on the western slopes of the Andes in Colombia, Venezuela, Ecuador and Peru. The plants referred to this subspecies have long been named C. thymifolia Humb. & Bonpl. ex Willd. Coriaria microphylla Poir. predates C. thymifolia by almost two years, however, and should be used if this group is main- tained as a separate species. Representative specimens examined: COLOMBIA. ANTIOQUIA: La Sierra, 18 km. north of Medellín, 2000 m., Archer 1307 (BM, COL, US). BOGOTÁ: Holton 27, 808 (GH); Triana 5596 (BM, COL); Cordillera de Bogotá, 2650 m., Triana s. n. (BM, MEXU). BOYACA: Grubb, Curry & Fernández-P. 519 (COL, US). CALDAS: Killip & Hazen 8956 (GH, US) ; Laguneta, Salento, 2500 m., von Sneidern 3149 (COL, MO). CAUCA: Cordillera Central, vertiente occidental, 2470 m., Cuatrecasas 19369 (A, 2 sheets, F); Puracé, Fernández-P. 6218A (COL); entre Popayán y Puracé, 2500 m., Yepes Agredo 390 (COL, F, US). CUNDINAMARCA: Salto de Tequendama, 2100 m., Alston 7409 (BM); García-Barriga 130, 17488 (COL); al sur de Usme, entre La Regadera y El Hato, 3000-3100 m., Idrobo, Jaramillo, Mesa-Bernal & Smith 399 (COL, MO, US). HUILA: Cordillera oriental, east of Neiva, 1300-1800 m., Rusby & Pennell 1008 (GH, MO, US). MAGDALENA: Van der Hammen 1165 (COL). NARIÑO: Ferndndez-P., et al. 1083 (coL); Pasto, Cebadal, 2700-2800 m., Schultes & Villarreal 7939 (COL, F, US). NORTE DE SANTANDER: Cuatrecasas & Garcia-Barriga 10064 (COL, US). PUTOMAYO: Garcia-Barriga 7766 (COL, US); Valley of Sibundoy, 1972] Coriaria — Skog 251 Sibundoy, 2225-2300 m., Schultes & Villarreal 7714 (COL, F). SANTANDER: Killip & Smith 16784, 19227 (A, GH, US). TOLIMA: Cuatrecasas 9361 (COL, US). VALLE: Cordillera Central, vertiente occidental, 2270-2320 m., Cuatrecasas 18140 (F). COSTA RICA. CARTAGO: Volcán Irazi: 10000-11300 ft., Allen 676 (F, MO); Four- nier 1202 (CRU); Potrero Cerrado road, 6500 ft., Lankester s. n., 1925 (F, US); 2400 m., Skog 1339 (CONN); Smith 4761 (GH, US, 2 sheets) ; 1400 m., Torres-R. 288, s. n., 1924 (su). ECUADOR. AzuAy: Rio Tarqui, 8300 ft., Camp E-3905 (F, GH, US). BOLIVAR: entre Guaranda y Vinchoa, 2800 m., Acosta Solis 5927 (F). CANAR: near the village of Marcos, Camp E-2488 (MO). CHIMBORAZO: 2450 m., Acosta Solis 5534 (F); Huigra, Rose & Rose 22182 (BM, US). IMBABURA: Shanchi- pamba, 2750 m., Acosta Solis 14514 (F, 2 sheets). LOJA: 2200 m., Dodson & Thein 567 (MO); near Loja, between 2300-2700 m., Skog 1162, 1167, 1168, 1169, 1172, 1192 (CONN). PINCHINCHA: Cord. occ., los Alpes, 2800-3000 m., Acosta Solis 7079 (F); vicinity of Quito, 3000 m., Asplund 6157 (LD, US); above Quito, 10300 ft., Balls B5846 (BM, E, MO, US); Quito, Couthouy s. n., 1855 (GH); Quito, Jameson 206 (BM); Alrededores de Quito, 2850 m., Paredes 23 (Q). TUNGU- RAHUA: entre Leito y la Coma, Cord. oriental, 2700-3000 m., Acosta Solis 8977 (F); Volcano Tungurahua, 2200-2400 m., Dodson & Thein 1886 (M0); Rio Ambato Valley, W. of Ambato, Fagerlind & Wiborn 965 (LD). LOCALITY NOT DEFINITE: Heinricks 634 (M); interandine highland, 2000 m., Rimbach 97, (A, F). GUATEMALA. CHIMALTEN- ANGO: Barranco de La Sierra, southeast of Patzüm, about 2100 m., Standley 61508 (A, F). EL PROGRESSO: slopes adjacent to Finca Piamonte, 2500-3000 m., Steyermark 43711 (A, F). ESQUINTLA: Texcuaco, 150(?) m., Morales 1061 (F). GUATEMALA: near Finca La Aurora, 1500 m., Agwilar 248 (F); near Guatemala, Kellerman 5252 (MEXU, US); Pinula, 4200 ft., J. Donnell Smith 1928 (M). HUE- HUETENANGO: San Juan Atitan, 8400 ft., Skutch 1166 (A, F). JALAPA: between Miramundo and summit of Montaña Miramundo 2000-2500 m., Steyermark 32753 (F). QUEZALTENANGO: Sunil, Hartweg 524 (BM, GH, LD); near Santa Maria de Jesus, 6400 ft., Skutch 901 (A, F, US). SACATEPEQUEZ: Near Antigua, 6000 ft., Kellerman 1510 (F, 2 sheets, US); near San Lucas, 2300 m., Williams & Molina 11833 (F, GH). SAN MARCOS: southwest of San Marcos, 8000 ft., Morley 725 (F, GH, US). SOLOLÁ: Volcán Atitlán, Beaman 4091 (GH); Volcán Santa Clara, 2100-3000 m., Steyermark 46991 (F). JUNCTION OF HUEHUETENANGO, TOTONICAPÁN AND QUEZALTENANGO: Sierra Madre mountains, 2400-2600 m., Williams, Molina & Williams 22608 (F). LOCALITY NOT DEFINITE: Fosberg 19055 (coL, Us). MEXICO. CHIAPAS: Puerto Viento, Rayón, Breedlove 9003 (BM); Mt. Pasitar, near San Cristobal, Matuda 461 (F, MEXU, MO, US); Mt. Tacana, 1000-2000 m., Matuda 2417 (A4, F, MEXU) ; near San Cristobal de las 252 Rhodora [Vol. 74 Casas, above 7000 ft., Pfeifer, Kremer & Abendroth 2501 (CONN, 2 sheets); Rio Prospero, Seler 2269 (GH, US). GUERRERO: Hinton 14201, 15398 (GH, US); Distrito Mina, Petlacala, Ynes Mexia 8972 (F, GH, MO). JALISCO: Rich canyons, mountains near Lake Chapala, Pringle 2437 (BM, F, GH, M, MEXU, MO, US). MEXICO: ravines near Ozumba, 8000 ft., Pringle 9713 (CU, F, GH, MEXU, MO, US); Pringle 11971 (CU, F, GH, US); along brooks, Amecameca, Purpus 1768 (F, GH, MO, US); Ozumba, Purpus 3057 (BM, E, F, GH, MO, US). MICHOACAN: San Juan Viejo, Beaman 4393 (GH); near Paracutin, along road into Angahuan, 7500 ft., Bratz M-806 (MEXU); Tancitaro, Uruapan, Hinton 15448 (GH, US); 2 km. N. & NE. of Puentacillar, Iltis, Koeppen & Iltis 387 (BM); above Tancitaro, 7000 ft., Leavenworth 270 (F, GH, MO). MORELOS: Jojutla a de Juárez, D. de Juárez, Salazar s. n., August 8 1912 (MEXU, 2 sheets). SINALOA: Along Mex. #40, between Durango and Mazatlan, steep mountain sides, Pfeifer & Skog 3073 (CONN). PANAMA. Volcán de Chiriqui Potrero Muleto, Boquete district, 10400 ft., Davidson 1013 (A, F, US); be- tween El Hato and Cerro Punta, Llanos, Ebinger 800 (Mo); Petrero Muleto to summit de Chiriqui, Woodson & Schery 416 (GH, US). PERU. APURIMAC: Prov. Abancay, arriba Abancay, 3300-3500 m., Ferreyra 9801 (USM); Abancay, Vargas 1954 (CUZ) ; between Huan- carama and Cochacaya, 3400 m., West 3761 (GH, MO). AYACUCHO: Aina, between Huanta and Rio Apurimac, 750-1000 m., Killip & Smith 23119 (F, US). cuzco: Machu-Piechu, 7600 ft., Gourlay 87 (E); Machu-Picchu, 2200 m., Skog 1144, 1146, 1147 (CONN); Machu- Picchu, 6000 ft., Stafford 1053 (BM, F); Urubamba, 2000 m., Vargas 1871 (cuz, GH); Pillahuata, cerro de cusilluyoc, 3000-3300 m., Pen- nell 14131 (F, GH, US); Prov. Quispicanchis, Dist. Marcapata, 3000 m., Vargas 9701 (^, F, MO); Ollantaytambo, about 3000 m., Cook & Gilbert 607 (^, F, GH, US, 2 sheets). HUANCAVELICA: east of Surcu- bamba, Prov. Tayacaja, 2500 m., Stork & Horton 10343 (A, F, MO). HUANUCO: Prov. Ambo, cerca a San Rafael, 2600 m., Ferreyra 1961 (USM, US); Pampayacu to Huanuco, Kanehira 230 (A, F); Chaglia, Woytkowski 5293 (F, MO). JUNIN: east of Huasahuasi, 2400-2500 m., Hutchinson 1115 (BH, F, GH, M, MO, US, USM). LA LIBERTAD: Prov. Otuzco, cerca a Usquil, 3000-3100 m., Ferreyra 7668 (US, USM). PUNO: Prov. Carabaya, Ollachea, 2700 m., Vargas 6918 (cuz). LOCALITY NOT DEFINITE: Jameson 577 (E); “Perou,” J. Jussieu s. n. (P-JU); Ruiz & Pavon s. n. (BM, MO). VENEZUELA. MÉRIDA: Mérida, Alston 6673 (BM, US); Plaza de Mucuqui, 2200 m., Bernardi 189 (MER, VEN); northeast cf Mérida along road to Valencia, 1850 m., Breteler 3300 (CONN, IFLA, 3 sheets, US); San Rafael, 3185 m., Gehriger 9 (^, GH, MO, Us); Distrito Campo Elías, San José de Acequias, 2400 m., López-Palacios 774 (CONN, MER); El Cucharito near Tabay Village, 1800 m., Skog 1247 (CONN); between Tabay and 1972] Coriaria — Skog 253 Mérida, along Rio Chama, 1820 m., Steyermark 55865 (F). TACHIRA: Distrito Uribante, San José, antes de la poblacion de Pregonero, 1440 m., Marcano-Berti 1680 (CONN, MER). TRUJILLO: Jajó, Burkart 16686 (VEN). ACKNOWLEDGEMENTS The author gratefully acknowledges the aid and advice of Dr. H. W. Pfeifer during the work, Dr. W. J. Dress for reading the manuscript and support from the National Sci- ence Foundation Grant to the University of Connecticut for graduate student travel, which enabled the author to visit herbaria and collect in Central and South America. Also, the author is grateful to the curators of the herbaria cited for allowing him to examine material of Coriaria. The abbreviations for herbaria are from Index Herbariorum, ed. 5 by Lanjouw and Stafleu, except for the following, which are not included: CRU — The herbarium of the Uni- versity of Costa Rica, San José, Costa Rica; csa — The personal herbarium of Professor Hugo Gunckel L. of San- tiago, Chile; IFLA — The herbarium of the Instituto Forestal Latino-Americano, Mérida, Venezuela. LITERATURE CITED BENTHAM, G. & J. D. HOOKER. 1862. Genera Plantarum . 1: 429. CANDOLLE, A. P. DE. 1816. Essae sur les propriétes medicales des plantes, ed. 2. p. 350-351. CRONQUIST, A. 1968. The Evolution and Classification of Flowering Plants. 396 p. HALLIER, H. 1905. Provisional scheme of the natural (phylo- genetic) system of Flowering Plants. New Phytologist 4: 151- 162. HUTCHINSON, J. 1964. The Genera of Flowering Plants. 1: 172- 173. LAWRENCE, G. H. M. 1951. Taxonomy of Vascular Plants. 823 p. MaxriMOWICZ, C. J. 1881. De Coriaria, Ilice et Monochasmate, hujusque generibus proxime affinibus Bungea et Cymbaria. Mém. Acad. Imp. Sci. Saint Pétersbourg, sér. 7. 29(3): 1-13. Russow, E. 1884. Uber den anatomischen Bau der Laubsprosse der Coriarieen. Sitzungsber. Naturf.-Ges. Univ. Dorpat 6: 88. L. H. BAILEY HORTORIUM CORNELL UNIVERSITY ITHACA, NEW YORK LIST OF THE CLADONIAE OF SOUTHEASTERN MASSACHUSETTS, WITH SPECIAL REFERENCE TO THE COLLECTION OF CHARLES ALBERT ROBBINS LAWRENCE B. MISH After Charles Albert Robbins died in 1930, S. F. Blake (1933) wrote of the long and methodical collecting and identifying done by this quiet man from Wareham, Massa- chusetts, compiling the largest private collection of the genus Cladonia up to that time. Robbins’ collection for the small town on Cape Cod was more extensive than the col- lection of Evans (1930) for the whole state of Connecticut. For almost forty years, Robbins’ Cladonias have been housed in the Farlow Herbarium and his approximately 46 species are unknown to the world except for those discussed in his five papers, published between 1924 and 1929. While surveying the plants of Southeastern Massachusetts during a sabbatical leave in 1968-69, it was found that some of the plants in the Robbins collection were not included in the work of Thomson (1967). It would seem wise, there- fore, to publish a list at this time. The numbers appearing in front of each taxon correspond to those of Thomson (1967). Where no connection could be made with the taxa in Thomson’s work, Zahlbruckner (1922-1940) was referred to. The letters following the taxa indicate the collection locality, viz.: B — Bournedale NF — North Falmouth pr — Bridgewater o | — Orleans pre — Brewster P — Plymouth c — Carver R — Rockland p — Duxbury S —Stoughton E — Eastham sh — Sharon H — Hingham w — Wareham M — Marshfield wf — Wellfleet Na — Nantucket wH — Woods Hole 254 1972] Cladoniae — Mish 255 Those citations marked with an asterisk (*) are for plants not collected by Mr. Robbins. SCIENTIFIC NAME 1. C. papillaria (Ehrh.) Hoffm. f. molariformis (Hoffm.) Schaer. f. papillaria Thoms. f. stipata Flk. 5. C. coccifera (L.) Willd. v. frondescens (Nyl.) Vain. v. stemmatina Ach. 9. C. deformis (L.) Hoffm. f. crenulata Ach. 11. C. pleurota (Flk.) Schaer. 12. C. incrassata Flk. f. clavata Robb. f. squamulosa (Robb.) Evans 14. C. floerkeana (Fr.) Flk. v. intermedia Hepp. f. fastigiata Laur. v. intermedia f. peritheta Robb. f. intermedia (Hepp.) Thoms. f. carcata (Ach.) Thoms. f. floerkeana Thoms. f. squamosissima T. Fr. 15. C. bacillaris (Ach.) Nyl. f. attenuata Robb. f. peritheta (Wallr.) Zahlbr. f. fruticaulescens Vain. f. obtusa Wallr. f. bacillaris Thoms. 16. C. cristatella Tuck. f. aurantiaca Robb. PLACE B * Un gam zzz nm p a um ue dU m Mo EUIS (S Uu TEE ive] W,S Widespread WwW 256 21. 22. 23. 25. 30. əl. 32. 36. 3T. 39. f. rh Ph bh Th kh rh Ph Rhodora lepidifera (Vain.) Robb. pleurocarpa Robb. squamulosa Robb. squamosissima Robb. cristatella Thoms. ochrocarpa Tuck. ramosa Tuck. vestita Tuck. C. macilenta Hoffm. f. macilenta Thoms. C. didyma (Fée) Vain. var. muscigena (Eschw.) Vain. C. leporina Fr. C. piedmontensis Merr. f. lepidifera (Vain.) Robb. phyllocoma Robb. obconica Robb. piedmontensis Thoms. intermedia Robb. squamossisima Robb. C. strepsilis (Ach.) Vain. f. f. f. f. abortiva Robb. strepsilis Thoms. coralloidea (Ach.) Vain. glabrata Vain. C. robbinsii Evans f. squamulosa (Evans) Evans C. capitata (Michx.) Spreng. f. capitata, Thoms. C. clavulifera Vain. C. apodocarpa Robb. C. polycarpoides Nyl. f. epiphylla (Robb.) Thoms. æ TEE ~ TELE Z fo ~ zzz" Q2QAQ9009 Ss EI æ TERREN [Vol. 74 Ua 1972] 40. 46. 4T. 49. 56. 5T. 58. Cladoniae — Mish f. squamulosa..(Robb.) Thoms. f. polycarpoides Thoms. C. cariosa (Ach.) Spreng. f. squamulosa (Müll. Arg.) Vain. C. mateocyatha Robb. f. squamulata Robb. C. gracilis (L.) Willd. var. dilitata (Hoffm.) Vain. f. dilacerata (Flk.) Vain. var. elongata (Jacq.) Fr. f. elongata Thoms. C. verticillata (Hoffm.) Schaer. f. pallida Robb. var. cervicornus Flk. f. phyllophora Flk. phyllocephala Flot. aggregata (Del.) Oliv. apoticta (Ach.) Vain. phyllocephala (Flot.) Oliv. f. verticillata Thoms. C. simulata Robb. C. pityrea Flk.) Fr. var. pityrea Thoms. pityrea Thoms, squamulifera Vain. subacuta Vain. cladomorpha (Flk.) Vain. scyphifera (Del.) Vain. C. pyxidata (L.) Hoffm. var. pyxidata Thoms. f. centralis Oliv. f. marginalis Hoffm. f. prolifera Wallr. 257 33 * = a: MEE. a a 33333 EEEE: og 5 m e len 258 Rhodora [Vol. 74 f. lophyra (Ach.) Korb. f. homodactyla Wallr. 59. C. chlorophaea (Flk.) Spreng. f. costata (Flk.) Arn. f. simplex (Hoffm.) Arn. f. prolifera (Wallr.) Arn. 60. C. grayi Merr. peritheta Evans cyathiformis Sandst. centralis Evans prolifera Sandst. squamulosa Sandst. pallida Robb. epiphylaa Robb. foliosa Robb. . clavata Robb. 64. C. fimbrriata (L). Fr. f. stenoscypha Evans f. conista (Ach.) Robb. f. pycnotheliza (Nyl.) Vain. f. simplex (Weiss) Vain. 66. C. nemoxyna (Ach.) Nyl. f. ambigua Wallr. f. phyllocephala Arn. f. scyphifera Robb. f. nemoxyna Thoms. s444 xz a a Ph rh rh rh rh r5 rh rh rh 2zzzzzzz22 a wees w e Ui 68. C. coniocraea (Flk.) Spreng. f. ceratodes (Flk.) Thoms. f. phyllostrota (Flk.) Vain. f. truncata (Flk.) DT. & Sarnth. f. actinota FIk. f. pycnotheliza (Nyl) Vain. f. peritheta Robb. 72. C. turgida (Ehrh.) Hoffm. f. conspicua (Schaer.) Nyl. TEREE vn 2 4444 * ise] e$ 1972] Cladoniae — Mish 76. C. furcata (Huds.) Schrad. var. furcata Fk. TT. 79. 81. 82. 83. 86. 88. C. scabriuscula (Del. in Duby) Nyl. a DU DEP led fissa (Flk.) Aigr. furcata Thoms. subclausa (Sandst.) Evans rigidula Mass. subulata (Ach.) Vain. . corymbosa (Ach.) Vain. var. palamaea (Ach.) Vain. var. paradoxa Vain. var. pinnata (Flk.) Vain. f. f: P^ Ph rh Ph Ph f. recurva (Hoffm.) Sandst. foliolosa (Del.) Vain. turgida Scriba ex Sandst. elegans Robb. squamulosa Robb. . adspersa Flk. farinacea (Vain.) Sandst. scabriuscula Thoms. . parasitica (Hoffm.) Hoffm. f. f. f. congesta Robb. stertlia Robb. parasitica Thoms. floridana Vain. beawmontii (Tuck.) Vain. caespiticia (Pers.) Flk. f. epiphylla (Arn.) Sandst. multiformis Merr. carassensis Vain. E subregularis Vain. f. digressa Vain. f. f. spectabilis Robb. regularis Vain. *3 4344 2443 43445 - 243223 3 Z3 4 8 JEEEEELTILLLE - æ is o Ua T 259 260 Rhodora [Vol. 74 f. obliqua Robb. m. spectabilis Robb. f. subregularis Vain. m. spectabilis Robb. f. subirregularis Robb. z 89. C. squamosa (Scop.) Hoffm. f. pityrea Arn. f. fascicularis (Del.) Nyl. f. denticollis (Hoffm.) Vain. f. squamosa Thoms. var. laevicorticata Sandst. f. degenerascens Anders. f. laevicorticata (Sandst.) Evans f. carneopallida Sandst. var. muricella (Del.) Vain. f. sarmentosa (Tayl.) Müll Arg. f. ventricosa (Del.) Oliv. var. phyllocoma Robb. f. muricella Sandst. pseudocrispata Sandst. rigida (Del.) Nyl. subesquamosa Nyl. subtrachynella Vain. f. turfacea (Arn.) Vain. 90. C. subsquamosa (Nyl.) Vain. f. subrigida Robb. f. luxurians (Nyl.) Vain. f. degenerata Robb. f. sublevis Robb. f. subrigida Robb. f. scabrida Robb. 91. C. crispata (Ach.) Flot. f. subascypha Savicz var. dilacerata (Schaer.) Malbr. W zzzzz xz a z zz zzz Ph rh rh rh 4 tt ot 2a 355 3434334344 = 1972] Cladoniae — Mish 261 f. elegans (Del.) Vain. f. infundibulifera (Schaer.) Thoms. var. divulsa (Del.) Arn. f. crispata Thoms. var. virgata Ach. f. kairamoi Vain. 92. C. cenotea (Ach.) Schaer. f. exaltata Nyl. f. cenotea Thoms. 93. C. glauca Flk. 97. C. uncialis (L.) Wigg. f. humilior Fr. 44 449 3 444 4 J f. turgescans (Del. E. Fries w f. obtusata (Ach.) Nyl. W, E f. uncialis Thoms. Ww f. setigera Anders S W f. soraligera (Robb.) Sandst. W f. subobtusata (Coem.) Arn. Ww 101. C. boryi Tuck. W, P, D, WH, E, Bre, Q f. prolifera Robb. Ww f. lacunosa (Bory) Evans W, S, D, Sh 102. C. caroliniana Schwein, ex Tuck. W 107. C. alpestris (L.) Rabh. W, Br, E H 108. C. impexa Harm. f. prolifera Robb. w f. impexa Thoms. Ww f. subpellucida Harm. W 109. C. terrae-novae Ahti W 110. C. tenuis (Flk.) Harm. W: Ss, R f. prostrata Robb. Ww f. prolifera Robb. Ww 111. C. subtenuis ( Abb.) Evans W, S, E 262 .. Rhodora [Vol. 74 113. C. mitis Sandst. f. pallida Robb. tenuis Sandst. divaricata Sandst. prolifera Sandst. soralifera Sandst. f. prostrata Sandst. 115. C. sylvatica (L.) Hofim. f. sphagnoides (Flk.) Oliv. f. congesta Robb. m. fissa Flk. f. gracilior Vain. f. arbuscula Wallr. f. decumbens Flk. f. scabrida Robb. m. sorediata Sandst. f. pygmaea Sandst. 116. C. rangiferina (L.) Wigg. Widespread f. incrassata (Schaer.) Anders W f. tenuior (Del.) Vain. wW R, Nonquit - rh rh rh Ph Z "i 4444446 [^ Uu. Z "rj aL zZ £5 44444444248 Ua Not included in Thomson's book: C. poculifera Robb. f. simplex Robb. C. pseudopyxidata Robb. f. clavata Robb. f. Simplex Robb. f. squamulosa Robb. f. lateralis Robb. f. prolifera Robb. C. pyxioides. (Wallr.) Britz. f. simplex Britz. == 222222 22 z C. spumosa Flk. 99 1972] Cladoniae — Mish 26 LITERATURE CITED BLAKE, S. F. 1933. Charles Albert Robbins, 1874-1930. Rhodora 35: 140-145. Evans, A. W. 1930. The Cladoniae of Connecticut. Trans. Conn. Acad. Arts & Sci. 30: 357-510. THOMSON, J. W. 1967. The Lichen Genus Cladonia in North Amer- ica. U. Toronto Press. ZAHLBRUCKNER, A. 1922-1940. Catalogus Lichenum universalis, vols. 1-10. Leipzig. DEPARTMENT OF BIOLOGY BRIDGEWATER STATE COLLEGE BRIDGEWATER, MASSACHUSETTS 02324 ON THE MARINE ALGAE OF KENT ISLAND, BAY OF FUNDY KENNETH L. KOETZNER and R. D. Woop The extreme tidal amplitudes found in the Bay of Fundy are world famous, and biologists are intrigued by the rela- tionship between organisms and altitudes of such extreme tides. Whereas much of the benthic region of the upper reaches of the bay consists of clean-swept mud, there are scattered rock outcrops along the bay where attached algae can be studied. One such spot is Kent Island, New Bruns- wick, a fairly small island in the mouth of the Bay of Fundy. It lies just southeast of Grand Manan Island and is one of the Three Island group, located at approximately 44°35’ N lat and 66°46’W long. The junior author (rdw) visited the Bowdoin College Biological Station on Kent Island from June 28 to July 3, 1957, to study the marine algae. The results were compiled, but publication was deferred at the request of a colleague. Now, after nearly a decade during which no apparent con- flict of interest was apparent, further delay seemed un- justified; and, at the request of other colleagues, the report is now issued. During the five days on the island, collecting was done around the entire coastline; but, as work was sharply limited by rains, effort was concentrated in four areas: — on the north shore (N), a boulder-strewn beach ; — on the east shore (E), a cobble beach with rocks and tide pools at the low water level; — on the south shore (S), a rocky cliff region; — on the west shore in the Basin (B), a cove ex- posed as mud flats at extreme low tides. Specimens, both dried and fluid (5% of neutral formalin in seawater), were prepared and are on deposit at the University of Rhode Island. Examination of the specimens and compilation of the data was the primary responsibility of the senior author (klk) ; while the field work, observations, and final prepa- ration of manuscript were the responsibility of the junior 264 1972] Marine Algae — Koetzner and Wood 265 author (rdw). Classification and nomenclature follow Tay- lor (1957). Perhaps the first report of the algae of the island was by Bowers (1942) who identified 35 species, 15 of which were not noted during the present study. Other early re- ports of algae from the region but not from the island itself are those by Hay (1882) of 33 species and Hay and McKay (1886) for the Bay of Fundy. In addition, there are records by Eaton (1873) for Maine, Klugh (1917) for New Bruns- wick, Roscoe (1931) for Nova Scotia, Bell and MacFarlane (1933a, b) of 66 species for the Bay of Fundy, Humm (1950) for Newfoundland, and Taylor (1937; 1957) who treated in detail the algae of the entire northeastern coast of North America, including the Bay of Fundy. The junior writer (rdw) is indebted to Dr. James Moul- ton of Bowdoin College for the idea and arrangements for the visit; Dr. Charles Huntington, Director of the Bowdoin College Biological Station, and his wife, for their kindness and hospitality; and to that fabulous down east captain who “punched” his way through the impenetrable fog and found Kent Island “by ear.” (E = east shore; B = Basin; N — north shore; S — south shore; see text for descriptions) Chlorophyceae Ulotrichaceae l. Ulothrix flacca (Dillw.) Thuret. E, S. Abundantly matted on rocks at + 11 ft. Ulvaceae 2. Enteromorpha clathrata (Roth) J. Ag. — N, E, S, B, N of Basin. Variable in habit. Not common. (2a. E. compressa (L.) Grev. — (Bowers, 1942) ). 3. E. erecta (Lyngb.) J. Ag. — E. Rare. 4. E. intestinalis (L.) Link — N, E, S, B, N of Basin. On rocks and stones. Common. 9. E. linza (L.) J. Ag. — E. Rare. (5a. E. marginata J. Ag. — (Bowers, 1942) ), 266 Rhodora [Vol. 74 6. E. minima Nag. — E. Attached to stones. 7. E. prolifera (Müll.) J. Ag. — E, B. Masses attached to rocks in lower littoral. (also Bowers, 1942). 8. Monostroma fuscum (Post. & Rupr.) Witt. (f. blytii (Aresch.) Collins — N. In wash, rare. 9. Ulva lactuca L.—N, E, S. Lower littoral. (also Bowers, 1942). Prasiolaceae 10. Prasiola stipitata Suhr — N, E. Found high in spray zone. Cladophoraceae 11. Chaetomorpha melagonium (Web. & Mohr) Kütz. — Found in midlittoral zone and lower tide pools. (11a. C. area (Dilw.) Kütz. — Bowers, 1942). 19. Cladophora gracilis (Griff. ex Harv.) Kütz. — N, E, S, B. Abundant throughout mid and lower littoral zone. 18. C.ruprestris (L.) Kütz. — N, B. Uncommon, in lower littoral. 14. C. refracta (Roth) Kütz. — N, B. Uncommon. (also Bowers, 1942). 15. Rhizoclonium tortuosum Kütz. — N. With Clado- phora sp. in midlittoral tidepools. 16. Spongomorpha arcta (Dillw.) Kütz.— E, S. Form- ing thick, stiff tufts up to 6 in. in diameter, abun- dant in lower littoral zone. 17. S. spinescens Kiitz. — S. Uncommon, in association with S. arcta. At + 3 ft. Phaeophyceae Ectocarpaceae 18. Ectocarpus confervoides (Roth) Le Jol.—N, B. Epiphytic on coarser brown algae. (18a. Giffordia granulosa (J. E. Smith Hamel — (Bower, 1942, as E. granulosus) ). 19. Pylaiella littoralis (L.) Kjellm. — B. On rocks, Fucus and Ascophyllum. 1972] Marine Algae — Koetzner and Wood 267 Ralfsiaceae 20. Ralfsia fungiformis (Gunn.) Setch. et Gard. —N, E, S. Forming crusts several inches in diameter on rock, upper littoral zone. Elachisteaceae 21. Elachistea fucicola (Vell.) Aresch. — N, E. S. At- tached in dense tufts to Ascophyllum. Chordariaceae 22. Chordaria flagelliformis (Müll) C. Ag.—E, B. Found awash, often mixed with Dumontia. 23. Sphaerotrichia divaricata (C. Ag.) Kylin. — N of Basin. Occasional, + 9 to + 13 feet. Desmarestiaceae 24. Desmarestia aculeata (L.) Lamour. — N, S, B. Occa- sional, found in lower littoral. 25. D. viridis (Müll.) Lamour. — E. On shells of lower littoral, + 4 ft. Punctariaceae (25a. Asperococcus echinatus (Mert.) Grev.— (Bowers, 1942) ). 26. Scytosiphon lomentaria (Lyngb.) C. Ag. —S. At- tached to rocks, + 2 ft. 27. Punctaria latifolia Grev. — E, S, B. Common at or below low water. Dictyosiphonaceae 28. Dictyosiphon foeniculaceus (Huds.) Grev. — E, B. Common near low water. Laminariaceae 29. Agarum cribrosum (Mert.) Bory — N. Occasional, in sublittoral region. (also Bowers, 1942). 30. Alaria esculenta (L.) Grev. — E, N, S. Fairly com- mon near or below low water. (also Bowers, 1942). 31. Laminaria agardhii Kjellm. — N, E, S. Common, found with Alaria. (also Bowers, 1942). 268 Rhodora [Vol. 74 32. L. digitata (L.) Lamour. — E, S. Common below low water. (also Bowers, 1942). (32a. L. intermedia Fosl. — (Bowers, 1942) ). 33. L. longicruris De La Pyl.— B. Found attached to weir. (also Bowers, 1942). 34. L. platymeris De La Pyl. — E, B. In tide pools on beach at + 2 ft. (34a. L. stenophylla (Kütz.) J. Ag. — (Bowers, 1942) ; — L. digitata fide Taylor (1957: 185). ] 35. L. sp. —B. A form similar to L. agardhii but with flat stipe, 0.5-2 em. thick; blade thinner and wider. Fucaceae 36. Ascophyllum nodosum (L.) Le Jol. — N, E, S, B. Common in the entire littoral zone in most areas. (also Bowers, 1942). (36a. A. mackaii (Turn.) Holms et Batt. — (Bowers, 1942) ). 37. Fucus edentatus De La Pyl. — N, E. Form with ex- tremely flattened and broad receptacles. (also Bowers, 1942). 38. F. evanescens C. Ag., prox. — S of Basin. Occasional on rocks in sand. (also Bowers, 1942). 39. F. filiformis Gmelin — N, E. Common in high tide pools. (also Bowers, 1942). 40. F. spiralis L. — N, E, S, B. Common in all areas of the upper littoral zone, often mingling with strand vegetation. 41. F. vesiculosus L. — N, E, S. B. Common throughout the littoral zone (also Bowers, 1942). Rhodophyceae Bangiaceae 49. Porphyra umbilicalis (L.) J. Ag. — E, S. Occasional in lower littoral. (also Bowers, 1942). Acrochaetiaceae 43. ?Aerochaetium daviesii (Dillw.) Nàg. — N. (43a. Kylinia secundata | (Lyngb.) Papenf. — (Bowers, 1942 as A. secundatum) ). 1972] 44, 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. Marine Algae — Koetzner and Wood 269 Dumontiaceae Dumontia incrassata (Müll.) Lamour. — E, S. At- tached to rock in tide pocls, commonly with Chor- daria. Squamariaceae Hildenbrandia prototypus Nardo — E. Common in tide pools in the upper littoral. Corallinaceae Corallina officinalis L.— E, S. Abundant at or slightly above low water level. Kallymeniaceae Euthora cristata (L.) J. Ag. —S, W. Found awash. Rhodophyllidaceae Cystoclonium purpureum (Huds.) Batt. — N, S. On rocks at low water, with Porphyra. (also Bowers, 1942). Phyllophoraceae Ahnfeltia plicata (Huds.) Fries — E, S. Common in the mid to lower littoral zone. Phyllophora brodiaei (Turn.) J. Ag. — E. S. Occa- sional. Gigartinaceae Gigartina stellata (Stackh.) Batt. — E, S. Common to abundant in the mid to lower littoral. Rhodymeniaceae Halosaccion ramentaceum (L.) J. Ag. —N, S. Occa- sional. Rhodymenia palmata (L.) Grev. — E. Common at low water, often attaining length of 1.5 m. (also Bowers, 1942). Ceramiaceae Ceramium rubrum (Huds.) C. Ag. — S. Uncommon. Ptilota serrata Kütz. — S, B. In wash. 270 Rhodora [Vol. 74 . Delesseriaceae 56. Phycodrys rubens (Huds.) Batt. — N, S, W. Attached in the lower littoral. Rhodomelaceae 57. Polysiphonia harveyi Bailey — N. Uncommon. 58. P. lanosa (L.) Tandy — E, S. Epiphytic on Asco- phyllum, forming dense masses. 59. P. nigrescens (Huds.) Grev. — E, S. Forming dense growths on rocks in lower littoral. 60. P. urceolata (Lightf.) Grev. — N. Uncommon. 61. Rhodomela confervoides (Huds.) Silva — N, S. At- tached to rocks at lower water. LITERATURE CITED BELL, H. P. and MACFARLANE, C. I. 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. BELL, H. P. and MACFARLANE, C. I. 1933b. The marine algae of the Maritime Provinces of Canada, II. A study of their ecology. Ibid. 9: 280-293. Bowers, C. H. 1942. Algae of Kent Island. Bowdoin Sci. St. Bull. 8: 36-37. EATON, D. C. 1873. List of marine algae collected near Eastport, Maine, in August and September, 1873, in connection with the work of the U. S. Fish Commission under Prof. S. F. Baird. Trans. Conn. Acad. Arts & Sci. 2: 343-350. Hav, G. U. 1882. Marine algae of the Bay of Fundy. Bull. New Brunswick Nat. Hist. Soc. 1: 62-68. Hay, G. U. and MacKay, A. H. 1886. Marine algae of the Bay of Fundy. Ibid. 1: 32-33. HuMM, H. J. 1950. Notes on the marine algae of Newfoundland. Jour. Tenn. Acad Sci. 25: 229. KLUGH, M. V. 1917. Marine algae of the Passamaquoddy region, New Brunswick. Contrib. Can. Biol, Suppl. 6th Ann. Rep. Dept. Naval Serv., Fisheries Bur., pp 79-85. 1972] Marine Algae — Koetzner and Wood 271 Roscoe, M. V. 1931. The algae of St. Paul Island. Rhodora 33: 127-131. TAYLOR, W. R. 1937. Marine algae of the northeastern coast of North America. Univ. Michigan Press, Ann Arbor. 427 p. TAYLOR, W. R. 1957. Ibid., 2nd ed. 509 p. NEW YORK STATE DEPARTMENT OF ENVIRONMENTAL CONSERVATION, 4175 VETERANS MEMORIAL HIGHWAY, RONKONKOMA, N.Y. 11779 and DEPARTMENT OF BOTANY, UNIVERSITY OF RHODE ISLAND, KINGSTON, R.I. 02881 A WHITE-FLOWERED FORM OF IRIS LACUSTRIS FROM ONTARIO. In an open dune hollow within fifty yards of the shoreline north of McNab Point in Bruce County, Ontario, a white-flowered form of the Dwarf Lake Iris, Iris lacustris Nutt., was discovered amongst a few hundred blooming plants of the typical form. It is proposed that this new form be named Iris lacustris Nutt. forma albiflora Cruise & Catling, forma nova; differt forma typica quad flores albos non caeruleos habet. Presently known only from the type collection. Type (Specimen and Color Photograph): Associating with many of the typical form and with Oryzopsis asperi- folia and O. pungens, in moist sand in first dune hollow behind bay north of McNab Point (44° 28’ N., 81° 24’ W.), U. T. M. G. zone 17, 692243, ca. 2 miles south of South- ampton, Saugeen Township, Bruce County, Ontario. 22 May, 1971, P. M. Catling & S. M. McKay, (TRT 168074). J. E. CRUISE & P. M. CATLING DEPARTMENT OF BOTANY UNIVERSITY OF TORONTO TORONTO 181, ONTARIO, CANADA NEOMIRANDEA ALLENII, A NEW EPIPHYTIC COMPOSITE OF THE AMERICAN RAIN FOREST R. M. KING and H. ROBINSON Members of the Eupatorian genus Neomirandea, ranging from southern Mexico to Eucador, might well be called the Orchids of the Asteraceae. These usually epiphytic plants with rather succulent stems and showy inflorescences have only recently been recognized as a distinct genus (King & Robinson, 1970). Collectors have often commented, some- times extensively, on the unusual habit of the plants, and Professor B. L. Robinson (1918) in his description of a Colombian species, N. sciaphila (B. L. Robinson) R. M. King & H. Robinson, mentioned the horticultural potential of the plant. It is rare that labels do not mention the vining habit, epiphytic nature, or showy reddish-purple flowers. We take occasion here to call attention to some of the interest- ing problems of this unique group and to describe a four- teenth species from Panama and Colombia. The ecology of the genus Neomirandea presents a par- ticularly important area for future investigation. Most of the species, even some of the largest ones, are referred to as epiphytic. The habit prompted the name Eupatorium parasiticum Klatt, and a collection of the type species of the genus, N. araliaefolia (Lessing) R. M. King & H. Robinson was described by Skutch, “shrub or small tree with branches 25 ft. long and 6 inches in diameter. Epiphytic on an oak tree, 8 ft. above ground, the roots clasping the trunk in the manner of those of a strangling fig, concrescent where touching each other. A single large root descends along the trunk to the ground. Flowers white, rare. More about this interesting plant in my journal under March 4, 1933." The new species described here is cited by Allen as “fleshy branching epiphytic shrub 114 meters. In tops of tallest trees." The question remains, what of the few remaining members of this well marked genus not known to be epiphytic? But for their relation- ship, these species might never be examined properly to see 272 1972] Neomirandea — King and Robinson 273 if their substrate is in any way specialized, possibly old stumps or rotten logs. Experiments with the seeds of these plants are also essential to determine what adaptation they have to their substrate. Among the Eupatorieae, Neomirandea is one of the few genera of particular interest cytologically. The chromosome picture in the Eupatorieae is for the most part rather simple. Most groups show a haploid number of 10 chromo- somes or multiples of that number. Fleischmannia normally has n — 10, and a few species with » — 4 (Baker, 1967). A whole related complex of genera which we call Agera- tinoid (Ageratina, Oxylobus, etc.) shows base numbers of x = 17, and x = 16. Thus far, two species of Neomirandea have been counted, (Turner & King, 1964), N. angularis (B. L. Robinson) R. M. King & H. Robinson, n = 25 and N. costaricensis R. M. King & H. Robinson, » — 17 as Eupatorium aff. eximium. 'These two counts suggest some cytological complexity in this genus which is in some ways between the Ageratinoid complex and other members of the Eupatorieae. The new species is represented by two collections which show a range from Central Panama to the foothills of the northern Andes in Colombia. The characters of the species are as follows: Neomirandea allenii R. M. King & H. Robinson, sp. nov. Frutex epiphyticus carnosus. Folia opposita majuscula subglabra integra, petiolo perbreve; laminae late ellipticae 7-9 cm. longae, 4-6 cm. latae, apice anguste obtusatae, nervis lateralibus prominulis pinnatis. Involucri squamae ca. 12 subaequilongae late lanceolatae extus parce pubescentes. Flores ca. 10 in capitulo ca. 4.5 mm. longi; corollae anguste infundibulares extus distincte glanduliferae, faucibus intus glabris; styli non nodulosi; achaenae setiferae et glandi- ferae inferne dense glanduliferae et setiferae; setae pappi scabrae, cellulis apicalibus setarum nonnullarum valde obtusis. Fleshy branching epiphytic shrubs. Stems terete, min- utely pubescent. Leaves opposite, petioled; petioles glabrous 274 Rhodora [Vol. 74 ca. 5 mm. long, blades fleshy, ovate, entire, glabrous on both surfaces, up to 9 cm. long and up to 6 cm. wide, lateral veins prominent. Inflorescence a corymbose panicle. Phyl- laries ca. 12, subequal, broadly lanceolate, pubescent, in 2-3 series. Receptacle flat or slightly convex, glabrous. Heads ca. 10 flowered, flowers purple or pink, ca. 4.5 mm. long (excluding style branches) corollas narrowly funnel- form, lobes about as long as wide, outer surface of corolla with numerous short stalked glands, cells broadly oblong with straight walls, inner surface of corolla glabrous; anther appendages large, anther collars slender, composed mainly of rectangular to quadrate cells, walls not orna- mented. Style base not enlarged, glabrous, stylar append- ages not enlarged, mamillose. Achenes prismatic, 4-5 ribbed, with a few short stalked glands and uniseriate multiseptate hairs, glands and hairs numerous at base and apex, car- popodia distinct, cells quadrate, thin walled, pappus of ca. 30 scabrous setae ca. 5 mm. long, some setae with very blunt apical cells. PANAMA: PROVINCIA DE COCLE: Region north of El Valle de Anton, alt. 1000 meters, August 21, 1946. Fleshy branching epiphytic shrub, 115 m. In tops of tallest trees. Leaves leathery. Flowers purple, showy. Paul H. Allen 3643 (Holotype NY). Additional collection: COLOMBIA: ANTIOQUIA: Above Llano Grande, elevation 2000 m. Earl L. Core 497 (us). The new species by its lack of hairs inside the corolla, and by the lack of an enlarged style base proves to be a member of the subgenus Critoniopsis related to N. eximia (B. L. Robinson) R. M. King & H. Robinson and N. scia- phila (B. L. Robinson) R. M. King & H. Robinson. Among the prominent distinctions of N. allenii are the larger leaves with prominent secondary veins, the distinct glands on the surface of the corolla, the hairs and glands near the base of the achenes, and the frequent interspersed pappus setae with very blunt apical cells. 1972] Neomirandea — King and Robinson 275 ACKNOWLEDGEMENT This study was supported in part by the National Science Foundation Grant GB-20502 to the senior author. LITERATURE CITED BAKER, H. G. 1967. The evolution of weedy taxa in the Eupatorium microstemon species aggregate. Taxon 16: 293-300. KING, R. M. & H. R. RoBINSON. 1970. Studies in the Eupatorieae (Compositae). XXI. A new genus, Neomirandea. Phytologia 19(5): 305-310. ON ROBINSON, B. L. 1918. A descriptive revision of the Colombian Eupatoriums. Contr. Gray Herb. 55: 264-330. TURNER, B. L. & R. M. KING. 1964. Chromosome numbers in the Compositae. VIII. Mexican and Central American species. Southw. Nat. 9: 27-39. DEPARTMENT OF BOTANY SMITHSONIAN INSTITUTION WASHINGTON, D.C. 20560 STUDIES IN THE GUTTIFERAE. III. AN EVALUATION OF SOME PUTATIVE SPONTANEOUS GARDEN HYBRIDS IN HYPERICUM SECT. MYRIANDRA' PRESTON ADAMS For many years several species of Hypericum have been in cultivation at the Arnold Arboretum, Jamaica Plain, Massachusetts. Some of these are woody shrubs of Sect. Myriandra native to eastern North America. Around 1909 Alfred Rehder, a long-time member of the staff, discovered in the Arboretum garden plants of Hypericum which ap- peared to him to be intermediate in many characteristics between some of these native American species. He thought that these “atypical” individuals were spontaneous hybrids and, being interested in new and different plants for horti- cultural purposes, gave them binomials. The specimens which he collected are now in the herbarium of the Arnold Arboretum. During monographic studies of the woody specimens of Hypericum in eastern North America (Adams, 1962), I examined Rehder’s specimens. Having observed little evi- dence of hybridization among these plants in nature, I welcomed the opportunity to study putative examples of spontaneous crossing under garden conditions. The present contribution is an evaluation of the possible hybrid nature of Rehder’s specimens. Hypericum X Arnoldianum Rehder, Mitteil. Deutsch. Dendrol Ges. 19: 253. 1910 [19112]. Rehder applied this name to plants which he believed had originated from a cross between H. lobocarpum Gattinger and H. galioides Lam., with the latter being the maternal parent. During 1910 Rehder made several collections of this putative ‘Financial support for this study was provided by the Graduate Council Research Fund of DePauw University and the Fernald Fund for Field Study in Systematic Botany at Harvard University, estab- lished by the late Mr. F. W. Hunnewell of Wellesley, Mass. I want to thank Dr. Tod F. Stuessy of Ohio State University for helpful comments on the manuscript. 276 1972] Hybrids in Hypericum — Adams 277 hybrid. These plants persisted in the Arboretum for many years, there being collections made in 1916, 1921, and 1927 in the herbarium of the Arboretum. Study of Rehder’s specimens of H. X Arnoldianum lend support to his hypothesis of a hybrid origin for these plants. The most convincing evidence is the presence of a very poor seed set in the mature fruits on a collection made by Rehder on October 9, 1910. In each capsule examined nearly all of the ovules had aborted early in their development and only a few well-formed seeds were present in each fruit. The exact parental origin of H. X Arnoldianum is prob- lematical. Some substantiation for Rehder's belief that H. lobocarpum may have served as one of the parents of this species is provided by analysis of the style numbers. The 5-styled condition is characteristic of the majority of fruits of H. lobocarpum: some 79% of 2014 capsules from 23 collections of wild plants had 5 styles (Adams, 1962). While the 5-styled condition was found in only 3% of the 63 capsules of H. X Arnoldianum examined, about 16% had 4 styles. The 4-styled condition was present in about 19% of the fruits of H. lobocarpum examined. In addition, the deeply lobed condition so characteristic of the capsules of H. lobocarpum is approached by the partially lobed fruits of H. Arnoldianum. It seems reasonable, therefore, to consider H. lobocarpum to have participated in the ancestry of H. X Arnoldianum. The identity of the maternal parent of H. X Arnoldianum is less clear. Study of Rehder's specimens of this species does afford support for his hypothesis that H. galioides was involved. Evidence against this hypothesis is provided by analysis of style numbers. Of a total of 63 mature fruits of H. X Arnoldianum examined, 16% possessed 4 styles, 3% had 5 styles, and the remaining 81% were 3-styled. Capsules of the putative maternal parent, H. galioides, are almost invariably 3-styled, with 4-styled fruits being seen very rarely. What might have been the maternal parent of H. X Arnoldianum? A clue may be provided by the striking 278 Rhodora [Vol. 74 resemblance between specimens of this species and plants of H. densiflorum Pursh, especially those obtained from populations of the latter species growing in eastern Ten- nessee. In both plants the leaves are 10-15 times longer than wide. In addition, the seed size and coat ornamenta- tion of H. X Arnoldianum compare very well with that present in H. densiflorum. Plants of H. densiflorum were in cultivation at the time that Rehder described H. X Arnoldianum and these could well have served as one of the parents of this species. The close resemblance of H. X Arnoldianum to plants from the eastern Tennessee popu- lations of H. densiflorum could be explained by postulating that some of the Arboretum’s living collections of the latter species might have been originally obtained from the Ten- nessee portion of the geographic range of H. densiflorum. The available evidence, therefore, suggests that the spon- taneous cross that is hypothesized to have produced H. X Arnoldianum most likely occurred between plants of H. densiflorum and H. lobocarpum. Rehder’s suggestion that H. galioides may have been involved may have been due to misidentification of specimens. Such an error is under- standable, since several collectors of Hypericum in eastern Tennessee, especially during the late 19th and early 20th centuries, identified specimens of H. densiflorum as H. galioides. However, both Svenson (1940) and Adams (1962) have shown that the latter species is a native of the lower Atlantic and Gulf Coastal plains. Further support for the hypothesis that H. lobocarpum and H. densiflorwm may have been the parents of H. X Arnoldianum is provided by chromosome analysis. In plants from the Arnold Arboretum identified as H. X Arnoldianum (perhaps the same individuals studied by Rehder), Hoar and Haertl (1932) found a gametic number of 9. They reported “no irregularity in chromosome be- havior nor morphological sterility of pollen" and concluded that this hybrid “apparently came from compatible par- ents.” Since they did not preserve voucher specimens, their identification cannot now be verified. Jf they did indeed 1972] Hybrids in Hypericum — Adams 279 study the chromosomes of true H. X Arnoldianum, then the observed chromosome regularity and viable pollen would constitute strong support for the hypothesis that the putative parents were indeed H. lobocarpum and H. densiflorum. 'These two species are very closely related. In fact, H. lobocarpum, a plant of Arkansas and adjacent portions of surrounding states, has been considered as a variety of H. densiflorum (Svenson, 1940), but the geo- graphic ranges of each of these species are not known to overlap, there being at least 80 miles distance between the nearest known populations (Adams, 1962). Hypericum X Dawsonianum Rehder l. c. The plants of this putative hybrid were hypothesized by Rehder to have orginated from a spontaneous cross between H. lobocarpum Gattinger and H. prolificum L., with the latter as the maternal parent. Two collections, both in mature fruit, were made by Rehder in the Arboretum gardens during the fall of 1910. That Rehder's plants of H. X Dawsonianum are most likely indeed of hybrid origin is strongly suggested by the very poor seed set in the fruits, with most of the ovules having failed to develop to maturity. Additional support for this hypothesis is provided by an array of features and eonditions more or less intermediate between the two putative parents, H. lobocarpum and H. prolificum. The seed size and coat ornamentation of the few seeds that are present in the mature fruits are distinctly intermediate between the two presumed parental species. The style number of Rehder's collection of October 9 of H. X Daw- sonianum is also intermediate between these two species: about 50% are 3-styled and 50% are 4-styled. Almost the same ratio of 3- and 4-styled capsules is present in Rehder's collection of October 10; a very few fruits, however, had 5 styles. The 5-styled condition eccurred in about 79% of 2014 capsules from 23 collections (Adams, 1962) of H. lobocarpum. Fruits with 3 styles are the rule in H. pro- lificum, with only an occasional capsule having 4 or, very rarely, 5 styles. In addition, the fruit size of H. X Daw- 280 Rhodora [Vol. 74 sonianum is well within the range of H. prolificum. The sulcate to deeply lobed capsules of H. X Dawsonianum, however, are strikingly similar to those of H. lobocarpum. Further substantiation of the hypothesis of hybridization between H. lobocarpum and H. prolificum is the observa- tion that interbreeding may occur occasionally between plants of these two species growing in the field. I have seen 3 collections from Ripley Co., Missouri (Steyermark 66885, F; Bush 282, GH, NY ; Makensie 395, NY) and one from Howell County (Steyermark 40010, F, MO) which I cannot assign definitely to either species. Several factors favor the occurrence of hybridization between these two species. The distributional range of the more southern H. lobocarpum overlaps that of the more northern H. pro- lificum in central Arkansas, forming a zone some 3 or 4 counties wide running from the southwestern corner of the state diagonally across to southeastern Missouri (Adams, 1962). Plants of both species flower at the same time, and there is little obvious difference in their habitat requirements. As living specimens of H. lobocarpum and H. prolificum had been cultivated in the Arnold Arboretum for many years, it seems reasonable to conclude that Rehder’s H. X Dawsonianum specimens did indeed origi- nate as the result of spontaneous crossing between these two species under garden conditions. The meiotic chromosome behavior of an individual plant of H. X Dawsonianum was reported by Hoar and Haertl (1932) as displaying no irregularity. Neither did they observe any evidence of pollen sterility. Based on their findings, they concluded that the putative parents of the hybrid were genetically “compatible.” As with the case of H. X Arnoldianum, however, no voucher specimens were preserved and, therefore, it is not now possible to verify the original identification. Hypericum X nothum Rehder, l. c., 254. On October 10, 1910 Rehder collected specimens in the Arboretum which he interpreted to be the product of a cross between H. kalmianum L. and H. densiflorum Pursh, with the latter as the maternal parent. There is little evidence, however, 1972] Hybrids in Hypericum — Adams 281 to support his hypothesis of hybrid origin of these plants. The few 4- and 5-styled fruits present on the specimens of the putative hybrid are suggestive of H. kalmianum, which is typically 5-styled although 4 styles are not infrequently encountered. However, most of the fruits on Rehder's specimens have only 3 styles, a characteristic of H. densi- florum. Other close resemblances to H. densiflorum include seed shape, size and coat ornamentation, inflorescence form, and fruit shape and size. These features strongly suggest that Rehder's specimens of H. X nothum are most likely not of hybrid origin. They can be readily assigned to H. densiflorum. Hypericum X VanFleetii Hort. ex Rehder, Man. Cult. Trees and Shrubs, ed. 2, 640. 1940. According to Rehder, this binomial refers to a horticultural form believed to be a hybrid between H. prolificum and H. frondosum Michx. There is no evidence that the several specimens of culti- vated plants in the herbarium of the Arnold Arboretum bearing this name are of hybrid origin. Instead, I believe that they are readily referable to H. prolificum. It is doubtful if any of these putative hybrids are still being maintained by horticulturists. There were none present in the gardens of the Arnold Arboretum in 1959. This is not surprising since there are several introduced species of Hypericum which make better garden plants, with larger and more showy flowers and other more de- sirable features. In the living collections at the Kew Gardens in England there are several woody shrubs of Hypericum Sect. Myriandra originally obtained from east- ern North America. According to Dr. N. K. B. Robson of the British Museum of Natural History (personal commu- nication), these plants *do nct seem to fit exactly into any of the described species." It is entirely possible, therefore, that when two or more of the woody species are brought into the garden, some hybridization may take place. In nature, however, there appears to be very little interspecific hybridization among the woody members of Hypericum Sect. Myriandra. 282 Rhodora [Vol. 74 LITERATURE CITED ADAMS, PRESTON. 1962. Studies in the Guttiferae. I. A synopsis of Hypericum Section Myriandra. Contrib. Gray Herb. 189, 1-51. Hoar, C. S. and E. J. HAERTL. 1932. Meiosis in the genus Hyperi- cum. Bot. Gaz. 93: 197-204. SvENSON, H. K. 1940. Plants of the southern United States. II. Woody species of Hypericum. Rhodora 42: 8-19. DEPARTMENT OF BOTANY DEPAUW UNIVERSITY GREENCASTLE, INDIANA 46135 A NEW SPECIES OF THE ORCHID GENUS HOFFMANNSEGGELLA H. G. JONES The genus Hoffmannseggella was established by the author (Jones, 1968) to accommodate the species which were formerly assigned to the section Cyrtolaelia (Schlech- ter, 1917) of the genus Laelia. In his review of Laelia published in 1952, the late Dr. F. C. Hoehne, who prob- ably knew these plants better than anyone else, wrote: *Indubitavelmente, elas se distinguem tanto das que con- stituem as outras seccoes, que poderiam ser agrupadas como género distincto"; and later in the same paper; “Acreditamos, entretanto, que haverá vantagem . . . e razao para separar a seccao Cyrtolaelia de Laelia como género distincto." Apart from Laelia, one other generic name has been associated with Hoffmannseggella cinnabarina (Batem.) H. G. Jones, which is the generic type of Hoffmannseggella: in 1843 Hoffmannsegg had transferred this species to his new genus, Amalias. Unfortunately, in the original pub- lication of Amalias (1842), Hoffmannsegg had made only one specific combination, and this was based upon Laelia anceps Lindl., which is a true species of Laelia, not belong- ing to the section Cyrtolaelia. Consequently, since Amalias anceps (Lindl) Hoff. automatically becomes the type- species of Amalias, this name must be treated as a generic synonym of Laelia, and was not available for use for the species of section Cyrtolaelia, when the latter were raised to generic rank. The genus Hoffmannseggella falls naturally into two fairly distinct sections: in the typical section (sect. Hoff- mannseggella), the pseudobulbs are short and thick — conical, subfusiform or ovoid in shape — and the flower- scapes quite tall, usually much longer than the leaves. In the species of sect. Pleurothalloides, however, the pseudo- bulbs are reduced to slender cylindrical stems; and the flower-scapes are much shorter, about equal to or very 283 284 Rhodora [Vol. 74 slightly longer than the leaves. This gives the plants a rather distinct resemblance to the genus Plewrothallis — hence the sectional name — a resemblance which has been commented upon by Reichenbach (1874) and by Hoehne (1930). In the course of delimiting the species of the latter section for my projected monograph of the genus, I have found it necessary to recognize one new species, which is described and named below. Hoffmannseggella brevicaulis H. G. Jones, sp. nov. Species affinis H. harpophyllae a qua caulibus multo brevioribus, floribus majoribus labello apice anguste atte- nuato facile distinguitur. Epiphytica, erecta, circa 24 cm alta; rhizomate valde abbreviato; radicibus albescentibus, filiformibus, flexuosis, glabris; caulibus cylindraceis, uni- foliatis, circa 7 cm longis; folio erecto oblongo, coriaceo, apice obtuso vel subacuto, circa 17 cm longo, medio circa 2 em lato. Racemus erectus vel suberectus, 2-3-florus, quam folio multo brevior; floribus luteis, circa 6.5 cm diametri- bus: sepalis oblongis, apice acutis, circa 3.5 cm longis, medio circa 9 mm latis; petalis quam sepalia aequimagnis vel paulo brevioribus; labello circa 3 cm longo, medio circa 1.5 em lato, parte superiore profunde trilobo, carinis 2 paralellis elevatis, acutis; lobis lateralis subovatis, acutis; lobo mediano elongato, anguste attenuato, apice multo acuminato. Columna crassa subcylindrica, circa 1 cm longa, medio circa 4 mm lata ; ovario pedicellato glabro, circa 4 cm longo. Type: Brazil habitat not recorded — flowered unde: cultivation in Barbados, Feb. 1967. Herb Jones. H (L) /19a. This taxon has been previously referred to by the author in another paper (Jones 1970). It first came to my atten- tion during the course of examining the specimens of H. harpophylla (Rchb.f.) H. G. Jones, preserved in Reichen- bach's herbarium (Naturhistorischen Museums, Vienna), when I noticed that two of the sheets included under this name were obviously atypical. These were subsequently found to agree with cultivated specimens from Brazil which 1972] Hoffmannseggella — Jones 285 undoubtedly represented an undescribed species, somewhat related to H. harpophylla, but easily distinguished by the characters described in the above diagnosis, which may be summarized as follows: — H. HARPOPHYLLA 1. Plants tall and rather loose; pseudobulbs longer than the leaves. 2. Leaves narrow, tapering to a slender, acute point. 3. Inflorescence numerous, 6-8-flowered. 4. Flowers rich orange-red. 5. Sepals and petals narrow, usually less than 6 mm wide. 6. Front lobe of the labellum broadening outwards from the base, apex obtuse. H. BREVICAULIS 1. Plants shorter and more compact; pseudobulbs shorter than the leaves. 2. Leaves broader, apex obtuse or subacute. 3. Inflorescence 2-3-flowered. 4. Flowers yellow. 5. Sepals and petals broader, 7-9 mm wide. 6. Front lobe of the labellum very narrow, tapering to a long, slender point. H. brevicaulis is also related to another quite recently- discovered species, H. Kautskyi (Pabst) H. G. Jones (Hoffmannseggella Kautskyi (Pabst) H. G. Jones, comb. nov. Basionym: Laelia Kautskyi Pabst in Orch. Rev. 78: 321. 1970). The latter species resembles H. brevicaulis in the color of the flowers, but the habit, the form of the labellum, and the flowering period are all different. All the species of Hoffmannseggella are confined geographically to Brazil. REFERENCES HoEHNE, F. C. 1930. Album de Orchidáceas Brasileiras. 264 pp., Secretario da Agricultura, São Paulo. HOEHNE, F. C. 1952. Algo concernente ao género Laelia Lindl. das Orchidáceas e uma nova espécie para éle do Estado de Minas Gerais. Arquivos de Botánica do Estado de Sáo Paulo 2: 157-167. 286 Rhodora [Vol. 74 HOFFMANNSEGG, G. 1842. Verzeichniss der Orchideen für 1842. 22 pp., C. Heinrich, Dresden. HOFFMANNSEGG, G. 1843. Verzeichniss der Orchideen für 1843. 45 pp. H. M. Goltschalck, Dresden. Jones, H. G. 1968. Studies in Neotropical Orchidology. Acta Botanica Academiae Scientiarum Hungaricae 14: 63-70. Jones, H. G. 1970. Orchidaceae Americanae. Caldasia 10: 491-495. REICHENBACH, H. G. 1874. New Garden Plants. Gardener’s Chroni- cle. 5: 542-543. SCHLECHTER, R. 1917. Die Einteilung der Gattung Laelia und die Geographische Verbreitung ihrer Gruppen. Orchis 11: 87-96. POST OFFICE BOX 111 BRIDGETOWN, BARBADOS WEST INDIES CHROMOSOME NUMBER IN LIQUIDAMBAR FRANK S. SANTAMOUR, JR.! The genus Liquidambar belongs to the Hamamelidaceae, although most authorities consider this genus so distinct that they place Liquidambar, along with some other genera such as Bucklandia and Disanthus, in the sub-family Liquidambaroideae. Cytologically, all genera of Hamameli- daceae (sub-family Hamamelioideae) reported thus far have a basic chromosome number of x—12 (Darlington and Wylie, 1955). Liquidambar is the only genus of the sub- family Liquidambaroideae that has been studied cytologic- ally. Anderson and Sax (1935) studied meiosis in L. styra- ciflua L. growing at the Arnold Arboretum in Jamaica Plain, Massachusetts. They concluded that L. styraciflua was a diploid (2»—30) with a basic number of z=15 chromosomes. They also noted marked meiotic irregu- larities and high pollen sterility in their material, and ascribed these irregularities to cultivation in an area north of the natural species range. Pizzolongo (1958) reported that the diploid number of L. styraciflua (from shoot apices) and L. orientalis L. (from root tips) was 2n—32, and concluded that the haploid (and basic) number was »—16. However, the only countable meiotic figure he found in L. styraciflua was an anaphase I, showing groups of 15 and 16 chromo- somes. Meiotic irregularities and consequent pollen sterility apparently depended “upon some univalent chromosomes which do not respect the metaphasic congression and cause an unequal chromosome distribution among the pollen grains." This study was conducted in Italy, where the trees, as in the Anderson and Sax work, were growing outside their native range. Even with the irregular meiotic divisions, "Research Geneticist, U. S. National Arboretum, Plant Science Re- search Division, Agricultural Research Service, U.S. Department of Agriculture, Washington, D.C. 20002. 287 288 Rhodora [Vol. 74 FA ‘fo V e., *a LI + Figure 1. Meiotic stages in Liquidambar styraciflua L.; (a) Diakinesis with »—16 (540 X) (b) Anaphase I with two groups of 16 chromosomes (1200 X). however, Pizzolongo did not believe that the species had two biotypes, with 2»—30 and 2n=32 chromosomes. Work with Liquidambar in the project on “Cytogenetics, Breeding, and Evaluation of Shade Trees" at the U.S. National Arboretum has stressed interspecific hybridiza- tion. We have obtained hybrids between L. stryaciflua and both Asiatic species, L. orientalis and L. formosana Hance. Thus, it was important to determine the number and meiotic behavior of the chromosomes of our parent trees, especially L. styraciflua, to decide if detailed cytological analysis of the progenies would be necessary. 1972] Liquidambar — Santamour 289 In the spring of 1971, inflorescences of L. styraciflua were collected from 4 native trees on the grounds of the Arboretum in Washington, D. C. Two of these trees were used as parents in the breeding work. The flowers were fixed in 1:3 acetic-alcohol for 24 hours and stored in 80% ethanol. Meiosis in the pollen-mother-cell was studied using standard aceto-carmine squash techniques. Pollen was col- lected from the same trees and pollen abortion was deter- mined from a random sample of 200 grains stained with aceto-carmine. Pollen size was based on measurements of 50 sound pollen grains from each tree. Mitosis in root tips of seedlings of L. styraciflua X L. orientalis and L. styraciflua X L. formosana parentage was also examined using aceto-carmine squash techniques. Accurate chromosome counts at first metaphase of meiosis were, as ncted by previous authors, impossible to achieve. The chromosomes at this stage are clumped to- gether and no more than 10 distinct bivalents could be observed. Usually one, and frequently two, bivalents were found outside the metaphase grouping and exhibited pre- cocicus separation. Pizzolongo attributed the high pollen abortion he observed to this irregular chromosome behavior. In my material it was not difficult to determine n=16 chromosomes at diakinesis or late anaphase I. Many diads were counted with this number in both nuclei. At the 4-nucleate stage (anaphase II), nuclei with n=16 chromo- somes could also be determined, but in no case could all four nuclei be counted. The major problem in documentation was obtaining figures in which all the chromcsomes were in the same plane for photographic purposes. The picture of diakinesis (Fig. 1a) shows 16 chromosomes, with at least three and perhaps four chromosomes attached to the nucleolus. This configuration explains the multinucleolate microspores noted by Pizzolongo. At anaphase I (Fig. 1b), two groups of 16 chromosomes are shown. 290 Rhodora [Vol. 74 Pollen abortion in the four trees studied ranged from 6% to 20% and averaged 11%. Pollen grain diameter averaged about 39 microns, and did not differ significantly among trees. Although abnormal meiotic behavior may explain the rather high pollen abortion sometimes found in this species, the degree of meiotic irregularities probably depends on individual tree characteristics. Chromosome counts on root tips of hybrids of L. sty:a- ciflua with L. orientalis and L. formosana gave 2n-—82. These facts imply that all three species have the same chromosome number. LITERATURE CITED ANDERSON, EDGAR & KARL Sax. 1935. Chromosome numbers in the Hamamelidaceae and their phylogenetic significance. Jour. Arnold Arb. 16: 210-215. DARLINGTON, C. D. & A. P. WYLIE. 1955. Chromosome Atlas of Flowering Plants (ed. 2) George Allen and Unwin Ltd., 519 pp. PIZZOLONGO, PAOLO. 1958. Ricerche cario-tassonomiche su alcune Hamamelidales. Ann. Bot. (Roma) 26: 1-18. RECENT DISCOVERIES IN THE NEW HAMPSHIRE FLORA A. R. HODGDON AND FREDERIC L. STEELE? Our field work in New Hampshire during the 1971 season consisted only of a few day-excursions, no real effort having been made to search for unusual plants. However, several discoveries were made of which the following are most significant. Scirpus robustus Pursh. Saltmarsh at Adams Point, Durham, Strafford County, A. R. Hodgdon, G. Brooke, J. Wicks, et al., September 29, 1971 #19145. It is often true that several rare and local species may occur in close proximity. It happens that the halophytic composite Iva frutescens var oraria, previously reported from Durham (Hodgdon & Krochmal 1950), reaches its northeastern known limit of range, aside from its disjunct occurrence in Nova Scotia, within a stone’s throw of the Bulrush. The ample size of the Scirpus colony shows that it must have been growing here for a long time. Thus it must have been overlooked by the senior author many times during the past twenty or thirty years, perhaps because attention was directed too much to the Iva. The nearest reported stations for Scirpus robustus are in Newbury and Cambridge, Massachusetts, making the Durham locality the initial record for New Hampshire and the first north of Massa- chusetts. It is a conspicuous tall sedge, leafy nearly to the inflorescence, with big ovoid spikelets of rich brown color, the plant in size somewhat resembling the also robust Scirpus maritimus var Fernaldi which, however, typically has elongated spikelets. In the herbarium of the New Eng- land Botanical Club there are specimens from coasta] areas of southwestern Maine which have regularly been called S. maritimus var Fernaldi, but which have ovoid spikelets or otherwise approach S. robustus in some characteristics. 'Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientifie Contribution No. 616. 291 292 Rhodora [Vol. 74 Presumably the separation of these two taxa is not always an easy matter. Pycnanthemum Torrei Benth. Southern slope of Jeremy Hill, Pelham, Rockingham County, A. R. Hodgdon and Frederic Stecle, August 4, 1971 #19148. Jeremy Hill lies within a mile or two of the Massachusetts line. The south- ern slope where we fcund the Pycnanthemum has been known for a long time as a rich collecting area; many species of Desmodium and Lespedeza, Phryma leptostachya and other plants of interest grow along the road which extends upward on the slope into the woods. Arthur Stanley Pease collected there and his success in finding unusual species so far north led the senior author also to visit the place a number of years ago. It has been known as the northeastern known limit of range for Desmodium mari- landicum (L.) DC., Lespedeza virginica (L.) Britton and Pycnanthemum incanum (L.) Michx. while at least ten other species are found on this slope that have not been reported in Maine. On our August visit we explored the rocky slope just above the road a little way into the woods and came upon a clump or two of a big mint with which we were unfamiliar but which we assumed to be a Pycnan- themum and which proved to be P. Torrei. The calyx-teeth of P. Torrei are acuminate and are about 1.5 mm long serving readily to separate it from P. verticillatum or P. virginianum. In addition the leaves are thin and mem- branaceous and the heads are few at the tips of somewhat elongated slender branches. The leaves of P. Torrei are considerably wider than those of P. tenuifolium. This is a somewhat remarkable extension of range all the way from New Haven County, Connecticut where there are two stations for it known in New England. It was col- lected in New Haven by C. H. Bissell and R. W. Woodward in August 1910 and in Cheshire also in New Haven County by E. B. Harger in August 1929. Specimens from these two localities are present in both the Gray and the New Eng- land Botanieal Club Herbaria. It seems unlikely that this mint, being scarce even in southern Connecticut, has moved 1972] New Hampshire Flora — Hodgdon and Steele 293 to Jeremy Hill in recent times. On the other hand, it is surprising that it was overlooked by the botanists who have visited the area unless indeed their attentions were too much diverted by all the other interesting plants. Lechea tenuifolia Michx. Along cleared trail to fire tower, Jeremy Hill, Pelham, A. R. Hodgdon and Frederic L. Steele #19146. Our success that August day, in finding a species of Pycnanthemum seemingly new to New Hampshire, en- couraged the first author to assume that other significant species might also be discovered on Jeremy Hill. It was decided therefore to follow a trail upward to the fire tower. Plants of Viola pedata were scattered along the trail and higher up, where the trail widened into a cleared roadway there were plants of Hypoxis hirsuta in good flower. A few depauperate plants of the pinweed, Lechea tenuifolia were growing nearby. In 1879 Kate Furbish found this species in Wells, York County, Maine. In the Herbarium of the New England Botanical Club one can find this Wells speci- men but none from New Hampshire. In the Gray Her- barium however, there are two specimens from New Hampshire both being F. W. Batchelder collections from Pelham, one from the Gumpas District, the other with no precise record of locality. It is likely then that ours is the first record from Jeremy Hill and since Batchelder’s more recent collection was made in 1902, a contemporary com- munication about it may be justified to show that it still is to be found north of the Massachusetts line. Another pinweed, Lechea villosa', is found on Jeremy Hill and indeed quite generally in townships close to the Massachusetts line in New Hampshire, Durham being a somewhat isolated outpost for it to the northeast. ‘It seems preferable to retain the binomial Lechea villosa of Elliot which is clearly typified and about which there is no question instead of the vague, poorly defined and dubious L. mucronata of Rafinesque who published several other names that might be interpreted as L. villosa and who demonstrated in his published study of the genus that he failed to understand the group. 294 Rhodora [Vol. 74 Cyperus Houghtonii Torr. Sandy roadside by pond near Silver Lake, Hollis, Hillsboro County, A. R. Hodgdon and Frederic L. Steele, August 25, 1971 #19144. This rare species is not new to New Hampshire, having previously been collected in Bath and Hinsdale in the Connecticut Valley. Its distribution in New England is somewhat un- usual, however, and therefore deserves comment. We found it growing along with Cyperus strigosus, Fimbristylis autumnalis, Bulbostylis capillaris and Panicum philadel- phicum at the edge of a road near a small pond not far from Silver Lake in Hollis. It was difficult to sort out the depauperate plants of Cyperus from the other plants with which it grew, indicating that it may be more common in suitable situations, obscured at times by associated plants, than it now seems. Its New England range includes Win- slow (Kennebec Valley), Maine where it was discovered by Ralph Bean on July 22, 1915, Bath and Hinsdale, New Hampshire, Chittenden and Rutland Counties in western Vermont and several scattered localities in Massachusetts, there being no records reported by Seymour (1969) from Connecticut and Rhode Island. Specimens of the four principal species discussed in this article have been deposited in the herbaria of the Univer- sity of New Hampshire and the New England Botanical Club. LITERATURE CITED Hopepon, A. R. and STANLEY KROCHMAL. 1950. Range — extensions and clarifications in New Hampshire. Rhodora 52: 162-163. SEYMOUR, FRANK C. 1969. The Flora of New England. Charles E. Tuttle Co., Rutland, Vermont. 596 p. DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM 03824 and ST. MARY’S-IN-THE-MOUNTAINS LITTLETON, NEW HAMPSHIRE SOME ALGAL INVADERS OF THE NORTHWESTERN FRINGES OF THE SARGASSO SEA WILLIAM J. WOELKERLING The Sargasso Sea encompasses a wide but not sharply de- limited area (biologically speaking) of the North Atlantic Ocean situated roughly between 20 and 40 degrees north latitude and 35 and 75 degrees west longitude (Marmer 1928) and owes its name to the presence of two species of brown algae: Sargassum fluitans Boergesen and S. natans (L.) J. Meyen. On a recent cruise (during May 1970), the author observed numerous plants of Ascophyllum nodosum (L.) LeJolis and Fucus vesiculosus L. adrift with Sargas- sum along the northwestern fringes of the Sargasso Sea. The area visited includes the region from 69°24’ west long- itude to 69°48’ west longitude and 38°53’ north latitude to 39°11’ north latitude (ca. 375 square miles) and lies rough- ly 125 miles south of Nantucket Island, 150 miles south of Cape Cod, Massachusetts, 170 miles east-southeast of Mon- tauk Point, Long Island, and 75 miles north of the Gulf Stream. The occurrence of macroscopic algae within the region appeared to be sporadic; densities as high as one plant per square meter were encountered in some places while at others virtually no weed was observed. Entangled masses of Aescophyllum and Fucus similar to those reported for Sargassum (Winge 1923) were not observed. Samples taken at several stations with the aid of a long- handled dip net revealed both epiphytized and unepiphytized plants of Ascophyllum and Fucus. In all, 14 species of al- gae were collected (Table 1) including 2 Chlorophyta, 10 Phaeophyta, and 2 Rhodophyta. None of the epiphytic spe- cies has been recorded from the Sargasso Sea previously although isolated plants of Ascophyllum and Fucus have been reported (Collins 1917, Collins and Hervey 1917, Winge 1923). Many individuals of Ascophyllum and Fucus 295 296 Rhodora [Voi. 74 examined appeared fresh and may still have been growing at the time of collection, but others bore very distended re- ceptacles and showed signs of vegetative decay. Most of the epiphytes appeared somewhat moribund and lighter in color than their mainland counterparts and probably were not actively growing. These observations suggest that at least the epiphytized plants encountered represent detached specimens which have drifted out from the east coast of the United States rather than true pelagic forms. Both Ascophyllum and Fucus, however, are reported to propagate vegetatively in the drifting state in European waters (Levring 1940, Oltmanns 1889, Reinke 1892, Sauva- geau in Collins and Hervey 1971, p. 78) and further study appears warranted to determine whether reproduction of any sort similarly occurs in the northern Sargasso Sea. The presence of apparently healthy, unepiphytized plants of As- cophllum and Fucus in the region visited certainly suggests the possibility of vegetative propagation. In fact, Collins and Hervey (1917, p. 79) have concluded (without experi- mental data) that Ascophyllum grows actively and occasion- ally reproduces sexually in the Sargasso Sea. Until more direct evidence comes to hand, however, it seems best to re- gard Ascophyllum, Fucus and particularly their associated epiphytes as invaders rather than permanent components of the Sargasso Sea flora. Sincere thanks are due Gordon Volkmann of the Woods Hole Oceanographic Institution for arranging passage on the WHOI research vessel ATLANTIS II. One of the prepared sets of herbarium specimens has been retained by the au- thor while duplicates have been deposited in the Farlow Herbarium of Harvard University and the University of California Herbarium at Berkeley. The work described here was supported by Grant GB-13250 from the National Science Foundation to the Systematics-Ecology Program. SEP Contribution No. 231. 1972] Algal Invaders — Woelkerling 297 REFERENCES COLLINS, F. S. 1917. The Sargasso Sea. Rhodora 19: 77-84. COLLINS, F. S. and A. B. HERVEY. 1917. The algae of Bermuda. Proc, Am. Acad. Arts. Sci. 53(1): 1-195. LEvRING, T. 1940. “Studien über die Algenvegetation von Blekinge, Südschweden". Dissertation. Lund. MARMER, H. A. 1928. The Sargasso Sea. U. S. Naval Inst. Proc. 54(1): 33-35. OLTMANNS, F. 1889. Beiträge zur kenntnis der Fucaceen. Bibl. Bot. 14: 1-94, pl. 1-15. REINKE, J. 1892. Uber giste der Ostseeflora. Ber. dt. bot. Ges. 10: 4-12. WINGE, O. 1923. The Sargasso Sea, its boundaries and vegetation. Rep. Dan. oceanogr. Exped. 1908-1910 3(2): 1-34. TABLE 1. Algae Collected Along Northwestern Fringes of Sargasso Sea, May 1970. SPECIES REMARKS Chlorophyta : Monostroma pulchrum On Fucus Farlow Spovgomorpha arcta (Dillwyn) Kuetzing On Fucus Phaeophyta : Ascophyllum nodosum Specimens up to 1.0 m long (L.) LeJolis collected Chordaria flagelliformis Immature; on Fucus (Müller) C. Ag. Elachistea lubrica On Ascophyllum and Fucus Rupr. 298 Rhodora [Vol. 74 Fucus vesiculosus L. Specimens up to .5 m long collected Isthomoplea sphaerophora On Polysiphonia lanosa in (Harvey in Hooker) turn on Ascophyllum Kjellman Punctaria latifolia Grev. On Fucus Punctaria plantaginea On Fucus (Roth) Grev. Pylaiella littoralis L. On Fucus Kjellman Sargassum natans L. Pelagic J. Meyen Scytosiphon lomentaria On Fucus (Lyngbye) C. Ag. Rhodophyta : Erythrotrichia carnea On Fucus (Dillwyn.) J. Ag. Polysiphonia lanosa L. On Ascophyllum Tandy DEPARTMENT OF BOTANY THE UNIVERSITY OF WISCONSIN MADISON, WISCONSIN 53706 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, particularly 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- atics 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 74, No. 798. including pages 161-299. was issued June 30, 1972 299 CONTENTS: — continued A White-Flowered Form of Iris lacustris from Ontario. J. E. Cruise and P. M. Catling crsscccsccccssssseeessecsesseereesennenens 271 Neomirandea allenii, A New Epiphytic Composite of the American Rain Forest. R. M. King and H. Robinson .sssesssrrrereserererereereererererereees 272 Studies in the Guttiferae. III. An Evaluation of Some Putative Spontaneous Garden Hybrids in Hypericum Sect. Myriandra. Preston Adams .........emnm 276 A New Species of the Orchid Genus Hoffmannseggella. H. G. JONCS ceccccssscecccsssssccecscessecseessssssscessseeeeeeeeenaeesssseseeegeceeesees 283 Chromosome Number in Liquidambar. Frank S. Santamowr, J.. eene nnne nnn nennen 287 Recent Discoveries in the New Hampshire Flora. A. R. Hodgdon and Frederic L. Steele .........../:2 291 Some Algal Invaders of the Northwestern Fringes of the Sargasso Sea. William J. Woelkerling |... eene 295 Instructions to Contributors. .............. eene 299 Hodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON > Associate Editors RADCLIFFE BARNES PIKE ELIZABETH ANNE SHAW STEPHEN ALAN SPONGBERG Vol. 74 September, 1972 No. 799 CONTENTS: Intraspecific Variation in Chromosomes of Sedum in the Southwestern United States. D IL de EST RR The Rachilla in Cape Cod Species of Carex with Notes on the History of the Perigynium and Rachilla. Henry K. Svensom verccsssssssssssssssrsrssesevsrssssstssssesssesssavscarasasscessesee Distribution of Native and Introduced Knapweeds (Cen- taurea) in Canada and the United States. EE NN E TNE E AEA o E E D A New Variety of Leucophyllum laevigatum (Scrophu- lariaceae) from Mexico. Robert W. Kiger wcscscssssssssscsssssssssssssssssscsssesesessssassessecesssssssessees (Continued on Inside Back Cover) The Nem England Botanical Club, Bue. 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 double-spaced throughout. Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver’s blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Albion R. Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire 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 Rbodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 74 September, 1972 No. 799 INTRASPECIFIC VARIATION IN CHROMOSOMES OF SEDUM IN THE SOUTHWESTERN UNITED STATES CHARLES H. UHL Many species of Sedum include plants with different chromosome numbers (Uhl, 1961, 1970b). This intraspe- cific variation in number occurs in two patterns, here desig- nated as established variation and casual variation. Estab- lished variation is represented by clear-cut geographical or ecological chromosome races, where any population clearly has a standard chromosome number, but populations in other areas or habitats may differ in the number of chromo- somes in their sets (dysploidy) or in their number of chromosome sets (polyploidy). Casual variation is more random and accidental and is represented by individual plants that depart from the locally established chromosome race in ways that usually reflect some cytological instabil- ity or irregularity. The commonest examples are trisomy (2n + 1) and the presence of a variable number of small, extra, accessory, or “B” chromosomes. Each established variant in chromosome number has a balanced, more or less stable and true-breeding outfit of chromosomes that has been tested and proved and that oc- cupies a geographical or ecological portion of the range of the species. Most casual variants, on the other hand, have an unstable outfit of chromosomes that does not breed true to its own chromosomal type. Casual variants repre- sent a sort of cytological “noise” or "static", and where 501 302 Rhodora [Vol. 74 they are common they may sometimes blur or obscure the pattern of established variation. They may originate as segregants from hybrids with related species or between different established variants (chromosome races), or their occurrence may simply indicate that the population has not yet evolved to chromosomal stability. Doubtless each estab- lished variant first arose as a casual variant that was suc- cessful and eventually stabilized, and this process must still be going on. A relatively high background of casual variation might be expected in species with a significant amount of established variation. On the one hand, this reflects the continuation of the processes (instabilities) that originally led to the chromosome races (established variants), and on the other hand it may result from the oc- currence of hybrids where the isolation was not yet com- plete. The species of Sedum in the southwestern United States offer good examples of both types of chromosomal varia- tion. This paper describes the chromosomes and the dis- tribution of the chromosome races in five of these species: S. cockerellii Britton (including S. griffiihsii Rose), S. niveum Davidson, S. stelliforme S. Wats., S. wrightii A. Gray, and S. havardii Rose. Probably all of these species, as well as other related species, occur in adjacent areas of northern Mexico, but material from there has been limited, except for S. wrightii. Living plants of these species are easy enough to identify, but many specimens of them are incorrectly identified in herbaria. In particular S. cocker- ellii, S. stelliforme and S. wrightii have been confused, and ranges given for these species in some Floras cannot be depended upon. Most plants were collected in the field but grown in cul- tivation for a time before study. Counts were made from conventional] aceto-carmine squash preparations of pollen mother cells. The photographs are from permanent prep- arations, all X 2000. Voucher specimens are in the Wie- gand Herbarium or Bailey Hortorium of Cornell Univer- sity. 1972] Sedum — Uhl 303 Clausen and Uhl (1943) classified S. cockerellii and S. griffithsii, as then known, into separate subsections, largely on the basis of a difference in chromosome number »—16 and n=—14 or 29, respectively). However, the two are very closely related, and in my opinion the additional chromo- somal and morphological variation known today make it impractical to separate them. Therefore S. cockerellii is conceived broadly here, to include also S. griffithsii and the various other binomials earlier considered synonymous with one or the other of them (Clausen and Uhl, 1943). Thus conceived, S. cockerellii has at least four chromo- some races (established variants), with 14, 16, 29 or 30 gametic chromosomes. Probably the lone Texas population, with n=15, represents a fifth chromosome race. Each of these races occupies a definite geographic area, and in no case was more than one chromosome race found in the same range of mountains (Fig. 29). One or more extra chromo- somes were found in plants from six widely separated lo- calities. These casual variants occurred in populations in which the standard numbers (as confirmed by counts from other plants in four of the six populations) were «—14 (2 populations), n=15 (1 population), and n=16 (3 pop- ulations). In all but one case the extra chromosomes were smaller than the regular members of the set; they some- times differed among themselves in size (e.g., Fig. 4) and occasionally paired with each other, although usually not with members of the standard set. Most of them seem to represent typical examples of accessory or “B” chromo- somes. Their origin and function, if any, are completely unknown. In general, plants of S. cockerellii from south central Arizona and northern Sonora have 14 pairs of chromo- somes (Fig. 1). These include typical S. griffithsii (C5176). One plant from the Huachuca Mountains (U887A) had two small extra chromosomes that formed a bivalent in 7 of the 23 metaphase I plates examined (Fig. 2) but remained unpaired in the remaining 16 in the same anther (Fig. 3). Another plant from the same population (U887B) had the 304 Rhodora [Vol. 74 standard 14 pairs. Still another plant, from Sycamore Canyon, southeast of Ruby (UC65.135), had three small, extra chromosomes (Fig. 4), but six other plants collected at various times from the same locality (U884, U1824) all had the standard 14 pairs. The lone Texas population, which I believe is a new spe- cies record for the state, has n=15 (Fig. 5), although one of the four plants studied had a small, extra chromosome, possibly an accessory (Fig. 6). One pair of chromosomes is large, perhaps the result of combining most of two chromosomes from an ancestor with n=—16. Plants of S. cockerellii ranging from Oak Creek Canyon in north central Arizona to the White Mountains in the east central part of the state, mostly along or near the Mogollon Rim, as well as most collections from New Mex- ico, have n—16 (Fig. 7). These include plants from the vicinity of the type locality (U1315, U1316, C5254). Con- strictions, apparently kinetochores, are detectable in some chromosomes of mitotic metaphases (2n—32, Fig. 8). Thus there is no reason to believe that diffuse or multiple kine- tochores are related to the dysploidy in this species. A plant from the White Mountains of east centra] Arizona had a small accessory chromosome (U1155, Fig. 9), but plants from six other localities in the same mountains all had the standard 16 pairs. One plant from the Jemez Mountains of north central New Mexico had a small ac- cessory chromosome (U1147G) and another from the same locality (C5281) had several (six in Fig. 10), but seven other plants from the same population had the standard 16 pairs (U1147A, B, C, D, E, F, H). Still another plant, from the Sangre de Cristo Range in northern New Mex- ico, had probably four extra chromosomes (U1133, Fig. 11). Since these were large enough to represent standard chromosomes, and since they appeared often to enter into multivalent formation with regular chromosomes (Fig. 12), the plant is possibly a quadruple trisomic (2n+4), per- haps descended from a triploid. In the Mule Mountains (near Bisbee), in the Chiricahuas, 1972] Sedum — Uhl 305 on Mount Graham, and on Kitt Peak in the Quinlan Moun- tains, all in southern Arizona, S. cockerellii has n=29 (Fig. 13). Plants from the Sierra Blanca Range in south- ern New Mexico have n—30 (Fig. 14). In these two tetra- ploid chromosome races univalents and multivalents are formed occasionally and make it difficult to detect any cas- ual variation in chromosome number. None was noted in the 18 tetraploid collections studied. S. niveum is closely related to S. cockerellii (Clausen and Uhl, 1943). High in the San Bernardino and Santa Rosa Mountains of southern California this endemic spe- cies has n—16 (Fig. 15), as does one race of S. cockerelli. S. niveum has recently been discovered about 200 miles to the south, high in the Sierra San Pedro Martir of north- ern Baja California (Moran, 1969). Here it is octoploid, with n—64 (Fig. 16), but occasionally forming a few mul- tivalents and univalents at meiosis. At one of its two type localities, Carr Peak in the Hua- chuca Mountains of southeastern Arizona, S. stelliforme has n=20, plus a variable number of accessory chromo- somes, two to seven in different plants, (Figs. 17, 18). An- other plant, probably this species, from more than 400 miles to the southeast in southern Chihuahua, Mexico, also had »—20 (Fig. 19), plus one or two extra, accessory chromosomes in some cells. Very similar plants from the White Mountains of east central Arizona and from the Black Range in southwestern New Mexico are considered for the present to be also the same species, but they all have n=12 (Fig. 20). The cause and significance of the two different, apparently unrelated, chromosome numbers (established variants) in the species is not at all clear; perhaps closer morphological study may support their tax- onomic separation. One plant from the Black Range (U1625A) had 8 small accessory chromosomes (six are shown in Fig. 21), but two other plants from the same population and plants from seven other populations in the same mountains all had »—12. S. wrightii includes plants with four levels of ploidy 306 Rhodora [Vol. 74 based on z—12, diploid through octoploid (n=12, 24, 36, and 48; Figs. 22, 24-26). No plants with accessory chromo- somes or other casual variants were definitely noted, but one collection (UC52.1680) clearly had 13 pairs of chromo- somes (Fig. 23), none of which was distinctively smaller than the others. Four plants collected later from what was presumed to be the same locality (U879) all had the stand- ard diploid n=12. It is not certain whether the plant with n=13 is better considered another established variant or a casual variant. Univalents, multivalents, and close second- ary associations are common enough to make exact counts difficult in some hexaploids (n=36) and octoploids (n= 48). Probably these are autopolyploids. Autopolyploidy has been reported in S. ternatum (Baldwin, 1936; Uhl, 1970b), and, on the basis of evidence from chromosome pairing in hybrids, it appears to be common in the Mexican Crassu- laceae (Uhl, 1970a, and unpub.). The four principal chromosome types (established vari- ants) of S. wrightii are not as discrete geographically (Fig. 30) as those of S. cockerellii and S. stelliforme (Fig. 29). Only diploids were found in southern New Mexico, and they occur also in the Davis Mountains of western Texas and in the vicinity of Saltillo, Coahuila, Mexico. Tetraploids (n=24) were found at the presumed type lo- cality (U882, near El Paso), near Marathon, Texas, in the Figs. 1-20. Sedum chromosomes in pollen mother cells at meta- phase I (except Figs. 8, 16), 32000. Arrows identify univalents and/or accessory (“B”) chromosomes. Figs. 1-14. S. cockerelli. Fig. 1. U883, »—14. Fig. 2. U887A, n=15. Fig. 3. U887A, n—14-- 2, Fig. 4. UC65.135, w—14--3B. Fig. 5. U880C, w—15. Fig. 6. U880B, »—15-4-1B. Fig. 7. U900, »—16. Fig. 8. U900, mitosis in anther wall, 25—32. Note constrictions (kinetochores) in some chromosomes. Fig. 9. U1155, w—16--1B. Fig. 10. C5281, n—16+ 6B. Fig. 11. U1133, »—16--4B. Fig. 12. U1133, 24, (identified by lines) --14&--2, Fig. 18. U888, »—29. Fig. 14. U1307, n—30. Figs. 15-16. S. niveum. Fig. 15. M345, n—16. Fig. 16. M15333, n— 64 (Metaphase II). Figs. 17-20. S. stelliforme. Fig. 17. U1680A, n—20--2B. Fig. 18. U1680C, n—20+7B. Fig. 19. U2051, n=20. Fig. 20. U1312, »—12. 1972] Sedum — Uhl 507 308 Rhodora [Vol. 74 Figs. 21-28. Sedum chromosomes in pollen mother cells at meta- phase I, X 2000. Arrows identify univalents and/or accessory (“B”) chromosomes. Fig. 21. S. stelliforme. U1625A, n—12--6B. Fig. 22-26. S. wrightii. Fig. 22. U879, n—12. Fig. 23. UC52.1680, »—13. Fig. 24. UC53.407, n—24. Fig. 25. U877, n—36. Fig. 26. M6317, »—48. Figs. 27-28. S. havardii. Fig. 27. UC52.1679, n—54, Fig. 28. U876, m—81. Sierra del Carmen (across the Rio Grande from Big Bend) and in the Sierra Madre Oriental in southern Nuevo Leon, Mexico. Hexaploids (»—36) were found at Big Bend and 135 miles to the east, near Del Rio, Texas, and octoploids (n—48) only in southern Nuevo Leon. S. havardii is apparently of Mexican affinities. It is known in the United States only in the Big Bend region of western Texas, where plants with 54 and 81 gametic chromosomes have been found (Figs. 27, 28). Presumably the basic karyotype of this species consists of 27 chromo- somes, but no collection with this number has yet been found. The only collection with n—81 (U876) was made only 11/2 miles from another (U875) with n=54. Thus it is not yet clear whether the plants with the different numbers really represent established chromosomal vari- 1972] Sedum — Uhl 309 1 S. cockerellit © n=14 ascen — — A nsi5 D n=!6 asa e n-29 D uU mn-3O mu di m S. stelliforme Dt . n=12 m by x n=20 n [ ] He E A x : Fig. 29. Distribution of chromosome races (established variants) of Sedum cockerellii (s. lat.) and S. stelliforme in New Mexico, Arizona and adjacent regions. 310 Rhodora [Vol. 74 nzl2 n=13 n=24 n=36 n=48 $ o NEW MEXICO -eonpo a TEXAS (2 Fig. 30. Distribution of chromosome races of Sedum wrightii in New Mexico, Texas and northeastern Mexico, 1972] Sedum — Uhl 311 ants (races) or only two samples from a large population containing casual variants in ploidy. Two of the most sim- ilar species are S. diffusum (x—19) of northeastern Mex- ico and S. alamosanum (x—18) of northwestern Mexico, both of which also have polyploid races. Plants of S. cockerellii with n=16 (U1690) have been found within inches of S. stelliforme with n—12 (U1691) in the White Mountains, and plants of S. griffithsii (here included in S. cockerellii) with n—14 on Carr Peak in the Huachuca Mountains (U1679) are no more than a mile from plants of S. stelliforme that have n—20 + accessories (U1680). At two localities on opposite sides of Red River Pass in the Sangre de Cristo Range of northern New Mex- ico, S. cockerellii with n=16 (U1134, U1138) was found within inches of the yellow-flowered, tetraploid S. lanceo- latum, also n=16 (U1135, U1139), with both species in flower together. This locality is near the northern limit of the range of S. cockerellii, near the southern limit for S. lanceolatum. S. cockerellii was never found close to S. wrightii, al- though perhaps the two might occur together in the Davis or Organ Mountains. S. cockerellii generally occurs in the Transition and Canadian Zones, nearly always near streams that flow all year, whereas S. wrightii is generally a plant of the warmer and drier Sonoran Zone (Clausen and Uhl, 1943). However, in the Sacramento Mountains about Clouderoft, New Mexico, diploid S. wrightii (n—12) oc- curs at higher elevations (four collections: 8000-8400’) than does S. cockerellii 20-30 miles north in the Sierra Blanca Range about Ruidoso (five collections: 7100-7600’). Each species was found in only the one range in this area and not found in the other. S. wrightii (hexaploid, U873) occurs with S. havardii (U875) in the Chisos Mountains of Big Bend National Park and with other species in Mex- ico. Each of the five species has two or more chromosome races (established variants). Polyploidy definitely occurs in four of the five, and it may also account in some degree 312 Rhodora [Vol. 74 for the difference between the races of S. stelliforme with n—12 and n=20 + B's. Extra chromosomes, usually ac- cessories, were found in some diploid plants in three spe- cies. Probably this sort of casual variation should be ex- pected where so much established variation also occurs. In the same region Graptopetalum rusbyi of Arizona also includes polyploids and variations from strict diploidy (Uhl, 1970a). Apparently quite a few structural changes have occurred in the chromosomes of these species. Structural rearrange- ments must have been involved in the evolution of the dif- ferent numbers (14, 15, 16) in diploid S. cockerellit. Chromosome bridges at anaphase, which usually indicate heterozygosity for an inversion, were not especially looked for, but they were common in two plants of this species (U887A and C5176). Differences in chromosome number interfere with free interchange of genetic material and thus they represent a means of reproductive isolation that may eventually lead to new taxonomic species. However, anointing each estab- lished variant (chromosome race) as a different taxonomic species, as some (e.g., Love, 1951) have favored for other groups, seems undesirable here. It would increase the num- ber of species reported on from five to at least 15, and many of the splinter species probably could not be distin- guished without cytological study. Thus it seems better to take a broad concept of these species. The chromosomal variants are considered to indicate that evolutionary changes are actively occurring in the five species, but the changes are considered not yet to have crossed the thresh- old of the practical, taxonomic species. More careful mor- phological study, such as that now being carried out by my colleague, Professor R. T. Clausen, may some day re- quire modification of this conclusion for one or more of the five species. The large proportion of chromosomal variants in these species finds a parallel in certain other species of Sedum in the United States, e.g., S. glaucophyllum and S. terna- 1972] Sedum — Uhl 313 tum (Uhl, 1970b), and also S. lanceolatum (Uhl, unpub.). However, it contrasts sharply with the situation in still others. For example, in 78 collections of S. spathulifolium, ranging from Vancouver Island to southern California and exhibiting considerable variation in such characters as leaf thickness, width, crowding and glaucosity, no established variants were found, and all plants but two had seemingly identical karyotypes with n—15 (Uhl, unpub.). The only two exceptions were both casual variants: a plant with n=14 in a population that also included a standard plant, and a triploid. Seventy-nine other collections, representing five other species related to S. spathulifolium, also all had n=15, with no variants, either established or casual. How- ever, one related species had n=30 and another n—45. It is not clear whether the relative scarcity of non-standard chromosome complements in these species means that they are produced less frequently or that they survive less well, or both. At any rate, the karyotype here has remained stable in number and probably also in form during the evolution of enough morphological diversity that six spe- cies are recognized and while spreading 1100 miles apart. This stability must reflect differences in the nature of the genetic and chromosomal changes that have contributed to evolution in S. spathulifolium and its relatives (subgenus Gormania), contrasted with those prevailing in most of the species described earlier. Summary Variation in chromosome number is found within each of five species of Sedum of the southwestern United States. Established variation is represented by polyploid and/or dysploid geographic chromosome races, two to five for each species. Casual variation, represented by the occurrence of occasional plants that are trisomic or that bear accessory chromosomes, was found in two, possibly three, species. Successful casual variants can become established variants (races). Although differences in chromosome number are accompanied by some degree of reproductive isolation, rec- 314 Rhodora [Vol. 74 ognition of each chromosomal variant as a different species does not seem appropriate in these actively evolving cases. Acknowledgement I thank those who have provided some of the plants re- ported on here, especially Dr. Reid Moran, of the Natural History Museum, San Diego (M nos.), and Mr. Paul Hutch- ison, formerly of the University of California Botanical Garden, Berkeley (UC nos.). Publication was assisted by the L. W. Sharp Fund of Cornell University. COLLECTIONS STUDIED S. cockerellii Britton Arizona: U894 n—14 Pima Co. 4 mi. N. of road summit, Mt. Lemmon, Santa Catalina Mts., 6240'. U893 n—14 Pima Co. 0.8 mi. below Mt. Lemmon Lodge, 7520’. UC52.1674 n—14 Same locality, (E. R. Blakely). U892 n—14 Pima Co., Bear Canyon, 15 mi. by road S. of Mt. Lemmon Lodge, 5480’. U883 m—14 Santa Cruz Co. Madera Canyon, Santa Rita Mts., 6400’. C5176 m—14 Santa Cruz Co. near summit, Baldy Peak, Santa Rita Mts., 8900’. ? Topotype of S. griffithsii Rose, (R. T. Clausen). U884 n=l4 (5 plants) Santa Cruz Co., Sycamore Canyon, 5 mi. S.E. of Ruby, Atasco Mts., 3960’. U1824 n=l4 Same locality, (A. M. Phillips III). UC65.135 n—14+3B Same locality, (D. M. Hutt). U885 n—14 (2 plants) Santa Cruz Co., Sonoita Cr., 3 mi. S.W. of Patagonia, 3920'. U886 «—14 Santa Cruz Co., Mile 16 N.E. of Nogales, 2.8 mi S.W. of Patagonia, 3900’. U887A m—14--2, Cochise Co. Ramsay Canyon, Huachuca Mts., 6000'. U887B n»—14 Same locality. U1681 14 Cochise Co., Reef Mine, Upper Carr Canyon, Hua- chuca Mts., 7100'. U1679 mn—14 Cochise Co., N. slope of Carr Peak, Huachuca Mts., 7700’. C5189 m—14 Cochise Co. Same locality, (R. T. Clausen). 1972] Sedum — Uhl 315 UC52.1678 n=l4 Cochise Co., Guadalupe Canyon, Guadalupe Mts., 4300’, (E. R. Blakely). Mexico: M14849 w—14 Sonora. Cerro Colorado, S.E. of Cananea, (C. Glass and D. Gold). Arizona: U900 n—16 Coconino Co, Oak Creek Canyon, 12 mi. N. of Sedona, 5600'. U1617 m—16 Coconino Co., Macks Crossing, E. Clear Cr., 15 mi. N.E. of Clints Well, 6260'. UC52.1673 n—16 Gila Co., N. peak of Four peaks Mt., Mazatzal Mts., 7645', (E. R. Blakely). U1688 m—16 Gila Co., Workman Creek Falls, 44 mi. N. of Miami, Sierra Ancha, 6600’. U1618 »—16 Apache Co. near dam, Hawley Lake, White Mts., 8200'. U1694 n—16 Apache Co., Snake Cr., 19 mi. E. of McNary, White Mts. U1158 m—16 Apache Co., 3.8 mi. N. of Greer, White Mts., 8400’. U1690 n»—16 Apache Co., 14 mi. W. of Alpine, White Mts., 8500'. U1157 m—16 Greenlee Co., 6.6 mi. S. of Alpine, 7950’. U1156 a—16 Greenlee Co., 10.3 mi. S. of Alpine, 7700’. U1155 m^—16--1B Greenlee Co., 2.7 mi. N. of Hannagan Meadows, White Mts., 8800'. New Mexico: U1152 5»—16 Catron Co., 10 1/2 mi. W. of Reserve, San Francisco Mts., 7200'. U1151 5—16 Catron Co. Tularcsa R., 1 mi. E. of Aragon, 6750’. U1150 n»—16 Catron Co. Tularosa R., 3 mi. E. of Aragon, 6900’. U1621 m»—16 Catron Co., Wall L., 9 mi. S. of Beavernead, Black Range, 6400’, U1619 m—16 Sierra Co. Poverty Can., 19 mi. N.W. of Winston, Black Range, 7500'. U1153 m—16 Grant Co., 4 mi. W. of Mule Cr., San Francisco Mts., 5800’. U1622 n—16 Grant Co. Black Canyon Forest Camp, 32 mi. N. of Mimbres, Black Range, 6770’. U1309 n=—16 Grant Co. 2.5 mi. N. of Pinos Altos, Pinos Altos Range, 6600’. U1310 n=16 Grant Co. Iron Cr. 6.6 mi. W. of Emory Pass, Black Range, 6800’. U1311 m—16 Grant Co., Iron Cr., 5.1 mi. W. of Emory Pass, 7200’. U1314 5—16 Torrance Co., Red Canyon Forest Camp, 7 mi. W. of Manzano, Manzano Mts., 7800'. 316 Rhodora [Vol. 74 U1148 »—16 Sandoval Co., Jemez R. at Soda Spring, 1.6 mi. N. of Jemez Springs, 6350’. C5281 n—16+3-6B Sandoval Co. 4 mi. N. of Battleship Rock, near LaCueva, Jemez Mts., 7700’, (R. T. Clausen). Ull47 n=16 (7 plants) »—16--1B (1 plant) Sandoval Co., La- Cueva, (Probably same locality as preceding). U1146 n—16 Sandoval Co. Jemez Falls, 7850'. U1145 n—16 Sandoval Co., Las Conchas Forest Camp, Jemez Mts., 8400'. U1149 a—16 Sandoval Co., Las Huertas Cr., 6 mi. S. of Placitas, Sandia Mts., 7830'. C4633b n—16 (2 plants) Sandoval Co., Ellis Brook, N. side Sandia Mts., (R. T. Clausen). C4633a n=16 Sandoval Co., Sandia Crest, Sandia Mts., 10,400’, (R. T. Clausen). U1315 n—16 Santa Fe Co., 6 mi. N.E. of Santa Fe, 7500’. U1316 n—16 Santa Fe Co., 9 mi. N.E. of Santa Fe, Black Canyon Forest Camp, 8300'. Ul1143 m—16 San Miguel Co., E. side Pecos R. at bridge 6.4 mi. N. of Pecos, 7200'. C5254 n—16 San Miguel Co., Pecos R., 5 mi. S. of Terrero, 7500’, (R. T. Clausen). Uli42 a—16 San Miguel Co. Gallinas Cr., 1/2 mi. W. of Mon- tezuma, 6750’. U1317 n=16 Rio Arriba Co. Rio Vallecitos, 5.5 mi. N. of La- Madera, Ortega Mts., 6900’. U1138 n=16 Taos Co., S. side Red River, 2.7 mi. E. of Questa, 7550’, U1137 n=16 Taos Co. S. side of Red River, 4.5 mi. W. of Red River village, 8150’. U1136 n=i6 Taos Co., Mouth of Mallette Canyon, village of Red River, 8650’. U1140 n—16 Taos Co., S. side Rio Pueblo at Comales Forest Camp, 7850’. U1141 n—16 Taos Co. N. side Rio Pueblo, 2.2 mi. N.W. of Tres Ritos, 8250'. U1134 a—16 Colfax Co., 2.6 mi. E. of Red River Pass, Sangre de Cristo Range, 8900'. U1133 n»—16--4 Colfax Co., Cimarron Cr., 5 mi. E. of Eagle Nest, 7850’. U1132 n=16 Colfax Co., Cimarron Canyon, 5 mi. W. of Ute Park, 7750’. U1131 n=16 Colfax Co, Cimarron Cr. 1 mi. W. of Ute Park, 7550’. 1972] Sedum — Uhl 317 Texas: U880 "—15 (2 plants) n=15+1 (1 plant) Jeff Davis Co., Ma- dera Canyon, 25 mi. W.NW. of Fort Davis, Davis Mts., 5800’. 1961. U1633 m—15 Same locality, 1967. Arizona: U1685 752—329 1.6 mi. SE of Columbine Ranger Sta., Mt. Graham, 9215’. U1684 m—29 Graham Co., Grant Creek, Swift Trail, Mt. Graham, 8750’. U1683 n=29 Graham Co., Near Snow Flat, Swift Trail, Mt. Gra- ham, 9000’. U1686 n=29 Graham Co., 1.7 mi. N.W. of Ladybug Saddle, Swift Trail Mt. Graham, 8800’. U1682 mn—29 Graham Co. 1 mi. below Turkey Flat, Swift Trail, Mt. Graham, 7300’. U1926 ^—29 Graham Co. Marijilda Canyon, Mt. Graham, 8200', (A. M. Phillips III). U1927 m—29 Graham Co., Marijilda Canyon, Mt. Graham, 4800', (A. M. Phillips III). C5224 n=29 Cochise Co. S. of Rustler Park, Chiricahua Mts., 8500', (R. T. Clausen). UC65.144 n=29 Cochise Co., 1/2 mi. below Onion Saddle, Chiri- cahua Mts., 7300', (D. Hutt 1260). U1678 n=29 Cochise Co., S. Fork, Cave Cr, 3 mi. S.W. of Portal, Chiricahua Mts., 5150’. U1253 n=29 Cochise Co., Chiricahua Mts., (C. English). U888 5»—29 (2 plants) Cochise Co., 1/2 mi. N. of Mule Pass, Mule Mts., N. of Bisbee, 5840’. New Mexico: U1629 n=30 Lincoln Co., Bonito Cr., 7.2 mi. W. of Angus, Sierra Blanea, 7500’. U1628 n=30 Lincoln Co., 3.1 mi. W. of Angus, 7100’, U1307 m—30 Lincoln Co. Eagle Cr., 2.5 mi. W. of Alto, Sierra Blanca, 7600’, U1304 n=30 Same, 1.5 mi. W. of Alto, 7450’. U1630 m—30 Otero Co., Carrizo Cr., Mescalero Apache Indian Res., 2.2 mi. W. of Ruidoso, Sierra Blanca, 7200'. S. niveum Davidson California: C4759 n—16 San Bernardino Co., Sugar Loaf Peak, S. Bernardino Mts., 7000', (R. T. Clausen). C4759b »—16 Same, Dollar Lake, 9300', (R. T. Clausen). 318 U919 M345 M15346 M15333 M15682 Rhodora [Vol. 74 n—16 San Bernardino Co., above Jenks Lake, 8000’, (D. Verity). 1n—16 Riverside Co., Santa Rosa Peak, Santa Rosa Mts., (P. A. Munz). n—16 Riverside Co., 1 mi. N.W. of Toro Peak, Santa Rosa Mts., 7450', (R. Moran). Mexico: n—64 Baja California, Observatory Peak, Sierra San Pedro Martir, 9300', (R. Moran). n—64 prob. Same, N. slope, 8400'. (R. Moran). S. stelliforme S. Wats. U1680 C5192 U1693 U1689 U1691 U1692 U2051 U1623 U1624 U1625 Arizona: n—20+2-7B (5 plants) Cochise Co., N.E. slope Carr Peak, Huachucha Mts., 8300. TOPOTYPE n—20+2B Same Locality, (R. T. Clausen). n—12 Apache Co., N. side Crescent L., White Mts., 9100’. n—12 Apache Co., Cienega Redondo, 16 mi. W. of Alpine, White Mts., 8950’. n—12 Apache Co. 11/2 mi, W. of Three Forks, Black R., 14 mi. W. of Alpine, 8550’. n—12 Apache Co. Williams Valley, 7.2 mi. W. of Alpine, 8650’. Mexico: n—20--1 or 2B Chihuahua, 55 mi. W. of San Francisco del Oro, (M. Kimnach and F. K. Brandt 1058). New Mexico: n—12 Grant Co., 1.4 mi. S. of Black Canyon Forest Camp, Black Range, 7000’. »—12 Grant Co., 2.6 mi. S. of same, 7300’. n—12 (2 plants) Grant Co., 3.8 mi. N. of Rocky Canyon Forest Camp, Black Range, 7800'. U1625A n—12-4-8B Same locality. U1626 U1627 U1312 U1313 U1620 n—12 Grant Co., 1.2 mi. N. of same, 7600’. 1—12 Grant Co., 2.9 mi. S. of same, 7400’. n—12 Grant Co. 2.8 mi. W. of Emory Pass, Black Range, 7600’. n—12 Sierra Co., 1 mi. N. of Emory Pass, Black Range, 9000’. n—12 Sierra Co., N. side of Boiler Peak, 3.2 mi. W. of Continental Divide, Black Range, 7900’. S. wrightii A. Gray U1631 New Mexico: n—12 Otero Co., Silver Springs Canyon, 7.8 mi. N.E. of Clouderoft, Sacramento Mts., 8000’. 1972] Sedum — Uhl 319 U1303 n=12 Otero Co., Silver Springs Canyon, 5 mi. N.E. of Cloudcroft, 8400’. U1302 n=12 Otero Co., Cox Canyon, 5.2 mi. S.E. of Cloudcroft, 8150’. U1632 n=12 Otero Co. Upper Penasco Canyon, 13 1/2 mi. S. of Cloudcroft, 8400’. C5241 n=12 Dona Ana Co, Canyon at La Cueva, Organ Mts., (R. T. Clausen). Texas: U882 n—24 El Paso Co, McKelligon Canyon, N. of El Paso, S. side of Franklin Mts., 5400’, TOPOTYPE. C364 . n—12 Jeff Davis Co., Fern Canyon, Davis Mts. UC52.1680 n—13 Jeff Davis Co., 20 mi. N. of Alpine, Davis Mts., 4900', (E. R. Blakely 1378). U879 n—12 (4 plants) Jeff Davis Co. 5 mi. S. of Fort Davis, Davis Mts., 4800', (Same locality as preceding?). U878 «—12 Brewster Co., Sunny Glen, 6 mi. W.NW. of Alpine, 4800'. U1676 m—24 Brewster Co., 3.4 mi. S. of US 90 at Marathon, 4100'. U877, U1675 n=36 Brewster Co. Green Gulch, N. of Panther Gap, Chisos Mts., Big Bend National Park, 5600’. U873 n—36 Upper Boot Canyon, Chisos Mts., 6900’. UC52.1672 m—36 E. side of Emory Peak, Chisos Mts., 7000’, (E. R. Blakely 1402). C5298 »n—36 Chinese Wall, Chisos Mts., (R. T. Clausen). U868 n—36 Val Verde Co., E. side of Devils River, 8 mi. N.W. of Del Rio, 1100’. UC54.1334 n=36 Same locality, (J. B. Perry). Mexico: UC53.407 n—24 Coahuila, 8 mi. S.W. of Piedra Blanca, Sierra del Carmen, 7000’, (Ward Russell). U1497 m—12 Coahuila, Chorro Canyon, 27 km. S.E. of Saltillo, ca. 6500'. U1501 n=12 Coahuila, Chorro Canyon, 33 km. S.E. of Saltillo, ca. 1000'. M7825 m—12 Coahuila, Chorro Canyon, (R. Moran). M6317 n=48 Nuevo Leon, San Jose de Raices, (R. Moran). U1494 n=45-48 Nuevo Leon, 6 km. S. of San Jose de Raices, 6100’. U1916 n—48 Nuevo Leon, 3 km. W. of Iturbide, 5400’. U1914 n—24 Nuevo Leon, 16 mi. N. of Ascension, 7900’. U1912 n=?24 Nuevo Leon, 30 mi. N. of Doctor Arroyo, 4 mi. N. of La Escondida, 5900'. 320 Rhodora [Vol. 74 Sedum havardii Rose Texas: U875 n—54 Brewster Co. Chisos Mts., Big Bend National Park: N. of Boot Spring, 6900’. U876 n—81 11/4 mi. S. of Boot Spring, 7000’. UC52.1679 n—54 E. side of Emory Peak, 7000’, (E. R. Blakely). U1674 m—54 Above E. side of Panther Pass, N. side of Casa Grande, 6200'. C363 n=54 Green Gulch, 5500’. LITERATURE CITED BALDWIN, J. T., Jr. 1936. Polyploidy in Sedum ternatum. J. Hered. 27: 241-248. CLAUSEN, R. T., and C. H. UHL. 1943. Revision of Sedum cocker- ellii and related species. Brittonia 5: 33-46. Love, A. 1951. Taxonomical evaluation of polyploids. Caryologia 3: 263-284. Moran, R. 1969. Sedum in Baja California. J. Cactus & Succu- lent Soc. Am. 41: 20-25. UHL, C. H. 1961. Some cytotaxonomic problems in the Crassula- ceae. Evolution 15: 375-877. 1970a. Chromosomes of Graptopetalum and Thomp- sonella (Crassulaceae). Am. J. Bot. 57: 1115-1121. 1970b. Heteroploidy in Sedum glaucophyllum. Rho- dora 72: 460-479. DIVISION OF BIOLOGY CORNELL UNIVERSITY ITHACA, NEW YORK 14850 THE RACHILLA IN CAPE COD SPECIES OF CAREX WITH NOTES ON THE HISTORY OF THE PERIGYNIUM AND RACHILLA HENRY K. SVENSON This account takes up the local occurrence of the pecu- liar bristle-like or spikelet-bearing structure within the perigynium of Carex, known as the “rachilla”. There is a review of various opinions, especially those of Duval- Jouve, Holttum, and Raymond. There is also a discussion of the relationship of Cymophyllus to Carex and to the West Indian Bisboeckelera. A plate with many figures il- lustrates the various interpretations of the rachilla. It was a surprise to find on Cape Cod, in southeastern Massachusetts, Carex with development of the rachilla. First detected was Carex Emmonsii, which appeared pecu- liar to the naked eye. It grew in rich soil with skunk cabbage along the so-called Marston Mills River. The micro- scope showed that spikelets, developing within the perigy- nium, had extended out of it, a characteristic of Schoenozi- phium. This is a genus of sedges mainly South African. Further observation showed that this condition is not rare in Carex Emmonsii in dry sandy fields. C. Emmonsii is the commonest Carex on Cape Cod, and easily recognized by the tufts of long recurving leaves, which remain over winter. It grows especially well where the soil has been disturbed. The same rachilla structure is found, though less abundantly, in robust specimens of the closely related Caiex pensylvanica. In a sandy field in Osterville, both species, together with rachilla-bearing C. scoparia, grew in a field being overrun by Robinia viscosa, Prunus serotina, and Ewonymus europaea. Further good examples of C. pensylvanica were found in sandy fill near Johns Pond in Mashpee. This axis within the perigynium assumes various forms, most commonly as a small bristle, but also as the schoenoxi- phoid condition mentioned above (see fig. 7). On Cape Cod I have seen the rachilla in C. scoparia, C. Swanii, C. debilis 321 322 Rhodora [Vol. 74 1972] Carex — Svenson 323 var. Rudgei, C. bullata, C. canescens, C. Emmonsii and C. pensylvanica. There are undoubtedly more. The figure of C. complanata, collected by me in Tennessee, was drawn by Miss Purdy about 1940. It has a rachilla reminiscent of a hookless Uncinia. The rachilla in Carex was first noted by Kunth (1788- 1850), who correctly interpreted it. The problem is illus- trated in figs. 11-15, where rachillas are shown in the series Schoenoxiphium to Carex, based on figures from Kukenthal (1909), Das Pflanzenreich: Cyperaceae-Caricoideae; and from R. S. Snell (1936). Anatomy of the spikelets and flowers of Carex, Kobresia, and Uncinia, in Bull. Torrey Club 63: 277-295. In Carex, as in fig. 15, the rachilla is commonly reduced; but see fig. 4, C. complanata, and fig. 5, C. scoparia. There are many examples in Boott’s (1858), “Illustrations of the Genus Carex". In many of his illus- trations the rachilla is flat with a serrated margin. That of C. Fraseri is strap-shaped without serrations. That of S. scoparia (my fig. 5) is perhaps closest to C. Camposii (t. 574) from the Sierra Nevada of Spain, which Kueken- Figs. 1-17. The rachilla (except in 4) is black. Axis of spikelet, a; scale (glume, b; perigynium, c; achene, d. Fig. 1. Uncinia unci- noides, X 10, Hawaii, Svenson, Sept. 5, 1960; 2. style-base enlarged. 3. Rachlla of Carex tonsa, Ostervile, Mass., Svenson, June 10, 1967. 4. Carex complanata, X 10, Altamont, Tennessee, Svenson, Oct. 3, 19835 (drawn by Maud H. Purdy). 5. Carex scoparia, X 10, Osterville, Svenson 2933, June 23, 1970. 6. Carex pallescens, X 5, from Duval-Jouve, pl. vii, f. 8. 7. Carex Emmonsii, X 5, Svenson, June 16, 1967 (lowest scales removed). 8. Same collection as 7 (all scales removed). 9. Carex pallescens, X 10, from Duval-Jouve, pl. vii, f. 9 (perigynium bisected longitudinally). 10. Carex debilis var. Rudgei, Osterville, Svenson, Aug. 18, 1967. 11-15. Diagrammatic sections to show rachilla of 11, Schoenoxiphium; 12. Cobresia; 13. Uncinia; 14. Carex microglochin; 15. Carex (with reduced rachilla) ; all from Snell (1936) and Kuekenthal (1909). 16. Schoenoxiphium rufum, X 6, from C. B. Clarke, t. 141, f.2 17. The same, with peri- gynium removed, f. 3. Dissections and samples of material from Cape Cod are filed on a sheet in the cover of Carex Emmonsii from Barnstable County in the herbarium of the New England Botanica] Club. 324 Rhodora [Vol. 74 thal places in the section Elatae, not represented in Amer- ica. But none of Boott's plates illustrate a rachilla in the Ovales. By Barros (1935), Cyperaceae of Argentina, in Anales Mus. Ciencias Nat. Hist., the rachilla is shown in 4 species. The best account of these contradictory structures is by Duval-Jouve (1864): Sur la signification morphologique d'une anomalie qué présentent les utricules de quelques Carex, in Bull. Soc. Bot. France. 1864. 19 pp. 1 pl. Since the structures are similar to those that I have found, and since many species treated by him are also in New England, I have given his account the most extended treatment. In the spring of 1864, Duval-Jouve first found Carex palles- cens with peculiar compound spikes, and spikelets issuing from the perigynia. Altogether he records a list of 24 species with a rachilla, obtained from the vicinity of Arles and Strasbourg. The anomalous spikelets were found most frequently toward the base of the spike, sometimes in the middle (C. acuta, C. pseudocyperus, but only rarely toward the apex. In C. flacca?, C. Oederi, C. flava, and C. hirta, compound spikelets were so abundant that in some local- ities it was almost impossible to find a plant without them. In C. flacca the rachilla appeared most often as the Schoe- noxiphium-type. In C. flava the perigynia were often split, 'N. J. Duval-Jouve (1810-1883). For biography see Flahault, Bull. Soc. Bot. France. His work is typical of the best of French botanists. He is perhaps best known for his work, illustrated in color, of Equi- setum hiemale. Allis as though inlaid with gold. His work on Carex is equally as good. As a forerunner of botanical work he wrote a "Traite de Logique, ou Essai sur la theorie de la Science", Paris (1848), and a second edition, which I have seen, in 1885. It treats of various kinds of observations, perceptions, evidence. etc., things which often today tend to be forgotten. Surely no other botanist was so well prepared for his scientifie task. I did not, however, find a history of the transition of nomenclature from logie to biology, in which I was particularly interested. "Now a very abundant and variable species throughout the gypsi- ferous region (about Windsor), often growing in grassy pastures near the gypsum cliffs. Roland, Fl. Nova Scotia (1947), p. 230. 1972] Carex — Svenson 325 resembling Elyna spicata. All species were in places highly modified by pastures, ditches, animals, and in many other ways. They were best developed on terrain newly dis- turbed, or where traces of modification were apparent. On Cape Cod, Carex pallescens? is common in grassy places, but I have yet to find a specimen with a rachilla. The achene in Carex represents the lowest element of a reduced axillary inflorescence, and the achene occupies a basal lateral position at the lower part of the perigynium. This axis (the rachilla) when present is commonly bris- tle-ike, or it may have several flowers as in Schoenozi- phiwm. Presence and development of the rachilla and the origin of the perigynium have been the subject of intensive study — one of the unfilled crossword spaces in the Cypera- ceous organization. It is generally conceded that the perigy- nium consists of a single scale, and not the union of two scales. Torrey, in North American Cyperaceae (1838), p. 404, noted in a specimen of C. Willdenowii that the perigy- nium of the lowest flower enclosed the peduncle of a sep- arate staminate spike. The two remaining pistillate flowers had within the perigynium, in place of a staminate spike, a green squamaceous rudiment. “This curious plant, there- fore, seems in a manner to connect Schoenoxiphium N. ab E. with Uncinia, and to explain the nature of the setaceous body in the latter genus". Theodore Holm (1854-1932) provides a bibliography of 55 items on the perigynium, together with an excellent account of terminology, in Am. Journ. Sci. ser. II. 214- 220 (1896) entitled "The cladoprophyllum and anthopro- "The var. neogaea Fernald (Rhodora 44: 306. 1942) is supposed to be without a short beak at the apex of the perigynium; but Boott (1858, pp. 39, 40) says the beak may be present or absent, and he shows a beak in t. 450, illustrating a specimen from Rhode Island. Kuekenthal (1909, p. 433) describes the perigynium as erostrate ex- cept in var. glaberrima which has “utriculae interdum brevissime rostratae”. The American var. neogaea therefore does not appear to amount to much. On Cape Cod, and otherwise in Massachusetts as I have seen it, Carex pallescens does not have the appearance of a native species. 326 Rhodora [Vol. 74 phyllum in Carex”, and the accompanying plate shows schoenoxiphoid spikelets in C. crinita and C. Sprengelii. Holm was one of the ablest of American botanists, who combined precise taxonomy with morphology; he was also an excellent artist. One of the latest accounts is by Holt- tum: “The spikelet in Cyperaceae” in Bot. Review, vol 14 (no. 8) 525-541. 1948. He, likewise, reviews the history of the perigynium. Much of the interest in the subject comes from the fact that Carex is one of the world’s largest genera, and because it is abundant in the north temperate region. 267 species are listed in Gray Man. ed. 8; 793 spe- cies in Kuekenthal’s world-wide treatment (1909). The development of the perigynium is almost like a sec- ond evolution of the seed plants. The developing embryo has been protected within the perigynium. Dispersal of mature perigynia is by floating, flying, attachment to men or animals, or in many other ways. This protection has emancipated Carex from the tropics, which (according to Holttum and to Raymond) is its primitive home, and al- lowed its penetration into areas freed from the Pleistocene ice. Raymond (1959): Carices indochinenses necnon siamenses. Mém. Jard. Bot. Montreal, no. 53, mentions the Careyanae and Pandophyllae as perhaps the most primitive elements of Carex. The Careyanae were segregated by him from the Laziflorae, which are characteristic sedges of the Appalachian forest of southeastern United States. As Fernald says (Rhodora 5: 249. 1903) the genera of the Cariceae are not very good: “Besides the genus Carex the members of the Cariceae have been grouped by differ- ent modern authors into various ill-defined genera varying with the personal equation". But that is also true of other genera in the Cyperaceae. The presence of the rachilla in widely diverse species of Carex led Duval-Jouve to suspect that the rachilla might not be of phylogenetic significance, but merely atavistic remnants appearing sporadically. Thus the occurrence of the rachilla in many species of Primocarex led to the early idea that Primocarex was prim- 9 1972] Carex — Svenson 327 itive. It is interesting in this respect to compare the rachilla of C. complanata (fig. 4) with Uncinia (fig. 1). Of interest is the lack of an articulation or joint in the style of the Vesiculares (in which the perigynium is highly adapted for floating), and in the Ovales* (in which it is wind-distributed). The Ovales have their great distribu- tion on the Atlantic coastal plain and in Pacific North America, cf. chart by Kuekenthal (1909) p. 14, and the recent fine treatment by Hermann (1970): “Carices of the Rocky Mountains and Colorado Basin". Nelmes: “Facts and Speculations on phylogeny in the tribe Cariceae" in Kew Bull: (1961) 427-436, explained that some unispicate species were of uncinoid origin, and he distributed most of Kuekenthal's Primocarex variously in the genus. And to emphasize the ill-defined genera, Kuekenthal (1940) de- scribed Schoenoxiphium kobresioides from material col- lected by Van Steenis in Sumatra, and termed it the “Ver- bindungslied zwischen Schoenoxiphium und Kobresia". Holttum thought Mapania to be the most primitive genus. See also my comments on Mapania macrophylla in Bull. Torr. Club 75: 95. 1948, with fig. 3. The breakdown of the tubular translucent corolla provides us with a possible ori- gin of the perigonial bristles in the Cyperaceae. I was incor- rect in ascribing M. macrophylla to Santo Domingo as well as British Guiana. Meyer’s plants all came from the Es- sequibo River in British Guiana. Mackenzie set up Cymophyllus, based on Carex Fraseri. By Pursh it was considered a Mapania, but I believe it to be a true Carex, illustrated by Boott (t 484) with a strap- shaped rachilla. There is a reduction series in the West Indies from Mapania and Hypolytrum to Bisboeckelera (Hoppia) and Calyptrocarya, which simulates Carex; but I think it is an unrelated parallel series, as shown by the various illustrations in Clarke’s “Illustrations of Cypera- *The style is said to be jointed by Mackenzie, and is so illustrated in North American Cariceae (1940), but I have not found it so. See fig. 5). 328 Rhodora [Vol. 74 ceae" (1909). In my illustration of Mapania sylvatica, the type of the genus, in “Cyperaceae of Panama", the corolla segments are inrolled around the stamens. This I am sure is the correct interpretation, and the flower is a normal flower of the Cyperaceae, and not an assemblage of “‘par- tial inflorescences”. (Ann. Missouri Bot. Gard., 1943) Another interesting structure in Carex is the style-base, which is sometimes swollen and articulated. Duval-Jouve described C. oedipostyla, i.e. with a “swollen foot” in Mém. Acad. Sci et Lettres de Montpellier (1870), including a plate with many figures. The articulating style-base is wide- spread in Eleocharis and some other genera, but has little real significance in their delineation. It is one of the his- torical straws grasped by the drowning botanist. Some decades ago I thought that the swollen style-base in the Laxiflorae might be a transition to broadleaved Cyperus (Mariscus) of the tropical rain forest, and especially to Cyperus diffusus. The structure of the achene and prophyl- lum in Carex (including perhaps Schoenoxiphium), is closer to Cyperus than to any other genus. In visiting Nic- aragua last spring, I found Cyperus diffusus to be a road- side and plantation weed, and not a rain forest type. It is probably different in the Old World. Schoenoxiphium is generally considered to be the most primitive of the Caricoideae, and the accompanying figures from Kuekenthal, Snell, and Clarke show the ideas pre- sented. It is a small genus centering on South Africa, and extending to the mountains of Madagascar, southern Si- beria just north of Afghanistan (Krechetovich in Flora Tadzhikskoi USSR 2: 10-12. 1963), and to Sumatra. D. M. Napper, in Journ. East Afr. Hist. Soc. and Corydon Museum, Nairobi, 24 (no. 2); 1963, p. 12, says *predomi- nantly South African, about 15 species. The normal habi- tats are damp forests and wooded or open grasslands. It is unusual for Schoenoxiphium to occur at the higher alti- tudes (over 7,000 or 8,000 ft.) where Carex usually oc- curs”. 1972] Carex — Svenson 329 The Caricoideae include Schoenoxiphium Nees (1832), Kobresia® Willd. (1809), Elyna Schrader (1806), Uncinia Persoon (1807), and Carex? Linnaeus (1753). Kobresia has according to Kukenthal (1909) its origin and central point in the high mountains of central Asia from Turkestan to Central China. 14 of the species are peculiar to the Himalayas, especially the western part, and 5 to the high mountains of Central China. The few North American species are well treated (with photographs) by Dunan in Bull. Torrey Club 83: 192-195. 1956; and in K. simpliuscula rachillas are shown. According to Mackenzie, Kobresia extends from Newfoundland and Greenland to Colorado and Alaska. K. elachycarpa Fernald was de- scribed in 1897 from banks of the Aroostook River at Fort Fairfield in Maine, but later treated by him as Carex elachy- carpa, which goes to show how tenuous are the limits of these genera. Uncinia has been well treated by Kuekenthal (1909) and consists of 24 species mainly of the southern hemisphere from the Antarctic Islands through the South American Andes to Mexico and Jamaica. The remainder is in Aus- tralia, Tasmania, New Zealand, New Guinea, and the Hawaiian Islands. The illustrated specimen (fig. 2) was collected by me in the Hawaiian Islands. Carex microglochin Wahlenberg is illustrated in fig. 4. It extends from Greenland to Alaska, southward to Quebec and Colorado. It is also known in the Patagonian region and in Eurasia. In describing the occurrence in Newfound- land, Fernald in Rhodora 28: 61. 1926, says “It is one of the rarest and taxonomically most interesting of species, having the primitive rachilla, which is obsolete in most "According to Kuekenthal, named for the botanophile Paul de Cobres of Augsburg. It is sometimes spelled with a “C”. *Duval-Jouve (1864, footnote p. 16): The word Carex was used by Virgil, Columella, Calpurnia, etc., to designate hard, sharp, or cutting herbs . . . perhaps tall Juncus, Typha, or Sparganiwm, or perhaps dry thickets at the borders of pastures, i.e. “Frondibus hir- sutis et carice pastus acuta" thickets which the shepherds of Proven- cal still call carigues. 330 Rhodora [Vol. 74 northern Carices, projecting from the top of the perigy- nium; in other words, C. microglochin is an excessively local relic-species now known only in a few remote corners of the globe, and about as well placed in the Antarctic genus Uncinia as in the genus Carex, which abounds in the northern hemisphere . . . its prickly little fruit had obviously been spread by pedestrians with their high skin boots." Now, with all these contradictory views, and the ques- tionable relationship of the various elements, it is hoped that the reader will go out to collect some rachilla mate- rial for himself, and will then turn to Fernald's amusing story of Carex microglochin in the twenty-eighth volume of Rhodora. OSTERVILLE, MASSACHUSETTS 02655 DISTRIBUTION OF NATIVE AND INTRODUCED KNAPWEEDS (CENTAUREA) IN CANADA AND THE UNITED STATES! R. J. MOORE The genus Centaurea (family Compositae, tribe Car- dueae) is a large and taxonomically complex group of at least 500 species, centered in the Mediterranean region. The great majority of species are Eurasiatic but two spe- cies are native to the southwestern United States and a few others are native in northern South America. A number of Old World species are naturalized, to varying degrees, in Canada and the United States. During the course of studies of the species found in Canada, data concerning the genus in the United States have been brought together from the literature and from herbarium collections examined. Floras of various areas of the United States treat the species found in their range but there is no complete inventory of United States spe- cies. Moreover, there is obvious disagreement between au- thors regarding the nomenclature and taxonomy of certain species and uncertainty as to the occurrence of a few spe- cies in the country. Boivin (1966-67) has listed the spe- cies found in Canada and we have discussed the morphol- ogy and distribution of these species in a study of the tribe Cardueae (Moore and Frankton, 1972). In the following key is given the diagnostic characters of the taxa which have been reported or have been found to occur in Canada and the United States. Brief notes re- garding taxonomy, common synonymy and a summary of the distribution are added. The identification and distribution of Canadian plants have been verified from herbarium collections, notably those of the following institutions: Department of Agri- culture, Ottawa (DAO); National Museum of Canada (CAN); Montreal Botanic Garden (MTJB) ; Botanical In- stitute , University of Montreal (MT) ; University of Brit- ‘Contribution no. 892 of the Plant Research Institute. 331 332 Rhodora [Vol. 74 ish Columbia (UBC); British Columbia Provincial Mu- seum, Victoria (V). Information regarding United States species has been compiled from the commonly known re- cent floras, some of which are cited here, and from her- barium specimens examined in the course of studies of the Cardueae of Canada. Specimens have been borrowed from the Gray Herbarium of Harvard University (GH), New York Botanical Garden (NY), and the University of Ver- mont (VT) to resolve certain specific problems. Verifica- tion of United States records which appear to be credible has not been attempted. The various North American in- troductions have been compared with European specimens of the taxa in our herbarium (DAO). Specimens of all the taxa discussed here have been seen, but in the cases of a few rarer species (C. eriophora, C. diluta, C. nervosa) only Old World specimens were available for study. Since most of the species are adequately described in available floras, descriptive comments are here appended only in cases where there is disagreement with the con- cepts expressed by most authors or to emphasize points of distinction between species. The genus Centaurea is here treated sensu lato, as is the most common practice in North America. As indicated by synonymy, some of the species are often placed in segre- gate genera. DESCRIPTIVE CHARACTERS Some discussion of the characters used in the following key may be helpful. The phyllaries (involucral bracts) take many forms in the genus and are very useful as diag- nostic characters since they are present and easily exam- ined in all flowering specimens. The innermost row of phyllaries is frequently unlike the outer rows and descrip- tions here refer to the more conspicuous outer 2 or 3 rows, unless otherwise stated. An apical spine is considered to be a rigid pointed apical structure that is distinctly longer than any lateral points that may be present on the phyl- lary. Phyllary margins (as distinct from the phyllary 1972] Centaurea — R. J. Moore 333 apex) may also bear spines, which are pointed rigid struc- tures, unlike the wiry, more or less flexible marginal proc- esses seen in the condition described as pectinate, or the serrations of membranaceous or chartaceous margins. In many species the apical portion of the phyllary is expanded into a structure (appendage) that is distinctly broader than the phyllary base (blade). In our species the appen- dage is either pectinate or chartaceous and with entire or lacerate margins. The flower heads may consist wholly of perfect tubular florets, all similar in size, or of shorter, perfect central florets surrounded by a peripheral row of longer, sterile tubular florets, which simulate ray florets. In the key, these conditions are described as eradiate (all florets similar) or radiate (short central and longer periph- eral florets — actually a falsely-radiate condition). KEY TO SPECIES a. Outer and mid involucral phyllaries terminated by a distinct spine (very short in C. salmantica and C. di- luta) ; with or without marginal spines as well. (b) b. Marginal spines or processes absent. (c) c. Apical spines more than 5 mm long (usually 10- 25 mm). (d) d. Stems winged (decurrent leaf bases) ; flowers yellow (Sect. Mesocentron DC.) (e) e. Apical spines pinnately spinose (bearing several lateral spinules on the shaft); phyllaries arachnoid. (f) f. Apical spines 5-10 mm long; involucre about 15 mm high 1. C. melitensis f. Apical spines more than 10 mm long; involucre approx. 20 mm high ...... p ) 2. C. eriophora e. Apical spine palmately spinose (without spinules on the shaft but with 2-5 pairs of palmately arranged basal spinules) ; phyllaries glabrous. (g) g. Heads 35-40 mm high, involucre 20- 334 Rhodora [Vol. 74 25 mm high, 15 mm broad; tips of in- ner phyllaries not dilated; achenes approx. 4.5 X 3 mm; pappus dark brown ............. 3. C. sulphurea £ Heads 20 mm high, involucre 15 mm high, 10 mm broad; tips of inner phyl- laries chartaceous and dilated; achenes approx. 3 X 1.5 mm; pappus white . MEM" 4. C. solstitialis d. Stems not winged; flowers pink to purple (Sect. Calcitrapa Cass.). (h) h. Involucres longer than broad (length 12- 17 mm; L/W approx. 2.5) ; heads sessile or subsessile; pappus absent .......... Lecce eee eee eee eae 5. C. calcitrapa h. Involucre globular, about as long as broad (length 11-14 mm) ; heads pedicel- late (pedicels 10-50 mm long); pappus about 2 mm long ......... 6. C. iberica c. Apical spines 1-5 mm long (i) i. Apical spines 3-5 mm; phyllaries sericeous, margin dark pigmented .... 7. C. muricata i. Apical spines 1-2 mm; phyllaries glabrate, margins not pigmented. (j) j. Phyllary tips dark, terminated by a 1 mm Spine or sometimes merely mucronate; margin entire ......... 8. C. salmantica j. Phyllary tips not pigmented, with a 1-2 mm spine and a small chartaceous, pal- mate-lacerate appendage; margin narrow- ly scarious ... 9. C. diluta b. Marginal spines or processes (at least 4 pairs) pres- ent, in addition to the apical spine (Sect. Acrolo- phus Cass.) (k) k. Heads radiate (but peripheral florets only slight- ly larger than the inner florets) ; marginal proc- esses straight or curved and somewhat flexible, 1972] Centaurea — R. J. Moore 335 phyllary tip and margin pigmented, forming a dark V-shaped mark. (1) l. Involucres 6-8 mm broad, globose and rounded at the base; heads borne in corymbs or corymbose panicles, the branches not strik- ingly straight and stiff in appearance . uri AG c GA, zu 10. C. maculosa l. Involucre 4-6 mm broad, distinctly longer than broad and the base usually tapering; heads borne in panicles on stiffly diverging branches Sp ass i oH cs DE 11. C. paniculata Heads eradiate; marginal spines straight and stiff, phyllary margins not pigmented in a V- shaped marking but occasionally with a dark ir- regular apical mark. (m) m. Heads 14-16 mm high; achenes 2.3-2.5 mm long, pappus absent or short (1 mm) ; flowers usually white but sometimes yellow, pink or mauve no ui M ME 12. C. diffusa m. Heads 12-13 mm high; achenes 2.5-3 mm long; pappus present (to 2.5 mm); flowers pink or maüve — 10 5 b SIL Les VAS M a 4 13. C. virgata var. squarrosa a. Outer phyllaries not terminated by a conspicuous, stiff spine which is longer and more prominent than any marginal processes (margins pectinate, lacerate or en- tire) (n) n. Phyllaries bearing an apical appendage (an abrupt expansion of the phyllary blade, broader than the base of the blade) (o) o. Heads small (to 4 cm broad) (p) p. Involucres approximately as long as broad; phyllary appendages broad and completely covering the bases of adjacent phyllaries (q) q. Appendages brown and papery, with lac- erate margins ............ 14. C. jacea q. At least some of the appendages pectinate 336 Rhodora [Vol. 74 (wiry marginal processes), appendages black or dark brown (r) r. Apex of appendages extended into a very long, recurved process and with numerous long lateral processes, feath- er-like in appearance (s) S. Plants rarely branched, bases of upper leaves truncate and almost clasping; involucres 20-24 mm high; achenes 4-5 mm, pappus 2.5- 3 mm long .. . 15. C. nervosa Plants usually branched; leaf bases tapering; involucres about 15 mm long; achenes 3-3.5 mm long, pap- pus 1 mm . 16. C. austriaca r. Apex of appendages appressed and not extended into a long recurved tip (t) t. t. Heads eradiate; pappus present (short, 0.5 mm but fully devel- oped); appendages black, wiry processes long, 2-3 X width of the blade, completely covering the ad- jacent phyllaries ... 17. C. nigra Heads eradiate or radiate; pappus absent or vestigial (few short bristles) ; appendages brown (not dark brown) and variable from pectinate to chartaceous-lacerate, appendages broad, covering adja- cent phyllaries . .......... Lee eee 18. C. X pratensis p. Involucre longer than broad (L/W = 1.5); phyllary appendages smaller and not com- pletely obscuring adjacent phyllaries (u) u. Appendages black or dark brown, small and usually triangular, not concealing the contrasting green phyllary blades; rameal 1972] Centaurea — R. J. Moore 337 leaves narrow and smaller than the cauline leaves Durum cnt 19. C. nigrescens u. Appendages dark brown, to twice the width of the phyllary base and almost covering adjacent phyllary bases; rameal leaves not clearly differing in size from the cauline leaves ........ 20. C. dubia o. Heads large (over 4 cm broad) (v) v. Phyllary appendages chartaceous and pal- mately lacerate; heads eradiate, flowers yel- low .. i . 21. C. macrocephala v. Appendages pinnately pectinate; heads radi- ate, flowers purplish (w) w. Phyllaries brown, appendages bearing 8- 12 pairs of cilia .. ... 22. C. rothrockii w. Phyllaries straw-colored and sometimes purple-tinged, appendages bearing 3-8 pairs of cia. 0 23. C. americana n. Phyllaries lacking an apical appendage (x) x. Phyllary margins papery, either entire or lacerate but not pectinate (y) y. Phyllary margins essentially entire, not lacerate (z) z. Heads to 2 cm high, eradiate .. 24. C. repens z. Heads to 4 cm high, radiate . 25. C. moschata Phyllary margins regularly dentate-lacerate (aa) (Sect. Cyanus DC.) aa. Leaves elliptie, oblanceolate, 10 mm or more wide; heads over 3 cm broad ............ NIRE sesto ae Oe. DE MOntAanNa aa. DT linear, lanceolate, to 3 mm wide; heads to 3 cm broad ... 21. C. cyanus x. Phyllaries pectinate (with wiry marginal processes) (Sect. Lopholoma Cass.) (bb) bb. Heads eradiate; foliage hoary .... ..... . 28. C. cineraria bb. Hos radiate; tod glabrous s 29. C. scabiosa io; oe Kee sie CAR RAN tel. ie: 6 vw. dps ea WW eal ee MP. 338 1. Rhodora [Vol. 74 NOTES ON SPECIES AND THEIR DISTRIBUTION C. melitensis L. Maltese Centaury, Tocalote, Croix de Malte; common: U. S.— west coast, Ariz., N. M., Texas, established in the western states, occasional and perhaps not persisting in the central and eastern states (Steyermark, 1963; Fernald, 1950) ; Canada — Vancouver Is., B. C. C. eriophora L. Rare: U. S.— Cal. Only one 1909 collection from Los Angeles is reported (Howell, 1959). C. sulphurea Willd. (C. sicula of Cal. authors, accord- ing to Howell (1959) ; rare: U. S. — Cal. C. solstitialis L. Barnaby's Thistle, Yellow Star-this- tle, Centaurée du Solstice; common: U. S. — probably in all states, weedy in the southwest; Canada — sw Ont., Man., Sask. C. calcitrapa L. Purple Star-thistle, Caltrope; com- mon: U. S.— southeastern states to N. Y. (Fernald, 1950), west coast (Abrams & Ferris, 1960), Ariz. (Howell, 1959), Ill. (Jones & Fuller, 1955) ; Canada — sw Ont., Vancouver Is., B. C. Howell (1959) records that a similar plant, possibly to be called C. calcitrapoides L., had been collected in Riverside Co., Cal. We have a collection from Hum- boldt Co., Cal. (Tracy 16769 — DAO) which may be C. caleitrapoides, since a sparse pappus is present. This species differs from C. calcitrapa in the taller habit and the presence of a short pappus. C. iberica Trev. in Spreng. Iberian Star-thistle; rare?: U. S. — Cal. Howell (1959) reports this spe- cies as rare in 3 counties of Cal., first collected in 1924. Several west coast floras list it for Cal. only. This species is similar to C. calcitrapa and speci- mens identified as C. calcitrapa may be C. iberica. Two such cases have been revised: Soil Conservation Col- 1972] Centaurea — R. J. Moore 339 lection, Sept. 10, 1955, Converse Co., Wyo. — DAO; Jones s.n., July 13, 1946, Labette Co., Kans. — DAO). To our knowledge, C. iberica has not been reported in Kansas or Wyoming. Iberian Star-thistle may occur in states other than California, in ranges reported for C. calcitrapa. 7. C. muricata L. (Amberboa muricata (L.) DC., Vol- untarella muricata (L.) Benth. & Hook.) ; rare: U. S. — Santa Barbara, Cal. 8. C. salmantica L. (Microlonchus salmanticus (L.) DC., Mantisalea salmantica (L.) Briq. & Cavill.); rare: U. S. — Cal. (Howell, 1959), Ariz., (Kearney & Peeb- les, 1960) 9. C. diluta Ait. Rare: U. S. — Cal. 10. C. maculosa Lam. Spotted Knapweed, Centaurée; common: U. S. — all states except possibly the south- eastern; Canada — N. S., N. B., Que., Ont., B. C. The plant is a serious weed in some areas, forming infes- tations. This taxon is treated in many different ways by European authors. Some take up the prior name C. stoebe L., which may or may not apply. Others treat maculosa as a variety of C. paniculata or divide macu- losa into three subspecies, which by other authors, are raised to species. Among North American collections the ssp. micran- thos (Gmel.) Gugler is distinguished by smaller heads (involucre 10-11 mm high) and fewer (4-6 pairs) and shorter phyllary processes and by the black or brown pigmentation of the phyllary margin and processes, contrasting with the green blade. The ssp. rhenana (Bor. Gugler and ssp. maculosa have larger heads (involucre 11-14 mm) and 5-10 pairs of longer mar- ginal processes which are black — dark brown in rhenana but brown and partially white in the typical subspecies. 340 11. 12. 13. 14. 15. Rhodora [Vol. 74 C. paniculata L. Rare: Canada Vancouver Is., B. C. This species is similar to C. maculosa and there has been dispute whether these B. C. plants are paniculata or maculosa. Howell (1959) has restored C. paniculata to the B. C. Flora and his decision seems to be justi- fied. C. diffusa Lam. Tumble Knapweed; common: U. S. — apparently widespread in the north, above 40°, probably common in western states and occasional elsewhere; Canada — Alta., B. C. C. virgata Lam. var. squarrosa (Willd.) Boiss. (C. squarrosa Willd.) ; rare: U. S. — Cal, Utah (Howell, 1959). C. jacea L. Brown Knapweed; common: U. S. — widely reported for northeastern and west coast states, probably midwest also; Canada — Que., Ont., B. C. Specimens of C. pratensis (nigra X jacea) are fre- quently identified as C. jacea. Probably both of these occur within the range commonly stated for C. jacea. Phyllary appendages are brown, chartaceous and en- tire or subentire but not deeply lacerate as in the hy- brids. Heads are radiate. C. nervosa Willd. Not known to occur in Canada or U. S. This species has been reported only by Fernald (1950) from one collection (Burnham, July 30, 1916, margin of grain- field, east of lower falls, N. Beaver Creek, Vaughns, north of Hudson Falls, Washington Co., N. Y.— GH!). We have revised this specimen to C. austriaca. Plants of C. nervosa are 10-40 cm tall, usually un- branched, with a single head, the stem leafy to the top; lower leaves oblanceolate, base tapering, middle and upper leaves oblong, base truncate or shallowly clasping; involucre 20-24 mm high; phyllaries lanceo- late to 2 cm long, recurved, plumose, black or dark brown. 1972] Centaurea — R. J. Moore 341 16. C. austriaca Willd. (C. phiygia ssp. austriaca (Willd.) Gugler); rare: U. S. — N. Y., Vt., O. This species has been reported only by Seymour (1969) on the basis of a collection from Dorset, Ben- nington Co., Vt. (Gilbert s.n., Sept. 22, 1966 — VT!). We agree with this identification, This collection and that of Burnham (see C. nervosa) are from adjacent counties of New York and Vermont, from locations ap- proximately 25 miles apart. Gilbert notes that the plants had persisted near farm buildings for several years but did not seem to mature seed. However, it seems that the introduction has survived for half a century in this area. An additional collection, probably this species, from Ohio (Webb 1525, Sept. 18, 1921, Portage Co., O. — GH!) has been seen. These plants are 15-80 cm tall, branched and bear- ing many heads per plant; cauline leaves narrow, up- per leaves not clasping the stem; heads smaller than in C. nervosa, ovate, approximately 14 X 16 mm; in- ner phyllaries rounded at tip, middle and outer phyl- laries long and plumose, black or dark brown, recurved. 17. C. nigra L. Black Knapweed, Centaurée noire; (C. jacea var. nigra Briq. C. consimilis sensu Piper & Beattie, Flora Southeastern Washington, 1914) ; com- mon: U. S. — widespread in the northern states, prob- ably south to about 37° N; Canada — Nfld., P.E.I., N: SS N. Ba One. Ont, B. C. The type specimen of C. nigra has eradiate heads (Marsden-Jones & Turrill, 1954) and it seems that plants that are typical in other characters as well are always eradiate. Descriptions and illustrations in some floras indicate that C. nigra may have radiate or eradi- ate heads. The radiate plants are, in our opinion, C. X pratensis, a hybrid between jacea and nigra which is highly variable and sometimes closely approaches the parental species. In typical C. nigra the phyllary appendages are 342 18. 19. Rhodora [Vol. 74 black (not dark brown), the wiry cilia very long, al- most matted, covering adjacent phyllaries. A white- flowered form (C. nigra f. pallens Spenn) has been collected in N. S. and Que., Canada. C. X pratensis Thuill. Protean Knapweed; common: U. S. — probably throughout the ranges reported for C. nigra and C. jacea (northern states); Canada — N. S., Que., Ont., B. C. This is a hybrid between C. nigra and C. jacea and is apparently included in the description of C. nigra in some floras. Abrams & Ferris (1960) and Howell (1959) report the occurrence of the species on the west coast of U.S.A. Specimens from Mass., Me., N. Y., Pa., R. I., Vt., (GH, VT) have been examined in this study. This hybrid shows great variation in the form of the phyllary appendage: wiry and pectinate, dark brown but not black, cilia long and covering adjacent phyllaries; some appendages pectinate and others part- ly pectinate and partly chartaceous-lacerate and show- ing many intermediate conditions between pectinate (nigra character) and chartaceous (jacea character) ; sometimes all chartaceous and subentire. Thus the ap- pendage form may closely approach those of the typ- ical parent species but usually shows some evidence of an intermediate condition. Heads are usually radiate but may be eradiate in specimens which closely ap- proach C. nigra. C. nigrescens Willd. Short-fringed Knapweed; (C. vochinensis Bernh., C. transalpina Schleicher p.p.) ; common: U. S. — probably widespread in the north- ern states; Canada — N. S., Que., Ont., B. C. Kerner (1881, pp. 81-85) selected the type of C. nigrescens in Willdenow’s herbarium and found it to be the same as the later-described C. vochinensis. Many U. S. floras recognize C. vochinensis, some take up C. nigrescens and a few list both species. Appar- 1972] Centaurea — R. J. Moore 343 ently C. dubia is included. There is confusion in the application of these names. Collections of C. nigres- cens from Conn., D.C., Ind., Mo., N. J., N. Y., Neb., Ore., Pa., Va., Wisc., (DAO, GH, NY) and from the above-listed provinces of Canada have been seen. The heads separate C. nigrescens from the general- ly similar C. nigra. The involucre of C. nigrescens is longer than broad and the small, triangular, black, short-fringed appendages contrast strongly with the green phyllary blades which are not covered by the appendages. The upper (branch) leaves tend to be noticeably smaller than the lower leaves. Heads of this species and of C. dubia are usually eradiate. 20. C. dubia Suter (C. nigrescens Willd. p.p., C. transal- pina Schleicher s. str., C. jacea var. transalpina Briq., C. dubia ssp. eu-dubia (Suter) Gugler & Thellung, C. nigrescens var. dubia Hermann); common: U. S.— Mass., N. J., N. Y., O., Ore., Va., Vt. According to Cronquist (1955; Gleason & Cronquist, 1963) who is the only author to recognize C. dubia in North America, the species is widely established in northeastern United States and southeastern Canada and occasional in northwestern U.S.A. We have seen specimens (NY, GH) from the above states ; none from Canada are known. The Species is closely re- lated to C. nigrescens and it is difficult to separate some specimens. These plants are similar to C. nigrescens but the phyllary appendages have longer cilia which almost, and sometimes completely, cover the adjacent phyllary bases. There is no distinct difference in size between upper and lower leaves and the latter may tend to be broader. 21. C. macrocephala Puschk. in Willd. Rare: Canada — Que. a casual adventive from cultivation, probably not persisting (Moore & Frankton, 1972). 344 22. 23. 24. 26. 27. 28. 29. Rhodora [Vol. 74 C. rothrockii Greenm. Native species: U. S.:— sw N. M., se Ariz. C. americana Nutt. Native species: U. S. — Ark. & Ia. to Ariz., occasionally an escape from cultivation in the northeast. C. repens L. Russian Knapweed; (C. picris Pall., Acroptilon repens (L.) DC., A. picris (Pall.) Boiss.) ; common: U. S.— reported in all states except the southeastern; Canada — Ont., Man., Sask., Alta., B. C. This species is frequently a serious weed in our area. There is cytological and morphological justifica- tion for segregating this species as Acroptilon repens. C. moschata L. Sweet Sultan; (Amberboa moschata (L.) DC.) ; rare: U. S. — northeast, Ill., Cal.; Canada — Vancouver Is., B. C. This garden escape seems to persist in the north- ern states but is never common. C. montana L. Mountain Bluet; rare: Canada — Nfld., N. B., Que., Ont., B. C. Gleason & Cronquist (1963) state that this species is a rare escape in the northern part of the range of their flora (northeastern states and adjacent Canada). C. cyanus L. Bluebottle, Cornflower, Barbeau, Casse lunette; (Leucacantha cyanus Nwd. & Lunnel, Cyanus segetum Bauh.) ; common: U. S. — probably all states : Canada — all provinces except Sask. This garden escape is reported in all regions. The variety denudata Suksdorf, reported for Wash., is a less pubescent variant. C. cineraria L. Dusty Miller; rare: U. S. — Cal. C. scabiosa L. Greater Centaurea, Hardheads, Cen- taurée scabieuse; rare: U. S.— O., Ia., Mont., Can- ada — Que., s. Ont. This escape is apparently not widely established; it 1972] Centaurea — R. J. Moore 345 is mentioned for only three states by several floras. The reported occurrence in N. B., Canada (Boivin, 1966-67) is an error due to a mislabelled specimen (Boivin, personal comm.). SUMMARY Twenty-six introduced and two native species of Cen- taurea are known to occur in the area Canada-United States. Sixteen of the adventive species are found in Can- ada and three of these in Canada alone. An additional ten introduced and two native species occur in the United States, making a total of 23 introduced and two native species in that country. Probably eleven of the introduced species can be classed as widespread: C. solstitialis, C. calcitrapa, C. maculosa, C. diffusa, C. jacea, C. nigra, C. X pratensis, C. repens, C. cyanus and perhaps C. nigrescens and C. dubia. With the exception of the last, these species are known from both countries and generally the ranges are continuous, although more extensive in the United States. A number of the rarer species are known only in the southwestern states, notably California. Five species (C. diffusa, C. jacea, C. maculosa, C. nigra, C. repens) are regarded as weeds in Canada (Frankton & Mulligan, 1970), C. repens being the most serious. Rus- sian Knapweed has been reported to cause sheep and horse poisoning in the Old World and C. repens and C. solstitialis have recently been shown (Young, Brown & Klinger, 1970) to cause Chewing Disease in horses in the western United States. ACKNOWLEGEMENTS Appreciation is expressed to the curators of herbaria who have made their collections available for study. LITERATURE CITED ABRAMS, L., & R. S. Ferris. 1960. Illustrated flora of the Pacific states. Vol. IV. Stanford University Press. 346 Rhodora [Vol. 74 Boivin, B. 1966-67. Enumeration des plantes du Canada. Natu- raliste canad. 93: 253-274; 371-437; 583-646; 989-1063. 94: 131- 157; 471-528; 625-655. CRONQUIST, A. 1955. Vascular plants of the Pacific Northwest. Part 5. University of Washington Press. FERNALD, M. L. 1950. Gray's Manual of Botany. Ed. 8. American Book Co. FRANKTON, C., & G. A. MULLIGAN. 1970. Weeds of Canada. Can- ada Dept. Agriculture Publ. 948. Queen's Printer, Ottawa. GLEASON, H. A., & A. CRONQUIST. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. Van Nos- trand. HowELL, J. T. 1959. Distributional data on weedy thistles in west- ern North America. Leaflets Western Botany 9: 17-29. Jones, G. N., & G. D. FULLER. 1955. Vascular plants of Illinois, University of Illinois Press. KEARNEY, K. H., & R. H. PEEBLES. 1960. Arizona Flora. Univer- sity California Press. KERNER, A. 1881. Schedae ad floram exsiccatam Austro-Hungari- cam. I. Vienna. Moore, R. J., & C. FRANKTON. 1972. Tribe Cardueae (Thistles) in Canada. Canada Dept. Agriculture (in press). MARSDEN-JONES, E. M., & W. B. TunmRILL. 1954. British Knap- weeds. Ray Society. London. SEYMOUR, F. G. 1969. The flora of New England. C. E. Tuttle Co., Rutland, Vt. STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univer- sity Press. YouwxG, S., W. W. Brown, & B. KLINGER. 1970. Nigropallidal en- cephalomacia in horses caused by ingestion of weeds of the genus Centaurea. J. Amer. Vet. Med. Assoc, 157: 1602-1605. PLANT RESEARCH INSTITUTE CANADA DEPARTMENT OF AGRICULTURE OTTAWA, ONTARIO K1A0C6 CANADA A NEW VARIETY OF LEUCOPHYLLUM LAEVIGATUM (SCROPHULARIACEAE) FROM MEXICO ROBERT W. KIGER In the course of field work in northern Mexico during the summer of 1971, we (J. L. Reveal, W. J. Hess and the author) collected Leucophyllum laevigatum Standl. from the Carneros Pass area of Coahuila. Our collection differs in several respects from the typical phase of the species as it is found in San Luis Potosi, Durango and Chihuahua. Aside from ours, the only other collection from Coahuila, based on a study of specimens at the National Museum of Natural History at the Smithsonian Institution and the herbarium of the Field Museum, is one made by Pringle in the same area in 1890. It also differs from the typical phase and resembles our collection closely. In view of the morphological differences found in the Coahuila specimens, their geographical separation from the rest of the known range of the species, and the evidence that the same geo- graphical and morphological correlation has existed over at least the past 80 years, a new variety of L. laevigatum is here proposed. This is the first intraspecific taxon to be described for the species, itself first described in 1924. Leucophyllum laevigatum Standl. var. coahuilense Kiger, var. nov. A var. laevigato foliis plerumque late oblanceolato- spathulatis, ad 16 mm longis ac 8 mm latis, frequente ac profunde emarginatis, nunquam acutis, saltem subtus per- sistenter stellato-tomentulosis, sepalis lineari-lanceolatis, corolla caerulea, 2-2.5 cm longa, tubo latius campanulato diagnoscenda. Low spreading shrub, the young branches densely white stellate-tomentulose with the pubescence thinning with age; leaves mostly broadly spatulate-oblanceolate, to 16 mm long and 8 mm wide, the apex rounded or more frequently markedly emarginate, never acute, often mucronulate, the 347 348 Rhodora [Vol. 74 young petioles densely stellate-tomentulose, the leaf-blades sparsely stellate-puberulent above and moderately so be- low when young, the pubescence persistent at least below ; pedicels 3-5 mm long, stellate-tomentulose ; calyx cleft near- ly to base, the sepals linear-lanceolate, to 4 mm long, acute, stellate-tomentulose outside, often glandular within and without; corolla blue, yellowish mottled low in tube, 2-2.5 cm long, campanulate, glabrous and semi-lustrous outside, the tube to 13 mm long, moderately to sparsely arachnose- pubescent inside, the limb unequally 5-lobed, the lobes obo- vate, the 3 larger broadly so, rounded at the apex, mod- erately villous-sericeous within, the margin cilate, the stamens didynamous, the filaments adnate to the corolla tube for 1/2 their length, to 7 mm and 8 mm long respec- tively, the stigma included in the corolla throat at anthesis about 1 mm below the mouth; mature capsules to 5 mm long. TYPE. MEXICO: Coahuila: Carneros Pass area, along Mex- ico Highway 54, 0.7 mile west of highway, about 0.5 mile south of Estación Carneros, 23 miles south of Saltillo, in low canyon associated with Yucca, Agave, Opuntia and Juniperus, 5 August 1971, Reveal, Hess & Kiger 2617. Holo- type US! Isotypes to be distributed from US. ADDITIONAL SPECIMENS EXAMINED. MEXICO: Coahuila: Carneros Pass area, Pringle 3460 (F, US). The typical variety of Leucophyllum laevigatum has leaves proportionately more narrowly spatulate-oblance- olate, to 20 mm long and 6 mm wide, more often rounded and less frequently and deeply emarginate, sometimes acute, and becoming glabrous or at least glabrate with age. The sepals are linear and usually 2-3 mm long, rarely reaching 4 mm in length. The most striking differences between the two varieties are associated with the corolla. In the typical variety, as in other species of Leucophyllum in Mexico, the corolla is purple to violet, whereas in var. coahuilense it is a definite blue. In the latter the corolla is consistently from 2 to 2.5 cm long at anthesis, most often 1972] Leucophyllum — Kiger 349 closer to 2.5 em, while in var. laevigatum it is from 1.3 to 2 cm long, usually about 1.5 cm. The tube is proportionate- ly longer in var. coahwilense and is more broadly cam- panulate than in the typical variety where it is narrowly so. Though capsules as long as those usually found in var. laevigatum (to 6 mm) have not been observed in the new variety, the difference is probably not significant since the specimens of the latter have only a few mature capsules. Too, while the capsules of the typical] variety reach 6 mm in length, they are often smaller and thus overlap the range observed for var. coahuilense. The type of Leucophyllum laevigatum var. laevigatum was collected by Nelson (4689) in Durango, Mexico, be- tween Ramos and Indé (holotype, Us!). Other specimens of this variety examined include — MEXICO: San Luís Potosí: Charcas, Lundell 5512. Durango: Huarichic, north- east of Pedricena, Pennell 18604; Indé, Reko 5207; 5 miles northeast of Yerbanis, Moran 6248; 10 miles north-north- west of Rodeo, Morley 630; 20 miles south of Zarca, Gen- try 8614. Chihuahua: near La Boquilla, Shreve 8072. All specimens examined are at US. Field studies in Mexico were supported by an NSF grant, GB-22645, to James L. Reveal for studies on the genus Eriogonum and the Intermountain Flora. DEPARTMENT OF BOTANY UNIVERSITY OF MARYLAND COLLEGE PARK, MARYLAND 20742 VARIATION WITHIN CALOCHORTUS VENUSTULUS (LILIACEAE) JAMES L. REVEAL AND WILLIAM J. HESS While collecting specimens of a new Hriogonum in the high mountains west of Hidalgo del Parral of northern Mexico, we made small collections of routine nature in order to better understand the local vegetation. In addi- tion to the new Eriogonum, we found an unknown Castil- leja which Dr. Noel Holmgren of the New York Botanical Garden will describe and a variant of Calochortus venus- tulus Greene which likewise is undescribed. Calochortus venustulus Greene var. imbricus Reveal & Hess, var. nov. A var. venustulo petalis 20-28 mm longis, pedunculis 10-15 cm longis, plantis 3-4.5(6) dm altis dif- fert. Stems 3-4.5(6) dm high; peduncles erect, slender, 10-15 em long; petals obovate, 2-2.8 cm long. TYPE. MEXICO: Durango: Sierra Madre Occidental, along the dirt road from Hidalgo del Parral toward El Vergel out of San Francisco del Oro, about 60 road miles west of Parral and 18.5 road miles west of Ojito, on forest floor and slopes near limestone outcrops, associated with Quercus reticulata, Pinus ponderosa and P. leiophylla, Arctostaphy- los, Arbutus and numerous other low shrubs and herbs, at about 8000 feet elevation, 11 August 1971, Reveal, Hess & Kiger 2741. Holotype, us! Isotypes to be distributed from US. The var. imbricus (from the Latin imbricus, rainy, al- luding to the rainy day that we discovered this plant) is a lovely addition to a genus already well noted for its beautiful members. Particularly, in the darkness of the rain and heavy clouds on that day, its large, golden flowers stood out in marked contrast from the dark greens and browns of the pine-oak forest floor. During our investigations of this taxon, we found that Calochortus venustulus, as defined by Ownbey (1940), should be further subdivided. The typical variety, var. venustulus, is common in the Sierra Madre Occidental west of Durango where it occurs 350 1972] Calochortus — Reveal and Hess 351 in the forests and dry meadows. However, to the north, west of Chihuahua and hence northward nearly to the United States, a second element is common in open plains and scattered oak forests. This phase was named C. mad- rensis by Watson just a few months after Greene proposed C. venustulus and has been placed in synonymy under the latter name. However, the northern population is consis- tently taller, with longer peduncles and shorter leaves in relationship to the inflorescence. This pattern of distribu- tion is somewhat similar to that found in Eriogonum atro- rubens Engelm. in Wisliz. (Reveal, 1967), and likely is expressed by other species as well. Thus, we feel the north- ern element should be recognized as C. venustulus var. madrensis (S. Wats.) Reveal & Hess, based on C. madren- sis S. Wats., Proc. Amer. Acad. Arts 23: 283. 1888. These variants may be distinguished as follows: A. Petals (10) 12-16 (18) mm long. (B) B. Plants 1-2 dm high; leaves mostly longer than the inflorescence; peduncles 3-6 cm long; Durango var. venustulus B. Plants (1.5) 2.5-4.5 dm high; leaves mostly as long to slightly shorter than the inflorescence; peduncles 5-8 cm long; Chihuahua var. madrensis A. Petals 20-28 mm long; plants 3-4.5 (6) dm high; leaves mostly shorter than the inflorescence; peduncles 10-15 em long; extreme northern Durango .. var. imbricus Field studies in Mexico were supported by an NSF grant, GB-22645 to Reveal for studies on the genus Eriogo- num and the Intermountain Flora. LITERATURE CITED OwNBEY, M. A. 1940. A monograph of the genus Calochortus. Ann. Missouri Bot. Gard, 27: 371-560. REVEAL, J. L. 1967. Notes on Eriogonum — II. Variations in Erio- gonum atrorubens. Sida 3: 82-86. UNIVERSITY OF MARYLAND COLLEGE PARK, MARYLAND 20742 and NEWARK STATE COLLEGE UNION, NEW JERSEY 07083 NEW FORM OF THUJA OCCIDENTALIS RESEMBLING KNOWN CULTIVARS Mary I. MOORE In 1963 the author visited the canyon of the Barron River in Algonquin Provincial Park, Ontario at latitude 45954' N and longitude 77°38’-39’ W. Normal Thuja occi- dentalis L. grows on slopes at the foot of cliffs in this mile long fault in granitie gneiss. On several talus formations there are dwarf forms, barely a meter high, with dead tops, compact, prostrate branching and normal leaf devel- opment. One dwarf shrub of very unusual form was found (Moore, 1964) on a north facing rock slide covered with a carpet of chasmophytic ferns, Cladonia and bryophytes such as Hedwigia and Polytrichum. A specimen was sent to the Plant Research Institute, Dept. of Agriculture, Ot- tawa. Drs. W. G. Dore and C. Frankton were unfamiliar with the shrub but established the anatomy of the stem as that of a conifer and the odor as that of Eastern white cedar (Frankton in litt., 1963). In 1968 Dr. R. E. Beschel, Queen's University, Kingston, Ont. visited the shrub and noted its resemblance to certain bizarre cultivars. Both he and Mr. E. Perem of the Wood Anatomy Group, Eastern Forest Products Laboratory, Ottawa considered that the wood anatomy corresponded with the description given for T. occidentalis (Arbor vitae). DESCRIPTION The shrub has a central stem 0.8 m high, with eight branches up to 2 m in length on the lower half. Of these, two come from beneath the rock and are in a prostrate position, The branches may have three orders of branch- ing and bear live or dead leaves. They are basically of three types: (1) About 75% are whip-like, sparingly ramified, with four rows of equal, appressed-decussate, ovate, keeled and pointed leaves 3-5 mm long (Fig. 1; 2A, 2B. Fig. 2A). The overall tetragonal habit of the branches resembles strongly some appressed species of Cassiope D. Don. 352 1972] Thuja occidentalis — M. Moore 353 Figure 1. Branch and leaf form of typical Thuja occidentalis and the dwarf conifer in the Barron canyon. A — side view of branchlet, B — end view of branchlet: 1 — typical T. occidentalis, 2 — tetrag- onal, decussate form on dwarf conifer, 3 — lycopodiaceous form on dwarf conifer. (2) Several branches bear fastigiate clusters of twigs with rows of divaricate, needle-shaped leaves 6-8 mm long. These appear juvenile but some are stiffer and more sharp- ly pointed. They may develop in a gradual sequence out of the first type and revert again to it (Fig. 2B). (3) A few small sprays bear leaves similar to but usually smaller than (1). These branchlets are flattened, without the constricted segments of normal T. occidentalis, and are intensely branched in the manner of Lycopodium trista- chyum Pursh (Fig. 1; 3A, 3B. Fig. 2C). DISCUSSION The shrub bears no external resemblance to a normal Thuja occidentalis. A search of botanical and horticul- Q9 or nS Rhodora [Vol. 74 Figure 2. Branches from dwarf conifer in Barron canyon. A — tetragonal, whip-like branch, sparingly ramified, B — fastigiate clus- ter with juvenile-type needles, C — lycopodiaceous form of branch- ing. tural literature reveals no such specific entity. The forma prostrata of Vict. & Rousseau has normal adult leaf struc- ture and differs mainly in habit. Var. ericoides Beiss. & Hochst., which is also a cultivar, bears only small resem- blance to parts of the shrub with juvenile-type needles. The cultivar, 'Douglasi? Rehder, given as a synonym for ‘Fili- formis’ Beiss., has some similarity with it and is described as a compact shrub with slender, pendulous branches often flattened at the tip. Bailey (1923) says these branches are partly four-angled with sharply pointed leaves, but his il- lustration shows very typical T. occidentalis foliage. Reh- der (1901), the authority for the variety, states that the stronger shoots are round, covered with widely-spaced, long, pointed and somewhat projecting leaves, but that there is 1972] Thuja occidentalis — M. Moore 355 no juvenile foliage. The illustration of Welch (1966) con- firms this. 'Ohlendorfi? has tetragonal branches with de- cussate leaves but the illustration in Welch shows a very compact shrub with upright branches. The leaves of the tetragonal and lycopodiaceous branches do show an interesting resemblance to those of T. orien- talis L., species of Cupressus and Chamaecyparis (Dalli- more & Jackson, 1948) as well as to forms of Juniperus. Horticultural forms of dwarf and other abnormal coni- fers are developed by mutation, self-pollination and hy- bridization. According to Welch most, if not all, cultivars of Arbor-vitae were originally the result of seed mutation. Such seedlings can be cultivated in a nursery, but in nature their chances of survival are not great. ‘Spaeth? (Anony- mous, 1893), a synonym of ‘Ohlendorfi’, was developed in this way from a seedling of T. occidentalis. CONCLUSION The question has been asked, is the shrub a monstrosity induced by the particular habitat in which it occurs, or is it a genetic form that arose as a result of self-fertilization, hybridization or mutation. The shrub is close to the river where in some years it could be subject to damage from ice and high water; a visit in 1970 showed that this had recently occurred with dam- age to many of the long, tetragonal branches. The climate of the canyon is, in part, abnormally cool; several rare arctic-alpine species occur there (Moore, 1964) and on the first visit in late May two large ice falls were still pres- ent. The lower branches of cedars along the river are browsed. However, no other known cedars have abnor- ma] leaves. Abnormal leaf forms may appear on young conifers, usually under thirty years of age, after which they usually outgrow all or most of the characteristics (A. R. Buckley, Horticulture Division, Plant Research Institute, pers. comm., 1969). The shrub in question must be at least fifty 356 Rhodora [Vol. 74 years old since Mr. Perem gave the age of a small branch- let 2.3 mm in diameter as more than twenty years. The plant presumably could not have been a hybrid be- cause no other species of Thuja grows naturally in On- tario. The canyon could not have been cultivated and what farms may have been in this area of the precambrian shield have returned to forest. It could have been due to self-pollination. However, it seems most probable that the shrub is the result of a spontaneous mutation. Unfortu- nately there is no evidence of flowering or cone develop- ment. Mr. John Santon, Petawawa Forest Experiment Sta- tion, has rooted tiny cuttings of both juvenile and tetrag- onal foliage. Growth the first winter was generally of the tetragonal, decussate form common to the shrub but pres- ent summer growth is mostly of the juvenile type. Mr. Santon considers that this alteration of form may be seasonal but commented that a return to juvenile growth is frequent in cuttings. The dwarf conifer in the Barron canyon appears to be a unique form. Its wood anatomy defines the shrub as Thuja occidentalis and the odor is characteristic of that species. It can be assumed that this Eastern white cedar was not induced by humans although the shrub does re- semble in part several bizarre cultivars of the species pre- viously described. ACKNOWLEDGMENTS I wish to thank all those who gave help with the manu- script and especially those who helped to identify the shrub. SELECTED BIBLIOGRAPHY ANONYMOUS. 1893. Thuya occidentalis Spaethi. Gartenflora, Ber- lin, 42: 539. BAILEY, L. H. 1923. The cultivated evergreens. MacMillan Co., New York. 224-25, 1944. The standard cyclopedia of horticulture. MacMillan Co., New York. 3335-36. 1972] Thuja occidentalis — M. Moore 357 CARRIERE, E. A. 1880. Revue du genre Retinospora. Revue Hor- ticole, Paris. 93-97. DALLIMORE W., and A. B. JACKSON. 1948. A handbook of coniferae. Edward Arnold Ltd., London. 686 pp. FERNALD, M.L. 1950. Gray’s manual of botany. American Book Co., New York. 1632 pp. HORNIBROOK, Murray. 1923. Dwarf and slow-growing conifers. Scribner, New York. 165-75. KRÜSSMANN, G. 1960. Die Nadelgehólze. Paul Parey, Berlin and Hamburg. 295-303. Moore, Mary I. 1964. Some interesting plants in the Barron can- yon in Algonquin Park. Canadian Field-Naturalist. 78: 125-27 REHDER, ALFRED. 1901. Thuya occidentalis var. Douglasi. Móeller's Deutsche Gártner-Zeitung. Erfurt, Germany. Vol. 16: 30. 1940. Manual of cultivated trees and shrubs hardy in North America. MacMillan Co., New York. 996 pp. SILVA Tarouca, E. and C. SCHNEIDER. 1923. Unsere Freiland- Nadelgehólze. Leipzig. 288-90 SYNGE, P. M. (editor). 2nd edition. Dictionary of gardening. Royal Hort. Soc., Oxford. 4: 2104. WELCH, H. J. 1966. Dwarf conifers. Faber and Faber, London. 334 pp. PETAWAWA FOREST EXPERIMENT STATION CHALK RIVER, ONTARIO SNOW COVER AND THE DIAPENSIA LAPPONICA HABITAT IN THE WHITE MOUNTAINS, NEW HAMPSHIRE! WESLEY N. TIFFNEY, JR. The alpine zones of the White Mountains, New Hamp- shire, provide a convenient study area for investigators interested in the ecology of alpine plants. The White Moun- tains are a series of peaks, heavily glaciated during the Pleistocene Epoch, 46 of which exceed 4,000 ft. The highest of these is Mt. Washington (6,288 ft.) in the Presidential Range. During the Wisconsin substage of Pleistocene glaciation the climate allowed arctic plants to become estab- lished in the area. The present harsh climate typical of the alpine zones of the White Mountains has maintained these plants as an arctic outpost in a temperate area. Treeline, here defined as the upper limit of continuous tree cover, is located at about 4,800 ft. on the west side of the Presi- dential Range and at about 5,200 ft. on the east (Antevs, 1932; Bliss, 1963). Prevailing winds are from the west. The best developed alpine zone in the White Mountains is found on the Presidential Range; it extends from Mt. Madison (5,363 ft.) in the north to Mt. Clinton (4,312 ft.) in the south. The area is about eight miles long and two miles wide at its greatest extent and encompasses about 7.5 square miles (Bliss, 1963). A smaller alpine zone exists in the Franconia Range some 20 miles west of the Presi- dential Range; it extends from Mt. North Lafayette (5,000 ft.) to Mt. Haystack (4,600 ft.), having as its highest point Mt. Lafayette (5,249 ft.). This zone encompasses about 1.5 square miles. The climate of the alpine zone has been summarized by several investigators (Antevs, 1932; Bliss, 1963; Pease, 1964). Weather information has been gathered by the Mt. ‘This paper constitutes a part of a doctoral dissertation submitted to the University of New Hampshire. 358 1972] Diapensia — Tiffney 359 Washington Observatory located on the summit of that peak. In general, the climate is characterized by high winds, low temperatures, high precipitation with heavy winter snow and extensive fog and cloud cover. This paper represents a portion of a larger investigation involving the distribution of White Mountain plants rela- tive to winter snow cover. The purpose of the present study is to define the ecological relationship between snow cover and a habitat typified by Diapensia lapponica L., an Angio- sperm cushion-forming plant, used by Bliss (1963) to characterize the association he termed the “Diapensia community". In both the arctic and in the alpine zones of the White Mountains Diapensia is reported in exposed and wind- swept areas (Antevs, 1932; Hadley and Bliss, 1964; Lóve and Lóve, 1966; Courtin, 1968, 1968a). Several investiga- tors have further suggested that the plant is limited to areas blown clear of winter snow (Hadley and Bliss, 1964; Courtin, 1968, 1968a). Bliss (1963) reports that Diapensia lapponica is a prime colonizer of bare ground and is found in ". . . areas that are subjected to frost action because of the lack of winter snow cover". Later, (1966) he reports, "Diapensia communities are found where winter snow cover is thin, or non-existent, and melts early in spring". Bliss (1969) recorded snow depth for two winters in three locations; it is not stated if any of these locations were in Diapensia areas. However, he states, “During much of the winter, large areas of Alpine Garden are blown free of snow, areas dominated by Diapensia communities". My objective in this study was to analyze the species composition of the Diapensia habitat and to determine the degree of interdependence among its component species. First, it was necessary to test the snow cover hypothesis described above by gathering extensive records of winter snow depths in Diapensia habitats. I observed a number of habitats over several seasons to find if the plants did inhabit snow-free areas and if these areas were sub- stantially free of snow throughout the winter season. A 360 Rhodora [Vol. 74 system of summer sampling was designed to accumulate data for statistical analysis of interdependence among com- ponent species of the habitat. Snow Depth and Winter Dessication For the winter snow depth study 14 plots were estab- lished and marked with wire stakes at the margins of the greatest Diapensia concentrations. Two plots were located in each of the following areas: near the fourth peak of Mt. Adams, at the Lake-of-the-Clouds Hut, on the summit cone of Mt. Washington, on the section of the Chandler Ridge termed Cape Horn, at the lip of Huntington Ravine in Alpine Garden, at Lion Head in Alpine Garden and at Cow Pasture on the Automobile Road. These plots were visited throughout the winters of 1966-1967, 1967-1968 and 1968-1969 on as nearly a regular basis as the harsh winter weather of the Presidential Range permitted. Each plot was visited at least once each winter and some aS Many as eight times. During the three winters 14 trips were made to check snow depths in plots. After a snowfall in the White Mountains the wind fre- quently rises, usually from the west. Most of the snow which has accumulated during the storm in the open alpine zone is blown away and collects below treeline on the east side of the mountains. This is particularly true on the Presidential Range and leads to the massive accumula- tions of snow common in the large glacial valleys on the east side of the range, such as Tuckerman and Huntington Ravines. Snow remaining in the alpine zone is packed into the sparce vegetation or in the lee of large wind obstruc- tions. In ascending the mountain increasing snow depth is encountered until treeline is reached; above treeline snow cover conforms to the microrelief of the alpine zone. In the Diapensia habitat there are few obstructions and virtu- ally all the snow is blown clear throughout the winter season (Fig. 1). A trip to check snow depth made on February 19-26, 1967, will serve as an example of the snow cover /wind 1972] Diapensia — Tiffney 361 Fig. 1. Snow-free ridge crest, about one-half mile long, near Lion Head in Alpine Garden, Presidential Range, New Hampshire. The area is dominated by Diapensia lapponica. March 3, 1969. relationship on the Presidential Range. On the night of February 20 a storm deposited a recorded 2.7 inches of snow at the summit of Mt. Washington. Drifts of two to three feet accumulated in the vicinity of the Weather Observatory. On the morning of February 21 the wind rose from the west and reached a sustained velocity of 60 to 70 MPH with a peak gust of 105 MPH. Virtually all the snow, including the large drifts, was removed from the summit area. On February 23-24 another storm developed and deposited a recorded 19 inches of snow at the summit. Winds rose during the evening of February 24 and reached a sustained velocity of 70 to 80 MPH. The wind persisted through February 26. Again, almost all ac- cumulated snow was removed from the summit area. Visits to six Diapensia habitats on February 26, varying from 6,000 to 4,000 feet elevation, showed that all were devoid 362 Rhodora [Vol. 74 SR ag Ge 777 0 v de iiim ae, - es Wr LI Fig. 2. Winter aspect of a Diapensia habitat at Cape Horn, Chandler Ridge, Mt. Washington. Surrounding areas are covered with snow. The ice axe is about 40 inches long. Feb, 26, 1968 of snow after a total fall of 21.7 inches over the previous six days (Fig. 2). Late wet snowfalls, although more dense than dry mid- winter snows, are also blown away by strong winds and do not accumulate in the Diapensia habitat nor do they delay flowering. On May 26, 1967, a 12.8 inch snowfall was followed by winds of 70 MPH. Visits to several habitats on May 30 and 31 disclosed no snow in the areas. Less than three weeks later on June 18, 1967, Diapensia plants were in full bloom throughout the Presidential Range (Fig. 3). Winds are not as strong where plants are actually grow- ing on the range as they are where recorded on a 39 foot instrument tower on the summit of Mt. Washington. A wind velocity of 100 MPH on the summit is equivalent, on the average, to a speed of 30 MPH at plant level (Courtin, 1972] Diapensia — Tiffney 363 Fig. 3. Plants of Diapensia lapponica in bloom, Cape Horn, Mt. Washington, New Hampshire. June 17, 1968. The lateral area covered by the photograph is about 18 inches long. 1968). Such wind velocities are stil] sufficient to remove snow from exposed areas and to promote desiccation in winter-exposed plants. In an average winter on Mt. Washington the Weather Observatory records 221.4 inches of snow. The winter of 1966-1967 totaled 260.6 inches, the winter of 1967-1968, 211.1 inches and the record winter of 1968-1969 yielded 566.4 inches of snow. These variations in annual snowfall have little effect on the total amount of snow adhering to the ground in the alpine zone. My observations indicate that the snow holding capacity of the zone is reached early in the winter. Additional snow is blown away by strong winds. This study spanned two relatively normal snowfall years and one record year. Amounts of snow in Diapensia habitats and other areas checked remained constant. This minimal snow cover melts rapidly once warm -weather arrives and melts at about the same time each year. Even 364 Rhodora [Vol. 74 Fig. 4. Winter condition of Diapensia lapponica plants, Cape Horn, Chandler Ridge, Mt. Washington, New Hampshire. Meltwater has formed ice about the bases of the plants but the crowns are fully exposed. Jan. 26, 1967. The ice axe is about 40 inches long. late-lasting drifts supporting specialized snowbank com- munities (Bliss, 1963) disappear at about the same time each season. The flowering time of Diapensia lapponica and other spring-blooming plants on the Presidential Range is not greatly influenced by heavy or light snow years or by heavy snowfalls late in May. In the three years included in this study alpine plants were at the peak of their bloom between June 15 and June 25 (Fig. 3). The examples above, drawn from the winter of 1966- 1967, are typical of results of other visits in 1967-1968 and 1968-1969. In some cases meltwater had formed ice about the bases of Diapensia plants but in all cases observed the major portion of the plant was exposed (Fig. 4). Areas dominated by Diapensia were those areas blown clear of snow throughout the winter. Accumulations of as little as a few inches of snow marked the margin of the Diapensia 1972] Diapensia — Tiffney 365 habitat and the beginning of a transition to another form of plant cover. Desiccation is a major factor in the distribution of alpine plants as Tranquillini (1964) notes when he states, “Ex- tremely short snow cover can only be tolerated by plants that are resistant not only to cold but also to long periods of drought . . .", and Sakai (1970) notes that “. . . desic- cation damage in winter constitutes the greatest limiting factor for growing plants in cold climates and in high mountains . . .". Plants in the Diapensia habitat appear well suited to resist the effects of desiccation. Several in- vestigators have noted that the low mat or cushion form exhibited by plants such as Diapensia is of survival value in these conditions (Holttum, 1922; Hadley and Bliss, 1964; Courtin, 1968a). The low profile of these plants offers less resistance to strong winds and their desiccating effect is minimized. Information on the freezing resistance of several plants of the Diapensia habitat has been accumulated by Sakai and Otsuka (1970). They note that several species acquire a resistance in the fall at —15? C. that allows them to with- stand winter temperatures of —70° C. After acquiring such resistance Vaccinium uliginosum could withstand tempera- tures of —50° C., V. vitis-idaea and Loiseleuria procumbens could survive at —70° C. The most cold resistant plant tested was Diapensia lapponica which, after full harden- ing, could survive immersion in liquid nitrogen at —-196? C. (Sakai and Otsuka, 1970). The lowest recorded tempera- ture for Mt. Washington is —44? C. (ATS EJ. Species Composition Twenty-two Diapensia habitats were chozen for sampling to determine species composition and major sample areas were located in them. Plots were placed in the center of the habitat. The sampling procedure used is similar to that of Bliss (1963). An area 4 by 8 meters (320,000 sq. cm.) was marked with surveying tape and stakes. The 4-meter axis was located parallel to the slope of the ground. 366 Rhodora [Vol. 74 1 2 3 4 5 6 7 8 TRANSECT LOCI - METERS Fig. 5. Arrangement of quadrats within a major sample area. The transects on which the quadrats are located are randomly chosen. The design is similar to that of Bliss (1963). Five rectangular quadrats 25 by 40 cm. (1,000 sq. cm.) were established along a sequence of transects based at four randomly chosen locations on the 8-meter axis. Alter- nate quadrat transects were begun on the 8-meter axis, and others 0.5 meters from it. Twenty rectangular quadrats were established in each major sample area for a total sampled area of 20,000 sq. cm. or 6.3% of the total area sampled (Fig. 5). Quadrat size and number corresponded with recommendations for plot size and sample design in alpine vegetation made by Eddleman et al. (1964). Measurements were taken of the ground area covered by all vascular plants in each 25 by 40 cm. quadrat. Measure- ments were taken also for cover of moss and lichen and for uncolonized ground and rock. These were expressed as percent cover for each sample area, and grand totals for all 22 areas sampled were compiled. Areas sampled for species composition were, in general, the same as those used for winter observations. Results are given in Table 1. My results expressed as percent cover are roughly com- parable to those of Bliss (1963) expressed as importance 1972] Diapensia — Tiffney 367 Table 1 Standard Type Mean Range Deviation Rock 31.1 1.4-66.6 19.5 Diapensia lapponica 27.6 1.8-54.0 15.0 Lichen (all types) 13.0 0—46.4 14.1 Vaccinium uliginosum 5.8 0-18.5 5.2 Juncus trifidus 5.3 0-23.0 6.4 Uncolonized Ground 4.3 0-18.2 4.6 Moss (all types) 3.2 0—31.1 6.7 Carex bigelowii EM 0—11.8 3.6 Salix uva-ursi 1.5 0-7.9 2.5 Vaccinium vitis-idaea 1.1 0-7.2 2.0 Potentilla tridentata 1:0 0—4.8 1.3 Loiseleuria procumbens 0.8 0-7.8 1.9 Rhododendron lapponicum 0.6 0—4.3 1d Arenaria groenlandica 0.5 0-2.5 0.7 Results of sampling major plant components of the Diapensia habitat, White Mountains, New Hampshire. Means, ranges and standard deviations for all types sampled are expressed as percent ground cover of the total area sampled. Other plants were present in quantities of less than 0.5% ; these minor components were not subjected to sta- tistical analysis. These were, together with their means: Solidago cutleri 0.4% ; Poa alpigena 0.3%; Scirpus cespi- tosus var. callosus 0.2%; Betula ssp. 0.1%; Vaccinium angustifolium 0.05% ; Abies balsamea 0.05% ; Empetrum nigrum 0.03% ; Prenanthes boottii 0.01% ; and Lycopodium selago 0.002%. These minor forms totaled 1.1% of the total area sampled. The nomenclature used is that of Fernald (1950). 368 Rhodora [Vol. 74 values. He recorded higher values for rock and uncolonized ground as well as for Juncus trifidus; slightly higher values for Potentilla tridentata, Loiseleuria procumbens, Rhodo- dendron lapponicum, and Arenaria groenlandica; similar values for Diapensia lapponica and Vaccinium uliginosum, and a lower value for lichen. Results of Bliss’s analysis are based on 2 to 9 sample areas. Results of my sampling indicate that 47.3% of the area in the Diapensia habitats was occupied by vascular plants, 35.4% was occupied by rock and uncolonized ground and 16.2% by moss and lichen. The typical plant of these habitats was Diapensia lapponica which was about five times as concentrated as the next ranking vascular plant. Wide ranges and large standard deviations were found for all component species tested (Table 1). This suggests that it is not possible to predict with accuracy the concen- trations of the components to be found in a given Diapensia habitat. The minimum concentration of Diapensia lap- ponica, 1.8% of the area sampled, was encountered in a plot located on the west flank of Mt. Madison. Other sample plots placed in nearby Diapensia habitats yielded results of 30.1%, 35.0% and 47.2% cover of D. lapponica. The maximum concentration encountered, 54.0% was in a plot located on the west side of Mt. Lafayette in the Franconia Range. Nearby sample plots in this area showed concentra- tions for D. lapponica of 15.6%, 22.0% and 33.4%. Values for other component species of the habitat were equally variable. Analysis of Component Species To assess the degree of interdependence existing among the component species of the Diapensia habitat statistical testing was necessary. Tests based on frequency data such as the “C.” correlation of Hurlbert (1969) were rejected due to the difficulty of collecting accurate frequency data in alpine vegetation. As Woodin (1959) has indicated, frequency information is difficult to obtain in alpine areas of mat-form vegetation as it is hard to tell where one plant 1972] Diapensia — Tiffney 369 ends and another begins. Goodall (1970) notes that, “Many plants, however, have methods of vegetative reproduction which make distinctions between individuals highly arbi- trary. Biomass per unit area is a measure which is more consistently objective’. For these reasons estimates of biomass in terms of percent ground cover were used here. The purpose of this part of the study is to assess the degree of interdependence present among plants of the Diapensia habitat. Sokal and Rohlf (1969) have said, “. .. when we wish to establish the degree of association between pairs of variables in a population sample, correlation analysis is the proper approach.” Hence, the product-moment cor- relation coefficient has been used here. In computing the correlation coefficient, data are ranked in two sets. The first set (Y,) is arranged in increasing rank order, the second set (Y.) is paired with the first. The correlation coefficient (7) indicates the degree of associa- tion between the two sets (Y, and Y.) and may be expressed in percent. The value of r may vary from -1 (minus 100%) to +1 (plus 100%). A value approaching -1 indicates negative association between Y, and Y,, a value approach- ing +1 indicates positive association between the two sets of variables. A value of r approaching 0 indicates little interdependence between Y, and Y.. Reciprocal tests yield the same results. For each correlation coefficient (value of r) significance tests are performed against the hypoth- esis that r — 0. Each of the 14 major species components of the Diapen- sia habitat (those having means greater than 0.5% of the total area sampled) was tested against the other 13 major components. This required a total of 91 correlation co- efficient tests, which were performed with the aid of a computer and using Leasco Systems and Research program no. CL-00001.023-00. Results are given in Table 2. Ninety-one tests were performed; only eight yielded positive or negative correlation coefficients of 50% or greater. The strongest relationship observed was between moss and Arenaria groenlandica with a value of 68.0%. [Vol. 74 Rhodora 370 ‘stsayjodady [[nu ayy Jeq} AzyIqeqoid ayy, 0 wunoruoddn) wo4puapopo Y IL Eg- SUIQuUNIOLd DILNI)ISLOT L6 06- 0 DID] Wapily 0]uuojoqd 967 8I II 6 Danpi-sijta WNLULIOD A G— 0 i a te ET IsUN-DAN xi]g T9 9 L6- cv 9— pE- 14Ío]aBiq LIMI 89 ST- LT SSON vs L 6€ puno.r) peziuo[oou n "19]1€29.L8 10 2/59p Jo 4 JO sanjea 10J (%0'¢) GQ’ SI ont, SI ‘Q = 4 "UA YIM *A Jo oouopuoadop.oqur yusdied se passoidxe JIL sq[nsey "393 -qeu 94} Jo 3uouoduroo sofe Tayyo Ad19A9 YAM (pe[dures vog? [v0 Jo ACO ULY} 197891? Sut?our Zursey sSoy}) yeyiqey msuədmqg ayy Jo 3uouoduroo olewu quovo TOF (4 JO San[VA) 4S9} quolOuJooo uoljv[9.Lt00 ƏY} JO synsəy LI- fe 9T L~ OF L CT SI- I GI- I2- eI L- r&- 6I- c 8 pZ- vc yoo i 8 Vh S RE * E 3 E S 6 PIL po- uaWqorT noiuoddo] nisuoadpnw] 66— 61— D23ipupnquoo4D mDi4Duo4V wmnoaucddn) wo4puopoponu y suaqumoo4d. p14n2]28107T 0)09uopi42 0j123u220d DIDP SIA UNIMIIID A ?s4n-Dàm. LYNG! umopabig LIMA sso punoir) pezruo[oou[] snpyi4j snounf WUNSOULBYN MUMU A uyr] nnuoddn) Msuədmqg 1972] Diapensia — Tiffney 371 None of the relationships tested could be considered strong (in excess of 90%). A negative correlation coefficient resulted from the test between rock and lichen (-66% ). Hence, rock and lichen covary negatively 66% of the time. Lichens sampled in this study were of the foliose and fruticose types. Crustose or rock-inhabiting forms were rare and were not included. A rock habitat is not suitable for the growth of lichens in the areas tested. With the exception of the negative rela- tionship with lichens mentioned here, the large quantities of rock present in the Diapensia habitat (31.1% of the total area sampled) do not greatly influence the plants growing in the habitat. Negative correlations resulted from tests of Diapensia lapponica with Vaccinium uliginosum (56%) and with Juncas trifidus (-54%). This indicates that D. lapponica is negatively associated with V. uliginosum and J. trifidus about 55% of the time. The latter two plants are com- ponent species of an aggregation termed the “dwarf-shrub- rush-heath community” by Bliss (1963). This aggrega- tion is commonly found at the margins of the Diapensia habitat. As these margins are reached snow cover increases slightly and V. uliginosum and J. trifidus begin to dominate. These two species also characterize areas of slightly greater snow cover, as in the lee of small obstructions dispersed throughout the Diapensia habitat itself. A gradient was established by Bliss (1963) with Diapensia at the snow- free extreme and the snowbank community at the other extreme. “Dwarf-shrub-rush-heath” is placed next in order with increasing snow cover to the Diapensia aggregation on this model. Rather than indicating that competitive exclusion is operating among D. lapponica and V. uligino- sum and J. trifidus, I conclude that the negative correla- tions indicate that the margin of the Diapensia habitat has been reached in these sample situations and that a change to dominance by other forms is occurring. In this case the optimum snow cover situation for Diapensia has 372 Rhodora [Vol. 74 been exceeded, and the optimum for the other two species is being approached. This situation poses some possibilities for future experi- mental work. Snow depth in selected Diapensia habitats could be increased artificially by erecting several wind- breaks in the habitat. Vegetation could be mapped prior to establishing windbreaks, then re-mapped in successive growing seasons. At question would be changes in the composition of plant aggregates growing in the altered habitat. A project of this nature is planned. The test between Vaccinium uliginosum and lichen yielded a positive correlation of 61%. This indicates positive inter- dependence between V. uliginosum and lichen 61% of the time. Lichens, lacking roots, are not as firmly anchored as other plants and are frequently blown away by strong winds, an occurrence noted also by Antevs (1932). Most plants in the Diapensia habitat are mat-forming or cushion plants. An exception is V. wliginosum which, particularly in areas of slightly greater snow cover, adopts a more up- right stature. Under these conditions V. uliginosum consists of a number of interwoven branches which provide anchor- age for lichens. Possibly, the vascular plant provides sup- port for lichens of several types and prevents them from being blown away by strong winds. Positive results were obtained from tests between Are- naria groenlandica and uncolonized ground (54%), and moss (68%) and Carex bigelowti (61%). The *sedge-meadow community” of Bliss (1963) was composed of two vascular plants, Carex bigelowii and Are- naria groenlandica. In addition to keeping snow cover records for the Diapensia habitat I accumulated informa- tion on winter snow depth in areas of sedge meadow on the summit cone of Mt. Washington. About one to two inches of snow accumulate in these areas, the rest being blown away. This amount is sufficient to cover the ground level meristems of C. bigelowii, although dead culms from the previous growing season frequently protrude from the thin snow cover. Sedge meadow occupies a similar posi- 9 1972] Diapensia — Tiffney 373 tion on the snow cover gradient to the *dwarf-shrub-rush- heath community” of Bliss (1963). It is a marginal form to the Diapensia habitat, being supported by slightly in- creased snow depth. Again, experimental work should be performed to see if this form increases in concentration as a result of induced increases in snow cover in selected Diapensia habitats. Carex bigelowii and A. groenlandica are associated about 60° of the time in my samples. There is doubt as to whether Arenaria groenlandica is ahnual or perennial (Gleason and Cronquist, 1963). The plant is described as being associated with mossy areas by Antevs (1932) and is regarded as being an early colonizer of bare ground by Antevs (1932) and by Bliss (1963). The plant is noted as growing on disturbed ground and particularly in hiking trails by Harris (1964). An annual or biennial life cycle would offer some explanation for the affinities of this plant for mossy areas and areas of open ground. In my tests it was positively associated with un- colonized ground 54% of the time and with moss 68% of the time. Open ground provides space to grow in the alpine zone otherwise occupied by perennial plants or rock. Mossy areas may promote seed germination and seedling survival due to greater water availability. Observations made in this study indicate that A. groenlandica is little controlled by snow cover, being distributed widely in a number of habitats which show wide variation in their amount of winter snow. Loiseleuria procumbens is a cushion-forming plant very similar in form to Diapensia lapponica. It is rarely found in other than Diapensia habitats in the White Mountains. But, the test of L. procumbens with D. lapponica yielded a correlation coefficient of only —22% indicating that the plants were negatively associated —22% of the time. This value of r (—22%) is not significantly different from 0 at the .05 (5.0%) conference level. Of ten vascular plants tested (Table 2) five showed association with other vascular plants. These were Dia- pensia lapponica, Vaccinium uliginosum, Juncas trifidus, 374 Rhodora [Vol. 74 Carex bigelowii and Arenaria groenlandica. One of these five, V. uliginosum covaried with lichen and one, A. groen- landica, showed interdependence with moss and with un- colonized ground. One group of non-vascular plants, lichens, covaried with rock. Five vascular plants showed no interdependence with other components of the habitat. These were Salix uva-ursi, Vaccinium vitis-idaea, Potentilla tridentata, Loiseleuria procumbens and Rhododendron lap- ponicum. Of ninety-one potential positive or negative inter- actions tested only eight occurred; none of these eight could be considered intense. Discussion The community concept involves the ideas that there are interactions among the component plants of the community and that the species structure of the community follows some predictable pattern. As Odum (1971) has said, “Com- munities not only have a definite functional unity with characteristic trophic structures and patterns of energy flow but they also have compositional unity in that there is a certain probability that certain species will occur together". Both Savile (1960) and Bliss (1962) have noted that competition in severe habitats is often secondary to the effects of the physical environment. The result ", .. may be the random occurrence of plants with few distinct associations" (Bliss, 1962). Savile (1960) sug- gests that arctic vegetation may better be described by habitats than by associations. I have used the term habi- tat, here defined as the place where a plant grows, in this analysis of an area dominated by Diapensia lapponica. Sakai (1970) has noted that winter desiccation is a major limiting factor to alpine plant growth. Sakai and Otsuka (1970) have shown that Diapensia lapponica, Vaccinium uliginosum, V. vitis-idaea and Loiseleuria procumbens can survive very low temperatures. I suggest that Diapensia and other plants associated with it can survive in this snow-free and exposed habitat because they can tolerate very low temperatures and resist desiccation from strong 1972] Diapensia — Tiffney 375 winter winds. Other plants are excluded because they are not adapted to survive in this harsh physical environment. Further investigation may provide more information on such relationships in the alpine zone. Conclusions The occurrence of the Diapensia habitat is predictable in relation to winter snow cover. Diapensia lapponica is found in areas blown clear of snow throughout the winter season. Increasing snow cover results in increasing domi- nance by other forms of plants. The Diapensia habitat is located at one extreme of the snow cover gradient, and this study suggests that aggregations characterized by Vac- cinium uliginosum, Juncus trifidus and Carex bigelowii may be next in rank with increasing snow cover. Wide variation was found among concentrations of all species components over the 22 habitats tested. These habitats do not consist of predictable percentages of species components. Interdependence among the species components of the Diapensia habitat are few. Of ninety-one potential inter- actions tested eight occurred at significant levels and none of these could be considered intense. Control in the habitat is primarily exercised by the harsh microclimate. Prime factors are low temperatures and high winds combining to promote severe winter desiccation in this snow-free area. Plants which are able to survive in this harsh microenviron- ment do so and grow here. Others are not fitted to survive in these conditions and may be excluded by the micro- environment. The unpredictable concentrations of plants in the areas tested, together with their lack of interactions, renders the use of the term habitat more appropriate than the term community to describe the aggregate found in this location. Acknowledgements I should like to give particular thanks to Dr. Albion R. Hodgdon for his continued support of this work. Thanks 376 Rhodora [Vol. 74 are also due to the Mt. Washington Observatory for pro- viding living facilities; and to S. Tiffney, M. Carriker, J. Roughgarden and J. Davis for their discussion and assist- ance. LITERATURE CITED Antevs, E. 1932. Alpine zene of the Mt. Washington Range. Mer- rill and Webber Co., Auburn, Maine. 118p. Buss, L. C. 1962. Adaptations of arctic and alpine plants to environmental conditions, Arctic 15: 117-144. 1963. Alpine plant communities of the Presidential Range, New Hampshire. Ecology 44: 678-697. 1966. Plant productivity in alpine microenvironments on Mt. Washington, New Hampshire. Ecol. Monog. 36: 125-155. 1969. Alpine community patterns in relation to en- vironmental parameters in Essays in plant geography and ecology. K. N. H. Greenridge, ed. Nova Scotia Museum, Halifax. pp. 167-184. CourTIN, G. 1968. Microclimate of Mt. Washington. Mt. Wash- ington Observatory News Bulletin 9: 2-6. 1968a. Evapotranspiration and energy budget of two alpine environments, Mt. Washington, New Hampshire. Ph.D. thesis, Univ. of Ill, Urbana. 172p. EDDLEMAN, L. E., E. E. REMMENGA and R. T. WARD. 1964 An evalu- ation of plot methods for alpine vegetation Bull. Torrey Bot. Club 91: 439-450. FERNALD, M. L. 1950. Gray’s manual of botany, eighth ed. Ameri- can Book Co., N.Y. 1632p. GLEASON, H. A. and A. CRoNQUIST. 1963. Manual of the vascular plants of Northeastern United States and adjacent Canada. D. Van Nostrand Co. Inc., N.Y. 810p. GoopaLL, D. W. 1970. Statistical plant ecology im Ann, Rev. Ecol. and Syst, R. Johnson, P. Frank and C. Michener eds. Ann. Revs. Inc., Palo Alto, Calif. pp. 99-124. HADLEY, E. B. and L. C. Briss. 1964. Energy relationships of alpine plants on Mt. Washington, New Hampshire. Ecol. Monog. 34: 331-357. Harris, S. K. 1964. Mountain flowers of New England. Appala- chian Mountain Club, Boston. 147p. HoLTTUM, R. E. 1922. 'The vegetation of West Greenland. Jour. Ecol. 10: 87-108. HURLBERT, S. H. 1969. A coefficient of interspecific association. Ecology 50: 1-9. 1972] Diapensia — Tiffney 377 LóvE, A. and D. LóvE. 1966. Cytotaxonomy of the alpine vascular plants of Mt. Washington. Univ. Colo. Series in Biol, No. 24. Univ. Colo. Press. 74p. PEASE, A. S. 1964. A flora of Northern New Hampshire. New England Botanical Club Inc., Cambridge, Mass. 278p. OpuM, E. P. 1971. Fundamentals of ecology, third ed. W. B. Saunders Co., Philadelphia. 574p. SAKAI, A. 1970. Mechanism of desiccation damage of conifers wintering in soil-frozen areas. Ecology 51: 657-664. . and K. OrsuKA. 1970. Freezing resistance of alpine plants. Ecology 51: 665-671. SAVILLE, D. B. A. 1960. Limitations of the competitive exclusion principle. Science 132: 1761. SoKAL, R. R. and J. RoHrr. 1969. Biometry. W. H. Freeman & Co., San Francisco, Calif. 776p. TRANQUILLINI, W. 1964. The physiology of plants at high altitudes. Ann. Rev. Plant Physiol. 15: 345-362. WooDIN, H. E. 1959. Establishment of a permanent vegetation transact above timberline on Mt. Marcy, New York. Ecology 40: 320-322. BIOLOGY DEPARTMENT UNIVERSITY OF MASSACHUSETTS — BOSTON 100 ARLINGTON STREET, BOSTON, MASS. 02116 NEW AND INTERESTING PLANTS FROM THE GREAT PLAINS RONALD L. MCGREGOR, 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 Kansas, Nebraska, South Dakota and Wyoming. A new form of Sida spinosa L. with white flowers that has been known in Kansas for several years is described. All speci- mens cited, unless indicated otherwise, are located at KANU. Paspalum distichum L. KANSAS: NEOSHO CO.: 5 1/4 min Erie on US 59, edge of farm pond in water, 18 October 1970, Holland 2854. This represents an extension of the range of the species northward from Oklahoma. Panicum philadelphicum Bernh. ex Trin. KANSAS: CHAU- TAUQUA CO.: 2 mi se Sedan, sandy open scrub oak area, scattered to common, 2 October 1969, McGregor 20708. GREENWOOD CO.: 1 mi ne Fall River, sandy open oak wooded area, abundant in local area, 2 October 1969, Mc- Gregor 20691. This represents an extension of the range of the species northward from Oklahoma and westward from Missouri. Andropogon intermedius R. Br. NEBRASKA: WEBSTER co.: 2 mi s, 3 w Red Cloud, reseeded prairie, common, 11 August 1965, MeGregor 19466. In Kansas this species is known from CLOUD CO.: 1 September 1952, Fraser 956, and REPUBLIC CO.: 10 September 1970, McGregor 23514. This species is widespread in tropical and subtropical areas of the Old World and is introduced in the United States. Eleocharis quadrangulata (Michx.) R. & S. var. crassior Fern. KANSAS: BOURBON CO.: Bourbon Co. St. Lake No.3, 1 mie & 2 n of Hiattville, common along se side of lake, soggy area of shoreline over limestone, 7 August 1970, Magrath & Weedon 5750. COFFEY CO.: 6.5 min & 378 1972] Great Plains — McGregor, Magrath & Weedon 379 1.5 w of Westphalia, side of a large pond, muddy loam, three small clumps, 19 September 1970, Magrath 6250, NEOSHO CO.: 2 mi s & 2 w of St. Paul, muddy margin of pond in native meadow, 13 July 1965, Holland 1065. Scirpus olneyi A. Gray. KANSAS: CHEROKEE CO.: 1.5 mi w, 0.5 s Baxter Springs, shallow water and muddy mar- gin of pond, large colony, 17 July 1970, Bare, Johnson & McGregor 2530; 6 August 1970, Magrath & Weedon 5719 (duplicates determined by T. M. Koyama). This extends the range of the species northward from Texas and westward from Missouri where it is known only from Jefferson County. Carex melanostachya M. B. ex Willd. KANSAS: DOUGLAS CO.: 0.5 mi n of Lone Star, roadside ditch, a few large colonies, 14 May 1964, Kolstad 1193; 25 May 1964, Kol- stad & Harms 2150 (duplicates determined by F. J. Her- mann); 24 May 1971, Weedon 6938. This is the first report of this species for North America. Komarov (1964) describes the habitat of this species as steppes, meadows and grassy places on saline soils, and gives its general distribution as Central Europe, the Mediterra- nean, Balkans, Asia Minor, Iran, Kulja, and the Caspian coast of the Caucasus. Rumex stenophyllus Ledeb. NEBRASKA. BUFFALO CO.: 1 mi s of Elm Creek, sandy flood plain of the Platte River, 14 July 1968, Bare, Johnson & McGregor 1670. CUSTER co.: 6.5 mi se Anselmo, prairie roadside bar ditch, in water, sandy soil, common, 20 July 1968, Stephens & Brooks 25182. DAWES CO.: 1.5 mi ne Crawford, wooded creek bank, wet sandy soil, weedy area, common, 14 July 1968, Stephens & Brooks 24637. DAWSON CO.: 2 mi s of Gothenburg, sandy flood plain of the Platte River, par- tially wooded, 14 July 1968, Bare, McGregor & Johnson 1594. GARDEN CO.: 1 mis & 0.5 mi e of Lewellen, low wet prairie in the North Platte River valley, 13 July 1968, Bare, McGregor & Johnson 1443. GREELEY CO.: 10 mi w of Spalding, sandy banks of Cedar River, common, 28 Au- 380 Rhodora [Vol. 74 gust 1967, Bare & McGregor 419. HARLAN CO.: 4 mis of Alma, sandy area near lake, locally common, 30 August 1965, McGregor 19568. HOLT CO.: 8 mi s of O'Neill, wet roadside area, sandy soil, 13 July 1966, Barker 2968. HOOKER CO.: 16 mi w Mullen, low spot in sandhill prairie, moist sandy soil, several plants, all small, 27 August 1968, Stephens 28267. KEITH CO.: 0.5 mi s of Brule, bank of South Platte River, locally abundant, 28 August 1966, McGregor 20175. KIMBALL CO.: 6 mi w of Kimball, mar- gin of lake, common in local area, 30 August 1966, Mc- Gregor 20269. LINCOLN co.: 4 mi w of Sutherland, sandy banks of the Platte River, 14 July 1968, Bare, Johnson & McGregor 1542. SCOTTS BLUFF CO.: 0.5 mi n of Melbeta, bank of North Platte River, sandy soil with some white alkali present, 27 September 1968, Weedon 4776. SHERMAN CO.: 3 mi se Litchfield, bank of Beaver Creek, small colony, 2 September 1965, McGregor 19784. WEBSTER CO.: 4 mi w of Red Cloud, bank of the Repub- lican River, large colony, 25 June 1953, McGregor 7545; 3 mi w of Red Cloud, 1 August 1964, McGregor 18676. Polygonum sawatchense Small. NEBRASKA: SIOUX CO.: 6 mi nw Fort Robinson, open pine forested hillside, 24 June 1970, McGregor 22358. This is an extension of the range of the species southward from the Black Hills of South Dakota. Suckleya suckleyana (Torr. Rydb. KANSAS: CHEYENNE co.: 0.5 mi w of St. Francis, sandy bank of river, locally common, 17 August 1971, McGregor 24027. HAMILTON CO.: 1 mi s of Syracuse, muddy bank of water hole, local colonies, 18 August 1971, McGregor 24091. KEARNY CO.: 3 mi ne Lakin, muddy shore of Lake McKinney, locally abundant, 18 August 1971, McGregor 24101. STAFFORD CO. : Quivira Natl. Wildlife Refuge, open areas in Distich- lis flats, locally common, 19 August 1971, McGregor 24129. Paronychia depressa Nutt. KANSAS: CHEYENNE CO.: 12 mi n St. Francis, rocky ledge, 9 June 1967, Stephens 1972] Great Plains — McGregor, Magrath & Weedon 381 11462. This is an extension of the range of the species southward from Nebraska and eastward from Colorado. Stellaria aquatica (L.) Scopoli. KANSAS: DONIPHAN CO.: 1 mi n White Cloud, willow thicket, bank of Missouri River, several large colonies, 29 August 1970, MeGregor 23467. This species is native to Europe. In North Amer- ica it is known from Quebec and Ontario to North Caro- lina and Louisiana, also in British Columbia. It is not known from any of the states adjacent to Kansas. Cerastium glomeratum Thuillier. KANSAS: CHEROKEE CO. : 6 mi e Baxter Springs, open rocky hillside, 13 Apri] 1946, McGregor 2735. NEOSHO CO.: 9 mi n, 0.5 e Erie, pasture, 3 May 1965, Holland 876. In the central plains states this species has been reported from South Dakota, Illinois and Texas. Draba lanceolata Royale. SOUTH DAKOTA: LAWRENCE CO.: 2 mi s, 8 w Walnut, Crooks Tower, top of peak, ca. 7180 ft, open woods, 12 August 1969, Stephens 35914. This is an extension eastward from the Rocky Mountains and Montana. Alyssum desertorum Stapf. SOUTH DAKOTA: FALL RIVER CO.: 2.5 mi e, 1 s Oelrichs, flat prairie at Limestone Butte, dry rocky gravel soil, few plants, 20 May 1970, Stephens 38265. WYOMING: CROOK CO.: 17 mi s Sun- dance, open prairie hillside, dry rocky clay soil, few plants, 13 June 1970. WESTON CO.: 7 mi se Osage, flat prairie pasture, dry gravel clay soil, 12 June 1970, Stephens & Brooks 39597. This plant is known from collections in Oregon, Washington, Idaho and Montana. This is a weed introduced from the Old World, which is now spreading in North America. Lesquerella gordonii (Gray) Wats. KANSAS: BARBER CO.: 4 mis Sun City, gypsiferous soil, grassland, 6 May 1956, Lathrop 2249; 11 mi w Medicine Lodge, red soil, pasture, 2 June 1968, Birkholz 2241. CLARK CO.: 6 mi n Ashland, sandy soil, 28 April 1939, Horr E250; e of Clark Co. St. 382 Rhodora [Vol. 74 Lake, grassland, dry sandy loam, common, 18 June 1951, Horr 3969. COMANCHE CO.: 4 mi se Coldwater, rocky level grassland, 28 April 1939, Horr, s.n.; 2 mi s of Cold- water, stationary sand dune bank, 28 May 1957, Lathrop & McGregor 3518. KIOWA CO.: 2 mi e, 3 n Belvidere, sandy grassland, common, 13 June 1951, Horr & Mc- Gregor 3808. MEADE CO.: 2 mi s, 11⁄4, e of Meade, over- grazed rocky pasture, common, 5 May 1957, Horr 4950; Meade Co. St. Park, sandy grassland, 16 June 1965, Bare 4. SEWARD CO.: 9 mi ne Liberal, sandy prairie, locally abundant, 27 April 1965, McGregor 19057. This is an extension of the range of the species northward from western Oklahoma. Saxifraga occidentalis Wats. var. occidentalis. SOUTH DAKOTA: LAWRENCE CO.: 2.5 mi e, 5 n Savoy at Bridal Veil Falls, Spearfish Canyon, moist steep rocky bank, 22 June 1970, Stephens 40481. According to Hitchcock, et al. (1961), the nearest other localities are northwestern Wyoming and western Montana. Croton capitatus Michx. NEBRASKA: SARPY CO.: 0.8 mi n&3 w (on St. Spur N 150) & 2 s on N 50 of Louisville, loess over sandstone, dry roadside ditch, rare 27 Septem- ber 1970, Magrath 6362. This extends the range of the species northward from Kansas. Croton glandulosus L. var. septentrionalis Muell. NE- BRASKA: ANTELOPE CO.: 1.4 mi nw of Clearwater on US 275, prairie roadside & railroad right-of-way, loam, extremely dry, one plant seen, 27 September 1970, Magrath 6390A. CASS CO.: 0.5 mi n Louisville, Louisville Park, flat area around small lake, dry sandy soil, abun- dant, 5 September 1970, Stephens 44634. These collec- tions extend the range of the species northward from Kansas. Croton monanthogynus Michx. NEBRASKA: ANTELOPE CO.: 1.5 mi w (on gravel rd) & 1.5 s & w (on paved rd) of Oakdale, dry roadside near oak woods, sandy loess, 1972] Great Plains — McGregor, Magrath & Weedon 383 scattered, 27 September 1970, Magrath 6381. FRANKLIN co.: 1 mi w on N 4 & 1.5 s of Campbell, roadside ditch, dry loam, rare, 29 September 1970, Magrath & Hays 6472. PAWNEE CO.: 1 mis, 5 e Burchard, weedy prairie pasture, dry rocky soil, 5 September 1967, Stephens 17523. RICHARDSON CO.: 1.4 mi e of Humboldt on N 4, roadcut & roadside ditch, rocky loam, common, 26 Sep- tember 1970, Magrath 6335. SARPY CO.: 0.8 min & 3 w on St. Spur N 150 & 2 s of Louisville on N 50, loess over sandstone, roadside ditch, uncommon, 27 September 1970, Magrath 6363. THAYER CO.: 2 mi w of Chester on N 8, roadside ditch, loam, rare, 29 September 1970, Magrath 6481. WEBSTER CO.: 1.4 mi e of Webster-Franklin Co. line on N 4, disturbed area in an open prairie hay mea- dow, loam, rare, 29 September 1970, Magrath & Hays 6475. These collections extend the range of this species northward from Kansas. Callitriche terrestris Raf. emend. Torr. KANSAS: MONT- GOMERY CO.: 12 mi sw Independence, at Camp Wilder- ness, along damp draw in oak woods, 20 June 1970, Hol- land 2812c, in part. This is an extension of the range of the species westward from Missouri and northward from Oklahoma. Anoda cristata (L.) Schlect. KANSAS: NEOSHO CO.: in South Mound, Lincoln Township, farmlot, 30 September 1970, Holland 2848. This species is native to South America and the southwestern United States. In the midwest, it is known to occur in Iowa and in McDonald Co., Missouri. Sida spinosa L. f. albiflora Magrath, form. nov. A forma spinosa differt petalis albis. TYPE: KANSAS: DOUGLAS CO.: West campus of Uni- versity of Kansas in Lawrence, growing along e edge of experimental greenhouse in loam soil, colony of 50-70 plants, 7 September 1970, Magrath 6147 (Holotype: KANU!). The white flowered plants were scattered in a large colony of plants that had typical yellow flowers. 384 Rhodora [Vol. 74 This white flowered form of S. spinosa was first observed in this same area by Vernon Harms in 1962. Harms (1965) performed crossing studies on this taxon and concluded that the white flowered form represented a homozygous recessive condition for a single gene. Hibiscus lasiocarpos Cav. KANSAS: BOURBON CO.: 1.5 mi e & 3 n of Hiattville, Bourbon Co. St. Lake, growing in marshy area along w side of lake, common, 25 August 1969, Magrath 4588; 7 August 1970, Magrath & Weedon 5746. CHEROKEE CO.: 2 mi s of Galena, in mixed woods of the flood plain woods of Shoal Creek, 5 August 1956, Lathrop 2863; 1 mi s of Riverton, low swampy area, abundant in local area, 12 August 1960, McGregor 15924; swampy area along Spring River, 26 August 1969, Bare 2169. The Bourbon Co. collections represent the first area in Kansas other than se Cherokee Co. where this species is known to occur. Thymelaea passerina (L.) Cosson & Germ. KANSAS: CLOUD CO.: 1 mi n Concordia, flood-plain of Republican River, sandy soil, willow thicket, scattered, 11 September 1970, McGregor 23572. This species has been reported for North America from west-central Iowa, northeastern Nebraska and extreme western Nebraska. European manuals indicate that the species is a common weed in dry soils and appears commonly in grain fields of south and central Europe and western Asia. Centunculus minimus L. KANSAS: MONTGOMERY CO.: 12 mi sw Independence, at Camp Wilderness, along damp draw in oak woods, 20 June 1970, Holland 2812c, in part. This represents an extension of the species westward from Missouri and northward from eastern Oklahoma. Glaux maritima L. NEBRASKA: SIOUX CO.: n edge of Agate, flat prairie swale near Niobrara River, sandy soil, 16 August 1970, Stephens 43966. This species occurs in North Dakota, but is not known from South Dakota. 1972] Great Plains — McGregor, Magrath & Weedon 385 Sabatia campestris Nutt. f. albiflora D. M. Moore. KAN- SAS: CHEROKEE CO.: 0.4 mi w & 1 s of Baxter Springs, in prairie in soil over limestone, abandoned rock quarry area, 17 July 1969, Johnson 2259, in part. MONTGOMERY co.: 2 mi s & 0.2 w of Sycamore, prairie hay meadow, sandy clay soil, rare, 12 July 1969, Weedon & Magrath 5516. WILSON CO.: 4.5 mi e Neodesha, sandy blue stem prairie, scattered, 4 July 1970, McGregor 22386. Eustoma grandiflorum (Raf.) Shinners f. album (Hol- zinger) Waterfall KANSAS: RICE CO.: 0.5 mi s of Sterling, heavily grazed pasture, sandy soil, one plant seen, 9 August 1970, Magrath & Weedon 5754. STAFFORD co.: T218, R11W, Sec 21, found in dense Spartina area, very rare, 13 August 1958, Ungar 666a. Apocynum medium Greene. KANSAS: DOUGLAS CO.: 14. mi s of Lone Star, thin soil over limestone, abundant on old slope of roadcut, 11 June 1969, Johnson 2081; 14 June 1970, Johnson 2496. This extends the range of this species westward from Missouri. Asclepias incarnata L. var. incarnata f. albiflora Heller. KANSAS: COFFEY CO.: 6.5 mi n & 1.5 w of Westphalia, growing near edge of large pond, moist loam, one plant, 5 September 1970, Magrath 6136. DOUGLAS CO.: in old roadbed at southern end of Baldwin Lake, 15 August 1961, Richards 3686; 1 mi s & 1.5 e of Baldwin City, growing at edge of lake, six plants, 19 August 1969, Bare 2114. NEBRASKA: DODGE CO.: Fremont Lakes Rec. Area, 3 mi w of Fremont on US 30, growing along se edge of lake, one white flowered plant in large colony of pink flowered plants, sandy loam, 21 August 1970, Magrath 5978. Hydrophyllum appendiculatum Michx. NEBRASKA: DOUG- LAS CO.: Omaha, wooded ravine s of Hummel Park, 6 June 1968, Sutherland 1689. This extends the range of this species northward from Kansas and northwest- ward from Missouri. 386 Rhodora [Vol. 74 Scutellaria brittonii Porter. NEBRASKA: KIMBALL CO.: 2 mi w Kimball, upland prairie, dry sandy-gravel soil, 14 June 1967, Stephens 11664; 8 mi sw, 1.5 w Bushnell, open pine-prairie hilltop, dry rocky clay soil, few plants, 5 June 1969, Stephens & Brooks 31479; 29 June 1970, Stephens & Brooks 40859. This species also occurs in Wyoming, Colorado and New Mexico. Leonurus marrubiastrum L. NEBRASKA: DOUGLAS CO.: 4 mi e of Venice on N 92, Elkhorn River, slip-off bank s of bridge, muddy silt with some sandy spots, rare, 21 August 1970, Magrath 5968. NEMAHA CO.: 8.1 mi s of Auburn on US 73 & 75, riverbed e of bridge, slip-off bank, silty-sandy clay, scattered, Magrath 5931. SOUTH DA- KOTA: HUTCHINSON CO.: 0.5 mi e Olivet, bank of James River, moist sandy soil, abundant, 8 September 1970, Stephens 44866. This extends the range of the species northward from Kansas and northwestward from Mis- souri. Lamium amplexicaule L. f. albiflorum D. M. Moore. KAN- SAS: DOUGLAS CO.: City of Lawrence, 2 blocks n of 23rd St. & 1 block e of Pennsylvania St., one plant, grow- ing in pasture, loam, 30 April 1971, W. & T. Vollmer, s.n. COFFEY CO.: 1 mi n, 7 w on US 50 of Waverly, roadside park, growing in ditch, loam, uncommon, 22 May 1972, Magrath 7407. Solanum sarrachoides Sendl. KANSAS: NEOSHO CO.: 4 mi s&be Erie, in farmyard near fence rows, 29 September 1970, Holland 2840. This is a South American species that has been introduced into North America. It is known otherwise only from Jasper and McDonald Coun- ties in Missouri and northeastern Oklahoma. Mimulus alatus Ait. f. albiflorus House. KANSAS: FRANK- LIN CO.: Ferndale, 1.5 mi s, 1 e & 1!4 s of Homewood, muddy creek floodplain near base of sandstone bluffs, one plant, 29 August 1970, Magrath 6125. LINN CO.: stream channel just n of Mound City, just e of bridge on K 7, 1972] Great Plains — McGregor, Magrath & Weedon 387 surfacing limestone, rare, 7 August 1970, Weedon & Magrath 6546. Agalinis heterophylla (Nutt.) Small. KANSAS: CHAU- TAUQUA CO.: 1.6 mi ne of Sedan, oak woods over sand- stone, rare, 14 September 1970, Magrath 6224. CHEROKEE CO.: 2.1 mi w of Jct of US 166 & 69 on 166, 1.6 s & 0.2 e, abandoned rock quarry, rocky limestone soil, common, 14 September 1970, Magrath 6243; 1 mi w Baxter Springs, scattered in prairie, 3 October 1970, MeGregor 22641. GREENWOOD CO.: 0.6 mi e of Sallyards on US 54, rocky prairie hay meadow, one plant found, 13 Septem- ber 1970, Magrath 6201A. This extends the range of the species westward from Lawrence and Dunklin Counties in Missouri and northward frcm Oklahoma. Orobanche multiflora Nutt. KANSAS: SCOTT CO.: 12 min & 3 w of Scott City, gravelly prairie hillsides with caliche outcrops, only one plant seen, 3 October 1970, Bare 2605. This extends the range of the species northward from Oklahoma and eastward from Colorado. Triodanis biflora (R. & P.) Greene. NEBRASKA: JOHN- SON CO.: 3.2 mis of Cook on N 50, grazed pasture, loam, rare, 28 June 1970, Magrath & Hays 5639A. PAWNEE CO.: 0.6 mi s of Pawnee-Johnson Co. line on N 50, prairie hay meadow, in disturbed area, loam, rare, 28 June 1970, Magrath & Hays 5642. This is an extension northward of this species from Kansas. Lobelia siphilitica L. var. siphilitica f. albiflora Britt. KANSAS: DOUGLAS CO.: 1.5 mi e of Lecompton, Kansas River bluff, moist ditch, 19 September 1968, Bare 1944. Iva axillaris Pursh. KANSAS: FINNEY CO.: river bank, 18 July 1912, Wilson & Miller, s.n. HAMILTON CO.: 1 mi s, 1 e of Coolidge, prairie roadside right-of-way, moist sandy soil, locally abundant, area 25 ft. across, 3 August 1971, Stephens 50368. These are extensions of the range of this species southward from Nebraska and eastward from Colorado. 388 Rhodora [Vol. 74 Helianthus mollis Lam. NEBRASKA: RICHARDSON CO.: 1.4 mi e of Humboldt on N 4, prairie pasture, loam, un- common, 26 September 1970, Magrath 6336A. WEBSTER co.: 2 mi s, 3 w Red Cloud, sandy prairie hillside, rare in the area, 11 August 1965, McGregor 19479. This is an extension of the range of the species northward from Kansas. Bidens comosa (Gray) Wieg. WYOMING: GOSHEN CO.: side channel of North Platte River w of city park in Torrington, sandy to clay loam soil, scattered, 28 September 1968, Weedon 4787; North Platte River on s edge of Torrington, both banks of river, sandy soil, scattered, 28 September 1968, Weedon 4789. WASHAKIE co.: Worland, in a wet saline area, 16 July 1966, Tresler 322 (RM). According to Sherff (1937), the distribution of this species is from Maine, Quebec and North Dakota southward and southwestward to North Carolina, Ten- nessee, New Mexico and Utah. It has also been reported from eastern Colorado. Artemisia annua L. KANSAS: DONIPHAN CO.: 1 mi n White Cloud, bank of Missouri River, willow thicket, several large colonies, 29 August 1970, McGregor 23467. This is the second known record for the state. LITERATURE CITED BARKLEY, T. M. 1968. A Manual of the Flowering Plants of Kansas. The Kansas St. Univ. Endowment Assoc. Manhattan, Kansas. Harms, V. L. 1965. Observations on the Breeding System of Sida spinosa (Malvaceae). Trans. Kan. Acad. Sci. 68(1): 123-24. HircHCOCK, C. L., A. CRONQUIST, M. OwNBEY, J. W. THOMPSON. 1961. Vascular Plants of the Pacific Northwest. Part 3: Saxi- fragaceae to Ericaceae. Univ. of Washington Press, Seattle. Komarov, V. L. ed. 1964. Flora of the U.S.S.R. Vol. III. pp. 328- 29. Pub. for Smithsonian Inst. and NSF by the Israel Program for Scientifie Translations, Jerusalem. SHERFF, E. E. 1937. The Genus Bidens. Field Mus. Nat. Hist. Bot. Ser. 16: 251. DEPARTMENT OF BOTANY UNIVERSITY OF KANSAS, LAWRENCE 66044 NEW RECORDS OF NORTH DAKOTA PLANTS O. A. STEVENS In 1961 I gave a ten-year supplement to my Handbook (Stevens 1950), which was incorporated as an (unpaged) appendix to the third printing of the book (1963). The intention was to provide a second supplement in 1970, but at that date active collecting was proceeding, especially in the western part of the state and this supplement was post- poned. In the meantime several partial lists have appeared and it seems desirable not to postpone this one longer. In addition to species which are reported for the first time (marked with *), a few significant additional loca- tions for other species are included. Mere changes in names are usually not mentioned. Botrychium lunaria L. was found in Burke County in 1971 by Glen Hegstad. *Equisetum palustre L. was reported by Barker (1971) for Ransom County. *Equisetum scirpoides Michx. was reported by Stewart (1970) from Barnes County. *Pinus banksiana Lamb. Several cone-bearing and seed- ling trees were found near McLeod, in Richland County in 1970 (Gerald Seiler 1775). This is presumably a forest planting but the record has not been found. *Bromus squarrosus L. Abundant in an abandoned, sandy field, Kidder County in 1971, Stevens 3292, 3294. *Chloris virgata Sw. This probably was an escape from S C S nursery at Mandan, Morton County, September 13, 1962, Geo. Karey. *Cinna arundinacea L. was collected at two locations in Ransom County, Stevens 2742, 2761. *Eragrostis spectabilis (Pursh) Steud. was reported by Barker (1971) from Ransom County. *Polypogon monspeliense L. is recorded from LaMoure County, H. Kantrud, September 30, 1970; Slope County in 1970, Zaczkowski 4000; Williams County in 1971, G. D. Hegstad. 389 390 Rhodora [Vol. 74 *Triplasis purpurea (Walt.) Chapm. was collected in two places in Richland County in 1964 (W. Wanek 124). *Sporobolus airoides Torr. is confirmed by N. Zaczkow- ski 3012, 3017, and 4257. *Carex alopecoidea Tuckerman was reported by Stew- art (1970). I had tentatively placed here a specimen from Richland County, Stevens 88. O. A. Kolstad at the Univer- sity of Kansas, who examined all of our North Dakota Carex in 1965, put this as C. stipata but part of a sheet of stipata in C. alopecoidea. Presumably all five stalks were from one clump (which proves nothing). Kolstad referred to alopecoidea one specimen from Barnes County, Stevens in 1950. *Carex capillaris L. Bottineau County in 1961, Stevens 2499. Carex davisii Schw. & Torr. Our specimens reported as C. gracillima were placed here by Kolstad. Stewart (1970) reported *C. buxbaumii, *C. lacustris, *C. lasiocarpa, *C. limosa and *C. vesicaria. *Iris missouriensis Nutt. A specimen was received, but not saved, from Mrs. Paul DeKrey, Tappen, Kidder County in 1968. G. D. Hoag visited the place and reported a num- ber of clumps in a depression of the prairie. Richard Wil- liams found it in northeastern Emmons County in 1971. *Juglans nigra L. can be regarded as naturalized because it is often planted and squirrels carry the nuts to other places. At least three seedlings appeared in our yard, pre- sumably from nuts from a tree across the street (one, re- planted about 15 years ago is still growing) and I have seen others at several places. *Juncus vaseyi Engelm. A specimen from Bottineau County was determined by L. J. Harms. Quercus ellipsoidalis E. J. Hill. Little (1971, map 163 E) shows this extending into Richland County but we have no knowledge of such (may be planted). Atriplex glabriuscula Edmonst. The specimens reported as this (Stevens, 1963) are now referred to A. hetero- sperma Bunge by Aellen and by Frankton (1968) who 1972] North Dakota Plants — Stevens 391 uses A. subspicata (Nutt.) Rydb. for what we had called A. hastata, but retained one specimen from Richland County, Bell 377, as A. hastata, also one from Clay County, Minnesota. *Lychnis chaleedonica L. We have two Specimens, pre- sumably escapes. Benson County, J. Lunell in 1917 “bor- dering a slough”, and Dickey County, K. M. Larson in 1965. *Salix pedicillata Pursh was collected in McHenry County, Stevens 3225, in the fen cited by Stewart (1970). Eriogonum gordoni Benth. was reported as E. trichopes (Stevens 1960) but apparently is E. gordoni or close to it. Further specimens are from Burke and Mountrail Counties in 1970 by Hegstad. Anemone multifida Poir. from Golden Valley County was a surprise, N. Zaczkowski 364, 3481. * Ranunculus cardiophyllus Hook. A specimen collected in McKenzie County in 1967 by C. A. Barr seems to be this species. Alyssum desertorum Stapf. apparently is the proper name for that reported (Stevens 1963) as A. alyssoides. Better specimens are from the same county by Zaczkowski. *Arabis glabra Gray. Two specimens from Dunn County, Stevens & Moir 2255 and Sept. 14, 1956, and one from Burke County, Stevens, July 20, 1966, seems to belong here. Chorispora tenella (Pall.) D.C. A second record is from Billings County, Zaczkowski 3333. “Erysimum repandum L. A single specimen collected on the campus in 1967 seems to be this species. *Berberis repens Lindl. was reported by Barker (1971) from Bowman County. *Ribes cereum Dougl. A sterile specimen was collected by Jack Nelson at Pretty Butte, western edge of Slope County, in 1960. Nick Zaczkowski collected flowering specimens there in 1970. *Potentilla palustris L. was reported by Stewart (1970) from McHenry County. *Astragalus purshii Dougl. was reported by Zaczkowski 392 Rhodora [Vol. 74 (1970). It was identified by Fernald from a specimen which I collected in McKenzie County July 15, 1943. An- other was collected in Emmons County in 1945. Walsh (1960) referred these to A. lotiflorus. Caragana arborescens Lam. Glen Hegstad, who has been studying our northwestern counties extensively, reports apparent establishment. This area had many farmsteads abandoned after 1934. Lotus corniculatus L. has become quite common along streets and lawns in the Fargo area. Polygala senega L. Three collections by Hegstad in Burke County, 4770, 5413, 8038, extend the known range of this species. * Euphorbia stictospora Engelm. A specimen, Bell 1072, said to have been collected in the dooryard of a ranch in 1907 in McKenzie County was determined as this by Rich- ardson (1968). It must have been a chance introduction. *Rhamnus davurica Pall. In 1965 I saw in Ransom County a windbreak of this, containing great numbers of seedlings. A second record is from O. J. Lung in 1964 from Emmons County (“came up in garden”). The fruits seem not usually eaten by birds. Acalypha rhomboidea Raf. A second specimen is from Stutsman County, Sept. 3, 1964, Mrs. Amy Rolling, “came up in flower bed". *Acer saccharinum L. is often planted as a street tree. Two to three year old seedlings were collected by a hedge near a large maple tree in 1970, Stevens 3285. *Lavatera cretica L. was collected in Divide County on Aug. 20, 1969 by L. M. Mitich and by Hegstad (6239) in Burke County in 1970; escaped from, or persisting from, gardens. Hypericum perforatum L. A second specimen, Stevens 3152, in Ransom County on the Sheyenne River bank by an old farm house, may have been planted. There were sev- eral plants but no new ones. *Gaura parviflora Dougl. reported by Barker (1971) 1972] North Dakota Plants — Stevens 393 from Golden Valley County is probably a railroad intro- duction. Conium maculatum L. was collected in Golden Valley and Billings Counties in 1969 and 1970 by Zaczkowski. Osmorhiza claytoni (Michx.) C. B. Clarke. A second record was by V. Facey, Grand Forks County in 1958. *Halenia deflexa (Sm.) Griseb. was reported by Barker (1971) from Pembina County (KANU). Phlox alyssifolia Greene was collected on Bullion Butte, Stark County, Zaczkowski 2329 in 1970. Physostegia parviflora Nutt. I am inclined to think that our plant should be considered a variety of P. virginiana L. It is uniformly leafy, the leaves 1-2 (—5) em wide, sharp- ly toothed, flowers 15 (—18) mm long. Specimens of P. formosior Lunell, so labeled by Lunell, do not differ mate- rially, but one from the same area labeled P. parviflora has the lower leaves narrowly oblong with blunt, low teeth. One from McLean County (V. T. Heidenreich 210) has leaves about 7 mm wide and some from Burke County (Hegstad 8458) have narrow leaves and blunt teeth but his 5943 from within ten miles of the same place agrees with our eastern specimens. *Lythrum salicaria L. is much planted and a consider- able colony develeped in a low, roadside area at Fargo (Stevens 2332). I found one plant at a similar place about one mile distant. Stachys palustris L. 'The prevalent form in open, usually moist ground, has rather densely, evenly pilose stems (S. scopulorum Greene?). Along wooded streams in the east- ern edge of the state the mid and lower stems often are nearly glabrous, with a few, retrorse, coarser, often some- what pustulate, hairs on the angles and a few or more soft hairs on the sides. The calyx varies from sparsely to more densely pilose, rarely with short, glandular hairs. *Collinsia parviflora Lindl. was found in Slope County, Zaczkowski 3072. *Pentstemon procera Dougl. was reported by Barker (1971) from Burke County. 394 Rhodora [Vol. 74 *Chaenorrhinum minus (L.) Lange was collected along a formerly used railway track at Valley City, Barnes County in 1963, Stevens 2670. The area was pastured and plants not seen in 1971. Veronicastrum virginicum (L.) Farwell. Roys Willen- bring, collecting in Pembina County 1969-71, found a few plants near Cavalier, Pembina County, 799. *Galium verum L. One specimen is from Dickinson, Stark County in 1912, C. H. Waldron 130. I recall that perhaps about 1935, Dr. L. R. Waldron showed me a frag- ment of it. In 1967 I discovered a colony of it in a yard near his house, and distributed several sheets, Stevens 3028. In 1970 I looked again but did not find it. *Sambucus pubens Michx. In 1964 I visited a farm grove near Colfax, Cass County, where this plant had apparently been introduced and spread through the grove and along fence lines. Some years ago it had spread through the uni- versity’s woody plantings but was eradicated. * Ambrosia (Franseria) tomentosa (A. Gray) Payne. A colony appeared in garden plots of the University at Fargo in 1963 (Stevens 2693). Another specimen is from Grand Rapids, LaMoure County, Bill Shak, Aug. 21, 1921. Aster ciliolatus Lindl. This name apparently should be used for what has been called A. sagittifolius L. I have been unable to make satisfactory conclusions. Bidens acuta (Wieg.) Britton. I still feel that this is a better “species” than many others. In our area it is quite striking, usually with sessile leaves, heads the size of those of B. vulgata with 2-3 cm long, leafy, outer bracts. The long-petioled leaves of B. comosa are lacking. Centaurea maculosa Lam. A large colony of an acre or more was found along State Highway 1 in Nelson County at the outlet of Stump Lake in 1962, Stevens 2962. *Chaenactis douglasii (Hook.) H. & A. This is another extension from the west in Bowman County, Zaczkowski. Crepis tectorum L. I had used the name C. capillaris, first on a specimen by Lunell from Bottineau County, but I believe it is C. tectorum which is quite common in the 1972] North Dakota Plants — Stevens 395 Cavalier-Pembina area and spreading more widely. It is also abundant in western Becker County, Minnesota. *Erigeron canbyi. Specimens from the Killdeer Mts., Dunn County, by C. A. Barr, May 8, 1963, were considered this form by Mr. Barr. Hymenopappus tenuifolius Pursh. Collections from Ben- son County, W. T. Barker 5567, make a notable eastern range extension. *Senecio vulgaris L. was received from Dickinson, Stark County in 1954; Stevens 1558 in 1955. Another garden lo- cation is at Fargo, Cass County in 1962, Stevens 2648, 2870. It had been found at Hawley, Clay County, Minne- sota. *Thelesperma marginatum Rydb. was collected in Wil- liams and Divide Counties, Hegstad 3095, 3552, 5'798, 8030, in 1969-71. REFERENCES BARKER, W. T. 1971. Vascular plants new to North Dakota. Rhodora 73: 58, 59. FRANKTON, C. and I. J. BassETT. 1968. The genue Atriplex in Canada. I. Three introduced species — Canad. Jour. Bot. 46: 1309-1313. LITTLE, E. J. 1971. Atlas of United States Trees. VoL 1. UTS. Dept. Agr. Misc. Publ. 1146. RICHARDSON, J. W. 1968. The genus Euphorbia in the high plains of Kansas and prairie plains of Kansas, Nebraska, South and North Dakota. Univ. Kan. Bull. 48: 45-112. STEVENS, O. A. 1950. Handbook of North Dakota Plants. Institute of Regional Studies, North Dakota State University, Fargo. - 1961. New records of North Dakota plants. Rho- dora 63: 39-46. 1963. Handbook of North Dakota Plants. Insti- tute of Regional Studies, North Dakota State University, Fargo. (Third printing) STEWART, R. E. 1970. New records of vascular plants in North Dakota. The Prairie Naturalist 3: 35-48, Warp, R, and W. T. BARKER. 1971. Cytotaxonomic studies of the Scirpus lacustris complex in North Dakota. Proc. N. Dakota. Acad. Sci. 25: 32. 396 Rhodora [Vol. 74 WELsH, S. L. 1960. Legumes of the north-central states: Galegea. Iowa St. Coll. Jour. Sci. 35: 111-249. ZACZKOWSKI, N. K. and W. T. BARKER. 1971 (abstr.). Floristic survey of southwestern North Dakota. Proc. N. Dakota Acad. Sci. 25: 32. NORTH DAKOTA STATE UNIVERSITY FARGO, N. D. 58102 PRIORITY OF THE BINOMIAL PYROLA CHLORANTHA ERICH HABER Some confusion exists in the literature concerning the usage and priority of the binomials P. chlorantha Swartz and P. virens Schreber in Schweigger & Koerte. The most commonly used binomial for the green-flowered shinleaf in Europe up to and including the present time, and in North America until 1941, was P. chlorantha (Swartz, 1810). Fernald's short note (1941) in which he cited P. virens as antedating P. chlorantha by six years resulted in the com- mon acceptance of the former name in most North Amer- ican publications. However, the Flora Erlangensis of 1804 does not mention Pyrola virens. It becomes clear, however, that P. chlorantha has priority over P. virens when the publication containing the orig- inal description of P. virens is closely examined ( Schweig- ger & Koerte, 1811). The 1811 publication, Flora Erlan- gensis, is composed of two parts plus an Addenda et Cor- rigenda to Part I. The pagination of the addendum is sequential to that in Part I which is taken directly from a doctoral dissertation by Schweigger (1804). A. F. Schweigger and F. Koerte published Flora Erlan- gensis in 1811 at which time the addendum as well as Part II were bound with a facsimile of the 1804 publication. The title page for the addendum bears neither of the au- thors’ names, whereas the author of Part I is clearly indi- cated as being Schweigger and the authors of Part II as being Schweigger and Koerte. In the preface to the com- 1972] Pyrola — Haber 397 plete volume written by Koerte, it is stated that all the additions which are preceded by an asterisk are the work of J. C. D. von Schreber, who had reviewed the manuscript. P. virens on p. 154 of the addendum is preceded by an asterisk. There appears to be no question that the valid date of publication of P. virens is 1811, one year after the publication of the binomial P. chlorantha. The correct ci- tation for the former is as follows: P. virens Schreber in Schweiger & Koerte, Fl. Erlang. I, add. p. 154. 1811. The author would like to express his appreciation to Dr. John W. Grear Jr. for his assistance in obtaining repro- ductions of pertinent sections of the origina] publications by Schweigger and Koerte. LITERATURE CITED FERNALD, M. L. 1941. Transfers in Pyrola, Rhodora 43: 167. ScHWEIGGER, A. F. 1804. Specimen Florae Erlangensis. Erlangen, Hilpert. ScHWEIGGER, A. F. & F. KoERTE. 1811. Flora Erlangensis. Erlan- gen, Palm. Swartz, O. P. 1810. Sv. Vet. Acad. Stockh. Handl. 31: 190. NATIONAL MUSEUM OF NATURAL SCIENCES BOTANY DIVISION OTTAWA, ONTARIO K1A0MS8 CHROMOSOME NUMBERS OF SCIRPUS PURSHIANUS AND S. SMITHII ALFRED E. SCHUYLER Two eastern North American aquatic bulrushes, Scirpus purshianus Fern. and S. smithii Gray, superficially resem- ble one another and have been confused in taxonomic lit- erature. Beetle (1942) stated that, “Apparently S. smithii is no more than an aquatic extreme of S. debilis [an earlier name applied to plants of S. purshianus]," while Fernald (1942) maintained, “. .. all living botanists who accurately know living plants in the field find them abundantly dis- tinct." Some botanists (Gleason and Cronquist, 1963) have continued to treat these species in accordance with Beetle's statement and others (Seymour, 1969) have con- formed with Fernald's statement. The characteristics used by Fernald to delineate S. purshianus, S. smithii, and their forms are accepted here and the following discussion is in accordance with his treatment of these taxa in Gray’s Manual (Fernald, 1950). Part of the taxonomic confusion with S. purshianus and S. smithii is probably caused by the similar pattern of in- fraspecific variation of each species. Each has ecologically differentiated forms characterized morphologically by the presence or absence of well-developed barbed bristles at- tached to the achenes. S. smithii, which often appears to be associated with somewhat alkaline substrates, has three described forms: f. levisetus (Fass.) Fern. (having nearly smooth bristles), f. setosus (Fern.) Fern. (having well- developed barbed bristles), and f. smithii (lacking bris- tles). S. purshianus, which often appears to be associated with more acid substrates, has two forms: f. purshianus (having well-developed barbed bristles) and f. williamsii (Fern.) Fern. (lacking bristles). In both S. purshianus and S. smithii, the presence or ab- sence of barbed bristles appears to be correlated with the stability of the habitat. For example in eastern New York, New Jersey, eastern Pennsylvania, and Delaware, S. smithii 398 1972] Chromosome Numbers — Schuyler 399 f. smithii is almost completely restricted to the specialized conditions of fresh inter-tidal zones of the Hudson, Raritan, and Delaware Rivers while in the same area f. setosus occurs in and around lakes, ponds, and quagmires (see Long, 1910, for a discussion of these forms in the vicinity of Philadelphia). S. smithii f. levisetus is common in the estuaries of the Kennebec River in Maine and the St. Law- rence River in Quebec. Although bristles are present in f. levisetus, they lack the prominent barb-like teeth char- acteristic of f. setosus. Thus it appears that plants of S. smithii which lack barbed bristles are better adapted to the periodic fluctuation in water level characteristic of the estuarine environment than are those which have barbed bristles. A similar pattern emerges for S. purshianus f. purshianus and f. williamsii, although tidal fluctuation in water level is not involved since neither form grows in tidal estuaries. However, f. williamsii is frequently found on the sandy shores of lakes in Cass, Kalamazoo, and Van Buren Counties in southwestern Michigan while the more widespread f. purshianus is infrequently found in this area. There is (or was) considerable water level fluctuation in these southwestern Michigan lakes from spring to fall, a factor which probably is a barrier to plants adapted to lake shores with more uniform water levels, This may be what excludes f. purshianus from many southwestern Michigan lakes since, throughout most of its range, it often grows along lake shores with relatively uniform water levels. Thus the forms of both species having barbed bris- tles appear better adapted to relatively stable conditions associated with little or moderate water level fluctuation while the forms lacking barbed bristles appear better adapted to relatively unstable conditions associated with substantial water level fluctuation. Despite the ecological and morphological resemblance be- tween forms of these two species, I have always found that the morphological criteria used by Fernald to distinguish them are reliable. In addition, the chromosome numbers determined for the forms of each species provide evidence [Vol. 74 Rhodora ‘(9224 Lajfinyog) ijs ‘S Jo soMOsOWOLYyD 9n019]p— "6 SIT '(e9et «4ojfim[og) snupu[S4nd “gS Jo səwosowoaryəð 2120I9]A T SA 1972] Chromosome Numbers — Schuyler 401 supporting Fernald’s treatment. 19 meiotic units (Fig. 1) were observed in pollen mother cells of S. purshianus f. purshianus and f. williamsii (Table 1). 20 meiotic units, with one unit consistently larger than the rest (Fig. 2), were observed in pollen mother cells of S. smithii, f. levi- setus, f. setosus, and f. smithii (Table 1). Ordinarily, chromosome numbers are consistent for a given species of Scirpus and provide evidence corroborating distinctions among them. The chromosome numbers reported here sup- port the specific recognition of S. purshianus and S. smithii and indicate that the infraspecific taxa are associated with the proper species in Fernald's treatment. TABLE 1. Chromosome Numbers Determined for Forms or Scirpus purshianus and S. smithii Number of Taxon Meiotic Units Voucher Specimens? S. purshianus f. purshianus 19 Massachusetts: Barnstable Co., 4265; Pennsylvania: Northamp- ton Co. 3819; Pennsylvania: Berks Co., 4088 S. purshianus f. williamsii 19 Michigan: Kalamazoo Co., 3903 S. smithii f. levisetus 20 Maine: Sagadahoc Co., 4276 S. smithii f. setosus 20 New Jersey: Sussex Co., 4201 S. smithii f. smithii 20 New Jersey: Burlington Co., 4203 ^A]] are collections by the author preserved at the Academy of Natural Sciences. ACKNOWLEDGMENTS I am grateful to George H. Lauff for use of facilities at the W. K. Kellogg Biological Station during the summer of 402 Rhodora [Vol. 74 1966. Vincent Abraitys, Florence M. Givens, Patricia R. Schuyler, and Henry K. Svenson helped in connection with field work, Karen Keil did chromosome squashes, and Helena Greenwood typed the manuscript. LITERATURE CITED BEETLE, A. A. 1942. Studies of the Genus Scirpus L., V. Notes on the Section Actaeogeton. Am. Journ. Bot. 29: 653-656. FERNALD, M. L. 1942. Transfers in Scirpus $ Actaeogeton. Rho- dora 44: 479-484. 1950. Gray's Manual of Botany. 8th ed. Ameri- can Book Co. New York. lxiv + 1632 pp. GLEASON, H. A. & A, CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. Van Nostrand Co., Ine., Princeton. li + 810 pp. LONG, B. 1910. Range Extension of Scirpus smithii var. setosus. Rhodora 12: 155-156. SEYMOUR, F. C. 1969. The Flora of New England. Charles E. Tuttle Co., Rutland. xvi + 596 pp. DEPARTMENT OF BOTANY ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA PHILADELPHIA, PENNSYLVANIA 19103 TUSSILAGO FARFARA L. (SENECIONEAE-ASTERACEAE) IN VIRGINIA MILES F. JOHNSON Tussilago farfara, the Coltsfoot, a perennial rhizomatous herbaceous native of the Old World with a single yellow head terminating a scaly bracted scape in early spring, and developing large cordate leaves later, was seen in a newly developed City Park in Richmond, Virginia, in April, 1971. A survey of the manuals for this area (Fernald, 1950; Gleason & Cronquist, 1963; Harvill, 1970) showed T. far- fara absent from Virginia. However, a survey of other literature and herbaria proved that the Richmond speci- men was not a new state record. This paper is based upon collections at herbaria of the following institutions: Virginia Polytechnic Institute and State University, Blacksburg, Lynchburg College, Lynch- burg, College of William and Mary, Williamsburg, Virginia Commonwealth University, Richmond, and the United States National Museum, Smithsonian Institution, Washing- ton, D.C. I wish to acknowledge aid of the directors and (or) curators at these herbaria. The first known collections of Tussilago in Virginia in- clude Freer 2627 (Lynchburg, US), from route 39 near McClung, Bath County, Freer 2628 (Lynchburg, US) col- lected from the west side of Lantz Mountain, Highland County, Freer 2686A (VPISU) from Rockbridge County and Gwynn s.n. (VPISU, US) from Warren County; all specimens were collected in 1962. See Fig. 1. The first known pub- lished report of Tussilago in Virginia is that of Freer (1968) from Rockbridge County. A general migration of Tussilago farfara in the United States from north to south is noted in Fernald (1.c.) and Gleason & Cronquist (1.c.). Fernald (1.c.) lists the species ranging from Newfoundland to its southern limits in New Jersey, Pennsylvania and Ohio though Reed (1951) re- cords Tussilago in Baltimore County, Maryland, as early 405 404 Rhodora [Vol. 74 as 1903 and Strausbaugh and Core (1964) report this spe- cies in West Virginia in 1933. Gleason and Cronquist (1.c.), publishing 13 years later than Fernald, indicate a southern range extension to Maryland and West Virginia. It is apparent that the southern migration of Tussilago is continuing at present. This species entered West Vir- ginia in 1933 in Monongahela County which borders on southwestern Pennsylvania and by 1964 was abundant in Kanawha County in southern West Virginia and probably in every county of the state (Strausbaugh and Core, 1.c.). Migration continued and Tussilago seems to have entered Virginia from the west in the early 1960’s (Fig. 1). The migration pattern within Virginia perpetuates the trend southward and eastward (Fig. 1). It is noted that the earlier collections were made in the mountains and that the majority of the recent ones have come from the Piedmont and Coastal Plain. The plant in Richmond was seen in sandy, disturbed soil immediately adjacent to the James River. The river, the headwaters of which rise in western Virginia, may well have transported the diseminule from which the specimen grew. Thus, the range of Tus- silago farfara in the United States should be extended to include Virginia and this interesting species should become part of the treatment of the Flora of Virginia now in prep- aration. Figure 1. Distribution of Tussilago farfara in Virginia. Dates indicate the earliest known collection from each county. 1972] Tussilago — Johnson 405 LITERATURE CITED FERNALD, MERRITT LYNDON. 1950. Gray’s Manual of Botany eighth edition. American Book Company. New York. 1632 p. FREER, R. S. 1968. Plants of the Central Blue Ridge: Supplement II. Castanea 33: 163-193. GLEASON, H. A., AND ARTHUR CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. Van Nostrand Company, Inc. Princeton. 810 p. HARVILL, A. M. 1970. Spring Flora of Virginia. McClain Printing Company. Parsons, West Virginia. 240 p. REED, C. F. 1951. An Early Record of Tussilago farfara in Mary- land. Castanea 16: 11. DEPARTMENT OF BIOLOGY VIRGINIA COMMONWEALTH UNIVERSITY RICHMOND, VIRGINIA 23220 INVESTIGATIONS OF NEW ENGLAND MARINE ALGAE V. THE ALGAL VEGETATION OF THE HAMPTON-SEABROOK ESTUARY AND THE OPEN COAST NEAR HAMPTON, NEW HAMPSHIRE! ARTHUR C. MATHIESON AND RICHARD A. FRALICK Most botanical studies of New England salt marshes have been restricted to Massachusetts (e.g. Chapman, 1940; Drum and Webber, 1965; Webber, 1967, 1968; Webber and Wilce, 1971) and Connecticut (e.g. Miller and Egler, 1950) and little is known of these valuable wetlands in New Hampshire salt marches (i.e. Davis, 1956; Vagenas, 1969). Davis sum- marized the general ecology of the Crommet Creek Estuary, which is a small tributary of Great Bay, New Hampshire. Vagenas studied the seasonal occurrence, composition and zonation of algae and flowering plants at five locations in the Hampton-Seabrook salt marsh. For the past seven years we have conducted a variety of floristic and ecological stud- ies of the marine algae on the open coast of New Hampshire and the Great Bay Estuary System (Mathieson, Hehre and Reynolds, in press; Mathieson, Reynolds and Hehre, in press; Hehre and Mathieson, 1970). The present investi- gation was initiated, at the request of Normandeau Associ- ates Incorporated, in order to provide a broad “baseline” of information regarding the species composition, distribution and ecology of the benthonic marine algae of the Hampton- Seabrook Estuary and the adjacent open coast of New Hampshire (Fig. 1). Collections and observations of marine algae were made at fifty-four stations in the Hampton-Seabrook Estuary and the adjacent open coast of New Hampshire (Fig. 1 and Table I) during the summer and fall of 1969. Prior to 1969 seasonal collections had been made on the open coast near ‘Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as scientific contribution number 534. 406 1972] Marine Algae — Mathieson and Fralick 407 Bound Rock, and limited collections were made in the es- tuary (during the fall and winter of 1966). The majority of the stations were established by Normandeau Associates in conjunction with their ecological survey of the estuary. The estuarine sites were identified by a pair of numbered stakes on opposite sides of the channel, and most of the col- lections were restricted to the side of the channel numbered on the map (Fig. 1) unless conspicuous differences were noted on the two sides. The locations were primarily shore sites, but a few offshore outcrops of rock were also studied (Fig. 1). An aluminum boat and outboard motor were used at most stations. Representative samples of all conspicuous algal species were obtained at each station. Collections were made in the littoral (on foot) and sublittoral zones (by SCUBA). Ex- tensive collections were made in the sublittoral zones on the open coast, but because of poor visibility, tidal currents and limited substrate for seaweed attachment, fewer sublittoral samples were available in the Estuary. A complete set of nearly 700 herbarium voucher specimens has been deposit- ed in the Algal Herbarium of the University of New Hamp- shire (NHA). The nomenclature of the Second Revised British Checklist (Parke and Dixon, 1968) has been applied in most cases. The vertical distribution of the major algal components described in this article is given according to the biological classification of Lewis (1964). A record of surface water salinity and temperature was made at most stations with a thermometer and a set of hy- drometers. All of the salinity values were corrected to 15 C. Diurnal variations of temperature, salinity and current speed were recorded at one location (station C-9) during a twelve hour period. The current speed was recorded with a Little Captain boat speedometer (Swift Instrument Com- pany of Boston, Massachusetts), which was modified with a six-foot well tube. Table II summarizes the surface water temperature and salinity values recorded (1966-1969). The hydrographic conditions on the exposed open coast (Bound Rock) are 408 Rhodora [Vol. 74 HAMPTON— SEABROOK AREA, NEW HAMPSHIRE 42°56 | . YS a A17 A9 T TN AB D Gad k S E m . - . eS . A3 b Al » f^ Pa PR, WI fy we Ep uA HAMPTON BEACH e x 2 Bil B2 We B7 B6 Z BIS a e oe * e FS) BIO B9 A AS BS 0 °B4 cS H4 e s - B8 *n | H3 ! ASS 82 HBIa* [LÀ 4 x *N e// se HAMPTON HARBOR 71 ES HB2*,* ES Hi e 2" "HB3 ^ FAJ . HB - B3 SS . BOUND x ROCK a" D2 Di = Rocks H2 Y M 7 p^ 4 S. & cio a 0.9 qe . f= a SEABROOK BEACH "3 (GJ e CI N e 0 is s : al E c 5 2 9» A c3 . le C4 C5 y sN . oe) ce ao d ! Mil N EN ile —— c8 c9 . 42 52 70 51 oot f 704 n 1 8 1972] Marine Algae — Mathieson and Fralick 409 more uniform than in the Estuary — particularly at the headwaters of the tidal rivers. Gross fluctuations (both daily and seasonally) of temperature and salinity are evi- dent at any estuarine location. Figures 2 and 3 summarize the results of a twelve hour study made at station C-9 on the Blackwater River. The location has a pronounced tidal current; thus, vertical mixing of the water column is evi- dent and there is no stratification. Diurnal variations of temperature and salinity varied from 13.2 to 16.0 C and 11.5 to 14.5 0/00, respectively (Fig. 2). The maximum sa- linity and the minimum temperature were recorded at slack water. Diurnal variations of tidal currents are shown in Figure 3. The current varied from 0-4.8 knots. A variety of substrates is evident at the stations. In gen- eral there is a gradient from sandy beaches with rock out- crops (i.e. Bound Rock which is on the exposed open coast) to sandy-muddy locations (i.e. at the mouth of the Estuary) to muddy and peat-like substrates (i.e. at the head of the tidal rivers). Only limited quantities of solid substrates (rock outcrops, boulders and breakwaters) are present in the Estuary. There are pronounced seasonal fluctuations of sand levels on the open coast near Bound Rock, and the abrasive action of sand limits both the species diversity and biomass of algae. Even so there are many more species at Bound Rock than within the Estuary. The conspicuous vegetation at Bound Rock is composed of Cladophora sericea, Enteromor- pha intestinalis, Pseudendoclonium marinum, Ulva lactuca, Fucus distichus ssp distichus, F. vesiculosus, Petalonia fas- cia, Ralfsia verrucosa, Scytosiphon lomentaria, Ahnfeltia Figure 1. The Hampton-Seabrook Estuary lying between latitudes 42° 51’ 30" to 42° 55’ 55" north and longitudes 70° 49’ 30" to 70° 51’ . 80" west within the townships of Hampton, Hampton Falls and Sea- brook, New Hampshire. Note that all of the tidal waters enter and leave via the Hampton Harbor Inlet and that five rivers (Taylor, Hampton Falls, Hampton, Brown and Blackwater) as well as many smaller creeks and brooks are present within the Estuary. Dots with numbers show the stations. 410 Rhodora [Vol. 74 DIURNAL VARIATION OF TEMPERATURE ANO SALINITY AT STATION C79 l6 4 4 c H 28 K l4 4 x m ~ = » E K - æ E : i = ri = 12 4 a ^ m 24 10 7 OCT.9, 1969 [ T T T T T T T T T T T T 7 9 n 13 15 7 TIME Figure 2. Diurnal variation of temperature and salinity at station C-9. DIURNAL VARIATION OF CURRENTS AT STATION C-9 5 4 4 4 a 931 c m m m z 2 4 A 2 14d o = wu O 4 OCT.9,1969 T T T T T T T T T T T T 7 9 1 13 15 17 TIME Figure 3. Diurnal variation of currents at station C-9. 1972] Marine Algae — Mathieson and Fralick 411 plicata, Chondrus crispus, Corallina officinalis, Hilden- brandia prototypus and Porphyra umbilicalis. Some species (e.g. Polyides rotundus and Ahnfeltia plicata) are restrict- ed to such sandy areas. Other plants which are not “sand loving” forms are often reduced in size, and they are only found in cracks and crevices. Several species are restricted to tide pools, which are somewhat protected from the fullest force of the sand and surf. Fucus vesiculosus domi- nates the eulittoral zone. Ascophyllum nodosum is con- spicuously absent. Several common sublittoral species such as Alaria esculenta, Laminaria saccharina, L. digitata and Desmarestia viridis are rare at Bound Rock. However, they are present in greater abundance in the offshore rocks near Hampton Beach (station HB1-HB4 in Fig. 1). A comparison of the common plants at Bound Rock with those of a “typical” estuarine site shows several differences. A variety of fucoid (Ascophyllum nodosum, A. nodosum f. scorpioides and Fucus vesiculosus var. spiralis) green (Cladophora sericea, Enteromorpha erecta, E. intestinalis, Monostroma oxyspermum, Percursaria percursa, Rhizo- clonium riparium and Ulva lactuca), blue green (Lyngbya spp, Oscillatoria spp and Gomphosphaeria spp.) and yellow green algae (Vaucheria sp.) dominate the littoral zone. In addition Spartina alterniflora, S. patens and Distichlis spicata are abundant in the upper portion of the shore. Results A total of 118 taxa of benthonic marine algae was col- lected from the fifty-four stations, including 30 Chlorophy- ceae, 35 Phaeophyceae and 53 Rhodophyceae. A detailed examination of the Cyanophyceae, Bacillariophyceae and Xanthophyceae was beyond the scope of the present investi- gation. Table III summarizes the number of seaweed taxa found at each of the stations. The largest number of taxa was found at Bound Rock with fewer towards the head of the Estuary. Gross fluctuations of species numbers were recorded at different locations within the Estuary. The following catalogue of Species includes some synony- 412 Rhodora [Vol. 74 my, notes on collection sites and habitats and noteworthy taxonomic features. The occurence of each species is de- signated numerically as follows according to its habitat (Hehre and Mathieson, 1970; Mathieson, Hehre and Rey- nolds, in press; Mathieson, Reynolds and Hehre, in press) : #1 Coastal — restricted to the open coast #2 Estuarine — restricted to the estuarine environment #3 Cosmopolitan — present in both estuarine and open coastal environments. Several of the cosmopolitan species may not occur in both habitats in the Hampton-Seabrook area, but they have been recorded from either the Great Bay Estuary or the open coast adjacent to Bound Rock. List of Species Chlorophyceae : Blidingia marginata (J. Agardh) P. Dangeard: equals Enteromorpha marginata J. Agardh in Taylor, 1957) Found once on rocks in the lower eulittoral zone at Bound Rock. #3 Blidingia minima (Nàgeli ex Kützing" Kylin: (equals Enteromorpha minima Nageli in Taylor, 1957) Found once in the estuary (B9) on a muddy bank in the upper littorcl zone. Abundant on the exposed open coast (Bound Rock) where it forms fringe zones. #3 Bryopsis plumosa (Hudson) C. Agardh Locally abundant in the estuary (A3, A5, A9, A15, A17, B1, B2, B4 to B6, B8, and H3) on muddy surfaces in the sub- littoral and lower eulittoral zones. Not found on the open coast. #2 Chaetomorpha aerea (Dillwyn) Kützing: Collected three times from the lower eulittoral (in tide pools) at Bound Rock. #1 Chaetomorpha linum (O. F. Muller) Kützing: (including Chaetomorpha atrovirens Taylor, in Taylor, 1957) 1972] Marine Algae — Mathieson and Fralick 413 According to our interpretation C. atrovirens and C. linum are not distinct for there is a continuous gradient of size and color between the two. Chaetomorpha linum is the old- er name and it should be retained. The plant is common as an entangled mass amongst various algae in the lower eulit- toral and sublittoral zones of the estuary (A2, All, A18, B1, B2, B4 to B6, B9, B10, B12, C1, C2, C6, C7, and C9) and the exposed open coast (Bound Rock, HB1 and HB4). £3 Chaetomorpha melagonium (Weber et Mohr) Kützing: Occasional on rocks in the lower eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1 and HB4). £3 Cladophora sericea (Hudson) Kützing sensu van den Hoek, 1963: Locally abundant in high tide pools in the estuary (A6, AT, A9, A10, B3, B7, B10, B12, C3 to C5, and H3) and on the exposed open coast (Bound Rock). £3 Codiolum pusillum (Lyngbye) Kjellman in Foslie: Locally abundant on rocks in the upper littoral zone of the exposed open coast (Bound Rock). Often mixed with Bangia f uscopurpurea and various blue green algae. It may be the sporophyte stage of one or more local species of Urospora (Scagel, 1966). £1 Enteromoprpha compressa, (L.) Greville: Found once on rocks in the lower eulittoral zone at Bound Rock. £3 Enteromorpha erecta (Lyngbye) J. Agardh: Abundant on muddy surfaces of the eulittoral zone through- out the estuary (Al to A9, A13, A14, A16, A17, B1 to B7, B9, B13, C1 to C6, H1 and H3) ; occasionally present as an epiphyte on fucoid algae and Spartina alterniflora. Uncom- mon on the exposed open coast (Bound Rock). #3 Enteromorpha intestinalis (L.) Link: Common on rocks and muddy surfaces of the eulittoral zone throughout the estuary (Al, A2, A4, A6 to A10, A13, A17, B1 to B5, B8, C1, C3, C5 to C7, C9, and H3) and on the open coast (Bound Rock). #3 414 Rhodora [Vol. 74 Enteromorpha linza (L.) J. Agardh: Found sporadically in the estuary (C9 and H1) and the ex- posed open coast (Bound Rock and HB1) ; present on mud, rocks and as an epiphyte in the lower eulittoral and upper sublittoral zones. #3 Enteromorpha plumosa Kitzing: Found twice in the estuary (A18 and B12) on muddy sur- faces in the lower eulittoral zone. #3 Enteromorpha prolifera (O. F. Müller) J. Agardh: Infrequent in the estuary (B2, C2, C3, C5, C6 and C11); present on muddy surfaces of the eulittoral zone. Mixed with Enteromorpha erecta but never occurring as abun- dantly. #3 Monostroma fuscum (Postels et Ruprecht) Wittrock: Found sporadically throughout the estuary (A15, B2, B4, B6, B8, C1 and C3) on mud and rocks in the lower eulittoral and sublittoral zones. Abundant in localized areas on the exposed open coast (Bound Rock and HB4), and with the same vertical distribution. #3 Monostroma grevillei (Thuret) Wittrock : Infrequent on rocks in the mid and lower eulittoral zones of the estuary (A5, A15, B6 and B10). Abundant on the ex- posed open coast (Bound Rock) and with the same vertical distribution. #3 Monostroma leptodermum Kjellman: Found once adrift at A3. #3 *Monostroma oxyspermum (Kiitzing) Doty: Locally abundant throughout the estuary (A2 to A4, A9, B1, B4, B6, B8, Cl, C2, and C4 to C7) — particularly in areas of low salinity. It forms a distinct band on high ver- tical (muddy) banks and occasionally occurs as an epiphyte on Spartina alterniflora and other vascular plants. #2 Monostroma pulchrum Farlow: Common (during the summer) on rocks and on various al- gae in the lower eulittoral and sublittoral zones of the ex- posed open coast (Bound Rock). #3 *Within range but not previously recorded for New Hampshire. 1972] Marine Algae — Mathieson and Fralick 415 Percursaria percursa (C. Agardh) Rosenvinge: Locally abundant in the estuary (Al, A2, A16, B1, B3, B6, B7, and C3 to C5) as free-floating masses in tide pools and attached to muddy surfaces in the upper eulittoral zone. Often mixed with Rhizoclonium riparium and Cladophora sericea. #3 Pseudendoclonium marinum (Reinke) Aleem et Schulz: (equals Protoderma marinum Reinke in Taylor, 1957) Abundant on rocks from the mid eulittoral to the sublittoral zones on the exposed open coast (Bound Rock) ; found once in the estuary (All). £3 Rhizoclonium riparium (Roth) Harvey: Abundant throughout the estuary (A1 to A5, A7 to A9, A15, B1 to B4, B6, B8 to B10, B11 to B13, C1 to C6, C8, C10, C11, and H3) as free floating masses in tide pools and attached to muddy surfaces in the upper littoral zone. Often mixed with Cladophora sericea and Percursaria percursa. #3 Rhizoclonium tortuosum Kützing : Abundant during the summer on the exposed open coast (Bound Rock) ; found once in the estuary (C9). Present as entangled masses amongst various algae (particularly Chondrus erispus and Gigartina stellata) in the lower eu- littoral and sublittoral zones. £3 Spongomorpha arcta (Dillwyn) Kützing: Abundant on rocks (rarely as an epiphyte) in the mid- lower eulittoral zones of the exposed open coast (Bound Rock) ; most conspicuous in the late winter and spring. #3 Spongomorpha spinescens Kiitzing: Abundant on rocks in the mid to lower eulittoral zones of the exposed open coast (Bound Rock) ; most conspicuous in the spring and summer. #3 Ulothrix flacca (Dillwyn) Thuret in Le Jolis: Locally abundant (during winter and spring) on rocks in the upper littoral zone of the exposed open coast (Bound Rock). #3 416 Rhodora [Vol. 74 Ulva lactuca L: Ubiquitous throughout the estuary (being found at all sta- tions except A6, A11, A15, A16, C8, D2, D3, H2, and H4) on mud and any solid substrates in the lower eulittoral and sublittoral zones. Present on the exposed open coast (Bound Rock and HB2 and HB4) but not as abundant as in the estuary. £3 Urospora collabens (C. Agardh) Holmes et Batters: Locally abundant (particularly during the winter and spring) on rocks in the upper littoral zone at Bound Rock. #3 Urospora penicilliformis (Roth) Areschoug: The abundance and distribution (both seasonal and verti- cal) of this species is essentially similar to that of U. collabens — except that it tends to appear later than U. collabens. #3 Urospora speciosa (Carmichael ex Harvey in Hooker) Leblond et Hamel: Occasional (particularly during the winter and spring) on rocks in the upper littoral zone at Bound Rock. £1 Phaeophyceae: Agarum cribrosum (Mertens) Bory: Common on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB1, HB3, and HB4) ; not found in the estuary. #1 Alaria esculenta (L.) Greville: Locally abundant on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB2 and HB3); not found in the estuary. #1 Ascophyllum nodosum (L.) Le Jolis: Common throughout the estuary (A2, A4, A11, A13, A14, A17, A18, B1, B2, B9, B10, C1 to C4, C10 and C11 and H3) on any solid substrate from the upper sublittoral to the mid eulittoral zones. It is a rare plant on the exposed open coast at Bound Rock. #3 Ascophyllum nodosum (L.) Le Jolis f. scorpioides (Horne- mann) Reinke: 1972] Marine Algae — Mathieson and Fralick 417 Common on high muddy banks throughout the estuary (Al to A3, A5 to AT, A12 to A14, A17, B1 to B3, B5, B6, B8, B9, B13, C1 to C3, C5, C6, C11, D2 and D3) ; entangled amongst Spartina alterniflora and other vascular plants. #2 Chorda filum (L) Stackhouse: Found once on pier pilings in the estuary (H3); another time on rocks and shells on the exposed open coast (Bound Rock). In both cases the plants were present in the sub- littoral zone. #3 Chorda tomentosa Lyngbye: Locally abundant (during the summer) on scattered rocks in the sublittoral zone of the exposed open coast (Bound Rock and HB3). #3 Chordaria flagelliformis (O. F. Müller) C. Agardh: Found twice in the estuary (H1 and H3) ; in both cases it was growing on pier pilings in the lower eulittoral — sub- littoral zones. The plant is common (during the summer) on the exposed open coast (Bound Rock and HB2) and has the same vertical distribution as in the estuary. £3 Desmarestia aculeata (L.) Lamouroux: Common on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB2 to HB4). #1 Desmarestia viridis (O. F. Müller) Lamouroux: Occasional on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB1, HB3 and HB4). #1 Dictyosiphon foeniculaceus (Hudson) Greville: Found once on mud covered rocks in the lower eulittoral zone of the estuary (C5). #3 Ectocarpus confervoides (Roth) Le Jolis: (equals E. siliculosus (Dillwyn) Lyngbye in Parke and Dixon, 1968) Occasional on rocks and larger algae in the eulittoral and sublittoral zones of the estuary (A14, A17, B9, C9 to C11 and H1) and the exposed open coast (Bound Rock, HB1 to HB4). The estuarine plants were more robust in stature than the plants from the open coast. #3 418 Rhodora [Vol. 74 Ectocarpus siliculosus (Dillwyn) Lyngbye: Found once as an epiphyte on Spartina alterniflora in the upper littoral zone of the estuary (C5). #3 Elachista fucicola (Velley) Areschoug : An occasional epiphyte on Ascophyllum nodosum and Fucus vesiculosus var spiralis in the estuary (Al, B12, C1, C3, C5 and H3); very common on Fucus vesiculosus on the exposed open coast (Bound Rock and HB2). #3 Fucus distichus (L.) emend Powell subsp. distichus Powell: Locally abundant in high tide pools of the exposed open coast (Bound Rock). £1 Fucus distichus (L.) emend Powell subsp. edentatus (De la Pylaie) Powell: Common on rocks in the lower eulittoral — sublittoral zones of the exposed open coast (Bound Rock and HB2). Found once in a similar habitat near the mouth of the estuary (H1). £3 Fucus distichus (L.) emend. Powell subsp. evanescens (C. Agardh) Powell: Locally abundant on rocks in the lower eulittoral — sub- littoral zones of the exposed open coast (Bound Rock). #3 Fucus vesiculosus L: Abundant on semi-exposed rocks from the mid to the lower eulittoral at Bound Rock. #3 Fucus vesiculosus L. var spiralis Farlow: Ubiquitous throughout the estuary (at all stations except A12, A15, A16, C8, H1, H2 and H4) on mud, rocks, shells and any other solid substrates in the mid to the upper eulit- toral zone. It is one of the most conspicuous species on the upper banks of the salt marshes, where it is associated with Ascophyllum nodosum f. scorpioides, Spartina alterniflora and various other vascular plants. #2 Giffordia granulosa (J. E. Smith) Hamel: Found once in the estuary (A7) on mud-covered rocks in the lower eulittoral zone. #2 1972] Marine Algae — Mathieson and Fralick 419 Laminaria digitata (Hudson) Lamouroux: Present on rocky substrate in the sublittoral zone of the exposed open coast (Bound Rock, HB1, HB3 and HB4) and at the mouth of the estuary (H1). Each specimen had a consistent +-— anatomy (i.e. mucilage ducts are present in the blade and absent from the stipe, Wilce, 1965). #3 Laminaria saccharina (L.) Lamouroux sensu Wilce, 1965: Its distribution was essentially similar to that of L. digitata except that it was found at one other coastal (HB2) and estuarine location (A18). All of the specimens were the —— ecotype of Wilce, 1965 (ie. L. agardhii Kjellman in Taylor, 1957). £3 Leathesia difformis (L.) Areschoug: A common epiphyte (during the summer) on Chondrus crispus and other algae in the lower eulittoral-upper sub- littoral zones of the exposed open coast (Bound Rock and HB2). £1 Petalonia fascia (O. F. Müller) Kuntze: Common throughout the estuary (A4, A5, A7, A9, A14, A15, A17, B2, B4, B6, B10, B13, C2, C6, C11, D2 and D3) and on the exposed open coast (Bound Rock). It is present in the eulittoral zone on rocks (often in tide pools), mud, and occasionally as an epiphyte on large algae. #3 Pilaiella littoralis (L.) Kjellman: Common on rocks and as an epiphyte on Fucus vesiculosus in the mid-lower eulittoral zones of the exposed open coast (Bound Rock). Occasionally present in the estuary (Cl, C5 and C6) and with the same vertical distribution. #3 *Pseudolithoderma extensum (Crouan frat.) S. Lund: (equals Lithoderma extensum (Crouan) Hamel in Tay- lor, 1957) Occasional on sublittoral stones and boulders of the ex- posed open coast (HB2 and HB4). #1 Ralfsia borneti Kuckuck: Occasional on stones and boulders in the lower eulittoral — sublittoral zones of the estuary (A8, A10, B10, B11, and C1). According to Edelstein, Chen and McLachlan (1970) 420 Rhodora [Vol. 74 R. borneti is a stage in the life history of Petalonia fascia, and it is not a valid taxa. #3 Ralfsia clavata (Harvey in Hooker) Crouan frat.: Found once in the estuary (A-8) on mud covered rocks in the lower eulittoral zone. It is also described (Edelstein, Chen and McLachlan, 1970) as a stage in the life history of Petalonia fascia. #3 Ralfsia fungiformis (Gunner) Setchell et Gardner: Found once on rocks in the sublittoral zone of the exposed open coast (HB4). #1 Ralfsia verrucosa (Areschoug) J. Agardh: Found twice on rocks and shells in the upper sublittoral and eulittoral zones of the estuary (B3 and C1). Abundant in the same zones on the exposed open coast (Bound Rock). #3 Saccorhiza dermatodea (De la Pylaie) J. Agardh: Occasional on rocks in the sublittoral zone of the exposed open coast (HB1 and HB3). #1 Scytosiphon lomentaria (Lyngbye) Link: Present on rocks (often in tide pools), mussels shells, mud and occasionally epiphytic on various plants in the eulittoral zone of the estuary (A5, A14, A15, A17, B1, B2, B4, B10, C9, C11 and D3) and the exposed open coast (Bound Rock). #3 **Sorapion kjellmanii (Wille) Rosenvinge: Found once on rocks in the sublittoral zone of the exposed open coast (HB2). £1 **Sphacelaria plumosa Lyngbye: (equals Chaetopteris plumosa (Lyngbye) Kiitzing in Taylor, 1957) Occasional on sand covered rocks in the mid-lower sublit- toral zone of the exposed open coast (HB1). #1 Sphacelaria radicans (Dillwyn) C. Agardh: Occasional on muddy or sandy surfaces in the mid-lower eulittoral zone of the estuary (A9, B1 and B10); also **Range extension and a new record for New Hampshire. 1972] Marine Algae — Mathieson and Fralick 421 present on sand covered rocks in the sublittoral zone of the exposed open coast (HB1, HB3 and HB4). £3 Spongonema tomentosum (Hudson) Kützing: (equals Ectocarpus tomentosus (Hudson) Lyngbye in Taylor, 1957) Found once as an epiphyte on Laminaria saccharina in the sublittoral zone of the exposed open coast (HB3). #1 Rhodophyceae : Ahnfeltia plicata (Hudson) Fries: Locally abundant on sand-covered rocks and boulders in the lower eulittoral — sublittoral zones of the exposed open coast (Bound Rock, HB1, HB2 and HB4). #3 Antithamnion floccosum (O. F. Müller) Kleen: Found once on sand covered rocks in the sublittoral zone of the exposed open coast (HB3). #3 Audouinella membranaceae (Magnus) Papenfuss: Epiphytic on species of Sertularia which in turn may be epiphytic (commonly on fucoid algae) or saxicolous in the eulittoral zone of the exposed open coast (Bound Rock). #3 * Asterocytis ramosa (Thwaites in Harvey) Gobi ex Schmitz: Found once as an epiphyte on Cladophora sericea in a high marshy tide pool (C4). Growing in association with Per- cursaria percursa and various blue green algae. #2 Bangia ciliaris Carmichael: Found once as an epiphyte on Cladophora sericea in a high marshy tide pool (C5) in the estuary. #3 Bangia fuscopurpurea (Dillwyn) Lyngbye: Abundant (particularly during the winter and spring) on rocks in the upper littoral zone of the exposed open coast (Bound Rock). #3 Callithamnion baileyi Harvey: Found once in the estuary (H3) on a styrofoam float. #3 Callithamnion corymbosum (J. E. Smith) C. Agardh: Found once in the estuary (A17) on rocks in the upper sublittoral zone. #2 422 Rhodora [Vol. 74 Ceramium rubrum (Hudson) C. Agardh: Present on rocks and epiphytic on large macroscopic algae in the lower eulittoral and sublittoral zones. Occasional within the estuary (B1 to B3, B7, C2, C5, and H3), but more abundant on the exposed open coast (Bound Rock, HB1 to HB4). #3 Ceramium strictum Harvey: Abundant throughout the estuary (A1 to A4, AT, A8, A10, A12, A17, B1 to B6, B9, C1, C3 to C7, C11 and H1) on muddy surfaces in the lower eulittoral — sublittoral zones. Uncommon on the exposed open coast (Bound Rock). #3 Chondrus crispus Stackhouse: Common throughout the estuary (Al to A3, A5, A9 to A15, A17, A18, B2, B4 to B6, B8 to B11, Cl, C9, C11, D3 and H1) and the exposed open coast (Bound Rock, HB2 to HB4); present on any solid substrate in the lower eulittoral and sublittoral zones. The stature of the estuarine plants is much larger than the plants from the open coast. #3 Clathromorphum circumscriptum (Strömfelt) Foslie: (equals Phymatolithon compactum (Kjellman) Foslie in Taylor, 1957) Abundant on rocks and shells in the lower eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1 to HB4). Found once in the estuary (A10) on a sub- tidal population of mussels. £3 Corallina officinalis L.: Locally abundant on rocks and boulders (often in tide pools) in the eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1 to HB4). £1 Cystoclonium purpureum (Hudson) Batters var. cirrhosum Harvey: Present on rocks and as an epiphyte on larger algae in the sublittoral zone of the exposed open coast (Bound Rock, HB1 to HB4). #3 Dermatolithon pustulatum (Lamouroux) Foslie: 1972] Marine Algae — Mathieson and Fralick 423 (equals Lithophyllum pustulatum (Lamouroux) Foslie in Taylor, 1957) An occasional epiphyte on various algae (particularly Chondrus crispus and Gigartina stellata) in the lower eulittoral-sublittoral zones of the exposed open coast (Bound Rock, HB2 and HB4). #3 Dumontia incrassata (O. F. Müller) Lamouroux: Abundant on rocks in the mid and lower eulittoral zones of the exposed coast (Bound Rock). Occasional throughout the estuary (A5, A14, A15, A17, A18, B6, B10, and D3) and with the same vertical distribution as on the open coast. #3 Euthora cristata (C. Agardh) J. Agardh: Present on rocks and occasionally epiphytic on various plants (e.g. Phyllophora spp.) in the sublittoral zone of the exposed open coast (Bound Rock, HB1, HB3 and HB4). #1 Gigartina stellata (Stackhouse) Batters: On rocks in the lower eulittoral and sublittoral fringe zones of the exposed open coast (Bound Rock). #3 Gloiosiphonia capillaris (Hudson) Carmichael ex Berkeley : Found twice on rocks in the sublittoral zone of the ex- posed open coast (HB1 and HB4). £3 Hildenbrandia prototypus Nardo: Common on rocks in the eulittoral and sublittoral zones of the exposed open coast (Bound Rock and HB2). Less com- mon in the estuary (A8, A10, A11 and B10) but with the same vertical distribution. #3 Kylinia secundata (Lyngbye) Papenfuss: A common epiphyte on various algae in the eulittoral zone of the exposed open coast (Bound Rock). #3 Lithophyllum corallinae (€rouan frat.) Heydrich: Found once in the sublittoral zone of the exposed open coast (HB4) ; a specific epiphyte on Corallina officinalis. #1 Lithothamnium glaciale Kjellman: Common on shells and rocks (often in tide pools) in the 424 Rhodora [Vol. 74 lower eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1 to HB4). #1 Melobesia lejolisti Rosanoff : (equals Fosliella lejolisii (Rosanoff) Howe in Taylor, 1957) An occasional epiphyte on Phyllophora spp. in the sub- littoral zone of the exposed open coast (Bound Rock and HB3). #3 Membranoptera alata (Hudson) Stackhouse: A common epiphyte on various algae (occasionally on rock) in the sublittoral zone of the exposed open coast (Bound Rock, HB1, HB3 and HB4). #1 Nemalion helminthoides (Velley in Withering) Batters: Uncommon on rocks in the lower eulittoral zone of the exposed open coast (Bound Rock). #1 Petrocelis middendorfii (Ruprecht) Kjellman: Present on rocks and shells in the sublittoral zone of the exposed open coast (HB2 and HB4). #3 Peyssonelia rosenvingii (Schmity in Rosenvinge) : Occasional on rocks in the sublittoral zone of the exposed open coast (HB1 and HB4). #1 Phycodrys rubens (L.) Batters: Common on rocks and as an epiphyte on various algae in the lowest eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1, HB3 and HB4) and at the mouth of the estuary (H1). #3 Phyllophora brodiaei (Turner) Endlich: Common on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB1 to HB4). #1 Phyllophora membranifolia (Goodenough et Woodward) J. Agardh: The vertical and horizontal distribution of P. membrani- folia is essentially similar to that of P. brodiaei. It was found at stations HB2, HB3 and Bound Rock. #3 *Phymatolithon laevigatum (Foslie) Foslie: Present on rocks and shells in the sublittoral zone of the exposed open coast (HB2 to HB4). #1 1972] Marine Algae — Mathieson and Fralick 425 Phymatolithon lenormandi (Areschoug) Adey: Present on rocks in the sublittoral zone of the exposed open coast (HB4). #3 Plumaria elegans (Bonnemaison) Schmitz: Common on vertical rock faces under overhanging fucoids in the lower eulittoral zone of the exposed open coast (Bound Rock). It was found once in the estuary (A14) with the same vertical distribution. #3 Polyides rotundus (Hudson) Greville: Occasional on sand covered rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB2 to HB4). #3 Polysiphonia denudata (Dillwyn) Greville ex Harvey in Hooker: Found twice in the estuary (B6 and B10) on muddy sur- faces in the sublittoral zone. #3 Polysiphonia elongata (Hudson) Sprengel: Found twice in the sublittoral zone of the estuary (A7 and H3). #2 Polysiphonia fibrillosa (Dillwyn) Sprengel : Common throughout the estuary (Al, A2, A4, A10, B2 to B6, C2, C4 to C6, C9 and H3) on muddy surfaces in the lower eulittoral and sublittoral zones. #2 Polysiphonia lanosa (L.) Tandy: Hemiparasitic on Ascophyllum nodosum on the open coast (Bound Rock) and at the mouth of the estuary (A2 and A4). £3 Polysiphonia nigra (Hudson) Batters: Present on rocks in the lower eulitoral-sublittoral zones of the estuary (A15, B2, B3, B6, B7 and H3) and the exposed open coast (Bound Rock). #3 Polysiphonia nigrescens (Hudson) Greville: On rocks and shells in the lower eulittoral and sublittoral zones of the estuary (A4, AT, All, A12, A15, B2, BA, B5, B10, C1, C2, C5, H1 and H3) and the exposed open coast (HB1 and HB4). £3 426 Rhodora [Vol. 74 Polysiphonia novae-angliae Taylor: Present on rocks in the sublittoral zone of the exposed open coast (Bound Rock, HB1 and HB2). #3 Polysiphonia subtilissima Montagne: Found once in the estuary (A12) ; growing on mud in the lower eulittoral zone. #1 Polysiphonia urceolata (Lightfoot ex Dillwyn) Greville: Common on rocks in the lower eulittoral and sublittoral zones of the exposed open coast (Bound Rock, HB1 to HB4). Found once in the estuary at (B6) with the same vertical distribution. #3 Porphyra miniata (C. Agardh) C. Agardh: Common (particularly during the summer) on rocks and epiphytic on various plants in the upper sublittoral zone of the exposed open coast (HB1 to HB4). #3 Porphyra umbilicalis (L.) J. Agardh: Common on rocks, mud and on various algae in the eulit- toral zone of the estuary (Al, A3 to A5, A8, A12, A14, A17, C3, C5, C6, C9, H1 and H3) and the exposed open coast (Bound Rock). #3 The forma epiphytica Collins was only found at Bound Rock. #3 Ptilota serrata Kiitzing: Present on rocks and epiphytic on various algae in the sublittoral zone of the exposed open coast (Bound Rock, HB1 and HB3). #1 Rhodochorton purpureum (Lightfoot) Rosenvinge: Present on vertical rock faces under overhanging fucoids in the midlower eulittoral zone of the exposed open coast (Bound Rock). #3 *Rhodophysema elegans (Crouan frat. ex J. Agardh) Dixon: (equals Rhododermis elegans Crouan in Taylor, 1957) Occasional on rocks and shells in the sublittoral zone of the exposed open coast (HB2 and HB4). #1 Rhodomela confervoides (Hudson) Silva: Locally abundant on sand-covered rocks in the lower eulit- 1972] Marine Algae — Mathieson and Fralick 427 toral and sublittoral zones of the exposed open coast (Bound Rock, HB1 and HB3). £1 Rhodophyllis dichotoma (Lepeschkin) Gobi: Found once adrift at Bound Rock. #1 Rhodymenia palmata (L.) Greville: Relatively common on rocks and various algae (particularly Laminaria spp.) in the sublittoral zone of the exposed open coast (Bound Rock, HB1 to HB4) and the mouth of the estuary (H1). #3 DISCUSSION Eight of the taxa of marine algae recorded in the present study represent new records for the state of New Hamp- shire (Monostroma oxyspermum, Asterocytis ramosa, Phymatolithon laevigatum, Polysiphonia fibrillosa, Rhodo- physema elegans, Pseudolithoderma extensum, Sorapion kjellmanii and Sphacelaria plumosa). Sorapion kjellmanii and Sphacelaria plumosa also represent extensions of known distributional range on the northeast coast of North America, for they were both previously recorded from Ellesmere Island to Nova Scotia (as summarized in Taylor, 1957 and Cardinal, 1968). It should be noted that both species were collected in the mid-lower sublittoral zones of the open coast. The primary factors restricting the growth and distri- bution of seaweeds in the Hampton-Seabrook Estuary are a lack of stable substrate and adverse hydrographic condi- tions. As a consequence gross fluctuations of species num- bers occurred at different locations, and a gradual reduction in species diversity and biomass was evident from the open coast to the head of the estuary. Green algae showed the widest distribution throughout the estuary, for they grew on a variety of substrates and in a wide range of tempera- tures and salinities. A greater diversity and biomass of seaweeds were re- corded in the Great Bay Estuary complex (Mathieson, Reynolds and Hehre, in press) than in the Hampton- 428 Rhodora [Vol. 74 Seabrook Estuary. Thus, 98 taxa of algae were found in the former area, while only 57 were found in the latter. The variety of species found in the estuarine environments of Great Bay is primarily due to its large size (over 15,000 acres of tidewater) and its greater variety of habitats (particularly tidal rapids) and substrates. Peak popula- tions of seaweeds were found at Dover Point and Adams Point in the Great Bay System (Mathieson, Reynolds and Hehre, in press). Both of the sites are exposed to strong tidal currents and their floras are “open coastal" in nature. More than half of the 76 cosmopolitan species found in the Great Bay area were either restricted to the two tidal rapid sites or they did not extend inland beyond Dover Point or Adams Point. Only one major tidal rapid was observed in the Hampton-Seabrook area (station C-9), and it was so shaded that few algae grew there. The dis- crepancy in species numbers between the two estuaries is primarily accounted for by the cosmopolitan component of Great Bay. All of the species found in the Hampton-Seabrook Estuary, except Monostroma oxyspermum, Chorda filum, Dictyosiphon foeniculaceus, Elachista fucicola, Ralfsia borneti, R. clavata, Sphacelaria radicans, Asterocytis ramosa, Plumaria elegans and Polysiphonia fibrillosa, were also found in the Great Bay area. The occurrence of P. elegans and S. radicans in estuarine waters was particularly striking, for both of the species are most abundant on the open coast. ACKNOWLEDGEMENTS The author wishes to thank the following individuals for their help: Mr. John Shipman, Dr. Norman Reynolds and Dr. Edward Hehre for their help during the collection and processing of specimens; Dr. Donald Normandeau, of Nor- mandeau Associates, who provided a boat, outboard motor and other support to complete the investigation. 1972] A-12 A-13 A-14 A-15 Marine Algae — Mathieson and Fralick 429 Table I Stations At the mouth of the first major tributary SE of Tide Mill Creek on the Hampton River, Hampton Falls, Hampton, N. H.* (1.3 miles inland) The northeast portion of an “island” formed at the mouth of the Taylor River, Blind Creek and the upper part of the Hampton River, Hampton Falls, Hampton, N. H. (1.91 miles inland) Approximately 1500 feet NE of the mouth of Tide Mill Creek and the Hampton River, Hampton, N. H. (1.3 miles inland) Approximately 1800 feet NW of Station A-2 on the Hampton River, Hampton-Hampton Falls, N. H. (2.3 miles inland) Opposite a small brook, which empties into the first tributary above the mouth of the Hampton Falls River, Hampton, N. H. (3.3 miles inland) At the junction of the first oxbow NE of station A-5 on the Taylor River, Hampton, N. H. (3.2 miles inland) Hampton Landing on the Taylor River, Hampton, N. H. (3.1 miles inland) At the mouth of Nudds Canal and Blind Creek, Hampton, N. H. (2.31 miles inland) Tide Mill Creek, by the route 101 bridge, Hampton, N. H. (2.56 miles inland) Hampton Falls River south of Depot Avenue and near the Boston and Maine Railroad bridge, Hampton Falls, N. H. (5.2 miles inland) Approximately 2000 feet SW of the Boston and Maine “sub- station” which is between Lafayette and Landing Roads. Station A-11 is adjacent to the B & M railroad tracks, and on the Taylor River, Hampton, N. H. (4.4 miles inland) End of Depot Avenue on the Hampton Falls River, Hampton Falls, N. H. (5.0 miles inland) Middle of the southernmost oxbow near the mouth of the Taylor River and the Hampton Town Line, Hampton, N. H. (3.0 miles inland) A bend in the first tributary above (north) the Hampton Falls River where the river crosses the railroad tracks, Hampton, N. H. (3.7 miles inland) Same as A-14, except the land side of the railroad tracks, rather than the harbor side as A-13, Hampton, N. H. (3.7 miles inland) * Distance inland from exposed open coast Rhodora [Vol. 74 A salt marsh on a point of land made by the Hampton River and the first tributary above the Hampton Falls River, Hampton, N. H. (3.5 miles inland) Hampton Landing, Taylor River, Hampton, N. H. (3.01 miles inland) The Willows; at the mouth of Tide Mill Creek and the Hamp- ton River, Hampton, N. H. (1.6 miles inland) On a small “island” opposite Eastman’s Slough and about 2500 feet west of Locke Point State Park area, Hampton Falls, Seabrook, N. H. (1.0 miles inland) The southernmost portion of Eastman Slough, near Halftide Rock and at the mouth of the Brown River, Hampton Falls- Seabrook, N. H. (1.7 miles inland) Just inside the mouth of Hunts Island Creek and at the junction of the Brown River, Seabrook, N. H. (1.8 miles inland) Approximately 500 feet NW of the first tributary past Hunts Island Creek, Hampton Falls-Seabrook, N. H. (2.0 miles in- land) Approximately 700 feet NE of the mouth of Swain's Creek, Hampton Falls, N. H. (2.2 miles inland) Robbins Point, Hampton Flats, Hampton Falls-Seabrook, N. H. (2.3 miles inland) Browns River, first tributary upstream from Swain's Creek, Hampton Falls, N. H. (2.5 miles inland) Approximately 800 feet upstream from Robbins Point, Hamp- ton Falls, N. H. (2.4 miles inland) Approximately 700 feet upstream from station B-8, Hampton Falls-Seabrook, N. H. (2.53 miles inland) End of Rocks Road, on the Browns River, Hampton Falls- Seabrook, N. H. (2.8 miles inland) Near the mouth of the first major tributary east of the head waters of the Browns River, Hampton Falls-Seabrook, N. H. (3.0 miles inland) Approximately 1500 feet upstream (west) from station B-11, just before a major oxbow, Hampton Falls-Seabrook, N. H. (3.3 miles inland) Swain’s Creek, neck of the first oxbow, Hampton Flats, Hampton Falls, N. H. (2.6 miles inland) At the mouth of the Blackwater River; near the first tributary SW of Mills Point, Seabrook, N. H. (1.5 miles inland) Approximately 1200 feet SW of the first tributary past River- side, Seabrook, N. H. (2.2 miles inland) Approximately 1200 feet south of C-2, Seabrook, N. H. (2.3 miles inland) 1972] Marine Algae — Mathieson and Fralick 431 C- 4 Approximately 500 feet SE of C-3, Seabrook, N. H. (2.4 miles inland) C- 5 Approximately 800 feet SE of C-4, Seabrook, N. H. (2.5 miles inland) C- 6 Approximately 800 feet SE of C-5 and near the first tributary southeast of C-2, Seabrook, N. H. (2.7 miles inland) C- 7 Approximately 1200 feet SE of C-6 and near a large white rock, Seabrook, N. H. (3.0 miles inland) C-8 Approximately 1500 feet SE of C-7, Seabrook, N. H. (3.3 miles inland) C-9 By the route 286 bridge which crosses the Blackwater River, Seabrook, N. H. (3.6 miles inland) C-10 Mills Point — mouth of the Blackwater River, Seabrook, N. H. (1.3 miles inland) C-11 Riverside, Seabrook, N. H. (1.7 miles inland) D-1 At the mouth of the creek between Knowles Island and main- land, Seabrook, N. H. (1.34 miles inland) D- 2 Before the first bend near the mouth of Mill Creek, Seabrook, N. H. (1.6 miles inland) l D- 3 Walton Landing, end of Walton Road, Seabrook, N. H. (2.1 miles inland) H- 1 At the junction of the middle piling of the toll bridge in Hampton Harbor and the tower at Hampton, N. H. (0.4 miles inland) H- 2 In the channel of Hampton Harbor near the mouth of the Blackwater River and at the junction of the imaginary line between the Seabrook Marina and Knowles Island, Seabrook, N. H. (1.0 miles inland) H- 3 Smith and Gilmore Marina, Hampton, N. H. (0.8 miles inland) H- 4 Hampton Marina — mouth of Hampton River, Hampton, N. H. (0.97 miles inland) HB-Bound Rock The area in the immediate vicinity of Beckman's Point, which is near the mouth of Hampton Harbor, Hampton, N. H. Ex- posed open coast. HB-1 A series of offshore rocks in the immediate proximity of the red marker flag, and in 35-40 feet of water, Hampton, N. H. Exposed open coast HB-2 The southernmost rock outcrop (a rookery) before the Hamp- ton Breakwater; in 15-20 feet of water and the sheltered side, Hampton, N. H. Exposed open coast. HB-3 The exposed (seaward) side of the same rock outcrop as HB-2; 30-40 feet of water, Hampton, N. H. Exposed open coast. HB-4 The southernmost outcrop which is opposite the Hampton State Beach Park Breakwater; in 20-25 feet of water, Hamp- ton, N. H. Exposed open coast. 432 Station Al A2 A3 A4 A5 A6 AT A8 A9 A10 A11 A12 A13 A14 A15 A16 A17 A18 B1 B2 B3 B4 B5 B6 Temperature (C) 16 - 20 19 - 19.5 16 - 20.5 19 - 20.8 18 - 21.5 20.5 20.5 - 29.5 15.0 - 19.8 (-1.0) - 22.5 24.0 21.5 9.0 - 10.0 8.0 7.0 8.0 9.0 0.0 0.0 16.0 17.0 - 18.0 17.0 - 19.0 16.5 16.0 - 16.5 2.0 - 17.0 18.0 18.0 16.0 - 16.5 16.5 - 20.0 16.0 - 19.0 16.0 - 19.0 Rhodora Table II Surface Water Temperatures and Salinities Salinities (0/00) 31 - 33.0 31 - 31.5 31 - 32.0 31 - 32.0 31 - 31.3 32.0 30.5 - 32.0 20.2 - 30.0 33.5 31.3 28.6 - 30.4 27.4 - 27.6 24.0 23.9 30.6 24.7 - 24.9 30.6 - 30.8 26.3 22.8 - 30.9 30.5 - 30.9 30.5 24.4 - 30.3 20.7 - 29.9 28.9 28.9 17.2 - 18.2 16.2 - 31.6 14.2 - 30.9 13.2 - 30.9 28.9 29.4 28.5 25.6 26.0 25.2 25.2 25.6 [Vol. 74 Dates* 12-21-69 10-04-66 11-18-66 11-18-66 11-18-66 11-18-66 11-13-66 11-13-66 12-06-69 12-06-66 1972] Marine Algae — Mathieson and Fralick 433 C8 15.5 24.5 C9 13.2 - 16.2 11.5 - 14.5 10-09-69 C10 10.0 27.2 10-13-66 C11 11.0 26.8 10-22-66 D2 13.0 23.6 11-11-66 D3 13.0 20.2 11-11-66 H1 16.5 30.5 H2 16.8 90.5 H4 1.0 30.5 12-21-66 Bound Rock 0.0 - 19.0 29.5 - 32.0 7-18-66 * All dates from August and September, 1969 except those specified. Table III. Number of Seaweed Taxa of Various Stations Stations Total Chlorophyceae Phaeophyceae Rhodophyceae Al 19 5 3 4 A2 14 1 3 4 A3 11 6 2 3 A4 13 5 3 5 A5 12 5 4 3 A6 5 3 2 -— AT 12 5 4 3 A8 10 4 3 3 A9 11 T 3 1 A10 11 3 3 5 A11 7 2 2 3 A12 7 1 1 5 A13 8 3 3 2 A14 12 2 6 4 A15 8 4 = 4 A16 2 2 Es = A17 15 4 6 5 A18 8 3 3 2 B1 15 8 5 2 B2 19 8 5 6 B3 13 6 3 4 B4 15 8 3 4 B5 E 5 2 4 B6 19 9 3 1 B7 10 6 2 2 B8 7 4 1 2 B9 11 5 4 2 B10 16 5 6 5 434 Rhodora [Vol. 74 Bll 5 2 2 1 B12 T 5 2 — B13 6 3 3 0 C1 17 7 7 3 C2 13 6 4 3 C3 14 8 4 2 C4 11 6 2 3 C5 19 8 5 6 C6 15 7 5 3 C7 6 4 1 1 C8 1 1 — — C9 10 5 2 3 C10 5 2 3 — C11 11 3 6 2 D1 — — — — D2 4 — 4 — D3 6 — 4 2 H1 14 3 5 6 H2 1 — — 1 H3 18 6 5 7 H4 0 - — — HB-Bound Rock 79 23 21 35 HB1 30 3 8 19 HB2 30 1 10 19 HB3 30 0 10 20 HB4 38 4 9 25 REFERENCES CARDINAL, A. 1968. Répertoire des Algues Marines Benthiques de L’est du Canada. Cahiers d’Information #48, Sta., Biol. Mar., Grand Riviere, mimeographed, 213 pp. CHAPMAN, V. J. 1940. Studies in salt marsh ecology. VI and VII. J. Ecol. 28: 118-152. Davis, R. B. 1956. An ecological study of a tidal salt marsh and estuary. M. S. thesis, University of New Hampshire, 102 pp. DRUM, R. and E. E. WEBBER. 1965. Diatoms from a Massachusetts salt marsh. Bot. Mar. 9: 70-77. EDELSTEIN, T. L., C. M. CHEN and J. MCLACHLAN, 1970. The life cycle of Ralfsia clavata and Ralfsia borneti. Can. J. Bot. 48: 527-531. 1972] Marine Algae — Mathieson and Fralick 435 HEHRE, E. and A. C. MATHIESON. 1970. Investigations of New Eng- land marine algae III: Composition, seasonal occurrence, distri- bution and reproductive periodicity of the marine Rhodophyceae in New Hampshire. Rhodora 72: 194-239. HOoEK, C. VAN DEN. 1963. Revision of the European species of Cladophora. xi 4- 248 pp, Leiden. LEWIS, J. R. 1964. The ecology of rock shores. xii + 323 pp. Eng- lish University Press Ltd., London. MATHIESON, A. C. E. HEHRE and N. REYNOLDS. Investigations of New England marine algae I. A floristic and descriptive eco- logical study of the marine algae at Jaffrey Point, New Hamp- shire. Nova Hedwigia (in press). MATHIESON, A. C., N. REYNOLDS and E. HEHRE. ibid II. The species composition, distribution and zonation of seaweeds in the Great Bay Estuary System and the adjacent open coast of New Hamp- shire. Nova Hedwigia (in press). MILLER, W. R. and F. E. EGLER. 1950. Vegetation of the Wequete- quock-Pawcatuck tidal marshes, Connecticut. Ecol. Mon. 20: 143-172. PARKE, M. and P. S. Drxon. 1968. Check-list of British marine algae — second revision. J. mar. biol. Ass. U. K. 48: 783-832. ScAGEL, R. F. 1966. Marine algae of British Columbia and northern Washington, Part I: Chlorophyceae (green algae). Nat. Mus. Canada, Biol. Ser. No. 74, Bull. No. 207; viii + 257. TAYLOR, W. R. 1957. Marine algae of the northeastern coast of North America. viii + 509 pp. Univ. Michigan Press, Ann Arbor. VAGENAS, G. 1969. An ecological study of the Hampton-Seabrook, New Hampshire salt marshes. M. S. Thesis, University of New Hampshire, 74 pp. WEBBER, E. E. 1967. Blue-green algae from a Massachusetts salt marsh. Bull. Torrey Bot. Club. 94: 99-106. 1968. Seasonal occurrence and ecology of salt marsh phanerogams at Ipswich, Massachusetts, Rhodora 70: 442-450. WEBBER, E. E. and R. T. WILCE. 1971. Benthic salt marsh algae at Ipswich, Massachusetts. Rhodora 73: 262-291. WILCE, R. T. 1965. Studies in the genus Laminaria III. A revision of the North Atlantic species of the Simplices Sections of Lami- naria. Botanica Gothoburgensia III, pp. 247-256. DEPARTMENT OF BOTANY AND JACKSON ESTUARINE LABORATORY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824 EDGEWORTHIA (THYMELAEACEAE) NEW TO THE WESTERN HEMISPHERE WILBUR H. DUNCAN AND MARIE MELLINGER Edgeworthia chrysantha Lindl, a shrub previously known to be native only from eastern Asia, has been found growing wild in Rabun County in northeastern Georgia in the Blue Ridge Mountains. Plants were first observed 15 September 1971 by the junior author. They are now known to occur at intervals along 2.2 miles of Wolf Creek in de- ciduous woods at elevations of about 1200-1600 feet. The woods have had selective cutting but have never been cleared. Many colonies of the Edgeworthia occur along Wolf Creek. One consists of at least 500 plants and extends along the banks of the stream for about 1/4 mile. The largest plant is 3.5 m tall and the stem 4 cm in diameter 15 cm above the ground. Plants vary in size, those on the stream bank being the most robust with smaller plants extending 6 m from the stream bank. Plants 75 cm or more tall usually bear inflorescences. Open flowers were found on some plants on 26 February 1972. These flowers and vegetative material collected be- fore frost enabled us to identify the plants positively to species. Specimens with full sized but immature fruits were collected 21 May 1972. All characteristics of the plants compare well with those of 23 specimens which we have seen from Japan. Edgeworthia chrysantha was intro- duced into Japan for cultivation as a source of paper and is now widely escaped there (Ohwi, 1965). Is Edgeworthia native to the Southeastern States? We have found no reports to substantiate or repudiate such a theory. It is especially significant that neither Small (1933) nor Nevling (1962) reports the genus for this area. Ap- parently it has been reported as native only from China to Himalaya (Hooker, 1885; Nakai, 1924; Hamaya, 1955; Makino, 1957; Hutchinson, 1967). Fairchild (1903) re- ports the introduction of the species into the United States from Japan and later (1908) says that planting trials at Washington, D.C., and elsewhere indicate it will prove 436 1972] Edgeworthia — Duncan and Mellinger 437 hardy as far north as the Carolinas and will seed freely. Bailey and Bailey (1941) state that E. chrysantha “— is introduced in N. Amer.” The Rabun County plants may be from one of these introductions, however, we have no data to indicate the species was introduced there or that it has escaped cultivation from elsewhere. Yet, it is incred- ible to think that the plants are an eastern Asiatic dis- junct. A few vascular plant species occur in both the Southeast- ern United States and China and/or Japan, e.g. Mitchella repens L. All are more widespread in both hemispheres than is the Edgeworthia in Rabun County. In addition, there are many genera in the Southeastern United States that have different species occurring in Asia, but no pair in any genus has the strong similarities as do our Edge- worthia plants and the 23 specimens we have seen from Japan. We are continuing our search for information which might indicate the Rabun County plants are introduced or escaped. If such data are lacking, then, because of the cir- cumstances under which the plants are growing, we think the colony could be native. We solicit any information that might be of help. Obviously descriptions of E. chrysantha are not readily available to most persons in the United States. Therefore, a short description is presented below along with a partial synonomy. Edgeworthia chrysantha Lindl. (E. papyrifera (Sieb.) Sieb. et Zucc., E. tomentosa (Thunb.) Nakai, E. gardneri of authors, not Meissner) Deciduous shrub to 3.5 m tall, usually ternately branched. Twigs yellow-brown, appressed-pubescent when young. Leaf scars raised about 45° at the lower end, with one crescent- shaped bundle scar. Leaves petioled, the blades lanceolate to sometimes oblong; acute to shortly acuminate or rarely obtuse at the apex; cuneate to acuminate at the base; 8-25 em long; thinly silky, especially below; the hairs .25-.75 mm long, appressed, and pointing toward the tip. Inflores- cences are in axillary peduncled heads which develop be- fore the leaves fall and are conspicuous during the winter. 438 Rhodora [Vol. 74 Flowers fragrant, 20-50 in each head. Heads subtended by several densely silky ovate to oblong bracts which usually fall before the flowers open. The heads droop early, at least by the time that all flowers have opened. Flowers silky hairy on outside, apetalous, 10-18 mm long, the calyx lobes 2-3 mm long and yellow inside. Stamens 8 in two series of 4 on inside of floral tube, filaments very short. Ovulary nearly sessile, hairy, 1-locular. Styles 3-4 mm long. Stigma clavate. The entire pistil 7-9 mm long. The flowers open as early as late February. The fruits mature in summer. They are hairy at the apex, 6-7 mm long, el- lipsoid to ovate, dry, green, and often surrounded by the somewhat persistent floral tube. Voucher specimens are in the University of Georgia Herbarium: Mellinger, 15 Sep- tember 1971; Duncan 23751, 21 May 1972 (fruiting); in deciduous woods along Wolf Creek, Rabun Co., Georgia, Blue Ridge Province. Edgeworthia as presently interpreted apparently consists of four species. They have been separated mostly on the basis of vestures or the lack of any, flower colors, and pe- duncle lengths. E. longipes Lace, which occurs in upper Burma, has peduncles 9-11 cm long. In other species they are only 1-2.5 em long. E. albiflora Nakai, which occurs in Yunan and Szechwan Province, China, may be recognized by the whitish interior of the calyx and glabrous under- sides of the leaves. In the remaining two species, E. gard- neri Meisner and E. chrysantha, the leaves are hairy beneath (sometimes faintly so) and the flowers are yellow inside. The flowers appear with the leaves in the former and before the leaves in the latter. A much more reliable character apparently has not been used previously; the in- volucral bracts are ovate to oblong in E. chrysantha and linear (up to 35 mm long) in E. gardneri. There seems to be a short-bracted and a long-bracted form of E. gardneri. These forms may deserve taxonomic recognition and should be studied. The long-bracted form occurs from Nepal into northern Burma, and the short-bracted form from eastern Nepal into southwestern China. Edgeworthia chrysantha occurs in the southern half of China and as an adventive in 1972] Edgeworthia — Duncan and Mellinger 439 Japan. It is considered by Nakai (1929) as two species which are separated by stoutness of twigs and size of leaf scars. We, however, agree with Hamaya (1955) that there is a gradual transition in sizes and there is no basis for division into two species. We acknowledge with appreciation the loan of specimens from several herbaria, and information provided by Drs. Lorin I. Nevling, Richard A. Howard, and Frederick G. Meyer. LITERATURE CITED BAILEY, L. H. and E. Z. BAILEY. 1941. Hortus Second. Macmillan Co. N.Y. FAIRCHILD, Davip. 1903. Mitsumata, a Japanese Paper Plant. U.S.D.A., Bureau of Plant Industry, Bull. No. 42: 9-11. 1908. Cultural Directions for the Mitsumata, a Japanese Paper Plant, U.S.D.A. Bureau of Plant Industry. Misc. Leaflet. 2 p. HAMAYA, ToSHIO. 1955. A Dendrological Monograph of the Thymelaeaceae Plants of Japan. Bull. of the Tokyo University Forests. No. 50: 246-248. HOOKER, Sir J. D. 1885. Flora of the British India, v. 5. Cheno- podiaceae to Orchidaceae. L. Reeve & Co., Ltd., Ashford, Kent, England. HUTCHINSON, JOHN. 1967. The Genera of Flowering Plants. Ox- ford at the Clarendon Press. 659 p. MAKINO, ToMiLARO. 1957. An Illustrated Flora of Japan with Cultivated and Naturalized Plants (Enlarged Ed.) The Hoku- ryukan C., Ltd., Tokyo. 1304 p. & indices. NAKAI, T. 1924. Some New and Noteworthy Ligneous Plants of Eastern Asia. Journal of the Arnold Arboretum 5: 81-83. NEVLING, LoRIN L, JR. 1962. The Thymelaeaceae of the Southeast- ern United States. Journal of the Arnold Arboretum 43: 428- 434. OHWI, JISABURO. 1965. Flora of Japan. Smithsonian Institution, Washington, D.C. 1067 p. SMALL, J. K. 1933. Manual of the Southeastern Flora. Published by the author. N.Y. 1554 p. DEPARTMENT OF BOTANY UNIVERSITY OF GEORGIA ATHENS, GEORGIA 30601 ROUTE 1, TIGER, GEORGIA 30576 REQUEST FOR UNNEEDED COPIES OF RHODORA FOR DECEMBER 1971, NO. 796 The error on the cover of the fourth number of Rhodora for 1971, indicating it as September 1971, no. 795, has resulted in many persons discarding it as a duplicate. Had they looked inside the cover they would have detected the error immediately. However so many requests have come in for replacements that Dr. Herman Sweet requests that any subscribers who have this December issue, and do not regularly keep a file of Rhodora, send this particular copy to him at the Botanical Museum, Oxford Street, Cambridge, Mass. 02138. 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, particularly 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- atics 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 Pot. Club 85: 277-300. 1958. Volume 74, No. 799, including pages 301-440, was issued September 29, 1972. 440 CONTENTS: continued Variation within Calochortus venustulus (Liliaceae). James L. Reveal and William J. Hess ..................... eee 350 New Form of Thuja occidentalis Resembling Known Cul- tivars. Mary- NIE. MERE T 352 Snow Cover and the Diapensia lapponica Habitat in the White Mountains, New Hampshire. Weasley N. Tifney; JY. ————— 358 New and Interesting Plants from the Great Plains. Ronald L. McGregor, Lawrence K. Magrath and Ronald R. Weedon ........ ,2 eee nennen ntn eet nennt 378 New Records of North Dakota Plants. O. A. Stevens eese eese sanete tesa asta stet e sess aae ee eee eese eee bes ee 389 Priority of the Binomial Pyrola chlorantha. Erich Haber PP S 396 Chromosome Numbers of Scirpus purshianus and S. smithii. VU ME ONES] 11771 1 RR 398 Tussilago farfara L. (Senecioneae-Asteraceae) in Virginia. Miles F. Johmsom crsccccccccssssesssscessscceccccevesenssecsssseeceesceeeeeeeseneeees 408 Investigations of New England Marine Algae V. The Algal Vegetation of the Hampton-Seabrook Estuary and the Open Coast near Hampton, New Hampshire. Arthur C. Mathieson and Richard A. Fralick .................. 406 Edgeworthia (Thymelaeaceae) New to the Western Hem- isphere. Wilbur H. Duncan and Marie Mellinger ............................ 436 Request from Editor ................... eere eene 440 Instructions for Contributors to Rhodora .................................. 440 Hovdova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON > Associate Editors RADCLIFFE BARNES PIKE STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY Vol. 74 December, 1972 No. 800 CONTENTS: Taxonomic Fern Notes, VI— New Species of American Cyatheaceae. Rolle, Tryon .srececssscseccssccscrsserssscececevsssesscscecececscsocsonscnesenossnssesesese 4. 441 New Species in the Phacelia crenulata Group (Hydrophyllaceae) N. Duane Atwood E AAA A E 451 Rediscovery of Tilia neglecta Spach. Henry K. Svenson aaa ana tte Des 469 The Ecology of Benthic Salt Marsh Algae at Ipswich, Massa- chusetts I. Zonation and Distribution of Algal Species. E. E. Webber and R. T. Wile ette 475 (Continued on Inside Back Cover) The Nef England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02138 STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION Title of Publication: RHODORA Date of Filing: Sept. 25, 1972 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 Names and Addresses of Publisher, Editor, and Managing Editor: Publisher: New England Botanical Club, Inc., Botanical Museum, Oxford Street, Cambridge, Mass. 02138 Editor: Albion R. Hodgdon, Dept. of Botany, University of New Hampshire, Durham, N. H. 03824 Managing Editor: Herman R. Sweet, Botanical Museum, Harvard University, Oxford Steet, Cambridge, Mass. 02138 Owner: No stockholders Known Bondholders, Mortgagees, and other Security Holders: None Total No. copies printed 875 875 Paid Circulation Sales through dealers, etc. None None Mail subscriptions 772 765 Total paid circulation 772 765 Free distribution 20 20 Total distribution 792 785 Office use, left-over, etc. 83 90 Total 875 875 I certify that the statements made by me above are correct and complete. Herman R. Sweet, Business Manager MANUFACTURED BY THE LEXINGTON PRESS, INC. LEXINGTON, MASSACHUSETTS Rhodora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 74 December, 1972 No. 800 TAXONOMIC FERN NOTES, VI — NEW SPECIES OF AMERICAN CYATHEACEAE. ROLLA TRYON The many ample collections of tree ferns made during explorations in South America by Julian Steyermark, Bas- sett Maguire, John Wurdack, Ramón Ferreyra, Benkt Sparre, and Paul Hutchison and Kenneth Wright are an essential source of information for studies of the Ameri- can Cyatheaceae now in progress. Several of the new species among them are described here in addition to the species of tree fern that is characteristic of the mossy forest of Puerto Rico. Some of the endemism in the Cyatheaceae in Guayana is reflected in the four new species from that region. Al- though endemism in the ferns in this area is not as strong as in many groups of flowering plants, it is significant especially with respect to the remarkable morphological innovations. The species are described under the genera recognized in my recent paper on the classification of the family (Tryon, 1970) and special terms employed are defined there. The illustrations have been prepared by Sally Landry, Mary Robbins, and Lydia Wunsch. Sphaeropteris. sipapoensis Tryon, spec. nov. Figs. 1-4. Petiolus sine spinis squamis multis persistentibus struc- tura conformi circa 1-2 cm longis brunneolo-albis manifeste ciliatis; lamina bipinnata pinna apicali subconformi; rhachis 441 442 Rhodora [Vol. 74 squamata; pinnae ad 40 cm longae petiolulatae; pinnulae integra vel aliquantum lobatae subcoriaceae articulatae breviter petiolulatae base late cordata apice subacuto vel acuminato pagina superiore glabra pagina inferiore squamis valde dissectis ; venae liberae; sori submarginales plerumque a margine circa 0.5 mm exindusiati paraphysibus longiori- bus quam sporangiis albidis circa 1 mm longis. Holotypus: Mixed forest at Intermediate Camp, Cerro Sipapo (Pará- que), Terr. Amazonas, Venezuela, 2 Feb. 1949, B. Maguire and L. Politi 28165, NY. Isotypus: GH. Sphaeropteris sipapoensis is closely related to S. margi- nalis (Kl) Tryon of British Guiana and Surinam. The two species share the following characters: large, brown- ish-white, long-ciliate petiole scales, stalked pinnules, a nearly conform terminal pinna at the lamina apex and the absence of an indusium. Sphaeropteris marginalis has the sori farther (mostly 1.2-2.0 mm) back from the margin than S. sipapoensis and the paraphyses are shorter (ca. 0.5 mm long) and tan to brown in color. Sphaeropteris marginalis also has smaller, less coriaceous, and more deeply lobed pinnules than S. sipapoensis. Further collections are required in order to assess the importance of these pinnule characters. The related S. marginalis has been reported from Mexico (for example, by Maxon and Morton, p. 70, 1948) but this is based on an erroneous association of a specimen of the species at Kew ! with the label of Jurgensen 915, Sierra San Nolasco, Mexico. Fournier (p. 135, 1872) cites Jurgensen 915 as Hemistegia decurrens [= Cnemidaria decurrens (Liebm.) Tryon], a species of Mexico and un- doubtedly the one properly associated with the label. Sphaeropteris Atahuallpa Tryon, spec. nov. Figs. 5-8. Petiolus parce tuberculatus squamis structura conformi ad 6 cm longis 1 cm latis brunneolo-albis basem versus densissimis persistentibus squamis minoribus longe ciliatis apice filiformi concolori squamellis multis valde dissectis etiam instructus; lamina bipinnato-pinnatifida vel juxta 1972] Cyatheaceae — Tryon 443 rhachim bipinnato-pinnatisecta apice sensim acuminato; pinnae ad 70 cm longae subsessiles; pinnulae subsessiles segmentis minime crenato-serratis costulis inferne squamis albidis; venae liberae; sori ad furcam venarum; indusium subfuscum sphaeropteroideum. Holotypus: Cerros Calla Calla, above Balsas on road to Leimebamba, 3000-3100 m, Prov. Chachapoyas, Dept. Amazonas, Peru, 14 Oct. 1964, P. C. Hutchison and J. W. Wright 6922, GH. Paratypus: Summit of Puma-urcu, southeast of Chachapoyas, 3100-3200 m, Prov. Chachapoyas, Dept. of Amazonas, Peru, 3 July 1962, J. J. Wurdack 1153, GH. Sphaeropteris Atahuallpa is a distinctive species, related by its sphaeropteroid (globose) indusium only to the species of the Sphaeropteris horrida group (Tryon, 1971). From these, however, it differs in the petiole scales which lack a dark apical seta and dark rigid marginal teeth. The largest petiole scales are five to six cells thick near their base. Atahuallpa, a son of Huayna Capac, was Lord-Inca of the Kingdom of Quito. He was captured at Cajamarca by Francisco Pizarro, then ransomed with silver and gold to the present value of perhaps fifty million dollars, and finally betrayed and murdered by Pizarro. Alsophila bryophila Tryon, spec. nov. Figs. 9-10. Petiolus brevissimus modice tuberculatus squamis struc- tura marginata brunneis apice seta minima brunnea fragili; lamina pinnato-pinnatisecta vel juxta rhachim bipinnata basem versus modice vel sensim deminuta apice sensim acuminato; rhachis pubescens plus minusve squamata; pinnae ad 15-25 cm longae sessiles costa pagina superiore pubescenti pagina inferiore pubescenti saepe squamis sub- planis sine setis fuscatis; segmenta integra coriacea utrin- que longe pubescentia costulis basem versus squamis paucis albidis bullatis sine setis; venae liberae; sori ad furcam venarum; indusium hemitelioideum vel meniscoideum. Holotypus: Near West Peak, Luquillo National Forest, Puerto Rico, 12 Nov. 1966, G. J. Gastony 12, GH. Para- typi (all from Puerto Rico): Howard, et al. 15645, 15722, 444 Rhodora [Vol. 74 EN US NN: NW Y a L / // ]] |] (IJ Wy XM NOI Figs. 1-4. Sphaeropteris sipapoensis, (Maguire and Politi 28765). Fig. 1, basal pinna, X 1/3, GH. Fig. 2, central pinnules of central pinna, X 1/3, Ny. Fig. 3, portion of fertile pinnule (paraphyses removed from receptacles to left), X 1, NY. Fig. 4, articulate pin- nule base, enlarged, NY. Figs. 5-8. Sphaeropteris Atahuallpa (Hutchinson and Wright 6922, GH). Fig. 5, apical portion of crozier, X 1/3. Fig. 6, portion of petiole, X 1/3. Fig. 7, central pinnules of central pinna, X 1/3. Fig. 8, fertile segments: sori and venation (left), with scaly indu- ment (right), X 1 1/3. Figs. 9-10. Alsophila bryophila (Gastony 12, GH). Fig. 9, cen- tral pinna, X 1/3. Fig. 10, fertile segments: indusia (upper right), with pubescence (upper to left), venation and receptacles (lower), x 1 1/3. 1972] Cyatheaceae — Tryon 445 15725, A; Howard and Nevling 15740, A; Shafer 3305, GH, MO; Shafer 3632, F, NY; Sintenis 1480, GH. Alsophila bryophila grows in wet forests in Puerto Rico at elevations of about 900-1000 m. The species is char- acteristic of the mossy (elfin) forest of the Luquillo Mountains and the specific epithet is derived from this association. Alsophila bryophila is close to A. minor (D. C. Eaton) Tryon, A. hotteana (C. Chr. and Ekman) Tryon and A. Brooksii (Maxon) Tryon, from which it can be distin- guished by the hemitelioid to meniscoid indusium and the absence of dark setae on the scales of the under surface of the pinnae. The three related species all have a cup-shaped indusium and some, frequently many, scales or squamules on the under surface of the pinnae with dark setae on the margins and a dark seta at the apex. They also have the upper surface of the segments glabrous to rarely sparingly pubescent, which in A. bryophila the segments are usually strongly pubescent above. Alsophila Brooksii, also in Puerto Rico, is fully bipinnate through most of the lamina. Some materials of A. hotteana have a meniscoid indusium but in these specimens it is not certain that the structure is com- plete. They do have the distinguishing dark-armed indu- ment on the axes beneath. The other species of American Alsophila: A. Abbott (Maxon) Tryon, A. capensis (L. f.) J. Sm., A. confinis (C. Chr.) Tryon, A. dryopteroides (Maxon) Tryon, A. Engelii Tryon, A. Nockii (Jenm.) Tryon, A. paucifolia Baker, A. Salvinii Hook. and A. Urbanii (Brause) Tryon, differ from A. bryophila in having a cup-shaped indusium and (or) in having a fully bipinnate or more complex lamina, and in different indument on the under surface of the pinnae. Alsophila bryophila has long been known under the name of Cyathea pubescens Kuhn. That species, however, is Nephelea pubescens (Kuhn) Tryon and is endemic to the Blue Mountains of Jamaica (Gastony, 1972). Cyathea 446 Rhodora [Vol. 74 pubescens has also been ascribed to Cuba and Hispaniola (Maxon, p. 386, 1926), but the specimens from those islands belong to Alsophila minor. Trichipteris Steyermarkii Tryon, spec. nov. Figs. 11-12. Petiolus basem versus acute tuberculatus squamis struc- tura marginata; lamina bipinnata pinna apicali conformi; pinnae circa 25-35 cm longae articulatae breviter petiolu- latae; pinnulae integrae vel subintegrae coriaceae articu- latae breviter petiolulatae base cordata apice subacuto vel acuto pagina superiore glabra pagina inferiore squamis pau- cis brunneolis subplanis; venae liberae; sori ad furcam vena- rum exindusiati paraphysibus brevioribus quam sporangiis. Holotypus: Bosque nublado virgen en la cumbre, Cerro de Humo, Peninsula de Paria, Estado Sucre, Venezuela, 2 Mar. 1966, J. A. Steyermark 94923, VEN. Isotypi: GH, U. Paratypus: Selva nublada en la cumbre plana, Cerro Patao, Peninsula de Paria, Estado Sucre, Venezuela, 19 Jul. 1962, J. A. Steyermark and G. Agostini 91048, VEN. Trichipteris Steyermarkii is related in its lamina archi- tecture and articulate pinnae and pinnules to T. sagittifolia of Trinidad and to T. cordata of the states of Aragua and Carabobo, Venezuela. Both species differ from T. Steyer- markii in having bullate scales on the lower surface of the pinnules (T. sagittifolia also has some flattish ones). Futher differences are the many long slender spines at the base of the petiole of T. sagittifolia and the pinnately lobed pinnules of T. cordata. Hooker (p. 37, 1865) questioned the source of the Cruger collection upon which Alsophila sagittifolia was based: "Trinidad ? (possibly the opposite coast of Venezuela)". The type collection (Kew !) has bullate scales on the pin- nules and corresponds with the Trinidad species (for ex- ample, Fendler 142 and 143). Trichipteris Cyclodium Tryon, spec. nov. Figs. 13-14. Petiolus sine spinis squamis structura marginata brun- neis albido-limbatis dense ciliatis; lamina 1-pinnata pinna 1972] Cyatheaceae — Tryon 447 apicali conformi articulata; pinnae circa 15-18 cm longae valde ascendentes integrae late crenatae vel minime pin- natilobatae valde coriaceae articulatae petiolulatae utrinque glabrae vel subglabrae base late cuneata apice obtuso; venae liberae utrinque elevatae; sori plerumque in venis simpli- cibus 3-4 seriati exindusiati paraphysibus longioribus quam sporangiis. Holotypus: Camp Savanna, Cerro Sipapo (Paráque), Terr. Amazonas, Venezuela, 6 Dec. 1948, B. Maguire and L. Politi 27541, NY. Isotypus: GH. Trichipteris Cyclodium is a very distinctive species. The lamina has a close resemblance to that of Cyclodium menis- cioides (Willd.) Presl (a species that can also be placed in Stigmatopteris or Dryopteris, depending upon how generic lines are drawn). Trichipteris Cyclodium is related to T. Williamsii (Maxon) Tryon of Panama and to the next species T. Maguirei by its simple, articulate and stalked pinnae and by its lamina with a conform terminal pinna. It differs from T. Williamsii in its very coriaceous, strongly ascending, obtuse pinnae and whitish-bordered petiole scales. Trichipteris Williamsii has the pinnae only moder- ately thickened, patent and long-acuminate and the petiole seales are entirely whitish or have a brown stripe toward the apex. A comparison with T. Magwuirei is made under that species. Trichipteris Maguirei Tryon, spec. nov. Figs. 15-16. Petiolus sine spinis squamis structura marginata brunneis albido-limbatis dense ciliatis ; lamina 1-pinnata pinna apicali conformi articulata vel apice binato; pinnae circa 8-12 cm longae integrae coriaceae articulatae utrinque glabrae pinnis basilaribus circa 7 mm petiolulatis base cuneata apice abrupte acuminato serrato; venae liberae vel aliquot junctae utrinque elevatae; sori plerumque in venis simpli- cibus 1-2 seriati exindusiati paraphysibus longioribus quam sporangiis. Holotypus: Frequent, terrestrial, talus forest near first escarpment, Cerro Sipapo (Paráque), Terr. Amazonas, Venezuela, 11 Dec. 1948, B. Magwire and. L. Politi 27597, NY. Isotypus: GH. Paratypi: (all from 448 Rhodora [Vol. 74 Figs. 11-12. Trichipteris Steyermarkü (Steyermark 94923, VEN). Fig. ll, central pinna, X 1/3. Fig. 12, fertile pinnule, with recep- tacles and scales, X 1. Figs. 13-14. Trichipteris Cyclodium (Maguire and Politi 27541, NY). Fig. 18, central pinnae, X 1/3. Fig. 14, portion of fertile pinna, with sori, X 2/3. Figs. 15-16. Trichipteris Maguirei (Maguire and Politi 27597, GH). Fig. 15, central pinnae, X 1/3. Fig. 16, portion of fertile pinna, with sori, X 1. Figs. 17-19. Cyathea Steyermarkii (Steyermark 105194, GH), Fig. 17, lamina apex, X 1/3. Fig. 18, central pinnules of central pinna, X 1/3. Fig. 19, portion of fertile pinnule, with receptacles, X 1. 1972] Cyatheaceae — Tryon 449 Terr. Amazonas, Venezuela) Terrestrial, uncommon, wet montane moss forest, Phelps Camp to North Savanna, 1400 m, Cerro Sipapo (Paráque), 17 Dec. 1948, B. Maguire and L. Politi 27752, NY; terrestrial, wet mixed montane forest, slope above Cano Grande, 1500 m, Cerro Sipapo (Paráque), 21 Jan. 1949, B. Maguire and L. Politi 28493, NY; dense woodland along right fork of Cano de Dios, 1900 m, Cerro Huachamacari, Río Cunucunuma, 13 Dec. 1950, B. Maguire, R. S. Cowan and J. J. Wurdack 30188, NY. Trichipteris Maguirei differs from the preceding species, T. Cyclodium, in its smaller pinnae that are cuneate at the base and abruptly acute to acuminate at the apex, and in the lamina that often has a pair of terminal pinnae. The larger pinnae of T. Cyclodium are broadly cuneate at the base and obtuse at the apex, and the lamina has a single terminal pinna. Tvichipteris Maguirei is perhaps more closely related to T. Williamsii, but differs from it in the smaller pinnae, ca. 8-12 cm. long, the shorter pinna stalks, those of the basal pinnae being ca. 7 mm long, and the petiole scales with whitish borders. In T. Williamsii, the pinnae are ca. 15-20 cm long, the stalks of the basal ones are ca. 13 mm long, and the petiole scales are entirely whitish or have a brown stripe toward the apex. Cyathea Steyermarkii Tryon. spec. nov. Figs. 17-19. Petiolus basem versus modice muricatus squamis struc- tura marginata brunneis brunneolo-albis-limbatis; lamina bipinnata pinna apicali conformi articulata; pinnae circa 25-30 cm longae articulatae petiolulatae apice sensim acuminato; pinnulae non articulatae crenatae vel aliquan- tum pinnatilobatae chartaceae base cuneata apice obtuso vel subacuto; venae liberae; sori plerumque in venis simplicibus inter costam et marginem medii; indusium parvum hemi- telioideum. Holotypus: Cumbre del Cerro Autana, 4° 52” N. lat. 67° 27” W. long., 1230-1270 m, Terr. Amazonas, Venezuela, 21-22 Sept. 1971, J. A. Steyermark 105194, GH. 450 Rhodora [Vol. 74 Cyathea Steyermarkii is related to three other species with a hemitelioid indusium: Hemitelia conformis Tryon, H. petiolata Hook. and H. Woronovii Maxon and Morton. I placed the first two in Cnemidaria (Tryon, 1970) ; how- ever, Robert Stolze, who is monographing the genus, in- forms me that these species and H. Woronovii are to be excluded and properly belong in Cyathea. Hemitelia petio- lata is the only one of these hemitelioid species that has been placed in Cyathea (as C. panamensis Domin). The three species have the sori close to the margin, while in C. Steyermarkii the sori are about half way between the costa and the margin. Hemitelia petiolata and H. Woro- novii have the apex of the lamina gradually reduced to the tip, while in C. Steyermarkii the apex is a conform termi- nal pinna. Hemitelia conformis is further distinguished by the pinnae that have a conform, articulate terminal pinnule. In C. Steyermarkii the pinnae are gradually re- duced at the apex. LITERATURE CITED FOURNIER, E. 1872. Mission Sci. Mex. Amér. Cent. Recherches Bo- taniques, I. Cryptogamie (Mex. Plantas, I. Cryptogamie). Paris. GASTONY, G. J. 1972. A revision of the fern genus Nephelea. Con- trib Gray Herb. 203. (in press) Hooker, W. J. 1865. Synopsis Filicum. London. Maxon, W. R. 1926. Pteridophyta of Porto Rico and the Virgin Islands. Sci. Surv. Porto Rico and V. I. 6 (3): 378-521. MAxon, W. R. and C. V. MORTON. 1948. Pteridophyta, i» B. Ma- guire, et al., Plant explorations in Guiana in 1944. Bull. Torrey Bot. Club 75: 66-80. Tryon, R. 1970. The classification of the Cyatheaceae. Contrib. Gray Herb. 200: 1-53. 1971. The American tree ferns allied to Sphaeropteris horrida. Rhodora 73: 1-19. GRAY HERBARIUM, HARVARD UNIVERSITY, CAMBRIDGE, MASS. 02138 NEW SPECIES IN THE PHACELIA CRENULATA GROUP (HYDROPHYLLACEAE)' N. DUANE ATWOOD While preparing a taxonomic revision of the Phacelia crenulata group it was necessary to investigate the rela- tionships of several taxa in southern Utah, northern Arizona and Mexico. These investigations demonstrated the presence of several previously undescribed taxa. These areas are well known for their varied geology, topography and climate. They have also been some of the least bota- nized and most poorly understood localities within the range of the P. crenulata group. Therefore, it is not sur- prising that new species are present, particularly in a genus which contains many taxa with restricted distribu- tion. The purpose of this paper is to present the diagnoses, descriptions, distributions, and illustrations of the pro- posed new taxa. The herbaria where specimens are deposited are abbre- viated according to Lanjouw and Stafleu (1964). Those herbaria not listed in Lanjouw and Stafleu are as follows: Dixie Junior College, St. George, Utah (DIX), Weber State College, Ogden, Utah (WSC), and West Texas State Uni- versity, Canyon, Texas (WTS). The author is grateful to the curators of herbaria that were visited or who sent loan material for this study, and to Mrs. Kaye Thorne for preparation of the illustrations. Phacelia constancei Atwood sp. nov. A P. palmeri differt caulibus brevioribus, foliis angusti- oribus et revolutioribus, et caulibus rubellis pilioribus et ramis lateralibus frondosioribus. Herba biennis erecta, 1.5-4.8 dm alta, frondosa omnino; folia petiolata (pro parte maxima), 1.0-10 cm longa, 0.3- "This study was supported in part by the Department of Botany and Range Science, Brigham Young University, Provo, Utah, and by Cost of Education Funds from a National Defense Education Act Title IV Fellowship. 451 452 Rhodora [Vol. 74 1.5 em lata, involuta, e undulata ad pinnatifida, lineares lanceolata, hirsutula supra, pilis glandularis dispersis; caules crasses, simplices vel ramosi omnino, rubelli, e hirsutuli ad hirsuti, et minute glanduliferi; inflorescentia, cymarum scorpiodorum compositorum; pediceli usque ad 1 mm longi; sepala e elliptica ad oblanceolata, 3-4 mm longa, hirsutuli ad hirsuti et stipitata glandulifera; corolla tubu- laris, 5-6 mm longa; stamina exserta 3-4 mm; stylus fur- catus pubescens infimus 1/3, exsertus 3-4 mm stamina longior; capsula subglobosa, hirsutula et glandulifera omnino, sepala brevior; semina 4, nigricantia, 2.5-2.8 mm longa, 1.0-1.2 mm lata, elliptica, margines corrugati, pagina ventralis foveolata, excavata et divisa porca prominens, porca lata una corrugata, pagina dorsalis foveolata. TYPE: Arizona: COCONINO COUNTY: 1 mi north of Fre- donia, on clay hills, growing with Sarcobatus, Lycium, Astragalus, and Phacelia corrugata, 27 May 1968, D. At- wood 1385A (HOLOTYPE: BRY; ISOTYPES: ARIZ, ASC, CAS, DIX, GH, NY, POM, RM, UC, US, UT, UTC, WSC). Additional materials examined: Arizona: MOHAVE COUNTY: ca 19 mi south of Kanab Indian Reservation (southern boundary) on road to Tuweep junction, 27 May 1969, D. Atwood 1765, (BRY); 5 mi northeast of Tuweep, 17 August 1946, K. F. Parker, et al. 6254, (GH, NY, POM, US); ca 4 mi north of Tuweep junction, 27 May 1969, D. Atwood 1761, (BRY, CAS); 1 mi north of Fredonia, 5 June 1968, D. Atwood 1529, (BRY, GH, UC) ; 1 mi north of Fredonia, 27 May 1969, D. Atwood 1785, (ARIZ, BRY, CAS, NY, US, UTC, WSC); Utah: KANE COUNTY: ca 30 mi east of Kanab along Hwy 89, mi post 30, 6 June 1968, D. Atwood 1532, (BRY, CAS, GH, NY, US); 15 mi east of Kanab, 5 June 1969, D. Atwood 1739A, (ARIZ, BRY, US, UT); 15 mi east of Kanab, 5 June 1942, Ripley & Barneby 4836, (CAS, RSA); 20 mi east of Kanab, 5 June 1969, D. Atwood 1792B, (BRY, CAS, GH, NY, POM, RM, US); ca 32 mi east of Kanab along Hwy 89, 6 June 1968, D. Atwood 1532A, (BRY, POM, RSA); ca 3.7 mi east of Skutumpah-Alton junction at head of Johnston Canyon (R.4. 5W., T.408.), 7 June 1969, D. Atwood 1801A, (BRY). Taxonomically, Phacelia constancei (Figure 1, Map 1) appears to be most closely related to P. palmeri Torr. ex S. Wats., but is distinguished from that species by the smaller growth form, narrower and more revolute leaves, the stems are reddish with shorter and fewer hairs and the lateral inflorescence branches are more leafy. 1972] Phacelia — Atwood 453, Fig. 1. Phacelia constancei. Habit and seed of plant. 454 Rhodora [Vol. 74 ^i l Map 1. Southern Utah and northern Arizona. Range of P. con- stancei. The latter occurs from Mohave County, Arizona, west to Washington and Iron County, Utah, and in Clark County, Nevada. The new taxon is known only from restricted areas in Mohave County, Arizona north to Kane County, Utah. Both species grow on alkaline clay bluffs and flats of the Moenkopi Formation, late May to mid August. Chromosome voucher specimen (n=11): Tuweep Valley, 5 mi ne. of Tuweep, 17 August 1946, K. F. Parker et al. 6254, (UC). This species is named in honor of Dr. L. C. Constance, University of California at Berkeley, who has made nu- merous important contributions in the genus Phacelia. Phacelia rafaelensis Atwood sp. nov. A P. utahensis differt corolla tubularo diluta et canali- culata lobis aliquatum expansis, staminibus et stylis exsertis tantum 3-5 mm, filis et antheris tristis, semine porca inter- dum corrugata. 1972] Phacelia — Atwood 455 Herba biennis, erecta, 0.8-5.4 dm alta; caules crassi simplices vel interdum ramosi basi, e olivacei ad brunnei glandiferi et hirsuti; folia basalia petiolata dentata ad pinnatifida, 2-7 cm longa, 0.5-1.5 em lata, strigosa ad hir- suta, folia caulina sessilia, undulati-crenata ad dentata, oblonga ad lanceolata, 1.0-10 cm longa, 0.5-3.5 cm lata, strigulosa ad hirsuta et sparsim stipitati-glandulifera ; inflorescentia terminalis proparte maxima interdum axil- laris, paniculata; flores fere sessiles; sepala oblonga ad spathulata, 3-4 mm longa in fructis, 1.0-1.7 mm lata, gland- ulifera et hirsuta; corolla tubularis pallida, 5-6 mm longa; stamen et stylus exsertus 3-5 mm; stylus 34 longitudis, pubescens infernus 14; capsula globosa, 4-5 mm longa, stipitati-glandulifera et hirsuta; semina 4, elliptica ad oblonga, 3.4-4.0 mm longa, 1.5-2.0 mm lata, pagina ven- tralis alveolata pallidior pagina dorsalis, excavata et divisa porca prominenti, porca lata una corrugata, margines integrae, pagina dorsalis brunnea et alveolata radose. TYPE: Utah: WAYNE COUNTY: Capital Reef National Monument, red clay hills (Moenkopi Formation) southeast of visitor center, 12 June 1969, D. Atwood & L. C. Higgins 1834, (HOLOTYPE: BRY; ISOTYPES, ARIZ, ASC, B, BRY, CAS, COLO, DIX, GH, JEPS, MEXU, NY, POM, RM, RSA, UC, US, UT, UTC). Additional materials examined: Arizona: COCONINO COUNTY: Lee's Ferry, 16 June 1890, M. E. Jones s.n, (POM); 2 mi w. of Navajo Bridge, 12 May 1955, Ripley & Barneby 12664, (CAS); MOHAVE COUNTY: 2 mi e. of Fredonia, 6 June 1942, Ripley & Barneby 4362, (RSA); ca 4 mi w. of Fredonia, mi post 22, 27 May 1968, D. Atwood 1390, (ARIZ, ASC, BRY, CAS, POM, US); 1 mi n. of Fredonia, 5 June 1968, D. Atwood 1530, (BRY, B, POM); ca 4 mi s. of Fredonia-Pipe Springs road, 13 May 1952, E. McKlintock 52-600, (CAS); Utah: EMERY COUNTY: Chinle Formation, at jct of Buckhorn Wash & San Rafael River Canyon north of river, 30 May 1970, Welsh, Atwood & Moore 9819, (ARIZ, ASC, B, BRY, CAS, GH, NY, RM, RSA, UC, US, UT, UTC, WSC); 2 mi s. of San Rafael River Bridge, 16 June 1969, D. Atwood 1843, (ARIZ, ASC, B, BRY, GH, NY, US, WSC) ; 11 mi n. of Goblin Valley turn off on Temple Mountain road from US 24, thence 43 mi s.w. on mining road, 17 June 1969, D. Atwood 1855, (BRY, RM); R.9E., T.26S., San Rafael Swell, 17 June 1969, D. Atwood 1847, (ARIZ, 456 Rhodora [Vol. 74 ASC, B, BRY, CAS, GH, NY, UC, US, UT, UTC); ca 13 mi e. of Buckhorn Wash, north of San Rafael River, 30 May 1970, Welsh, Atwood & Moore 9846, (BRY); ca 5 mi e. of Buckhorn Wash along road north of San Rafael River, 30 May 1970, Welsh, Atwood & Moore 9844, (ARIZ, B, BRY, CAS, GH, NY, US); ca 2 mi w. of San Rafael River Bridge at jet. of Buckhorn Wash and San Rafael River, 1 June 1970, Welsh, Atwood & Moore 9903, (BRY, NY, US, WSC); WASHING- TON COUNTY: 11 mi w. of Hurricane, 2 May 1942, F. Gould 1681, (CAS, UC); near Virgin, 10 May 1941, Eastwood & Howell 9200, (CAS); e. of Bloomington, 10 April 1970, Welsh, Atwood & Matthews 9535, (BRY, NY, UC, WSC); 2 mi e. of Hurricane, 27 May 1968, D. Atwood 1409, (BRY, CAS); Rockville, 12 May 1941, Eastwood & Howell 9327, (CAS, GH); 4 mi s.w. of Saint George, 27 May 1968, D. Atwood 1417, (BRY); Moenkopi Formation at Bloomington, 3 May 1970, Welsh, Atwood & Matthews 9694, (BRY); clay hills n. of Bloomington, 25 May 1969, D. Atwood 1698, (ARIZ, ASC, B, BRY, CAS, GH, JEPS, NY, RM, UC, UTC, WSC); ca 5 mi s.w. of St. George, 7 April 1968, Higgins 1244, (BRY); WAYNE COUNTY: Capital Reef Wash 2 mi s.e. of Fruita, 6 May 1940, B. Maguire 18146, (CAS, NY, UC, UTC) ; 3 mi s. of Highway 24 on Notom Road, 17 June 1969, D. Atwood 1860, (ASC, BRY, CAS, GH, RSA, UC, UT, US, WSC); Moenkopi Formation at Fruita, 10 May 1969, Welsh Atwood & Higgins 8968, (BRY, CAS, RM, RSA, UC, US, UT); 3 mi w. of Fruita, 4 June 1953, R. McVaugh, 14438, (CAS, NY, UC). Phacelia rafaelensis appears to be related to P. utahensis but differs from that species as indicated in the diagnosis. P. utahensis is endemic to the Arapien Shale Formation in Sevier and San Pete County, Utah, whereas P. rafae- lensis (Figure 2, Map 2) occurs from Emery County south to Kane County, Utah and Mohave County, Arizona and east to Washington County, Utah. It grows on clay hills. of the Moenkopi Formation. mid April to mid June. This species is named for the San Rafael Swell, a floristic- ally unique area, in Emery County, Utah. Phacelia howelliana Atwood sp. nov. A. P. corrugata differt corollis pusillioribus et tubo albo, seminibus diluti-marginatis dorsalibus laevibus, plantis brevioribus et plerumque ramosis et frondosis basi. Plantae annuae, 0.9-2.3 dm altae; caules pro parte max- ima et frondosi ad basin versus, glandulifera et aliquot pili longiores simplices praesenta ; folia e late oblonga ad ovales, Phacelia — Atwood 457 1972] Phacelia rafaelensis. Habit and seed of plant. Fig. 2. [Vol. 74 Rhodora 458 2” =. = = o m mo ae m m aa a e e m ml tS Lj r 1 I l / ——— ——————————————-—h co m--——-—-—---—------2—----2--2-2-2--4---- eee eee ee Southern Utah and northern Arizona. Range of P. rafae- Map 2. lensis. 1972] Phacelia — Atwood 459 2-6 cm longa, 1.0-2.5 cm lata, irregulariter crenata ad lobata, strigosa et leviter glandulifera, petiolus usque ad 5.0 cm longa; inflorescentia cymarum scorpiodorum compositorum, pedicelli usque ad 2.0 mm longi; sepala linearia ad anguste oblanceolata, 3.5-4.0 mm longa, 1.0-1.5 mm lata, glanduli- fera et hirsuta; corolla 5-6 mm longa, 6-7 mm lata, rotata ad infundibuliformis, lobi pallidi violacei, tubus albus, stamina et stylus exsertus 3-4 mm, stylus breviores stamina furcatus 34, longitudo pubescens infirmus !4; capsula ob- longa ad subglobosa glandularis et hirsutulosa apprime versus apex; semina 4, brunnea, 3.2-4.0 mm longa, 1.4-1.8 mm lata, elliptica, margines corrugati usque involuti ad complanati, pagina ventrali foveolata, excavata et divisa porca prominens, interdum curva lata una et vix corrugata, pagina dorsalis prophyreus laevis et margo pallida circum- tineta. TYPE: Utah: SAN JUAN COUNTY: ca 0.4 mi n. of Bluff, Utah on highway 163, in mouth of canyon on rocky-sandy- clay soil, 13 May 1970, D. Atwood 2454, (HOLOTYPE: BRY; ISOTYPES, ARIZ, ASC, B, BRY, CAS, GH, NY, RM, UC, US, UTC). Additional materials examined: Utah: GRAND COUNTY: 13 mi n. of Moab, 15 June 1944, A. H. Holmgren & S. Hansen 3319, (NY); Moenkopi Formation, T.25S., R.23E., in Castle Valley, 5 June 1970, Welsh, Atwood & Moore 9957, (ARIZ, BRY, CAS, WTS) ; Arches National Monument, vicinity of Courthouse Wash, 2 May 1963, G. L. Pyrah, et al. 71, (BRY); Potash Plant, 20 April 1962, S. L. Welsh, 1670A, (BRY); SAN JUAN COUNTY: Bluff, near the Navajo Twins, 29 April 1961, D. Bright 55, (BRY); Bluff, 24 May 1919, M. E. Jones s.n., (POM); along road just w. of Monument Valley Hospital, 15 May 1970, D. Atwood 2511, (ARIZ, ASC, B, BRY, CAS, DIX, GH, JEPS, NY, POM, RM, RSA, UC, US, UTC, WSC); Goulding Trading Post); 26 June 1948, J. T. Howell 24687, (CAS); canyon of the San Juan River between Bluff and the Colorado River, P. Thompson s.n., May 1941, (CAS); dark canyon-Cataract Canyon, 1 July 1938, E. Clover & M. Jotter 2143A, (GH); ca 2 mi e. of Bluff, 12 May 1944, A. H. Holmgren 3187, (UTC). This entity is related to P. corrugata A. Nels. which ranges throughout most of Utah and extends into Colorado, northern New Mexico and northern Arizona. It is distin- guished from P. corrugata by its low much branched growth 460 Rhodora [Vol. 74 form, smaller corolla and white tube. The leaves are mostly basal and the seeds are different as indicated in the diag- nosis and general description. P. howelliana (Figure 3, Map 3) is known only from San Juan and Grand Counties, Utah where it grows in red sandy, gravelly or sandy-clay soils at ca 4500 to 5000 feet. It probably occurs in Colo- rado near Moab and undoubtedly in Monument Valley in - Fig. 3. Phacelia howelliana. Habit and seed of plant. 1972] Phacelia — Atwood 461 a ee ee ee —— — — ee o Bluff y —?e Map 3. Southern Utah, Grand and San Juan Counties, Range of P. howelliana. 462 Rhodora [Vol. 74 Arizona, although no specimens have been seen from either area. This species is named in honor of John Thomas Howell at the California Academy of Sciences, who has published many important works within the genus Phacelia. Phacelia vossii Atwood sp. nov. Plantae perennis, 2.1-7.3 dm altae; caules erecta e caudice ligneo, hirsutis ad hirsutulis et stipitati-glanduliferis pilis; folia lineara ad lanceolata, 2.0-11 cm lata, revoluta, pagina ventralis glandulifera dense, pagina dorsalis hirsuta et stipitate-glanduliferis dispersis, margines dentatae ad ir- regulariter dentatae, petiolata, petiolus usque ad 1.5 cm longa; inflorescentia terminalis vel secus caules dispersa usque ad 14 longitudo vel, minus, simplex ad cymarum scorpiodorum compositorum, racemi individua usque ad 7.5 cm longa per fructus, pedicelli usque ad 2.0 mm longa per fructus, glandulifera et hirsuta; corolla tubularis (campanulata ?), diluta purpurascens ad albida (?), 5.5 mm longa ; sepala oblanceolata ad spathulata, 4-6 mm longa, 1.5-2.0 mm lata, glandulifera et hirsuta; stamina exserta ; stylus exserta circa 2.0 mm stamina longior, circa 9.0 mm longa, furcatus 94 longitudo, inferna 14, pubescens; capsula ovata, 3.7 mm longa, 2.5-2.7 mm lata, glandulifera et stri- gosa; semina 4, elliptica ad oblonga, 2.5-3.1 mm longa, 1.0- 1.4 mm lata, margines integra ad corrugata parte, pagina dorsalis foveolata, pagina ventralis foveolata et porca prominentia centrale corrugata, porca excavata utrinque. TYPE: Mexico: NUEVO LEON: Sierra Madre Oriental, caleite and limestone hills beyond Pablillo toward Santa Clara, about 15 mi s.w. of Galeana, sparse in arroyos, flowers light purple, 18 July 1934, C. H. & M. T. Mueller 1075, (HOLOTYPE: GH; ISOTYPE, MEXU). Additional materials examined: Mexico: NUEVO LEON: Hacienda Pablillo, Galeana, 13 August 1936, Mary Taylor 156, (UC, NY). Phacelia vossii (Figure 4, Map 4) is known only from the type collections by Galeana in the State of Nuevo Leon, Mexico. It apparently grows in calcite and limestone soils. 463 Phacelia — Atwood 1972] A Fig. 4. Phacelia vossii. Habit and seed of plant. 464 Rhodora [Vol. 74 COAHUILA ;^ \ p EN / ^ € } E 1 \ I t { ) J A r ^ . ` \, oN "d IN t "a " e MU -^77N i S N ` [ { \ \ - \ / t l p -2 v Map 4. Mexico, States of Coahuila and Nuevo Leon. Range of P. vossit. The flower color is not apparent in the specimens at hand, but the label of the type material indicates that it is a light purple. Other members of the crenulata group which pos- sess a pale corolla are often also whitish at times. This taxon appears to be related to P. pinnatifida Griseb. According to Brand (1913) the latter occurs in the Andes of South America in Peru, Boliva, and Argentina, from 3000 to 3800 meters. The specimens cited were identified as P. integrifolia Torr., however P. vossti is easily distin- 1972] Phacelia — Atwood 465 guished from the former by its evident perennial habit, pale more or less tubular corollas, smaller corrugated seeds and more densely pubescent herbage. This species is named in honor of John Voss who revised the crenulata group in 1987. Phacelia welshii Atwood sp. nov. Plantae annuae, 1.0-4.5 dm altae; caules plus minusve flavo-virentes, simplices vel ramosi frondosi, hirsuti et dense tecti multi-cellularis stipitatis, glandibus; folia oblonga ad lanceolata, 1.5-8.0 em longa, 0.5-2.7 cm lata, hirsuta et dense glandulifera, margines undulatae crenatae vel dentatae saepe revolutae, folia basales fasciculata, petiolata, petiolus 2.0 cm longus or minus, folia caulina sessiles vel fere sic, saepe cordata ad basim; inflorescentia cymarum scorpiodorum compositorum, terminalis ad ex- trema principalis caulis et rami lateralis, dense glandulifera et hirsuta, racemi individui congesta, sed laxe per fructus, usque ad 1.0 dm longa, pedicelli usque ad 1.5 mm longa; sepala spat lata glandulifere; corolla campanulata, pur- purascens ad violacea, 5.0-6.0 mm longa et lata, pubescentia ; anthera lutea; stylus exserta circa 8.0 mm, furcatus 94 longitudo, inferna 1⁄4, hirsutulosa et glandulifera; semina 4, oblonga, ventraliter brunnea ad prophyreus dorsale, foveo- lata, 2.8-3.4 mm longa, 1.3-1.5 mm lata, pagina ventrali pagina dorsalis pallidior quam et porcis prominentis, porca corrugata lata una, margines corrugatae et plus minusve revolutae. TYPE: Arizona: COCONINO COUNTY: along highway 89 north of Gray Mountain, on red shale (also common south of town on red shale), 19 May 1970, D. Atwood 2608, (HOLO- TYPE: BRY; ISOTYPES, ARIZ, ASC, B, BRY, CAS, COLO, DIX, GH, NY, POM, RM, RSA, UC, US, UT, UTC, WSC, WTS). Additional] materials examined: Arizona: COCONINO COUNTY: 2.5 mi w. of highway 89-64 jct, red shale, 19 May 1970, D. Atwood 2611, (ARIZ, ASC, B, BRY, CAS, COLO, DIX, GH, NY, UC, US, WSC, WTS); 10 mi n. of Wuptaki National Monument jet with highway 89, 19 May 1970, D. Atwood 2605, (ARIZ, ASC, B, BRY, CAS, GH, NY, POM, RM RSA, UC, US, UT, UTC, WTS) ; ca 1.5 mi n.w. of Headquarters of Wuptaki Na- 466 Fig. 5. Phacelia welshii. Habit and seed of plant. [Vol. 74 1972] Phacelia — Atwood 467 FLAGSTAFF Map 5. Coconino County, Arizona. Range of P. welshii. 468 Rhodora [Vol. 74 tional Monument along highway on red shale, growing with Ephedra, Hilaria, Oryzopsis, blue grama, Atriplex and apache plume, 18 May 1970, D. Atwood 2601, (ASC, BRY, CAS, US, WSC); rocky limestone areas above the Little Colorado River, 6 May 1961, D. Demaree 44391, (CAS, UC); red shale hills along road ca 0.5 mi n. of headquarters of Wupatki National Monument, 18 May 1970, D. Atwood 2598, (ARIZ, B, BRY, CAS, COLO, DIX, GH, NY); Willow Springs n. of Cameron, 7 June 1942, Ripley & Barneby 4875, (CAS) ; red shale 11 mi s. of Cameron, 20 June 1948, Howell 24397, (CAS). P. welshii (Figure 5, Map 5) is probably most closely related to P. corrugata and P. utahensis. It can be distin- guished from the former by its broader, dark brown seeds, long (up to 1.3 mm long), flattened, multicellular, stipitate glands and different leaves. Typically P. corrugata is a more northern plant but is not uncommon in Coconino County. The new taxon can be differentiated from P. uta- hensis by its more open inflorescence, corrugated seeds, and broader, shorter, densely glandular leaves. P. utahensis is endemic to the Arapien Shale Formation in Utah. This species is named in honor of Dr. Stanley L. Welsh, Brigham Young University, for his rich contributions toward the flora of the Intermountain area, and for his encouragement and invaluable help throughout the course of this study. DEPT. OF BOTANY AND RANGE SCIENCE BRIGHAM YOUNG UNIVERSITY PROVO, UTAH 84601 REDISCOVERY OF TILIA NEGLECTA SPACH HENRY K. SVENSON Tilia neglecta has always been a questionable species of Linden. It was described in 1834 from “ambulacris Hort. Bot. Paris” and was stated to be of North American origin. The range is from Montreal to Washington and southwest- ward. Due to variability, and absence of a “type” with which the name could be correlated, it has remained one of the least tangible species in a taxonomically difficult a ; delis y ee Fig. 1. Four views of Tilia neglecta, Sandy Neck, Cape Cod, Massachusetts. 469 470 Rhodora [Vol. 74 genus. In Rhodora 72: 339-344. 1970, I described a “forest” of Tilia neglecta in a dune hollow on Cape Cod, Massachu- setts (fig. 1), and reviewed its subspecific synonyms, but at that time I had not seen the fine treatment of the Ameri- can species by G. N. Jones in Illinois Biological Monographs no. 39, 1968. In the spring of 1970, on a visit to the herbarium of the Jardin des Plantes I found in the European folders a speci- men of T. neglecta labeled in Spach’s hand, dated 1834, and from the “Menagerie”. Then, after making detailed sketches of this specimen, I went over to the western en- trance of the zoological garden. Just a short distance within the gate and opposite the old viper pit, stood a gigantic weatherbeaten linden, which in its leaves corresponded exactly with the herbarium specimen, evidently the long-lost Tilia neglecta. There were no records, but Dr. H. Heine said that it was not a European linden, and suggested that the tree originated from seeds sent by Michaux, which would have produced flowers and fruit by 1834. From my visit in 1937 I remember vividly the rocky viper pit. These vipers are now in elegant quarters in the new reptile house, and the rocks are given over to some harmless snakes. The exhibit, I think, has suffered from wear, and there is a sign in French (a universal language) “Please do not throw stones at the snakes." At the opposite entrance of the *Menagerie" there is a large unlabeled tree of T. heterophylla of about the same vintage as T. neglecta. In the summer of 1970 Dr. Heine made for me a fine series of herbarium specimens of the Menagerie tree (see fig. 2), showing foliage, flowers, and sprout growth. These I have given to the Gray Herbarium. They confirm in microscopic detail the Tilia neglecta specimen mentioned above, and Dr. Heine has labeled this tree as the “holotype”. Accompanying the specimen to be retained by the Gray Herbarium, I have placed the detailed sketches I made of the 1834 specimen. André Michaux (1746-1802) ranged far in the eastern United States. He had two stations for assembling seeds 1972] Tilia — Svenson 471 ee E e a EE EL Fig. 2. Leaf (upper surface) and flower cluster from the type tree of Tilia neglecta (Jardin des Plantes, Paris), X ca. 2/3 (actual size of leaf 11.5 X 9.5 em). and roots; one at Charleston, South Carolina, the other on the east bank of the Hackensack River at New Durham in New Jersey. An account of the New Jersey garden, which covered 8 or 10 acres now occupied by the Hoboken Ceme- tery, is given by Rusby in Bull. Torrey Club 11: 88-90. 1884. Zenkert in Flora of the Niagara Frontier Region, 1934 says that the younger Michaux (1770-1855), having on his first expedition (1801-1803) traveled in the terri- tory west of the Alleghenies, during the three years of his second journey explored the Atlantic Coastal States from Maine to Georgia and made five trips to the interior, one 472 Rhodora [Vol. 74 of which, undertaken in 1806 or 1807, took him from New York to Lakes Ontario and Erie. The Michaux Herbarium at Paris consists of selected plants to illustrate the “Flora boreali-americana" published in 1803. The Paris herbarium is also well represented by other early collectors. In addition to the Michaux Herbarium, there are four folders of American lindens, in which there are many Michaux specimens. I saw in the third folder the specimen collected by W. Riehl in Missouri, labeled as T. neglecta by Spach. It was sent to Jones and correctly annotated as T. americana. Among the early collectors are Tuckerman*, Pearson, Cooper, Torrey, Oakes, and others. In the European folders are three additional sheets of T. neglecta, so named by Spach “fleurs 12 juillet, 1850; fr. Sept. 1851; cultivée dans les promenades de Carlsruhe (grand duché de Baden)". There are also the varieties of Tilia published by A. Braun. Both the American and Euro- pean collections would be well worth going over. For help at the Jardin des Plantes I am also greatly indebted to the Director, Dr. Jean Jacques Leroy, and to Dr. Alicia Lourteig who has charge of the Michaux Herbarium. To return to a survey of American Literature on the subject, we find that Hough, Handbook of Trees... (1907, and reprint 1947), says that T. Michauxii is similar to T. heterophylla, and also confounded with T. pubescens *Edward Tuckerman (1817-1886) a graduate of Union College in Schenectady, was the first, and I think the only curator of their her- barium. In his fine classification of the genus Carex, published in Schenectady and dedicated to John Torrey, he mentions himself as «Mus. et Herb. Concord Cur.". While I was teaching at Union, I came upon a trunk or two filled with Tuckerman specimens, and wrote an account of Tuckerman in the Union Alumni Bulletin (about 1924). The plants consisted primarily of collections by Jona- than Pearson and Lewis Beck from the Schenectady region; plants collected by Tuckerman in Europe, chiefly in the Epping Forest of England; and a large series of European plants given to him by Grisebach. Of Tuckerman's Carices and other interesting plants of eastern America, I found none, and suppose that he took these col- lections with him on leaving Union. I understand that the Union College Herbarium is now dismantled. 1972] Tilia — Svenson 473 Aiton, and not well understood. T. neglecta is not treated by Deam in Flora of Indiana, but L. Braun in The Woody Plants of Ohio (1961) publishes a map, and considers it “a wide-ranging species or species-complex, perhaps inter- mediate between 7. americana and T. heterophylla, but not synonymous with either.” House in Annotated list of the ferns and flowering plants of New York State, 1924, men- tions T. neglecta as frequent in central New York; the lower Hudson River Valley, Long Island, and Staten Island. Wiegand and Eames in Flora of the Lake Cayuga Basin, New York, 1926, always careful in their judgments, note that “the pubescence on the lower surface of the leaf is highly variable, and on it several species have been founded. The pubescence, though always close, fluctuates very gradu- ally in density, and no other structural characters occur. It seems better, therefore, to recognize this closely pubescent form as simply a variety, T. americana var. heterophylla Loudon. Sargent’s T. neglecta is transitional to typical T. americana’. Jones’ detailed treatment recognizes only three species in eastern United States: T. americana, T. heterophylla, and T. caroliniana. Sargent, in Bot. Gaz. 66: 494-496. 1918 states “I now consider T. neglecta to have been the T. Michauxii of Nuttall, which is the T. argentea of Michaux . . . The pubescence on the lower surface of the leaves is so constant and so persistent throughout the season, it seems best to consider it a species rather than a pubescent form of T. glabra. The base of the style in T. neglecta is furnished with long hairs, and that of T. glabra appears to be quite glabrous". In the Paris tree the petals are 7.5 X 3 mm with nar- rowed base; the staminodia about 4.0-4.5 X 1.0 mm. In a microscopic comparison of the Cape Cod and Paris ma- terial, we see that the calyx is of the same shape, but in the Paris specimen the pubescence is not dense. Pedicels. are thin and elongate, about 10 X 0.5 mm (see fig. 2) in the Paris material; but in the Cape Cod specimens densely stellate and about 4-5 X 0.7 mm. For pubescence of the under surface of the leaf, I have gone over the Cape Cod 474 Rhodora [Vol. 74 material more carefully. Under a magnification of 40X, boiled fragments 2 mm square were cut with a razor blade directly on the 0.5 mm grid eyepiece. Simple hairs could then be readily counted for 1 sq. mm. The count of hairs or trichomes from an area, not dense, for 5 mm. square of lower leaf surface: 4-pronged 15-38 (10-25%) ; 3-pronged 1-6 (1-5%); 2-pronged (Malpighian) 6-10 (1%); 6-8- pronged (none); simple 100-150 (90%). This analysis (cf. the footnote on p. 343 of Rhodora, vol. 72) shows that there is a considerable amount of variation. For the Paris tree (in several counts) there was an average of about 375 hairs on 5X5 mm of lower leaf sur- face: 4-pronged (5%); 3-pronged (1%); 2-pronged (396) ; 5-pronged (less than 1%); simple (90%). In this Paris material the 2-pronged hairs are merely bifurcate; the simple hairs are mainly set at an angle on the veins, giving the veins a fuzzy appearance, whereas in the Cape Cod material the larger veins are practically glabrous. As an exeuse for this statistical treatment, I may cite the statement of Jones (p. 110) that much of the practical identification and classification of Tilia must be made on the amount and kind of pubescence, particularly on the underside of the leaves. Probably this statistical method is not the best one, but cutting and counting is not as diffi- cult as it might seem. Perhaps the pattern of veining of the upper surface of the leaf would be more adaptable. At any rate, I have begun at the beginning, as Fernald would say, and the different opinions of capable botanists and the variations observed by them in these leaf struc- tures, shows how difficult the subject is. I have accumu- lated notes and sketches of several specimens in the Paris Herbarium, and I have gone over casually the specimens of T. neglecta in the Gray Herbarium. I do not have the inclination to pursue the subject further. — Osterville, Mass. 02655. THE ECOLOGY OF BENTHIC SALT MARSH ALGAE AT IPSWICH, MASSACHUSETTS I. ZONATION AND DISTRIBUTON OF ALGAL SPECIES E. E. WEBBER AND R. T. WILCE In a previous publication (Webber & Wilce, 1971) we treated the salt marsh algae at Ipswich in light of their oc- currence, morphology, cytology, seasonal and reproductive periodicities, based on both field and laboratory investiga- tions. In this paper the descriptive ecology of these plants is discussed. The ecological nomenclature used to describe vertical and horizontal distributions of algae has varied considerably. Rather than adopt new terms describing the zoned character of the salt marsh algal vegetation, we have selected estab- lished terminology common to terrestrial ecological studies which applies also to flowering plants of the marsh environ- ment (Weaver & Clements, 1938; Feldmann, 1951; Oosting, 1956; den Hartog, 1959). Accordingly, we have attempted to describe both the physiognamy and the vegetationa] units at each station, the latter in relation to their successional character. Superficially, the zoned arrangement of an algal salt marsh vegetation often is not easily detected. The flat ex- panse of varying shades of green which cover the salt marsh surface contrasts sharply with the distinct and frequently colorful patterns of algal zonation of the rocky coast (see Frontispiece, Lewis, 1964). In order to discuss algal zona- tion and distribution at the Ipswich salt marsh, it was necessary to identify levels on the marsh surface with which to relate the presence and vertical range of algal species. Marsh phanerogams, owing to their abundance, distinctive- ness, and regular and sharp zonation, served as reference points to this end. From the seaward edge of the marsh extending landward, the following seed plant associations 475 476 Rhodora [Vol. 74 were recognized: Spartina alterniflora var. glabra, Spar- tina patens, Spartina patens-Scirpus americanus (or Spar- tina patens-Distichlis spicata) mixture, and Juncus gerardi. Algal zonation in the salt marsh becomes more apparent and more readily understood when studied in relation to the distribution of these phanerogams. As stressed by Lewis (1964) and suggested earlier by the Stephensons (1949), a discussion of zonation along any shore should be based upon biological criteria, rather than relying strictly upon tidal limits. These limits may not be coincident with natural biological zones, especially when one considers the upper limits of vegetation in relation to tide levels. On shores exposed to wave action and spray, the distribution of those organisms which typically form the uppermost biological zones will be extended even higher than where a splash and spray zone are not present (c.f., Lewis, 1964, fig. 52). In the salt marsh environment at Ipswich, one encounters an approximation of a spray zone only near Station 1 (Webber & Wilce, 1971, p. 265) ; here, the presence of wood pilings provides a steep vertical face which contrasts with the gently sloping character of the surrounding marsh surfaces. This change in topography results in a stronger local wave splash, as compared with wave action on the marsh surfaces; thus, the upper dis- tributional limit of Calothrix crustacea, for example, oc- curs at a higher level on these pilings than on the surround- ing marsh. Therefore, using essentially biological criteria in relation to tide levels we recognize in the marsh environ- ment a sublittoral, a littoral, and a less well defined, but nonetheless recognizable, supralittoral zone. The sublittoral zone of the Ipswich salt marsh, when compared with the same zone characteristic of the open coast, is essentially non-existent. The marsh drains twice daily through a series of canals which, at low tide level, are mostly devoid of water. Where the marsh is closest to the sea and in the major trunk canals, water with salinity characteristics of open coastal water remains at low tide. It is in these salt marsh areas, the openings to the sea and 1972] Benthic Algae — Webber and Wilce ATT the major canals, where a true sublittoral zone can be rec- ognized, and, correspondingly, a sublittora] flora. We stress the fully marine character of the sublittoral benthic vegetation. Also, in our list of species from the Ipswich salt marsh (Webber & Wilce, 1971), we cite a num- ber of taxa commonly represented in the marsh sublittoral as drifting plants, e.g., Chaetomorpha melagonium and Phycodrys rubens, to name but two; these plants, of vari- able longevity in the marsh environment, are transients from the open coast sublittoral. Their frequent appearance in the marsh sublittoral in conjunction with the attached marsh sublittoral vegetation is a manifestation both of the marine character of the marsh subtidal zone and the prox- imity of the marsh entrance to the open coast. The littoral zone at the Ipswich marsh contains those crganisms which are regularly submerged and exposed by the rise and fall of the tides. Biologically, this zone ex- tends from the upper limit of the sublittoral, as indicated by Fucus distichus ssp. evanescens, to that uppermost por- tion on the shore populated by Juncus gerardi. Algae of the littoral zone show a zoned pattern of distribution as con- trolled by local factors of, e.g., habitat competition, avail- ability of substrates, and topographic differences resulting in degree of exposure to insolation, desiccation, tidal action, and fresh water influence. These algal assemblages of pre- dictable major constituents we interpret as associations, folowing Borgesen, 1905, Davis, 1913, Feldmann, 1951, Wi!ce, 1959, den Hartog, 1959 and Jorde Klavestad, 1963. However, as Chapman (1956) points out, one characteristic of salt marsh vegetation is its relative instability; thus, the term “associes” is a more apt designation of the vegeta- tional status in these specific environments. The supralittoral zone is characterized by those species which are predominantly terrestrial and which are not merely extensions of upper littoral marine species. The or- ganisms which are regularly submerged and exposed by the tide are in fact, considered “maritime” (Lewis, 1964) rather than strictly marine. While a comparatively clear-cut dis- 478 Rhodora [Vol. 74 tinction between those plants and animals of the upper lit- toral and those of the supralittoral may be possible on a rocky coast, this distinction is not always apparent at the Ipswich salt marsh. For example, Calothrix crustacea clear- ly occurs on the marsh surface (Station 1) in the upper lit- toral zone. In addition, this species along with the upper littoral green alga Pseudendoclonium submarinum, colonize wood pilings near Station 1. Owing to moderate wave splash on these pilings, both algal species extend vertically higher than the level of the surrounding marsh surface. Yet, at Ipswich, the position of C. crustacea and P. sub- marinum on the pilings near Station 1 is comparable to the position on the marsh surface populated by such terrestrial or “maritime” plants as Juncus balticus var. littoralis, Pani- cum virgatum, and Solidago sempervirens. Indeed, on the pilings with Calothrix and Pseudendoclonium one always finds the lichen Lecanora chlarotera which, according to Hale (personal communication), is not marine in its dis- tribution, but rather is a common terrestrial species. Thus, the vegetation at this level is dominantly maritime-terres- trial, and does not represent solely an upward extension of the marine flora. We recognize, therefore, a supra-littoral zone biologically distinct in its composition of vascular plants and associated cryptogams which lies invariably im- mediately above the uppermost limit of salt marsh phanero- gams, i.e., above Juncus gerardi. ZONATION AND DISTRIBUTION OF ALGAL SPECIES Before describing the species composition at each sta- tion, it is appropriate to identify and describe briefly the stations from which the algal collections and related data were taken (see also Webber & Wilce, 1971). Station 1: typically marine, little influenced by fresh water runoff; shoreline a vertical mud bank with substra- tum consisting of mud, small stones, and the Spartina grasses; seasonal salinity range of 18-30°/o0. Station 2: Similar to Sta. 1, but with a gently sloping 1972] Benthic Algae — Webber and Wilce 479 shoreline; substrates predominantly cobble, shells of Midio- lus demissa and Spartina grasses. Station 3: Tidal ditch outflow, 0.3-1 m deep, seasonally influenced by fresh water runoff; substrates of small stones and wood pilings; seasonal salinity range 3-33? /oo. Station 4: Section of a major tidal zreek beneath a highway bridge panning Fox Creek, moderately influenced by fresh water runoff; substrate mostly of stone and shell; seasonal salinity range 13-329/00. Station 5: The innermost extremity of the Ipswich salt marsh, consisting of two substations due to the habitat character in the tidal creek in this region of the marsh; seasonal salinity range for both substations 0-27°/00. Substation 5a: A transect through the tidal creek near Northgate Rd. where at high water the bottom is cov- ered to a depth of 0.6 m, and mostly exposed at low water; markedly influenced by fresh water runoff ; sub- strates of small stones, mud, and plants of Ruppia maritima; this area is characterized also by a twice daily rapid flow of water. Substation 5b: Differs from the latter by having a mud bottom, continual water cover at a 0.6 m depth, and a generally slow water displacement downstream. Sublittoral zone: Stations 1 and 2. Species common to the sublittoral of these stations were not unlike those one might encounter in the sublittoral on a rocky coast. These algae included: Enteromorpha intestinalis Enteromorpha flexuosa Enteromorpha linza ssp. pilifera Enteromorpha linza Kornmannia leptoderma var. oblanceolata Ulva gigantea Enteromorpha ahleriana Chaetomorpha linum Enteromorpha clathrata Ralfsia verrucosa Enteromorpha flexuosa Ralfsia clavata ssp. flexuosa Scytosiphon lomentaria 480 Rhodora [Vol. 74 Petalonia fascia Chorda filum Laminaria saccharina Dumontia incrassata Hildenbrandia prototypus Agardhiella tenera Gracilaria foliifera Gracilaria verrucosa Chondrus crispus Ce:amium diaphanum Ceramium rubrum Ceramium rubriforme Polysiphonia denudata Polysiphonia lanosa Polysiphonia nigra Polysiphonia nigrescens The majority of these species were of late spring and summer occurrence, while the following attained maxi- mum vegetative and reproductive development during the winter and early spring months: Enteromorpha linza var. oblanceolata Kornmannia leptoderma Ralfsia clavata Scytosiphon lomentaria Petalonia fascia Laminaria saccharina Dumontia incrassata Ralfsia verrucosa Conspicuous perennials at the sublittoral-littoral inter- face were Fucus distichus ssp. evanescens, Fucus vesiculo- sus and Chondrus crispus. Station 3: There was a decrease in numbers of sublit- toral species at Station 3 as compared with Stations 1 and 2. This reduction in taxa is due likely to the comparative lack of rocky substrate, the pronounced fresh water in- fluence in the winter and spring months, and the shallow water at this station. Those algae common to the sublit- toral of Station 3 throughout the summer were: Enteromorpha intestinalis Enteromorpha flexuosa ssp. pilifera Ulva rigida Cladophora liniformis Bryopsis plumosa Ceramium diaphanum Ceramium fastigiatum Polysiphonia denudata The winter and early spring algal vegetation was dom- inated by four species: Kornmannia leptoderma Scytosiphon lomentaria Petalonia fascia Melosira juergensii 1972] Benthic Algae — Webber and Wilce 481 The only sublittoral perennials in this station were Ralf- sia clavata, Ralfsia verrucosa, and Chondrus crispus. Station 4. The submerged algal vegetation here was scant. From November through March Kornmannia lepto- derma, Pentalonia fascia, and Dumontia incrassata com- prised the macroscopic algae at this station in the sublit- toral zone. These taxa were continually vegetative and greatly reduced in size, as compared with their larger and normally reproductive counterparts at Stations 1-3. The sublittoral perennials were the same as those at Station 3, i.e., Ralfsia clavata, Ralfsia verrucosa, and Chon- drus crispus. Station 5,a and b. The vegetation in the sublittoral here had the smallest number of algal species of any station. We have considered this to be a reflection of the reduced salinities occurring here through most of the year. During the spring and autumn, the following species dominated: Capsosiphon fulvescens Ectocarpus confervoides Enteromorpha intestinalis var. dasycarpa Polysiphonia urceolata In early spring only the stalked diatom, Gomphonema olivaceum, constituted the macroscopic algal vegetation, covering the numerous small stones in this tidal creek. Owing to a thick ice cover, samples of winter algae were not obtained from this station. Littora! Zone Stations 1 and 2. Fucus vesiculosus and Ascophyllum nodosum occurred in the littoral zone wherever stone, rock, or other solid substrate was present. The landward extent of these species differed, however, between Stations 1 and 2. For example, at Station 1 the marsh surface is essen- tially 1 meter higher above high water neap tide levels than is that of Station 2, owing to the presence of a ver- tical mud bank at the former station. Fucus vesiculosus became established on this bank, the plants extending about 3/4 of the height of this vertical surface. At this 482 Rhodora [Vol. 74 uppermost level, Fucus thalli tended toward a spiral growth habit, and they were smaller than those plants in the mid- littoral. By contrast, the lower level and more sloping top- ography of Station 2 resulted in both Fucus and Ascophyl- lum extending onto the marsh surface, where they devel- oped as typical marsh fucoids, i.e, F. vesiculosus var. spiralis and A. nodosum f. scorpioides, both at the bases of Spartina alterniflora var. glabra. Immediately beneath the seaward marsh edge, a dark green algal zone, the Enteromorpha associes (Fig. 1), was conspicuous on the mud at both stations. The summer and autumn algae comprising this associes were: Enteromorpha ahlneriana Cladophora sericea Enteromorpha flexuosa Rhizoclonium riparium ssp. paradoxa f. riparium Enteromorpha prolifera f. validum Percursaria percursa Throughout the winter and spring months this Entero- morpha associes assumed a pronounced brown hue owing to increased numbers of Pylaiella littoralis, with lesser amounts of Ectocarpus confervoides var. arcta. Associated with these plants at this time were irregularly distributed clusters of: Enteromorpha flexuosa Percursaria percursa ssp. paradoxa Vaucheria intermedia Monostroma oxyspermum Covering the mud in the Spartinetum alternifloretum at Station 1 were conspicuous green tufts of Rhizoclonium riparium (f. riparium and f. polyrhizum), Rhizoclonium implexum, and Vaucheria intermedia. While these algae were similarly located at Station 2, their distribution at the latter station was limited to the more seaward edge of the Spartinetum. We attribute this local restriction in vertical distribution to competition for habitat; marsh fucoids and the ribbed mussel, Modiolus demissa, thoroughly blanketed much of the available mud substrate beneath Spartina alterniflora var. glabra, thus Benthic Algae — Webber and Wilce 483 1972] D aq Ssnasiu) Wü1V9HIA SNYONOHI WNIJINYd $4 108vu29 snonne dd d — snnvarwawy snauias — = SN31Vd VNILYWdS 4 34 VH8V19 'HVA ad saloons HSUVW nani SIIIOSSY VHdHOWOH3.1N3 SN32S3NVA3 'dsS U SNHIILSIA SNINA WNSOGON Wn11AHd02SV SNSOINIISIA Y sn3n4 VNIHVH22VS VIBVNIWVT UJA X23N 311sv5 SNOTV 3Nl'380H$ Jo 1N3N93$ S"3413W S5! 303131 tI A— 54343H Sl ^ 484 Rhodora [Vol. 74 prohibiting any further upshore spread of Rhizoclonium riparium, R. implexum, and Vaucheria intermediwm. A variety of filamentous bluegreens was collected throughout the year from the mud of this Spartinetum at both Sta- tions 1 and 2 (Webber, 1967). Moving landward, the next algal associes cecurred be- neath Spartina patens and consisted almost entirely of Vaucheria arcassonensis (Webber, 1968). Growth of this alga, with lesser amounts of V. intermedia, was particularly evident during the summer and autumn months, forming dense, felt-like mats on the soil. Numerous filamentous bluegreens also were common at this time on the sandy peat soil of the Spartinetum patentis. Old culms of Spartina patens were epiphytized through- out the year mainly by Calothrix confervicola, as tufts to 3 mm tall Often entangled with C. confervicola were plants of Microcoleus tenerrimus ; the latter species never occurred alone as a dominant. Blum (1968), working on several Cape Cod (Massachusetts) marshes, concluded that a species of Calothrix (likely C. confervicola) is the most important colonizer in the moist micro-habitats on dead S. patens culms. He also demonstrated a marked interde- pendence between specific salt marsh grasses and their un- derlying algal species. Blum further elucidated the rela- tionship between the mature form of the graminoid zone and such environmental parameters as light penetration and intensity in the zone, filtration and detritus retention, and drainage, as all these events are influenced by the morphology of a specific stand of grass. The brownish-green tubular thalli of Capsosiphon fulves- cens were present also in the Ipswich Spartinetum paten- tis. While mocroscopic individuals of this species were epilithic in the sublittoral at Substation 5a (Webber & Wilce, 1971, p. 269), the macroscopic thalli of Capsosiphon (the type commonly encountered elsewhere) grew only in a single, small, water-retaining marsh depression at Sía- tion 1. The discovery of two populations of Capsosiphon, distinctly different in their morphologies, seasonality, and 1972] Benthic Algae — Webber and Wilce 485. habitat requirements, suggests further questions as to the character of the species and the role of the environment in determining the expression of that character. Labora- tory and field experiments designed to elucidate this mat- ter are forthcoming. The highest band of salt marsh phanerogams at Stations ] and 2 was the Juncus gerardi association; the bases of Juncus and its associated algal species were wetted by sea water only during periods of spring tides. The algal veg- etation on the sandy soil here was predominantly Tolypoth- rix tenuis and Rhizoclonium riparium f. validum. On patches of soil not colonized by Tolypothrix and Rhizoclonium, Calothrix crustacea formed brownish-blue masses through- out the year, being especially abundant during the autumn, winter, and spring months. Station 3. The littoral zone of Station 3 consisted of portions of the marsh surface and several wood pilings lo- cated in the outflow of the tidal ditch at this station. Bright green patches of Pseudendoclonium submarinum were always apparent, occurring essentially in the top 20 cm of these pilings throughout the year. Monostroma oxy- spermum occurred just beneath P. submarinum on the pil- ings’ surfaces; this species first appeared in September as small foliose clusters, and developed abundantly through- out the winter until late April, after which it was no longer apparent at this station. The lowermost extent of M. oxy- spermum coincided with low water neap tide levels. Ulo- thrix subflaccida, mixed with Monostroma, was first evident in December, and most common during the spring months. Similar to M. oxyspermum, U. subflaccida was absent dur- ing the summer (July through September). The algal vegetation of the marsh littoral at Station 3 consisted of few species; these were Vaucheria intermedia, Rhizoclonium implexum, Percursaria percursa, and a mix- ture of filamentous bluegreens. It was quite similar in its species composition to the associes of comparable zonation at Stations 1 and 2, being essentially a telescoped version of the same zone at these stations. 486 Rhodora [Vol. 74 Station 4. Wooden bridge supports over Fox Creek and the small stones and larger rocks in the creek bed imme- diately beneath the bridge were substrates in the littoral zone of this station. A scant algal vegetation characterized the littoral zone here, with only Pseudendoclonium sub- marinum and a few filaments of Calothrix crustacea appar- ent on the bridge supports through the year. Ralfsia clav- ata and R. verrucosa were yearly inhabitants also, as epi- lithic crusts in the most damp and most shaded areas of the creek bed along with large quantities of the barnacle, Balanus balanoides. The creek bed and that portion of the littoral zone with Pseudendoclonium and Calothrix could also be characterized as the Balanus zone. Balanus was present on virtually all solid substratum at this station, and was the dominant competitor for substrate coloniza- tion. Perhaps this is the reason for so few species of at- tached algae here. In other upper shore locations one usually finds numerous algal species in the Balanus zone. While the obvious factors contributing to this associes of so few species appear to be competition for substrate, ex- posure to insolation and to desiccation, the specific ecologi- eal factors responsible for this causal relationship are yet to be discovered. Station 5, a and b. The littoral zone of Station 5 con- sisted of muddy tidal creek banks which contained an algal flora throughout the spring, summer, and autumn months of Vaucheria compacta var. koksoakensis mixed with fila- mentous bluegreens. Owing to an uneveness in the creek bed coupled with an irregular accumulation of variously sized stones, a segment of Substation 5a drained completely at low tide. Thus, a small and well-defined littoral region was present in the bed itself. Small stones here were covered with attached plants of Monostroma oxyspermum from September through December. This species also was encountered at Stations 1-3 from late October through the winter, per- sisting until June. Conover (1958) reported a similar au- tumn-winter (October to January) reoccurrence of M. 1972] Benthic Algae — Webber and Wilce 487 oxyspermum in addition to its spring (March to June) growth in Great Pond Estuary, Massachusetts. His data shows comparable spring and autumn light intensities, and he suggested this factor as responsible for his observed pattern of seasonal distribution. While Conover’s hypothe- sis might well be involved in an explanation of the pres- ence of M. oxyspermum in Substation 5a at Ipswich, our field data indicate that salinity levels may also affect the seasonal appearance of these plants. Salinities at Substa- tion 5a during the September to December period ranged from 25-27?/oo, a variation similar to that, 24-309/oo, measured at Stations 1-3 during the period of maximum spring growth of Monostroma oxyspermum. Thus, the seasonal patterns of occurrence of this species at Ipswich coincide with low light intensities and high salinity levels. Winter collections were not possible at Station 5 be- cause of a thick ice and snow cover. Supralittoral zone. This zone was recognized only in conjunction with the wood pilings near Station 1. The algal vegetation which occurred in and on these pilings was uniform in species composition throughout the year, and consisted of: Pseudendoclonium Urococcus foslieanus submarinum Branched filamentous Chry- Calothrix crustacea sophyte undescribed Schizothrix calcicola SUMMARY COMMENTS In this paper we have attempted to summarize our de- scriptive ecological data in an overview interpretation of the ecology of the salt marsh algae at Ipswich. The algal taxa are characterized relative to their horizontal and ver- tical zonation on the marsh, and the seasonality of the dominant species is described. LITERATURE CITED BLUM, J. L. 1968. Salt marsh Spartinas and associated algae. Ecol. Monogr. 38: 199-221. BQRGESEN, F. 1905. The algal-vegetation of the Faeróes. coasts, 488 Rhodora [Vol. 74 with remarks on the phytogeography. In: Warming, Botany of the Faróes. 2: 679-845, pls. 18-24. CHAPMAN, V. J. 1956. The marine algae of New Zealand. J. Linn. Soc. Lond. (Bot.) 55: 333-501. Conover, J. T. 1958. Seasonal growth of benthic marine plants as related to environmental factors in an estuary. Inst, Mar. Sci, Univ. of Texas. Port Aransas. 5: 97-197. Davis, B. M. 1913. General characteristics of the algal vegetation of Buzzards Bay and Vineyard Sound in the vicinity of Woods Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fish. 31: 443- 544. (1911). DEN HarTOG, C. 1959. The epilithic algal communities occurring along the coast of the Netherlands. Wentia. I: 1-241. FELDMANN, J. 1951. Ecology of marine algae. 313-333. In: G. M. Smith (ed.), Manual of Phycology. Chronica Botanica, Wal- tham, Mass. HALE, M. (Personal communication. Smithsonian Institute. Wash- ington, D.C.). JogpE, L, AND N. KravEsTAD. 1963. The natural history of the Hardangerfjord. 4. The benthonic algal vegetation. Sarsia 9: 1-99. Lewis, J. R. 1964. The Ecology of Rocky Shores. The English Universities Press, Ltd. London. 323 pp. OosTING, H. J. 1956. The Study of Plant Communities. 2nd ed. W. H. Freeman Co. San Francisco. 440 pp. STEPHENSON, T. A., and A. STEPHENSON. 1949. The universal fea- tures of zonation between tide marks on rocky coasts. J. Ecol. 37: 289-305. Weaver, J. E., and F. E. CLEMENTS. 1938. Plant Ecology. 2nd ed. McGraw-Hill Book Co., Inc. N. Y. 601 pp. WEBBER, E. E. 1967. Bluegreen algae from a Massachusetts salt marsh. Bull. Tor. Bot. Club, 94: 99-106. 1968. New England salt marsh Vaucherieae. Rho- dora. 70: 274-277. , AND R. T. Witce. 1971. Benthic salt marsh algae at Ipswich, Massachusetts. Ibid. 73: 262-291. WincE, R. T. 1959. The marine algae of the Labrador Peninsula. and northwest Newfoundland (ecology and distribution). Bull.. Nat. Mus. Can. 158, iv + 103. DEPT. OF BIOLOGY KEUKA COLLEGE KEUKA PARK, NEW YORK 14478 and DEPT. OF BOTANY UNIVERSITY OF MASSACHUSETTS AMHERST, MASSACHUSETTS 01002 LUPINUS MEXICANUS CERV. EX. LAG. Davip B. DUNN Since Lupinus mexicanus Cerv. ex. Lag. (1816) was the first taxon in the genus Lupinus named for Mexico it is imperative to determine which lupine the name was applied to. As the earliest name it would take priority over all the later names. In all of Charles Piper Smith’s works on the lupines of Mexico he literally ignored the name. In my own studies it has been a perplexing problem for quite a number of years. The curator of the herbarium at Madrid, Spain, searched, but was unable to locate a speci- men of Lupinus mexicanus of any vintage. Since seeds were sent to England by Lagasca, and the plants grown were utilized to prepare a much more detailed and accurate description for Edwards Botanical Register (no. 457, in 1820) than that originally provided by Lagasca, Dr. Walters at Cambridge, England, also searched the herbarium there but was unable to find a specimen of the material used in either of the original descriptions provided by Lagasca and Edwards. In 1832, Maund was attempting to identify Lupinus mexicanus and stated that the species had been lost to science, but that he thought that it had been re- gained. He supplied an illustration as had Edwards. Edwards suggested that L. mexicanus was probably a biennial and that it had flowered in the “stove” (glass- house) in February, which suggests that they had planted the seeds the previous season, but flowering had not oc- curred until February. Maund’s plant material is indicated as a perennial, which was probably another taxon, and nothing is given which helps identify L. mexicanus. The curators of the herbaria of Kew, Paris, Berlin, Zurich and others searched but were unable to find any specimens of the period involved. Since there is apparently no type specimen and since the description of Lagasca could apply to several taxa it is necessary to select some element to fix the application of the name until such time as an 489 490 Rhodora [Vol. 74 authentic specimen may be discovered. The illustration in Edwards Botanical Register (no. 457, 1820) serves this purpose and I select it to serve in lieu of a type specimen for Lupinus mexicanus Cerv. ex Lag. It should also be pointed out that there is no accurately labeled material of Lupinus mexicanus in any of the herbaria and the material distributed by Pringle as L. mexicanus has been a major source of error, since he applied the name to, at best, a variant of L. aschenbornii, which is an alpine perennial. During the preparation of the treatment of Lupinus for the ‘Flora de Valle de Mexico” (for Dr. Rzedowski) the lupines of the area have been studied in considerable detail. Several field trips to the area have been conducted and numerous collections made. Among the collections are plants collected along the toll road between Mexico City and Queretaro, the latter, the site of the palace of the Emperor Maximilian. This route is probably not far from the original route the Spaniards traveled, since it is on the most direct route between the two early cities. In the area near Tula, a number of collections of lupines were made, which closely match the detailed description in Edwards Botanical Register. 'The collection made in July by Giles Waines contained both flowering and fruiting specimens. Plants collected along the toll road a few miles northwest of the Tula turnoff by both Harmon and Dunn on three different years in late December were in full fruit, as well as having a few branches in flower. Seeds from both the Waines collection and the Dunn collection were grown in the research greenhouse at the University of Missouri. What had been assumed to be the same taxon turned out to be two different taxa of the same complex — one an annual flowering in the fall and early winter and the second a biennial germinating at the same time as the annual but requiring cold treatment during the winter before flowering the following season. By planting the seeds of the annual in January and the biennial in the early fall of the previous year the two were brought into flower at the same time. Attempts to self-pollinate the biennial all failed, so the 1972] Lupinus — Dunn 491 taxon is obligate outcrossing. Most of the attempts to cross the annual and the biennial failed but a few produced some seeds. Thus there are two distinct breeding populations isolated primarily by their phenology, which remain dis- tinct in nature, but which have not fully achieved inter- sterility, which is not uncommon in plants. After becoming aware of this difference, the colonies along the toll road were examined and found to contain many plants with only caespitose clusters of basal leaves with no flowering shoots, in late December, while the annuals were in fruit with the lateral branches still in flower. The biennial is Lupinus mexicanus and the annual is Lupinus bilineatus Benth. The following translations of the Latin descriptions of both Lagasca and Edwards are given for determining the identity: “288. Lupinus mexicanus Cerv. ex. Lag. Gen. et Sp. Nov. 22, 1816. Calyces alternately bracteolate (appendaged) upper-lip semibifid, lower lip obscurely tridentate. Lupinus mexicanus Cerv., near L. termis L. Leaves ternate at the base, remains of 5-7 leaflets; leaf- lets lanceolate, mucronate, upper surface glabrous, lower surface pilose. Stipules setaceous, pilose as the stems. Peduncles opposite the leaf, spicate-racemose above. Bracts setaceous, deciduous. Flowers alternate (scattered), short stalked, blue. Calyx bracteoles setaceous, short. Legumes pilose. Habitat in New Spain (Mexico). Seeds sent with D. Vincent Cervantes." From Edwards Bot. Reg. 457. 1820, the following trans- lation: (Note, familial and generic portions are not given and interpretations are included in parentheses). “Plants, except for the corolla, entirely shaggy-pilose. Leaflets 3?- 5?, 7-8, elongate-cuneate, narrow, short pointed, tapered for a considerable way down, glabrous above, rendered white beneath by shaggy-pilose pubescence, longest about 2 inches, shorter than the long-piled petioles; stipules linear-subulate, erect, long-piled. Racemes elongate spi- cate, laxly many-flowered, flowers scattered, sparse, la- 492 Rhodora [Vol. 74 vender to purplish-blue, with darker streaks (veins), bracteose before anthesis; peduncles opposite the leaves (true of all lupines that have the top lateral bud elongate as a branch) ; pedicels hirsute, shaggy, ascending, shorter than the calyx, bracts linear-sublate, very narrow, filiform, longer than the calyx, caducous. Calyces shaggy-hirsute, green, with divaricate lips, the upper spilt at the end into a notch. Banners folded, reflexed, astride the upper margins of the wings by a deep sulcus; wings hatchet shaped, pointed, cohering by the front edges; keels pale ascending upward, narrow subulate-falcate, as long as the wings; with a long deep-purple tapered point, — etc.” The distinctive traits from which L. mexicanus can be determined are the shaggy-pilose hairs, the upper surface of the leaflets glabrous, while the lower is shaggy-pilose, and the corolla is glabrous as intimated by Edwards, the long setaceous bracts and short setaceous bracteoles at the lateral sinuses of the calyx, the shaggy-hirsute pedicels, the deep sulcus of the banner, the narrow subulate-falcate keel with the long tapered point, and Edward's term of pointed hatchet-shaped wings. His discussion of flowering time and suggestion that the taxon is a biennial apply. The field location is that of the most direct route between the two early major cities of that time. All of the traits apply to the biennial plant material collected by Giles Waines. The greenhouse plants grown from seed more closely resemble the illustration provided in Edwards, as would be expected, since greenhouse plants were used in preparing the description. The field material, in nature, has denser foliage and a denser raceme, which would represent the effects of high light intensity at high eleva- tions. The specimen collected by Giles Waines Lupin #2 (UMO- 88263) (Fig. 1) is considered as typical material of Lupinus mexicanus Cerv. ex. Lag. A field duplicate has been sent to Madrid (MA) and a duplicate grown in the greenhouse has been sent to Cambridge (CGE) ; others will be distrib- uted as available. The close relatives are Lupinus bilineatus 1972] Lupinus — Dunn 493 id eo 2* Lapin dI ce ld act e fact. ged Asper Quest As i per- os Q4 p i Tum pole [Mesie - Ga wu 1 ds "CA. ea owt e Ale i ke Tase pila en Dek, i / i rdi trad . : Vlas, Dt bee. i fy í i WM L i H due got ead tal Z id Bel he pu ur bola E guy uet zy a riw Plants Figure 1. A field specimen of Lupinus mexicanus Cerv. ex. Lag., a biennial collected in July by Giles Waines, Lupine #2 (deposited at UMO; duplicate sent to MA; a greenhouse grown duplicate sent to CGE). 494 Rhodora [Vol. 74 Benth (1839), L. hartwegii Lindl. (1839), and L. persis- tens Rose (1905). The taxon which has been misinterpreted as Lupinus mexicanus by several authors is Lupinus aschenbornü. While the latter taxon has glabrous flowers and a glabrous upper leaflet surface and pilose hairs, the hairs do not match Edward's term of shaggy nor are the leaflets shaped right nor is the keel slender, long-faleate. The keel is almost straight on the upper edge and the banner is narrow and without the deep sulcus. In addition L. aschenborni is alpine and it is a perennial, which goes dormant every winter. It is a most unlikely candidate for a greenhouse plant or a plant that could be grown outside in Madrid, Spain, at the Botanical Garden. In addition the flower size and shape are entirely wrong for the material illustrated by Edwards from seeds sent by Lagasca. DIVISION OF BIOLOGICAL SCIENCES UNIVERSTY OF MISSOURI-COLUMBIA COLUMBIA, MISSOURI 65201 NOMENCLATURAL AND TAXONOMIC NOTES ON MEXICAN COMPOSITAE ROGERS McVAUGH The following notes have accumulated during the prepa- ration of a treatment of the family Compositae for a pro- posed Flora Novo-Galiciana. Included are one new com- bination, discussions of points of nomenclature, mention of notable range-extensions, and comments on taxonomic decisions, all of which are somewhat out of place in a formal floristic treatment. The area covered by the pro- posed flora includes the Mexican States of Jalisco, Colima, and Aguascalientes, and some adjoining territory; for a fuller description of it see Brittonia 13: 145-147. 1961, Contr. Univ. Mich. Herb. 9: 1-7. 1966, or Contr. Univ. Mich. Herb. 9: 207-357. 1972. Another paper preliminary to the Flora Novo-Galiciana, including descriptions of a number of taxa of Compositae new to science, appeared in Contr. Univ. Mich. Herb. 9: 359-484. 1972. For general support of the field-work and other activities on which this paper is based, I am grateful to the National Science Foundation (Grant no. GB-5218X). In the course of revision of the various groups of Com- positae for the Flora, I have seen most of the relevant types (or in some instances isotypes). When I have seen and studied a specimen, this is indicated in the text below in the conventional way [!]. To the many persons and insti- tutions that have permitted and assisted my studies of the valuable collections in their charge, my sincere thanks. Many of the remarks on nomenclature refer to the names published by Kunth in the fourth volume of Nova Genera et Species Plantarum (1818), or those published by De- Candolle in the fifth and sixth volumes of his Prodromus (1836, 1838). To the authorities at Paris, where I have studied most of the types of Humboldt & Bonpland, and at Geneva, where similarly I have seen and studied the types of DeCandolle, I am most grateful. 495 496 Rhodora [Vol. 74 It is necessary to comment on the dates of publication of the fourth volume of the Nova Genera et Species, which dealt wholly with the Compositae. Folio and quarto editions of this were published simultaneously (as they were for the other 6 volumes of the set). Volume 4 appeared in 1820, in 5 installments, on 17 Apr, 22 Mai, 31 Jul, 18 Sep and 26 Dec, respectively. Apparently the quarto edition was not available to the public before these dates, nor was the folio edition generally distributed. The printing of the folio text was completed, however, in September 1818, and at least 4 copies were distributed and became more or less accessible to the botanical public by 1 December 1818. The authors presented a copy to the Institut in Paris on 26 October 1818. Under the code of nomenclature now in force, this constitutes effective publication as of that date, of all the new names proposed in the volume. The fact that the authors considered the work unpublished until it was generally distributed in 1820, is irrelevant. In citations of names published in the Nova Genera. et Species, botanical custom has been to cite the page-numbers of the quarto edition, as this has been more generally available. Some authors have cited the pages of both quarto and folio edi- tions, on the assumption that the two were published simul- taneously. Names of Compositae, however, are correctly cited as from the folio edition only, because publication of the quarto pages was delayed for a year and a half to two years. For summary of the dates of publieation of both editions and the cireumstances surrounding the distribu- tion of volume 4, see Stafleu, F., Taxonomic Literature (Regnum Veg. 52: 225-226. 1967). Many of the circum- stances were first made publie by Henri Cassini, who was given a copy of the folio text by Kunth, and who criticized it in his own published articles before the dates of general distribution in 1820. Cassini's remarks, originally pub- lished in the Journal de Physique, de Chimie, d'Histoire Naturalle et des Arts, early in 1819, were reprinted in his Opuscules Phytologiques, vol. 1, pp. 324, 339, etc. 1826. 1972] Mexican Compositae — McVaugh 497 Ageratella microphylla (Sch. Bip.) A. Gray in S. Wats. Proc. Am. Acad. 22: 419. 1887. Ageratum microphyllum Sch. Bip. in Seem. Bot. Voy. Her- ald 298. 1856. Ageratella microphylla var. seemannii and var. palmeri A. Gray in S. Wats. Proc. Am. Acad. 22: 419. 1887. Ageratella palmeri (A. Gray) Rob. Proc. Am. Acad. 41: 272. 1905. The type of A. microphylla (from the “Sierra Madre", i.e. perhaps from Cerro del Pinal, Sinaloa, Seemann 2043!, at K) was taken from a plant with unusually broad leaves, as well shown in the illustration published by Hemsley (Biol. Centr. Am. Bot. 5: pl. 42. 1881). The specimen evidently represents a portion of a plant in which several long leafy branches arose from a larger stem. As is often the case in the Compositae, the leaves are opposite at the lower nodes of such branches, but sometimes alternate above. Similarly abnormally branched specimens, of other- wise typical “palmerv’, have the same leaf-arrangement but with fewer opposite leaves (e.g. Gaiser 62, at MICH, from near Guadalajara). Nothing exactly like the type of microphylla has been found by any subsequent collector, and we can but surmise what it may represent, until it can be found again in the field. Gray (1887) distinguished what he called two varieties on the basis of the broad, incised-dentate leaves and subspicate inflorescence of var. seemannii (i.e. typical microphylla), and the narrow, subentire leaves and looser inflorescence of var. palmeri. Robinson (Proc. Am. Acad. 41: 271-272. 1905), and Blake, in the Trees and Shrubs of Mexico, noted these features, and stated also that the leaves were alternate in palmeri but opposite in microphylla. Both these authors treated the two taxa as distinct species. Blake's key is based entirely on leaf-characters. In fact it seems that neither position nor shape of leaves is completely diagnostic. In the original specimens of var. palmeri (Palmer 587!, at GH, from near Guadalajara), and in most 498 Rhodora [Vol. 74 flowering specimens, the narrow leaves of sterile axillary shoots are much in evidence, giving the impression of a narrow-leaved plant. There is much individual variation in this respect, but the main cauline leaves subtending axillary shoots are seldom very narrow, and often much toothed or lobed. The differences between “spikelike” and “loose” inflores- cences also appear to be related to individual differences in branching. When vigorous lateral leafy branches (e.g. those 10-30 cm long) produce a number of short-peduncled axillary clusters of flower-heads, the effect is of a spike; when (as in most specimens), the lateral branches are reduced in length, and in number of nodes, and bear few or no heads except small clusters at the tips, the effect is of a small loose panicle. Various intermediates can be demonstrated. Archibaccharis hieraciifolia Heering, Jahrb. Hamburg. Wiss. Anst. 21, Beih. 3: 40. 1904. Baccharis hieraciifolia Hemsl. Biol. Centr. Am. Bot. 2: 129. 1881, not B. hieracifolia Lam., 1783. Hemibaccharis hieraciodes Blake, Contr. U.S. Nat. Herb. 20: 547. 1924. Archibaccharis hieracioides (Blake) Blake, Jour. Wash- ington Acad. Sci. 17: 60. 1927. Blake (1927) discussed the nomenclature of this species and that of Archibaccharis hirtella. Both were assigned names by Heering in 1904, but both were called ''n.spec." in spite of the fact that both names were derived from previously published names. As Blake said, “it seems ad- visable to treat these two names of Heering as representing new combinations". Under the International Code of Botanical Nomenclature (Art. 72) a new combination derived from a previously published but illegitimate name (e.g. Archibaccharis hieraciifolia from Baccharis hieracit- folia Hemsl.) may be treated as a new name. It is there- fore proper to use the name originally proposed by Heering, not Archibaccharis hieracioides, the substitute proposed by Blake. 1972] Mexican Compositae — McVaugh 499 Baccharis sulcata DC. in DC. Prodr. 5: 419. 1836. Baccharis potosina A. Gray, Proc. Am. Acad. 15: 33. 1879. Blake (in the Trees and Shrubs of Mexico, 1926) treated Baccharis sulcata as a synonym of B. thesioides H.B.K., but the type of B. sulcata (Villalpando, Méndez! in G-DC) certainly represents another species. The narrow leaves (up to 3-3.5 mm wide) vary from quite entire to toothed, with up to 4 or rarely 6 short teeth per cm of margin. In other respects, including the characters of heads and in- florescence, and the habit, the plants are typical of what has been called B. potosina (San Luis Potosí, Parry & Palmer 410! at GH, the type). Bidens acrifolia Sherff, Bot. Gaz. 94: 591. 1933. Bidens polyglossa Sherff, Brittonia 16: 61. 1964. Southern Sinaloa (Concordia, Dehesa 1532!, K, the type), and western Jalisco (northwest of Cuautla, McVaugh 13633!, MICH, type of B. polyglossa). The type of Bidens acrifolia is an immature and incom- plete specimen, but in all observable details it agrees pre- cisely with the abundant available material of B. poly- glossa. The apparent differences mentioned by Sherff (Brittonia 16: 62. 1964) are insignificant or (those per- taining to the leaf-pubescence and to the phyllaries) non- existent. Chaptalia runcinata H.B.K. Nov. Gen. & Sp. 4 [ed. fol.]: 5. pl. 303. 1818. Sonora (Pennell 19647), western Durango (Maysilles 7452, 7809, 7848; Cronquist 9560), northern Nayarit (Rose 2022, at US). Costa Rica; Venezuela and Colombia (Hum- boldt & Bonpland!, at P, the type) ; Bolivia to southeastern Brazil and northern Argentina. Not previously reported from north of Costa Rica, but abundant on the summer- wet, high pine plains of western Durango (cited specimens from Sonora and Durango all at MICH). The seasonal forms with reduced or filiform marginal flowers, reported by Burkart for South American representatives of this species, have not been found in Mexico, but otherwise the 500 Rhodora [Vol. 74 Mexican material agrees perfectly with that from further south. Conyza viscosa Mill. Gard. Dict. ed. 8. Conyza no. 8. 1768. Conyza lyrata H.B.K. Nov. Gen. & Sp. 4 [ed. fol.] : 55. 1818. The type, from Veracruz (Houstoun!, BM), is a pale- pilose plant resembling what has been called Conyza lyrata var. pilosa Fernald, Proc. Am. Acad. 36: 506. 1901 (Chia- pas, Seler 1879!, GH, the type), whereas the type of C. lyrata H.B.K. (Guayaquil, Hwmboldt & Bonpland!, P) is much less hairy, with more conspicuous glands. Erigeron longipes DC. in DC. Prodr. 5: 285. 1836. Erigeron scaposum DC. in DC. Prodr. 5: 287. 1836. Erigeron scaposum p latifolium DC. in DC. Prodr. 5: 287. 1836. Erigeron affine DC. in DC. Prodr. 5: 289. 1836. From Coahuila and perhaps Durango southward nearly throughout southern Mexico to Central America, variable in habit, originally described as subscapose. The leaves, especially in some parts of eastern Mexico, may be grouped toward the base of the stem, whereas in Nueva Galicia the leafy part of the stem is usually elongated and the leaves well spaced along it. I have not seen the type (Karvinskt, s.n., at M), which is from some unknown locality in Mexico, but judging from fragments in DeCandolle's herbarium (G-DC!), it represents the same species as the other names cited above. Subscapose forms seem to be especially abundant in Oaxaca, and it may be that Karvinski's speci- mens came for that state. Hrigeron scaposum and E. longipes were described by DeCandolle as perennials, whereas E. affine, the type of which was a plant with elongated leafy stems, was erroneously described as an annual. The type of E. affine, from near the City of Mexico (Berlandier 522!, G-DC), almost certainly represents a perennial species. The basal parts of the plant are much like those of E. scaposum, and in fact the two supposed species are essentially similar except that in E. affine the hairs are longer on both leaf-surfaces. The type of E. sca- 1972] Mexican Compositae — McVaugh 501 posum came from near Toluca (Andrieux 277!, G-DC), and that of B latifolium from near Mexico City (Berlandier 375!, G-DC). Erigeron velutipes Hook. & Arn. Bot. Beech. Voy. 434. 1841. Erigeron alamosanum Rose, Contr. U.S. Nat. Herb. 1: 102. 1891. Western Mexico, from southern Sonora (Palmer 348!, vs, the type of E. alamosanum) to Chihuahua, Sinaloa, southern Zacatecas, Nayarit, Jalisco, and Michoacán. The type (Sin- clair s.n.!, at K) was collected between San Blas and Tepic. Plants of E. velutipes have often been misidentified with Erigeron tenellum DC. in DC. Prodr. 5: 288. 1836, the type of which came from Matamoros, Tamaulipas. (Ber- landier 2129! in G-pc, lectotype). E. tenellum, like E. velu- tipes, is a slender branched annual, but the herbage is eglandular and the base of the plant is only moderately stiff- hairy. The name E. alamosanum was based on specimens a little larger than average for the species. Similarly vigorous plants are not infrequent along the Pacific Slope from Sonora to Nayarit. Eupatorium albicaule Sch. Bip. ex Klatt, Leopoldina 20: 89. 1884. Eupatorium albicaule var. laxius Rob. Proc. Am. Acad. 35: 330. 1900. Eupatorium leucoderme Rob. Proc. Am. Acad. 41: 274. 1905. Eupatorium ymalense Rob. Contr. Gray Herb. II. 75: 14. 1925. Lowlands, sea-level to 300 m. in elevation, Sinaloa (Ymala, Palmer 1474!, GH, type of var. laxius and of E. ymalense), Nayarit, Jalisco, Colima, Michoacán (Chuta, Langlassé 183!, GH, type of E. leucoderme) ; Tabasco and Chiapas; Yucatán Peninsula; north to Veracruz (Papantla, Liebmann 88!, P, an isotype), San Luis Potosí, and Tamau- lipas. This geographical range is like that of many other species of tropical Mexico; see the maps in Arboles Tropi- cales de México, by Pennington & Sarukhan (published by 502 Rhodora [Vol. 74 the Instituto Nacional de Investigaciones Forestales, and the FAO, pp. vii, 413. México, 1968). Plants growing in western Mexico were distinguished from the original Eupatorium albicaule, first as the var. laxius and later as an independent species, E. ymalense. The only significant difference between the two populations seems to lie in the shape of the phyllaries, which are acute or attenuate in the plant of the Pacific lowlands, and usu- ally, but not always, blunt or subacute in the plant of eastern Mexico and the Yucatán Peninsula. Robinson re- ported the heads in E. albicaule as *about 7 or 8-flowered", but they are usually 10-13-flowered, as in the supposed E. ymalense. Eupatorium collinum DC. in DC. Prodr. 5: 164. 1836, var. collinum. Eupatorium stillingiaefolium DC. in DC. Prodr. 5: 160. 1836. This variety appears to be restricted to eastern Mexico. The type (Berlandier 2162! in G-DC), from near Tantoyuca in the Atlantic lowlands of Veracruz, apparently does not differ significantly from the type of E. stillingiaefolium (Berlandier 2142! in G-DC), which was collected in Tamau- lipas. In these plants, and in modern specimens from the same region, the phyllaries are narrow and attenuate or acute (the middle ones 1 mm wide), often nearly all equal, sparingly ciliate, evidently resinous-dotted and more or less densely short-pubescent, but scarcely if at all arachnoid- tomentose. In a second variety ranging widely through central and western Mexico into Central America, the plants are more pubescent and less conspicuously glandular, the phyllaries are more strongly graduated, and some or all of them obtuse and evidently closely fimbriate-ciliate. I can detect no great variation within the limits of this second population, except that rather densely pilose indi- viduals seem to be more frequent in Chiapas and Central America than from Guerrero westward. The second variety is the following: 1972] Mexican Compositae — McVaugh 503 Eupatorium collinum DC., var. mendezii (DC) McVaugh, comb. nov. Eupatorium mendezii DC. in DC. Prodr. 5: 160. 1836. Eupatorium neaeanum DC. in DC. Prodr. 5: 160. 1836. ?Eupatorium nigrescens Hook. & Arn. Bot. Beech. Voy. 297. 1838. Sonora to Jalisco, Guanajuato (León, Mendez!, in G-DC, the type), Morelos, Guerrero (Acapulco, Née !, in GDC; type of E. neaeanum), Oaxaca, Chiapas and Central America. DeCandolle grouped E. neaeanum, E. mendezii and E. stillingiaefolium with other species having the heads 25- 30- flowered, but E. collinum was described as having the heads 90-flowered, and accordingly was not closely associated with the others in the Prodromus. I suspect that some error was involved here, although Robinson (in the Trees and Shrubs of Mexico, p. 1448) states that the heads in collinum may have as many as 46 flowers. The number of flowers usually varies from 22 to 28; it is rarely as low as 18 or 20, and even more rarely 30 or more; I have seen only one specimen in which the heads were about 36- flowered, and none with a larger head. Robinson separated E. collinum from E. mendezii and E. neaeanum partly on the basis of pubescence and leaf- shape. The latter seems quite useless as an indicator of Specific lines in this group of species. Pubescence in the group is highly variable, as noted above, but the types of E. mendezii and E. neaeanum are not strongly pubescent in comparison with other specimens from central and western Mexico, and the differences between them and typical collinum are no more than might be expected of regional populations in one species. Eupatorium hebebotryum (DC.) Hemsl. Biol. Centr. Am. Bot. 2: 95. 1881. Critonia hebebotrya DC. in DC. Prodr. 5: 141. 1826. Hebeclinium tepicanum Hook. & Arn. Bot. Beech. Voy. 434. 1841. 504 Rhodora [Vol. 74 Eupatorium tepicanum (Hook. & Arn.) Hemsl. Biol. Centr. Am. Bot. 2: 101. 1881. Nayarit (Tepic, Sinclair!, K, type of H. tepicanum), Jalisco, Michoacan, Morelos, Guerrero (Haenke!, in G-DC, the type probably from this state) ; Central America. The distinctions used to separate the two supposed species, namely that between sessile and pedicellate heads, and that in the color of the dried leaves, appear to be inconse- quential. Eupatorium ovaliflorum Hook. & Arn. Bot. Beech. Voy. 297. 1838. Eupatorium. bertholdii Sch. Bip. in Seem. Bot. Voy. Herald 299. 1856. Southern Sonora, ? western Durango (*Sierra Madre", Seemann 2011! at K, isotype of E. bertholdii), Sinaloa, Nayarit (Tepic, Beechey!, at K, the type), Jalisco, Michoa- cán. According to Robinson (in Trees and Shrubs of Mexico, p. 1433), Eupatorium bertholdii is distinguished from E. ovaliflorum by having the heads 10- to 13-flowered and the involucre 2-2.6 mm thick, as against heads 20- to 40-flowered and the involucre 4-5 mm thick. In the material at hand these differences seem not to hold; I have not seen any head with more than 22 flowers, and none as much as 4mm thick. The flowers are usually about 17-20, and the in- volucre 2.5-3 mm thick. Eupatorium polybotryum DC. Prodr. 5: 174. 1836. Nothites ovatifolia DC. in DC. Prodr. 5: 187. 1836, not Eupatorium ovatifolium Hieron., 1908. Ophryosporus ovatifolius (DC.) Hemsl. Biol. Centr. Am. Bot. 2: 79. 1881. Eupatorium petraeum Rob. Proc. Am. Acad. 41: 2775. 1905. Ophryosporus petraeus (Rob.) Rob. Contr. Gray Herb. II.. 75: 4. 1925. Decachaeta ovatifolia (DC.) King & H. Rob. Brittonia 21: 282. 1969. Southeastern Jalisco, western Michoacán, Edo. de México,. 1972] Mexican Compositae — McVaugh 505 Guerrero (Langlassé 565!, GH, type of E. petraeum). A related species ranging from western Michoacán north- westward to Sinaloa and Chihuahua is Eupatorium sca- brellum Rob. Proc. Am. Acad. 35: 339. 1900 [E. micro- cephalum A. Gray, Proc. Am. Acad. 21: 384. 1886, not of Regel; Ophryosporus ovatifolius sensu Rob. in Standl. Contr. U.S. Nat. Herb. 23: 1469. 1926, not Nothites ovati- folia DC.]. E. scabrellum and E. polybotryum are separable as follows: 1. Phyllaries 12-15, 2- 4-seriate, the outer gradually shorter, the innermost (2-5 in each head) longer and narrower than the others and completely or partially modified into pales, deciduous with the achenes or before; flowers (7-) 10-12 (-15). .... D TOR AEHROE COUPES OTT E. polybotryum. 1. Phyllaries 6-8 in 1-2 nearly equal series (in addition to 2-3 smaller outer bracts), the innermost neither prolonged and resembling pales, nor deciduous; lowers 4.9... oaa m as E. scabrellum. The nomenclature of these species has become confused. In the Trees and Shrubs of Mexico, Robinson (1926, p. 1469) took up the name Ophryosporus ovatifolius but mis- applied it to the species correctly known as Eupatorium [Ophryosporus] scabrellum, at the same time recognizing as independent species O. scabrellus (Rob.) Rob. and O. petraeus (Rob.) Rob. Robinson said of his Ophrysporus ovatifolius, “Typical material of this species, early collected by Haenke in Mexico but without indication of locality, has. never been precisely matched except by specimens collected by Seemann, also without recorded locality". Presumably Robinson referred to two collections studied by him at the Gray Herbarium; the Seemann collection is the type of Eupatorium microcephalum A. Gray (= E. scabrellum Rob.). The Haenke specimen consists of a few heads and the tip of a branch of an inflorescence, annotated by Gray as “Nothites ovatifolia DC.', and by Robinson as Eupa- torium polybotryum DC. [ie. Ophryosporus ovatifolius 506 Rhodora [Vol. 74 of Robinson’s treatment in 1926]. Unfortunately Robin- son seems to have misinterpreted this fragment, which certainly represents Eupatorium petraeum, not E. micro- cephalum. The heads are only T- 8-flowered, but the in- volucre is several-seriate, and the narrow paleaceous inner phyllaries are unmistakably those of "E. petraeum". The original Haenke specimens at G-DC, the types of Eupa- torium polybotryum and Nothites ovatifolia, respectively, I have seen through the courtesy of Prof. J. Miége. These represent one and the same species, as Robinson long ago suggested, but that species is “Eupatorium petraeum”, not Ophryosporus ovatifolius in the sense of Robinson. Both the Haenke collections represent a species with 12-15 graduated phyllaries of which the inner are deciduous at maturity; with 1-3 narrow pales on the receptacle; and the flowers 8-11 in each head. King and H. Robinson (1969) first formally combined Eupatorium polybotrium [sic] and Nothites ovatifolia, under the name Decachaeta ovatifolia. The correct epithet for the combined taxa is therefore ovatifolia except in Eupatorium where the name is preoccupied. Gnaphalium canescens DC. in DC. Prodr. 6: 228. 1838. Gnaphalium wrightii A. Gray, Proc. Am. Acad. 17: 214. 1882. There seems to be no significant difference between the plants of nothern Mexico and southwestern United States that have been called G. wrightii (western Texas, Wright 394!, the type), and the plants of central Mexico, from Durango and Aguascalientes to Jalisco, Guanajuato, Hidalgo and the Valley of Mexico, that have been called G. canescens (León, Méndez, in G-DC, the type). The leaves in this species tend to be broadest near the middle or above. A very similar species is Gnaphalium roseum H.B.K. Nov. Gen. & Sp. 4 [ed. fol]: 63. 1818, in which the leaves tend to be oblong or broadest at base. The type (from Guana- juato, Humboldt & Bonpland!, P), is a coarse woolly plant with immature heads. 1972] Mexican Compositae — McVaugh 507 Gnaphalium inornatum DC. in DC. Prodr. 6: 225. 1838. This species was based upon four collections made by Berlandier, viz. no. 1195 (*ad montem de Las Cruces"), no. 740 (from the Valley of Mexico), no. 309 (between Tampico and Real del Monte) and no. 1146 (from Guchi- laque, between Mexico and Cuernavaca). DeCandolle de- scribed it as a woolly plant with narrow erect shortly decurrent leaves, pale reddish (‘‘refescentibus”) phyllaries, about 10 hermaphrodite flowers and 30-40 pistillate flowers. Judging from the specimens in the Prodromus herbarium (cf, also IDC microfiche 1066), and from duplicates (at P) of all the cited numbers, it seems that three different species are represented by the syntypes of G. inornatum. The description in the protologue does not exactly fit any of the species. Nos. 1146 and 1195 apparently represent the same species, in which the number of flowers in a head is 100 or more and the perfect flowers are 7-11 or more. DeCandolle's report of 10 perfect flowers presumably was based on one of these specimens, and one of them may appropriately be designated as lectotype. No. 740 may be eliminated from consideration as the heads are about 50- flowered, and the perfect flowers are 4 in number. De- Candolle's specimen is so immature that it seems unlikely that he dissected a head (cf. Field Mus. neg. 28713). The remaining syntype, no. 309, seems to represent a third species, also one in which the heads are about 50-flowered and the perfect flowers about 5. Berlandier 1146 is designated as lectotype since in the Prodromus herbarium it appears to be a somewhat better specimen (than no. 1195) ; since from the position of speci- mens in the herbarium it seems to have been the one on which DeCandolle's report of 10 perfect flowers was based; and since from Berlandier's notes it appears this collection was more widely distributed than his no. 1195. Gnaphalium sphacilatum H.B.K. Nov. Gen. & Sp. 4[ed. fol.] : 67. 1818. Gnaphalium pedunculosum I. M. Johnst. Contr. Gray Herb. II. 68: 99. 1923. 508 Rhodora [Vol. 74 Durango (Palmer 411! in GH, the type of G. pedunculo- sum), northern Jalisco, San Luis Potosi, D.F., Edo. de México (Teotihuacán, Hahn; beween the City of Mexico and Huehuetoca, Humboldt & Bonpland!, the type, at P). Similar, narrow-leaved plants with uniformly gray thin tomentum occur in various other areas both in North and South America, and have been called by various other names, either treated as distinct species or as varieties of Gnaphalium purpureum. The type of G. sphacilatum is an immature but otherwise typical plant with the charac- teristic bracts, pappus, glabrous acute phyllaries, and linear thinly silky leaves. Gnaphalium stramineum H.B.K. Nov. Gen. & Sp. A[ed. fol]: 66. 1818. Gnaphalium chilense Spreng. Syst. Veg. 3: 480. 1826. Gnaphalium sprengelii Hook. & Arn. Bot. Beech. Voy. 150. 1833. Gnaphalium berlandieri DC. in DC. Prodr. 6: 223. 1838. Material from central and eastern Mexico can usually be distinguished from California specimens of Gnaphalium “chilense” because of the somewhat narrower leaves and the tendency for the uppermost leaves to become bractei- form. The types of G. stramineum (between Moran and Omitlán, Humboldt & Bonpland!, at P), and G. berlandieri (D.F., Berlandier 471!, in G-DC) are representatives of the Mexican population. The Humboldt & Bonpland specimens were apparently badly wilted before drying, and perhaps taken originally from slender or starved plants, but the heads have the characters of this species (flowers 175 or more, hermaphrodite flowers 15-16, phyllaries obtuse, the inner ones about 23). An isotype at Paris (Bonpland 4108) seems clearly to represent the same taxon. The character- istic yellow color of the phyllaries can no longer be seen in the original material of G. stramineum. Gnaphalium viscosum H.B.K. Nov. Gen. & Sp. 4[ed. fol.]: 64. 1818. Gnaphalium hirtum H.B.K. Nov. Gen. & Sp. 4[ed. fol.] : 64. 1818. 1972] Mexican Compositae — McVaugh 509 27Gnaphalium gracile H.B.K. Nov. Gen. & Sp. A[ed. fol.]: 65. 1818. Gnaphalium tenue H.B.K. Nov. Gen & Sp. 4[ed. fol.]: 65. 1818. Gnaphalium leptophyllum DC. in DC. Prodr. 6: 226. 1838. A plant primarily of the Central Plateau of Mexico, widely distributed from western Texas to northeastern Sonara, south at moderate elevations except in the extreme deserts to Oaxaca and Central America. Cronquist (in Vasc. Pl. Pacific N.W. 5: 204. 1955; and Man. Vasc. Pl. N.E. U.S. & Can. 737. 1963) has applied the name Gnapha- lium viscosum very broadly, so as to include the plant of temperate North America that has been better known as G. decurrens Ives, or G. macounii Greene. This seems to be unrealistic; G. viscosum has a characteristic Texano- Mexican range distinct from that of G. macowunii and, com- pared with other currently accepted species of Gnaphalium, it is morphologically very distinct. It differs from G. ma- counii vegetatively in its far more numerous and narrower leaves. The inner phyllaries in G. macounit are usually about 21, the flowers fewer (125-150 in a head) but often larger than those of G. viscosum. At least until the com- plex of glandular, decurrent-leaved fragrant species of Gnaphalium can be revised as a whole, G. viscosum may well be circumscribed narrowly. Nothing exactly referable to G. macounii has been found in Nueva Galicia, but nu- merous collections from the Sierra Madre Oriental suggest that G. macounii ranges well south into eastern Mexico. In Chihuahua G. viscosum may be confused with G. leuco- cephalum A. Gray, in which the phyllaries are pearly white, relatively dull and opaque and symmetrically gradu- ated in length. This is the only Mexican species having the herbage strongly glandular-pubescent throughout, the leaves very numerous (100 or more in well developed plants) and linear or narrowly sagittate, the flowers 200 or more in a head, the mature receptacle 3-4 mm wide, and the plants strictly annual, not forming a basal rosette. The types of G. viscosum, G. hirtum, G. tenue and G. leptophyllum seem 510 Rhodora [Vol. 74 without question to represent the same taxon. The type of G. gracile (Guanajuato, Humboldt & Bonpland! at P) is apparently from a weak, depauperate plant with relatively few leaves and numerous heads. Heterotheca inuloides Cass., var. rosei Wagenknecht, Rho- dora 62: 69. 1960. Heterotheca leptoglossa DC. in DC. Prodr. 5: 317. 1836. In Wagenknecht’s revision, Heterotheca leptoglossa is maintained as a distinct species, distinguished from H. inuoides by “its annual habit, narrow leaves, smaller capi- tular [sic], and linear phyllaries". The type of H. lepto- glossa. (León, Méndez!, in G-DC and cf. Intern. Doc. Center microfiche, Prodromus herbarium, no. 857), is the upper part of what appears to be a vigorous plant of H. inuloides, with leaves normal for that species. As many plants of H. inuloides flower the first year, the differences between annual and perennial habit in this group are of question- able significance. In the type of H. leptoglossa neither phyllaries nor ligules are atypical of H. inuloides. The ligules are not linear as described by DeCandolle but ob- lanceolate, 2.5 mm wide by 10-11 mm long. The heads are of about average size for H. inuloides, and the phyllaries are not unusually narrow. Montanoa [*Montagnaea"] karvinskii DC. in DC. Prodr. 5: 565. 1836. Montanoa olivae Sch. Bip. ex. K. Koch, Wochenschr. Gaertn. 7: 406. 1864. Montanoa gracilis Sch. Bip. ex K. Koch, Wochenschr. Gaertn. 7: 407. 1864. Montanoa subtruncata A. Gray, Proc. Am. Acad. 22: 424. 1887. ?Montanoa affinis Blake, Contr. U.S. Nat. Herb. 22: 612. 1924. A common and widely distributed plant, from Sinaloa to Jalisco, Guerrero and Oaxaca, easily recognized by the wingless petioles, the glabrous lobes and throat of the disk-flowers, and the pilose anthers. It was long known 1972] Mexican Compositae — McVaugh 511 as M. subtruncata, until Blake (Contr. U.S. Nat. Herb. 26: 246. 1930) confirmed the identity of that plant with M. olivae and M. gracilis. After examination of the type of M. karvinskii (without locality, Karvinski s.n. in G-DC), I can confirm Blake's tentative suggestion that this is a still older name applying to the same species. Apparently the plant called M. affinis is merely a nearly glabrous form of the same species. Oxypappus scaber Benth. Bot. Voy. Sulph. 118. 1845. Chrysopsis scabra, Hook & Arn. Bot. Beech. Voy. 434. 1841. not C. scabra, Ell., ?1823. Pectis seemannii Sch. Bip. in Seem. Bot. Voy. Herald 309. 1856. Oxypappus seemannii (Sch. Bip.) Blake, Contr. U.S. Nat. Herb. 26: 261. 1930. The name Oxypappus scaber was a new combination based on the illegitimate name Chrysopsis scabra, Hook. & Arn. According to the International Code of Botanical Nomenclature (Art. 72) such a combination is not to be rejected, but is to be treated as a new name. See also above under Archibaccharis hieracifolia. Tragoceros americanus (Mill.) Blake, Contr. U.S. Nat. Herb. 26: 240. 1930. Calendula americana Mill. Gard. Dict. ed. 8. Calendula no. 10. 1768. Tragoceras schiedeanum Less. Linnaea 9: 269. 1834. The original material of Calendula americana was sent to Miller from Veracruz by William Houstoun, between 1729 and 1733. Blake (1930) reported, after examination of the type, that it represented what had been called Tragoceros microglossus DC. (DC. Prodr. 5: 533. 1836), a rather local species of Jalisco and Guanajuato. According to Torres, however (Brittonia 15: 290-302. 1963), the only species of Tragoceros known from the Atlantic lowlands of Mexico is the one called T. schiedeanus. The most east- erly localities known for T. microglossus (T. “americanus” ) are more than 500 km from Veracruz, in quite a different 512 Rhodora [Vol. 74 vegetational zone. It seemed highly unlikely that a dis- tinctive inland species like T. microglossus should have been found near Veracruz about the year 1730, but never have been recollected there. In April, 1970, therefore, I reexamined the type of Calendula americana (Houstoun, at BM). It is true, as Blake remarked, that the heads are sessile or practically so, as in T. microglossus, but the ligules of the ray-flowers are the narrow, tapering and conspicuously bifid ones of T. schiedeanus, not the obtuse or barely emarginate ones of T. microglossus. As such bifid rays occur in this genus only in T. schiedeanus and the related T. zinnioides (which has larger heads), there can be little doubt that the Houstoun specimen represents what has been called T. schiedeanus. It is therefore un- fortunately necessary to use the name T. americanus for this species, and relegate the well-known T. schiedeanus to synonymy. The original spelling of the generic name was Tragoceros, and according to the International Code of Botanical No- menclature this spelling must be kept, although various authors have changed it to Tragoceras in an attempt to conform to the spelling of the classical Greek word for horn, from which the -ceros part of the name was derived. The Code recommends (Rec. 75A) that names ending in -ceras be treated as neuter in the future, but this does not affect names already published, and in no way authorizes anyone to alter the original spelling of any such name. There is ample biological precedent for names ending in -ceros (not -ceras), as pointed out by Bentham a century ago (Gen. Pl. 2, pt. 1: 356. 1873). Such names as Antho- ceros and Rhinoceros have been consistently treated as masculine since the time of Linnaeus. Tragaceros has been used in its original form by some authors (e.g. Hemsley and Asa Gray), who have treated it as masculine. Others (notably Lessing, DeCandolle and most recently Torres) who have used the -ceras spelling have treated the name as neuter. There would seem to be no justification for this latter course under the Code, and the neuter forms of 1972] Mexican Compositae — McVaugh 513 specific epithets published under Tragoceras are to be treated as orthograhic errors. Trigonospermum adenostemmoides Less. Syn. Gen. Comp. 214. 1832. Since the appearance of the note on this species by Mc- Vaugh and Laskowski (Contr. Univ. Mich. Herb. 9: 498- 500. 1972) I have seen the type of the name, through the courtesy of the Director of the Institut für Systematische Botanik und Pflanzengeographie of the Martin Luther University, Halle (HAL). The specimen bears Schiede's original label: “Composita. Herba annua caule ultraorgyali. Jun. 29", and an annotation in the hand of Lessing: “Trigonospermum n[.] g[.] adenostemoides n[.] sp [.]" The ample fruiting and flowering specimen clearly repre- sents the taxon treated as T. adenostemmoides in our paper cited above. Vernonia triflosculosa H.B.K. Nov. Gen. & Sp. 4[ed. fol.]: 31. 1818. Vernonia barbinervis Sch. Bip. in Seem. Bot. Voy. Herald 297. 1856. Vernonia (?) palmeri Rose, Contr. U.S. Nat. Herb. 1: 101. 1891. Vernonia chacalana Blake, Contr. Gray Herb. II. 52: 19. 1917. Southern Sonora (Palmer 387!, at US, type of V.(?) palmeri), Sinaloa, Durango (Chacala, Goldman 333!, at GH, type of V. chacalana), Nayarit, Jalisco, Colima, Guer- rero (Acahuizotla, Humboldt & Bonpland!, the type) ; Central America. The type of V. barbinervis (Seemann 1998!, at P), came from the “Sierra Madre", i.e. probably from Durango, Sinaloa, or northern Nayarit. According to both Gleason and Blake, Vernonia palmeri is a distinct species ranging from Sonora to Tepic, char- acterized by the (usually) relatively abundant pubescence of the lower leaf-surface. Vernonia barbinervis, known only from the type-region, has the foliage almost glabrous except that the leaves are densely tomentose along the 514 Rhodora [Vol. 74 midvein beneath. In typical Vernonia triflosculosa, sup- posed to range from Colima to Costa Rica, the leaves are said to be glabrous to “thinly tomentulose" beneath. Both glabrous- and pubescent-leaved plants occur in Nueva Galicia, but I do not find any other features correlated with differences in pubescence, and I believe only one species can be recognized. Viguiera puruana Paray, Bol. Soc. Bot. Méx. 22: 4. 1958. Viguiera blakei McVaugh, Contr. Univ. Mich. Herb. 9: 454. 1972. My attention was called to the similarity between Viguiera puruana and V. blakei by a specimen collected in Michoacán (6 km south of Tuxpan on the road to Zitá- cuaro, Rzedowski 25153, MICH) and correctly identified as V. puruana. Since then, through the kindness of Dr. Ramón Riba y Nava Esparza, I have seen the type of V. puruana (Michoacán, San José de Purta, Paray 1780, MEXU). There can be no doubt it represents the taxon more recently described under the name of V. blakei. Xanthocephalum sericocarpum A. Gray, Proc. Am. Acad. 15: 31. 1879. Two very similar species have long been confused under this name. One is a perennial of arid grasslands and their borders, ranging from Querétaro and San Luis Potosi to northern Jalisco and the plains near the City of Durango. The other is an annual, chiefly of pine forests, from western Chihuahua to Aguascalientes and northern Jalisco. The ranges of the two overlap, as far as known, only in Aguas- calientes and Jalisco: 1. Phllaries 16-18 (-25); disk-flowers commonly about 80-85; plants perennial, from a woody taproot. .. MNT X. sericocarpum. 1. Phyllaries 35-50; disk-flowers 100-200; plants annual. peep e ween peessdessaveesaseneess X. conoideum. The type of X. sericocarpum (San Luis Potosi, Parry & Palmer 369!, GH) evidently represents the perennial species. The heads are small, the phyllaries 20-25, broad 1972] Mexican Compositae — McVaugh 515 and relatively firm (not lanceolate or narrowly rhombic with broad hyaline margins as in X. conoideum). A col- lection by Schaffner, mounted on the type-sheet by Gray, apparently represents the same species. The annual species is X. conoideum Hemsl. Biol. Centr. Am. Bot. 2: 110. 1881. The type (Coulter 299!, at K) was collected somewhere in central Mexico, “between Real del Monte and Zacatecas". Zinnia bicolor (DC.) Hemsl. Biol. Centr. Am. Bot. 2: 153. 1881. Mendezia bicolor DC. in DC. Prodr. 5: 533. 1836. Zinnia tenella Rob. Proc. Am. Acad. 63: 39. 1907. Western Chihuahua, Durango (Tejamen, Palmer 500!, at GH, type of Z. tenella) , southern Zacatecas, northern and eastern Jalisco, Guanajuato (León, Méndez! in G-DC, the type), San Luis Potosí. The ligules of the ray-flowers are white or yellow (the two colorforms sometimes mixed in the same population), or yellow with a red spot at base. A plant with yellow ligules red-spotted at base formed the basis for Z. tenella. According to Torres Z. tenella differs from Z. bicolor also in having “lanceolate to elliptic’ rather than “linear to lance-oblong" leaves, and “obovate to cuneate” rather than "linear-elliptic" ray-achenes. In Nueva Galicia the differ- ences in leaf-shape, in achene-shape, and in ray-color, appear to represent individual variations, and it seems futile to try to distinguish more than a single species in the group. Zinnia violacea Cav. Ic. 1: 57. pl. 81. Dec 1791. Zinnia elegans Jacq. Ic. P1. Rar. 3: 15. pl. 589. 1793; Coll. Bot 5: 152 791; Essentially all authors have taken up the name Zinnia elegans in preference to Z. violacea. The place of publica- tion of Z. elegans was correctly cited by a few contemporary authors, as by Willdenow (Sp. Pl. 3, pt. 3: 2140. 1803) and by Sims (in Bot. Mag. pl. 527. 1801). In DeCandolle’s revision of the Compositae, however, through a typo- 516 Rhodora . [Vol. 74 graphical error the reference was given as “Jacq. Coll. 3. p. 152” (DC. Prodr. 5: 536. 1836). This error has been perpetuated by subsequent authors who have merely copied from DeCandolle without looking up the original reference. Thus in Hemsley’s treatment of Zinnia in the Biologia Centrali-Americana, and in the Index Kewensis, the cita- tion is of volume 3 of the Collectanea, not volume 5. As volume 3 was published late in 1791 (cf. Stafleu, F., Taxonomic Literature, p. 232. 1967) it may well have en- joyed priority over the first volume of Cavanilles' Icones, which appeared in December of the same year. The fifth volume of the Collectanea, however, did not appear in print until 1797. It seems clear that the name Zinnia violacea Cav. has priority of more than a year over Z. elegans Jacq., which was published first in 1793 and again in 1797. UNIVERSITY HERBARIUM UNIVERSITY OF MICHIGAN ANN ARBOR, MICHIGAN 48104 THE COMBINATION PELTANDRA VIRGINICA (L.) SCHOTT & ENDLICHER: The indigenous eastern North American genus Peltandra was described by Rafinesque (J. Phys. Chim. Hist. Nat. Arts 89: 103, 1819) when he based it on a simultaneously described new species, P. un- dulata Raf. in a somewhat confusing manner, Rafinesque states that “Calladium sagittaefolium" (= Arum sagittae- folium L., a member of the genus Xanthosoma according to Fernald, Rhodora 50: 59, 1948) and “C. virginicum” (pre- sumably — Arum virginicum L.) are related to Peltandra, but he apparently did not intend their inclusion in the genus. Rafinesque (New Flora and Botany of North Amer- ica 1: 87, 1836) later conceded that P. undulata was probably identical to Arum virginicum. However, he re- 1972] Peltandra virginica — Blackwell 517 tained the name P. undulata, failing to employ the correct combination P. virginica, Index Kewensis wrongly at- tributes the combination P. virginica to Rafinsque. Most authors who have dealt with Peltandra virginica credit the name to Kunth (Enumeratio Plantarum 3: 43, 1841). Included in this list are: Morong (Mem. Torrey Bot. Club 5: 102, 1894), Tidestrom (Rhodora 12: 48, 1910), Blake (Rhodora 14: 104, 1912), Small (Manual of the Southeastern Flora 246, 1933), Barkley (Madrofio 7: 133, 1944), Gleason (New Britton and Brown Illustrated Flora 1: 368, 1952), Huttleston (Taxon 2: 33, 1953), Fassett (A Manual of Aquatic Plants, second ed., 164, 1957), and Radford et al. (Manual of the Vascular Flora of the Caro- linas 257, 1968). On the other hand, Fernald (Rhodora 50: 56, 1948; Gray's Manual of Botany, eighth ed., 383, 1950) and Merrill (Index Rafinesquianus 81, 1949) recog- nize Scott and Endlicher (Meletemata Botanica 19, 1832) as having made the combination. Clearly, authorship of this combination is a point of nomenclatural confusion. A study of the original literature of concern revealed that the combination Peltandra virginica was first used in print by Steudel (Nomenclator Botanicus 1: 603, 1821) when he mistakenly attributed the combination to Rafin- esque. Steudel should not receive credit for a new com- bination in this instance as he considered P. virginica a member of the genus Arwm, indicating it as a synonym of Arum virginicum. The only species accepted by Steudel as belonging to Peltandra is P. undulata. Steudel (Nomen- clator Botanicus, second ed., 1: 249, 1840) later discon- tinued recognition of the genus Peltandra and included it in the synonymy of Caladium. Merrill (1949) notes Steudel’s (1821) use of the name P. virginica and rightly does not credit him with the combination. Schott and Endlicher (1832) accord Peltandra generic status and accept P. undulata and P. virginica as distinct species, attributing both names to Rafinesque. Kunth (1841) cites Schott and Endlicher’s treatment and similarly recognizes both species, also crediting Rafinesque with the names. At 518 Rhodora [Vol. 74 a considerably later date, Schott (Synopsis Aroidearum 50, 1856) maintains the same opinion concerning the taxa of Peltandra and their authorship. Although there can be little or no support for regarding P. virginica and P. undulata as distinct, Schott and End- licher (1832) validly make the combination Peltandra virginica for the first time, even though they erroneously attribute the combination to Rafinesque. Fernald (1948, 1950) and Merrill (1949) are thus vindicated in their designation of authorship as Peltandra virginica (L.) Schott & Endlicher. By the new wording of Recommenda- tion 46C of the International Code of Botanical Nomen- clature, as proposed by Yeo (Regnum Vegetabile 60: 62, 1969, Proposal 281) and amended and accepted by the Eleventh International Botanical Congress (see Taxon 19: 48, 1970), the citation of authorship as Peltandra virginica (L.) Rafinesque ex Schott & Endlicher would be equally correct, though in my opinion less desirable since the ascription to Rafinesque is apparently a mistaken one. I wish to gratefully acknowledge the American Philo- sophical Society for grant support of my current systematic investigations of the genus Peltandra. WILL H. BLACKWELL, JR. DEPARTMENT OF BOTANY MIAMI UNIVERSITY OXFORD, OHIO 45056 SPONTANEOUS HYBRIDS BETWEEN CERASTIUM TOMENTOSUM LINN. AND C. ARVENSE LINN. J. K. MORTON The “White Rock" or “Snow in Summer" of our gardens — Cerastium tomentosum Linn. is a native of southern Europe, now commonly cultivated as a rock garden plant throughout the temperate world. It is a very vigorous peren- nial which rapidly spreads and takes over the whole of the rock garden, hence its rhizomes are frequently dug up and thrown onto roadsides, river banks and rubbish dumps where it sometimes becomes established and competes suc- cessfully with the native flora. C. arvense Linn. is a native species in North America where the most widely occurring form is diploid. However the common European form is tetraploid (2n = 72 as in C. tomentosum) and has been introduced into northeastern North America where it fre- quently becomes a weed along roadsides, riverbanks, and in old pasture and quarries etc. Whether its introduction to this continent was deliberate or accidental is uncertain, though the former is not improbable for it makes an at- tractive rock garden plant, apart from its propensity to spread too vigorously. In the mid and upper reaches of the Grand River valley in southern Ontario, both these species occur; C. tomen- tosum as a common garden plant persisting in derelict gardens and occasionally established as a garden throwout; and C. arvense as a well established weed on road, rail and river banks etc. Hybrids between the two species have been discovered at Fergus, Waterloo and Galt. At Fergus the hybrid plants are growing amongst concrete slabs on a grassy roadside bank at the Elora end of the town. The area probably originated as a rockery but has long been neglected, and both parent species are growing alongside the hybrids. Further plants of the hybrid occur below the road, on old ballast used in constructing the road and dumped on the river bank. At Waterloo the plant was 519 520 Rhodora [Vol. 74 found in an old rockery of a long since derelict farm house, and has since been transplanted into several rock gardens in the town. Neither parent now grows in the immediate vicinity of the old farm house and the nearest colony of C. arvense is several miles away. At Galt the hybrid plants are growing amongst grass on cindery ground on an old rubbish tip, and in the lawn of an adjoining house on the Brantford side of the town between route 24 and the Grand River. Neither parent is growing in the immediate vicinty but C. arvense occurs along the Grand River about a mile distant. The Fergus and Waterloo plants appear to be the F; hybrid and are intermediate between the two parents, but somewhat closer to C. tomentosum in general appearance, looking like a rather poorly developed plant of that species in which the white indumentum is less dense. The Galt colony apparently consists of F» segregates probably result- ing from selfing of the F, hybrid. They are smaller, very variable plants and several of them are nearer to the C. arvense parent in general appearance. These hybrids flower copiously and produce a number of short capsules, the teeth of which barely exceed the calyx. Most of the ovules in these abort, but frequently one or two, occasionally more, very large seeds are formed. These are fertile and the progeny show extensive segregation, ranging from close to one or other of the parents, to various combinations of characters intermediate between the parents. It is probable that these seeds result from selfings rather than pollination from either parent. Hybrids between C. tomentosum and C. arvense are rarely encountered in Europe where the species are native and sometimes grow in close proximity. The only reference in the literature to such hybrids, that I have been able to locate, is Ascherson and Graebner 1919 where they are referred to C. maureri and C. rigoi without authority or supporting information. Neither of these names appears to have been taken up by subsequent workers, perhaps be- cause of the very rare occurrence of this hybrid. 1972] Cerastium Hybrids — Morton 521 REFERENCE ASCHERSON, P. and GRAEBNER, P. 1919. Synopsis der Mitteleuro- paischen flora. 5, 1:636. DEPARTMENT OF BIOLOGY UNIVERSITY OF WATERLOO WATERLOO, ONTARIO, CANADA AMPHIPRORA ORNATA BAILEY — A SECOND STATION IN NEW ENGLAND L. GC. COLT, Jg. During continuing studies of the phytoplankton of the Connecticut River I collected a single specimen of Amphi- prora ornata Bailey. As the accompanying figure shows the species is distinct enough in structure so as to make it readily recognizable. The specimen was recovered from material taken by a vertical haul with a Wisconsin-type sampler in 46 feet of water. The collecting station is located 0.1 miles north of the dam at Vernon, Vermont. All of the stations being utilized during this study are within five miles of Vernon. The collection was made just before 12 Noon on June 5, 1970. At this time the air temperature was 68 degrees F., and the water temperature was 66 degrees F. Although the collecting station is physically closer to Vermont than New Hampshire, ownership of the Con- nectieut River below high water mark is vested in the State of New Hampshire. Thus, Amphiprora ornata Bailey is reported for the Town of Hinsdale, Cheshire County, New Hampshire. This appears to be only the second collection for New England. The previous station being somewhere in the vicinity of New Haven, Conn., reported by Terry (1907). Other reported stations include the type from Florida by 522 Rhodora [Vol. 74 I-/2 u~ Amphiprora ornata Bailey. Camera lucida drawing by the author. 1972] Amphiprora — Colt 523 Bailey (1850), Boyer (1827) in Pennsylvania, and Whit- ford (1969) in North Carolina. LITERATURE CITED BAILEY, J. W. 1850. Microscopial observations made in South Carolina, Georgia, and Florida. Smithsonian Contributions to Knowledge, Volume II, Art. 8. Boyer, C. S. 1927. Synopsis of North American Diatomaceae. Proc. Acad. Nat. Sci. Phila., Vol. 79(2): 229-583. TERRY, W. A. A partial list of Connecticut diatoms with some account of their distribution in certain parts of the state. Rho- dora 8(104): 125-140. WHITFORD, L. A., & G. J. SCHUMACHER. 1969. A Manual of the Fresh-water Algae in North Carolina. North Carolina Agric. Exp. Station Tech. Bull. 188. BOSTON STATE COLLEGE BOSTON, MASS. 02115 THE YELLOW MANDARIN, A NOTABLE ADDITION TO THE FLORA OF ARKANSAS PAUL L. REDFEARN, JR. Although the yellow mandarin, Disporum lanuginosum (Michx.) Nickols, is widespread in the eastern United States, it was, according to Robert G. Johnson (personal communication), not known west of Davidson County, Tennessee, except for one record of its occurrence in Ar- kansas which was most likely in cultivation. Dr. E. B. Smith (personal communication) also informs me that a single specimen from Texarkana, Miller County, Arkansas, is present in the herbarium of University of Arkansas and is marked “probably cultivated." The discovery of a new locality for the yellow mandarin in Arkansas is therefore notable for two reasons. First, because of the area where it was discovered, there can be no doubt now that it is a native, though rare, member of the flora of Arkansas. Sec- ond, it adds to the growing list of southern and Appalach- 524 Rhodora [Vol. 74 ian species present in the relic mesophytic forests com- mon throughout the Boston Mountains (Braun, 1950). This list includes such vascular plants as Fagus grandi- flora, Liquidambar styraciflua, Magnolia tripetala and Ly- copodium lucidulum, and bryophytes such as Hookeria acutifolia, Brotherella tenuirostris, Schwetschkeopsis fab- ronia, Sciaromium lescurii, Sphagnum capillaceum var. tenerum and Bryoxiphium norvegicum. Except for this last species, which occurs in several nearby ravines, all of these taxa are located in the immediate vicinity of the hab- itat of Disporum lanuginosum. Another notable species in this area is Dodecatheon frenchii. Like D. lanuginosum, it also occurs along the base of a north-facing sandstone bluff; otherwise, this species is known from one other lo- cality in Arkansas (Olah & DeFillipps, 1968) and in the Ozarks of southern Illinois (Voigt & Mohlenbrock, 1964). The specific collection data for Disporum lanuginosum in the Boston Mountains of Arkansas is: Newton County. On rocky soil at base of north-facing sandstone bluff, upper reaches of Terrapin Branch, sect. 26, T. 14 N., R. 23 W., alt. ca. 700 ft., Redfearn 27381 (ARK, SMS, NCU, UMO). LITERATURE CITED BRAUN, E. Lucy. 1950. Deciduous forests of eastern North America. The Blakiston Co., Philadelphia. p. 170. MOHLENBROCK, RoBERT H. and Joun W. VoicT. 1959. A Flora of Southern Illinois. Southern Illinois University Press, Car- bondale. p. 266. OLAH, L. V. and R. A. DEFiLLIPPS. 1968. A cytotaxonomic study of French's shooting star. Bull. Torrey Bot. Club. 95: 168-198. DEPARTMENT OF LIFE SCIENCES SOUTHWEST MISSOURI STATE UNIVERSITY SPRINGFIELD, MISSOURI 65802 RHAPIDOPHYLLUM HYSTRIX IN MISSISSIPPI J. RAY WATSON Rhapidophyllum hystrix (Pursh) Wendland & Drude, the Needle Palm, is a rare shrub of the southeastern Coastal Plain. It occurs sporadically from central Florida and southern Georgia westward to Mississippi (Small, 1923; Harper, 1928; Small, 1933). Recent field studies have extended its known distribution in Mississippi; herbarium specimens will be deposited in MISSA. The northernmost site known in Mississippi is in Lauder- dale County, approximately 10 miles south of Meridian (Watson 7314), where it was discovered by Leslie Hu- bricht. Southward, it has been found in Clarke (Watson 7439) and George (Denier 1273) counties and reaches its southern limit in Jackson County (Watson 7870). It ex- tends westward to Simpson County where it was noted by R. B. Channel in the early 1960’s (personal communica- tion). A recent search of this site to obtain herbarium specimens was, however, unsuccessful. Recently it has been found in Forrest County (Watson 8115, October 25, 1969), approximately 60 miles east of the Simpson County local- ity. Within the state, the Needle Palm is known to occur either along the flood-plains of small streams or along the drainages of rich, wooded ravines. LITERATURE CITED Harper, R. M. 1928. Economic Botany of Alabama, Part 2. Cata- logue of the trees, shrubs and vines of Alabama, with their economic properties and local distribution. Geol. Surv. Ala. Monogr. 9. SMALL, J. K. 1923. The needle palm — Rhapidophyllum hystrix. Jour. N. Y. Bot. Gard. 24: 105-114. 1933. Manual of the Southeastern Flora. The author, New York. DEPARTMENT OF BOTANY MISSISSIPPI STATE UNIVERSITY STATE COLLEGE, MISSISSIPPI 39762 525 EQUISETUM FROM PENINSULA FLORIDA DON R. REYNOLDS AND KEN CALHOUN The first report of Equisetum in Florida was made in 1932 when E. praealtum was discovered in Northern Florida (Small, 1938). E. robustum is recorded from 1929 growing along the Apalachicola River at Alum Bluff. Cor- rell (1938) lists E. praealtum as occurring along wet sandy stream banks in Liberty County, Florida. Schaffner (1939) - maps the locality of Equisetum in Florida as occurring in the pan-handle of the State. Wherry (1961) lists E. hyemale var. elatum as occurring in Gadston county along the Georgia border and westward to Texas. He lists the habitat as, “sandy shores and seemingly barren areas where moisture and nutrients can be found at depth. In- vading disturbed soil, fills . . ." Heretofore Equisetum is known only from the pan-handle area of Northwest Florida. We wish to report Equisetum from peninsula Florida. Two collections have been identified as Equisetum hyemale var. affine, following Hauke (1963). This determi- nation has been confirmed by a comparison with the speci- men of G. Engelmann in the Missouri Botanical Garden, St. Louis, which is the type of this variety. SPECIMENS EXAMINED: Florida Technological Uni- versity Herbarium: 8 February 1971. Ken Calhoun and Don R. Reynolds, Brooksville Limestone Quarry, Liberty County, Florida. The specimens were growing in moist clay in several places in the quarry. The quarry is approximately 14 mile long and 300 yards wide and 50 feet deep. It was abandoned in 1967. There was little competition from other plants for the large Equisetum clones which flourished along a large water filled pit that ran the length of the quarry. The area immediately surrounding the quarry and other likely areas within a 25 mile radius were searched on several 526 1972] Equisetum — Reynolds and Calhoun 527 occasions. Although several apparently suitable habitats were located, Equisetum seems to be restricted to the con- fines of the Brooksville Quarry. 9 May 1971. Don R. Reynolds and Ken Calhoun, Sand Quarry, approximately 10 miles east of Clermont on US Highway 50, Lake County, Florida. The specimens were growing in sandy soil which was intermittently wetted from water being pumped into the quarry as part of the dredging operation. The quarry covered about one square mile; a large part was covered with water. No vegetation was growing in the immediate proximity of the Equisetum clone. Hauke (1963) has demonstrated the clinal variation be- tween the southern states of E. hyemale var. affine, based on teeth retention, height of plants and ridge angularity. The clines of Mississippi, Alabama and Northern Florida are similar to the clines of Georgia except that the per- centage of teeth retention in the Georgia clines is much lower. According to the percentage of teeth retention in the two specimens cited (80-100%), the Equisetum from Liberty and Lake Counties represent a southerly extension of the Mississippi, Alabama, North Florida clines rather than the Georgia one. A third collection of Horsetail was recently given to Dr. Robert Long (Biology Department, University of South Florida) which was found growing in a ditch in Dunedin, Pinellis County, Florida. Dr. Long (personal communica- tion) identified the plants as Equisetum hyemale. This collection represents the southernmost record on the Florida peninsula. LITERATURE CITED CoRRELL, D. S. 1938. A county check-list of Florida ferns and fern allies. American Fern Journal 28: 11-15. HAUKE, RICHARD. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. J. Cramer. Stuttgart. 123 p. SCHAFFNER, J. H. 1939. Distribution of exclusively North American species of Equisetum. American Fern Journal 29: 45-47. 528 Rhodora [Vol. 74 SMALL, J. K. 1938. Ferns of the southeastern states. Hafner Co. N. Y. Reprint Edition. 517 p. Wuerry, E. T. 1961. The southern fern guide. Southern, eastern and midland United States and adjacent Canada. Doubleday and Co., Garden City, N. Y. 318 p. DEPARTMENT OF BIOLOGICAL SCIENCES FLORIDA TECHNOLOGICAL UNIVERSITY ORLANDO, FLORIDA 32816 WIND AND THE WINTER-EXPOSED PLANT. In his paper “Snow Cover and the Diapensia lapponica habitat in the White Mountains, New Hampshire” (Rhodora 74: 358- 377), W. N. Tiffney, Jr. makes repeated reference to the prevalence of high winds in the alpine zone which, in his belief, *. . . promote desiccation in winter-exposed plants". The idea that winter desiccation is significantly influenced by high winds is shared by at least one other ecologist (Lind- say, 1971), but this conclusion seems to be based on the application of summertime wind effects to the wintertime situation. The energy budget of the winter-exposed plant is, however, quite different from that of the more freely transpiring plant in a summer microclimate, and the preva- lence of high winter winds may instead forestall damaging water loss. Transpiration from a leaf surface, at any time, is directly proportional to the water vapor concentration gradient be- tween the leaf and air, and is inversely proportional to the diffusive resistances offered by the leaf and boundary layer of air adjacent to the leaf (Gates, 1965). Hence T DL 7 = 1 a where T is the transpiration rate, c; — c, is the water vapor concentration difference between the intercellular spaces of the leaf and the bulk air outside the leaf boundary layer, and r, and r, are respectively the leaf and boundary re- sistance to gaseous transfer. It should be noted that the driving force for transpiration, c; — Ca, is strongly influenced by the temperature difference between the leaf and air, since saturation vapor pressure is a function of tempera- ture. The greater the elevation of leaf temperature above 1972] Winter-exposed Plants — Marchand 529 air temperature, the greater the vapor concentration gradi- ent between the leaf and air. Leaf resistances to the diffusion of water vapor are pro- vided by the stomates and cuticle, the relative magnitude of these being somewhat species dependent. The minimum stomatal resistance of the alpine species Ledum groenlandi- cum is for, example, less than 2 sec cm-t, as determined for plants growing in both a northern bog (Small, 1972) and above timberline on Mt. Washington, New Hampshire (Marchand, unpublished data). The stomatal resistance of conifers may be somewhat higher, perhaps near 20 sec cm- (Waggoner and Turner, 1971). In contrast, cuticular resistance is usually several times greater in magnitude, having been reported (Holmgren et al., 1965) as high as 460 sec cm"! for the European species Quercus robur. In addition, cuticular resistance has been found to increase sharply with decreasing temperature (Holmgren et al., 1965). Boundary layer resistance is provided by a transfer zone of air in contact with (and influenced by) the leaf. The thinner this surrounding layer, the more rapid will be the heat convection or vapor transfer through this zone, since heat and vapor concentration gradients between the leaf and bulk air will be steeper. Single leaf boundary layer resistances have been found to be generally less than 0.9 sec cm-! for severa] deciduous woodland species (Holmgren et al., 1965), although this will vary directly with leaf size and shape and inversely with windspeed (Gates, 1965). It is through reduction of boundary layer resistance that the influence of wind currents is important. The effect of wind is two-fold; (1) it facilitates removal of moist air from the leaf surface, thereby increasing the rate of trans- piration, and (2) it increases the rate of heat transfer from the leaf, through forced convection, thus tending to main- tain temperature equilibrium between the leaf and air. In the latter case, c; — c, is reduced and consequently transpira- tion is decreased. The relative importance of these two processes will depend on other microenvironmenta] factors as well as on the physiological behavior of the plant. 530 Rhodora [Vol. 74 During the summer growth period, the most significant leaf resistance to loss of water (under non-stress condi- tions) is that of the open stomates. Cuticular transpiration is negligible as long as the stomates remain open. At this time of year then, the boundary layer resistance is closer in magnitude to the leaf resistance and any reduction of r, by turbulent exchange becomes significant in terms of in- creasing transpiration. Under these circumstances, the ef- fect of wind on leaf temperature seems to be of minor importance. In the wintertime, the relative importance of the diffusive resistances is changed significantly. Stomatal opening has not been reported to occur during the winter (Tranquillini, 1964; Schulze et al., 1967) as apparently it is prevented by low temperatures (Staefelt, 1962). As a result, vapor dif- fusion is largely via cuticular pathways and the leaf re- sistance thus becomes very high.* It is on this point that my argument is based. As seen from the above transpiration model, when r, is high, r, becomes negligible and any re- duction of the boundary layer resistance by high winds is unimportant in terms of increasing vapor transfer. So the dominant effect of wind in the wintertime is, in my opinion, related to the consequences of forced convection, and these are (1) maintenance of leaf temperatures below the freez- ing point of cell water when air temperatures are very low, and (2) reduction of temperature differences between the leaf and air with consequent reduction of e, — ca. In the first case, water loss will be limited to the process of sublimation from frozen tissues, requiring greater energy input than for the evaporation of free water. In both cases, the net effect of wind would be to reduce, rather than increase water loss. Where reference has been made to the work of Tranquil- lini (1964) and Sakai (1970), citing conditions of winter drought and desiccation damage in alpine areas, it should *The discussion here is pertinent only to evergreen species, and since heavy cutinization is characteristic of evergreens, the value of cuticular resistance is assumed to be very high relative to r.. 1972] Winter-exposed Plants — Marchand 581 be noted that such conditions are the combined result of frozen soils and the strong heating of plant parts above freezing, due to high direct and reflected radiation load on exposed plants. Thus it is most likely that winter-exposed plants will experience damaging water loss on days which are clear and windless. Whether or not winter desiccation in the alpine areas of the White Mts. of New Hampshire can be substantiated, the conclusion that “. . . low tempera- tures and high winds combine to promote severe desiccation in this snow-free area" seems unwarranted on the basis of simple physical considerations. LITERATURE CITED Gates, D. M. 1965. Energy, plants, and ecology. Ecology 46: 1-13. HOLMGREN, P., P. G. JARVIS, and M. S. JARVIS. 1965. Resistances to carbon dioxide and water vapor transfer in leaves of different plant species. Physiol. Plant. 18: 557-573. LINDSAY, J. H. 1971. Annual cycle of leaf water potential in Picea engelmannii and Abies lasiocarpa at timberline in Wyoming. Arctic and Alpine Res. 3: 131-138. SAKAI, A. 1970. Mechanism of desiccation damage of conifers win- tering in soil-frozen areas. Ecology 51: 657-664. ScHULZE, E. D., H. A. Mooney, and E. L. DUNN. 1967. Wintertime photosynthesis of bristlecone pine (Pinus aristata) in the White Mountains of California. Ecology 48: 1044-1047. SMALL, E. 1972. Water relations of plants in raised sphagnum peat bogs. Ecology 53: 726-728. STALFELT, M. G. 1962. The effect of temperature on opening of the stomatal cells. Physiol. Plant. 15: 772-779. TIFFNEY, W. N., JR. 1972. Snow cover and the Diapensia lapponica habitat in the White Mountains, New Hampshire. Rhodora 74: 358-377. TRANQUILLINI, W. 1964. The physiology of plants at high altitudes. Ann. Rev. Plant Physiol. 15: 345-362. WAGGONER, P. E. and N. C. TURNER. 1971. Transpiration and its control by stomata in a pine forest. Conn. Agr. Exp. Sta., Bull. 726, 87 p. PETER J. MARCHAND DEPARTMENT OF BOTANY UNIVERSITY OF NEW HAMPSHIRE DURHAM 03824 XII INTERNATIONAL BOTANICAL CONGRESS JUNE 23-30, 1975 The Closing Plenary Session of the XI International Bo- tanical Congress held at Seattle, U.S.A., in 1969, accepted an invitation issued by the Academy of Sciences of the U.S.S.R. to convene the XII International Botanical Con- gress in the City of Leningrad in 1975. In 1971 the Organiz- ing Committee was appointed: consisting of a chairman (A. L. Takhtajan) ; four vice-chairmen (A. A. Prokofiev, A. A. Theodorov, N. V. Tsitsin, A. A. Yatsenko-Khmelev- Sky) ; a secretary-general (O. V. Zalensky) ; a scientific sec- retary (N. S. Snigirevskaya) ; and a number of members at large. The XII International Botanical Congress is in- tended to facilitate interdisciplinary communication among botanists as well as an informal exchange of ideas. A num- ber of sections are planned, including special ones to accom- modate mycologists (also lichenologists), phycologists, and bryologists. The Congress will be divided between organized half-day symposia and half-day contributed paper sessions. In ad- dition to the opening and closing plenary sessions, two even- ing lectures are being scheduled. All special interest groups wishing to apply for space and time during the Congress should do so by writing as soon as possible to the secretary- general, Dr. Oleg Zalensky, Komarov Botanical Institute of the Academy of Sciences of the U.S.S.R., 2, Popov Street, Leningrad 197022, U.S.S.R. The sessions of the Nomenclature Section will take place, as usual, immediately before the opening of the Congress — in this instance June 20-23. Four days are set aside in order to enable the Section to convene for six to eight ses- sions of two to four hours each. A meeting of the International Association of Botanic Gardens (President Academician N. V. Tsitsin) will be held in Moscow at the Main Botanical Garden of the Academy of Sciences of the U.S.S.R. on June 20. A tentative schedule of scientific field trips has been 532 1972] Botanical Congress — Takhtajan 533 planned for the immediate pre-Congress and post-Congress periods. The principal purpose of these trips is to acquaint visiting botanists with as many interesting and unique fea- tures of the flora and vegetation of various regions of the U.S.S.R. as possible. Some specialized trips for phycologists, lichenologists, bryologists, and palaeobotanists are also planned. The double postcards announcing the XII International Botanical Congress will be mailed during the last months of 1972. Those who wish to receive further information on the Congress should return their interest cards by March 1, 1973, so that they will be placed on the mailing list for the First Information Circular expected to be published June- July 1973. CHAIRMAN, ORGANIZING COMMITTEE A. TAKHTAJAN REQUEST FOR UNNEEDED COPIES OF RHODORA FOR DECEMBER 1971, NO. 796 The error on the cover of the fourth number of Rhodora for 1971, indicating it as September 1971, no. 795, has resulted in many persons discarding it as a duplicate. Had they looked inside the cover they would have detected the error immediately. However so many requests have come in for replacements that Dr. Herman Sweet requests that any subscribers who have this December issue, and do not regularly keep a file of Rhodora, send this particular copy to him at the Botanical Museum, Oxford Street, Cambridge, Mass. 02138. Dovora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL RALPH CARLETON BEAN ROBERT CRICHTON FOSTER ROLLA MILTON TRYON >Associate Editors RADCLIFFE BARNES PIKE STEPHEN ALAN SPONGBERG GERALD JOSEPH GASTONY VOLUME 74 1972 Che Nef England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge, Mass. 02138 INDEX TO VOLUME 74 New scientific names and combinations are printed in bold face type Abies balsamea 367 Abronia Fendleri 83 Acalypha rhomboidea 392 Acanthospermum australe 203; humile 205 Acer rubrum, var. drummondii 147; f. tomentosum 147; sac- charinum 392 Acrochaetium catenulatum 92, 93; compactum 92; crassipes 92, 93; daviesii 268; microfilum 93 Acroptilon picris 344; repens 344 Actinella argentea 84; Richard- sonii, var. floribunda 84 Actinomeris heterophylla 97-100; pauciflora 97-100 Adams, Preston, Studies in the Guttiferae III. An Evaluation of some Putative Spontaneous Garden Hybrids in Hypericum sect. Myriandra 276 Aesculus glabra 147; var. leuco- dermis 147 Aethusa cynapium 156 Agalinis heterophylla 387 Agardhiella tenera 480 Agarum cribrosum 267, 416 Ageratella microphylla 497; var. palmeri 497; var. seemannii 497; palmeri 497 Ageratum microphyllum 497 Agoseris purpurea 84 Ahnfeltia plicata 269, 409, 421 Alaria esculenta 267, 411, 416 Alcina minor 176; ovalifolia 175; ovatifolia 175; perfoliata 12, 194, 195 Algae of Kent Island, Bay of Fundy. On the Marine 264 Algal Invaders of the Northeast- ern Fringes of the Sargasso Sea, Some 295 Algal Vegetation of the Hamp- ton-Seabrook Estuary and the Open Coast Near Hampton, New Hampshire, The 406 Alsophila Abbottii 445; Brooksii 445; bryophila 443-445; capen- sis 445; confinis 445; dryopte- roides 445; Engelii 445; hot- teana 445; minor 445, 446; Nockii 445; pauciflora 445; sagittifolia 446; Salvinii 445; Urbanii 445 Alyssum alynoides 391; deserto- rum 381, 391 Amalias anceps 283 Amaranthus caudatus 143 Amberboa moschata 344; muri- cata 339 Ambrosia tomentosa 394 Amphiprora ornata 521, 522; Bailey, A Second Station in New England 521 Andropogon intermedius 378 Anemone multifida 391 Anoda cristata 383 Antithamnion floccosum 421 Apocynum medium 385 Aquatic Plants in New England, Range Extensions of Vascular 131 Arabis Fendleri 83; glabra 391 Arceuthobium cryptopodium 83 Archibaccharis hieraciifolia 498, 511; hieracioides 498 Arenaria groenlandica 367-370, 372-374 Aristida Fendleriana 83; longi- seta 83 537 538 Artemisia annua 388; arctica 110, 111; borealis 110; frigida 110; norvegica 111; saxicola 102, 110 Arum sagittaefolium 516; virgin- icum 516, 517 Asclepias incarnata, var. incar- nata, f. albiflora 385 Ascophyllum mackaii 286; nodo- sum 268 295, 297, 411, 416, 418, 425, 481, 483; f. scorpi- oides 411, 416, 418, 482; ecad. Mackaii 130 Ascyrum hypericoides, var. hy- pericoides 145; var. multicaule 145 Asperococcus echinatus 267 Aster adscendens 104; chilensis 104; ciliolatus 394; ericoides 104; exilis 104; foliaceus 104; hirtifolius 104; lateriflorus 104; occidentalis 104; puniceus 104; sagittifolius 394 Asteraceae, Somatic Chromo- some Numbers for Some 102 Asterocytis ramosa 421, 427, 428 Astragalus ceramicus 83; loncho- carpus 83; lotiflorus 392; purshii 391 Atriplex glabriuscula 390; has- tata 391; heterosperma 390; subspicata 391 Atwood, N. Duane, New Species in the Phacelia crenulata Group (Hydrophyllaceae) 451 Audouinella catenulatum $85, 92, 96; crassipes 85, 86, 96; mem- branacea 421; microfilum 85, 89, 92, 96; microscopica 85-93, 96; (Naeg.) Woelk. Complex (Rhodophyta), Studies on 85 Austin, Daniel F. Interactions between Apis mellifera (Hy- menoptera: Apidae) and Tribu- lus cistoides (Zygophyllaceae) 117 Baccharis hieracifolia 498; hie- Rhodora [Vol. 74 raciifolia 498; potosina 499; sulcata 499; thesioides 499 Bangia ciliaris 421; fuscopur- purea 413, 421 Beardsley, Richard L. and Edw. T. Browne, Jr., Lathyrus aphaca L. New to Tennessee and the Southwest 155 Berberis repens 391 Bidens acrifolia 499; acuta 394; comosa 388, 394; polyglossa 499; tenuisecta 84; vulgata 394 Blackwell, Will H., The Combina- tion Peltandra virginica (L.) Schott & Enlicher 516 Blidingia marginata 412; minima 412 Botanical Congress, XII Interna- tional 532 Botrychium lunaria 389 Bowdoin College, The Parker Cleaveland Herbarium of 80 Brasenia schreberi 137 Bromus squarrosus 389 Brotherella tenuirostris 524 Browne Edw. T., see Beardsley, Richard L. Bryopsis plumosa 412, 480 Bryoxiphium norvegicum 524 Bulbostylis capillaris 294 Camutia perfoliata 195 Caragana arborescens 392 Calendula americana 511, 512; de- cumbens 21, 206 Calhoun, Ken, see Reynolds, Don R. Calladium sagittaefolium 510; virginicum 516 Callithamnion baileyi 421; corym- bosum 421 Callitriehe terrestris 383 Calochortus madrensis 351; venu- stulus 350, 351; (Liliaceae), Variation Within 350 var. im- bricus 350, 351; var. madrensis 351; var. venustulus 350, 351 1972] Calothrix confervicola 484; crus- tacea 476, 478 485-487 Cape Cod Species of Carex, The Rachilla in, With Notes on the History of the Perigynium and Rachilla 321 Capsosiphon fulvescens 481, 484 Carex acuta 324; alopecoidea 390; bigelowii 367, 370, 372-375; bul- lata 323; buxbaumii 390; Cam- posii 323; capillaris 390; canescens 323; complanata 323, 327; crinita 326; davisii 390; debilis, var. Rudgei 321, 323; elachycarpa 329; Emmonsii 321, 323; flacca 324; flava 324; Fraseri 323; gracillima 390; hirta 324; hystricina 149; lacus- tris 390; lasiocarpa 390; limosa 390; melanostachya 379; micro- glochin 322, 329, 330; Oederi 324; oedipostyla 328; pallescens 323-325; var. glaberrima 325; var. neogaea 325; pensylvanica 321, 323; pseudocyperus 324; scoparia 321, 323; Sprengelii 326; squarrosa 148; stipata 390; Swanii 321; tonsa 323; vesicaria 390; Willdenowii 325 Carex, the Rachilla in Cape Cod Species of, with Notes on the History of the Perigynium and Rachilla 321 Catling, P. M., see Cruise, J. E. Cedrela discolor 125; oaxacensis 124-126; C. DC. & Rose, Re- definition of 124; poblensis 126; salvadorensis 124, 126; saxatilis 126; tonduzii 124 126 Centaurea americana 337, 344; austriaca 336, 340, 341; calci- trapa 334, 338, 339, 345; calci- trapoides 338; cineraria 337, 344; consimilis 340; cyanus 337, 344, 345; var. denudata 344; diffusa 335, 340, 345; diluta Index to Volume 74 539 332-334, 339; dubia 337, 343, 345; ssp. eu-dubia 343; erio- phora 332; iberica 334, 338; jacea 335, 340-342, 345; var. nigra | 340; var. transal- pina 343; macrocephala 387, 343; maculosa 335, 339, 340, 345, 394; ssp. maculosa 339; ssp. micranthos 339; ssp. rhe- nana 339; melitensis 333, 358; montana 337, 344; moschata 337, 344; muricata 334, 339; nervosa 332, 336, 340, 341; nigra 336, 340-342, 345; f. pal- lens 342; nigrescens 337, 342, 343, 345; var. dubia 343; panic- ulata 335, 339, 340; picris 344; phrygia, ssp. austriaca 341; X pratensis 336, 340-342, 345; repens 337, 344, 345; rothrockii 337, 344; salmantica 333, 334, 339; scabiosa 337, 344; sicula 338; solstitialis 150, 334, 338, 345; squarrosa 340; stoebe 339; sulphurea 334, 338; transalpina 342, 343; virgata, var. squar- rosa 335, 340; vochinensis 342 Centunculus minimus 384 Ceramium diaphanum 480; fasti- giatum 480; rubriforme 480; rubrum 269, 422, 480; strictum 422 Cerastium arvense 519, 520; glo- meratum 381; maureri 520; rigoi 520; tomentosum 519, 520; Linn. and C. Arvense Linn., Spontaneous Hybrids be- tween 519 Chaenactis douglasii 394 Chaenorrhinum minus 394 Chaetomorpha area 266, 412; atrovirens 412, 413; linum 412, 413, 479; melagonium 266, 413, A477 Chaetopteris plumosa 420 540 Rhodora Chantransia collopoda 89, 92; mediterranea 92, 93; minutis- sima 92, 93; moniliformis 92; secundata 88; trifila 92, 93 Chaptalia ruminata 499 Cheilanthes Fendleri 82 Chloris virgata 389 Chondrus crispus 411, 415, 419, 422, 423, 480, 481, 483 Chorda filum 129, 417, 428, 480; tomentosum 417 Chordaria flagelliformis 267, 297, 417 Chorispora tenella 391 Chromosome Numbers from some Asteraceae, Somatic 102 Chrysopsis fulcrata 102, 104, 105; hispida 105; mariana 152; In New England 152; scabra 511; villosa 105 Chrysothamnus nauseosus, ssp. nauseosus 105 Cinna arundinacea 389 Cladonia alpestris 261; apodo- carpa 256; bacillaris 255; f. attenuata 255; f. bacillaris 255; f. fruticaulescens 255; f. ob- tusa 255; f. peritheta 255; beaumontii 259; bonyi 261; f. lacrimosa 261; f. prolifera 261; caespiticia 259; f. epiphylla 259; carassensis 259; f. digres- sa 259; f. obliqua, m. specta- bilis 260; f. regularis 259; f. spectabilis 259; f. subirregu- laris 260; f. subregularis 259; f. subregularis m. spectabilis 260; capitata 256; f. capitata 256; cariosa 257; f. squa- mulosa 257; caroliniana 261; cenotea 261; f. cenotea 261; f. exaltata 261; chlorophaea 258; f. costata 258; f. pro- lifera 258; f. simplex 258; clavulifera 256; coccifera 255; var. frondescens 255; var. [Vol. 74 stemmatina 255; coniocraea 258; f. actinota 258; f. cera- todes 258; f. peritheta 258; f. phyllostrota 258; f. pycnothe- liza 258; f. truncata 258; crispata, f. crispata 261; f. elegans 261; f. infundibulifera 261; f. subascypha 260; var. dilacerata 260; var. divulsa 261; var. virgata, f. kairamoi 261; cristatella 255; f. auran- tiaca 255; f. cristatella 256; f. lepidifera 256; f. ochrocarpa 256; f. pleurocarpa 256; f. ramosa 256; f. squamosissima 256; f. squamulosa 256; f. ves- tita 256; deformis, f. crenulata 255; didyma, var. muscigera 256; fimbriata, f. conista 258; f. pycnotheliza 258; f. simplex 258; f. stenoscypha 258; floe- rheana, f. carcata 255; f. floe- rheana 255; f. intermedia 255; f. squamosissima 255; var. in- termedia, f. fastigiata 255; f. peritheta 255; floridana 259; furcata 259; f. corymbosa 259; f. fissa 259; f. foliolosa 259; f. furcata 259; f. rigidula 259; f. subclausa 259; f. subulata 259; f. turgida 259; var. furcata 259; var. palamaea 259; var. paradoxa 259; var. pinnata, f. recurva 259; glauca 261; gra- cilis, var. dilitata 257; f. dila- cerata 257; var. elongata, f. elongata 257; grayi 258; f. cen- tralis 258; f. clavata 258; f. cyathiformis 258; f. epiphylea 258; f. foliosa 258; f. pallida 258; f. peritheta 258; f. proli- fera 258; f. squamulosa 258; impexa, f. impexa 261; f. proli- fera 261; f. subpellucida 261; incrassata 255; f. clavata 255; f. squamulosa 255; leporina 1972] 256; macilenta 256; f. maci- lenta 256; mateocyatha 257; f. squamulata 257; mitis 262; f. divaricata 262; f. pallida 262; f. prolifera 262; f. prostrata 262; soralifera 262; f. tenuis 262; multiformis 259; nem- oxyna 258; f. ambigua 258; nemoxyna 258; f. phyllocephala 258; f. scyphifera 258; papil- laria 255; f. molariformis 255; f. papillaria 255; f. stipata 255; parasitica 259; f. congesta 259; f. parasitica 259; f. sterilia 259; piedmontensis 256; f. in- termedia 256; f. lepidifera 256; f. obconica 256; f. phyllocoma 256; f. piedmontensis 256; f. squamossisima 256; pityrea, var. pityrea 257; f. cladomor- pha 257; f. pityrea 257; f. scyphifera 257; f. squamulifera 257; f. subacuta 257; pleurota 255; poculifera 262; f. simplex 262; polycarpoides 256; f. epi- phylla 256; f. polycarpoides 257; f. squamulosa 257; pseudo- phyxidata 262; f. clavata 262; f. lateralis 262; f. prolifera 262; f. simplex 262; f. squamu- losa 262; pyxidata 257; f. centralis 257; f. homodactyla 258; f. lophyra 258; f. margin- alis 257; f. prolifera 257; var. pyxidata 257; pyxioides, f. simplex 262; rangiferina 262; f. incrassata 262; f. tenuior 262; robbinsii, f. squamulosa 250; scabriuscula 259; f. ad- spersa 259; f. elegans 259; f. farinacea 259; f. scabriuscula 259; f. squamulosa 259; simu- lata 257; spumosa 262; squa- mosa 260; carneopallida 260; f. denticollis 260; f. fascicularis 260; f. laevicorticata 260; f. Index to Volume 74 541 muricella 260; f. pityrea 260; f. pseudocrispata 260; f. squa- mosa 260; f. subtrachynella 260; f. turfacea 260; f. ven- tricosa 260; var. laevicorticata, f. degenerascens 260; var. muricella, f. sarmentosa 260; var. phyllocoma 260; strepsilis 256; f. abortiva 256; f. coral- loidea 256; f. glabrata 256; f. strepsilis 256; subsquamosa, f. degenerata 260; f. luxurians 260; f. scabrida 260; f. sublevis 260; f. subrigida 260; subtenuis 261; sylvatica 262; f. arbuscula 262; f. congesta 262; m. fissa 262; f. decumbens 262; f. graci- lior 262; f. pygmaea 262; f. scabrida 262; f. sphagnoides 262; m. sorediata 262; tenuis 261; f. prolifera 261; f. pro- strata 261; terrae-novi 261; turgida 258; f. conspicua 258; uncialis, f. turgescans 261; f. humilior 261; f. obtusata 261; f. setigera 261; f. soraligera 261; f. subobtusata 261; f. un- cialis 261; verticillata 257; f. aggregata 257; f. apoticta 257; f. pallida 257; f. phyllo- cephala 257; f. verticillata 257; var. cervicornis, f. phyl- lophora 257 Cladoniae of Southeastern Mas- sachusetts with Special Refer- ence to the Collection of Charles Albert Robbins, List of the 254 Cladophora gracilis 266; holsat- ica 130; liniformis 280; re- fracta 266; rupestris 266; seri- cea 409, 411, 413, 415, 421 Clathromorphum circumscriptum 422 Cleaveland Herbarium of Bow- doin College, The Parker 80 542 Cnemidaria decurrens 442 Codiolum pusillum 413 Coleman, James R., Nomencla- tural Clarification of Two Spe- cies of Verbesina (Compo- positae) Endemic to Florida 97 Collinsia parviflora 393 Colt, L. C., Jr., Amphiprora or- nata Bailey — A Second Sta- tion in New England 521 Compositae, Nomenclatural and Taxonomic Notes on Mexican 495 Conium maculatum 397 Conopholis americana 141 Conyza lyrata 500; var. 500; viscosa 500 Corallina officinalis 269, 411, 422, 423 Corallorhiza maculata 141 Coriaria atropurpurea 249; (Cori- ariaceae) in the Western Hem- isphere, The Genus 242; cune- ifolia 250; japonica 246; mi- crophylla 249, 250; myrtifolia 244, 246; nepalensis 246; phyli- cifolia 249; ruscifolia 245, 246, 248; ssp. microphylla 245, 249; ssp. ruscifolia 246, 248; ter- minalis 245, 247; thymifolia 249, 250 Crepis capillaris 394; tectorum 394 Critonia hebebotrya 503 Croton capitatus 382; glandulo- sus, var. septentrionalis 382; monanthogynus 382; texensis 144 Cruise, J. E. and P. M. Catling, A White-flowered Form of Iris lacustris from Ontario 271 Cyanus segetum 344 Cyathea panamensis 450; pubes- cens 445; Steyermarkii 448- 450 pilosa Rhodora [Vol. 74 Cyatheaceae, New Species of American 441 Cyclodium meniscioides 447 Cymopterus Fendleri 83 Cynosciadium digitatum 148 Cyperus diffusus 328; esculentus 143; engelmannii 142; erythro- rhizos 142; Fendlerianus 83; ferruginescens 143; Houghtonii 294; iria 142; lancastriensis 142; strigosus 143, 294 Cystoclonium purpureum 269; var. cirrhosum 422 Daphnidostylus Fendleriana 83 Decachaeta ovatifolia 504, 506 Dermatolithon pustulatum 422 Desmarestia aculeata 129, 267, 417; viridis 267, 411, 417 Desmodium marilandicum 292 Diapensia lapponica 359, 361, 363-365, 367, 368, 370, 371, 373-375, 528; Habitat in the White Mountains of New Hampshire, Snow Cover and 358 Dictyosiphon foeniculaceus 267, 417, 428 Diplopappus ericoides, var. hir- tella 84 Disporum lanuginosum 523, 524 Distichlis spicata 411, 476 Dodecatheon frenchii 524; radi- catum 83 Draba lanceolata 381; neomex- icana 83 Dumontia incrassata 269, 423, 480, 481 Duncan, Wilbur H. and Marie Mellinger, Edgeworthia (Thy- melaceae) New to the Western Hemisphere 436 Dunn, David B., Lupinus mexi- canus Cerv. ex Lag. 489 Dysodium divaricatum 12, 175, 176; radiatum 176 162, 1972] Eastman, Lesley M., A Second Occurrence for Triphora trian- thophora (SW.) Rydb. in Maine 141 Eaton, Richard, Chrysopsis ma- riana in Nw England 152 Eclipta pusilla 60 Ectocarpus confervoides 266, 417; var. dasycarpa 481; siliculosis 417, 418; tomentosus 421 Edgeworthia albiflora 438; chry- santha 436-438; gardneri 438; longipes 438; papyrifera 437; (Thymelaeaceae) New to the Western Hemisphere 436; to- mentosa 437 Egeria densa 135, 140 Elachistea fucicola 267, 418, 428; lubrica 297 Elaeagnus angustifolius 148; um- bellatus 148 Eleocharis quadrangulata, crassior 378 Elephantopus carolinianus 103 Eleutheranthera ruderalis 206; divaricata 177 Elodea Nuttallii 136, 137 Elyna spicata 325 Empetrum nigrum 367 Enteromorpha ahleriana 479, 482; clathrata 265, 479; com- pressa 265, 413; erecta 265, 411, 413, 414; flexuosa, ssp. flexuosa 429; ssp. paradoxa 482; ssp. pilifera 479, 480; in- testinalis 265, 409, 411, 413, 479-481; linza 265, 414, 479; var, oblanceolata 479, 480; marginata 265, 412; minima 266, 412; plumosa 414; pro- lifera 266, 414 Epifagus virginiana 141 Equisetum from Peninsula Flor- ida 525 Equisetum hyemale 324, 527; var. affine 526, 527; var. ela- var. Index to Volume 74 543 tum 526; palustre 389; praeal- tum 526; robustum 526; scir- poides 389 Eragrostis spectabilis 389 Erigeron affine 500; alamosanum 501; canbyi 395; cinereus 83; longipes 500; macranthus 106; scaposum 500; Jf latifolium 500, 501; speciosus 106; var. macranthus 106; tenellum 501; velutipes 501 Eriogonum atrorubens 351; gor- donii 391; harperi 154; Good- man in Tennessee 154; tri- chopes 391 Erysimum repandum 391 Erythrotrichia carnea 298 Euonymous europaea 321; for- tunei 147 Eupatorium albicaule 501, 502; var. laxius 501, 502; bertholdii 504; coelestinum 103; collinum 503; var. collinum 502; var. mendezii 503; aff. eximium 273; Fendleri 83; hebebotryum 503; leucoderme 501; mendezii 503; microcephalum 505, 506; neaeanum 503; nigrescens 503; ovaliflorum 504; ovatifolium 504; parasiticum 272; petraeum 504, 506; polybotryum 504-506; scabrellum 505; stillingiaefo- lium 502, 503; tepicanum 503; ymalense 501, 502 Euphorbia Fendleri 83; stricto- spora 392 Eustoma grandiflorum, f. album 385 Euthora cristata 269, 423 Evans, K., see Mohlenbrock, Robert H. Fagus grandifolia 524 Fimbristylis autumnalis 294 Fosliella lejolisii 424 544 Fralick, Richard A., see Mathie- son, A. C. Franseria tomentosa 394 Fucus distichus, ssp. distichus 409, 418; ssp. edentatus 418; ssp. evanescens 418, 477; 480, 483; edentatus 268; evanescens 268; filiformis 268; spiralis 268; vesiculosis 268, 295, 298, 409, 411, 418, 419, 480, 481, 483; var. spiralis 411, 418, 482 Fundy, On the Marine Algae of Kent Island, Bay of 264 Galeana pratensis 205 Galium verum 394 Galpinsia Fendleri 83 Garay, Leslie A., In Memoriam Charles Schweinfurth, 1890- 1970 71 Gaura parviflora 392 Giffordia granulosa 266, 418 Gigartina stellata 269, 415, 423 Glaux maritima 384 Gloiosiphonia capillaris 423 Glyceria striata 148 Gnaphalium berlandieri — 508; canescens 506; chilense 508; de- currens 509; gracile 509, 510; hirtum 508, 509; inornatum 507; leptophyllum 509; leuco- cephalum 509; macounii 509; pedunculosum 507, 508; pur- pureum 508; roseum 506; spha- cilatum 507; sprengelii 508; stramineum 508; tenue 509; viscosum 508, 509; Wrightii 506 Gomphonema olivaceum 481 Gracilaria foliifera 480; verru- cosa 480 Graptopetalum rusbyi 312 Great Plains, New and Interest- ing Plants from the 378 Grindelia aphanactis 106 Haber, Erich, Priority of the Bi- Rhodora [Vol. 74 nomial Pyrola chlorantha 396 Halenia deflexa 393 Halosaccion ramentaceum 269 Haplopappus clementis 107; exi- mius 106; laceratus 107; lyalli 102, 106, 107; peirsonii 106; pygmaeus 107 Harriman, Neil A., Records on the Flora of Wisconsin 156 Hebeclinium tepicanum 503, 504 Hehre, Edw. J., Lomentaria clavellosa (Turner) Gaillon. An Addition to the Marine Algal Flora of New Hampshire 158; A. R. Hodgdon and R. B. Pike, The Parker Cleaveland Her- barium of Bowdoin College 80 Helianthus mollis 388 Hellquist, C. Barre, Range Ex- tensions of Vascular Aquatic Plants in New England 131 Hemibaccharis hieraciodes 498 Hemistegia decurrens 442 Hemitelia conformis 450; petio- lata 450; Woronovii 450 Hemizonia minima 205; durandi 204 Hess, William J., see Reveal, James Heterocladus caracasanus 244, 249 Heterophylleia caracasana 249 Heterotheca inuloides, var. rosei 510; leptoglossa 510 Hibiscus lasiocarpa 384; syria- cus 145; trionum 150 Hidalgoa ternata 204, 206 Hildenbrandia prototypus 269, 411, 423, 480 Hodgdon, A. R. and Frederic L. Steele, Recent Discoveries in the New Hampshire Flora 291; see also Hehre, Edw. J. Hoffmannseggella, A New Spe- cies of the Orchid Genus 283 Hoffmannseggella brevicaulis 284, 1972] 285; cinnabarina 283; harpo- phylla 284, 285; Kautskyi 285 Hookeria acutifolia 524 Hydrophyllum appendiculatum 385; Fendleri 83 Hymenopappus tenuifolius 395 Hymenoxys richardsonii 109; var. floribunda 109 Hypericum Arnoldianum 277; X Arnoldianum 276-280; X Daw- sonianum 279, 280; densiflorum 146, 278-281; denticulatum 146, 147; dolabriforme 146, 147; frondosum 281; galioides 276- 278; Kalmianum 280, 281; lobo- carpum 146, 276-280; X no- thum 280, 281; perforatum 392; prolificum 279-281; punctatum, var. pseudomaculatum 145, 146; var. punctatum 146; Sect. Myriandra, An Evaluation of Some Putative Spontaneous Garden Hybrids In 276; spathu- latum 146; sphaerocarpum, var. sphaerocarpum 147; var. turgi- dum 146, 147; X Van Fleetii 281 Hypochoeris glabra 149; radicata 149 Hypoxis hirsuta 293 Illinois Field Studies 142 Ipswich, Massachusetts, The Ecol- ogy of Benthic Salt Marsh Algae at, I. Zonation and dis- tribution of Algal Species 475 Iris lacustris 271; From Ontario, A White-flowered Form of 271; f. albiflora 271; missouriensis 390 Isthmoplea sphaerophora 298 Iva axillaris 387; frutescens, var. oraria 291 and Herbarium Jaegeria hirta 170 Index to Volume 74 545 Johnson, Miles F., Tussilago far- fara L. (Senecioneae-Astera- ceae) in Virginia 403 Jones, H. G., A New Species of the Orchid Genus Hoffmann- seggella 283 Juglans nigra 390 Juncus balticus, var. littoralis 478; gerardi 476-478, 483, 485; trifidus 367, 368, 370, 371, 373, 3875; vaseyi 390 Kansas, The Yellow Mandarin, A Notable Addition to the Flora of 523 Kiger, Robert W., A New Vari- ety of Leucophyllum laeviga- tum (Scrophulariaceae) from Mexico 347 King, R. M. and H. Robinson, Neomirandia allenii A New Epiphytic Composite of the American Rain Forest 272 Knapweeds (Centaurea) in Can- ada and the United States, Dis- tribution of Native and Intro- duced 331 Kobresia elachycarpa 329; sim- pliuscula 329 Koetzner, Kenneth and R. D. Wood, On the Marine Algae of Kent Island, Bay of Fundy 264 Kornmannia leptoderma 479-481 Kovanda, Miloslav, Somatic Chromosome Numbers for some Asteraceae 102 Kylinia collopoda 85, 91, 93, 96; compacta 85, 89, 93, 96; monili- formis 85, 93, 96; scapae 92, 98; secundata 268, 473 Laelia anceps 283 Laminaria agardhii 267; digitata 127, 129, 130, 268, 411, 419; intermedia 268; longicruris 546 268; platymeris 268; Sinclairii 127; saccharina 129, 411, 419, 421, 480, 483; stenophylla 268 Lamium amplexicaule, f. albiflo- rum 386 Lavatera cretica 392 Lathyrus aphaca 155; L. New to Tennessee and the South- west 155 Leathesia difformis 419 Lecanora chlarotera 478 Lechea tenuifolia 293; villosa 293 Ledum groenlandicum 529 Leonurus Marrubiastrum 386 Lespedeza virginica 292 Lesquerella gordonii 381; lasio- carpa 77, 79; var. ampla 78; ssp. berlandieri 78, 79; var. heterochroma 78; var. hispida 78; var. berlandieri 78, 79; in- termedia 83; var. hispida 79; ssp. lasiocarpa 77; Nomenclatu- ral Changes in 76 Leucantha cyanus 344 Leucophylum laevigatum 347, 348; (Scrophulariaceae) From Mexico, A New Variety of 347; var. coahuilense 347-349; var. laevigatum 349 Liatris punctata 103; var. ne- braskana 104; var. mexicana 104; var. punctata 104; var. typica 104 Linaria dalmatica 156 Liquidambar, Chromosome Num- bers in 287; formosana 288, 290; orientalis 287, 290; sty- raciflua 287, 288, 290, 524; X L. formosana 289; X L. ori- entalis 289 Lithoderma extensum 419 Lithophyllum corallinae pustulatum 423 Lithothamnion glaciale 423 156, 423; Rhodora [Vol. 74 Lobelia siphilitica, var. siphilitica, f. albiflora 387 Loiseleuria procumbens 365, 367, 368, 370, 373, 374 Lomentaria clavellosa 158; (Turner) Gaillon: An Addi- tion to the Marine Algal Flora of New Hampshire 158 Lotus corniculatus 150, 392 Lupinus ashenbornii 490, 494; bilineatus 491, 492; hartwegii 494; mexicanus 489-494; Cerv. ex Lag. 489; persistens 494; termis 491 Lychnis alba 149; 391 Lysimachia terrestris 150 Lycium pallidum 83 Lycopodium lucidulum 524; se- lago 367; tristachyum 353 Lythrum salicaria 393 chalcedonica McGregor, Ronald L., Lawrence K. Magrath and Ronald Wee- don, New and Interesting Plants from the Great Plains 378 Macrocystis pyrifera 130 McVaugh, Rogers, Nomenclatu- ral and Taxonomic Notes on Mexican Compositae 495 Madia minima 204, 205 Magnolia tripetala 524 Magrath, Lawrence K., see Mc- Gregor, Ronald Mandarin, The Yellow, A Notable Addition to the Flora of Kan- sas 523 Mantisalea salmantica 339 Mapania macrophylla 327; syl- vatica 328 Marchand, Peter, Wind and the Winter-exposed Plant 528 Mathieson, Arthur C., Emery Swan and Richard A. Fralick, An Abnormal Specimen of 1972] Laminaria digitata (L.) La- mouroux 127; and Richard A. Fralick, Investigations of New England Algae V. The Algal Vegetation of the Hampton- Seabrook Estuary and the Open Coast Near Hampton, New Hampshire 406 Melampodium achillaeoides 203; americanum 2, 6, 9, 12, 15, 16, 20-23, 31, 210; angulatum 202; angustifolium 21, 206; anoma- lum 203; appendiculatum 7, 18, 56, 61, 210; var. leiocarpum 61; var. sonorense 61; areni- cola 64, 65; argophyllum 6, 9, 13, 35, 40, 41, 45, 46, 210; ar- vense 188; aureum 7, 9, 12, 17, 191, 193, 211; australe 203; baranquillae 203; berterianum 176; bibracteatum 7, 14, 59, 187, 210; bonairense 203; brachyglossum 170; camphora- tum 4, 204; cinerascens 44; cinereum 6, 9, 35, 39, 41, 44, 46, 172, 210; var. cinereum 6, 14, 39, 40, 210; var. hirtellum 6, 14, 40, 42, 210; var. ramosissimum 6, 14, 40, 43, 210; (Compositae; Heliantheae), Revision of the Genus 1, 161; connatum 195; copiosum 176; coronopifolium 51, 53; cornutum 184, 185; cos- taricense 7, 18, 181, 183, 210; cupulatum 7, 17, 20, 56, 59, 60, 64, 65, 210; dichothomum 202; dicoelocarpum 4, 7, 10, 18, 30, 172, 182, 184, 210; digynum 204; diffusum 6, 8, 15, 25, 27, 210; var. lancelatum 26; divari- catum 5, 7, 8, 10, 12, 18, 64, 69, 172, 175, 178, 195, 202, 210; var. macranthum 176; diversifolium 202; dombeyanum 204; durandi 204; flaccidum 176; geminatum 204; glabri- bracteatum 7, 16, 23, 67, 210; Index to Volume 74 547 glabrum 7, 11, 19, 30, 199-201, 211; gracile 7, 14, 20, 164-166, 168, 210; var. oblongifolium 164; heterophyllum 21; hidal- goa 204; hirsutum 205; hispi- dum 20, 47, 53; humile 205; kunthianum 22; lanceolatum 25, 167; laxum 169, leucanthum 5, 6, 9, 11, 13, 34-36, 40, 41, 46, 172, 210; var. argophyllum 45; liebmanii 190; linearilobum 6, 11, 15, 21, 31-33, 202, 210; longicorne 6, 19, 20, 54, 56, 210; longifolium 7, 11, 12, 19, 27, 161, 210; longipes 6, 16, 29-32, 202, 210; longipilum 7, 11, 18, 20, 27, 68, 69, 185, 210; manil- lense 25; microcarpum 164; mi- crocephalum 7, 15, 20, 165-168, 210; mimulifolium 7, 20, 27, 163, 164, 210; minimum 205; minutiflorum 205; montanum 4, 7, 9, 12, 189, 190, 193, 202, 211; var montanum 7, 17, 190, 191, 211; var. viridulum 7, 17, 191, 211; nayaritense 6, 15, 58, 59, 210; nelsonii 22; nutans 7, 15, 30, 200, 201, 211; oblongi- folium 164; ovatifolium 176; paludicola 205; paludosum 176; panamense 177; paniculatum 7, 14, 166, 168-172, 210; parvulum 162; perfoliatum 3, 7, 8, 12, 18, 194-197, 200-202, 210; pilosum 6, 15, 27, 28, 210; pinnatum 202; pringlei 6, 16, 20, 23, 50, 51, 210; pumilum 176; ramosis- simum 39, 43; repens 7, 14, 59, 188, 211; rhomboideum 162; rhombifolium 177, 202; rosei 7, 18, 59, 64, 65, 67, 210; var. sub- integrum 64; ruderale 205; sericeum 6, 16, 20-22, 46-48, 51, 53, 210; var. brevipes 47; var. exappendiculatum 47; var. longipes 29; sinaloense 7, 19, 52, 186, 211; sinuatum 7, 9, 17, 548 Rhodora 20, 56, 59, 63, 64, 210; strigo- sum 6, 19, 51, 52, 55, 56, 210; suffruticosum 206; tenellum 7, 17, 59, 66, 67, 69, 210; var. flaccidum 176; tepicense 7, 19, 52, 175, 184, 185, 211; terna- tum 204; villicaule 68 Melampodium canescens 31, 32 Mellinger, Marie, see Duncan, Wilbur H. Melobesia lejolisii 424 Melosira juergensii 480 Membranoptera alata 424 Mendezia bicolor 515 Mertensia Fendleri 83 Microcoleus tenerrimus 484 Microlonchus salmanticus 339 Mimulus alatus, f. albiflorus 386 Mish, Lawrence B., List of the Cladoniae of Southeastern Mas- sachusetts with Special Refer- ence to the Collection of Charles Albert Robbins 254 Mitchella repens 437 Mohlenbrock, Robert H. and Dan K. Evans, Illinois Field and Herbarium Studies 142 Moore, Mary L, New Form of Thuja occidentalis Resembling Known Cultivars 352 Moore, R. J., Distribution of Na- tive and Introduced Knapweeds (Centaurea) in Canada and the United States 331 Monostroma fuscum 414; f. blytii 266; grevillei 414; leptodermum 414; oxyspermum 411, 414, 427, 428, 482, 485-487; pulch- rum 297, 414 Montanoa affinis 510, 511; graci- lis 510, 511; karvinskii 510, 511; olivae 510, 511; subtrun- cata 510, 511 Morton, J. K., Spontaneous Hy- brids between Cerastium to- [Vol. 74 mentosum Linn. and C. arvense Linn. 519 Myriophyllum alterniflorum 139; exalbescens 150; heterophyllum 139; humile 136; spicatum, var. exalbescens 132; verticillatum, var. pectinatum 137 Najas flexilis 133 Nemalion helminthoides 424 Neomirandea allenii 273, 274; A New Epiphytic Composite of the American Rain Forest 272; angularis 272; araliaefolia 272; costaricencis 273; eximia 274; sciaphila 272, 274 Nephelea pubescens 445 New Hampshire, The Algal Veg- etation of the Hampton-Sea- brook Estuary and the Open Coast near Hampton 406 New Hampshire Flora, Recent Discoveries in the 291 New Hampshire, Snow Cover and the Diapensia lapponica Habi- tat in the White Mountains of 358 Nigella damascena 143 North Dakota Plants, New Rec- ords of 389 Nothites ovatifolia 504-506 Nuphar variegatum 137 Nymphaea alba, f. rosea 188, 139; odorata 137, 138; var. gigantea 138; f. rubra 139; tuberosa 135- 139 Ontario, À White-flowered Form of Iris lacustris from 271 Ophryosporus ovatifolius 504- 506; petraeus 504, 505; scabrel- lus 505 Oreocarya fulvocanescens 83; multicaulis 83 Orobanche multiflora 387 1972] Oryzopsis asperifolia 271; pun- gens 271 Osmorhiza claytoni 393 Oxypappus scaber 511 Panicum philadelphicum 294, 378; virgatum 478, 483 Paronychia depressa 380 Paspalum distichum 378 Pectis seemannii 511 Pellaea atropurpurea 220-225, 228-233, 237-240; Spores, Chromosomes and Relations of the Fern 220; longimucronata 232; notabilis 220-225, 229-234, 237-240; ovata 240; ternifolia 220, 226, 234, 237, 239; var. ternifolia 221, 224, 226, 227, 229-232, 234, 235, 2371-239; var. Wrightiana 221, 224, 226-228, 232, 234-239; truncata 232, 239 Peltandra undulata 516-518; vir- ginica 517, 518; (L.) Schott & Endlicher, The Combination 516 Penstemon procera 393 Percursaria percursa 411, 415, 421, 482, 485 Percome caudata 109; glandulosa 109 Petalonia fascia 409, 419, 420, 480, 481 Petrocelis middendorfii 424 Peyssonelia rosenvingii 424 Phacelia constancei 451-454; cor- rugata 456, 459, 468; crenulata 451; Group (Hydrophyllaceae), New Species in the 451; how- elliana 456, 460, 461; integri- folia 464; palmeri 451; pinnati- fida 464; rafaelensis 454, 456- 458; utahensis 454, 468; vossii 462-464; welshii 465-468 Phlox alyssifolia 393 Phoradendron juniperinus 83 Phryma leptostachya 292 Index to Volume 74 549 Phycodrys rubens 270, 424, 477 Phyllophora brodiaei 269, 424; membranifolia 424 Phymatolithon compactum 422; laevigatum 424, 427; lenor- mandi 425 Physalis Fendleri 83; pendula 150; similas 83 Physostegia formosior 393; parvi- flora 393; virginiana 393 Pike, R. B., see Hehre, Edw. Pilaiella littoralis 419 see also Pylaiella Pinus banksiana 389; leiophylla 350; ponderosa 350 Plumaria elegans 425, 428 Poa alpigena 367; Bigelovii 83; Fedleriana 82 Polygala senega 392 Polygonum sawatchense 380 Polyides rotundus 411, 425 Polymnia perfoliata 194; revo- luta 195 Polypogon monspeliense 389 Polysiphonia denudata 425, 480; elongata 425; fibrillosa 425, 427, 428; harveyi 270; lanosa 270, 298, 425, 480; nigra 425, 480; nigrescens 270, 425, 480; novae-angliae 426; subtilissima 426; urceolata 270, 426, 481 Porphyra miniata 426; umbili- calis 268, 411, 426; f. epiphy- tica 426 Potamogeton Berchtoldii 135; var. tenuissimus 135; capillaceus 135; confervoides 135, 138, 139; crispus 132; epihydrus, var. ramosus 135, 136; filiformis 131; var. borealis 131; var. Macounii 131; Friesii 133; gemmiparus 135; gramineus, var. gramineus 135; var. my- riophyllus 135; Hillii 134; longiligulatus 133, 134; natans 135; pectinatus 133; praelongus 550 137; Spirillus 135; strictifo- lius, var. rutiloides 133; zosteri- formis 134, 137 Potentilla palustris 391; propin- qua 83; tridentata 367, 368, 370, 374 Prasiola stipitata 266 Prenanthes boottii 367 Protoderma marinum 415 Prunus serotina 321 Pseudendoclonium marinum 409, 415, 478, 485-487 Pseudolithodermum 419, 427 Pterophyton heterophyllum 99, 100; pauciflorum 99 Ptilimnium costatum 148 Ptilota serrata 269, 426 extensum Punctaria latifolia 267, 298; plantaginea 298 Pyenanthemum incanum 292; tenuifolium 292; Torrei 292; verticillatum 292; virginianum 292 Pylaiella littoralis 266, 298 Pyrola chlorantha 396, 397; Priority of the Binomial 396; virens 396, 397 Quercus ellipsoidalis 390; nove- mexicana $83; palustris 148; reticulatus 350; robur 529 Ralfsia borneti 419, 420, 428; clavata 420, 428, 479-481, 486; fungiformis 267, 420; verrucosa 409, 420, 479-481; 486 Ranunculus cardiophyllus 391; carolinianus 149; repens, var. pleniflorus 148; septentrionalis 149 Redfearn, Paul L., Jr., The Yel- low Mandarin, A Notable Addi- tion to the Flora of Kansas 523 Reveal, James L. and William J. Rhodora [Vol. 74 Hess, Variation within Calo- chortus venustulus (Liliaceae) 350 Reynolds, Don R. and Ken Cal- houn, Equisetum from Penin- sula Florida 426 Rhamnus davurica 392 Rhapidophyllum hystrix 525; in Mississsippi 525 Rhizoclonium implexum 482, 484, 485; riparium 411, 415, 484; f. polyrhizum 482; f. riparium 482; f. validum 485; tortuosum 266, 415 Rhodochorton purpureum 426 Rhododendron lapponicum 367, 368, 370, 374 Rhododermis elegans 426; Rho- domela confervoides 270, 426 Rhodymenia palmata 269, 427 Rhodophyllis dichotoma 427 Rhodophysema elegans 426, 427 Ribes cereum 391 Robinia viscosa 321 Robbins, Charles Albert, List of the Cladoniae of Southeastern Massachusetts with Special Reference to the Collection of 254 Robinson, H., see King, R. M. Rogers, Ken E., see Watson, J. Ray, Jr. Rollins, Reed C. and E. A. Shaw, Nomenclatural Changes in Les- querella 76 Rorippa islandica, var. fernald- iana 144; var. islandica 144; sessiliflora 144 Rubus idaeus, var. strigosus 144; occidentalis 144; procerus 144 Rudbeckia hirta 108; var. angus- tifolia 108; var. corymbifera 108; var. floridana 108; var. hirta 108; var. pulcherrima 108; serotina 109 Rumex stenophyllus 379 1972] Ruppia maritima 479 Sabatia campestris, f. albiflora 385 Saccorhiza dermatodea 420; poly- Schides 129 Salix Fendleriana 83; irrorata 83; pedicellata 391; uva-ursi 367, 370, 374 Sambucus pubens 394 Santamour, Frank J., Jr., Chro- mosome Numbers in Liquidam- bar 287 Sargassum fluitans 295; natans 298 Saxifraga occidentalis, var. occi- dentalis 382 Sciaromium lescurii 524 Scirpus americanus 476, 483; cespitosus, var. callosus 367; debilis 398; maritimus, var. Fernaldii 291; olneyi 329; purshianus 398, 400, 401; and S. smithii, Chromosome Num- bers of 398; var. purshianus 398, 399, 401; f. williamsii 398, 399, 401; robustus 291; smithii 398, 400, 401; f. setosus 398, 399, 401; f. levisetus 398, 399, 401; f. smithii 398, 399, 401 Schizothrix calcicola 487 Schoenoxiphium kobresioides 327; rufum 323 Schuyler, Alfred E., Chromo- some Numbers of Scirpus purshianus and Scirpus smithii 398 Schweinfurth, Charles, In Memo- riam 71 Schwetschkeopsis fabronia 524 Sclerocarpus columbianus 203 Scutellaria brittonii 386; ovata, var. ovata 148, 149; var. rugosa 148, 149; var. versicolor 148, 149 Index to Volume 74 551 Scytosiphon lomentaria 267, 298, 409, 420, 479, 480 Sedum alamosamum 311; cocker- ellii 302-306, 309, 311, 312, 314; diffusum 311; glaucophyllum 312; griffithsii 302, 303, 308, 311; havardii 302, 308, 311, 320; in the Southwestern United States, Intraspecific Variation in Chromosomes of 301; lanceolatum 3811, 312; niveum 302, 305, 317; spathuli- folium 312; stelliforme 302, 305, 306, 308, 309, 311, 312, 318; ternatum 306, 312; wrightii 302, 305, 306, 308, 310, 311, 318 Selaginella Underwoodei 82 Senecio filifolius 111; longilobus 111; vulgaris 395 Shaw, Elizabeth A., see Rollins, Reed C. Sida spinosa 378, 384; f. albiflora 383 Sigesbeckia flosculosa 204 Skog, Laurence E., The Genus Coriaria (Coriariaceae) in the Western. Hemisphere 242 Smith, Earle C., Jr., Redefinition of Cedrela oaxacensis C. DC. & Rose 124 Spartina patens 411, 476, 483, 484; alterniflora 411, 413, 414, 417, 418; var. glabra 476, 482, 483 Sphacelaria plumosa 420, 427; radicans 420, 428 Sphaeralcea Fendleri 83 Sphaeropteris atahuallpa 442- 444; horrida 443; marginalis 442; sipapoensis 441, 442, 444 Sphaerotrichia divaricata 267 Sphagnum capillaceum, var. tene- rum 524 Spilanthes guatamalensis 177 552 Spongomorpha arcta 266, 297, 415; spinescens 266, 415 Spongonema tomentosum 421 Sporobolus airoides 390 Solanum sarrachoides 386 Solidago cutleri 367; flexicaulis 107; multiradiata 107; var. scopulorum 107; sempervirens 108, 478; var. mexicana 108; var. sempervirens 108 Sorapion kjellmani 420, 427 Stachys palustris 393; scopulo- rum 393 Steele, Frederic L., see Hodgdon, A, R. Stellaria aquatica 381 Stevens, O. A., New Records of North Dakota Plants 389 Stuessy, Tod F., Revision of the Genus Melampodium (Com- positae: Heliantheae) 1, 161 Suckleya suckleyana 380 Svenson, Henry K., The Rachilla in Cape Cod Species of Carex with Notes on the History of the Perigynium and Rachilla 321; Rediscovery of Tilia neg- lecta Spach 469 Swan, Emery F., see Mathieson, A. C. Synthlipsis berlandieri 78, 79; var. hispida 78, 79; hetero- chroma 78 Takhtajan, A., XII International Botanical Congress, 528 Thelesperma marginatum 395 Thuja occidentalis 352-354, 356; f. prostrata 354; Resembling Known Cultivars, A New Form of 352; var. ericoides 354 Thymelaea passerina 384 Tiffney, Wesley N., Jr, Snow Cover and the Diapensia lap- poncica Habitat in the White Rhodora [Vol. 74 Mountains of New Hampshire 358 Tilia americana 472, 473; var. heterophylla | 473; argentea 473; caroliniana 473; glabra 473; heterophylla 470, 472, 473; Michauxii 472, 473; neg- lecta 469-474; Spach, Redis- covery of 469; pubescens 472 Tolypothrix tenuis 485 Tragoceros americanus 511, 512; microglossus 511, 512; schie- deanum 203, 511, 512; zin- nioides 512 Tribulus cistoides 117, 120 Trichipteris cordata 446; cyclo- dium 446-449; Maguirei 447- 449; saggittifolia 446; Steyer- markii 446, 448; Williamsii 447, 449 Trifolium pratense, f. leucochra- ceum 150 Trigzonospermum adenostemmoi- des 513 Triodanis biflora 387 Triphora trianthophora 141; (Sw.) Rydb. in Maine, A Sec- ond Occurrence for 141 Triplasis purpurea 390 Tryon, Alice F., Spores, Chro- mosomes and Relations of the Fern Pellaea atropurpurea 220 Tryon, Rolla, Taxonomie Fern Notes, VI. New Species of American Cyatheaceae 441 Tussilago farfara 403, 404; L. (Senecioneae-Asteraceae) in Virginia 403 Uhl, Charles H., Intraspecific Variation in Chromosomes of. Sedum in the Southwestern United States 301 Ulmus procera 143 Ulothrix flacca 265, 415; flaccida 485 sub- 1972] Ulva gigantea 479; lactuca 266, 409, 411, 416; rigida 480 Uncinia uncinoides 323 Unxia camphorata 204, 205; digyna 204; hirsuta 205; suf- fruticosa 206 Urococcus foslieanus 487 Urospora collabens 416; penicilli- formis 416; speciosa 416 Utricularia geminiscapa 136 Vaccinium angustifolium 367; uliginosum 365, 367, 368, 370- 375; vitis-idaea 365, 367, 370, 374 Vallisneria americana 133 Vaucheria arcassonensis 484; compacta, var. koksoakensis 486; intermedia 482, 484, 485 Verbena hastata X urticifolia 150 Verbesina chapmanii 100; (Com- positae) Endemic to Florida, Nomenclatural Clarification of Two Species of 92; heterophyl- la 98, 99; pauciflora 99, 100; warei 98-100 Veronica barbinervis 513; chaca- lana 513; palmeri 513; triflos- culosa 513, 514 Veronicastrum virginicum 394 Vesicaria lasiocarpa 77 Viguiera blakei 514; multiflora 109; puruana 514 Villanova achillaeoides 203 Viola pedata 293 Virginia, Tussilago farfara L. (Senecioneae-Asteraceae) in 403 Index to Volume 74 553 Voluntarella muricata 339 Watson, J. Ray, Rhapidophyllum hystrix in Mississippi 525; and Ken E. Rogers, Eriogonum harperi Goodman in Tennessee 154 Webber, E. E. and R. T. Wilce, The Ecology of Benthic Salt- marsh Algae at Ipswich, Mas- sachusetts I. Zonation and Dis- tribution of Algal Species 475 Wedelia minor 175; ovatifolia 175; perfoliata 194 Weedon, Ronald R., see Mac- gregor Ronald L. Wilce, R. T., see Webber, E. E. Wind and the Winter-exposed Plant, 528 Wisconsin, Records on the Flora of 156 Woelkerling, Wm., Some Algal Invaders of the Northwestern Fringes of the Sargasso Sea 295; Studies on the Audouinella microscopica (Naeg.) Woelk. Complex (Rhodophyta) 85 Wolffia punctata 136; columbiana 136 Wood, R. D., see Koetzner, Ken- neth Xanthocephalum conoideum 514, 515; seriocarpum 514 Zarabellia rhomboidea 12, 162 Zinnia bicolor 515, elegans 515, 516; tenella 515; violacea 515, 516 Volume 74, No. 800, including pages 441-553, was issued December 29, 1972. 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, particularly 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- atics 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. vmm CONTENTS: — continued Lupinus mexicanus Cerv. ex. Lag. IEE AIMÉ AAE E E E E 489 Nomenclatural and Taxonomic Notes on Mexican Compos- itae. Rogers MeVOWughy EE E E E enotas 495 The Combination Peltandra virginica (L.) Schott & Endlicher. Will H. Blackwell, Jv. ..ccccccccccssssccssesssccsssseccccccessssascesssssssessesssnes 516 Spontaneous Hybrids Between Cerastium tomentosum Linn. and C. arvense Linn. JURE IVE OTE os. oo de ct taea Cerere evoss coe donsensseceacuscesdtwasnsviervetesseetess da 519 Amphiprora ornata Bailey — A Second Station in New Eng- land. Ts CS CH LOU ROTE EA S ONDE MC ante eee o t e teer 521 The Yellow Mandarin, A Notable Addition to the Flora of Arkansas. tuos DB Pd E BIT PAA I A EE A E a a UU: ERES 523 Rhapidophyllum hystrix in Mississippi. PANELA Ta 1010: O E E E a A E E E 525 Equisetum from Peninsula Florida. Don R. Reynolds and Ken Calhoun ................. eee 526 Wind and the Winter-exposed Plant. EEG EU ERO! USE CERRAR D E DEUM REC 528 XII International Botanical Congress. ELT E N a E T inii MM nL 532 ee E OEO EE aana ieser antaiaheuuiees 583 Index to Volume 74 oi..c.ccccccccccccccccceccssscccecsscscceseeeseetaceceesteceeceeeens 5387