Volume 51 =: 1970 JOURNAL OF THE ARNOLD ARBORETUM HARVARD UNIVERSITY Dates of Issue No. 1 (pp. 1-132) issued 16 January, 1970. No. 2 (pp. 133-256) issued 14 April, 1970 No. 3 (pp. 257-430) issued 13 July, 1970 No. 4 (pp. 431-561) issued 19 October, 1970 Contents of Volume 51 The systematics and Breeding System of Gelsemium ( Loganiaceae ). ROBERT ORNDUFF The Genera of Scans in the Seutheatiers Waited States, THOMAS S. ; : The Ecology —; an Elfin Forest i in Puerto Rico. 10. Notes on Two Species of Marcgravia. RICHARD A. HOWARD 11. The Leafy Hepaticae of Pico del Oeste. MARGARET FULFORD, BARBARA CRANDALL, and RAY STOTLER . Types of Blind Vein- -endings i in the Dichotomous Venation of Circaeaster. ADRIANCE S. FOSTER Comparative Morphological ‘Studies i in . Dilleniaceae, V. Leaf Anatomy. WILLIAM C. DICKISON . The Morphology and Germination of the Seed of Elephantorrhiza elephantina. H. P. VAN DER SCHIJFF and L. SNYMAN . A New Species of Ormosia from Brazil. G. P. YAKOVLEV . , A Revision of the Boraginaceae of West Pakistan and Kashmir. S. M. A. KAzZM The Genera of the = nee in ‘the Southeastern United States. NorTon G. MIL Notes Relating to on Floras of Norfolk and Lord Howe Islands. P. S. GREEN . The Ecology of an Elfin F orest i in Puerto Rico, 12. A New Species of Gonocalyx — Lorin I. NEVLING, JR. Analysis of the Complex Vascularity i in Stems of Dioscorea composita. Epwarp S. AYENSU . The Taxonomic Status of Protocyrtandra ( ape ig Haploid ner Diploid ‘Pollen j in Hypericum patulum JOAB re THOMAS . Studies in the Leguminosae, 11. A New Species of Derris from the Solomon Islands. Ki > PROTHATGRE 5 ea 1970 129 133 185 Statement of Ownership . The Genera of Acanthaceae i in the Southeastern United States. ROBERT W. LONG Floral Morphology and Vascular Anatomy of the Hamamelidaceae: The Apetalous Genera of Hamamelidoideae. A. LINN BOGLE . : A aa of the Boraginaceae of West Pakistan and Kashmir (continued). S. M. Comparative Morphological Studies in Dilleniaceae, VI. Stamens and Young Stem. WILLIAM C. DICKISON . Additional Notes on the Malesian Species of Zanthoxylum (Rutaceae ). THoMAS G. HARTLEY . ; Notes on the Genus Galipea (Rutaceae) i in Central America. THomas §. ELIAs . The Genera of Molluginaceae and Aizoaceae in the Southeastern United States. . LINN BOGLE Seedling Morphology in the J uglandaceae, the Cotyledonary N Louis F. CONDE pe DONALD E. STONE . The Vascular System in the Axis of Dracaena fragrans (Agavaceae) 2. Distribution and Development of Secondary Vascular Tissue. M. H. ZIMMERMANN and P. B. TOMLINSON . Relationships in the Piriqueta caroliniana— P. cistoides Complex (Turneraceae ). ROBERT ORNDUFF A Revision of the Boraginaceae of West Pakistan and Kashmir (continued). S. M. A. KAZMI . : The Genera of Chrysobalanaceae i in the Southeastern United State GHILLEAN T. Sein A New Species of Parietaria. a (Urticaceae) from oe Mexico NorTON G. MILLER Chromosome i eae in 1 the J uglandaceae. A. S. HANs The Ecology of an Elfin Forest i in Puerto Ric 13. Phytochemical Screening and Literature Sure G. J. PErstNos, S. K. Curistie, J. M. BIDINGER, and M. J. LAPIANA . INDEX . 492 499 529 534 540 547 A ip : WY ES Z- : ; i y, INGA aN N/A SV hi mn i " Voluane’ st Number 1 pers ) Us if iF \ 2 re _ JOURNAL | ae OF THE ee ARNOLD ae _ HARVARD UNIVERSITY a i (2) | Mresount Botanical Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subscription price $10.00 per year. Subscriptions and remittances should be sent to Miss Dulcie A. Powell, Arnold Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Volumes I-XLV, reprinted, are available from the Kraus Reprint Cor- poration, 16 East 46th Street, New York, New York 10017, U.S.A. EDITORIAL STAFF B. G. Schubert, Editor C. E. Wood, Jr. T. G. Hartley CIRCULATION D. A. Powell Printed at the Harvard University Printing Office, Boston, Massachusetts COVER: The Golden Larch, Pseudolarix Gordon WirH volume 51 the Journal of the Arnold Arboretum begins its second half century of continuous publication. To celebrate the occasion we are pleased to present a new cover format. The cover design is the work of Norman Comeau of the Art Department, Thomas Todd Company, Printers, Boston. We hope from time to time to have drawings of other plants on the cover. The Table of Contents will appear on the back cover. Eds. . Second-class postage paid at Boston, Massachusetts JOURNAL OF THE ARNOLD ARBORETUM VoL. 51 JANUARY 1970 NUMBER 1 THE SYSTEMATICS AND BREEDING SYSTEM OF GELSEMIUM (LOGANIACEAE) Bie vast yyzto ROBERT ORNDUFF THE LOGANIACEOUS GENUS Gelsemium exhibits a pattern of distribution that has interested plant geographers and systematists for over a century (Gray, 1846, 1859; Hara, 1952, 1956; Miranda & Sharp, 1950; Li, 1952). The genus is represented by one species in southeastern Asia, a second species in the southeastern United States, and a third species in the southeastern United States and the highlands of southern Mexico and Guatemala. This distribution pattern is similar to that which occurs in a number of other taxa of flowering plants. Although this disjunction dates back to the close of the Tertiary, Ge/semium has continued to main- tain a relatively high degree of morphological and genetic homogeneity. The purpose of this paper is to present a general systematic account of Gelsemium and to discuss the breeding system which is associated with heterostyly in the genus. I am indebted to D. E. Breedlove, J. F. M. Cannon, S. C. K. Chan, J. E. Ewan, R. K. Godfrey, A. Lourteig, L. I. Nevling, Jr., G. B. New- comb, W. T. Stearn, and G. L. Webster who assisted this study in various ways. I am also grateful to the National Science Foundation, whose grants have provided financial support for this investigation. The paper was written during the tenure of an appointment in the Miller Institute for Basic Research in Science at the University of California. Specimens deposited at A, BM, DUKE, E, F, FSU, GH, K, LAF, MISSA, MO, NCU, NO, NSC, Ny, SMU, Uc, and usF have been examined and I am grateful to the staffs of these herbaria for making the specimens available to me. RELATIONSHIPS The relationships of Gelsemium (n = 8, see below) to other genera of Loganiaceae are not at all clear. In their description of the Asian species in 1849, Gardner and Champion suggested an affinity with the African Usteria (n = 11; Miége, 1960) and the South American Antonia based on similarities of capsule structure. Bentham (1876) placed Gelsemium, 2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the African-South American Mostuea (n = 10; Gadella, 1962, 1963), and the Central American Plocosperma together in the tribe Gelsemieae. The occurrence of distyly (Leeuwenberg, 1961) and indole alkaloids (Leen- houts, 1962) in both Gelsemium and Mostuea supports the morphological evidence indicating that these genera are allied, but the cytological data do not conclusively support this affinity. Moore (1947) indicated that Gelsemium has an isolated position in Loganiaceae and might have closest affinities with Apocynaceae, although he favored retention of the genus in the first family. Despite the number of generic names that have been applied in Gel- semium, the only attempt to discriminate between New World and Asian species at the generic level was that of Gardner and Champion, who dis- tinguished the Asian Medicia from Gelsemium because the former has “imbricated, not quincuncial, aestivation of the corolla, inflated capsule, and numerous peltate, compressed seeds surrounded on all sides by a broad inciso-dentate membranous wing.” The differences in aestivation are slight and the seeds of G. sempervirens are asymmetrically winged, thus leaving the capsule character as the sole one which might be used for a generic distinction. I am, therefore, inclined to follow the tradition of other workers in recognizing only a single genus. Nevertheless, the Asian species is considerably more different from the two American species than these are from each other. Chromosome numbers of 2” = 16 have been obtained for the three species of Gelsemium (Taste 1; Moore, 1947; Gadella, 1963). All chromosome counts reported in this paper were made on mitotic tissue of shoot apices with the exception of a single meiotic count obtained from microsporogenous tissue in G. sempervirens. Recently Duncan and De- Jong (1964) have reported 2m = 8 for G. rankinii based on sectioned material in which counts of 2” = 9, 10, and 16 were also noted. Because of the variation in the number of chromosomes they observed, the diffi- culties in interpreting sectioned material (see, for example, Epling, Lewis, & Raven, 1962), and the consistent chromosome number of 2” = 16 that I have obtained for this species from several localities, Duncan and De- Jong’s report and the evolutionary conclusions based upon it are open to question. Attempts to produce artificial hybrids from legitimate crosses between Gelsemium sempervirens and G. rankinii resulted in low seed production (TABLE 2). All hybrid seed was planted, but the germination percentage was also very low. A total of eleven hybrid plants has been obtained, but these are considerably less vigorous than seedlings of the parental species. Most of the hybrids died a few months after germination and the re- mainder have been slow to produce flowers. The only hybrid which has flowered is one individual obtained from a cross between Breedlove 9005 (G. sempervirens, Chiapas, Mexico) and Godfrey 62671 (G. rankinii, Lib- erty County, Florida). This plant produced a single flower with pollen having an 81% viability, based on the stainability of a sample of 200 pollen grains mounted in aniline blue-lactophenol. However, when it was 1970] ORNDUFF, GELSEMIUM 3 backcrossed to both parents only five seeds matured in the capsule. In pedicel, sepal, and capsule characters the hybrid resembled G. rankinii, but the winged seeds were similar to those of G. sempervirens. Thus, these two species are separated by barriers to crossing, by low germinability of hybrid seed, and by reduced viability and fecundity of the hybrids. Some plants of the Guatemalan and Mexican populations of G. semper- virens are indistinguishable from those in the United States, but others have leathery, dark green, glossy, involuted leaves, as well as a tendency for higher numbers of flowers in each inflorescence. In addition, these Meso-American plants apparently have unscented flowers, whereas flowers of this species in the United States have a very strong fragrance. TABLE 1. Chromosome numbers of Gelsemium CHROMOSOME COLLECTION DATA LOCALITY NUMBER G. SEMPERVIRENS: L. E, Anderson, s.n. Durham County, North Carolina n=3 Ornduff 6376 Hoke County, North Carolina 2n = 16 Ornduff 6707 Richland County, South Carolina in = 16 Breedlove 9005 Pueblo Nuevo Solistahuacan, 2n = 16 Chiapas, Mexico G. RANKINII G. B. Newcomb, in 1965 Tammany Parish, Louisiana 2n = 16 R. Godfrey 62667 Liberty County, Florida an = 16 R. Godfrey 62671 Liberty County, Florida 2n = 16 R. Godfrey 62673 Franklin County, Florida 2n = 16 G. ELEGANS: S.C. B. Chen, oH. Hong Kong 2n = 16 A few hybrids have been obtained from a cross between some plants of G. sempervirens from Chiapas, Mexico, and others from Durham County, North Carolina. These parental populations are from the extreme ends of the range of this species. The pollen viability of the two hybrid plants that have flowered at the time of writing is 100% (based on a sample of 100 grains mounted in aniline blue-lactophenol). This suggests that the somewhat divergent morphological and biochemical characters (i.e., lack of flower odor) of the geographically disjunct parental populations are not associated with sufficient genetic discontinuities to result in lowered pollen. viability of hybrids obtained from a wide geographical cross. TOXIC PROPERTIES The herbage of Gelsemium contains several toxic alkaloids (Debay, 1950; Martinez, 1959; Kingsbury, 1964). Gelsemium sempervirens causes death and abortion in livestock which feed upon its leaves (Hardin, 1961; Kingsbury, 1964). Ingestion of nectar and honey produced from Gelse- 8-9 1A 4 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 mium flowers reportedly has caused death in humans and bees in the south- eastern United States (Kingsbury, 1964). Information concerning G. elegans indicates that this species has been used in China, Viet Nam, and Borneo as a suicidal poison which is either ingested or smoked. A note with Khant 15216 («) states that this species is used as a fish poison in Burma. The label of Dickason 9789 (GH) reports that the flowers of G. elegans are “poisonous to smell! Kills butterflies on the flowers.” It has been stated (Gibbs in 1910, Bm) that in Borneo this Species is ‘much used for feeding pigs . . . but is very poisonous to man.’ Despite its toxic properties, Gelsemium sempervirens is frequently grown in the gardens of the southeastern United States and of California, where it is prized for its heavily fragrant yellow flowers. It has been cul- tivated in Europe since the mid-17th century. TAXONOMY Gelsemium Jussieu, Gen. Pl. 150. 1789. Jeffersonia Brick., Med. Rep. N.Y. 1: 555. 1800. Medicia Gardn. & Champ., Hook. Jour. Bot. 1: 324. 1849. Leptopteris Blume, Mus. Bot. Lugd. Bat. 1: 240. 1850. “Type species: G. sempervirens (L.) Jaume Saint-Hilaire. Twining woody vines; leaves opposite, simple, entire, petiolate; stipules represented by stipular lines; flowers pentamerous, distylous or homo- stylous, 1 to many, in axillary or terminal inflorescences: ila funnel- form, the lobes imbricated in bud, bright yellow or orange-yellow; stamens five, epipetalous; style quadrifid at apex; ovary two-locular; fruit a cap- sule; seeds flattened, usually winged; » = 8. KEY TO THE SPECIES —_ Flowers 2.0-3.5 cm. long, borne in inflorescences of 1-8 flowers; capsules not inflated; seeds ‘wingless or with a strongly asymmetrical entire bia 2. Sepals obtuse, deciduous in fruit: ‘upper half of pedicels bracteolate; os sules terminated by a very short inconspicuous beak; seeds with a strongly asymmetrical, essentially unilateral wing; flowers usually erties scented. eee enema Rte rns Alii tt tuys cane as rere cotta ney 1. G. sempervireng. Sepals repr persistent in fruit; upper half of pedicels Eitneiae: capsules terminated by a conspicuous beak; seeds = wings; flowers usually ae See gett ae ene ces ane ea y ee 2. G. rankinii. . Flowers 1.2-1.7 cm. long, borne in inflorescences of numerous flowers; cap- sules inflated; seeds surrounded by an inciso-dentate wing. ... 3. G. elegans. » — Gelsemium sempervirens (L.) Jaume Saint-Hilaire, Expos. Fam. Nat. 1: 338. 1805 (Feb.-Apr.). 258- 1V \Bignonia sempervirens L. Sp. Pl. 2: 623. Liss: ae ast Gelsemium nitidum Michx. Fl. Bor. Am. 1: 120. 1803. Holotype: P (photo- | graph! GH). 1970] ORNDUFF, GELSEMIUM 5 4 (aus af Gelsemium es reg Encycl. Méth. Bot. Suppl. 2: 714. “1811” (3 July Holot rayb- gab bLisianthus ae) Mill. ex Steud. Nomencl. (ed. 1) 358. 1821. Leaves lanceolate to narrowly ovate, the blades 3-7 cm, long, more or less evergreen, the petioles 0.3-0.5 cm. long; flowers 1-8, corolla 2.5-3.5 cm. long, including the lobes 0.8-1.2 cm. long, bright- ie orange-yellow, darker in throat, strongly fragrant (rarely odorless); sepals elliptic, ob- tuse, 3-5 mm. long, mostly deciduous in fruit; pedicels 0.2-1.2 cm. long, usually densely bracteolate throughout; capsules broadly oblong, 1.5—2.5 cm. long, terminated by a very short beak 0.1-0.2 cm. long representing the base of a deciduous style, woody, hard, tardily dehiscent; seeds bro ish, 1.2-1.5 cm. long, including an asymmetrical wing t 0 1.0 cm. long; n = 8. Flowers (January) F aber ues (May) and eaaeunny again in late summer or early autumn. Fic. la—d. DIsTRIBUTION. Sea level to 6000 feet (in Mexico and Guatemala). Climbing on fences, low bushes, and trees, or prostrate on ground. Pine or deciduous woodlands in damp to very dry soil, southeastern Virginia to Florida to eastern Texas; mountains of Mexico and Guatemala (Fic. 3). Representative specimens: “United States. ARKANSAS. Bradley Co.: Warren, Demaree 14364 (Gu, ny), Demaree e (ny). Calhoun Co.: Hampton, De- maree 14407 (Ny). Cleveland Bs , Demaree 13568 (Ny). Faulkner Co.: Near Republican, Demaree in May ae (a); near Holland, Palmer 26534 (A). Hot Springs Co.: Malvern, Demaree 14493 (GH, NY). Miller Co.: 14 miles east of Texarkana, Harrison in 1946 (cH). Monroe Co.: Little Rock, Manning 85 (GH). Pulaski Co.: Near Ledwidge, Demaree 8890 (A, NY).~FLorwa. Desoto Co.: 6 miles north of Arcadia, Ward & Ward 2825 (FLA). Hernando Co.: Brooksville, Jones in 1919 (A); Brooksville, Jones in 1920 (Ny, Us). Highland Co.: Sebring, Hunnewell 9029 (GH). Hillsborough Co.: No definite locality, Fredholm 6245 (cu). Manatee Co.: Manatee, Simpson in 1898 ( Us) ; vicinity of Manatee, Simpson 5 (us). Orange Co.: No definite locality, Smith in 1886 (MIcH). Pasco Co.: Ehren, Riegler in 1961 (FLA); no definite locality, Barnhart 2518 (Ny). Pinellas Co.: No definite locality, Frank in 1900 (Ny). Polk Co.: Near Lake Marion, McFarlin 4505 (micH); Bartow, McFarlin 6637 yet Sarasota Co.: Sarasota, Perkins in 1943 (GH). “TENNESSEE. Hamilton Co.: miles from Sole Creek. Prather 284 (Ny). No definite riage Leonard pes (us).“Texas. Angelina Co.: 4 miles northwest of — Cory 52589 (GH, NY); 7 miles southwest of Lufkin, Cory 49746 (GH, NY, Uc, US). Hardin Co.: South- west of Hooks Switch, Cory 52693 (NY); adobe “Bailey 941 (US); north of Sour Lake, Lundell & Lundell 10882 (uc, us). Harris Co.: Houston, Fisher 14 (us), Hall 512 (cH, NY, US). Montgomery Co. - Wes r Conroe, Palmer 33330 (A). Nacogdoches Co.: 5 miles south of Nacogdoches, ‘Lundell & Lundell 9761 (miIcH, us). Newton Co.: 22 miles south of Newton, Cory 10914 (cH). Polk Co.: Indian Reservation, Girvin on Jan. 31, 1940 (GH, uc); Livingston, Palmer 5192 (A, Micu, us), Palmer 6795 (A, US); near Rye, Traverse 300 (GH). San Jacinto Co.: Without definite locality, Tharp in 1935 (arcu, vs). Shelby Co.: Five miles north of Center, Rowell, York & Tharp 47243 (DUKE, GH, NY, UC, US). Tyler Co.: Hyatt Bog, 16 miles south of Woodville, Cory 52710 (cH, NY). Walker Co.: Near Huntsville, Tharp in 1946 (DUKE, NY, UC). VIRGINIA. Greens- JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Fic, 1. Characters of Gelsemium sempervirens and G. rankinii 3 . a-d, G. 5 pervirens. a, a of flowering stem, X 0.5; b, partly dissected short-styled ; » X 3; e-h scence, e, portion of — showing rounded leaf-bases often found i in the species, X 0.5; f, seed, < 3; g, partly dissected long-styled flower, 1.5; h, san prior to polly x 2, 1970] ORNDUFF, GELSEMIUM 7 ville Co.: Skipper’s, Fernald & Long 7143 (cH). Isle of Wight Co.: South of Lee’s Mill, Fernald, Long & Abbe 14224 (GH); south of Zuni, Fernald & Mooré 15146 (GH, NY, US). James City Co.: Hampton, Miller in 1903 (us); 5 miles west of Toano, Baldwin 101 (GH); southwest of Williamsburg, Grimes 2565 (Ny); near Williamsburg, Maxon 10614 (BM, MICH, Us). Nansemond Co.: Near Crismond, Fernald & Long 7142 (Gu, Us); edge of Dismal Swamp, Glea- son 8617 (NY). Northampton Co.: Savage Neck, Fernald, Long & Fogg 5411 (cH). Princess Anne Co.: Cape Henry, Egler 40-180 (Ny); Dismal Swamp, Coville & Kearney 101 (us), Morong in 1877 (Ny), Seaman s.n. (Ny, us), Ward in 1877 (us) ; vicinity of Norfolk, Jensen in 1907 (A, us); Virginia Beach, Richins in 1908 (cH), Hollick & Britton in 1890 (ny, us), Britton in 1892 (NY); south of Virginia Beach, Britton & Small in 1893 (Ny); near Virginia Beach, Coville & Kearney 17 (us); Virginia Beach, Fernald & Griscom 2876 (GH), Stevens in 1894 (us); without definite locality, Kearney 1168 (A, US). Southampton Co.: Meherrin River, Fernald & Long 7141 (GH, NY). Surry Co.: West of Bacon’s Castle, Fernald & Long po (GH). Sussex Co.: Near Moore’s Mill, Fernald & Long 7140 (pm, GH); Spring Grove, Eaton in 1938 (cH); northeast of Sussex Cities Fernald, Long & Abbe 14223 (GH, NY, US). “Guatemala. ALTA VERAPAZ: Cobar. von Tiirckheim 881 (GH, K, NY, Us), von (A, NY, Us). Mexico. “Curapas: Lagunas de Monte Bello, magne 697 (cH) ; Pueblo Nuevo Solistahuacan, Breedlove 9005 (ps, uc). “HIpDALco: About 4 miles from Zacualtipan, Moore 2337 (GH, uc). ‘Oaxaca: Talea, Galeot- ti 1605 (us). Puesta: Zacatlan, Salazar on Apr. 3, 1913 (us). VERA CRUZ: Jalapa, Halsted s.n. (NY), Pringle 7766 (GH, MICH, uc), Smith 1876 (NY, UC); Orizaba, Botteri 934, s.d. (BM, K). In his protologue dealing with this species, Linnaeus (1753) does not cite any specimens except one in Van Royen’s herbarium; all other cita- tions are of illustrations and written descriptions. Van Royen’s specimen at Leiden lacks flowers and fruits and is, therefore, not unequivocally referable to G. sempervirens rather than to G. rankinii. Since Linnaeus described fruits, he obviously had more in mind than Van Royen’s material alone. It is quite clear that the Linnaean protologue describes what we now call Gelsemium sempervirens. Furthermore, there is no evidence that G, rankinii had been collected before the early 19th century, whereas G. sempervirens was well known as a cultivated ornamental in Europe as early as the mid-17th century. Although the Van Royen specimen most likely is G. sempervirens, I am reluctant to designate it as a lectotype since vegetatively G. sempervirens and G. rankinii are not easily distinguishable (see Duncan & DeJong, 1964). The Catesby illustration (1731) is too generalized to be referable to one of these species, but his written description (mentioning rig siti winged seeds, and geographical distribution) clearly indicates that he was illustrating and describing G. sempervirens. So far as I can presen no herbarium material of this species known to have been seen by Catesby exists (see Dandy, 1958), and Catesby himself indicates that he drew largely from nature. Ray’s Historia Plantarum has a description also 8 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 mentioning odor and winged seeds, but there is no indication that a Specimen (or illustration) exists. The Plukenet drawing (1696) is very crude, although the winged seeds indicate that he was illustrating G. sempervirens. I am designating the illustration by Plukenet as the lecto- type of G. sempervirens since it is supported by a specimen in the Sloane Herbarium (H.S. 90: 55) at the British Museum which was probably collected by Banister. - I consider G. nitidum to be a synonym of G. sempervirens. In the Michaux Herbarium at the Muséum National d’Histoire Naturelle in Paris there are three sheets of Gelsemium that bear specimens which were pre- sumably collected by Michaux. Two of these sheets bear specimens of G. rankinii and the third has G. sempervirens. The only sheet annotated as G. nitidum is one of those with specimens of G. rankinii. Although Michaux had a mixed collection available when he described G. nitidum, his characterization of the flower odor indicates that he had G. semper- virens in mind. For this reason I do not believe that G. nitidum can be considered as the earliest correct name for G. rankinii and would suggest that there are good grounds for considering G. nitidum as a nomen con- fusum. {¢ 2. Gelsemium rankinii Small, Addisonia 13: 37. 1928. “Swamps of the Waccamaw River near Hallsboro, North Carolina, collected by H. A. Rankin, April 17, 1928.” Holotype: Nv! qs Gelsemium sempervirens (var.) inodorum Nutt. Gen. N. Am. Pl. 171. 1818. “Ao lotype: Bm! ‘Collected near Savannah, . Georgia. z Leaves lanceolate to narrowly ovate, the blades 3.0~7.5 cm. long, more or less evergreen, the petioles 0.3-0.6 cm. long; flowers 1—8; corolla 2.0— 3.5 cm. long, the lobes 0.8-1.5 cm. long, bright- to orange-yellow, darker in throat, usually odorless; sepals lanceolate, acuminate, 3-6 mm. long, persistent in fruit; pedicels 0.2-1.0 cm. long, ebracteolate on upper half; capsules oblong, 1.0-1.6 cm. long, terminated by a persistent stylar beak 0.2 cm. long, woody, hard, tardily dehiscent; seeds brownish, 3-4 mm. long, wingless; m = 8. Flowering (February) March to April and rarely again in early autumn. Fic. le-h. DIstRIBUTION, At low elevations, climbing on fences, low bushes, and trees, or prostrate on ground. Wet woodlands or swamps of the coastal plains of North Carolina (rare), southern Georgia, northern Florida, southern Mississippi, and southeastern Louisiana (Fic. 3). Specimens examined: “ALABAMA; Escambia Co.: Atmore, Blanton 206a (micH). Mobile Co.: Citronelle, Milligan in 1903 (us). FLorIDA. Bay Co.: Between Southport and Westbay, Hume in 1930 (ny). Calhoun Co.: Apalachi- cola, Mohr & Sargent in 1898 (us), Parker s.n. (NY); near Apalachicola, Chap- man herbarium 474a (A, GH, NY); Chipola River, east of Clarksville, Godfrey 56344 (FLA, GH). Escambia Co.: Escambia River near Pensacola, Sheppard in 1932 (Ny). Franklin Co.: Apalachicola River, Godfrey & Redfearn 55343 (FLA, 1970] ORNDUFF, GELSEMIUM 9 GH, NY); Crooked River, north of Carrabelle, Godfrey 61749 (ria). Holmes Co.: Between Chapley and Bonifay, Godfrey 56366 (DUKE, FLA, GH, NY, UC, US); west of Millers Crossroads, Godfrey 56403 (FLA, GH). Jackson Co.: West of Cot- tondale, Godfrey 56362 (FLA, GH, UC). Leon Co.: West of Tallahassee, Godfrey 58780 (FLA). Liberty Co.: 6 miles south of Telogia, Godfrey 59373 (FLA, GH); southeast of Telogia, Godfrey 61752 (FLA). Wakulla Co.: West of Crawford- ville, Godfrey 58229 (FLA, GH); north of Crawfordville, Godfrey 59372 (PLa, GH, NY, UC); 30 miles southwest of Tallahassee, Clewell & Hebb 586 (FLA). Walton Co.: Near Cluster Springs, Godfrey & Harrison 55399 (FLA, GH, UC, NY); south of De Funiak Springs, Godfrey 58256 (FLA, GH, NY), Godfrey 59020 (ria). “Grorcia. Pierce Co.: Without definite locality, Godfrey 63251 (FLA). Xoursiana. St. Martin Parish: St. Martinville, Langlois in 1886 (us). St. Tam- many Parish: Slidell, Lemaire 675 (FLA). Tangipahoa Parish: Hammond, Wil- son 346 (FLA); east of Hammond, Innes & Warnock 722 (cH). Washington Parish: Northeast of Pine, Correll 9231 (cu). VMississipp1. Jackson Co.: Biloxi, Tracy 6754 (GH, K); north of Escatawpa, McDaniel 4055 (ria). Pear River Co.: West of Picayune, Sargent 7943 (FLA). Stone Co.: East of Wiggins, Kral 16520 (ria).“Norta Carouina. Cumberland Co.: Fayetteville, perhaps cultivated, Rankin in 1930 (FLA). Although the existence of an odorless Gelsemium was recognized early in the 19th century, G. rankinii was not fully described until 1928. In 1927, H. A. Rankin, a nurseryman in Hallsboro, North Carolina, sent the first of a series of collections of living material of G. rankinii to J. K. Small at the New York Botanical Garden. Rankin had noted, among other things, that this ‘““Waccamaw river variety of Gelsemium”’ differed from G. sempervirens in its inodorous flowers, later flowering season, and pref- erence for a wetter habitat. Small noted additional differences between these two species in the distribution of bracteoles on the pedicels, shape of the sepals, size and shape of the capsule, and seed characters. Small, in general, accurately characterized G. rankinii in his original description of this distinctive species, but perhaps because of the attenuated descrip- tion and slightly inaccurate key (the leaf-base and anther characters he cites as distinguishing G. rankinii and G. sempervirens are unreliable) in his Manual of the Southeastern Flora (1933), few subsequent workers in the southeastern flora have recognized two species in the region. The majority of collections of G. rankinii in herbaria initially have been iden- tified as G. sempervirens, but in the late 1950’s R. K. Godfrey accurately distinguished between these two species in his herbarium annotations and extensive field collections. Later, Duncan and DeJong (1964) discussed the morphological, ecological, and phenological differences between the two. Because of its occurrence in very wet sites and the ease with which it is confused with the often sympatric G. sempervirens, it is probable that G. rankinii is more common in the field than might be indicated by its relatively low representation in herbaria. q5} 3. Gelsemium elegans (Gardn. & Champ.) Benth. Jour. Linn. Soc. 1: 90. 1857. q5 g Medicia elegans Gardn. & Champ., Hook. Jour. Bot. 1: 324. 1849. Holotype: kK! 10 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 45 Leptopteris sumatrana Blume, Mus. Bot. Lugd. Bat. 1: 240. 1849. ave Gelsemium sumatranum Boerl. Handl. 2: 448. 1899. Vine to 3.5 m. tall; leaves elliptic to ovate, the blades 6-13 cm. long, sometimes cuspidate, evergreen, the petioles 0.5-1.2 cm. long; flowers numerous, inflorescence terminal or axillary; corolla 1.2-1.7 cm. long including the lobes 0.3-0.8 cm. long, bright- to orange-yellow, odorless (or rarely malodorous?); sepals lanceolate, acuminate, 3-4 mm. long; pedicels 0.3-1.0 cm. long, ebracteolate or with a single subtending brac- teole; capsules ovate-elliptic in outline, 0.8—-1.5 cm. long, inflated; seeds brownish, 3—4 mm. in diameter, including an inciso-dentate wing ca. 1-2 mm. wide; = 8. Flowering September to December and occasionally at other times. In fruit March and April. Fic. 2a-f. DisTRIBUTION. Sea level to 6000 feet. Climbing on shrubs and trees in scrub and open woodlands. Assam, northern Burma, northern Thailand, Laos, Viet Nam, southern and southeastern China, Sumatra, and northern Borneo. Fic. 3. Specimens examined: “Borneo. British NortH Borneo: From A. B. C. Francis, resident of the interior, Gibbs 3130 (gm, us). Burma. Hkinlum, King- don-Ward 21632 (A, BM); Kachin Hills, Kingdon-Ward 9020 (a); Hookum Valley to Ava, Hb. Griffith 3732 (x); Myitkina district, Lace 6041 (kK); Pang- yang, Manglon State, Dickason 9789 (a); Sumpra Bum, Kingdon-Ward 20566 (a, BM); Wa States, Maung Po Khant 15216 (x).~China. FuKIEN: Foochow, Warburg 5780 (a); Inghok, Chung 3219 (uc). HAInAN: Dung Ka to Wen Fa Shi, Chun & Tso 43755 (a, NY, UC); without definite locality, Liang 64088 (K, ny), Liang 64203 (Ny), Tang 450 (A), Wang 35189 (Ny, us), Wang 36650 (A, Ny). Kwanost: Pin-lam, Ko 55585 (a); Po Yam Shan, Tsang 22925 (A); Shap Man Taai Shan, Tsang 22029 (a, pm), Tsang 24597 (a, NY); Tou Ngok Shan, Tsang 23174 (A); Tung Loo, Ching 5645 (A, uc, us); Tsin Hung Shan, Ching 6860 (A, NY, UC, Us); Yao Shan, Wang 40048 (A); without definite local- ity, Chen 91254 (a). Kwancrunc: Au Tsai, McClure 3534 (nv); Chung Tung, Tai Tsan, Ying Tak, Tsang & Wong 3188 (uc); Kwai Shan, Tsang 28553 (A); Lin Fa Shan, Tsang 25748 (A); Lofan Mts., Ford s.n. (Ny); Lofoushan, east river region, Ho 60162 (ny); Lofoushan Mts., Tsiang 1777 (A); Naam Kwan Shan, Tsang 20260 (k, Ny); Nam Shan, Tsang 28874 (a); Ngong T’in Lo Shan, Taam 347 (a); Shaan Sam village, McClure 13291 (uc); Sha Lo Shan, Taam 198 (A); Sinnei district, T. Ving 2705 (k); Tung Koo Shan, Tsang 21607 (K, NY, uc); Wan Tong Shan, Tai Tsan Ying Tak, Tsang & Wong 2703 (uc): Wat Shui Shan, Wang & Ling 7425 (uc); Weishang, Sunyi, Tsiang 2705 (Ny); Wong Chuk I and vicinity, Law 2173 (a); Wu Kan Tin, Tsiang Ying 83 (a, uc); Wui Shui, Tang & Fung 31 (Ny); Yang Shan and vicinity, Tsui 693 (K, NY, UC, US); Yun-fou, Wang 354 (a, uc, us); Yung-Yun city and vicinity, Lau 799 (a, NY), Lau 915 (a, Ny). Yunnan: Fo Hai, Wang 76247 (a), Wang 77396 (a); Hsian- meng-young, Wang 80994 (a); Keng-Ma, Wang 72863 (a); Meng-soong, Dah- meng-lung, Che-li Hsien, Wang 77992 (a); Mengtz, Hancock 243 (Kk); Meng- tsze, Henry 10452 (A, K, NY, US); Mar-li-po, Sze-tai-po, Feng 13954 (a); Mar- li-po, Tung-ting, Feng 13457 (a); Ping-pien Hsien, Tsai 55136 (a), Tsai 61880 (A); Si-chour-hsien, Faa-doou, Feng 12118 (A); Si-chour-hsien, Shiang pyng- shan, Feng 11485 (a); Tsing-pian, Tsai 52403 (a); without definite locality, 1970] ORNDUFF, GELSEMIUM Nia ber i elegans. a, portion of flowering stem, 65 b ee as Pr eemrestdh 5: ‘gd short-homostyle flower, X 2; e, pene Bic flower, X 2; f, short-styled flower, x .: 12 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Forrest 9214 (a). Hong Kong. N. K. Chun 40251 (K); Kow-loon, Lamont 467 (a); Lan-tao Island, Lamont 467 (pm); Lo Fan Shan, Anonymous in 1883 (cH); New pera Hb. Hongkong 1818 (a); without definite locality, Wright 609 (GH, K, us).Mindia. Assam: Jabocka (= Zoboka), Naga hill, Prain’s col- 1oo° | 77? G.sempervirens ° ry i on ey , | ehoae eee oo* ws* Ho* NS*E Fic. 3. Distribution of Gelsemium species. Top, distribution of G. semper- virens in the southeastern United States and (1NSET, upper left) Mexico and Guatemala. Lower Lert, distribution of G. elegans in southeastern Asia. LOWER RIGHT, distribution of G. ramkinii in southeastern United States. 1970] ORNDUFF, GELSEMIUM 13 lector 959 (A); Lushai Hills, Parry 322 (kK). Indonesia. Sumatra: Maranti, Rahmat Si Boeea 6116 (A); Mt. Singalan, Beccari 7 (kK), Beccari 355 (BM). aos. Muang Huang, Wiengchan, Kerr 20816 (x), Kerr 200116 (pm Thailand. Pu Huat (Nan), Kerr 4996 (pm, kK). “Viet Nam. ANNAM: Thanhhoa, Poilane 1893 (A). ToNKIN: Hoa Binh, Petelot 1906 (Ny, us); Kau Nga Shan, Tsang 30578 (A, K); Kun-Me-Tawng-Mé-Ping, Garrett 1258 (K); Ouonbi, Anonymous in 1885 (a); Taai Wong Mo Shan, Tsang 27326 (a, K), Tsang 29611 CA. HE), BREEDING SYSTEM Gelsemium sempervirens and G. rankinii are exclusively distylous, but G. elegans has both distylous and homostylous races. The distylous condi- tion in the genus seems first to have been observed by Walter (1788), although it was overlooked by a number of subsequent workers (see Gray, 1873). Distyly in Gelsemium is characterized by the occurrence of two types of plants in populations of each species. Some individuals have long- styled flowers, short stamens, and small pollen; others, have short-styled flowers, long stamens, and larger pollen grains (TABLE 4). These two forms also differ in their compatibility relationships (TABLE 2). TABLE 2. Results of interspecific and intraspecific pollinations in Gelsemium PER CAPSULE A NUMBER OF NUMBER VERAGE FLOWERS OF SEEDS NUMBER OF Cross USED PRODUCED SEEDS Interspecific pollinations: G. rankinii X G. sempervirens ae 5 0.2 G. sempervirens * G. rankinii 18 86 4.8 Intraspecific pollinations: G,. SEMPERVIRENS: long selfed, or long 99 73 0.7 short selfed, or X short 40 53 1.3 AVERAGE 0.9 long short 58 765 13.2 short X long 28 307 11.2 AVERAGE 12.4 G. RANKINII long selfed, or X long 12 0 0.0 short selfed, or X short 25 56 2.2 AVERAGE é 8. long X short 11 157 14.3 short * long 14 369 26.4 21.0 AVERAGE 14 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 The breeding system of Gelsemium rankinii and G. sempervirens was investigated by conducting a series of self-pollinations, pollinations among long-styled plants, and pollinations among short-styled plants (termed “G]legitimate pollinations” by Darwin, 1877), as well as pollinations be- tween long- and short-styled plants (“legitimate pollinations”). The re- sults of the illegitimate pollinations (TaBLe 2) indicate that a strong incompatibility system is present in both species since such pollinations result in a seed production that is much lower than seed production follow- ing legitimate pollinations. The long-styled form of both species seems to exhibit a stronger self-incompatibility than does the short-styled form. Legitimate pollinations produced a high seed set and it is assumed that in nature the reciprocal differences in position of stigmas and anthers of the two forms enhance legitimate pollinations by insects. The breeding system of the distylous race of G. elegans is unknown, but since the morphological differences between the long- and short-styled forms of this species parallel those that exist in the American species, I assume it has an incompatibility system similar to that of its New World relatives. The homostylous race of G. elegans is a short-homostyle, i.e., the flowers have the style length of the short-styled form and the anther TABLE 3. Population samples of Gelsemium sempervirens showing representation of plants with long- and short-styled flowers NUMBER OF NUMBER OF PLANTS WITH PLANTS WITH COLLECTION LONG-STYLED SHORT-STYLED NUMBER LocaLity FLOWER FLOWER Ornduff 6376 Hoke Co., N. Carolina 30 20 6388 Duplin Co., N. Carolina 24 27 6389 Duplin Co., N. Carolina 24 21 6394 Bladen Co., N. Carolina 22 15 6702 Orangeburg Co., S. Carolina 13 25 6704 Bamberg Co., S. Carolina 55 56 6705 Calhoun Co., S. Carolina 15 17 6707 Richland Co., S. Carolina 61 44 6708 Wake Co., N. Carolina 30 37 Breedlove 9005 Chiapas, Mexico 14 11 TOTAL 288 273 position and pollen grain size of the long-styled form (Fic. 2d; TaBLE 4). The only living plants of G. elegans that were available to me belonged to this homostylous race and were self-compatible, as would be expected from the position of the reproductive organs, This occurrence of a short-homo- style race in a normally distylous species is unusual. In the majority of heterostylous species in other plant families (such as Primulaceae, Rubi- aceae, and Turneraceae) in which homostylous plants occur, the latter are generally long-homostyles. 1970] ORNDUFF, GELSEMIUM 15 Data concerning the pollination biology of Gelsemium are scant. Two collections of bees visiting G. sempervirens have been made. My collection was made in North Carolina and was identified by T. B. Mitchell. The most common visitors to the flowers were males and females of the large- bodied Emphoropsis laboriosa (Fabr.) (Anthophoridae). In addition, smaller-bodied bees belonging to the genera Dialictus and Ceratina were also collected. In Virginia, Hurd (pers. comm.) has noted E. laboriosa as well as the bumblebee Bombus impatiens Cresson (Apidae) visiting flowers of G. sempervirens. It is probable that the larger-bodied bees are the chief pollinators of Gelsemium, since the shape and size of the corolla tube as well as the relative position of anthers and stigmas require rela- tively large-bodied insects to effect pollen transfer, Population samples of G. sempervirens indicate that long- and short- styled plants are present in a 1:1 ratio (TABLE 3). This is probably also the case for G. rankinii (Duncan & DeJong, 1964). Elsewhere in the Loganiaceae, heterostyly has been reported for Mos- tuea (Leeuwenberg, 1961) and has also been suggested for Buddleia (Leenhouts, 1962), although the latter genus requires further study. Otherwise, heterostyly is unknown in this family, although it occurs in the related Oleaceae and Rubiaceae. Despite the rarity of heterostyly in the Loganiaceae and its possibly independent evolutionary origin in this family, the morphological expression of heterostyly in Gelsemium is re- markably similar to that which exists in other unrelated distylous genera (Darwin, 1877; Vuilleumier, 1967). LITERATURE CITED BENTHAM, G. 1876. Loganiaceae. /n: G. BENTHAM & J. D. Hooker, Genera Plantarum. London. Catessy, M. 1731. Natural history of Carolina, Florida, and the Bahama Islands, Vol. I. London. Danpy, J. E. 1958. The Sloane herbarium. London. Darwin, C. 1877. The different forms of flowers on plants of the same species. ondon. Desay, A. 1950. Etude du Gelsemium sempervirens. These. Fac. Pharm. Univ. Paris, ser. U-137. p. 72. ‘Duncan, W. H., & D. W. DeJonc. 1964. Taxonomy and heterostyly of North American Gelsemium (Loganiaceae). Sida 1: 346~357. Eptinc, C., H. Lewis, & P. H. RAvEN. 1962. Chromosomes of Salvia: section " Audibertia. Aliso 5: 217~221. ; ' GADELLA, T. W. J. 1962. Some cytological observations in the Loganiaceae. Acta Bot. Neerl. 10: 51-55. . 1963. Cytological studies in the Loganiaceae. Proc. Nederl. Akad. Wet. C. 66: 265-269. a Gray, A. 1846. Analogy between the flora of Japan and that of the United States. Am. Jour. Sci. Arts 2: 135, 136. 1859. Diagnostic characters of phaenogamous plants . . . with obser- vations upon the relations of the Japanese flora to that of North America, [vox. 51 JOURNAL OF THE ARNOLD ARBORETUM 16 TABLE 4. Measurements of style length, stamen Jength, and pollen size of heterostylous and homostylous Gelsemium flowers STYLE SAMPLE STYLE LENGTH STAMEN LENGTH POLLEN SIZE SPECIES LENGTH SIZE (with range in cm.) (with range in cm.) (with range in «; sample size) ; Long 36 1.8 (1.22.3) 0.8 (0.7-0.9) 33.9 (31.5~36.4; 260) G. sempervirens Short 38 0.8 (0.5~1.3) 1.6 (1.5-2.3) 37.0 (32.8-40.1; 200) Long 10 2.2 (1.8~2.4) 0.7 (0.6-0.9) 31.5 (28.1-34.2; 200) G. rankiniu Short 10 0.7 (0.6-0.8) 1.8 (1.7-2.0) 36.1 (34,2-38.1; 200) Long 10 1.2 (1.0-1.4) 0.5 (0.4-0.6) 27.7 (22.8-33.2; 180) G. elegans Short 10 0.7 (0.5-0.8) 1.1 (1.0-1.4) 30.8 (25.6-38.0; 180) Homostyle 7 0.7 (0.6-0.8) 0.6 (0.5-0.7) 26.7 (24.0-30.8; 60) 1970] ORNDUFF, GELSEMIUM 17 and of oy parts of the northern temperate zone. Mem. Am. Acad. Arts Sci. ee —452. . 18 ae on Gelsemium. Am. Jour. Sci. Arts 5: 480 Hara, H. eis Contribution to a study of variations in the Japanese plants closely related to those of Europe or North America. I. Jour. Fac. Sci. Univ. Tokyo Bot. 6: 29-96. 1956. Contribution to the study of variations in the Japanese plants closely related to those of Europe or North America. II. /bid. 6: 343-391. Harpin, J. 1961. Poisonous plants of North Carolina. Agr. Expt. Sta. N. C. State Coll. oe 414: 1-128 Kincssury, J. M. 1964. Poisonous plants of the United States and Canada. Englewood Cliffs, New Jersey. Lees, P. W. 1962. Loganiaceae. Jn: C. G. G. J. VAN STEENIS, Fl. Males. 93-387. Lenvwenntne A. J. M. 1961. The Loganiaceae of Africa. II. A revision of stuea Didr. Meded. Landb. Wageningen Nederl. 61: 1-31. Li, H, L. 1952. Floristic relationships ee eastern Asia and eastern North America. Trans. Am. Philos. Soc. 42: 371-429. LINNAEUS, C..1753. Species Plantarum. na m. MarTINEz, M. 1959. Las plantas medicinales de México, 4th ed. Mexico City. MiécE, J. 1960. Nombres chromosomiques de plantes d’Afrique Occidentale. Revue Cyt. Biol. Vég. 21: 373-384. Mrranpa, F., & A. J. SHARP. 1950. Characteristics of the vegetation in certain temperate regions of eastern Mexico. Evolution 31: 313-333. Moore, R. J. 1947. Cytotaxonomic studies in the Loganiaceae. I. Chromosome numbers and phylogeny in the Loganiaceae. Am. Jour. Bot. 34: 527-538. PLUKENET, L. 1696. Almagestum botanicum. London. SMALL, J. K. 1933. Manual of the southeastern flora. k. VurmLteumier, B. S. The origin and sages pad development of heterostyly in the angiosperms. Evolution 21: 210-226. Wa tter, T. 1788. Flora caroliniana. London. DEPARTMENT OF BOTANY UNIVERSITY OF CALIFORNIA BERKELEY, CALIFORNIA 94720 18 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 THE GENERA OF ULMACEAE IN THE SOUTHEASTERN UNITED STATES 1 Tuomas S. ELIAs ULMACEAE Mirbel, Elém. Phys. Vég. Bot. 2: 905. 1815, nom. cons. (ELM FAmILy) Trees or shrubs with watery sap; leaves with conduplicate vernation, alternate |rarely opposite], simple, the blade usually oblique at base, entire to variously serrate, petiolate; stipules paired, lateral or intra- petiolar, caducous; flowers perfect to imperfect (often by abortion of either androecium or gynoecium), actinomorphic to slightly zygomorphic, solitary, cymose, or in axillary fasciculate aggregations, usually borne on branchlets of the previous season (Ulmeae) or of the current season (Celteae). Perianth herbaceous, subcampanulate, (2—)4-8(—9)-lobed, the lobes free or variously fused, imbricate [valvate], persistent. Stamens erect in bud, hypogynous, arising from the base of the perianth, usually the same number as and opposite the perianth lobes [or sometimes more], the filaments distinct, the anthers bilocular with longitudinal dehiscence, extrorse or introrse; pollen 2—5(—6) aperturate (-porate, -colpate or -rupate), oblate to oblate-spheroidal. Gynoecium composed of 2 connate carpels; styles 2, linear, stigmatose along the upper inner surface, ovary superior, usually 1-loculate (occasionally 2-loculate in Ulmus), sessile to stipitate, the ovule 1, anatropous or amphitropous, pendulous from the apex of the locule, with 2 integuments. Fruit a samara, dry or thinly fleshy, often winged or appendaged, the seed with a straight embryo and without endosperm, or fruit a drupe, the seed with a curved embryo with "Prepared for a generic flora of the southeastern United States, a project of the Arnold Arboretum and the Gray Herbarium of Harvard University made possible through the support of the National Science Foundation (Grant GB-6459X, principal investigator Carroll E. Wood, Jr.). This treatment follows the format established in the first paper of the series (Jour. Arnold Arb. 39: 296-346. 1958). The area cov- ered includes North and South Carolina, Georgia, Florida, Tennessee, Alabama, Missis- sippi, Arkansas, and Louisiana. The descriptions are based primarily on the plants of this area, with additional information in brackets. References which I have not seen are marked by an asterisk. I am grateful to Dr. Wood for his thorough editing and valuable suggestions in the course of this study. Dr. John W. Thieret has kindly read the manuscript and has Mrs, N the remaining items and the illustration of Ulmus are by Arnold D. Clapman. The ry late Dr. G. K. Brizicky supervised the drawings and made the dissections from ma- terials collected by Dr. Woo 1970] ELIAS, GENERA OF ULMACEAE 19 folded or rolled cotyledons and usually with scanty endosperm. Embryo sac of the Polygonum type (Celtis) or of the Adoxa and Drusa types (Ulmus). Chromosome numbers, x = 10, 11, 14. (Celtaceae Link.) Typr GENUS: Ulmus L. A family of about 15 genera and 150-200 species distributed through- out much of the Northern Hemisphere and reaching into subtropical and tropical areas. Four genera are represented in the continental United States: Ulmus, with six species in the southeastern United States; Celtis, with five species in our range; Planera, a monotypic genus of the south- eastern United States; and Trema, largely tropical, but with two species reaching into southern and central Florida. Additional New World genera include Lozanella Greenman, ae a Klotzsch, and Miranda- celtis A. J. Sharp, all essentially tropica The Ulmaceae, one of five families in rie order of Urticales, is distin- guished from its nearest new world relatives the Moraceae, Cannabaceae, and Urticaceae by watery sap, leaves usually oblique at the base, flowers usually perfect, stamens erect in bud, fruit a samara or drupe, and seed with little or no endosperm. Using floral morphology and anatomy, Bechtel demonstrated the Ulmaceae to be the most primitive family in the order Urticales. In Ulmus, the flowers show evidence of suppression of a whorl of stamens and of one of the perianth whorls. Although the floral parts are cyclic in most ulmaceous genera, those of Ulmus are spirally arranged. In the gynoecium of Ulmaceae the many abortive vascular strands suggest suppression of carpels, the bicarpellate condition possibly having been derived from a polycarpellate condition. Tippo, using data from wood anatomy, confirmed this phylogenetic arrangement. It was pointed out that ring porosity appeared more commonly in the Ulmaceae (essentially temperate) than in the Moraceae (essentially tropical). Dumortier divided the family into tribes Celteae (incorrectly Celti- deae) and Ulmeae, but Engler raised the two divisions to subfamilial status. A third subfamily infrequently recognized is Barbeyoideae, but many workers place the monotypic genus Barbeya Schweinf. in a family of its own, Barbeyaceae Rendle. The two tribes (subfamilies) can generally be separated by their pollen, the Ulmeae usually having 4- or 5-porate, -colpate, or -rupate grains, the Celteae usually having 2- or 3-porate grains. The only exception is Zelkova (Celteae) which typically has the Ul/mus type of pollen. Fairly abundant in the fossil record, fruits, as well as leaves, of Ulmus and Celtis have been found in large numbers. Zelkova, Hemiptelea, and Pteroceltis also are known as fossils. The wood of Ulmus is of limited importance in the manufacture of furniture. The bark of U. rubra, high in mucilage content, has been used in medicinal preparations. Numerous cultivars of Ulmus, many resulting from efforts to produce disease resistant forms, are aban domestically as ornamentals. 20 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 REFERENCES: Bartton, H. Ulmacées. Hist. Pl. 6: 137-216. 1877. (English ed. 6: 140-219. 1880 Becutet, e a The floral anatomy of the Urticales. Am. Jour. Bot. 8: 386- 410. . [Ulmaceae 388-394, 399-401. pls. 15-18.] oats i & J. D. Hooxer. Ulmeae. Gen. Pl. 3: 351-356. 1883. [Under BERNARD, C. Sur la distribution a des lone Bull. Herb. Boiss. II. 5: 1097-1112. 1905. [See also 6: 23-38. 1906. ] Berry, E. L. Tree ancestors. vi + 270 pp. ore 1923. [Ulmaceae 146- 1 : “The Middle and upper Eocene floras of southeastern North America. U.S. Geol. Surv. Prof. Pap. 92: 158, 159. pl. 50, figs. 4, 5. 1924. ENGLER, A. Ulmaceae. Nat. Pflanzenfam. III. 1: 59-66. 1888. Fowetts, H. A. Silvics of forest trees of the United States. U.S. Dep. Agr. Handb. 271. vi + 762 pp. Washington, D.C. 1965. [Celtis, 139-145; Ulmus, 724-742.] Grsson, H. H. American forest trees. 708 + xv pp. Chicago. 1913. [Based on a series of articles in the Hardwood Record, 1905-1913. Ulmus, 379- 394; Planera, 397-399; Celtis, 403-406. ] Gray, J. Temperate pollen genera in the Eocene (Claiborne) flora, Alabama. Science 132: 808-810. 1960. [Zelkova. GrupzinskayA, I. A. Ulmaceae and reasons for distinguishing Celtoideae as a separate family Celtidaceae Link. (In Russian; English summary.) Bot. Zhur. 52: 1723-1748. 1967. Hovutpert, C. Phylogénie des Ulmacées. Revue Gén. Bot. 11: 106-119. pls. oy 5, TO99: Joncmans, W. Ulmaceae. In: W. Joncmans, ed., Fossil. Catal. IT. 10: 1-84. 1922. Kurz, H., & R. K. Goprrey. Trees of northern Florida. xxxiv + 311 pp. Gainesville, Florida. 1962. [Ulmaceae, 105-115. LESQUEREUX, L. Contributions to the fossil flora of the western territories. Part I]. The Tertiary flora. 7m: Report of the United States Geological Survey of the Territories, vol. 7. 1878. [Ulmaceae, 187-191. pls. 26, 27.]; Part III. The Cretaceous and Tertiary floras. /bid., vol. 8. 1883. [Ulma- ] he comparative internal posed of seeds. Am. Midl. Nat. 36: 513-660. 1946. [Ulmaceae, 636-638, 646 MeEtcHior, H. Ulmaceae. Engler’s Syllabus der Pflanzenfamilien. ed. 12. 2: 52-54. fig. 16. 1964. NeEvLInG, L. I., Jk. Ulmaceae. Jn: R. E. Woopson, R. W. Scuery, et al., Flora of Panama. Ann. Missouri Bot. Gard. 47: 105-113. 1960. PLancHon, J. E. Ulmaceae. DC. Prodr. 17: 151-210. 1873. Porsson, J. Sur un nouveau genre des Celtidées. Bot. Jahrb. 11: 52. 1890. [Samaroceltis. | PrIEMER, F. Die anatomischen Verhiltnisse der Laubblatter der Ulmaceen (einschl. regerr aa und die Beziehungen zu ihrer Systematik. Bot. Jahrb. 17: 419-475. pls. 10, 11. 1893. Seymour, F. Hs Notes on Moraceae and Ulmaceae. Am. Midl. Nat. 48: 249, 250. 1952. 1970] ELIAS, GENERA OF ULMACEAE 21 Tippo, O. Comparative anatomy of the Moraceae and their presumed allies. Bot. Gaz. 100: 1-99. 1938. [Ulmaceae, 23-28. Vines, R. A. Trees, shrubs and woody vines of the Southwest. xii + 1104 pp. Austin, Texas. 1960. [Ulmaceae, 203-214. West, E., & L. ARNoLD. The native trees of Florida. xx + 212 pp. Gainesville, Florida, 1946. [Ulmaceae, 55-60.] KEY TO THE GENERA OF ULMACEAE A. Flowers usually perfect; fruits either flattened or + dry with fleshy pro- tuberances; embryo straight, cotyledons flat or longitudinally folded (ULMEAE). B. Fruit a flat, winged samara; leaves usually biserrate; flowers si stk either before or after the leaves RA IMy anaes $a eee BF, Rae Py. . Ulm B. Fruit ovoid, wingless with numerous fleshy protuberances; ideas fone serrate; flowers appearing with the leaves. ................ 2. Planera. A. Flowers usually imperfect; fruits drupaceous; embryo curved, cotyledons variously folded (CELTEAE). C. Carpellate flowers aie or in few-flowered fascicles; anthers extrorse; TONITE OCDE SPUIWIGR 6 oo or ke wk es ee ees 3. Colts C. Carpellate flowers in axillary, subsessile cymes; anthers introrse; plants 4. Trema. Tribe ULMEAE [Dumort. ] 1. Ulmus Linnaeus, Sp. Pl. 1: 225. 1753; Gen. Pl. ed. 5. 106. 1754. Trees (or infrequently shrubs) of various habitats, the See usually deeply furrowed; branches unarmed, slender, terete, often with corky wings; leaf buds axillary, covered with numerous ovate to ecoita chest- veined, deciduous to subpersistent [rarely persisting until new growth]; stipules lateral, linear-lanceolate to obovate, entire, free or connate at the base, scarious, ‘inclosing the leaf in bud, caducous. Flowers perfect [rare- ly imperfect, the plants then polygamous], vernal to autumnal, minute, articulate on slender 2-bracteolate pedicels, produced in axillary subsessile or pedicellate cymes or racemes. Perianth uniseriate, campanulate, slight- ly to deeply (4—)5(—9)-lobed, membranaceous, marcescent. Stamens with the filaments filiform to + flattened, exserted after anthesis; anthers ob- long, emarginate, subcordate at the base, dorsified below the middle, ex- trorse. Style deeply 2-lobed, divergent, the lobes papillate and stigmatic on the inner face, ovary sessile or stipitate, compressed, glabrous or hir- sute, usually 1-locular by abortion, rarely 2-locular; ovule amphitropous, micropyle extrorse, superior, Fruit a + flattened samara, ovate to oblong, often oblique, sessile or stipitate, surrounded at the base by the marcescent calyx, often beaked by the remnants of the persistent styles, occasionally marked vertically by the thickened line of union of the two carpels, and 22 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 with a thin reticulate-venulose, membranaceous, light brown, broad or narrow wing [+ unwinged], the wing often ciliate on the margin. Seed ovate, compressed, marked on the abaxial edge with the thin raphe, without endosperm; seed coat membranaceous, light or dark brown, Fic. 1. Ulmus. a-h, U. americana: a, mature leaf X %; b, branchlet with flowers, X Ya; c, flower at time of shedding of pollen, X 5; d, flower with recep- tive stigmas, X 5; e, gynoecium in partial section to show ovule, X 5; f, mature fruit, X 1; g, seed, X 4; h, embryo, X 4. i-k, U. rubra: i, mature leaf, X 1%; j, branchlet with nearly mature fruit and expanding leaves, X /; k, mature fruit, x 1. 1, U. crassifolia, leaf, * 1. m, U. alata, mature leaf, X ™%. rarely produced into a narrow wing. Embryo straight; cotyledons flat or slightly convex, fleshy, much longer than the superior radicle, the hilum oblong-linear, pale. (Including Microptelea Spach, Chaetoptelea Liebmann.) Type species: U. campestris L. = U. glabra Hudson; see 1970] ELIAS, GENERA OF ULMACEAE 23 R. Melville, Jour. Bot. 76: 261-265. 1938, and A. Rehder, Bibl. Cult. Trees & Shrubs 135. 1949.2 (Ancient Latin name for elm.) — E_m A genus of 25-30 species widely distributed through temperate and boreal regions of the Northern Hemisphere, with central and northern Asia the apparent center of distribution. Six species occur in eastern North America, but the genus is noticeably absent from western North America. Ulmus is found throughout Europe, extends south to the mountains of northern Africa, east to Turkey and the Caucasus Mountains, genera, sections, and series (Spach, 1841; Planchon, 1848; Schneider, 1904); in the most recent subgeneric classification, Rehder (1949) recog- nized five sections, each represented in our range by at least one species. KEY TO THE SECTIONS OF ULMUS IN THE SOUTHEASTERN UNITED STATES A. Flowers ene appearing before the leaves; perianth with short, often un- equal lobe B. Foes. subsessile to very short pedicellate, not pendulous; samaras not eiiate ‘on: the-maryine:..oiis. 2oigmeal, eeisicias Fiee. Sect. ULmus. . Flowers long aging often unequally so, becoming pendulous; sa- maras marginally cilia C. Flowers ce ae floral axis only slightly elongating; samaras glabrous on the sides; branches without corky wings. .............. eg ieig putin. ies te Seti Ai ene l eonua eck. t. BLEPHAROCARPUS. C. Flowers racemose, the floral axis slender and elongating; samaras pubescent on the sides; branches often with corky wings. wo n Weare ici te eran ay ine rs a ect. CHAETOPTELEA. A. Flowers autumnal or appearing paste after the leaves; perianth with the lobes deeply divided, often to the bas D. Flowers fasciculate, not pendulous, ‘ian leaves subpersis- tent: t0de pereistent.;.. 6. a-cac0 fa be Saal oe GER ee Sect. MICROPTELEA. D. Flowers in pendulous racemes, 6—9-merous; leaves deciduous. .......... Sect. TRICHOPTELEA. westward through southern Ontario to eastern North Dakota, southward to western Oklahoma and central and southeastern Texas, and eastward to central Georgia and northwestern Florida, U. rubra is commonly found in lowlands, especially in rich soil along stream banks and on low, *The Linnaean name for the type species of the name Ulmus is U. campestris L. aia : fs been used for three — gre (U. glabra Hudson, U. carpinifolia Rup- pius ex Suckow, and U. proce Salisbury) this name is now considered a nomen ambiguum to be re a puny Art. 69 of the Code (see Report of Standing Committee on Shesrae gee boas Prop. Bot. Nomencl. XI Internat]. Bot. Congr., peep! 1969. Append. E, p. 1969.). The correct name for this species appears e U. glabra Huds. (cf. sisietis 1938, and Rehder, 1949). 24 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 rocky hillsides. In addition to the sectional characters, U. rubra is char- acterized by the ciliate leaves being scabrous above and by the very mucilaginous inner bark. Sterile, scabrous-leaved specimens of U. amert- cana may be confused with this species but are distinguishable by the hardly mucilaginous bark. Section BLEPHAROCARPUS Dumort. comprises a small group of Euro- pean species and a single North American one, Ulmus americana L., American, soft, water, or white elm, 27 = (28) 56. It occupies a broad range from Newfoundland west to southern Manitoba and eastern Sas- katchewan, south to North Dakota, southeastern Montana, western Nebraska, western Oklahoma and central Texas, and in the east to Florida. It is readily identified by the flowers borne in close fascicles with elongating, pendulous pedicels, and a marginally ciliate, elliptic samara. The leaves may be smooth to scabrous above. Fernald recognized four forms based on combinations of two pairs of vegetative characters: f. pendula (Ait.) Fern., with pubescent branchlets and the leaves smooth above; f. /aevior Fern., with the branchlets as well as the upper leaf surfaces smooth; f. alba (Ait.) Fern., with pubes- cent branchlets and the leaves harshly scabrous above; and f. imter- cedens Fern., with glabrous branchlets and the leaves harshly scabrous above. All four can be found in our range, but attempts to distinguish them can end in frustration, for both smooth and scabrous leaves can be found on the same tree (Thieret). Varietas floridana (Chapm.) Little (U. floridana Chapm.), Florida elm, differing from var. americana only in degree in several quantitative characters (cf. Kurz & Godfrey, p. 113) is found on the Coastal Plain from eastern North Carolina to central Florida. Natural hybrids of Ulmus americana are rare. Artificial hybrids have been made between this species and both U. pumila L., 2n = 28, and U. laevis Pallas (cited as U. pedunculata Fougeroux by Winieski), but seed production from the crosses was very low and germination of the hybrid seeds even lower. Sax (1933) postulated that U. americana rarely crosses successfully with other Ulmus species because of an inability of the chromosomes to pair, since most species are diploid (2” = 28), while most American elms are tetraploids. Much of the genetic re- search with the elms, especially U. americana, has been concerned with the development of various races and hybrids which would hopefully be resistant to the Dutch elm disease and/or phloem necrosis. The closest relative of U. americana is the European U. laevis Pallas, 2n = 28. Section CHAETOPTELEA (Liebm.) Schneid., a New World section of only three species, is represented in our range by Ulmus Thomasii Sarg. and U. alata Michx., both of which frequently have corky-winged branch- lets. Ulmus Thomasii, rock elm, 2n = 28, is known from western Ver- mont west to southern Ontario, central Michigan, Wisconsin, northern Minnesota, and southeastern South Dakota, south to northern and eastern Nebraska an1 eastern Kansas, east to Missouri and Tennessee (and lo- cally in northwestern Arkansas), and northeastward to Ohio and New 1970] ELIAS, GENERA OF ULMACEAE 25 York. Ulmas alata, wahoo, winged, or cork elm, occurs from Virginia to Kentucky, southern Indiana, southern Illinois, and Missouri, south to central Oklahoma, eastern and southeastern Texas, and east to central Florida. Both species are commonly found on dry, nicky uplands, slopes, or cliffs. The two are easily distinguished by the broadly elliptic samaras and larger, distinctly petiolate leaves of U. Thomasii, in contrast with the lanceolate-ovate samaras and small, subsessile lexees of U. alata. The third member of this section is U. mexicana (Liebm.) Planch. (Chaetoptelea mexicana Liebm.), which occurs in mountainous areas from southern Mexico southward to Panama. Section MicropreLreA (Spach) Benth. & Hook. includes only the closely related but widely separated Ulmus parvifolia Jacq., of eastern Asia, and U. crassifolia Nutt., cedar, basket, or southern rock elm, of the southern United States. The latter is found in southwestern Tennessee, Arkansas, and southern Oklahoma, south to central and southern Texas, Louisiana, and western Mississippi. An isolated station was recently dis- covered in northern Florida (Suwanee County), and intervening stations should be looked for in Alabama and eastern Mississippi. It is ques- tionably known from Nuevo Leon, Mexico. The older branches fre- quently have corky wings. This species, along with its Asian counterpart, is usually a tall tree reaching to 25 m. and is often free of branches for half its height. The frequently thick bark usually peels off in scales, exposing the brownish inner bark. Although described as a member of the following section, the poorly known U. monterreyensis C. H. Muller of Nuevo Leon, Mexico, is vegetatively very similar to U. crassifolia, but fruits and additional flowering material are needed before it can be placed properly. Section TRICHOPTELEA Schneid. contains Ulmus serotina Sarg., Sep- tember elm, probably the sole member of this section, which ranges from southern Illinois, Kentucky, and Tennessee, south to northern Alabama, northwestern Georgia, Arkansas, and eastern Oklahoma. Flowering and fruiting later than U. crassifolia, U. serotina is usually found in bottom lands and on limestone hills. Of the numerous species and cultivars in cultivation (see Green), three appear to be naturalized and spreading from cultivation. The English elm, Ulmus procera Salisb. (§ Ulmus), naturalized from south- ern New England and New York southward to Virginia, has been re- ported by Crutchfield from Roanoke Island, North Carolina. The other European introduction in eastern North America, U. glabra Huds. (U. campestris L., pro parte) (§ Ulmus), Wych elm, has not been reported as being naturalized in our range. The third species, U. pumila L., (§ Ul- mus), dwarf or Siberian elm, introduced from Asia and apparently sta- bilized from Minnesota to Kansas and in Utah, has also been reported from North Carolina by Crutchfield. This species has been planted in the northeastern and midwestern United States in parks, along boule- vards, and on farms for windbreaks. It has mistakenly been called U 26 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 parvifolia, Chinese elm, a species of eastern Siberia, northern China, and Japan which is not cultivated to any extent in the United States. Pollination in Ulmus is clearly by wind, and, within our range, the elms contribute to both the early spring hayfever or tree-fever and the late-summer hayfever seasons. Ulmus alata, U. americana, and possibly U. rubra, all producing prodigious amounts of pollen, cause hayfever mainly in February and March in the Virginias and Carolinas (Wode- house). Flowering about the same time as the ragweeds, U. crassifolia usually complicates late summer hayfever. Ulmus serotina, does not ap- pear to be significant as a hayfever plant. Although the flowers of Ulmus are perfect, cross pollination is ap- parently necessary to insure good seed production. It has not been clearly established whether the plants are partially or wholly self-incompatible or whether the proterogyny or proterandry reported in several species accounts for the lack of self fertilization. Further study of the floral maturation patterns is needed. The base chromosome number for Ulmus appears to be 14 (2n = 28). Chromosome numbers for at least 13 species have been reported, with most U. americana and possibly some U. glabra at the tetraploid level, 2n = 56. According to Sax, U. americana is probably an autotetraploid. Ehrenberg, in a study of the pollen of U. glabra, reported partial sterility in 47.2 per cent of the plants studied. The embryo sacs are tetrasporic and of the Adoxa or Drusa types or a modified form of the two. First appearing in the Upper Cretaceous, Ulmus became abundant and widespread in the rocks of the early Tertiary (over 28 Eocene species described). Apparently reaching its maximum distribution during the Miocene (ca. 30 species), Ulmus was widespread in Europe, both eastern and western North America, and Asia. Pleistocene deposits have yielded some eight species, most of which are still extant. Although many species of elms have corky flanges (wings) on the branches, the nature of these has been studied only in U. x hollandica Mill. in which the wings are produced by a symmetrical longitudinal splitting of the cork into bands which project like wings from the stem surface (Smithson). The corky wings found in U. alata and U. crassi- folia have not been investigated. Of the many diseases of elms, none has received as much attention as the Dutch elm disease, so called because much of the early work on it was done in Holland. Discovered in France in 1918 and now occurring over most of western Europe, it reached the United States about 1930, probably in a shipment of elm burl logs imported for veneer. It has now spread throughout the northeastern United States and has reached On- tario and Quebec and at least as far west as Illinois and as far south as Tennessee. Ulmus species vary in susceptibility, but U. americana, wide- ly planted as a street tree in northeastern North America, is especially vulnerable and is possibly threatened with extinction. The disease is caused by an ascomycete, Ceratocystis ulmi (Ceratostomella ulmi), a yeast-like stage of which spreads through the vessels. It both produces 1970] ELIAS, GENERA OF ULMACEAE 27 a toxin and stimulates the production of tyloses in the vessels, blocking water flow and causing wilting and death of the infected parts. The fungal spores are carried by the bark beetles Scolytus multistriatus and Hylur- gopinus rufipes (and perhaps others), which lay their eggs in dead or injured bark of the trunk and branches of elms. The larvae tunnel under the bark in a characteristic feather-like pattern. The fungus sporulates in cracks in the bark, beneath dying bark, and in the beetle tunnels, and the sticky spores are carried by the emerging adult beetles to uninfected branches of the same or other trees where the beetles feed for a time before breeding. The spores enter the healthy tissues through the feed- ing wounds made by the beetles in the thin bark of young shoots. No sat- isfactory control has yet been found, but control of the beetles through spraying and removal and burning of diseased trees is widely practiced in cities. Several clones have been reported to be resistant to the disease: a highly regarded one is U. carpinifolia ‘Christine Buisman’, which, although showing considerable resistance to phloem necrosis as well, is unusually susceptible to a dieback caused by Nectria cinnabarina. Phloem necrosis, caused by the virus Morsus ulmi, is even more deadly and has been known to kill an elm in a single season. The virus attacks the phloem, eventually destroying the phloem elements, interrupting the food supply to the roots, and causing in the more advanced stages small and even large roots to die. It is transmitted by the elm leaf-hopper, Scaphiodeus luteolus, which can be controlled by various insecticides. An American disease, cephalosporium wilt, caused by the imperfect fungus Dothiorella ulmi, causes foliage wilting and dieback of the young branches. Judicious pruning can usually control and eradicate this threat. Elms are attacked by numerous insect pests, including the elm leaf- curl aphid, Ariosoma ulmi; the leaf beetle, Galerucella luteola; the elm borer, Saperda tridentata; the elm lace bug, Corythucha ulmi; and the gypsy moth, Porthetria dispar. The wood of elm is heavy, tough, strong, often flexible, durable under water, often difficult to split, and fairly attractive in appearance. The sapwood is usually yellowish or cream colored with brown or grayish- brown heartwood. The wood of all the species in our range is ring por- ous. A minor lumber source, elm wood is used in shipbuilding, panelling, crates, and in some furniture. At one time it was commonly used for wheel hubs. High in mucilage content, the inner bark of U. rubra, and to a lesser extent that of U. americana, has been used in various decoctions in folk and Indian medicine in the southeastern United States. REFERENCES: Many popular or semipopular articles about U/mus and numerous references concerning the Dutch elm disease have been omitted. Under family references see BAILLON, 187; BERNARD, 10861112: Berry, 146-153; Fowe ts, 724-742; GIBSON, 379-394: Joncmans, 23-66; Kurz & Goprrey, 109-115; VinEs, 208- 214; West & ARNOLD, 58-60. pte AHLGREN, C. E. Phenological observations of nineteen native tree species in 28 JOURNAL OF THE ARNOLD ARBORETUM [vor. 51 northeastern Minnesota. Ecology 38: 622-628. 1957. [Includes U. ameri- cana. BARINGER, J. W. Phloem necrosis of elms. Minutes Natl. Pl. Board Annual Meeting 19: 15, 16. 1944.* Barton, L. Me Storage of elm seeds. Contr. Boyce Thompson Inst. 10: 221-233. pls. 1, 2. 1939. BEATTIE, R. oe How the Dutch elm disease reached America. Proc. Natl. Shade Tree Conf. 9: 101-105. 1933.* BILLINGS, oe H. Precursory leaf-serrations of Ulmus americana. Bot. Gaz. 40: 224, 225, dis. 1,2: 1905. speomsi H. A. Light effects on tree growth and seed germination. Ohio Jour. Sci. 57: 357-364. 1957. [U. americana. | Bretz, T. W. Leaf-bud cuttings as a means of propagating disease-resistant elms. Pl. Disease Rep. 33: 434-436. 1949.* BroapHurRST, J. Self-pruning in the American elm. Torreya 17: 21-24. pls. 1, 2. 1917. Brown, N. A. Tumors on elm and maple trees. Phytopathology 31: 541-548. pls. 1-3. 1941. Brown, R. L. Wood structure of elms, maples and oaks as a means of identify- ing species. Rep. Mich. Acad. Sci. 4: 109-112. 1904 Carter, J. C. Inheritance of foliage variegation in variegated English elm (UI- mus procera argenteo-variegata). Trans. Illinois Acad. Sci. 39: 43-46, pls. 1,2. 1946; Cuase, S. S. A case of albinism and of presumptive self- compatibility in UI- mus L. Rhodora 70: 294-296. 1968. es a hae E. M. The recent naturalization of nee elm (Ulmus pumila n Utah. Great Basin Nat. 24: 103-106. 196 ee D. B. The Cephalosporium disease of ae Contr. Arnold Arb. 10: 1-91. pls. 1-16. 1937. CRUTCHFIELD, P. J. Taxa collected from Roanoke Island new to the flora of North Carolina. Castanea 29: 129-137. 1964. [U. procera, U. pumila. | DALLIMorE, W. Elms and elm timber. Quart. Jour. Forestry 14: 109-118. 1920. D’Amarto, F. Embriologia de Ulmus campestris L. Nuovo Giorn. Bot. Ital. I]. 47: 247-263. 1940. DeSmipt, W. G. Studies of the distribution and volume of the wood rays in slippery elm, Ulmus fulva. Jour. Forestry 20: 353-362. 1922. The American elm (Ulmus americana). Am. Forestry 22: cama srs Cc. E. ” Studies on elm pollen. Bot. Not. 3: 308-316. 1953. [U. glabra ExpaAHL, I. Die Entwicklung von Embryosack und Embryo bei Ulmus glabra Huds. Sv. Bot. Tidskr. 35: 143-156. 1941. Evert, R. F., C. M. Tucker, & J. D. Davis. Aspects of sieve-tube member ontogeny and structure in Ulmus americana. (Abstr.) Am. Jour. Bot. 55: 1968. - 716 FERNALD, M. L. Botanical specialties of the Seward Forest and adjacent areas of southeastern Virginia. Rhodora 47: 93-142. 1945. [The indigenous variations of Ulmus americana, 132, 133. Georce, E. J. Storage and dewinging of American elm seed. Jour. Forestry 35: 769-772. 1937. GittHaM, J. K., & T. E. Trmett. The hemicellulose of white elm (Ulmus 1970] ELIAS, GENERA OF ULMACEAE 29 americana). I. Canad. Jour. Chem. 36: 410-413. 1958; II. Jbid. 1467- 1474. GREEN, P. S. Registration of cultivar names in Ulmus. Arnoldia 24: 41-80. 1964 GRUDZINSKAIA, I. A. De genere Ulmus L. (sect. Blepharocarpus Dumort.) no- tulae systematicae. (In Russian.) Not. Syst. Leningrad 18: 51-57. 1957. ae F. G. Is the American elm (Ulmus americana) injured by natural gas? Pl. Physiol. 25: 433-440. 1950. Henry, A. On elm-seedlings showing Mendelian results. Jour. Linn. Soc. Bot. 39: 290-300. 1910. Hiccrns, D. J., & T. Artsumi. Time of floral differentiation in Ulmus ameri- cana, U. pumila, and U. carpinifolia. Bot. Gaz. 120: 177-180. 1959. Hotmes, F. W. Recorded Dutch elm eit distribution in North America as of 1959. Pl. Disease Rep. 45: 74, 1961. Jounson, L. P. V. Fertilization in eee with special reference to hybridization procedure. Canad. Jour. Res. C. 24: 1-3 46. LANcuUCcKA-SrRoponiowa, M. Two new genera: Hemiptelea Planch. and Weigela Thunb. in the younger Tertiary of noes (In English; Polish summary.) Acta Palaeobot. 8: 1-20 ig Liminc, F. G. A preliminary study of the lengths of the open vessels in the bratiches of the American elm. Ohio Jour. Sci. 34: 415-419. 1934. Linper, D. H. Some diseases of elm trees. Missouri Bot. Gard. Bull. 19: 61- 69. pls. 16, 17. LorsEau, J. E., & A. Hort. Sur la phyllotaxie et les anomalies des rameaux vévétatifs "de Vorme (Ulmus campestris L.). Compt. Rend. Acad. Sci. Paris 248: 2785-2787. 1959.* McCoy, E. E. A study of nutritional Aces of the American elm. New Jersey Dep. Agr. Circ. 394. 62 McDaniet, S., & C. Swirt. Ulmus emer (Ulmaceae) in Florida. Sida 3: 115, 116. Map. 1967 McLean, D. M. Histo-pathologic changes in the phloem of American elm af- fected with the virus causing phloem necrosis. Phytopathology 34: 818- MELVILLE a“ Typification and ee “ gs smooth-leaved my had car- pinifolia Gleditsch. Jour. Linn. Soc. Bot. 53: 83-90. pl. 3. «Law. pose bleeds Ruppius ex Sucko Ww] OUELLET, C. E. Factors able to influence the multiplication of the American elm (Ulmus americana L.) by cuttings of leafy branches. (In French.) Canad. Jour. Pl. Sci. 42: 150-162. 1962.* Peace, T. Pathology of trees and shrubs: with special reference to Britain. xx 753 pp. Clarendon Press, Oxford. 1962. [Diseases of elm, 417-427, life cycle of Dutch elm disease. ] —— E. B., & A. W. H. Daman. Convection plumes from Ulmus ameri- na L. Science 148: 392, 393. 1965. enue” P. P., B. O. Dopce, & H. W. Ricxett. Diseases and pests of ornamental plants. x + 775 pp. Ronald Press Co., New York. 1960. [Ulmus, 699- 710. ee F. C., & L. Frepertck. A comparative study of the nodal anatomy Ulmus americana and U. pumila. (Abstr.) ASB Bull. 11: 54. 1964. ie R. H. Studies on Ulmus: I. The range of variation of East Anglian elms. Watsonia 3: 138-153. 1955. [U. carpinifolia, U. glabra, U. procera.] SARGENT. C. S. Ulmus. Silva N. Am. 7: 39-58. pls. 311-315. 1895. 30 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 . Manual of the trees of North America (exclusive of Mexico). ed. 2. xxvi + 910 pp. Map. Boston & New York. 1922. [Ulmus, 308-316.] Sax, K. Chromosome numbers in Ulmus and related genera. Jour. Arnold Arb. 14: 82-84. pl. 56. 1933. [Undocumented chromosome counts. | SEYMOUR, F. = The type of Ulmus americana L. Rhodora 54: 138, 139. 1952. SHatruck, C. H. A morphological study of Ulmus americana. Bot. Gaz. 40: 209-223. pls. 7-9. 1905. SMALLEY, E. B., & A. J. Rrker. Tropical members of the Ulmaceae resistant to Dutch elm disease. Forestry Res. Notes 77. 4 pp. 1962.* [With refer- ence to breeding. | SmitH, L. M. Key to the Ohio elms in the winter condition. Ohio Nat. 5: 316. 1905. SmirHson, E. Development of winged cork in Ulmus & hollandica Mill. Proc. Leeds Philos. Lit. Soc. Sci. 6: 211-220. 1954. Straus, R. Some variations in fruit, seed, and seedling characteristics in the American may Ulmus americana L., in relation to geography of seed source. Diss. Abstr. B. 28(2): 511B, 1967.* STEINBAUER, C. E., & G. P. Srermvpaver. Effects of temperature and desicca- tion during storage on germination of seeds of the American elm (Ulmus americana L.). Proc. Am. Soc. Hort. Sci. 28(1931): 441-443. 1932. Taytor, L. A. Plants used as curatives by certain southeastern tribes. xi + 88 pp. Botanical Museum, Harvard University. 1940. [U. americana, 18, 19; Touw, A., & C. G. G. J. vAN SreeNnts. Note on Ulmus in Malesia (Ulmaceae). Blumea 16: 84. 1968. [U. lanceifolia Roxb. in Sumatra & sw. Celebes. | TRUE, R. P., & S.$. Scoutinc. Scouting and sampling elms with symptoms com- monly associated with Dutch elm disease as an aid in eradicating Ceratosto- mella ulmi. Phytopathology 29: 529-537. 1939. Tucker, C. M. Seasonal phloem development in Ulmus americana. (Abstr.) Am. Jour. Bot. 55: 716. 1968. TyLer, L. J. The influence of temperature on the Dutch elm disease in potted American elm. Phytopathology 35: 302-304. 1945. U.S. Dep. Acr., Forest Serv. Woody plant seed manual. U.S. Dep. Agr. Forest Serv. Misc. Publ. 654. 1948. [Ulmus, 364-367. | VERRAL, A. F., & T. W. AM. The transmission of Ceratostomella ulmi through root grafts. Phytopathology 25: 1039, 1040. 1935. The longevity of Ceratostomella ulmi in soils. Phytopathology 28: 763-765. 1938, Wa ker, R. I. ier AL rag and embryo development in Ulmus fulva. Bot. Gaz. 99: 592-598. . Chromosome numbers in “Ulmas. Science 75: 107. 1932. Megasporogenesis and development of megagametophyte in U/mus. Am. Jour. Bot. 37: 47-52. 1950. Went, J. C. Elm nomenclature. (In Dutch.) Nederl. Boschbouw-Tijdschr. 21: 106-108. 1949.* WinteskI, J. A. Artificial hybridization and grafting methods with Ulmus ameri- ana. Proc. Northeast. Forest Tree Improv. Conf. 7: 48-51. 1960. tham, Mass. 1945. (Ulmus, 86-90, pls. 41, 42: also 166, 172, 182.) Wyman, D. Elms grown in America. Arnoldia 1: 65-80. 1941; also 11: 79-93. 1951. [Mainly concerned with cultivars. ] 1970} ELIAS, GENERA OF ULMACEAE 31 2. Planera J. F. Gmelin, Linn. Syst. Nat. ed. 13. 2: 150. 1791. Deciduous trees or large shrubs of wet habitats, usually less than 20 m. tall, with scaly bark, the inner bark reddish brown; branchlets slender, terete, unarmed, puberulous; lenticels pale, scattered; leaf scars small, + orbicular; buds axillary, minute, subglobose, covered with thin, closely imbricate scales. Leaves petiolate, ovate-oblong, coarsely serrate, acute to rounded at apex, oblique to rounded at base, pinnately veined, the veins + conspicuously anastomosing beneath; stipules free, lateral, ovate, caducous. Plants polygamous, flowers perfect or often functionally im- perfect, fasciculate at the nodes of the one-year-old branches, articulate, short-pedicellate, the pedicels ebracteolate, the lower flowers often stami- nate. Perianth campanulate, 4- or 5-lobed from midway to near the base, the lobes rounded apically, greenish yellow, subscarious, submarcescent. Stamens as many as the perianth lobes in staminate flowers, in carpellate flowers often reduced or wanting; filaments filiform, exserted; anthers introrse, ovate, emarginate apically, cordate basally, dorsifixed below the middle; pollen 4—5-porate, -colpate or -rupate. Gynoecium often rudi- mentary in staminate flowers; styles 2, elongate, papillo-stigmatic on the inner faces, united at the base, wanting in staminate flowers; ovary ovate, stipitate, tuberculate, 1-loculate, the ovule anatropous to shortly amphi- tropous. Fruit obliquely ovate to oblong, + compressed, keeled on the back, with numerous fleshy protuberances, often inclosed basally by the marcescent calyx, often beaked by the persistent style. Seeds plano- compressed, ovoid, often oblique, testa membranaceous, endosperm ab- sent. Embryo straight; cotyledons + thick, flat, slightly unequal, the apex of the larger hooded and slightly infolding the smaller, the radicle minute, directed toward the linear hilum. Type species: P. aquatica La Gmelin (Anonymos aquatica Walt., nom. invalid., cf. ICBN 1966. Art. 20, 43.) (Named for the German botanist Johann Jakob Planer, 1743-1789.) — WATER ELM, PLANER TREE. A monotypic genus of the southeastern United States ranging along the Coastal Plain from southeastern North Carolina south to northern Florida, west to eastern Texas, and north in the Mississippi Embayment to southeastern Oklahoma, southeastern Missouri, southern Illinois, western Kentucky, and western Tennessee. Planera aquatica appears to be restricted to swamps, margins of sluggish streams or rivers and alluvial flood plains. It is frequently found growing in permanent water up to a meter deep, but can withstand long periods of inundation at greater depths (McAtee). Flowering begins in February and continues into April, with the fruits maturing approximately one month following anthesis. The seed viability has been reported to be very high (McAtee). The seeds are a major food source for waterfowl, especially mallard ducks. A study conducted in Louisiana from October through December on 171 mallard ducks re- vealed that 45.5 per cent of their total subsistence consisted of seeds of 32 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 P. aquatica (McAtee). Other species of ducks occasionally feed upon the seeds, but not as a major food supply (McAtee, Mabbott). The seeds apparently remain viable in standing water, thus supplying water- fowl with an overwintering food. Several species of Planera have been described on the basis of fossil leaves and leaf fragments. Planera longi folia and P. Ungeri are attributed to the Upper and Middle Miocene in the central Rocky Mountain area (Lesquereux, 1878, 1883), and P. Ungeri has been reported by Heer from Alaska. Berry (1924) considered the genus to be an old one, with four Upper Cretaceous species distributed from western Greenland to North Carolina. Also three or four Eocene species are known only from North America and the Arctic localities. Planera is also represented in the Miocene of Europe and North America and in the Pliocene of Europe and Asia. The wood of Planera is considered to be of little or no use economically. In comparison with the wood of U/mus, Gibson found that of Planera to be lighter in weight (specific gravity 0.53), poorer in fuel value, weaker, and more brittle. The wood is described as being soft, light brown in color with almost white sapwood. Planera differs from Ulmus in that the annual rings lack the rows of large open vessel-pores common in U/mus. The genus is closely related to Ulmus but has partially imperfect flowers and a characteristic wingless fruit with irregular fleshy protuberances. Unfortunately little is known of the genus in regard to its overall biology, i.e., floral anatomy, embryology, cytology, and ecology. REFERENCES: Under family references see also BERRY (1924, pp. 158, Rode Grisson (pp. 397-399), JONGMANS (pp. 16-18), Kurz & Goprrey (pp. 107, 108), LEsQuE- REUX (1883, p. 161; 1878, p. 189), HEER (p. 34), PriEMER es 453, 454), and West & Aeeetin (p. 56). Berry, E. L. The lower Eocene oe es southeastern North America. US. Geol. Surv. Prof. Pap. 91: 193, 916. Massortt, D. C. Food habits of seven aw s of American shoal water ducks. U.S. Dep. Agr. Bull. 862: 1-67. 1920. [P. aquatica, 52.] McAteer, W. L. Eleven important wild-duck foods. U.S. Dep. Agr. Bull. 205: 1-25. 1915. [Planera, 9-12.] SARGENT, C. S. Planera. Silva N. Am. 7: 61. pl. 316. 1891 arD, D. B. The genus Anonymos and its nomenclatural survivors. Rhodora 64: 87-92. 1962. [P. aquatica, 131.] Witsur, R. L. The identity of Walter’s species of Anonymos. Jour. Elisha Mitchell Sci. Soc. 78: 125-132. 1962. [P. aquatica, 131.] Tribe CeLTEAE Dumort., “Celtideae” 3. Celtis Linnaeus, Sp. Pl. 2: 1043. 1753; Gen. Pl. ed. 5. 467. 1754. Small to moderate-sized or large trees or shrubs of variable habitats, the bark smooth or often fissured and conspicuously warty; branches un- 1970 | ELIAS, GENERA OF ULMACEAE 33 armed or spinose; buds scaly or naked. Leaves distichous, serrate or en- tire, often oblique at the base, pinnately 3(—5)-veined, petiolate, mem- branaceous to coriaceous, deciduous [or persistent]; stipules lateral, free, usually scarious, caducous. Plants monoecious or rarely polygamomonoe- cious, the flowers vernal, small, pedicellate on the branches of the year, usually functionally imperfect, rarely functionally perfect, the usually lower staminate flowers cymose or fascicled, the carpellate flowers soli- tary or in few-flowered fascicles from the axils of the upper leaves. Perianth imbricate, slightly to deeply 4(5)-lobed, the lobes deciduous. Stamens as many as the perianth lobes, inserted on the usually densely pilose receptacle; filaments subulate, sometimes incurved in bud and spreading elastically, exserted after anthesis; in carpellate flowers usually shorter and included and often nonfunctional, rarely wanting; anthers ovate, dorsifixed just above the emarginate base rag together and face to face in bud, extrorse. Gynoecium in staminate flowers minute and rudimentary; in carpellate flowers style short, pprniere divided into two divergent, elongate, reflexed lobes, the lobes entire or bifid, papillate- stigmatic on the inner face, deciduous; ovary ovate, sessile, 1-loculate; ovule anatropous, with two integuments. Fruit a fleshy drupe, ovoid or globose, the outer part of mesocarp thick and firm, the inner thin and fleshy, the stone thick-walled, bony, smooth or rugose. Seed filling the locule; endosperm scanty, gelatinous, nearly inclosed between the folds of the cotyledons, or wanting; seed coat membranaceous, the chalaza col- ored, close to the minute hilum. Embryo curved; cotyledons broad, fo- liaceous, conduplicate or rarely flat, variously folded, corrugate, incumbent on or embracing the short superior radicle. (Mertensia HBK. non L., Momisia F. G. Dietr., Solenostigma Endl.). Type species: C. australis L., of Europe and Asia. (Name the classical Latin one given by Pliny for an African lotus with sweet berries, possibly Celtis australis.) — HACKBERRY, SUGARBERRY, NETTLE-TREE A genus of 50-60 species widely distributed in temperate and tropical regions of the world, with five species in our range. Because of the com- mon occurrence of intergrading forms, as well as ecological variations, the species are at best poorly defined and are much in need of careful bio- systematic study. The species enumerated here should be considered as tentative. Many of the minor variants have been described at various subspecific levels. ombining mainly vegetative and floral characters with geographical distribution Planchon divided Celtis into four subgenera, two of which, CrLTIs and MomisiA, are represented in our area. Subgenus SPONIOCELTIS, composed of three species with linear, entire stigmas, the flowers borne in cymes with the staminate and/or polygamous flowers borne near the apex of the branchlets, is disjunctly distributed in tropical America and Malesia. Celtis Swartzii Planch., of the Greater Antilles, is closely related to the Malesian C. cinnamomea Lindl., but C. trinervia Lam., attributed to this subgenus by Planchon appears to be placed better in 34 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 subgen. CeLtis. Subgenus SOLENOSTIGMA Planch., characterized by two- lobed or emarginate stigmas and the flowers borne in cymes, is restricted to southern Asia and Australia. The monotypic genus Mirandaceltis A. J. Sharp was established for C. monoica Hemsl., of Mexico, which differs in leaf venation and drupe size and which is perhaps somewhat inter- mediate between Celtis and the Asiatic genus Zelkova. Subgenus CELTIs (subgen. Fuceltis Planch.), widespread, mostly throughout the mild-temperate regions of the world, is characterized by linear, entire stigmas and staminate flowers fasciculate in the axils of new leaves. Three species occur in eastern North America. Celtis laevi- gata Willd. (C. mississippiensis Bosc), 2n = 20, with drupes 5-8 mm. long and narrow entire to somewhat serrate leaves, ranges from south- eastern Virginia, south through Florida, and west to southern and western Texas, northeastern Mexico and southeastern Oklahoma, southern Kansas, Missouri, southern Illinois, southern Indiana, and central Kentucky. Varietas Smallii (Beadle) Sargent (C. Smallii Beadle), of questionable status, differs only in its constantly serrate leaves. Shorter, thicker leaves and more pubescence dubiously separate var. texana (Scheele) Sarg. (C. texana Scheele) from var. laevigata. A shrub or small tree, generally of exposed habitats, with drupes 5-8 mm. long and relatively small, often entire leaves, C. tenuifolia Nutt. is found from southeastern Pennsyl- vania to Indiana, northern Illinois, Missouri and eastern Kansas and south to eastern Oklahoma, Louisiana and northern Florida. Fernald & Schubert distinguished var. georgiana (Small) Fern. & Schub. (C. georgi- ana Small) on the basis of its pubescence and leaves which are coriaceous and scabrous above. A third species in our range, C. occidentalis, 2n = 20, 28, has a more northern distribution, ranging from Massachusetts and New Hampshire to New York, southern Quebec, southern Ontario, cen- tral Michigan, southern Manitoba, Minnesota and central North Dakota, south to western Nebraska, northeastern Colorado, southeastern Wyoming, western Kansas and western Oklahoma, east to Arkansas, Alabama, and Georgia. Varietas pumila (Pursh) Gray (C. pumila Pursh) is based on plants with submembranaceous to membranaceous leaves which are conspicuously inequilateral at the base. Boivin (1967) described var. Soperi from southeastern Ontario and from northern Illinois on the basis of the leaves being regularly dentate. Other species within the continental United States which are assignable to this subgenus are C. Lindheimeri Engl., of central and southern Texas, and C. reticulata Torrey (C. Douglasti Planch.), of the western states (Wyoming and Idaho, west to western Washington, south to Oregon, eastern and southern California, east to Arizona, northern Mexico, New Mexico and western Texas, wes- tern Oklahoma, and Colorado). : Subgenus Momisia (F. G. Dietr.) Planch. with about 15 species in the Neotropics, is centered in Brazil and attains its northernmost limits in southern Florida and the southwestern United States. The flowers have linear, bifid or twice bifid stigmas, and they are borne in racemes. Typically shrubs, the species usually have solitary or paired axillary spines 1970] ELIAS, GENERA OF ULMACEAE 35 in the stipular position. Celtis iguanaea (Jacq.) Sarg. (Momisia igua- naea (L.) Rose & Standl., M. aculeata (Swartz) Klotzsch) is reported from the southwestern coast of Florida and the Coastal Plain of Texas. The smaller- and scabrous-leaved C. pallida Torr. (Momisia pallida (Torr.) Planch.), common in central, western and southern Texas, New Mexico, Arizona, and in Mexico from Chihuahua to Baja California south to Oaxaca, is also known from the western coast of peninsular Florida. Chromosome numbers reported for Celtis (six species) are 2n = 20, 22, 26, 28, and 40. Sax noticed in C. occidentalis very irregular meiotic divisters with little pairing and with the univalent chromosomes irregularly distributed, Although 80 per cent of the pollen was sterile, a large num- ber of fruits were produced. As Sax indicated, if the species contains two different basic sets of chromosomes with only a few homologous chromo- somes or parts of chromosomes it could breed true only by some form of apomixis, a problem in Celtis that has not ee Sie but one which may be crucial in the taxonomy of the g Flowering of Celtis species begins with or oly ‘after the appearance of the leaves in early April to late May. The drupes, ripening in Septem- ber and October, are dark red to purple and are often sought after by animals. The fruits are eaten principally by birds, which in turn aid in the dissemination of the hard stones. Numerous fossilized stones of the drupes of Celtis have been dis- covered in Europe, in Asia (including Japan), and in the western, mid- western, and southern United States. The first records are apparently from the early Eocene in Wyoming and the late Eocene in Georgia (United States) with only one European species. Continuing through the Oligocene, Celtis species are best known from the Miocene (at least nine species). Fossilized material of Celtis from the Pliocene is sparse, and even less is known of the genus in the Pleistocene Perhaps the most obvious disease of hackberries is the characteristic “witches-broom” produced by numerous proliferations at the branch tips. Although the cause is not clearly understood, a mite, Eriophyes, and the powdery mildew Sphaerotheca phytophylia are almost consis- tently associated with outbreaks of the deformations. No effective con- trol is known. Although at least 17 types of insect galls occur on Celtis occidentalis, four caused by species of the genus Pachypsylla (Hemiptera) appear to be most prevalent. Damage to the trees is probably slight, even though the adults overwinter in the cracks of the bark. More serious. threats, when abundant, are the spiny elm caterpillar, which is the larva of the mourning cloak butterfly, Nymphalis antiopa, and the larva of the- hackberry butterfly, Chlorippe celtis, both of which feed upon the leaves.. The hackberry engraver beetle, Scolytus muticus, often is found on the: dead or dying branches. The wood of both species is clear, light yellow with a thick, lighter- colored sapwood, but that of C. occidentalis is coarse grained, while that of C. laevigata is described as close grained. The secondary xylem is 36 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 also characterized by thin medullary rays and numerous small groups of small ducts arranged in intermediate concentric rings. In a comparison of the wood of temperate and tropical species of New and Old World Celtis, Cox and Grumbles independently showed that the species of temperate areas were ring-porous, while the tropical species were diffuse porous, an obvious reflection of the effects of seasonality on the trees. Only C. occidentalis and C. laevigata, of the species in our range, are large enough for the rather soft, weak wood to be utilized to any extent. The wood is used largely for posts, inexpensive furniture, boxes, crates, barrels, and other miscellaneous items. Described as a good tree for shade, species of Celtis are planted as yard and street trees. At least seven taxa, mainly from southeastern Asia and China, have been intro- duced into the United States. REFERENCES: Under family references see also BERNARD, BERRY (1923, pp. 154-156), JONG- MANS (pp. 4-15), FowELts, Gipson (pp. 403-406), Kurz & Goprrey (pp. 105- 107), Vines (pp. 203-207), and West & ARNOLD (p. 55). Avakov, G. S. On the find of Celtis glabrata Stev. and Lithospermum in lower quaternary lacustrine deposits of south Soe. (In Russian.) Dokl. Akad. Nauk SSSR 135(2): 443, 444. 1960. BAEHNI, C. Les Celtis sud-américains. Candollea 7: 189-214. 1936. Berry, E. W. Stones of Celtis in the tertiary of the western United States. Am. Mus. Novit. 298: 1-5. pls. 1-6. 1928. Borvin, B. Les Celtis du Canada. Nat. Can. 94: 621-624. 1867. [C. occt- dentalis & C. tenuifolia.} Brooks, B. W new species of fossil hackberry (Celtis) from the ‘Loup Fork” Miocene of Philips County, Kansas. Ann. Carnegie Mus. 18: 299- 304. pl. 17. Cuaney, R. W. Hackberry ii nies the Pleistocene loess of northern China. Am. Mus. Novit. 283: 927. Cox, M. J. The ee ee of the secondary xylem % five American species of Celtis. Am. Midl. Nat. 25: 348-357. pls. 1-6. 1941. Exsas, M. K. T rien grasses and other prairie vegetation from ae high plains of North America. Am. Jour. Sci. 29: 24-33. pl. 1. 1935. [Stone of Celtis cf. wilson from probably late Pliocene. | Fernatp, M. L., & B. G. Scouspert. Studies of American types in British her- aria. Rhodora 50: 150-176, 181-208, 217-236. pls. 1097-1117. 1948. [The type of Celtis Gresasvres L., 155-162, pls. 1097, 1098. Reprinted as Contr. Gray Herb. 167. Fox, W. S., & J. H. Soper. oe gf eraat of some trees and shrubs of the Carolinian Zone of southern Ontario. Part II. Trans. Roy. Canad. Inst. 30: 3-32. 1953.* [C. tenuifolia Nutt. Gray, J. Temperate Syess genera in the Eocene (Claiborne) flora, Alabama. Science 132: 808— 1960. Grou, H. Paes # in and Satie ss to the Province of Quebec. Canad. Field- Nat. 61: 141, a 1947. [Northern extension of C. occidentalis. | Grumestes, T. L. ee anatomy of the secondary xylem of four oriental species of Celtis. Lloydia 4: 145-152. pls. 1-5. 1941. 1970] ELIAS, GENERA OF ULMACEAE 37 Hit, E. J. Celtis pumila Pursh, with notes on allied species. Bull. Torrey Bot. Club 27: 496-505. pls. 1-4. 1900. Leroy, J. F. La conception synthétique de l’espéce: étude morphologique et biogéographique sur un groupe de Celtis de la section Solenostigma (Ulmacées-Celtidoidées). Bull. Inst. Franc. Afr. Noire 10: 212-234. 1951. Monyjavuze, A. Le groupement a ayes erg! (Celtis australis L.) en Algérie. Publ. Serv. Foréts Algérie. No. 3. 75 pp. pls. 1-7. 1958. Mutter, C. H. Mechanically died bark growth in Celtis. Am. Midl. Nat. 22: 436, 437. 1939. Reip, E. M. Celtis eocenica n. sp. from Réals near Béziers (Hérault). Bull. Soc. Etude Sci. Nat. Béziers 40: 1-18. 2 pls. 1936. [1st record of fossil Celtis from Eocene deposits in Europe. | Rorurock, J. T. Ha “ea sugar-berry (C. occidentalis L.). Forest Leaves 11: 136, 137. 1 pl. 1 SARGENT, C. S. Nettle a Garden ee 39-42. pls. es 1890. ——. Celtis. Silva N. Am. 7: 5. pls. 317-319. 1893. . Notes on North American eee Bot. Gaz. ee 208-242. 1919. [17 taxa of Celtis treated, 10 subspecific taxa described as new. SCHAFFNER, J. H. Twigs of the common hackberry. Ohio Nat. 5: 215, 216. 1904, SCHNECK, J. The hackberries as ornamental and shade trees. Meehan’s Monthly 7: 231, 232. pls. 1-4. 1897. SMITH, : - Hackberry — Celtis occidentalis, Linnaeus. Mich. Conserv. 20: Os 1951.* ee R., & E. B. Hrmeticx. Observations on witches’-broom of hack- berry (C. occidentalis). Pl. Disease Rep. 41: 541-544. 1957.* WELLs, B. W. The comparative morphology of the zoocecidia of Celtis occi- dentalis. Ohio Jour. Sci. 16: 249-290. pls. 12-19. 1916. Wotr, F. A. Formation of adventitious roots by the hackberry tree. Pl. World 13: 174, 175. 1910. 4. Trema Loureiro, Fl. Cochinch. 562. 1790. Evergreen trees or shrubs of tropical forest margins and disturbed areas, the bark usually lightly fissured; branchlets terete, unarmed, variously pubescent, usually with conspicuous lenticels; buds axillary, ours, petiolate, variously serrate [to subentire], pinnately veined, usually conspicuously 3-veined at the base, the veins prominent beneath; stipules free, lateral, subpersistent to persistent [caducous?]. Plants monoecious [subdioecious], the flowers imperfect or polygamous (but then functionally imperfect), articulated with the pedicel, in axillary, subsessile cymes Staminate flowers with the perianth 4- or 5-lobed, the lobes united near the base and induplicate-valvate or slightly imbricate; stamens 4 or 5, the filaments flattened, included to equalling the perianth lobes, the anthers introrse, ovate, dorsifixed below the middle; pollen 2-porate, suboblate, a rudimentary ovary usually present with numerous hairs at its base. Carpellate flowers with 4 or 5 flattened perianth lobes, the lobes united near the base, + imbricate; style central, divided to the base, the two branches linear, minutely plumose-stigmatic on the inner faces; ovary sessile with numerous pilose hairs inserted at the base, 1-loculate, ovule 38 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Fic. f same, vertical section, semidiagrammatic, X 10; g, oat fruit, < 5; h, seed, x 10; i, same, in partial section to show embryo, Xx pS ale Sage td Lamarckiana, leaf, x Y. amphitropous; stamens wanting. Drupe small, ovoid or subglobose, in- closed basally by the persistent perianth, often beaked by the persistent style, smooth [pitted]; exocarp + fleshy, endocarp hard; seed with fleshy endosperm. Embryo curved or nearly involute; cotyledons narrow, ra- dicle folded inward and ascending. (Sponia Comm.) Type species: T. cannabina Lour.; for discussion of the identity of the type and appro- priate synonyms see E. D. Merrill, A Commentary on Loureiro’s “Flora Cochinchinensis” 132, 133. 1935. (Name Greek, trema, hole, in reference to the sometimes pitted drupe.) A primarily pantropical genus of some 45 to 55 species with two, Trema Lamarckiana (Roem. & Schult.) Blume and T. micrantha (L.) Blume (T. 1970] ELIAS, GENERA OF ULMACEAE 39 floridana Britton) occurring in our area. Trema Lamarckiana is found in southern Florida (Dade and Monroe counties), including the Florida Keys, and from Bermuda and the Bahamas to the Greater Antilles and into the Lesser Antilles from Saba to St. Vincent. Readily distinguished from T. Lamarckiana by its larger leaves which are not scabrous beneath, T. micrantha is widespread in the West Indies and coastal Mexico south- ward to Central and South America and occurs in our area from the Florida Keys northward to central peninsular Florida. The species of Trema are highly variable and often confusing, and the taxonomy of the genus is poorly understood. Characterized by their spreading crowns and horizontal to slightly drooping branches, species of Trema are generally secondary forest trees and usually indicators of secondary succession. In a study of succession in old clearings on Barro Colorado Island (Panama), Kenoyer (1929) found that 7. micrantha, along with Cecropia, Apeiba, Ochroma, and Cordia were the first trees to establish themselves. Other species of Trema have been reported as secondary forest indicators or major components in various stages of secondary succession in western Africa and Malaya. The rapidly growing trees are also found along forest margins and in disturbed areas, both natural (e.g., gravel bars) and man-made (road- sides). Cytologically the genus is poorly known. The chromosome number of Trema orientalis Blume has been reported as 2n = 20, 36, 40 and that of T. cannabina as 2n = 40. Anatomically Trema is closely related to Celtis. Of the ulmaceous genera tested by Stake (1931) only Trema and Celtis had cystoliths in the upper epidermis, mainly in association with the hairs. Mucilage cells are common in the mesophyll of all species of Celtis and of many of Trema, but are lacking in the two Trema species in our range. Little is known of the embryology and floral biology of rema. Flowering and fruiting appear to occur throughout most of the year with the flowers often noticeable in early spring (late March and early April). The minute greenish flowers are soon followed by tiny orange to bright yellow (?) drupes. The wood has been described by Little & Wads- worth (1964) as light brown, soft, weak, lightweight and of limited use. In Puerto Rico the wood is occasionally used for fence posts and fuel and the strong fiber in the bark for cordage. The inner bark is light brown or pinkish and is tasteless to slightly bitter. REFERENCES: Under family references see also NEVLING, 108-110; PRiEMER, 467-470; and WEstT & ARNOLD, 51. Coompse, D. E. An analysis of the growth of Trema guineensis. Jour. Ecol. 48: 219-231. 1960. GARDINER, R. C. Studies in the leaf anatomy and geographic distribution of Trema. Unpublished master’s thesis, Univ. Connecticut. 59 pp. pls. 1-5. 1965. 40 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Kenoyver, L. A. General and successional ecology of the lower tropical rain- forest at Barro Colorado Island, Panama. Ecology 10: 201-222. 1929. LauTERBACH, C. A. Die Ulmaceen Papuasiens nebst einer Revision der Trema- Arten des Monsun-Gebietes. Bot. Jahrb. 50: 308-327, 1913. [Taxonomic treatment of 13 taxa of se. Asia. | Littte, E. L., Jr., & F. H. WapswortH. Common trees of Puerto Rico and the Virgin Islands. U.S. Dep. Agr. Aer. Handb. 249. 1964. [T. Lamarckiana, T. micrantha, 56, 58, pls. 15, 16. PENNINGTON, T. D., & J. SaRUKHAN. Arboles tropicales de Mexico. vii + 413 pp. Mexico. 1968. a micrantha, 118, 119. illus. & map. | SATAKE, Y. Systematic and anatomical studies on some Japanese plants. I. Systematic importance of spodograms in the Urticales. Jour. Fac. Sci. Univ. Tokyo Bot. 3: 485-511. 1931. [T. orientalis, T. amboinensis, 493, 494.] ARNOLD ARBORETUM OF HARVARD UNIVERSITY 1970] HOWARD, ELFIN FOREST, 10 41 THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 10. NOTES ON TWO SPECIES OF MARCGRAVIA RicHaArp A. Howarp THE TYPE GENUS Marcgravia of the Marcgraviaceae is familiar to col- lectors in the American tropics for its conspicuous heteroblasty and for the development of pedicellate saccate nectaries at the apex of the dense race- mose inflorescence, the flowers of which usually form a distinctive umbel- like cluster. The plagiotropic juvenile shoots of all species of Marcgravia examined in the West Indies have distichous leaves arranged on flattened stems which spread over the ground, over and on boulders, and up the trunks of trees (Ficure 1). Although the juvenile leaves vary in form and size, they have characteristics which permit them to be identified with particular species. In most species the juvenile leaves are subsessile and often overlap the stem (FicuRE la). Numerous adventitious roots develop in rows from the stem beneath the leaves and afix the young shoot to the substratum (Ficure 1b). Sir Hans Sloane in 1707 illustrated the climbing shoots and indicated that they were sterile and that he was unable to as- sociate them with known plants. He listed them with ferns and aroids. With increasing age and with unknown factors of maturity, freely arching orthotropic branches lacking adventitious roots develop from the axils of the juvenile leaves (FicuRE 1c). These axillary stems bear leaves of en- tirely different size and shape. The apex of the climbing juvenile shoot may also change suddenly, or with transitional phases, to the production of leaves of the mature form, usually when freed by weight or other forces from the substratum. The terminal free shoot or the axillary branch may, upon contact with the trunk of a tree or other substrata, revert to the juvenile leaf form. Examples have been found, and even reproduced in a greenhouse, of the repeated transfer of a single stem from juvenile to ma- ture form and back again (Ficure 1d) Stems of Marcgravia may reach the canopy of an elfin forest and extend over the crowns of other plants or ascend the trunks of forest trees, forming masses in the branches of the crown, and produce long axillary shoots, pendent or reaching nearly to the ground. The inflorescence of species of Marcgravia in the West Indies is de- veloped terminally on the axillary branches. After flowering and producing fruit, the flowering branch may die back a few internodes before an axillary bud on the same branch develops into a new shoot, continuing the elongation of the primary axis. The apical meristem of Marcgravia is surrounded and protected by overlapping leaves which form a slender pointed bud, enlarged at the base (FicurE 3a). In primordial stages the leaf is terminated by a relatively large multicellular gland often con- spicuous in the terminal bud (Ficures 2c, 4c, 5c). Similar glands may be found along the margin of the lamina (Ficure 2c). These glands are con- 42 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ma \ 3 i j nf (9 — <=. = Ficure 1. A plant of Marcgravia sintenisii grown in the greenhouse of the Arnold Arboretum in Jamai lain, Massachusetts. a. Plagiotropic stem wi e rm : An orthotropic stem which changed from the basal juvenile leaf form to the adult leaf form and reverted to the juvenile leaf form on plagiotropic stems when appressed to the wall. Ficure 2, left. Marcgravia sintenisii. a. Orthotropic stem apex with flower bud. b. Leaves removed to show young inflorescence. c. The inflorescence is 4 raceme. The nectary-bearing pedicels are contiguous with the flowering pedicels at this stage of development. Marginal hypophyllous glands are shown on the leaf. d. Young flower bud showing large marginal glands of the incipient peri- anth parts. e, f. Cleared nectaries. g. Two young nectaries showing the aborted flower bud. Ficure 3, right. Marcgravia rectiflora. a. Orthotropic stem apex with flower bud enclosed in leaves. b. The young inflorescence with nectary-bearing pedicels clearly differentiated at the apex of the raceme. c. The young inflorescence. d A cleared young nectary showing the early stages of invagination and distortion. [OL61 OT ‘LSHYOA NIATA ‘davMOH 44 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 spicuous in the juvenile leaf form and appear to remain functional for a long period of time. In the adult leaf form the terminal and marginal glands become brown and hard at an early stage of development. Large hypophyllous glands are found in characteristic patterns on the leaf blade, associated with the midvein or localized in particular spots on the lamina (Ficures 4, 5). The terminal bud containing a young inflorescence is only slightly more swollen initially than the vegetative bud. There is only a slight reduction in leaf size immediately below the inflorescence. In the raceme the flowering pedicels are borne in a dense spiral, but the axis is terminated with the development of pedicels dominated by the lateral saccate nectary which has its orifice at the base (FicurEs 2c, 3c). The development of the ter- minal nectary-bearing pedicels is precocious in comparison with the flower- ing pedicels. The flowering pedicels are ascending in relation to the axis in the young buds and the flowers are borne apically on the pedicel. With continued development the flowering pedicels tend to diverge from the inflorescence axis, in some species to a 90-degree angle, or even to be- come slightly reflexed. The apical flower primordium retains the same axis as the pedicel in the subgenus ORTHOTHALAMIUM (Wittmach, 1878). In other species forming the subgenus PLaciorHALamiuM the flower bud is offset from the apex and may be at a 90-degree angle to the axis of the pedicel. In many species, as in Marcgravia rectiflora, the flowering axis remains erect and the flowers ascend (Ficure 6). The terminal nectary-bearing pedicels, therefore, recurve to varying degrees so that the basal opening of the nectary is oriented upward. In other species, as represented by Marcgravia sintenisii, the inflorescence axis is pendent so that the flowering pedicels form an umbel with the flowers pointed downward (FicuRE 7). The axis of the nectary-bearing pedicels, therefore, is not distorted and the basal opening of the nectary is directed upward. This is the characteristic “wheel” of hanging flowers so frequently illustrated as typical of Marc- gravia, which has given rise to suggestions and speculations that the plants are pollinated by humming birds hovering beneath the open flowers and becoming dusted with pollen while obtaining nectar from the saccate ovaries. Within the many species of Marcgravia nectary-bearing pedicels show considerable variation. The nectary may be nearly as long as the pedicel and is then often described as sessile, or it may range from three-fourths to only one-fourth the length of the pedicel. The majority of species have elongate cylindrical nectaries with an asymmetrical orifice and lip, a nar- rowed neck, and a swollen base. A few species have flattened and broad- ened nectaries, described as galeate but to be considered so only when the inflorescence axis is ascending, The terminal pedicels producing nectaries may also bear a minute or abortive flower, suggesting that they are but modifications of the lower, nectary-less, flowering pedicels. The nectary-bearing pedicels do not persist long after the flowers have shed their calyptrate corollas. No fruiting specimens were found in the 1970] HOWARD, ELFIN FOREST, 10 45 wild or in herbarium collections with the nectary-pedicels remaining at- tached. Since the shape of the nectary is a character of taxonomic im- portance, the need for collections of flowering material is apparent. The morphology of the nectary and its role in pollination are contro- versial and still undetermined in the available literature for many of the species. Additional field observations are needed. Patrick Browne, in The Civil and Natural History of Jamaica, published in 1756, observed on Marcgravia that “this curious plant is frequent in the woods of Jamaica; and appears in such various forms, that it has been often mistaken for different plants, in the different stages of growth.” Of the inflorescence he reported, “The flowers are sustained by long foot-stalks, and disposed in the form of an umbella, about the extremities of the branches: but the sum- mit, or crown of the supporter, is constantly adorned with four, five, or more hollow, divergent, glandular bodies, that occupy the center of the umbella: these are of an arched oblong form, obtuse and roundish; they are hollow within, and affixed by very short foot-stalks, that rise im- mediately from one side of the aperture, or opening of the gland; which is 5 WR Ficure 4, left. a, b. Leaf of safe eha ute s ee athe the several hypo- ee glands near the base of the lam The terminal gland of young lea ft pgp ani section of the leaf tiie Hilal a gland. Fic cul right. a, b. Leaf lela ie rectiflora showing the two large glands near ir base of the lam c. Apex of young leaf. d. Diagrammatic section of the leaf blade facet a age nd. so disposed, as to receive the water that dribbles down along the branch in rainy weather. What the real use of these may be, is not easily determined: it is, however, remarkable, that the leaves of the branches are plain, of an oblong oval form, with a smooth membranous edge; while those of the younger plants are always observed to have many little glands, set gradually round the margin.” Thomas Belt, author of The Naturalist in Nicaragua, published in 1874, 46 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 is FAC a Wy lom. Ficure 6. a. The erect inflorescence of Marcgravia rectiflora. Flowers arin the ra bec subtending a four-parted calyx, the calyptrate corolla, and t numerous stamens which have shed pollen before the fall of the corolla. Bee : eee wn. The unaltered bract com- parable to the modified nectary bract is also shown. is considered to be the originator of the humming-bird pollination legend. He reported of Marcgravia nepenthoides, “The flowers of this lofty climber are disposed in a circle, hanging downwards, like an inverted candelabrum. From the centre of the circle of flowers is suspended a number of pitcher- like vessels, which, when the flowers expand, in February and March, are filled with a sweetish liquid. This liquid attracts insects, and the insects numerous insectivorous birds, including the species I have mentioned and many kinds of hummingbirds. The flowers are so disposed, with the stamens hanging downwards, that the birds, to get at the pitchers, must brush against them, and thus convey the pollen from one plant to another 1970] HOWARD, ELFIN FOREST, 10 47 A second species of Marcgravia that I have found in the woods around Santo Domingo has the pitchers placed close to the pedicels of the flowers, so that the birds must approach them from above; and in this species the flowers are turned upwards, and the pollen is brushed off by the breasts of the birds.” This story has been rewritten, applied to other species without verification in many subsequent volumes, although most recently qualified by Meeuse (1961) who states, “In all likelihood birds are the normal pollinators in Marcgravia, although bat pollination is suspected in a few cases.” A paper by I. W. Bailey (1922) entitled “The pollination of Marcgravia: a classical case of ornithophily?”, which questioned the gen- eral acceptance of humming-bird pollination, has been generally overlooked. Bailey examined several species of Marcgravia in British Guiana and con- cluded the flowers are self-fertile or autogamous, instead of being protan- drous and cross-pollinated by birds. He found that inflorescences would set fruit when enclosed in protective cloth which excluded any birds, and he reported that other inflorescences produced fruit under water. During a recent period of field study in eastern Puerto Rico two species of Marcgravia representing the two subgenera and matching the types de- scribed by Belt were available in quantity for frequent observation. One species endemic to Puerto Rico, Marcgravia sintenisii (FIGURE 7), occurs frequently at higher elevations in the Luquillo Mountains and was in the Pico del Oeste study area (Howard 1968). A second species, Marcgravia rectiflora (FicurE 6), has a wider range of distribution including Cuba and Hispaniola, and is reported from South America; it occurs at a lower elevation in Puerto Rico, primarily along the Rio Cubuy in the Luquillo Mountains. The species may be contrasted in other ways. Marcgravia sintenisii occurs in and on the canopy of the elfin forest. It has a pendent inflores- cence developed at the apices of short axillary branches. The flowers, therefore, hang and the stamens are exposed from below. The nectaries are flattened and galeate in normal position but appear flat-topped at the orifice and obtuse at the base, as observed in the flowering condition (Ficure 7d). The nectaries are pale yellow-orange in color until the corolla cap is ready to break free, when they turn a brilliant red. The flowering pedicels number 30 to 60 and the nectary-bearing pedicels about nine to eleven. The corolla caps fall in the early morning hours and the stamens diverge from the pistil by adaxial growth or elongation. The stigma appeared to be receptive, i.e., glistening and dusted with pollen at the time of abscission of the corolla cap, and within an hour or two after opening the anthers had shed all pollen and begun to wither. The flowers did not mature in sequence according to their position on the axis, but one radial group appeared to shed the corollas, followed by another segment often separated from the first by flowers slower in development. The whitish filaments persisted but were easily dislodged at this stage. Although it appears that the flowers are pollinated in bud, the inflorescence as a whole had still unopened flowers at the same time other flowers had already lost their stamens. In the course of observations made over a period of three 48 JOURNAL OF THE ARNOLD ARBORETUM {vou. 51 Ficure 7. Marcgravia sintenisii. a. A young erect inflorescence showing the elongation of the axis between the flower-bearing pedicels and the nectary-bear- ing pedicels. The young flower buds are oriented with the axis of the pedicel. Ab i and the flower is now at right angles to the axis of the pedicel. Further shift in orientation of the nectary is evident. c. Diagram of cleared young nectary showing the aborted flower bud. d. The inverted galeate nectary split to show the glandular tissue. years we noticed that the inflorescences were visited by the green-throated hummingbird (Sericotes holosericeus), by Todies (Todus mexicanus), and by bees seeking nectar from the orifice of the nectary. The most frequent visitor, however, was the bananaquit (Coereba flaveola) which grasped and hung from the brilliantly colored nectaries and pecked at the base (morphological apex) creating sizeable holes in and through the tissue of the nectary. Subsequently the bees also visited these injured areas in- stead of approaching the nectaries at the natural orifice. Marcgravia rectiflora is a stronger climbing plant of greater volume, which does not reach the canopy of the forests of lower elevations but seems to dominate many of the host trees by growing luxuriantly on the lower branches. The lateral branches of this species are exceedingly long and pendent, some reaching 15 feet and showing as many as 25 inflorescence scars. The inflorescence in Marcgravia rectiflora also develops terminally, but instead of hanging the tips of the branches turn upwards (FicurRE 6). The raceme, therefore, is morphologically erect and the flowering pedicels ascending. The nectaries developed on the terminal pedicels are strongly recurved and hang between the flower-bearing pedicels so that the opening of the nectary points skyward. The nectaries of Marcgravia rectiflora are elongate saccate structures larger at the base than at the orifice and nar- 1970] HOWARD, ELFIN FOREST, 10 49 ba< (> { 2 Mes Ficure 8, above. Four diagrams to represent the transitions in form of the intenisii. IGURE 9, below. Marcgravia rectiflora. a. A young nectary-bearing pedicel showing the paired vascular strands, one to the aborted flower bud, the other showing the recurved path associated with invagination of the bract. b. A dia- gram of the vascular pattern found in the clone of Marcgravia rectiflora show- ing the early departure of the vascular supply to the nectary-bract, contrasted with the vascular supply to the normal bract and the flower bud. rowed to a neck in the middle. The orifice is asymmetrical and the lip slightly recurved and pointed. The flowers number about nine to eleven and the nectary-bearing pedicels about four. The flowers are three to four times the size of those of Marcgravia sintenisii and shed the calyptrate corolla in a similar haphazard pattern, unassociated with the position of the flower on the inflorescence axis. Again, the stigmatic area was glisten- ing and possibly receptive when the corolla-cap fell. The stamens also 50 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 spread widely, but the anthers shriveled within a few hours and became dark brown. The filaments were easily detached. The position of the nectaries of Marcgravia rectiflora in relation to the flowers is such that pollination can not be effected by a hovering humming- bird in the theoretical manner but could occur in the way that Belt de- scribed birds acquiring pollen on their breasts from erect flowers. In fact no hummingbirds or todies were seen to visit the inflorescences of M. rectiflora in the course of many hours of observation. The bananaquit, however, was a frequent visitor, here landing in the middle of the inflores- cence, forcing its way to the recurved nectary-bearing pedicels, and tip- ping its beak into the nectary. The bananaquits also managed to tear the nectaries from the orifice to the base, laying open the glandular tissue, or literally removing the globular end of the nectary. In the process of this activity the bananaquits brushed against many of the open flowers, and stamens were dispersed in all directions. It is possible that cross-pollination between newly opened flowers could be effected in this manner. Subse- quently, bees which visited the inflorescence took advantage of the injured nectaries and approached the damaged tissue area directly from well below the flowers. In contrast to those of Marcgravia sintenisti, the nec- taries of M. rectiflora do not develop a brilliant color but remain green and brown Bees, wasps, and hornets which visited the young inflorescences of Marcgravia rectiflora did not approach the cluster of flowers from above but landed below the flowering pedicels and worked their way up the axis to the nectary glands. In many cases the bees, wasps, and hornets did not bother with the nectary orifice but confined their feeding to the “corky” excrescences developed on the pedicels or on the outer surface of the sac- cate nectary. During the course of frequent observations it was noted that the hanging axillary branch and the inflorescence of Marcgravia rectiflora were the preferred territory of an anolid lizard (Anolis evermanni), which spent many hours on successive days in the middle of the inflorescence, but scrambled back up the stem at times while foraging. Like the bananaquit, the lizard was seen to crawl over open flowers and to break off stamens. It is interesting to speculate that a few flowers of Marcgravia rectiflora were pollinated by an anolid lizard. The anolid found the inflorescence 4 good hunting area and was seen to catch small insects apparently approach- ing the flowers. On one occasion the anolid caught a butterfly, but the other insects remain unidentified. Mr. Henry Draper, with the aid of a telephoto lens, was able to record an anolid lizard with its snout in the orifice of one of the nectaries. Again the lizard probably was not supping on the nectar but catching small red- brown ants and mites which have been found within the nectaries in sub- sequent dissection. The lizard was missing from his favorite territory one afternoon when I planned to watch his behavior again. Nearly an hour of observation time passed before I realized that his usual haunt was then occupied by a snake (Alsophis portoricensis or Dromicos exiguus). This 1970] HOWARD, ELFIN FOREST, 10 51 snake was stretched along the full length of the branch and its head was in the center of the inflorescence. Neither the snake nor the lizard was seen on this branch in succeeding days. I. W. Bailey (1922) observed the flowering characteristics of several species of Marcgravia in British Guiana and found that bagged inflores- cences set fruit, as did some stems that flowered under water. The natural fertility of the two species in Puerto Rico was also high. Several inflores- cences that were covered tightly with bags of cheese cloth during the summers of 1966 and 1967 produced developing fruit on as high a per- centage of the flowers as did unbagged inflorescences. Swamy (1948) sup- ported Bailey’s observation that the anthers of Marcgravia split and re- lease pollen grains before flower anthesis. He also noted that a large per- centage of the pollen had germinated at anthesis and that pollen tubes of varying length were found. Some pollen tubes were observed on pollen collected from open flowers of both species occurring in Puerto Rico. It seems unlikely that pollen grains which had developed pollen tubes could be transported any distance in the process of cross-pollination. Swamy reiterated Bailey’s conclusion that the two species they studied were autogamous or self-pollinated. This seems to be true of the two species studied in Puerto Rico, M. sintenisii and M. rectiflora. In both species the anthers were open and pollen was shed within the bud before the calyptrate corolla had fallen. The stigmatic area, however, did not appear to be receptive until after the loss of the corolla, at which time it was glistening and covered with adhering pollen. Recently Meeuse (1961) suggested that Marcgravia may be pollinated by bats. No bats have been observed in the Pico del Oeste area by the individuals who participated in that study. Although the participants spent relatively few nights in this cloudy and cool area, it is hard to believe that bats would do so either. There are bats in the area where Marcgravia rectiflora was studied, and a reasonable at- tempt was made to observe the few open flowers of M. rectiflora in the pre- dawn and late evening hours. No bats were seen to visit the inflorescences under observation. Bailey pointed out in 1922, “There has been considerable speculation concerning the origin and morphological significance of these nectarif- erous appendages. Are they metamorphosed bracteoles, abnormal pedicels, modified bracts, or appendages sui generis? Most recent students of the Marcgraviaceae have accepted Planchon and Triana’s (1863) conclusion that they are evaginated bracts. It must be admitted that there is con- siderable evidence in favor of this view.’ He then noted, as had Goebel (1905), that “the concave or inner surface of the nectariferous appendages is the morphological equivalent of the under surface of the leaves. The outlets or pores of the hypophyllous glands are located in this dorsal sur- face and, accordingly, discharge their sugary excretions into the con- cavities of the bracts.’”’ However, Bailey also concluded, ‘That the nec- taries of Marcgravia are not abnormal pedicels, as maintained by Seemann (1870), but are compound structures resulting from the fusion of a nectariferous bract and a sterile pedicel, is indicated, not only by their 52 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ficure 10, left. A diagrammatic reconstruction of the nectary-bearing pedi- cel of the clone of Marcgravia rectiflora which develops a flower bud. Prepara- tion was made from cleared pedicels and from transverse sections. Two elon- gated hypophyllous glands are evident. The secondary vascular supply to the nectary area is derived from the full length of the bundle which ascends, curves over, and descends into the outer area of the flask-shaped nectary. The ab- i of a bract above the nectary contrasts with the presence of the opposing ract. Ficure 11, right. The galeate nectary of Marcgravia sintenisit diagrammatical- ly reconstructed from cleared nectaries and from transverse sections. The small vascular strand to the aborted flower bud is shown. An ascending trace with secondary branches curves at the top of the nectary before descending to the morphological apex of the bract where a rudiment of the apical gland is shown. bs flattened nectary has a wide orifice. Glandular tissue and its orifice are shown. 1970] HOWARD, ELFIN FOREST, 10 53 external morphology, but also by their internal anatomy.” Bailey found “two distinct systems of fibro-vascular bundles in the nectaries, one belong- ing to the sterile pedicel and the other to the adnate bract.” This state- ment is not completely accurate when applied to the two species used in the current study. The floral structure of the two species of Marcgravia studied in Puerto Rico shows a calyx consisting of two pairs of overlapping, decussate sepals subtended and appressed by two smaller bracts (FicurEs 6, 7). In the flowering pedicel the vascular traces to these appendages depart from the central stele at about the same level and immediately below the vascular bundles which supply the corolla, androecium, and gynoecium. In the nectary-bearing pedicels, however, the flower is usually aborted and repre- sented only by a small rudiment which has a very reduced vascular supply (Ficures 8c, 9a). The bulk of the vascular tissue is directed to the nectary with the median bundle arching over at the apex of the nectary and de- scending to the lip and its terminal gland (Ficures 8-11). Branch traces depart from both sides along the full length of this bundle, branching as they diverge into the body of the nectary. The vascular pattern can be observed readily in cleared nectary-bearing pedicels of Marcgravia recti- flora. (Ficures 9b, 10). Numerous branched idioblasts and sclereids develop early in the nectaries of Marcgravia sintenisii, and in such pro- fusion that only the main vascular bundles can be seen. The secretive tissue is developed extensively in these nectaries. Bailey suggested that these were comparable to the hypophyllous glands found on the leaves, and the present observations support this conclusion. Two ex- tensive areas of secretive tissue open through two pores in Marcgravia rectiflora (FicurES 10, 12). Additional areas of secretory tissue, each with pores opening internally to the cavity, have been found in the nectaries of M. sintenisii (FicureE 13). The presence of rudimentary flowers at the apex of the pedicels which also bear the saccate nectaries has been noted frequently. Both Wittmack and Bailey illustrate pedicels in which the flower is of significant size and in some cases separated by a respectable distance from the nectary. One clone of Marcgravia rectiflora found and observed in Puerto Rico had well- developed flowers at the ends of nectary-bearing pedicels (Ficures 6, 9, 10). The vasculature and glandular tissue of a nectary of this plant is illustrated (FicurE 10). This clone was observed many times over a period of three years and there was no variation in the production of large flower buds. During development the flowers of the nectary-bear- ing pedicels and those without nectaries progressed at comparable rates until each was about two-thirds the mature size. At this point the flowers with subtended nectaries failed to increase further. The regular flowers enlarged, the corolla was shed, and fruit was set. The nectary-bearing pedicels fell off without the terminal flowers maturing or shedding the corolla. Examination of the flowers subtended by nectaries showed nor- mal floral structure with well-developed stamens, the anthers of which contained mature pollen grains. When these structures were cleared and 54 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 2% oie HRS 13 cel of Marcgravia rectiflora showing the opening into the cavity. : Ficure 13, below. A median transverse section of the nectary-bearing pedi- cels of Marcgravia sintenisii. Three areas of glandular tissue are shown. the vascular structure was examined, it was evident, as shown in the dia- gram (FicurE 10), that a major vascular supply extended to the flower and that a branch of this system departed early, ran parallel to it, and ascended into the nectary area. This was conspicuous in contrast to nectary-bearing pedicels with abortive flowers where the vascular sup- ply to the nectary surpassed in volume the vascular supply to the aborted flower. In each of the types of pedicels a single unit of vascular stele was present at the base of the pedicel, and the vascular supply to the nectary or the bract was but a branch of this central stele. The normal flower of both species of Marcgravia has a four-parted calyx closely subtended by two bracts. The pedicels which bear nec- taries, by contrast, possess but one normally developed bract opposite the 1970] HOWARD, ELFIN FOREST, 10 55 nectary area (FicurE 10). The conclusion of Planchon that the nectary is a development of one of the bracts is verified. Although Planchon suggested this as an evaginated bract, the vocabulary of “fusion” or “adhering” remains in the literature. We have been able to examine a series of inflorescences of increasing age. Ficures 2, 3, 8, and 9 show the adjustment in form that takes place in the maturation of the nec- taries of both M. rectiflora and M. sintenisii. The distortion of form is gradual but consistent, leading to the respective saccate and galeate mature nectaries. The development pattern observed also supports the Planchon description of an invagination of one of the bracts which involves a distortion of the major part of the pedicel. ACKNOWLEDGMENTS I wish to acknowledge again the support of this study which was made possible primarily by a grant from the National Science Foundation (GB: 3975). A portion of the study was conducted on the property of Mr. Joseph B. Martinson in Puerto Rico; to him the collaborators in this study and I are grateful for generous hospitality. Patient observa- tions over many hours were made by Judy Appenzeller; Mr. and Mrs. Henry Draper, and Madelon Gauer. Dr. and Mrs. Richard Wagner col- lected special materials following frequently repeated requests. Mrs. Helen Roca Garcia prepared the technical preparations and the draw- ings. LITERATURE CITED BAILEY, I. W. The pollination of Marcgravia: A classical case of ornithophily? m. Jour. Bot. 9: 370-384, 1922. Bett, T. The Naturalist in Nicaragua. London. 1874. BROWNE, P. The Civil and Natural History of Jamaica, p. 244. pl. 26. 1756. ice K. Organography of Plants. Balfour translation. Oxford. 1905. WARD, . The ecology of an elfin forest in Puerto Rico, 1. Introduction and composition studies. Jour. Arnold Arb. 49: 381-418. Jue, H. O. Beitrage oe ae der Marcgraviaceen. Bihang. Sv. Vet.-akad. Handl. Afd. 3. 12: MEeusg, B. J. D. The lon of pollination. Ronald Press, N.Y. 1961. PLANCHON, J. E., & J. Trtana. Sur les bractées des Marceraviées. Mém. Soc. Sci. Nat. Cherbourg 9: 68-88. 1863 SEEMANN, B. C. A new pitcher plant from Central America. Seemann, Jour. Bot. London 8: 245, 246. 1870. StoaNeE, H. A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica etc. London. : Swamy, B. G. L. A contribution to the embryology of the Marcgraviaceae. Am. Jour. Bot. 35: 628-633. 1948. Wirtmack, L. Marcgraviaceae, in Martius, Flora Bras. XII. 1: 215-258. 1878. ARNOLD ARBORETUM Harvarp UNIVERSITY 56 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 11. THE LEAFY HEPATICAE OF PICO DEL OESTE* Marcaret FuULFoRD, BARBARA CRANDALL, AND RAY STOTLER THE SUMMIT oF Pico pEL Oxrste in the Luquillo Mountains of Puerto Rico, with an elevation of 1050 meters, is covered by an elfin forest or thicket from 2 to 5 meters high which in places, especially those covered by the low shrub, is almost inpenetrable, while in areas of taller trees the going is less difficult. In any case, the canopy is more or less closed throughout and even during sunny periods the light is diffuse. The rain- fall is about 200 inches per year. The area in which the collections were made has been under intensive eee || : FIGURE 1. A diagram indicating the habitats and substrata available to leafy Hepaticae in the elfin forest: I, leaf; II, twigs and branches; ILI, trunk of tree; IV, bromeliad epiphyte; V, shrub; VI, tree fern; VII, the ground; VIII, log. = ui “4 mt _ a . 1 ges y —— a Weata E> 1 - ; ; Work relating to a project supported by National Science Foundation Grant, No. GB 7138, Margaret Fulford, principal investigator. “I 1970] FULFORD ET AL., ELFIN FOREST, 11 5 B leaf covered with several species of leafy Hepaticae. FicurE 2A. Portion of a : trunk covered with a loose mat of leafy Figure 2B. Portion of a tree Hepaticae. 58 JOURNAL OF THE ARNOLD ARBORETUM [vox. 51 investigation by Dr. R. A. Howard for a number of years and has al- ready been described in detail (Howard, 1968). Most of our collections were made in the vicinity of his study trail. As pointed out by many investigators (see Howard, 1968) such a type of woody vegetation has been variously described as elfin forest, elfin woodland, montane thicket, and mossy forest. At least in Puerto Rico, this “mossy forest” on Pico del Oeste, El Yunque, Mt. Britton, and El Toro, is mossy only in appearance, for leafy Hepaticae comprise 90 percent or more of the bryophyte flora of these areas. While the general aspect of the vegetation of these four peaks is similar (Howard, 1968), the actual composition of the leafy hepatic flora is different for each of them. This will be demonstrated in a later paper. These leafy hepatics occupy all sorts of niches and substrata (FIGURE 1), and each may support several growth forms. The plants cover the ground, often in cushions up to a foot thick, cover the tree trunks and branches in large or small appressed mats, loose mats, deep tufts, large “balls” to 6 inches or more in diameter, or hang in festoons to two feet or more in length (Ficurrs 2B, 3), or cover the leaves of the trees and shrubs, herbaceous plants, and ferns (FicuRE 2A). In a general way, the variety of substrata and growth forms may be classified in the following manner: I. LEAVEs. a. Species with a tightly appressed growth habit, — in particular, young plants developing from discoid gemmae (gemmalings), in the genera Colura, Cyclolejeunea, Diplasiolejeunea, and Radula, or from tiny small-leaved branches in the genera Drepanolejeunea, Prionolejeunea, and others. b. Loose mats, usually on older leaves. Large colonies of the plants named under “a” show this form. Additional gemmae and tiny branches are caught in such mats. c. Tufts to 1 cm. high often become established within the older mats, €S- pecially the genus Adelanthus. II. Twics AND BRANCHES. E a. Appressed mats—of Ceratolejeunea spp., Prionolejeunea spp., Frullania, Radula, etc. . b-c. Loose mats to deep tufts — of Trachylejeunea, Bazzania longa, B. longi- stipula, Lepidozia patens, Plagiochila spp., etc. ; d. “Balls” to 6 inches across —of Bazzania stolonifera, B. breuteliana, Lepr dozia patens, Plagiochila spp. e. Pendent forms, to a foot or more long—of Herberta, Cystolejeuned, Plagiochila spp. and Syzygiella. III. TREE TRUNKS. These support a variety of growth forms, limited only by the time and space available. The forms and species are similar to those of twigs and branches. IV. BROMELIAD EPIPHYTES. a. Appressed forms, especially small plants of Colura, probably developed from gemmae. V-VI. SHRUBS AND FERNS. a. Appressed forms — on the leaves and petioles. The stems support the same FIGURE 3. Tree trunks in the elfin forest with leafy Hepaticae in loose mat s or pendent. L6T Lo “LSHUOA NIATA “IV LH GaOATAA IT Ut \o 60 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 range of growth forms and species as are to be found on the tree trunks and branches. VII. THE GROUND. a. Appressed forms or low tufts — occupy spots of recently exposed soil, as Alobiellopsis. b. Deep tufts and mounds — cover most of the area and form a deep, spongy, soggy, cushion over the forest floor. VIII. Logs. a. Appressed forms —a few slender forms as Nowellia, Telaranea and Cepha- lozia soon become established but they are soon replaced by the rapidly growing larger species of the tree trunks and ground. Most of the species are tinged with brown and some become bronze. A few are reddish to deep purple-black. Many individual plants, especially those on leaves, are subjected many times a day, to a deluge of rain followed immediately by high wind, sun, and a rapid drying out, so that they survive frequent periods of xerophy- tism. This is particularly true of the unistratose, discoid gemmae and the gemmalings of the leaf-inhabiting species of Colura, Cyclolejeunea, and Diplasiolejeunea which are so abundant. Those plants in colonies are not so exposed. Leafy Hepaticae may reproduce by spores, gemmae of various sorts, by specialized deciduous branches, by short, small-leaved branches, and by fragmentation of a stem or leaf. The first arrivals on a new leaf are, for the most part, the discoid gemmae of Colura, Cyclolejeunea and Diplasiolejeunea. Both Colura clavigera (FicurE 4) and Cl. rhynchophora (Ficure 5a) have highly specialized leaves, for most of the leaf is an inflated water- sac with a special valve opening. Numerous insects are caught in these traps. The superficial unistratose, discoid gemmae develop from cells of the leaf (FicurE 5a, G). The details of development of similar gemmae in other genera have been described by Evans (1911, 1912). There are usually three specialized cells on the margin which serve as ‘“Haftorgane” or organs of attachment (Ficure 5b, A). Such cells project at right angles to the surfaces of the gemma and the slime secreted from the tips serves to attach the gemma to the leaf. The new leafy plant de- velops through the activities of an apical cell which develops from a cell of the margin. The gemmae of Diplasiolejeunea (Ficure 6a, b) are of the same general form. In the genus Cyclolejeunea, perhaps the most widely occurring of the gemmiparous leaf-inhabiting taxa, gemmae develop from the marginal cells of the leaf (FicurE 7a). The large discoid gemma of C. convexistipa (Ficure 8), is unistratose and watch-glass shaped, and according to Evans (1904), apparently falls on the leaf with the concave side down and the margin appressed to the leaf surface, thus creating a water-hold- ing space in the middle. Three or four long rhizoids and numerous short branched ones develop from the marginal cells. The latter are slime se- creting and serve to anchor the gemma to the leaf (Ficure 8, R). The 1970] FULFORD ET AL., ELFIN FOREST, 11 61 leafy stem develops from an apical cell on the margin. In C. accedens the gemma has only two long unbranched rhizoids (FicurRE 7b, R), bent over the lower surface, and an abundance of short, branched, slime-se- creting marginal ones. The apical cell of the gemma gives rise to an oblong protonema or gemmothallus (FicuRE 7c, T), and the leafy plant develops from the margin of this latter thallus. Still another highly specialized means of vegetative reproduction among the leaf-inhabiting species occurs in the very common Drepanolejeunea inchoata. The plants are relatively large, with broad, coarsely toothed leaves, and in addition to the branches with adult leaves, there are some with smaller leaves of the same sort, and some with very slender lanceolate leaves. These latter branches produce even smaller easily detached branches, the propagula. The propagulum (FicurRE 9) is a minute, slender branch with several leaves and underleaves. The first underleaf (FicurE 9, U) is the largest and has, over much of its surface, a circular “suction disc” with slime secreting rhizoids from the marginal cells. This disc serves as an anchoring mechanism for the branchlet. Such propagula have been described by Evans (1902) for several Puerto Rican genera. In a study of epiphyllous hepatics of El Salvador, Winkler (1967) has also described the leaf-adaptions of many of these species. Small-leaved microphyllous or flagelliform branches are developed in many species. In Ceratolejeunea flagelliformis (FicurE 10 a vs 10 b) and Prionolejeunea aemula (FicurE 13 a vs 13 b), the whole branch is readily detached and blown about, while in Bazzania longistipula (Ficure 11 a vs 11 b) and Calypogeia cellulosa (Ficure 12 a vs 12 b), the small-leaved branches are brittle and tend to break into short lengths. Both types form new plants and colonies in favorable surroundings. Large numbers of fragments of leaves and stems are also to be found in any collection. In most species it has been demonstrated that the cells are totipotent and that new plants are produced in considerable numbers from such fragments, especially in such genera as Bazzania, Plagiochila, Radula and many genera of the Lejeuneaceae. All of the above described methods of reproduction occur in abun- dance in the area of Pico del Oeste. This is indicated from the occur- rence on the pollen-trap slides and the numbers of sporelings, gemmalings, specialized branches, and stem and leaf fragments found in our collec- tions. They are carried by the wind, strong up- and down-drafts, and by driving rains. But these are examples of short-distance dispersals and it must be kept in mind that even on the four peaks of the Luquillo Mountains there is a considerable difference in the hepatic flora. How much of this difference is due to a matter of dispersal and how much to habitat diffierences is an open question, since the hepatics require only microhabitats within any area. There is also a considerable difference in composition between the hepatic flora of Pico del Oeste and that at lower elevations, namely the Rain Forest, at 500 to 625 meters, or the several collecting localities at 700 to 750 meters. 62 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 7a Ficures 4-9. Some leaf-inhabiting species. Fic. 4. Colura clavigera, show- ing the strongly inflated leaves (L), X 85. Fic. 5 a. Cl. rhynchophora, an in- flated leaf with discoid gemmae (G), on the long-cylindrical tip, * 85. Fic. 5 b. A gemma with three attaching organs (A), X 240. Fic. 6 a. Diplasiole- jeunea unidentata, leaf with discoid gemmae (G), X 60. Fic. 6 b. A gemma with three attaching organs, X 240. Fic. 7 a. Cyclolejeunea accedens, diagram a portion of a leaf margin with discoid gemmae, 60. Fic. 7 b. Gemma with two large attachment cells at its base, and a pair of long rhizoids (R) on the lower side. Fic. 7 c. Gemmaling, with the gemma below and the gemmothallus T) with a leafy plant above. Fic. 8. C. convexistipa, gemma with short, branched, marginal rhizoids (R), and a young leafy plant. Fic. 9. Drepanole- jeunea inchoata, a tiny propagulum with three leaves and two underleaves (U); e first underleaf bears a large “suction disc” with marginal rhizoids (R), which serve to attach the propagulum to a leaf, X 240. Ficures 7 b, c, and 8 after Evans (1904). 1970] FULFORD ET AL., ELFIN FOREST, 11 63 There is evidence that certain of the species from these areas may have been repeatedly blown into the area of Pico del Oeste, but apparently they have not become established. In any patch of hepatics here, there are one to several dominants and as many as 18 additional species, some of which are tiny fragments which have been caught in the tangled growth. No one of the collections con- tained only a single species. The fragments of stems with growing tips had two or three to a dozen leaves, often smaller or conspicuously re- duced in size. The number of such stems which actually survive and grow into typical plants is not known. In all, 95 collections of leafy Hepaticae were made on Pico del Oeste. Mr. Stotler examined all of the packets in great detail and made a per- manent slide of each species in each packet, more than 650 slides in all. These include 15 families, 34 genera, and 62 species. We wish to thank Drs. Carroll E. Wood and Richard A. Howard for collecting material (38 packets), and Dr. Howard for his hospitality, and for guiding us through the area. I also wish to thank the National Science Foundation for some financial aid. The specimens are housed at the University of Cincinnati. In the following list an asterisk (*) denotes species reported from Puerto Rico for the first time. The numbers following each species in- dicate the number of times that species occurred in the total number of packets, In the distribution data “known” indicates that the data is taken from a monographic treatment of the genus, and “reported” indi- cates that the records are from the older literature and from lists. TRICHOCOLEACEAE Trichocolea flaccida (Spr.) Jack & Steph. 2/95. With other hepatics on a tree trunk and on a leaf. Known from through- out the West Indies; Mexico, Guatemala; Colombia, Venezuela, Ecuador, Peru, Bolivia. IsOTACHACEAE a aa ay aug (Steph.) Grolle. 1/95. Soil bank. Known from Puerto Rico, Guadeloupe, Martinique. HERBERTACEAE Herberta divergens (Steph.) Herzog. 20/95 In tufts or pendent on trees and shrubs, and on leaves, including Ocotea, Eugenia, oS and Calyptranthes, and on leaves of Anthurium. Known from Puerto Rico; Costa Rica; Brazil, Bolivia. Herberta saneilis (T. Tayl.) Spr. 7/95. In tufts or pendent on trees, shrubs, and scala including Calyptranthes and Grammadenia. Known from Cuba, Puerto Rico, Dominica; Costa Rica; Colombia, British Guiana, Brazil, Ecuador, Peru, Bolivia. LEPIDOZIACEAE Bazzania gh hanes (Lindenb. & Gott.) Trevis. 1/95. A large “ball” on a tree trunk. Widespread in the West Indies, Central America, and the gene ie half of South America. 64 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Figures 10-13. Some species with flagelliform or microphyllous branches. Fic. 10 a. Ceratolejeunea flagelliformis, adult stem, X 60. Fic. 10 b. Flagelli- form branch of this species, X 60. Fic. 11 a. Bazzania longistipula, adult leaf, xX 60. Fic. 11 b. Portion of a microphyllous branch of this species, < 60. Fic. 12 a. Calypogeia cellulosa, an adult leaf (L) and an underleaf (U), x 60. Fic. 12 b. Portion of a small-leaved branch of this species, X 60, Fic. 13 a. Prionolejeunea aemula, portion of an adult stem, X 60. Fic. 13 b. A short microphyllous branch of this species, X 60. Drawn from plants collected on Pico del Oeste. 1970] FULFORD ET AL., ELFIN FOREST, 11 65 Bazzania longa (Nees) Trevis. 27/95. Abundant on tree trunks, twigs, and leaves, including Tabebuia and Wal- lenia, and on Anthurium and ferns. Known from Cuba, Jamaica, Puerto Rico, St. Kitts, Guadeloupe, Dominica, Martinique, and Trinidad. (Frag- ments of stems are common in many collections.) Bazzania longistipula (Lindenb.) Trevis. 14/95. In mats or deep tufts on tree trunks and branches, including Eugenia, Ilex, and Miconia. Known from throughout the West Indies, Central America, and tropical South America. (Fragments of stems are very common in collections in the mountains). Bazzania roraimensis (Steph.) Fulf. 5/95. epatics on trees. Known from Jamaica, Puerto Rico; Co lombia, British Guiana, and Brazil. (All of the collections were depauperate fragments. Bazzania aoe (Sw.) Trevis. 1/95. orming a “ball” on a tree trunk. Common throughout tropical America. presence ts carinatum (Grev.) Reim. 5/95. With other hepatics on tree trunks and twigs. Known from Puerto Rico, Guadeloupe, Dominica, Martinique; Venezuela. dig patens. Lindenb. 16/95. In deep tufts on trunks and branches of trees and shrubs, including Eu- genia, re Tabebuia, and Wallenia, Known from Cuba, Jamaica, Haiti, Puerto Rico, St. Kitts, Guadeloupe, Dominica, Martinique; Guatemala; Colombia, Venezu ela. Telaranea sejuncta (Angstr.) S. Arnell. 1/95. Decayed log. Known from oan eastern United States; Puerto Rico, Guadeloupe, Dominica; Mexico, Guatemala, Costa Rica, Honduras; Co- lombia, Venezuela, Brazil, ake Peru, Bolivia CALYPOGEIACEAE Calypogeia cellulosa (Spreng.) Steph. 11/95. oe or among other hepatics on tree trunks and branches, leaves and over f from Cuba, Puerto Rico, Guadeloupe, and Dominica. Capes elliottit Steph. 4/95. In mats on tree branches, tree base, and old leaves. Known from Puerto Rico, Dominica. Calypogeia kate Nees & Mont. 1/95. On nthurium, Widespread and abundant throughout tropical Nea (tt most commonly grows on moist banks and rocks, logs, a and tree bark. CEPHALOZIACEAE Cephalozia caribbeania Fulf. 1/95. On bark, tree base. Known from Cuba, Jamaica, Puerto Rico, Guadeloupe, Dominica, Trinidad; Mexico, Guatemala, Honduras; Venezuela. *Nowellia caribbeania Fulf. 1/ On mound on ee of she Known from Puerto Rico; Guatemala, Hon- duras, Costa Rica; Vene Nowellia pasieteae Asean ” 3/98. (N. bicornis (Spr.) Fulf.) Among tufts of other hepatics. Known from Puerto Rico, Guadeloupe, Dadian 66 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ODONTOSCHISMACEA Alobiellopsis Eee (Spr.) Fulf. 1/95 n compact soil along path. Known from Puerto Rico, Dominica, Mar- tinique, Trinidad. Odontoschisma prostratum (Sw.) Trevis. 1/95. Near the base of a tree trunk. Known from eastern United States; Cuba, Jamaica, Puerto Rico, Guadeloupe, Martinique. ZOOPSIDACEA Zoopsis ‘aes Steph. 1/95. Among other hepatics on a tree trunk. Known from Cuba, Puerto Rico, Dominica, Trinidad. LOPHOCOLEACEA Pike elliottii (Steph.) Pagan 1/95. Loose mat on a tree branch. Reported from Puerto Rico, Dominica. Leptoscyphus sue (T. Tayl.) Mitt. 4/95. In mat tree bark, occasionally on the ground. Known from Puerto Rico, Guadeloupe, Martinique. LopHOZzIAc Syeysele perfalit (Sw.) Spruce. 5/95. In ark and leaves or hanging from tree branches. Known from Sad eG Rico; Costa Rica; Brazil, Peru. (Most of the stems have most leaves conspicuously bifid.) PLAGIOCHILACEAE Plagiochila bidens Gott. 20/95. On tree bark and living leaves including Calyptranthes, Grammadenia, Mi- cropholis, Ocotea, Trichilia. Reported from Cuba, Puerto Rico, Dominica, Guadeloupe; Bolivia. Plagiochila bursata (Desv.) Lindenb. 6/95. Reported common in the mountains of the American tropics. Plagiochila chinantlana Gott. On tree is and leaves. Reported from Puerto Rico; Mexico, Costa Rica; Colombia, Peru. pee p ensctint a T. Tayl. 6/95. In ra loose tufts on trees and over ferns. Reported from Cuba, Puerto Rico, Dominica; Mexico; Colombia, Venezuela *Plaiochia jamaicensis Lindenb. & Hpe. and branches. Reported from Jamaica, Puerto Rico. Plagiochile ad Lindenb. 1/95. On trees. Reported from Cuba, Puerto Rico, eee Dominica, Trini- dad; Costa Rica; Venezuela, ea Guiana, Brazi *Plagiochila subbidentata T. Tayl. 5. On bark and leaves of trees aoe sities Reported from Jamaica, Puerto Rico. **Adelanthus brevicaulis Steph. 13/95. much information is available as to the nature of the shoot/sporophyte re- bathonaite or the characteristics of the sporophyte in this species. I have tentatively placed it in this family. 1970] FULFORD ET AL., ELFIN FOREST, 11 67 On trunks, branches and leaves of trees and shrubs, including Grammadenia, Ocotea, Tabebuia, Wallenia. Reported from Puerto Rico, Guadeloupe; the Andes. SCAPANIACEAE Scapania portoricensis Hpe. & Gott. 3/95. On soil bank, and among other hepatics on tree bases and branches. Re- ported from Cuba, Jamaica, Puerto Rico; Colombia, Venezuela, Brazil, Ecuador. RADULACEAE Radula fendleri Gott. ex Steph. 5/95. and tufts on leaves and branches. Known from Puerto Rico, Guadeloupe, Martinique, Surinam, Venezuela. Radula saccatiloba Steph. 15/95. In tufts and mats on tree trunks, branches, and leaves including Calyp- tranthes, Ocotea, Tabebuia, and over ferns. Known from Cuba, Jamaica, Puerto Rico, Guadeloupe, Dominica, Martinique. FRULLANIACEAE *Frullania subtilissima (Nees & Mont.) Lindenb. 6/95. mats on twigs and living leaves, including Jlex and Micropholis. Re- ported from Dominican Republic, Puerto Rico, Guadeloupe, Dominica; French Guiana, Brazil. LEJEUNEACEAE Ceratolejeunea flagelliformis (Steph.) Fulf. 6/95. On bark of trees, shrubs, and on living leaves. Known from Puerto Rico, Martinique; Costa Rica. (Detached flagelliform branches are frequent.) Ceratolejeunea maritima (Spr.) Steph. 9/95. On bark of trees and shrubs, living leaves, and over ferns. Common through- out tropical America. Ceratolejeunea patentissima (Hpe. & Gott.) Evans 16/95. In mats on living leaves including Ardzsza, Calyptranthes, Ilex, Micro- pholis. (Most of the collections belong to var. acutifolia Fulf., with acute leaves.) Known from Cuba, Jamaica, Puerto Rico, Guadeloupe, Mar- tinique; Colombia, Venezuela. Ceratolejeunea valida Evans. 45/95. On bushes, twigs, and living leaves. Known only from Puerto Rico. *Colura clavigera (Gott.) Jovet-Ast. 1/95 With other hepatics, on bark. Known from Puerto Rico, Guadeloupe. Colura rhynchophora S. Jovet-Ast. 18/95. On leaves including, Guzmania and Vriesia, or as short stems among other. hepatics. Known from Puerto Rico, Guadeloupe. Cyclolejeunea accedens (Gott.) Evans. 17/95. : On living leaves, more rarely as fragments among other bryophytes. Knowm from Puerto Rico, St. Kitts, Guadeloupe, Dominica, St. Vincent; Venezue- la, Peru, Bolivia. Cyclolejeunea convexistipa (Lehm. & Lindenb. ) Evans 16/95 : ; On living leaves, rarely on bark, often fragments mixed with other hepatics. Both species reproduce by large, unistratose, discoid | gemmae which are of frequent occurrence on leaves. Widely distributed in tropical America. 68 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 icf eben ali (Lehm. & Lindenb.) Evans. 41/95. On n festoons, on banks among other hepatics. Known from sion Rico. St Kitts, Guadeloupe, Dominica, Martinique, St. Vincent; Colombia, French Guiana. Brazil. Diplasiolejeunea unidentata (Lehm. & Lindenb.) Schiffn. 11/95. and twigs, including Jlex, Miconia, Trichilia. Known from Jamaica, Puerto Rico, Guadeloupe, Dominica, Martinique, St. Vincent; El Salvador. The species produces abundant unistratose, discoid gemmae on the leaves. ng cg cena cor dopeei (Spr.) Steph. (D. pie Steph.) 15/95. On living leaves but more characteristically as small stems among 0 hepatics. Known pees Cuba, Jamaica, Puerto RG. St. Kitts, nto Venezuela, Brazil, Peru. Drepanolejeunea crucianella (T. Tayl.) oe 1/95. On leaves. Widespread in tropical America Drepanolejeunea evansii Bischl. (D. spinatus (Spr.) Evans) 1/95. On leave own from Puerto Rico, Guadeloupe, Martinique. gc aia nel ss (Meissn.) Steph. 29/95 ixed with other hepatics. Widespread throughout tropical America. This fede and probably others in this Sie produce very spe- cialized short branches for vegetative reproductio Drepanolejeunea fragilis Bischl. (D. araucariae en 17/95. aves and as fragments among other hepatics. Known from Jamaica, Puerto Rico, Guadeloupe; Guatemala, Costa Rica; Colombia, Surinam, Brazil. Harpalejeunea heterodonta Evans. 12/95 On leaves and as fragments among other hepatics. Known from Puerto Rico, Guadeloupe, Martinique; Colombia, British Guiana Harpalejeunea uncinata Steph. 13/95. On bark and leaves and as fragments among other hepatics. Known from Cuba, Dominican Republic, Puerto Rico, Guadeloupe, Martinique, Trini- dad; Panama. Neurolejeunea breutelii (Gott.) Evans 1/95 On a | nown throughout tropical America. Prionolejeunea aemula (Gott.) Evans. 19/95 On leaves and among other hepatics. Known from Puerto Rico, St. Kitts, Guadeloupe, Dominica; Colombia, Ecuador, Peru. Small branches with smaller leaves are readily detached and are frequent among other hepatics. Prionolejeunea aequitexta Evans. 1/95. On living leaves. Known from Puerto Rico. sir cies ese exauriculata Evans. 1/95. a living leaf. Known from Puerto Rico. aaa imnovata Evans. In small mats on twigs, with other hepatics. Known from Puerto Rico. Rectolejeunea phyllobola (Nees & Mont.) Evans. 1/95. On a lea Bos uthern Florida, widely distributed in the West Indies; Mexico, osta Ric iSeroeaieical involuta Steph. k and | 3/95. eaves. Reported from Puerto Rico, Dominica, St. Vincent; Brazil. 1970] FULFORD ET AL., ELFIN FOREST, 11 69 sg ab apt Spr. 4/95 pendent on trees and leaves, and over banks and logs. oe in gonial America. Trachylejeunea inflexa Steph. 21/95. n mats and tufts on trees and among other hepatics. Said to be common in the American tropics. LITERATURE CITED Evans, A. W. Hepaticae of Puerto Rico. I. The species of Leptolejeunea, in- cluding an account of their vegetative reproduction. Bull. Torrey Bot. Club 29: 496-510. Pls. 22-24. 1902. Hepaticae of Puerto Rico. II. Drepanolejeunea. Bull. Torrey Bot. Club 30: 19-41. Pls. 1-6. 1903. epaticae of Puerto Rico. IV. Odontolejeunea, Cyclolejeunea and Prionolejeunea. Bull. Torrey Bot. Club 31: 183-226. Pls. 8-12. 1904. ‘ aticae of Puerto Rico. X. Cololejeunea, Leptocolea and Aphano- lejeunea. Bull. Torrey Bot. Club 38: 251-286. 3 figs. Pls. 11, 12. 1911. . Hepaticae of Puerto Rico. XI. Ce ser hee Bull. Torrey Bot. Club 39: 209-225. 2 figs. Pls. 16, 17. Howarp, R. A. The ecology of an elfin Diet in ne Rico. 1. Introduction and composition studies. Jour. Arnold Arb, 49: 381-418. 14 figs. 1963. WrnEter, S. Die epiphyllen Moose der Nebelwalder von El Salvador C, A. Rev. Bryol. Lichénol. 35: 303-369. 20 figs. 2 tab. DEPARTMENT oF BroLocicaL ScIeENcEs [M. F.] UNIVERSITY OF CINCINNATI CINCINNATI, OHIO 45221 DEPARTMENT oF Botany [B. C. & R. S.] SOUTHERN ILLINOIS UNIVERSITY CARBONDALE, ILLINOIS 62901 70 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 TYPES OF BLIND VEIN-ENDINGS IN THE DICHOTOMOUS VENATION OF CIRCAEASTER ADRIANCE S. FOSTER A SALIENT FEATURE of the predominantly reticulate venation patterns of many dicotyledons as well as certain monocotyledons is the presence of blind vein-endings within the numerous areoles of the vascular system. Pray (1963) has treated in great detail the morphology and histogenesis of such vascular terminations in the leaves of Liriodendron, Aucuba, and Smilax. He concluded that “it seems probable that development of the minor venation of all angiosperms which have reticulate pattems similar to those investigated here involves similar ontogenetic processes.’ In the highly contrasted dichotomous venation patterns of Kingdonia and Circaeaster, however, the majority of the ultimate veinlets terminate in marginal teeth, vein anastomoses are infrequent and when areoles are formed, they are devoid of included vein-endings (Foster 1959, 1963, 1966, 1968; Foster & Arnott 1960). In describing this type of venation, I used the expression “blind vein-endings” to designate those veinlets which terminate in various positions and levels without reference to any of the distal marginal teeth. Do such blind vein-endings represent the vascular supply of “arrested” marginal teeth, or should they be considered as “sup- plementary veins” which are characteristic of “open” dichotomous venation in general? The present article describes and seeks to interpret morphologically the various types of blind vein-endings encountered in my extensive surveys of cultivated plants of Circaeaster agrestis Maxim. My original investiga- tion of herbarium material revealed very few examples of blind vein- endings, although it was admitted that their apparent rarity might have been the result of insufficient leaf sampling (Foster 1963, pp. 310, 311). In the leaves obtained from successive populations of Circaeaster raised in the growth chamber and in the Botanical Garden of the University of California during the past four years, however, blind vein-endings were relatively frequent. Not only were unbranched marginal and central end- ings found, comparable to some of those previously described, but a totally unexpected type of marginal, blind, dichotomously branched ending was discovered. This novel aspect of the venation of Circaeaster has been very carefully studied and constitutes a point of particular emphasis in the present paper. MATERIALS AND METHODS Ripe fruits, collected from the 1967 population of cultivated plants, were sown in large pots and 128 plants were raised from seedlings to maturity, 1970] FOSTER, VENATION IN CIRCAEASTER 71 in a growth chamber in which a photoperiod of 16 hours and a constant temperature of 50° F. were maintained. This temperature, which was about 10° higher than that employed in previous years, appeared to be a very significant factor in the unusually vigorous growth of the 1968 plants. At maturity, the plants were harvested and the leaf-rosettes preserved in 70% alcohol. A total of 6,179 leaves (including the larger floral bracts) were dissected from these specimens, examined for anastomoses and blind vein-endings, and the data recorded in tabular form. Leaves which were selected as the basis for the illustrations in this paper were cleared in NaOH followed by concentrated chloral hydrate, thoroughly dehydrated in the alcohol-series, and stained in a 1% solution of safranin in equal parts of absolute alcohol-xylene. Some of the leaves however were very difficult to clear satisfactorily in low concentrations of NaOH (i.e., 2.5%-— 5%) because of peculiar “amorphous” inclusions in many of the paren- chyma and epidermal cells. Prolonged treatment in cold 10% NaOH (for 3 weeks or longer) followed by chloral hydrate in most instances improved the appearance of such leaves prior to staining. In order to secure evenly stained preparations for subsequent photography, the leaf-specimens were left in the safranin for a relatively short period, approximately 144 to 2 hours rather than overnight as in the case of most of the leaves processed in previous years. This shorter period in the stain gave excellent prepara- tions, especially when the destaining was done in a mixture of 3 parts of xylene to 1 part of absolute alcohol. This solution prevented the too rapid leaching of safranin which always tends to occur when a mixture of equal parts of xylene-alcohol is used for destaining. I am grateful to Mrs. Irene Baker for her invaluable assistance in dealing with the germination and culture of the plants in the growth chamber. Thanks are also due Mr. Alfred A. Blaker, who made the photomicrographs, and Mrs. Emily E. Reid for preparing and arranging the line drawings. Iam also very appreciative of my wife’s assistance in proof-reading the manu- script. Symmetrical Patterns of Dichotomous Venation Comprehensive surveys of leaf morphology in herbarium collections and cultivated plants of Circaeaster have clearly shown the considerable fluctua- tion in (a) the shape of the lamina, (b) the number of teeth at the distal margin, and (c) the degree of symmetry of the venation patterns (Foster 1963, 1966, 1968). In many of the leaves obtained from herbarium ma- terial, the lamina is flabelliform or broadly ovate in contrast to the fre- quently narrower obovate or spatulate contour of the leaf-blade developed in the various populations of cultivated plants. Despite the variation in form and symmetry of the lamina, however, it is evident that usually all the vein-endings in a given leaf terminate in corresponding marginal teeth. This type of one-to-one correlation tends to predominate regardless of whether an even or an odd number of marginal teeth is present. It is, 72 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Fic. 1. Cleared leaf with symmetrical open dichotomous venation. Bifurcation of the petiolar bundle at base of lamina yields two “identical” types of vein- hee ig each composed of five veinlets terminating in corresponding margina teeth, X 8.6. 1970] FOSTER, VENATION IN CIRCAEASTER 73 however, particularly striking i in those cases where the associated pattern of dichotomous venation is symmetrical (Foster 1963, Pl. 1, fig. 5). An unusually clear example of ‘“‘isotomous” venation is shown i in FIGURE 1. In this leaf, the bifurcation of the petiolar bundle is followed by the dichotomy of each branch, resulting in two symmetrical pairs of veins at the base of the lamina. From each pair of basal veins, by appropriate dichotomy at successive and approximately opposite levels, two “identical” systems of veins are produced. Each system consists of a long unbranched marginal strand and two pairs of additional veins all of which enter cor- responding marginal teeth. The two halves or “sectors” of the lamina are thus vascularized by corresponding vein-systems which are separated from each other by a long, well defined “neutral space.” This term was used by Bock (1962, pp. 9 and 10, Figs. 15, 17) to designate the non-vascularized area between two vein-systems in leaves with symmetrical dichotomous venation. In this connection, the basic morphological resemblance between the venation pattern shown in Ficure 1 and that of the leaf of Ginkgo and a pinnule of Adiantum is indeed remarkable (see Bock 1962, Fig. 13a, b). Unbranched Blind Vein-endings Blind vein-endings in the leaves of Circaeaster may be conveniently classified into two main “types” viz.: branched and unbranched. The first type, to be described later, is relatively infrequent and has only been ob- served in a few leaves of cultivated plants (Fics. 16-22, 29). In contrast, unbranched blind endings are rather commonly seen and occur in a great many leaves of the plants which were raised in the growth chamber during 1968 (Fic. 2). It must be emphasized in the first place that the presence or absence of blind vein-endings is obviously not correlated with the size or shape of a leaf or its relative position in the leaf-rosette. Comparatively small primary leaves with only a few teeth may develop one or more blind endings (Fic. 3) and similar endings may occur in floral bracts (Fic. 4) and larger foliage leaves (Fics. 5-8). With reference to their general position in the venation pattern, un- branched vein-endings apparently occur most commonly in the lateral marginal region of the lamina. In some cases there is only a single blind ending in a leaf (Fics. 4, 6, 7, 8, 10) but a number of examples have been found where a blind ending is present in each margin of the lamina (Fics. 3, 5, 11). Considerable variation exists with respect to the level of diver- gence and the relative length of unbranched marginal blind endings. Often the ending is very short and diverges at a relatively high level in the lamina (Fic. 2, at right). But a good many examples have been found of marginal endings which diverge at the very base of the lamina. In these cases the marginal veins may terminate blindly after extending only a short distance along the margin (Fics. 3, 5). In other leaves, however, the blind ending may be more highly developed and extend nearly the full length of the lateral margin (Fics. 10, 11). Marginal blind endings, more or less inter- 74 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 mediate in length between these extremes also occur and are shown at the left of FicurEs 7 and 8. Less frequently, unbranched blind vein-endings occur in the upper cen- tral region or even in the distal margin of the lamina. In certain leaves which were studied, blind vein-endings are restricted to the central portion of the lamina (Fic. 13), while in others, both marginal and central blind endings occur in the same lamina (Fics. 2, 12, 14). One of the most re- markable examples I have seen of a distal blind vein-ending is represented in Figure 15. In this leaf, the short acroscopic branch of a very sym- metrical vein-dichotomy ends blindly, as shown by the arrow, between two adjacent teeth. It is also evident that central blind endings, as represented in Ficures 2, 13, 14, commonly terminate at varying distances below the narrow sinuses between adjacent marginal teeth. During the course of this investigation a very careful search was made to discover whether the development of unbranched blind vein-endings was ever correlated with the presence of corresponding vestigial marginal teeth. A priori it might be anticipated that since the sister-branch of a marginal or central blind ending enters a corresponding tooth, or by further dichot- omous branching vascularizes several successive teeth, the blind ending itself might represent the vascular supply of an undeveloped or rudimentary tooth. But only a single possible correlation of this sort was encountered (Fics. 6, 9). However, the peculiar massive cluster of tracheary elements found at the base of this vestigial tooth raises certain problems which will be more appropriately considered later in the discussion section of this paper. Marginal Blind Dichotoms As mentioned in the introduction of this paper, one of the most unex- pected results of my venation survey was the discovery of a dichotomously branched type of blind vein-ending in the lateral margin of the lamina of certain leaves (Fics. 16-22). In order to relate such ramified endings to the overall pattern of dichotomous venation in Circaeaster, I have adopted Bock’s (1962) term ‘“‘dichotom.” This term, which is brief and convenient for descriptive purposes is defined by Bock (1962, p. 6, Fig. 1) as follows: “As a general principle, dichotomy should be considered a growth pattern in which a base member develops two branch members, both advancing in a direction different from that of the base member. . . . The two angles formed by the base and the two branches may be equal, but also may vary. They are called the base angles, being always paired, while the angle formed by the two branches is the branch angle. Such a single dichotomy is called a dichotom.” In applying Bock’s general concept to the venation patterns shown in Ficures 16 to 22, the following interesting morphological differences be- tween the various forms of marginal blind dichotoms are quite evident, viz: 1. POINT OF ORIGIN OF THE BLIND DICHOTOM. In the leaves depicted in Ficures 19 and 22, the “‘base member” of the dichotom is a relatively long 1970| FOSTER, VENATION IN CIRCAEASTER 75 major lateral vein which bifurcates in the lower marginal region of the lamina. In contrast, the ‘base member” of the blind dichotom in the other five leaves represents the outer branch of a lateral vein-dichotomy which originates near the lamina base. In each of these cases, the sister-branch of the blind dichotom, by means of further dichotomous branching, forms two or three vein-endings which vascularize corresponding lamina teeth (Fics. 16, 17, 18, 20, 21). 2. LENGTH AND SYMMETRY OF DICHOTOM-BRANCHES. In one of the leaves, the two “branch members” of the dichotom are extremely sym- metrical and approximately equal in length (Fics. 18, 25). But in all the other examples, the inner (i.e. submarginal) “branch member” is more or less conspicuously longer than its sister-branch. This remarkable asym- metry is obvious in dichotoms with relatively short branches (Fics. 16, 19, 20, 24, 26) but is particularly striking when the inner branch terminates near the dentate margin of the lamina (Fics. 17, 21, 22, 23). 3. MObDE OF TERMINATION OF DICHOTOM-BRANCHES. As in the case of unbranched vein-endings, close attention was given to the possible associa- tion of blind dichotoms with corresponding rudimentary marginal teeth. Careful study of the venation under high magnification showed that in six of the leaves, the terminal tracheary elements of the dichotom-branches end freely in the mesophyll and that no histological evidence exists of vestigial teeth above such endings (Fics. 23, 24, 26). In one leaf, however, as shown in detail in Ficure 25, a peculiar “‘shelf-like” projection of the lamina margin lies directly above the outer branch of the blind dichotom. Whether this “shelf” should be regarded as a “rudimentary tooth” is ex- tremely problematical. When the projection was carefully examined under higher magnification than is represented in FicuRE 25, it was found com- pletely devoid of tracheary elements and in this respect quite unlike the apparently ‘authentic’ vestigial tooth shown in FicureE 9. DISCUSSION The varied types of blind vein-endings which have just been described, collectively represent a puzzling aspect of the dichotomous venation pattern of Circaeaster. Since usually all the vein-endings in a given leaf enter cor- responding marginal teeth, the blind endings which may additionally occur in various regions of the lamina pose interesting morphological and onto- genetic questions. From the standpoint of a morphological analysis of the mature venation, an unbranched blind ending in most instances appears to represent the shorter and usually less well-developed branch of a typical vein-dichotomy, In the simplest case the “dichotom” which yields a blind ending is very symmetrical in organization, one of its branches entering a marginal tooth, the other terminating at various levels in the lateral margin of the lamina (Fics. 2, 11, at right). More complex patterns result when the sister- branch of the blind ending forks once or twice, forming terminal veinlets which enter two or three adjacent teeth (e.g. Fics. 4, 7, 8, 10). Comparable 76 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 variations in venation pattern and in the degree of development of the blind ending also occur in the central or distal portions of the lamina (Fics. 2.12=25). It is tempting, in the case of unbranched blind vein-endings, to speculate that they may represent the persistence of the vascular supply of ‘“sup- pressed” marginal teeth. The present investigation, however, gives little real support to such an assumption. In the first place, the majority of the marginal blind endings are more or less strictly parallel with the edge of the lamina and terminate without reference to any protuberance which would qualify as the “rudiment” of a tooth (Fics. 2, 4, 5, 7, 8, 10, 11). Only one leaf was encountered in my entire survey in which a diminutive tooth is situated above the tip of a long marginal blind ending. (Fic. 6). However, since the longitudinal course of differentiation of xylem in the veins entering “normal” leaf teeth has not been determined, the significance of the isolated cluster of tracheary elements at the base of this vestigial tooth remains obscure (Fic. 9). Such an irregular and exaggerated development of the primary xylem does not occur, for example, at the point of entrance of a vein into a well developed tooth, nor is it matched in those puzzling cases where a tooth is vascularized by a basally unconnected strand (Fic. 19, right margin). Secondly, some of the vein-endings terminate very near the branch of a vein-dichotomy, suggesting that they may represent the incomplete devel- opment of Type V anastomoses rather than the vascular supply of “ar- rested” teeth (Fic. 11, at left margin; Fic. 12, upper center). A further example of this interesting condition has already been described in a recent paper on anastomoses in Circaeaster (Foster 1968, p. 61 and Fig. 15). Most of the objections which have just been raised against interpreting unbranched blind endings as the vascular supply of “arrested” teeth are also valid with respect to the morphological nature of marginal blind dichotoms (Fics. 16-22, 29). In the comparatively few leaves which had formed such ramified endings, there was no evidence of paired vestigial teeth and only one instance, as shown in Ficure 25, of a peculiar shelf-like extension of the margin above the dichotom. This ‘‘shelf,’’ however, lacks vasculature and does not seem to qualify as a rudimentary tooth; its morphological “‘significance,” if any, is highly problematical. From a comparative viewpoint it is important to note the occurrence of blind vein-endings in the dichotomous venation of other plants. Kingdonia is a good example for comparison because of the abundance and consistent development of blind endings in the dichotomously veined segments of its relatively large lamina (Foster 1959, Foster & Arnott 1960). Although the majority of the terminal veinlets enter corresponding marginal teeth, as in the smaller lamina of Circaeaster, 14.2% of all vein-endings studied in K ingdonia terminated blindly at various levels and positions in the lamina segments. Very commonly, the blind ending reaches the side or base of the sinus which separates adjacent marginal teeth (Foster & Arnott 1960, Fig. 14). But one of the most interesting points of similarity between Circaeaster and Kingdonia is the occurrence in both genera of marginal 1970 | FOSTER, VENATION IN CIRCAEASTER 77 blind dichotoms. FicurE 27 represents the enlargement of a small part of the margin of a lamina segment of Kingdonia. The branches of the con- spicuous blind dichotom shown in this figure are unequal and, as in Circaeaster, the inner branch is much longer than the outer (compare Fic. 27 with Fics. 22 and 23). It is also notable that each of the lateral veins of the small tepals of the flower of Kingdonia may dichotomize and form two unequal freely terminating branches very much like the branching pattern of some of the marginal dichotoms of Circaeaster (Foster 1961, p. 402, Figs. 4a, 5). When one seeks for additional examples of blind vein-endings in dichot- omous venation patterns, the genus Adiantum comes to mind as worthy of study. In A. capillus-veneris L. the great majority of the vein-endings ex- tend into corresponding marginal teeth, as in Circaeaster, and blind-endings are apparently rare in occurrence (Knobloch & Correll 1962, Plate 13, Fig. 1). My own extremely limited investigations on A. cooperi Baker, a species native to Costa Rica, revealed striking examples of marginal blind dicho- toms, very similar to those in the leaves of Circaeaster and Kingdonia (Fic. 28). This is clearly a remarkable example of parallel evolution, comparable to the similarity between the types of vein-anastomoses in Circaeaster and those in such unrelated taxa as Anemia, Zamia and Ginkgo (Foster 1968, pp. 63-65). In this connection, it is of interest to note that Arnott (1959, p. 408 and Fig. 9), in his intensive study of dichotomous venation in Ginkgo, found examples of “veins ending a considerable dis- tance from the margin.” Such veins seem entirely comparable to the central blind endings which may develop in Circaeaster (compare Arnott’s Fig. 9 with Fics. 2 and 13 of the present paper). In conclusion, the need for thorough ontogenetic studies on the time of origin and mode of differentiation of blind vein-endings in dichotomous vasculature should be strongly emphasized. It would certainly be of mor- phological interest, in the case of Circaeaster, for example, to know (1) how strictly the initiation and development of marginal teeth is coordinated with the pattern of differentiation of dichotomous venation and (2) the time of origin and the longitudinal course of development of the various types of blind vein-endings. Whether detailed information on these prob- lems would also shed light on the phylogenetic significance of dichotomous venation in Circaeaster remains, however, in my opinion, a completely open question. [Postscript to original manuscript | After the manuscript of the present paper had been completed, a re- markable type of marginal blind dichotom was found in which the outer, rather than the inner branch, was conspicuously longer than its sister branch (FicurE 29). Whether this type of asymmetrical vein dichotomy is “exceptional” or is as frequent in occurrence as the marginal dichotoms illustrated in Ficures 16-26 of this paper, remains to be determined by further surveys of leaf venation in Circaeaster. 78 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 — Ficure 29. Leaf with conspicuous asymmetrical blind dichotom which r earls ‘the bifurcation of the basal vein at the /eft of the lamina. Note that the basal vein at the right has formed three veinlets which terminate in correspond- ing cae ae the distal margin of the lamina. The scale at a side of the drawing equals LITERATURE CITED Arnott, H. J. 1959. Anastomoses in the venation of Ginkgo biloba. Am. Jour. t. 46: 405-411. Bock, W. 1962. Systematics of dichotomy and evolution. as Center Research ann Vol. 2. Geological Center, North Wales, Foster, A. S$. 1959. The morphological and taxonomic Seutecaaks of dichoto- mous Dalen in Kingdonia uniflora Balfour f. et W. W. Smith. Notes Bot. Gard. Edinb. 23: 1-12. . 1961. The floral cau Uta and relationships of Kingdonia uniflora. Jour. Arnold Arb. 42: 397-4 . 1963. The morphology a relationships of Circaeaster. Jour. Arnold Arb, 44: 299-32 1966. nik of anastomoses in the dichotomous venation of Circaeaster. Am. Jour. Bot. 53: 588-599. . 1968. Further morphological studies on anastomoses in the dichotomous venation of Circaeaster. Jour. Arnold Arb. 49: 52-67. . ARNOTT. 1960. Morphology and dichotomous vasculature of the leaf of Kingdonia uniflora. Am. Jour. Bot. 47: 684-698. Kwnosiocy, I. W., & D. S. CorrELit. 1962. Ferns and sic allies of Chihuahua, exico. Texas Research Foundation, Renner, Tex Pray, T. R. 1963. Origin of vein endings in angiosperm re Phytomorphology 0-81. DEPARTMENT OF BOTANY UNIVERSITY OF CALIFORNIA BERKELEY, CALIFORNIA 94720 1970| FOSTER, VENATION IN CIRCAEASTER 79 EXPLANATION OF PLATES PLATE I Fic. 2. Leaf with two ae vein- sup one in upper central region of lamina, the other in the margin at right. Note that the petiolar bundle reals far below the lamina-base, forming two eae vein-systems, X 1 PLATE II Fics. 3-8. Leaves with marginal blind vein-endings, Fic. 3. Primary lea with a blind vein- ending at each margin. Fic. 4. Floral bract with relatively long blind ending at right margin. Fic. . Leaf with eight symmetrically-vas- sane teeth ple a blind vein-ending at each margin. Note particularly the very low level of divergence of the blind db at right. Fic. 6. Leaf with long sie beet -ending and very rudimentary tooth (indicated by arrow) at left The structural details of the a of tracheary elements found at base ae this ‘tooth are shown in Fic. 9. Fic. 7. Leaf with eight symmetrically-aligned teeth and an unbranched vein-ending at left margin. Fic. 8. Leaf with two well- defined vein systems, the one at left with a long marginal blind ne Note also vein- oie aa at lower left of lamina. The scale at the side of each drawing equals 1 m PLATE III Highly magnified view of the cluster of tracheary elements at the base of the vestigial lateral tooth of the leaf shown in Fic. 6, * 250. PLATE IV Fics. 10-15. Leaves illustrating diversity in position and number of blind ndings. Fic. 10. Leaf with ten symmetrically vascularized teeth and a lo and two marginal blind vein- -endings. Fic. 12. A leaf with t endings, one in central region of lamina, the other at the right mar, IG i two unequal blind vein-endings, both in upper central portion of lamina. Fic Asymmetrical lamina with three blind endings, o t left n, the other two in upper central area. Fic. 15. Leaf with asymmetrical lamina showing two blind vein-endings, one very short and ru ain tary in upper central region, the | ther, indicated by arrow, terminating between two teeth at left. Note conspicuous Type I anastomosis in this leaf and ion Teaches’ xylem strand (represented as a pee isolated black line aed distal margin of lamina). The scale at the side of each drawing equals 1 PLATE V Fics. 16-22. Leaves oe marginal blind dichotoms. Fic. 16. oe at left margin, with nearly equal br anches. Details shown in Fic. 24. aT. ight : ~ ae t ind dichotom at left margin. See Fic. 25 for details. Fic. 19. Dichotom, at left margin, with unequal branches represented in sig detail in Fic. 26. Note that outermost tooth at right is vascularized b unc connected strand (US). each drawing a 1mm. 80 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 PLATE VI Marginal Her dichotoms. Fic. 23. Structural details of dichotom of lea shown in Fic. 17, X 49. Fic. 24. Magnified view of dichotom of leaf sales in Fig. 16, X 40. PLATE VII Marginal blind dichotoms. Fic. 25. Structural details of the very symmetrical marginal dichotom of the leaf shown in Fic. 18. Note that the dichotom termi- nates a short distance below a shelf-like projection of Noae mina-margin, X 54. Fic. 26. Greatly enlarged view of dichotom of leaf paced in Fig, 19, 67; PLATE VIII Fic. 27. Portion of marginal region of lamina segment of Kingdonia uniflora showing blind dichotom similar in type to that in leaf of Circaeaster, represented in Fic. 23, & 15.5. Fic. 28. Marginal region of pinnule of Adiantum cooperi, showing at lower left, a blind dichotom. Note remarkable similarity to blind dichotoms of Circaeaster shown in Fics. 17, 19, and 22, X 15.5. Jour. ARNOLD Ars. VOL, 351 PrLaTE I Foster, VENATION IN CIRCAEASTER Jour. ARNOLD Ars. VoL. 51 PiaTe II 2 | 5 a | | . 8 Foster, VENATION IN CIRCAEASTER Jour. ARNoLp Ars. VOL. 51 Pirate III Foster, VENATION IN CIRCAEASTER Jour. ARNOLD Arp. VOL. 51 PiaTE IV FosTER, VENATION IN CIRCAEASTER Jour. Arnotp Ars. VoL. 51 PLATE V 7 i I6 8 I 2\ ne | 20 | Foster, VENATION IN CIRCAEASTER Pirate VI Jour. ARNoLD Ars. VoL. 51 VENATION IN CIRCAEASTER ? FOSTER Jour. ARNOLD Ars. VOL, 51 PraTE VII FosTER, VENATION IN CIRCAEASTER Jour. Arnotp Ars. VoL. 51 PuaTeE VIII Foster, VENATION IN CIRCAEASTER 1970] DICKISON, DILLENIACEAE, V 89 COMPARATIVE MORPHOLOGICAL STUDIES IN DILLENIACEAE, V. LEAF ANATOMY* WILLIAM C. DICKISON N THE PRECEDING PAPER in this series (Dickison, 1969) I discussed the significance of nodal and leaf vascularization. The present article repre- sents an extension of that study surveying additional features of leaf anat- omy. It is hoped that the new information presented will help clarify the taxonomy, evolution, and generic definitions in this tropical family. All specimens studied have been cited with their respective collection data in my last paper. Though the lack of a complete collection of liquid-preserved specimens often prevented comparative ye wera several new anatomi- cal te are reported for the first time in the fam ntinued examination of leaf vascularization in = family has revealed a oe impressive range of variation, particularly in Hibbertia. In addi- tion to differences in vein diameter and presence or absence of sheathing sclerenchyma pointed out earlier, the pattern and orientation of the veins is often diagnostic. For example, H. glaberrima can be distinguished by the orientation of veins at right angles to the long axis of the leaf (FIGURE 46), whereas H. tomentosa is characterized by a closed venation pattern, i.e., very few free vein endings. A careful investigation of this variation using a wide range of material might prove to be of unusual taxonomic im- portance. I have already drawn attention to the characteristic leaf venation of many needle-leaved hibbertias, viz., a prominent midvein with recurved laterals. The massive nature, in some species, of recurved secondary veins, as they terminate at the leaf base is illustrated by Ficures 31 and 33. The extensive development of tracheary elements in this region of the leaf is not only of anatomical interest, but raises questions about possible physiological factors involved. FOLIAR IDIOBLASTS Foster (1956), in his review of plant idioblasts, recognizes three major categories of idioblastic cells — excretory, tracheoid, and sclerenchymatous. Examples of each class are found in leaves of the Dilleniaceae. Excretory idioblasts. A common component of this heterogeneous assemblage are crystalliferous idioblasts. The occurrence of enlarged, raphide-containing cells is frequently cited as a primary character dis- tinguishing the Dilleniaceae from other putatively related families (e.g., Cronquist, 1968). Since the distribution of raphides is limited in dicotyle- * Partial cost of publication me been covered by a grant from the Smith Fund, The University of North Carolin 90 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 dons, their presence assumes greater taxonomic importance than other more common crystal types (Metcalfe & Chalk, 1950). Gibbs (1958) is of the opinion that the presence of raphides is a primitive feature. Raphides have been observed in the leaves of all dilleniaceous genera except Schumacheria. Crystals were found with equal frequency in both the petiole and lamina; however, within the lamina, there is a tendency for raphides to be restricted to either the spongy or palisade mesophyll. Those found in the palisade layers are usually orientated in an upright manner (Ficure 27). The needle-like raphides are often produced in enlarged, thin-walled idioblasts containing an abundance of mucilage (FIcUREs 5, 21, 27). Calcium oxalate is deposited early in ontogeny as shown by the occurrence of crystals in the hypocotyl and cotyledons of Dillenia, Hib- bertia, and Tetracera (FiGURE 20). Crystals intermediate in shape between raphides and crystal sand are prevalent in some members of the family, e.g., Curatella. Examples of this condition may be seen in the enlarged pith parenchyma cells of the midrib in Doliocarpus olivaceus (FicuRE 22). The “crystal complement” of the family also includes cuboidal and nearly prismatic crystals as observed in the petiole of Doliocarpus guianensis (FIGURE 56) and Hibbertia pancheri. The latter types have not previously been reported. The distribution of crystalliferous cells in the phloem parenchyma of the petiole and midrib was noted by Metcalfe and Chalk (1950) in Hibbertia scandens (Ficure 23). This condition seems to be characteristic of the family as a whole, having now been observed in Curatella, Didesmandra, Dillenia, Doliocarpus, and Tetracera. The presence of silica containing cells in the Dilleniaceae has been known for some time. Workers have noted silicified cells in Curatella, Davilla, Doliocarpus, Hibbertia, and Tetracera. Silica deposition occurs throughout the parenchymatous regions of the leaf mesophyll and epidermis as either fine granular particles within the cell lumen (Ficure 54), or larger bodies impregnating the cell wall and intercellular cavities (FicurE 55). The abundance of these silica deposits, combined with crystals and sclerified trichomes, makes it possible to use the leaves of Curatella, Davilla, and Tetracera as sandpaper (Standley & Williams, 1961). Representing a third type of excretory idioblast of Dilleniaceae are the widely distributed “mucilaginous cells” (Ficure 57). It is of particular interest that Parmentier (1896) referred to the presence of “cellules oléigé- nes” in the mesophyll of Hibbertia scabra and suggested their occurrence was an excellent character to show the affinities of the Dilleniaceae to the Magnoliaceae. Although the chemical nature of the cell mucilage is un- known, these idioblasts do not appear comparable to the “ethereal oil cells” of many ranalian families. The large oil sac or drop, as well as scent, de- scribed by Wilson (1965) as being characteristic of ranalian oil cells is lacking. Structurally, the enlarged mucilage cells differ considerably in size and form. Intergradations with raphide-containing idioblasts are com- on. Tracheoid idioblasts. Cleared leaves of Hibbertia montana (E. H. 1970] DICKISON, DILLENIACEAE, V 91 Wilson 476), H. vestita (NSW 55998), and H. subvaginata (C. L. Wilson 764) revealed idioblasts which closely resemble the tracheoid type (FIGURE 47). The cells in question resemble typical tracheids in their spirally- thickened walls, general size and form, but are found completely isolated from other venation. They never occur in abundance. No evidence of un- differentiated procambium is present between these cells and the leaf vein. It is assumed, accordingly, that they arise in the lamina from the ground meristem. The best classification of these cells would probably be “storage tracheids” although there is no evidence to support their water-storing capacity. The occurrence of elements of this sort in Hibbertia is of particular interest. Emphasis has already been put on the highly diverse vein endings in many species of this genus and a varied foliar sclereid composition will be described subsequently. It is evident, therefore, that within this single taxon is present a wide diversification of idioblast morphology. Foster (1956) pointed out that the morphological relationship between the various foliar idioblasts represents a complex problem which merits further careful study. I would suggest that Hibbertia represents extremely favorable material for such an investigation. Sclerenchymatous idioblasts. Sclerenchymatous vein sheathing is present in the leaves of many Dilleniaceae (Dickison, 1969). This scler- enchyma, which often becomes massive (FicuRE 32), and encloses the terminal tracheids, is encountered in the petiole in the form of perivascular fibers (see Esau, 1965, p. 209) either capping vascular bundles or forming a continuous cylinder. In the larger veins of the lamina, sheathing extends from the adaxial to the abaxial surface. The first report of sclereids in leaves of Dilleniaceae was by Steppuhn (1895) who described the occurrence of thick-walled, branched “spicular” cells in two species of Hibbertia. Further comprehensive examination, however, has revealed a much wider range of foliar sclereid morphology and distribution. Although sclereids are wide-spread in the family, they reach greatest diversity in Hibbertia. On the basis of their distribution, it is convenient to distinguish four types: (1) sclereids diffuse in the mesophyll (and pith and cortex of petiole), (2) sclereids terminal at vein endings, (3) epidermal sclereids, and (4) sclereids subtending trichomes. With regard to the first category, various forms of sclereids are present: clusters of unbranched brachysclereids (‘stone cells”) are occasionally found in large numbers in the cortex and pith in petioles of Davilla, Doliocarpus (Ficure 56), Hibbertia and Schumacheria; Ficurr 9 shows an abundance of these sclereids in the leaf base of Hibbertia altigena; in addition to the elongate sheathing fibers, the midvein of several species may also be enclosed by short, unbranched sclereids. In one collection of Doliocarpus major (von Wedel 2860) such sclereids are located along the periphery of the lamina (FicurE 7). The conspicuous sclereids in Hib- bertia lucida (Ficurr 3), H. oubatchensis (FicuRE 2), H. pancheri, and H. patula (Ficure 1) all tend to be irregularly branched. These “armed,” thick-walled sclereids occur in clusters throughout the mesophyll and 92 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 parenchymatous regions of the petiole. Very distinctive osteosclereids are present in the lamina of H. salicifolia (Ficure 4). The latter cells, for the most part, are oriented with their long axis parallel to the long axis of the leaf. A fourth form of diffuse idioblast is the truly stellate sclereid (astrosclereids) found in the lamina of H. crenata (Ficure 13). In contrast to the above conditions, sclereids in Hibbertia lasiopus, H. miniata, H. montana, H. obtusifolia, and H. saligna are prevailingly as- sociated with terminal veinlets. In the leaves of H. lasiopus (C. L. Wilson 781), H. miniata (C. L. Wilson 782), H. montana (C. L. Wilson 838), and H. obtusifolia (Pullen 3882) the idioblasts are parenchymatous, mostly unbranched, or provided with only rudimentary branches, and occur in clusters at, or in the vicinity of, the terminal tracheids (FicurE 10). Along the margin of the lamina, however, are located extensively branched, stellate sclereids (FicurE 11). The terminal idioblasts of Hibbertia saligna (Constable 43107), on the other hand, are highly branched and occur singly at virtually all vein end- ings (FicuRE 12). In his paper on the terminal sclereids in Mouriria, Foster (1946) describes terminal cells intermediate in character between normal tracheary elements and sclereids. I have observed similar “transitional cells” in Hibbertia, again indicating the possible ontogenetic relationship between terminal sclereids and terminal tracheary elements. When the various terminal idioblastic cells described in my previous paper (Dickison, 1969) are considered in conjunction with the terminal sclereids now re- ported, the tremendous diversity of vein endings in this genus becomes evident. Next to be considered is epidermal sclerification, which, once again, is present in Hibbertia. Of particular note are the massive, thick-walled cells in the upper epidermis of H. nana. The striking nature of these isolated sclereids may be seen in surface view in Ficure 8. In sectioning, these cells are usually torn free leaving behind large spaces in the hypodermis (Ficure 26). In many of the needle-leaved hibbertias, the entire epider- mis becomes sclerified. Depending upon the species, the epidermal cells have wavy anticlinal walls (e.g., H. mucronata, Ficure 6), or straight walls with overlapping ends (e.g., H. acicularis, Figure 48). Sclerification of the epidermis in these species appears to be a specialization related to the xerophytic habit. Another unique feature observed only in Dillenia ochreata was the intensive sclereid development in the petiole wings (FicuRE 5). The last diagnostic sclereids in the leaves of Dilleniaceae are those which subtend the spiny, sclerified trichomes in Tetracera. This condition, which was first reported by Solereder (1908), has been observed only in the African species T. leiocarpa, the New World species T. ovalifolia and T. portobellensis, and an unnamed Krukoff collection from Brazil (Krukoff 6430). As seen in Ficures 14, 15, and 17, these columnar cells traverse the mesophyll immediately under the compound trichomes. Usually only a single sclereid is present under each trichome fascicle, but in the Krukoff collection a large cluster of sclereids is present. Although these sclereids 1970] DICKISON, DILLENIACEAE, V 93 are typically unbranched, they may become branched to varying degrees (FicuRE 16). The relationship between trichomes and sclereid formation in these plants is not clear. Their limited distribution, however, suggests they are of good systematic value. Moreover, the entire range of sclereid morphology in the family appears to be of potential taxonomic importance. TRICHOMES The scabrous nature of dilleniaceous leaves is well known to those familiar with the family. Solereder (1908) and Metcalfe and Chalk (1950) describe the trichome composition as consisting of simple unicellular, stellate, and rarely peltate hairs. The complete trichome complement of the family, however, has never been adequately illustrated. Simple unicellular trichomes are the most common form in the family, being present in all genera. Significant fluctuations are found in the length, form, and distribution of these hairs. Gradations in length occur from the narrow, very elongate (up to 1600) trichomes in species of Acrotrema (Ficure 41), Didesmandra and Dillenia, to the very short, pointed type seen in Doliocarpus (FicuRE 44). These hairs may be thick-walled, straight and highly sclerified (Ficures 13, 26), or, thin-walled and curled (FicurEs 19, 59). Distinctive short, thick-walled, curved hairs are present on the lower epidermis of Hibbertia baudouinii (FicuRE 28). Simple uni- cellular trichomes also may become silicified (e.g., Davilla). Stellate trichomes are present in Curatella (FicurE 45) and Hibbertia (Ficures 30, 34). Various forms of fasciculate trichomes with unicellular rays are common in the family. In Curatella (FicurE 45), Didesmandra (Ficure 43), and Tetracera (Ficurrs 14, 15) the rays are reduced to spines. Gradation between spiny fascicles and the stellate condition can be seen in Figures 17 and 58. The walls of these cells are sclerified and/or incrusted with silica. Although peltate hairs have been reported for Hibbertia lepidota, I have not observed true peltate trichomes during this study. I do not believe, furthermore, that the drawing by Solereder (doc. cit., p. 22) represents a truly peltate condition but rather an asteroid modification of the fasciculate type. A short, highly silicified, blunt-tipped, papillate trichome is prevalent in many species of Dillenia (Ficures 40, 60, 61). In surface view these trichomes are irregular in outline (FicuRE 61). In some species this may be the only type of trichome present. In species which possess both unicellular and papillate trichomes, their distribution seems to be of diag- nostic value (see Ficures 60, 61). Although the Dilleniaceae have long been thought to lack glandular trichomes, I am now reporting this feature for the first time in the family. Trichomes with typical glandular morphology, i.e., stalk and multicellular head, were found in Dillenia philippinensis and D. reifferscheidia. FIGURE 39 shows their location in depressions on the abaxial surface of the lamina. 94 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 EPIDERMIS AND STOMATA Following the early work of Parmentier (1896), the Dilleniaceae have been regarded by subsequent investigators (e.g., Solereder, 1908, and Metcalfe & Chalk, 1950) as possessing stomata of the ranunculaceous (anomocytic) and rubiaceous (paracytic) types. A lack of adequate liquid- preserved specimens has prevented critical observations on the epidermis of Acrotrema, Doliocarpus, and Schumacheria; nevertheless, my prelimin- ary studies reveal a wider range of structural variation than previously suspected. Although the systematic significance of this variation cannot be assessed at present, the type of stomatal apparatus, size of guard cells, size and undulation of ordinary epidermal cells, and presence of cuticular striations appear to be of potential taxonomic value. tomata are confined without exception to the abaxial leaf surface (Ficure 27). In many hibbertias they are located in stomatal crypts or grooves. Stomatal crypts are prevalent among many New Caledonian species, e.g., H. baudouinii (Ficure 28), H. nana (Ficure 26), H. pancheri (Ficure 25), H. pulchella, H. tontoutensia, H. trachyphylla, and H. wagapii. Stomatal grooves extending the length of the lamina are characteristic of most xerophytic, needle-leaved Australian species. Differ- ences occur in the shape of the groove and in the trichome distribution. Hibbertia pungens (Ficure 35) illustrates a condition where hairs com- pletely fill the groove. In H. uncinata (Ficure 37) trichomes are located only at the opening whereas in H. pachyrhiza (FicurE 36) the grooves lack trichomes. FiGuRE 36 also shows the thick cuticle formation in many of these species along with prominent striations. The leaves of H. gilgiana are distinguished by the anticlinal walls of both upper and lower epidermal cells being arched at the point of contact (Ficure 29). It can also be seen that the stomata are noticeably raised. A conspicuously papillate epidermis is also present in many species (e.g., H. baudouinii, Figure 28). Reference has previously been made to elongate, sclerified epidermal cells in xerophytic hibbertias (FicureE 48). The outer wall of these cells is frequently thickened and provided with papillae. Epidermal cells sur- rounding the spiny, fasciculate trichomes become differentiated by their thickened walls and lack of undulation. The epidermal cells of Doliocarpus olivaceus appear very distinctive in being circular and highly undulate. Un- fortunately I did not have good material for study of this species. Silici- fication of the epidermis has been noted in Curatella (FicurE 45), Davilla, Didesmandra, Dillenia, Hibbertia, and Tetracera. With regard to stomatal apparatus, Tetracera and Didesmandra possess the rubiaceous type. The two parallel subsidiary cells are distinguishable from the surrounding epidermal cells by their shape and dark contents (FIGURE 50). Guard cells vary in length from 20 to 354. The epidermis of Didesmandra is perhaps the most distinctive in the family. The anticlinal walls of epidermal cells are undulate, guard cells lack chloroplasts, and 1970] DICKISON, DILLENIACEAE, V 95 cuticular striations are orientated at right angles to the obliquely orienta- ted subsidiary cells (F1cuRE 49). The stomatal structure in Curatella, Davilla, Dillenia, and Hibbertia is predominantly ranunculaceous, since no subsidiary cells are evident (Ficure 53). Considerable variation does occur, however, in the undula- tion of epidermal cells and size of guard cells. Within the family guard cells range in length from 16 to 36u. Uniformly small guard cells (under 20) were found in Curatella, Davilla, Dillenia ovata, and Doliocarpus major. Of particular interest is the stomatal apparatus encountered in Hibbertia tetrandra. A careful examination showed it to be unlike all other Dillen- iaceae examined, as it could properly be classified as “gordoniaceous” (Ficure 51). The gordoniaceous stomatal apparatus (sensu Keng, 1962) is described as consisting of guard cells surrounded by two to four sub- sidiary cells which differ from ordinary epidermal cells in narrower cell shape and in their reaction to chemical reagents. In Hibbertia the num- ber of subsidiary cells varies from four to six. Keng (doc. cit.) tentatively named this stomatal apparatus for the theaceous genus Gordonia in which it was first described by Solereder (1908), who has subsequently con- sidered it as characterizing the subfamily Camellioideae of the Theaceae. In view of the apparent restricted distribution of the gordoniaceous type of stomatal apparatus, its presence in the Dilleniaceae seems to be of special significance in relating these two families. MESOPHYLL The mesophyll of dilleniaceous leaves displays considerable differenti- ation at both generic and specific levels. Each genus has a distinctive combination of characters. In addition to the specialized idioblasts al- ready mentioned, anatomical features of the mesophyll which demon- strate variation are: thickness, presence or absence of a hypodermis and its location on the adaxial or abaxial surface, bifacial or isolateral nature, number of palisade layers, and presence or absence of intercellular spaces in the spongy layers. A comparison of these features in all genera exclud- ing Hibbertia is listed in Taste 1. It should be noted that a hypodermis is found in some species of Dillenia. Also, the correlation between the coriaceous texture of the leaf, its multiple palisade layers, and a complex petiolar anatomy is noteworthy in Doliocarpus. The largest number of anatomical specializations, however, is found in the leaves of Hibbertia where extreme adaptations to environment have occurred. A few selected species listed in TaBLE 2 exemplify the range of diversification. DISCUSSION In response to extremes in the habitat, the leaves of Dilleniaceae exhibit marked differences in form, texture and thickness. Correlated with external 96 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 TABLE 1. Anatomical Comparison of Leaf Mesophyll in Dilleniaceae (excluding Hibbertia). Taxon NUMBER OF NUMBER OF LAYERS IN LAYERS IN. NUMBER OF CELLULAR AX PALISADE SPACES D. ABAXIAL HYPODERMIS HYPODERMIS LAYERS Acrotrema (adequate material not available for study) Curatella (Fic. 45) —- — , + isolateral sparse : 240, -FieG..759: ‘ coriacea (Humbert 5866), transverse section of lam showing abundance of curled, unicellular trichomes on lower surface, Fic. 60, Dillenia pete ena (Van Royen & Sleumer 5812), cleared leaf show- ing papillate trichomes restricted to vein : philippinensis (M ; ~ BiG. Ob k. : . Lagrimas s.n., Makiling, Philippines), cleared leaf showing irregular outline of papillate hairs and their general distribution over surface of lamina, < 160. Puate I Vv. a" Jour. Arnotp Ars. VOL. DICKISON, DILLENIACEAE, V Jour. ARNoLD Ars. VOL. 51 PuateE IT DICKISON, DILLENIACEAE, V Jour. ARNoLD ARB. VOL. 51 PLATE Ill DicKIson, DILLENIACEAE, V Jour. ARNOLD Ars. VOL. 51 Piate IV DICKISON, DILLENIACEAE, Vv PLATE V OLD ARB. VOL. Jour. ARN And ‘980 , = 9: a * * ok Ws: , Fe! Ce) By 1 eS g ioe 4 aX. = , See oh oat! ~. . is a “eet, ® 6 ee A me ha ® Be YY | , 19@ ea? + ay DIcKISON, DILLENIACEAE, V Jour. ARNOLD Ars. VOL. 51 PiaTE VI DICKISON, DILLENIACEAE, V Jour. ArRNoLp Ars. VOL. 51 Pirate VIT DICKISON, DILLENIACEAE, V Jour. ARNOLD Ars. VOL. 51 Pirate VIII DICKISON, DILLENIACEAE, V Jour. ARNOLD Ars. VOL. DICKISON, DILLENIACEAE, V Prare xX f DICKISON, DILLENIACEAE, V Jour. ARNOLD Ars. VoL. 51 PLate XI DIcKISON, DILLENIACEAE, V Jour. ARNOLD Ars. VOL. 51 PLATE XII |. pe ‘ SP : oe - 114 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 THE MORPHOLOGY AND GERMINATION OF THE SEED OF ELEPHANTORRHIZA ELEPHANTINA H. P. VAN DER SCHIJFF AND L. SNYMAN* Tue ceNus Elephantorrhiza Benth., described in 1842, belongs to the family Mimosaceae and consists, according to Phillips (1922, 1951), of only seven species with Elephantorrhiza elephantina (Burch.) S. Keels, the well known “elandsboontjie,” as the type species. All the representa- tives of the genus are small shrubs that vary in height from about one foot, as in F. elephantina, to small trees of ten feet and higher in E. burkei Benth. These plants are interesting in that they have thickened underground bases. It is a feature that generally occurs and is known in the Monocoty- ledoneae, but which is probably relatively rare in the Dicotyledoneae. However, this characteristic occurs in more South African Dicotyledoneae than is generally realized. Neither is it limited to a single or a few families, but it is found in a variety of families. The genera Talinum (Portulaca- ceae), Gyrocarpus (Hernandiaceae), Elephantorrhiza (Mimosaceae), Bau- hinia (Caesalpinaceae), Neorautanenia, Dolichos, Mucuna, and Erythrina (Papilionaceae), Rhoicissus and Cissus (Vitaceae), Adenia (Passiflora- ceae), Begonia (Begoniaceae), Cussonia (Araliaceae), Adenium and Pa- chypodium (Apocynaceae), Raphionacme and Ceropegia (Asclepiadaceae) , Ipomoea (Convolvulaceae), Pterodiscus, Harpagophytum, and Sesamo- thamnus (Pedaliaceae), and Coccinia (Curcurbitaceae) are particularly important in this respect. Knowledge of the morphological character and development of these thickened bases is to a large extent still only speculative, while little is known of the morphology and germination of the seed. The aim of this study is an attempt to gain a better insight into the morphology and germination of the seed and the establishment of the seedling, the latter being very interesting and unusual for the Dicotyledoneae. MATERIAL AND METHODS The plant which we studied, Elephantorrhiza elephantina, is a small shrub up to 2 feet high (Ficure 1) of which the underground parts are perennial while the stems above the ground partially or wholly die down annually during winter. The leaves are bipinnately compound and the inflorescence consists of * Most of this work was done while the senior author was a Mercer Research Fellow at the Arnold Arboretum, Harvard University, and was supported by a grant from the C.S.1.R., Republic of South Africa. 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 115 oa Ficure 1, Single iN a elephantina plant to show bipinnately com- pound leaves and infloresc small cream-colored flowers in a spike-like cluster that is up to 3 inches long. Although the ripe fruit is fairly hard, it readily absorbs water and disintegrates completely when wet, as a result of the softening of the peri- carp flaps that break away from the remaining seams. It has been observed in the veld, that a large percentage of the seeds germinate on the surface of the soil within these moist, disintegrating fruits. At this stage it is not yet clear whether this is the ideal condition for the germination of the seeds. The fleshy underground parts are very rich in tannic acid, a substance which is not only of medicinal value, but is also used in the tanning of hides. According to MacOwen (1897) an extract of the fleshy parts is used to treat sunburn, while Bryant (1909) maintains that the core of the “root” is used by Zulus as a cure for dysentery and diarrhoea. Thonner (1915) states that it is used as a fish poison and as a medicine, while it is known that an extract of the fleshy “roots” is an effective remedy for haemorrhoids. 116 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 FLAPS OF COTYLEDONS RADICLE COTYLEDONAL TUBE 4 DAYS Cen naa Fic 2. Drawings to illustrate seed a pags and the elongation of the ship (petiole) tube during germination 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 117 Steyn (1949) is not in agreement with Burtt-Davy that the plant is not toxic and is of the opinion that the seed contains an irritant toxin, which is harmful to animals. Elephantorrhiza elephantina occurs fairly generally in the Eastern Cape Province, the Orange Free State, Natal, Transvaal, South West Africa, dish seed disc cylinder paper FicurEs 3 AND 4. Fic. 3. ig pen used to investigate the mode of ger mination of the seeds. Fic. 4. Diagrammatic representation of the elongation of the cotyledon (petiole) tube and “taproot” during germination. 118 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Rhodesia, and Portuguese East Africa. In parts of the Transvaal highveld during spring it is often very common in veld which has been burned. The seeds used in this investigation were collected in the vicinity of Pretoria, where the plant occurs fairly frequently in grassveld. For the study of the morphology of the seed, dry seeds as well as seeds that had been soaked overnight were used. After drawings of the different seed forms were made (Ficure 2) the hard leathery testa was carefully removed in order to study and to make drawings of the embryo, which is very brittle when dry. Most of the material used for microtome sections was prepared accord- ing to standard methods and cast in wax. In some instances the material was fixed in acrolin, treated according to the method of Feder (unpubl.) and embedded in methacrylate resin. Although the latter method is more effective for the preparation of very thin sections, it is disadvantageous in that no serial sections can be made. The seeds used in the germination tests and for the study of the seedlings were first wounded with a small knife to make an opening in the testa for moisture to penetrate the seed. They were then soaked overnight, after which they were placed on wet filter paper in petri dishes and left to germinate at a room temperature of about 25° C. In cases where the development of the seedling was followed, the seed was germinated in pure vermiculite in glazed pots about 18 inches high. The young seedlings, of which specimens were removed daily to be studied, fixed, and to be used in the making of drawings, were easily washed out of the vermiculite by means of a slow stream of water. To trace the locality of elongation of the seedlings, germinated seeds were placed in holes made in a hardboard disc, which fitted into a glass cylinder impermeable to light. This was filled with moist crumpled filter paper and care was taken throughout to ensure that the roots did not come into contact with it (FicurRE 3). The pattern of elongation of the axis of the seedling was traced by means of Indian ink markings, 1 mm. apart on the developing seedlings (Miller & Wetmore, 1945). The first mark was made on the tip of the radicle immediately after its appearance at the micropyle. Thereafter, serial marks were made on the elongating seedling axis and schematic and numerical drawings were made on a piece of paper. Where serial marks moved farther apart, new marks were made in between to show the precise region of elongation (FIGURE 4). RESULTS Morphology of the seed. The seeds of Elephantorrhiza elephantina (Ficure 2) exhibit a wide variation in regard to size and form. How- ever, the majority of the seeds are spherically flattened, being slightly tapered in the region of the micropyle. Some of the seeds are also tapered on the side opposite the micropyle. If the seeds are densely packed to- 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 119 eA Sere, Os AS ce Ficures 5 and 6. Fic. 5. Longitudinal section through the base of embryo to show the position of the radicle and plumule. Fic. 6. Longitudinal section through base of seed during initial stages of germination to show the plumule in the still very short cotyledon (petiole) tube. 120 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 gether in the pods, they are flattened laterally. The seeds which are relatively large are up to 2 cm. broad and 2.5 cm. lon The embryo is surrounded by a hard leathery testa oa the two thick and fleshy cotyledons are very brittle and are easily broken in the dry condition. They lie close together to protect the radicle and plumule between them. The cotyledons are oval in outline and asymmetrical at the base. Each cotyledon has two flaps at its base that fold over one another and protect the relatively large radicle (FicurE 2). The cotyle- dons have short petioles which are fused to form a cotyledon tube or cylinder which surrounds the plumule and protects it (Ficures 5-10). In the fusion area between the cotyledon tube and hypocotyl there are indications of meristematic tissue (intercalary meristem) which, together with cell elongation, is responsible for the elongation of the cotyledon tube during germination (FiIcuRES 6-8). Germination of the seed. During the epigeal germination which usually occurs while the seed lies on the surface of the soil, the cotyledon flaps move apart and the radicle is forced out through the micropyle. According to the results obtained from the seedlings marked with Indian ink, it is clear that it is initially the cotyledon tube which elongates to carry the radicle out and that the apical growth of the radical only begins one or two days after germination has started (Ficurr 4). Root growth is fairly rapid and after one week tap roots of up to 9 inches in length were recorded. Longitudinal sections of the embryo confirmed without question that the cotyledon tube elongates not only to carry the radicle out, but also to implant the plumule into the ground (FicurEs 6-8). These sections also clearly show the meristematic zone as well as the cylindrical form of the cotyledon tube. In cross section it can be seen that the tube in the earlier stages of germination is flattened (FicurE 9) but later be- comes cylindrical as can be seen in Ficurre 10. Under favorable condi- tions this cotyledon tube may be up to 4 cm. long, which is probably of particular ecological importance as it apparently ensures that the seed- ling becomes established deeply enough. The plumule at first develops more slowly than the radicle. Between two to three weeks elapse from the time the radicle appears until the plumule becomes visible. It breaks through the cotyledon tube at the point of fusion between the cotyledon tube and the hypocotyl (FIGURE 11). At this stage the “root” has already undergone considerable sec- ondary thickening and has formed a conspicuous loose periderm which peels off in brown flakes (FicurE 13). The first sign of secondary thick- ening of the “root” (hypocotyl) is already visible three days after the radicle makes its appearance. No lateral roots were found on the upper and thickest part of the ‘root,’ while many lateral roots were present lower down (FiGurE 13). Each cotyledon stalk has three collateral vascular bundles (F1GuRE 11). It is interesting to note that these bundles in the petioles of the cotyledons undergo secondary thickening, and also form a periderm (FIGURE 12). 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA Wa FicurEs 7 and 8. Fic. 7. Longitudinal section through embryo to illustrate elongation of petiole tube. Fic. 8. Longitudinal section of petiole tube of about 2 cm. length. baz JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 DISCUSSION The type of germination encountered in Elephantorrhiza elephantina is not only very interesting, but also extraordinary for a representative f of the Dicotyledoneae. It is the characteristic type of germination o representatives of the Monocotyledoneae, of which species of the South oe neces .eeas +, :@t. 2@55 &, 0 SO. 86. aencesec.Seesss caeebe enntarels eseee 5.003500 e 52268. ware nO: 1h eee, & eS. s, SRLS oh 4% “8 238 a Pt Oe & Pe te gt oe s0 Sat see, anes ee ® AS i td ‘ame bt t-4rt 4¢ 4° ote, i 32 ee % Pa i Eikte ae » “SP er ee, oo Ficures 9 and 10. Fic. 9. Transverse section through petiole tube to illustrate the three vascular bundles of each petiole and the flattened petiole cavity. Fic. a 10. Transverse section through cylindrical petiole tube with young stem and leaf. 1970| VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 123 African genera Phoenix, Jubaea, and Hyphaene are probably the most elephantina is also interesting in that the cotyledons remain within testa, a feature which not only agrees with what is found in the Mono- cotyledoneae, but which is also characteristic of certain ppealiaine of certain Monocotyledoneae, the radicle and plumule are also carried out by the elongating cotyledon stalks. Ficure 11. Transverse section through petiole tube and young plumule at the stage when the plumule breaks through the tube. The same type of germination is also observed in Elephantorrhiza burkei, and according to Hofmeyr (1921-22) it is encountered in a repre- sentative of the Cucurbitaceae, Asa Gray (Hofmeyr, 1921-22), in his Structural Botany, maintains that it occurs in the American species Megar- rhiza californica of the Cucurbitaceae. [ VOL. JOURNAL OF THE ARNOLD ARBORETUM 124 Spores r” a : Sera “pened § ae ar Se Mae ce ae a fe aa: See a ME. oo ; ; Abi . Oa e = sent a Ot o ” a 4 pS Sine no eee - AE afin : Ax 5 CMe A ee . f young stem to illus- ar bundles. the petiolar collateral vascul section through petiole tube and ing of FIGURE 12. Transverse trate secondary thicken 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 125 Whether the germination of the seed and the development of the seed- ling of Elephantorrhiza elephantina is of any phylogenetic value is not clear. The phylogenetic importance of the germination morphology in the Polycarpicae can hardly be disputed, but here it appears as though it could be a case of convergence. Nevertheless, the facts that have been brought to light are most interesting and it can only be determined through further research how many of the South African plants exhibit this type of germination and whether it is of any phylogenetic significance. SUMMARY Elephantorrhiza elephantina the well known “elandsboontjie” of the family Mimosaceae is a small shrub of which the underground bases are thickened and fleshy. The germination of the seed of this species is interesting in that the radicle and plumule are carried out by the cotyledon petioles which elongate and which are fused to form a cotyledon tube. This cotyledon tube can be up to 4 cm. long and probably contributes to the successful establishment of the seedling. The elongation of the cotyledon tube is the result of cell elongation, and probably an intercalary meristem which is situated at the base of the cotyledons. The tap root develops very rapidly, while the main stem breaks through the cotyledon tube about two weeks after the radicle has made its appearance. Although this type of germination is characteristic of certain Mono- cotyledoneae and the cotyledon petiole of certain of the representatives of the Polycarpicae also elongate during germination, evidence Is as ye incomplete to evaluate the phylogenetic importance thereof. As far as our present knowledge goes it appears as though it may be a case of convergence. LITERATURE CITED Bryant, T. F. 1909. Zulu medicine and medicine men. Ann. Natl. Govt. Mu- seum 2. Fever, M. [Unpubl.] Plant histology: Techniques 1n use at Harvard. Hormeyr, J. 1921-22. Note on the germination of the seed of Elephantorrhiza burchellii Benth. S. Afr. Jour. Nat. Hist. 3: 215, 21 MacOwen, P. 1897. Elephantorrhiza burchellii Benth. Good Hope 10: 591. ee Miter, H. A., & R. H. WETMORE. 1945. Studies in the developmental anatomy of Phlox drummondii Hook. Am. Jour. Bot. 32: 628-634. ’ Puiiuips, E. P. 1922. Species of Elephantorrhiza in the South African herbaria. Bothalia 1: 187-193. ' : ; Puiiirs, E. P. 1951. The genera of South African flowering plants. p. 392. Pretoria. Government Printer. 6. Agric. Jour. Cape of 126 JOURNAL OF THE ARNOLD ARBORETUM [von. 51 ‘ in : : S it yaya 4 fe _* 3M Wits (i: pa 4 7 \ Wy J x A Ai wR, aN ney | fori, i K\\\ itil th at RYE uy. Saft SP LZ Ficure 13, a and b. Different stages in the germination of the seeds and a development of the seedlings to illustrate secondary thickening of the root” and lateral root development. 1970] VAN DER SCHIJFF & SNYMAN, ELEPHANTORRHIZA 127 Shy, | ra \) iy 128 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Srryn, D. G. 1949. Vergiftiging van mens en dier met gifplante, voedsel en drinkwater. Pretoria. Van Schaik. Tuonner, F. R. 1915. The flowering plants of Africa. p. 247. London. Dulau. WerrTsTEIN, R. 1962. Handbuch der systematischen botanik. p. 591. Amsterdam. Repr. Asher. DEPARTMENT OF GENERAL BOTANY UNIVERSITY OF PRETORIA SouTH AFRICA 1970] YAKOVLEV, NEW ORMOSIA FROM BRAZIL 129 A NEW SPECIES OF ORMOSIA FROM BRAZIL G. P. YAKOVLEV WHILE sTUDYING some unidentified material of South American Or- mosia in the herbarium of the Komarov Botanical Institute, two specimens of a collection made by Riedel and Luschnath interested me greatly. These proved to represent an undescribed species which, according to the detailed and thorough treatment of Dr. Velva E. Rudd, is most closely related to Ormosia friburgensis Taub. ex. Harms, but differs from that species in many characters, as cited below. It agrees with O. fri- burgensis in having large yellow seeds (Fic. 1) each with a brown spot, Fic. 1. Ormosia ruddiana, seed, X 1. a character which distinguishes both these species from the majority remaining which have red, blackish red, or black seeds. I am pleased to name this species for Velva E. Rudd, Curator, Depart- ment of Botany, Smithsonian Institution, who is an authority on the Leguminosae. Ormosia ruddiana Yakovlev, sp. nov. Fic. 2. Ormosia friburgensi affmis; arbor ad 6 m. alta; ramuli novelli fulvo- velutini; folia 7—13-foliolata, axi 12-20 cm. longo, laminis ellipticis vel oblongis, apice et basi acutis, marginibus leviter revolutis, subcoriaceis, supra subnitidis et glabris, 8-13 cm. longis, 2.5-3.5 cm. latis; venis se- cundariis 8-10, fere parallelis, conspicuis; flores ignotae, fructus inde- hiscens, sublignosus, leviter rugosus, fulvo- vel ferrugineo-velutinus, | 1- spermus, 4.5—6.5 cm. longus, 3-3.5 cm. latus, 2—-2.5 cm. crassus, semina subglobosa, flava, vel brunneola vel brunnea macula notata, 22-25 mm. longa et lata, ca. 20 mm. crassa; hilo 7-8 mm. longo, 2.5—3 mm. lato. Typus: Arbor 15-20 ped., mont. Iatocolum | Itacolomi], Riedel & Luschnath [Iter Brasiliense 1831-35] 649 (holotypus, isotypus LE). This locality is presumably Itacolomi, a mountain southwest of Ouro Préto in the state of Minas Gerais, Brazil. JOURNAL OF THE ARNOLD ARBORETUM & e 3 * 3 is ve ty VOL. 51 1970] YAKOVLEV, NEW ORMOSIA FROM BRAZIL 131 The two species, Ormosia friburgensis and O. ruddiana, differ in the characters listed here. O. FRIBURGENSIS O. RUDDIANA Leaf axis 2 cm. 2-20 cm. (including petiole) (excluding petiole) Leaflets 2-8 & 1-3. cm. 8~13 & 2.5-3.5 cm. Petiolules 1-2 mm. 3~4 mm. Length of legume 2.5—4.5 cm. 4.5-6.5 cm. Hilum 4-5 & 1-1.5 mm. 7~8 X 2.5-3 mm. DEPARTMENT OF BOTANY CHEMICAL-PHARMACEUTICAL INSTITUTE Pror. Popov Str. 14, LENINGRAD, P-22 = ’ ——, — os —— . Pea = an _ 7 oo e boa ites ; i aml eee i - Journal of the Arnold Arboretum Published quarterly in January, April, July, and October by the Arnold Arboretum, Harvard University. Subscription price $10.00 per year. Subscriptions and remittances should be sent to Miss Duicie A. Powell, old Arboretum, 22 Divinity Avenue, Cambridge, Massachusetts 02138, US.A. Volumes I-XLV, reprinted, are available from the Kraus Reprint Cor- poration, 16 East 46th Street, New York, New York 10017, U.S.A. EDITORIAL STAFF B. G. Schubert, Editor C. E. Wood, Jr. T. G. Hartley CIRCULATION D. A. Powell Printed at the Harvard University Printing Office, Boston, Massachusetts COVER: The Golden Larch, Pseudolarix Gordon WirH volume 51 the Journal of the Arnold Arboretum begins its second half century of continuous publication. To celebrate the occasion we are pleased to present a new cover format. The cover design is the work of Norman Comeau of the Art Department, Thomas Todd Company, Printers, Boston. We hope from time to time to have drawings of other plants on the cover. The Table of Contents will appear on the back cover. Eds. Se ee . Lye agra y . class postage 5 Boston, Massachusetts JOURNAL OF THE ARNOLD ARBORETUM VOL. ou Aprit 1970 NUMBER 2 A REVISION OF THE BORAGINACEAE OF WEST PAKISTAN AND KASHMIR S. M. A. Kazi THE TREATMENT OF THE BORAGINACEAE for Hooker’s Flora of British India (4: 137-179. 1883) was prepared by C. B. Clarke. The area then comprised the present states of India, Pakistan, Kashmir, Nepal, Burma, and Ceylon. Many other authors have since contributed to the flora of West Pakistan and Kashmir but most of their publications are of regional nature or only enumerations of plants collected by them from different parts of the country. Since C. B. Clarke’s work no comprehensive revision of Boraginaceae of West Pakistan and Kashmir has been prepared. The additional informa- tion we have on this area is scattered in the following general works or in the revisions of this family for some of the adjoining areas. In 1924 I. M. Johnston reviewed the Old World genera of the Borag- inaceae, subfamily Boraginoideae (Contr. Gray Herb. 73: 42-73. 1924). His observations on some genera and species of our area appeared in scat- tered papers (Contr. Gray Herb. and Jour. Arnold Arb.) published between 1924 and 1957. Brand, from 1921 to 1931 (in Engler, Pflanzenreich IV. 252 (Heft 78): 1-183. 1921; IV. 252 (Heft 97): 1-236. 1931) revised several groups of Boraginaceae, including many genera and species from West Pakistan and Kashmir. Later the family was revised by Popov (Fl. URSS 19: 97-691. 1953). His revision is confined to the plants of URSS, but gives information on those species with distribution extended to West Pakistan and Kashmir. Recently, H. Riedl (in Rechinger, Fl. Iranica 48: 1-266. 1967) dealt with the Boraginaceae of the Iranian Highlands, an area including the parts of West Pakistan westwards to the Indus; but the areas lying east- wards to the Indus and adjoining Kashmir and Jammu have been excluded as not being a part of the Iranian Highlands. His work is a valuable con- tribution to the study of Boraginaceae of West Pakistan as it gives the western limits of distribution of West Pakistan taxa and indicates their relationship with allied species of westerly distribution. In the present paper an attempt has been made to revise the Borag- 134 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 inaceae of West Pakistan, east and west of the Indus and the whole State of Jammu and Kashmir, with a special interest in determining the eastern limits of distribution of the taxa of this area and their eastern relatives. This revision is the result of a critical examination of the material ac- cumulated at the Herbaria of the Arnold Arboretum and Gray Herbarium of Harvard University, including most of the collections of Stewart, Koelz, and Webster from India, Pakistan, and Kashmir and many other collec- tions received from the Royal Botanic Gardens, Calcutta, the Forest Re- search Institute, Dehra Dun, and St. Xavier’s College, Bombay, which were the object of studies of the late I. M. Johnston for over thirty years. My own collections, especially those made with Professor K. H. Rechinger and Miss J. Lamond in West Pakistan in 1965, which are preserved at the North Regional Laboratories, Peshawar, and the material I received on loan from other herbaria. The following abbreviations’ (from Lanjouw & Stafleu, Index Herbariorum, ed. 5) are used in the citation of specimens. Arnold Arboretum (A) British Museum (Natural History) (pm) Royal Botanic Garden, Edinburgh (£) Pakistan Forest Institute, Medicinal Plant Branch, Peshawar (PFI-M) Conservatoire et Jardin Botaniques, Genéve (G) Gray Herbarium (cH) Islamia College, Peshawar (icp) Karachi University (KAU) Punjab University, Lahore (LAH University of Michigan, Ann Arbor (micH) North Regional Laboratories, Peshawar (PES) Peshawar University (PEU Gordon College, Rawalpindi (raw) Naturhistorisches Museum, Wien (w) U.S. National Museum, Washington, D.C. (us) Other abbreviations, used i in the citation of TYPE specimens, are also according to Lanjouw & Stafleu. I am greatly obliged to Dr. Richard A. Howard, Director of the Arnold Arboretum, for the facilities of work provided for me at the Harvard Uni- versity Herbaria under a Mercer Research Fellowship. I am also thankful for very courteous treatment, which I have received from the members of the staffs of the Arnold Arboretum and the Gray Herbarium. I am ex- tremely indebted to Mr. B. L. Burtt of the Royal Botanic Garden, Edin- burgh, and Dr. R. R. Stewart of the University of Michigan Herbarium for their encouragement, advice, and some very useful suggestions. I wish to express my thanks to Dr. Robert C. Foster for correcting my Latin diagno- sis, to Dr. Bernice G. Schubert for making editorial corrections and to the Directors and Curators of the herbaria, who were kind enough to send material on loan for my studies. ‘The symbols for the five herbaria in West Pakistan not included in Index Her- bariorum, ed. 5, were suggested by Dr. F. A. Stafleu in a personal communication. 1970 | KAZMI, BORAGINACEAE 135 In the systematic account which follows, I have cited all the material I have examined. I have tried to arrange the collections in geographical sequence, grouping them under West Pakistan and Kashmir and their respective districts arranged alphabetically. The collections which, due to lack of information, could not be placed in any district, have been listed under Miscellaneous. I have not changed the spelling of the localities or the language of the collector’s notes. In cases where the sheets were not numbered by collectors, I have given the herbarium number or any other indication, which might be useful in recognizing the sheet. BORAGINACEAE Juss. Gen. Pl. 128. 1789 “Borragineae” Syn.: Boraginaceae Lindl. Nat. Syst. ed. 2. 274. 1836; G. Don, Gen. Syst. 4: 306. 1838. Asperifoliae L. Ord. Nat. Pl. 489. 1792. Asperifoliaceae Reichenb. Con- spect. Regni Veg. 118. 1828. Herbs, shrubs or trees, usually scabrous, hispid hairy, sometimes glabrous. Leaves generally alternate, lowermost sometimes opposite, simple, usually entire, exstipulate. Inflorescence usually composed of 1 or more scorpioid or helicoid cymes that uncoil as the flowers open, or some- times glomerate-racemose or spicate, sometimes loosely cymose or with solitary flowers, bracteolate or not. Flowers mostly bisexual, actinomorphic or rarely zygomorphic, hypogynous. Calyx usually regular, sometimes irregular, sepals 5, distinct or basally connate, imbricate or rarely valvate. Corolla 5-lobed, imbricate or contorted in bud, rotate, salverform, funnel- form or campanulate; corolla tube with folds, sometimes terminated or partially closed by faucal appendages (scales). Stamens 5, epipetalous, equal or less commonly unequal, alternating with corolla lobes; anthers 2-celled, dehiscing longitudinally, basifixed or basally dorsifixed, introrse; annular nectariferous disc present or absent. Ovary superior, bilocular but becoming 4-locular at maturity; ovules anatropous or hemitropous; style terminal or gynobasic, long or short; stigma capitate or 2-lobed or some- times style twice bifid. Fruit drupaceous or dividing into two bilocular or 4 unilocular (or one 4-locular) nutlets; seeds erect or oblique, testa membranous, albumen fleshy, embryo curved, radical superior. A family of wide distribution, composed of about 100 genera and 2,000 species. About 60 genera are represented in the Old World including 36 in West Pakistan and Kashmir. KEY TO THE GENERA 1, Trees or shrubs. a CRG RE yO A ed hte SNe ek Dit Dea 1. Cordia. a a ae 6k i ele eae kas 3 2. Ehretia. 1. Herbs, annual, biennial or perennial, rarely undershrubs. 3. Style terminal on the entire ovary, not inserted between the pee Ey eS hk ba ee Rh eZ Coldenia. 4. Style 1 te OP ALANS hata ote te Oa A re 4. Peers 136 ; JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 3. Style arising from between the carpels. Ovules 2, always 2 nutlets developed. ................ 18. Rochelia. 5. Ovules 4, nutlets 4 at least when immature, rarely 1-3 abortive (but at younger stage always 6. Apices of anthers subulate and contorted. ...... 30. Trichodesma. 6. Apices of anthers obtuse or apiculate, not subulate or contorted ay aaeag nici ue the upper ones several times e low longer ee ee a re: 31. Caccinia. ip rane Sarah, all of equal size 8. Faucal appendages (scales) absent, corolla throat sometimes villous, corolla tube usually longitudinally plicate 9. Anther ped elongated, sterile; nat lobes re- 29. On VIE TOME. ci ss ceed Pew ce cues nosma. 9. Anther neta not elongated; corolla lobes distinct. 10. Gynobase elevated, pyramidal, more than half the length of the nutlets (corolla in W. Pakistan species strongly zygomorphic). .......... 7. Echiochilon. . Gynobase plane; corolla actinomorphic. 10 11. Corolla throat ie a filled with a style undivided, stigma 1. ........ 5. Sericostoma. 11. Corolla throat raked sometimes aii: style divided, sti 12. Corolla sata glabrous, never gibbous; ae 2-4-fid, rarely undivided; stigmas 28. Arn 42. Corl throat gibbous; style ~ fid, stigmas ee eee tere 27. Lithospermum. 8. Faucal ie (scales) present, oe represented only by hairs or minute glands (except in P. parviflora and P. echioides). 13. Appendages 10, in two tiers, _ w 5 alternating with the corolla lobes, the other 5 alternating with the stamens in the corolla tube. ............ 34. Decalipidanthus. . Appendages 5, in one tier, reed with the stamens at. or corolla lobes in the corolla 14. Nutlets neither seacata. appendicula nor prickly, smooth, granulate, verruculose, uberult or variously rugulose, glabrous or hirtulou 15. Calyx irregular, divided to the middle, den- tate between the lobes, much enlarged at fruit- ing, compressed, subbilabiate, gon dentate or DIAS 5 nk Bias a eo oe ak 9. Asperugo. Calyx regular, lobes eg not dentate between the lobes, not much enlarged at fruit- ing, neither compressed nor bilabiate, lobes not dentate or Faucal apperidages velvety or fimbriate areola of nutlet prominently oe out by a thickened girdle. ...... . Nonea. Faucal appendages usually none or apace never fimbriate, areola of nut- let various. — wn —" o> ad 1970] KAZMI, BORAGINACEAE 137 i. — ~I Areola of nutlet concave with dis- tinct smooth or slightly dentate but not thickened girdle 18. Areola of nutlet basal, if sub- ventral corolla via Lepper . Anchusa. Sonne of nutlet ental cor- a tube sie GLeCtt atu nb pees eee 4 Areola of riutlet te concave nor girdled. 19. Lower flowers distant, axillary, ai veloped. .. 6. Bothriospermum. inflo- rescence bracteate or ebracteate; corolla various; areola without rim or collar. 20. sina of nutlet affixed sub- ntrally Py the gynobase. ee ee . Lasiocaryum. 20. Areola 5s nutlet affixed ventrally, suprabasally or medially to the gynobase. 21. Corolla salverform, \o a — ° = Oo at wn 3 oO < oO t roe) al = = i) | me ally. eee ee aie a“ Anop Corolla tubular with NS pot suprabasally. tube, usually short- er; nutlets keeled. 8. Pseudomertensia. 138 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 22. malin sky-blue white, lobes at ne equalling the tube, usually long- M yosotis. 14. Nutlets at the margin and in “the middle area glochidiate-appendiculate, usually winged (in Lap- pula sometimes reduced to tubercles or prickles). 23. Gynobase flat; nutlets micas | tetrahedral, attached at one corner. .... 14. Trigonotis. Gynobase conical, pyramidal or le nut- lets not tetrahedral, variously attache 24. Nutlets with an elongate total lateral at- tachment to the gynobase, commonly de- current on s 25. Wings of millet. incurved, middle area partly covered. 21. Paracaryum. 25. Wings of nutlets not or rarely slight- aS incurved, middle area never cov- en dott See 22. Mattiastrum. i eae. variously attached to the gyno- base, not decurrent on style. 26. Nutlets with straight or uncinate, unicellular hairs, never tuberculate nor with multicellular glochidiate ap- pendages, rarely smooth and gla- brous (but if so always with stamens included). ....... 20. Omphalodes. 26. Nutlets with some or more multi- cellular glochidiate appendages or evidently tuberculate, rarely smooth and glabrous (but then with stamens always exserted). 27. Middle area of nutlet distinctly ac rey WwW to os ovate, apex acute, inner above areola keeled throughout Te le So a a Se ree 6. Lepechiniella. 27. Middle area of es if distinct, oblong-ovate, trapezifo 0 narrowly triangular, inner face above the areola not al age throughout its length. 8. Nutlets inseparable from the gynobase, bases of the marginal glochidiate appen- dages usually confluent. .. re 17. Heterocaryum. . Nutlets separable from the gynobase, bases of marginal nN co 1970] KAZMI, BORAGINACEAE 139 glochidiate — usu- ally not confluen 29. gy Linea. semiglobose or napiforn ments evidently longer than ca ae attached above faucal ap- uae, anthers and filaments exceeding the corolla. .............. site ae lo Gentle ahs PIO Ru a en Be Bh 3. Solenanthus. 30. ee equalling the anthers, not inserted above the faucal ap- s, anthers included or rarely slightly exceeding the corolla. al Mutlets with very few glochidiate appendages and with a well de- veloped wide cartilaginous margin. ............ 26. Actinocarya. Nutlets with many glochidiate appendages, marginless or with a weakly developed margin, thick, and evidently formed by the basal fusion of the lateral appendages. 32. Anthers at maximum 3 times longer than broad. .......... pass EOL Aae snes oh eeaWwea ky ae Coa med 25. Cynoglossum. 32. Anthers often more than 3 times longer than broad. ........ 24. Lindelofa. oi, _ 29. Nutlets more or less pyramidal to truncate-pyramidal-oblon 33. Nutlets about equalling the elongate gynobase, attached for nearly the whole length, (with rare exceptions) exceeded by style, margins of nut- lets toothed or lacerate, frequently glochidiate- Kamnenainr Qe, ee Oe Pr ee Ty er ee eee er Lappula. 33. Nutlets much surpassing the stout pyramidal paneer attached obliquely and supramedially by the deltoid or ovate areola, surpassing the style. 34. Nutlets Lilies or all of each fruit) with a thick medial, dorsal, an- HAE Chest ats 5 os th a cee eee ora ene ee 13. Microula. 34. Nutlets a a circumdorsal, commonly lacerate or Se margin. Fruiting calyx reflexed; rank tufted plants hgh her- Re pk us ee et kak nes Hackelia. 35. Fruiting calyx erect or ascending; low densely tees plants Wit Sitiall Hitt eaves. s..22 ei dasene geese: 10. Eritrichium. 1. Cordia L. Sp. Pl. 190. 1753; Gen. PI. ed. 5. 87. 1754 TYPE SPECIES: C. myxa L. Trees or shrubs. Leaves small to large, vege 4 evidently petiolate, style and stigmas much reduced or completely undeveloped); corolla campanulate to funnelform, small to large, white, yellow-orange or red, usually 5-merous but occasionally 4—8-merous, lobes ascending to recurved, tube short to long, cylindric or spreading; stamens exserted to included, filaments often hairy towards the base, usually well developed; style terminal on ovary, dichotomous, a simple column at base then dividing into 2 branches, which in turn fork to produce the 4 ultimate branches each bearing a stigma; stigmas 4, clavate to spathulate or capitate. Fruit a drupe with watery or glutinous mesocarp or rarely without fleshy 140 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 mesocarp and hence a nut; endocarp bony, 1-4-seeded. Seeds without endosperm. About 250 species, mainly of the American tropics. KEY TO THE SPECIES a. Leaves once and a half to twice as long as broad. b. Corolla tube longer than corolla lobes, lobes + 3 mm. long.........---: SIP Oy to! Dee Ce A ere eee ee ee 1. C. gharaf. b. Corolla tube shorter than corolla lobes, lobes 5-6 mm. long. ........... ekg oP Per er an rey Perret eee Tee 2. C. dichotoma. a. aor less than one and a half times as long as broad, frequently as broad c. Calyx not ribbed, glabrous externally. .................. 3. C. obliqua. c. Calyx ribbed, pubescent or tomentose externally. d. Leaves cordate or rounded at the base; corolla tube glabrous within. Fy RE yt eee ye ee ee eee ee, eee . C. macleodii. d. Leaves cuneate or rounded at the base; corolla tube hairy within. e. Pedicel tomentose; calyx + 10 mm. long. ......... 5. C. vestita. e. Pedicel subglabrous; calyx 3-4 mm. long. .......... 6. C. myxa. — . C. gharaf (Forssk.) Ehren. ex Asch. Sitz-ber. Ges. Naturf. Freunde 1879: 46. 1879; Verh. Bot. Ver. Brandenb. 21(2) : 69. 1880; Musch- ler, Man. Fl. Egypt 2: 781. 1912. Cornus gharaf Forssk. Fl. Aegypt.-Arab. xcv. 1775. Cornus sanguinea Forssk. 7bid. 33. 1775, non L. Cordia sinensis Lam. Encycl. Méth. Bot. i: 423. 1791; I. M. Johnston, Jour. Arnold Arb. 32: 11. 1951. Cordia rothii Roem. & Schult. Syst. Veg. 4: 798. 1819; DC. Prodr. 9: 480. 1845; Brandis, Forest Fl. India 338. 1874, Indian Trees 480. 1921; C. B. Clarke in Hook. f. Fl. Brit. India 4: 138. 1883; Cooke, Fl. Bombay Presi- dency 2: 202. 1908; yee Forest Fl. Punjab 360. 1924; Riedl in Rech- inger, Fl. Iranica 48: 6. 1967. Cordia angustifolia Roxb. Fl. Indica, ed. Carey & Wall. 2: 338. 1824, non Roem. & Schult. Cordia subopposita DC. Prodr. 9: 480. 1845. Type: Arabia: Hadie, Forsskdl, s.n. (as Cornus sanguinea Forssk. C.) Icon.: Brandis, Indian Trees fig. 172. 1921; Wight, Icon. Pl. Indiae Orient. 4: ¢.1379. 1848, under C. rothii Roem. & Schult. A large or small tree, twigs glabrous or nearly so. Leaves subopposite, 5-10 cm. long, 2—2.5 cm. broad, oblanceolate oblong, entire or undulate, apices usually rounded, rough and glabrous above when mature, more or less pubescent below; petioles 8-13 mm. long. Inflorescence short panicu- late-corymbose, subtomentose; peduncles 1.2—2.5 cm. long, puberulous to subtomentose. Flowers 5 mm. across, white, usually tetramerous, pedicel short, slender. Calyx + 5 mm. long, minutely pubescent externally, silky within, campanulate-tubular, lobes small, obtuse. Corolla tube equalling the calyx lobes, corolla lobes short, + 3 mm. long, obtuse, reflexed, fila- 1970] KAZMI, BORAGINACEAE 141 ments glabrous. Drupe usually 1-seeded, about 12 mm. long, ovoid, mucro- nate, striate, yellow or reddish brown when ripe DIsTRIBUTION: Pakistan, India, Ceylon, Arabia, and North Africa. West ane JHELUM Dist.: Langarpur, R. R. Stewart 544 (kK, RAW). Karacui Dist.: Jemadar ka Landa near Karachi Stocks 427 (x); Karachi, Jafri s.n. (KAU). LaworeE Dist.: Lahore, Thomson s.n. (K), R. R. Stewart 59. (K). Reported from Sind and Baluchistan. Kashmir: Reported from Jammu. Forsskal called the plant collected by him from Beit el Fakih and Hadie by its local name Cornus gharaf. This name has sometimes been rejected by many authors, most recently by Meikle (Israel Jour. Bot. 18: 141, 142. 1969), as a nomen nudum because no description was given on page xci, xclil, xcv, xcvii, and xcix of Forsskal Fl. Aegypt.-Arab. 1775, but on page xcix there is a clear reference to the description under Cornus sanguinea.” The epithet sanguinea was not available in the genus Cornus but could have been used by Ascherson for his new combination under Cordia in- stead of gharaf. However, since the name gharaf was selected by him, used later by many authors, and of course has no problems of priority, it should be accepted. Cordia gharaf is said (Parker l.c.) to be wild in Jaipur State on the Delhi Ridge and probably also in the hills of southeast Punjab, and planted elsewhere. 2. C. dichotoma Forster, Prodr. 18. 1786. Varrenia sinensis Lour. Pl. Cochinch. 138. 1790. Cordia indica Lam. Encycl. Méth. Bot. 1: 422. 1791. Cordia lourieri Roem. & Schult. Syst. Veg. 4: 466. 1819. C, suaveolens Blume, Bijdr. oe Wetens. 14: 843. 1826. C. brownii DC. Prodr. 9: 499. 1845. C. ixiocarpa F. Muell. Frag. cae he 1: 59. 1856. C. lowriana Brandis, Indian Trees 479. Type: “New Caledonia” without citation of collector’s name (BM). Trees 3-10 m. tall. Leaves elliptic to ovate or obovate, usually 8-12 cm. long, 4-8 cm. broad, apices obtuse, base rounded to acute, margins entire to rarely sinuate above the middle, upper surface sparingly puberulent, glabrescent or at times dotted, lower surface paler, puberulent, usually glabrescent; petioles slender, 2—4 cm. long. Inflorescence cymose, dichot- omous, bractless, loosely branched; peduncles 1-3 cm. long. Flowers of two sorts, male and hermaphrodite, produced, apparently, on separate trees. Male flowers shorter than the hermaphrodite, similar in form or at times with somewhat expanded corolla tube; filaments 3.5—4.7 mm. long, spar- *On xcix we find “Cornus gharaf. C. II. 10. In cephalagia.”” This refers to Centura I species no. 10 [on p. 33], which is Cornus sanguinea. Therefore, there o doube about what is ccsiet by Cornus gharaf and C. sanguinea. 142 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 ingly hairy below the middle; anthers 2—2.8 mm. long; ovary abortive, globose, glabrous, bearing a minute terminal papilla representing the undeveloped style. Hermaphrodite flowers with calyx 5—6 mm. long, ob- long in bud, not striate, campanulate, 4-6 mm. in diameter at summit, base rounded, sessile, lobes recurved, unequal, somewhat triangular; corolla 8-10 mm. long, lobes recurving, 5-6 mm. long, 2—2.5 mm. wide, tube 3-5 mm. long, usually shorter than the lobes, hairy only below the stamen- attachment; filaments 1-2 mm. long, sparingly hairy below the middle, styles with basal column 1—1.5 mm. long, the first branch about 1 mm. long, the ultimate stigmatiferous branches 3-6 mm. long, flattened, broad- ened, and not usually minutely lobulate above the middle. Drupe with viscous mesocarp, globose, 10-15 mm. in diameter, yellow or somewhat orange or reddish, supported by an accrescent indurate saucer-shaped calyx, 8-12 mm. in diameter. DIsTRIBUTION: Pakistan, Kashmir, India, southern China, Formosa, Hainan, Indochina, New Caledonia, and northeastern Australia. West Pakistan: Reported from Karachi, Sind, Lower Baluchistan, Salt Range, Multan, and Rawalpindi, both wild and cultivated. Kashmir: Reported from Mirpur, Billawar, and Jammu, both wild and culti- vated. The thinner more elongate, frequently acuminate leaves of Cordia dichotoma are usually once and a half to twice as long as broad, they are never subcordate at base as is frequently the case with the leaves of C. myxa and relatives. Cordia myxa, C. obliqua and their allies differ from C. dichotoma in their rounded thickish nonacuminate leaves, as well as in their coarser flowers, and very much larger drupes. Typically their leaves are less than once and a half as long as broad, frequently they are about as broad as long. 3. C. obliqua Willd. Phytogr. 1(4): t. 4, fig. 1. 1794. Willd., Linn. Sp. Pl. 1: 1072. 1797; DC. Prodr. 9: 479. 1845; C. B. Clarke in Hook. f. Fl. Brit. India 4: 137. 1883; Parker, Forest Fl. Punjab 359. 1924; Kashyap, Lahore Dist. Fl. 167. 1936. C. latifolia Roxb. Fl. Indica, ed. Carey & Wall. 2: 330. 1824; Brandis, Forest Fl. India 336. 1874, pro parte. Type: Habitat in Malabaria, ig sn, (B). Icon.: Willd. l.c. ¢. 4. fig. 1 A medium-sized deciduous tree; bark with numerous shallow longitud- inal fissures; branchlets glabrous or more or less brown-tomentose when young, Leaves 7-13 cm. long, 6-12 cm. broad, variable, orbicular, broadly ovate, elliptic or obovate, obtuse or more or less abruptly acuminate, base rounded, rarely cuneate, margins entire or slightly sinuate-dentate, gla- brous or nearly so above but rather harsh, without white discs, more or 1970] KAZMI, BORAGINACEAE 143 less pubescent, especially in the axils of the nerves beneath, basal nerves 3 rarely 5; petioles 17-35 mm. long. Inflorescence large, lax, of terminal or axillary pedunculate cymes; peduncles 2.5—5 mm. long. Flowers white, 5-10 mm. across, polygamous; pedicels short; buds pyriform. Calyx about 2.5 mm. long, glabrous externally or scarcely ie at the margins, pubes- cent within, splitting irregularly on the opening of the flower. Corolla tube as long as the calyx; lobes 5, as long as the tube, oblong-obtuse, re- curved. Filaments hairy at base. Drupe ovoid, 12~25 mm. long, apiculate, yellowish brown, pink or nearly black when ripe, shining, minutely rugose, supported by the saucer-shaped, faintly striate calyx; pulp nearly trans- parent, viscid, sweetish, edible. DIstRIBUTION: Pakistan, India, Ceylon (and North Africa ?). 3a. Var. obli Adult itis See beneath; calyx glabrous externally, scarcely villous on the margins. West Pakistan: Reported from Rawalpindi and Salt Range, often cultivated. 3b. Var. tomentosa (Wall.) Kazmi, comb. nov. C. tomentosa Wall. in Roxb. Fl. Indica, ed. Carey & Wall. 2: 339. 1824. C. wallichti G. Don, Gen. Syst. 4: 379. 1838; DC. Prodr. 9: 479. 1845; Cooke, Fl. Bombay Presidency 2: 200. 1908; Brandis, Indian Trees 479. 1921. c. ee var. wallichii (G. Don) C. B. Clarke in Hook. f. Fl. Brit. India 4: 137, 1383. Type: India in Dr. Heyne’s collection, under C. obliqua var. (K). Adult leaves densely stellate fulvous or white tomentose beneath; calyx glabrous externally. West Pakistan: Lahore Dist.: Lahore, Kazmi 2469 (PES). C. myxa sensu C. B. Clarke in Hook. f. Fl. Brit. India 4: 136. 1883, is a misidentification. 4. C. macleodii (Griff.) Hook. f. & Thoms. Jour. Linn. Soc. Bot. 2: 128. 1858; C. B. Clarke in Hook. f. Fl. Brit. India 4: 139. 1883; Brandis, Forest Fl. India 337. t. 41. 1874, Indian Trees 479. 1921; Cooke, FI. Bombay Presidency 2: 200. 1908; Parker, Forest Fl. Punjab 359. 1924. Hemigymnia macleodii Griff. Calcutta Jour. Nat. Hist. 3: 363. 1843. Type: Sylvae Jubbulpore vicinae, plerumque cum Tectona consociata, Herb. Griff. (kK). Icon.: Brandis, l.c. t. 41. 1874. A small tree with smooth gray bark, twigs clothed with dense white pubescence. Leaves alternate or sometimes almost subopposite, 7-18 cm. long, 5—15 cm. broad, orbicular or broadly ovate, obtuse, base cordate or 144 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 rounded, margins sinuate, firm and hard when mature, glabrous above with impressed nerves and raised discs, clothed beneath with short, dense, gray tomentum, basal nerves usually 3-5 or the nerves subbasal or sometimes the lowest pair of the secondary nerves arising at some distance from the base; petioles 2.5—5 cm. long, stout, tomentose. Inflorescence of terminal and axillary paniculate cymes, tomentose. Flowers white, 12 mm. across, subsessile, polygamous; buds obovoid. Calyx 6-8 mm. long, thick, ribbed, densely tomentose externally, glabrous within, lobes usually 6, as long or longer than the tube, spreading, spathulate-oblong, obtuse, veined. Fila- ments hairy at the base. Drupe 12-18 mm. long, ovoid, acute, supported by the cup-shaped, slightly ribbed calyx, not edible. DIstTRIBUTION: Pakistan and India. West Pakistan: SraLkoT Dist.: Sialkot Proper (seeds collected from Sialkot and grown at Dehra Dun), /shtiag Hussain s.n. (GH). Cultivated eastwards to Ra- walpindi. 5. C. vestita (DC.) Hook. f. & Thoms. Jour. Linn. Soc. Bot. 2: 128. 1858; Brandis, Forest Fl. India 338. 1874, Indian Trees 480. 1921; C. B. Clarke in Hook. f. Fl. Brit. India 4: 139. 1883; Parker, Forest Fl. Punjab 38. 1924. C. incana Royle, Illus. Bot. Himal. Mount. 1: 306. 1839, nomen nudum. Gynaion vestitum DC. Prodr. 9: 468. 1845. Type: In Himalaya occidentali tropica, Garhwal ad Pau, alt. 3,000— 4,000 ped. Edgeworth s.n. (G—vc). A small deciduous tree, bark greenish gray, exfoliating in large woody flakes, branchlets densely gray-tomentose when young. Leaves alternate, 7.5-15 cm. long, 6-13 cm. broad, sometimes larger, orbicular, broadly ovate or obovate, apices rounded or acuminate, base rounded or cuneate near the petioles, margins undulate, scabrous above with rather prominent white discs, tomentose beneath until mature, basal nerves 3-5, arising usually from a little above the base; petioles 2-4 cm. long, stout. Inflorescence of dense compound cymes, male in unilateral racemes; peduncles tomentose. Flowers yellowish white, 12 mm. across, polygamous; pedicels tomentose; buds pyriform. Calyx 10 mm. long, pubescent, more or less ribbed out- side, slightly hairy near the base inside, lobes 5. Corolla tube equalling the calyx, very hairy within, lobes 5, slightly shorter than the tube, spreading, obovate, crenulate, notched at the tips. Filaments hairy at base. Drupe 1.7 cm. long, ellipsoid, acute, supported by the accrescent calyx, which is campanulate, 12 mm. long, thick and strongly ribbed; pulp gelatinous, edible. DistTRIBUTION: Pakistan and India. West Pakistan: S1ALKoT Dist.: Mt. Tilla, Aitchison 45 (x). Salt Range and ted. Sub-Himalayan tracts, not common, often cultiva Kashmir: Reported from Mirpur and Jammu. 1970] KAZMI, BORAGINACEAE 145 6. C. myxa L. Sp. Pl. 190. 1753; DC. Prodr. 9: 479. 1845; Cooke, FI. Bombay Presidency 2: 200. 1908; Brandis, Forest Fl. India 366. 1874, excluding synonyms. Type: “Habitat in Aegypto, Malabaria” ngs oe 253.1 (LINN). Icon.: Wight, Illustr. Indian Bot. 2: t. 169. A moderate-sized deciduous tree reaching about 13 m., or a large shrub; bark dark, rough, fissured; branchlets usually glabrous. Leaves alternate, broadly ovate to suborbicular, obtuse, ca. 5 cm. long and 4 cm. broad, entire or with slightly uneven undulate margins, upper surface glabrous, the lower pubescent. Inflorescence paniculate or paniculate-corymbose, usually subglabrous. Calyx 3-4 mm. long, campanulate-tubular, usually pubescent, lobes short. Corolla tube hairy inside, lobes 5, short, 2.5-3 mm. long. Fruits up to 18 mm. long, subacute, ovate. DiIstRIBUTION: Tropical Asia, Pakistan, India, Iran, and Australia. West Pakistan: BALUCHISTAN: Shah Bilaweel, Stocks s.n. (kK). Reported from Nasirabad, Rindli and Drabbi in Baluchistan 2. Ehretia L. Syst. ed. 10. 936. 1759 Traxilum Raf. Sylva Telluriana 42. 1838. Type species: E. linifolia L. Trees or shrubs. Leaves small to large, usually distinctly petiolate, margins entire or serrate. Corolla white or yellowish, tube campanulate or elongate, lobes spreading or recurved; filaments elongate, anthers usually exserted; style terminal on the ovary, cleft above the middle, stigmas 2, small, capitate or elongate. Drupe subglobose, mostly yellow, orange or reddish, glabrous; endocarp at maturity breaking apart into either 2-seeded or 1-seeded pyrenes. Species 50, all tropical, principally in the Old World, except 3 species in America. KEY TO THE SPECIES ACB eaVeES ASE EET sol cities ae See ie Ae ae sacs Oe ek 1. E. serrata. a. Leaves entire. Leaves not seis 7.5 cm. in length, hairy — when mature; . Bini RI ts Se eg ek eh ee ete ahs obtusifolia. b. Leaves ee larger, aati up to 12 cm. in length, labrow beneath erties FGREEEOS RIAN AO 3S aS oe Se ee . E. laevis. 1. E. serrata Roxb. (Hort. Bengal. 17. 1814, nomen nudum) FI. Indica, ed. Carey & Wall. 2: 340. 1824, ed. Carey 1: 596. 1832; DC. Prodr. 9: 503. 1845, excluding vars.; Brandis, Forest Fl. India 339. 1874. E. acuminata R. Br. var. serrata (Roxb.) I. M. Johnston, Jour. Arnold Arb. 2 25-05, 146 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Type: “in Bengalo orientali, Silhet etc.” Roxburgh s.n. he Icon.: Lindl. Bot. Reg. 13: ¢. 1097. 1827; Wight, Ilustr. Indian Bot. 2: t. 170. 1841. A medium-sized deciduous tree with rough not deeply fissured gray bark, twigs usually brownish-pubescent when quite young. Leaves 8—15(-—20) cm. long, (3—)4-7(—10) cm. broad, usually lanceolate, sometimes elliptic- oblong, usually slenderly acuminate, margins finely and regularly serrate, the teeth ascending, usually with slender, prolonged, thickened, strict or incurved tips, base narrowed rarely rounded, midrib sparsely adpressed- hairy above, nerves slightly hairy in the axils beneath, otherwise glabrous; petioles 12-30 mm. long. Inflorescence of large terminal puberulous panicles with the lower branches axillary, 10-20 cm. long, 5-13 cm. broad. Flowers white, fragrant, sessile, 2.5 mm. across. Calyx sessile, 1.5—2 mm. long, lobes 0.8 mm. long, rounded, ciliate. Corolla 3-4 mm. long, nearly twice as long as the calyx, lobes 2-3 mm. long, about 1.5 mm. broad, blunt, spreading, tube about 1 mm. long. Filaments 2-3 mm. long. Style bifid, stigmas small, capitate. Drupe globose, nearly black when ripe, 3.5 mm. in diameter; pyrenes 2, each 2-celled, cells 1-seeded. DISTRIBUTION: Assam to Kashmir and West Pakistan. West Pakistan: Hazara Dist.: Balakot, Jmayat 21993 (k); Haripur, Kazmi 2474 (pes); Maksud near Haripur, 660 m., R. R. Stewart 12561 (GH, K, RAW); Abbottabad, Dickason s.n. (MIcH). RAWALPINDI Dist.: Rawalpindi-Murree Road, R. R. Stewart 6120 (a). Swat STATE: Saidu Sharif, Kazmi 2464 (PES); Margzar, R. R. Stewart 24306 (K, RAW), Kazmi s.n. (PEs). Kashmir: Pooncu Dist.: Nawal Nadi, R. R. Stewart s.n. (K, RAW). Miscel- laneous: Kashmir, Falconer Herb. East Ind. Co. s.n. (GH, K); Himal. Bor. Occid. Thomson, Hook. f. & Thoms. Herb. Ind. Orient. s.n. (GH, K). Reported from Jhelum Valley Road, Kishtwar, Udhampur and Jammu etc. 2. E. obtusifolia Hochst. ex DC. Prodr. 9: 507. 1845; C. B. Clarke in Hook. f. Fl. Brit. India 4: 142. 1883; Riedl in Rechinger, Fl. Iranica 48: 7. 1967. E. obovata R. Br. in Salt. Voyage 64. 1814, nomen nudum; DC. Prodr. 9 507. 1845, im synon, Type: Abyssinia Distr. Medschara, Schimper 2652 (s). A deciduous shrub, young twigs clothed with brown pubescence and gray spreading hairs. Leaves 3-8 cm. long, 2-4 cm. broad, sometimes longer, variable in shape, usually obovate, rounded or retuse at the apices; covered thinly on both surfaces with usually scattered, sometimes dense, adpressed gray hairs, persistently hairy beneath; entire; petioles 5-18 mm long. Inflorescence when young of close paniculate cymes, later lax and laterally corymbose, apparently terminal. Flowers pale bluish-white, 7-13 mm. across. Calyx 2.5 mm. long, externally hairy, lobes ovate-ob- long, acute or subacute, ciliate. Corolla 6-8 mm. long, tube exceeding the 1970] KAZMI, BORAGINACEAE 147 calyx, lobes about as long as the tube, blunt, spreading. Drupe globose to depressed globose 6 mm. in diameter; pyrenes usually 4 DisTRIBUTION: Pakistan, Kashmir, India, and Abyssinia. West Pakistan; Attock Dist.: Campbellpur, Kala Chitta hills, 600 m., R. R. Stewart 13620 (GH). Gujrat Dist.: Pabbi hills, R. N. Parker 3327 (a). Ha- ZARA Dist.: Mirpur, Kazmi 847 (pes). Kouat Dist.: Thal, Aitchison 515, 529 (GH). LanHore Dist.: Changa Manga ane, Kazmi 2498 (PEs). QUETTA Dist.: Harnai, Lace s.n. (K). RAWALPINDI Dist.: Murree Road, R. R. Stewart 1671 (A); near Rawalpindi, R. R. Stewart 13777 (GH). Miscellaneous: Shah Belaweel, Stocks 539 (kK). Reported from Salt Range, Kalat: Pab, Wahir and Las Bela, and Sind. Kashmir: Kotli beyond Mirpur, A. Rashid in R. R. Stewart 27243 (BM). 3. E,. laevis Roxb. Coromandel Pl. 1: 42. t. 56. 1795, Fl. Indica ed. Carey & Wall. 2: 341. 1824, ed. Carey 1: 597, 1832; C. B. Clarke in Hook. f. Fl. Brit. India 4: 141. 1883; Brandis, Forest Fl. India 340. 1874, Indian Trees 481. 1921; Parker, Forest Fl. Punjab 362. 1924; rome Lahore Dist. Fl. 167. 1936; Riedl in Rechinger, FI. Tranica 48: 7. 1967. E. affinis Wall. Cat. No. 900. 1829, nomen nudum. E. dichotoma Rottl. in Wall. Cat. No. 904. 1829. E. punctata Roth, Nov. Pl. Sp. Indiae Orient. 126. 1821. Beurreria laevis G. Don, Gen. Syst. 4: 390. 1838. Beurreria punctata G. Don, Gen. Syst. 4: 390. 1838. E. floribunda Benth. in Royle, Illustr. Bot. Himal. Mount. 1: 306. 1839. E. sari var. floribunda (Benth.) Brandis, ree Fl. India 340. 1874; C. B. Clarke in Hook. f. Fl. Brit. India 4: 143. E. ea var. platyphylla Merrill, Lingnan Sch adn 14: 55. 1935. Type: “Circar Mountains” without citation of collector’s name (CAL?). Icon.: Roxb. l. c. ¢. 56. 1795; Wight, Icon. Pl. Indiae Orient. 4: ¢. 1382. 8 A small or large deciduous shrub to 10 m. tall tree, bark smooth gray, twigs glabrous or nearly so. Leaves ovate-elliptic or obovate or even suborbicular, 7-12 cm. long, 5-11 cm. broad, rounded or angled at bases and apices, when mature somewhat lustrous above, when young minutely glandular pubescent below, but except for some hairs in the vein axils becoming glabrous at maturity; petioles 1-2.5 cm. long. Inflorescence appearing just before leaf renewal, terminal and axillary, loosely dichoto- mous, with few bracts on the primary axis, otherwise naked, ultimate branches bearing sessile flowers unilaterally in two crowded ranks, hence somewhat scorpioid, particularly just before anthesis. Flowers white, sessile or subsessile. Cal 1-2 mm. long, tawny-tomentose. Corolla 2.5-3.5 mm. long, subrotate, not conspicuous, tube open, 0.7—-1.5 mm. long; lobes 2—2.5 mm. long, recurving. Style 2-3 mm. long, lobes as much as 1 mm. long. Drupe yellow or orange, 3-4 mm. diameter, endocarp rugose, breaking up at maturity into 4 single-seeded parts. 148 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 DISTRIBUTION: Iran, Pakistan, Kashmir, India, Burma, Hainan, Poly- nesia, Indochina, and Australia. West Pakistan: Atrock Dist.: Kala Chitta hills, R. R. Stewart 13608 (GH). Hazara Dist.: Balakot, Kazmi 2467 (pes). JHELUM Dist.: Mount. Tilla, Kabir 20420 (rAW). LAHORE Dist.: Shahadra near Lahore, R. R. Stewart 15375 (GH). PESHAWAR Dist.: near Peshawar, R. R. Stewart 300b (A). QueETTA DIstT.: Torkhan Pass, above Harnai, Kazmi 2488 (pes). Miscellaneous: Shah Bilaweel, Stocks 607 (kK Kashmir: Reported from Jhelum Valley, Mirpur, and Jammu. S. Coldenia L. Sp. Pi 12%. 1753; ‘Gen, Pl ed. 5.61, 1754. Type sPEcIES: C. procumbens L. Usually branched, diffuse or prostrate herbs. Leaves small, crisped. Flowers white or yellow, sessile or nearly so. Calyx 4—5-partite, segments linear or lanceolate. Corolla tube short, rather broad, naked or with 4-5 appendages within, lobes 4—5, spreading, imbricate in bud. Stamens 4—5, inserted on the corolla tube, included; filaments short; anthers ovate. Ovary ovoid, slightly 4-lobed, 2-celled with 2 ovules or 4-celled with 1 ovule in each cell; styles 2, distinct, cohering from the base to the middle but easily separable, terminal; stigmas capitate. Drupe almost dry, of 4 subconnate, 1-seeded pyrenes. Seeds exalbuminous or with scanty albu- men; cotyledons flat, broad; radical short, straight, incurved or incum- bent on the cotyledons. Species about 11, all American kei the following, which is widely dispersed throughout the warmer region C. procumbens L. Sp. Pl. 125. 1753; Aitch. Cat. Pl. Punjab & Sind 93. 1869; C. B. Clarke in Hook. f. Fl. Brit. India 4: 144. 1883; Cooke, Fl. Bombay Presidency 2: 205. 1908. Type: ‘Habitat in India” Herb. No. 174. 1. (LINN). A procumbent herb, usually lying quite flat on the ground, stems reach- ing up to 45 cm. in length, shaggy with white hairs, branches often numerous, young parts silky with white hairs. Leaves crisped, 12-38 mm. long, 6-18 mm. broad, obovate-oblong, rounded at the apices, coarsely serrate or subpinnatifid, very hairy on both surfaces, base tapering; petioles shaggy, 3-9 mm. long. Flowers pale yellow, solitary, axillary, nearly sessile, Calyx divided to the base or nearly so, very hairy, segments 4, ovate, acute, ciliate, 2.5 mm. long. Corolla naked within, 2.5 mm. long. Stamens 4, scarcely higher than the corolla tube. Fruits dry 4-lobed pyramids, about 3 mm. long, 4 mm. broad at the widest part, grooved on two and ribbed on the other two sides, with a sharp central double beak, hairy, ultimately separating into 1-celled beaked pyrenes. DisTRIBUTION: Pakistan, India, Ceylon, and tropics. West Pakistan: Sind, Stocks 549 (xk). 1970] KAZMI, BORAGINACEAE 149 4. Heliotropium L. Sp. Pl. 130. 1753; Gen. Pl. ed. 5. 63. 1754. TYPE SPECIES: H. europaeum L. Annual or perennial herbs or undershrubs, villous or scabrous. Leaves alternate. Spikes terminal, rarely axillary, dichotomous, branches short or long, scorpioid. Flowers usually white, light bluish, or rarely yellow, bracteate, ebracteate, or bracts much reduced. Calyx 5-partite or -lobed, segments lanceolate or linear, free, rarely coherent at the base. Corolla tubular, cylindrical or infundibuliform, usually hairy outside, sometimes al- so inside, lobes 5, plicate, subvalvate or imbricate, inflexed or patent, linear to roundish. Stamens 5, anthers sessile or with short filaments, inserted on the corolla tube, included, ovate to linear-lanceolate, rarely apiculate. Ovary completely or imperfectly 4-celled, 4-ovuled; style terminal, short or long, ending in a stigmatic disc, with a short or long, more or less conical appendage. Fruits dry, 2—4-lobed, of 4 or less nutlets. Seeds straight or curved, albumen small or none. Species 275, in tropical and temperate zones of both hemispheres. KEY TO THE SPECIES a. Prostrate herbs; calyx divided up to 1/4 of its length, lobes short. Fruit CHIR TP COIR 5 hes Pe ont Sas oe tes 2 24. H. supinum. a. Prostrate, decumbent or erect herbs or undershrubs; calyx divided nearly to the base, lobes free, long. Fruit not enclosed by the calyx. b. Plants completely glabrous Peet oe ee ee eee 23. H. curassavicum. b. Plants variously hairy. c. Stigma with 2 linear, filiform, recurved arms, ee from the corolla. . A. ophioglossum. c. Stigma short, minutely lobed or not, always i included i in corolla. d. Inflorescence partly or fully bracteate, bracts sometimes much educed. e. Pree cig On sca ring fleshy and thick; corolla 1.5-2.5 ong, internally sparsely pilose from 0.5 mm. above re pane cl Sette este et ies 1. H. rarifolium. Stigma elongate conic, stigmatic ring not fleshy, thin; corolla 2.5-4 mm. long, cienadie slightly or densely hairy only at the throat. f. Plants decumbent or prostrate; leaves lanceolate or elliptic; flowers sessile, closely set on the inflorescence. .......... 3. H. marifolium. f. Plants usually erect, sometimes decumbent; est linear to linear-lanceolate ; flowers usually subsessile to pedicellate, rarely sessile; flowers distantly set on the inflorescence. g. Leaves linear, up to 45 mm. long, up to 3 mm. broa inflorescence bracteate ORaL Om os ee HF. seylonicum. g. Leaves linear-lanceolate not exceeding 25 mm. in length, 3 mm. in width; lower flowers of the inflorescence evi- dently bracteate, in the upper flowers — much re- duced or sometimes absent. ......... . H. strigosum. oO 150 JOURNAL OF THE ARNOLD ARBORETUM (wi, 34 d. Inflorescence ebracteat h. Corolla variously ao within Corolla inside clearly with. two hairy zones separated by glabrous zone in between. ....... 17. H. baluchistanicum. Corolla hairy inside, hairs neither in zones nor separated by any glabrous zone. j. Corolla 2.5-3 mm. long; stigma sessile. k. Inflorescence of closely set flowers; calyx lobes un- equal; leaf bases obtuse to acute; anthers 0.5-0.6 mm. long; stigma 0.3-0.4 mm. long. ............ 3 _e . 22 H. ovalifolium. k. Inflorescence of distantly set flowers; calyx lobes equal; leaf bases roundish or sometimes truncate; anthers 1 mm. long; stigma 0.9-1 mm iit ida Gia 16. motiflorum. j. Corolla 4-8 mm. ag stigma ergs or rea style up to 0.7 mm. long l. Corolla 4-5 mm. long, lobes sublinear, plicate, in- flexed into the corolla tube; stigma elongate-conical. 11 ee gillianum. 1. Corolla 5-8 mm. long, lobes ovate or roundish, im- bricate, patent; stigma compressed-conical. 5 Leaves lanceolate to ovate-lanceolate with short petioles, loosely covered with short. more or less erect hairs arising from tubercles, lower surface sometimes glabrous; flowers 3-7, loosely ar- ranged; corolla uniformly hairy inside. ........ ee ee ee eee ee ee 14. H. cabulicum. Leaves ovate to ovate-oblong with long petioles densely covered with long hairs apparently not arising from tubercles; flowers 7-20, closely arranged; corolla thickly hairy inside above the anthers and below the wee lobes, and thinly i between, ...<...2+4: 3 biannulatum. h. Corolla glabrous within. o. Plants grayish-white; inflorescence up to 12 cm. long with loose uniseriate flowers; calyx deciduous. ........ AE SRS SRA eet) By: OMe a ae 19. H. calcareum. o. Plants green to grayish-green; inflorescence 2—5(-8) cm. long with closely set biseriate flowers; calyx persistent. p. Nutlets pubescent to often densely tomentose. .... 20. H ee ee ee ee eee fa > ie ngage p. Nutlets glabrous, Pied one -rugose or istinctly verruculose ei dcava Bank i: s aan. n. Plants perennial q. Corolla lobes broad, oblong, ovate or roundish, subval- vate or imbricate r. Leaves ovate, oblong-ovate or oblong- mene calyx 3.5-4 mm. long; corolla more or less m. long; nutlets free from the beginning . Plants semnene herbs; leaves palmately lobed or simple; two-armed hairs present on stems and petioles: carpellate inflorescences pendent; gynoecium loosely subtended by a bracteole with short-stalked and cupulate or sessile MUSINNTAE RRP AC Lanes an 85 nen te eee ak aes 2. Humulus. 1. Cannabis Linnaeus, Sp. Pl. 2: 1027. 1753; Gen. Pl. ed. 5. 453. 1754. Tall (to 5 m.), erect, branched or unbranched, coarse, tap-rooted, weedy annual herbs occurring in waste places, disturbed ground, and al- luvial flats; stems obtusely polygonal or round in cross section, + ribbed or furrowed, often hollow at maturity, scabrous and sparsely covered with resinous dots, especially on the young parts of the axis. Leaves decussate near the base, alternate toward the apex and on branches, palmately com- pound with 3-15 (usually 5-9) coarsely serrate, long, lanceolate leaf- lets; the lower surface whitish green, with widely spaced yellow to brownish resinous dots and strigose hairs, the upper surface darker green with large, stiff, bulbous-based conic trichomes; stipules narrowly triangular. Plants dioecious, rarely monoecious except in certain cultivated strains and experimental plants; dimorphic, staminate plants tall, slender, with few leaves in the terminal inflorescence, dying soon after flowering; car- 1970] MILLER, GENERA OF CANNABACEAE 189 pellate plants stocky, with a dense crown of leaves associated with the in- florescence, living for several months after pollination. Staminate in- florescences loose cymose panicles, long, if originating from the axils of foliage leaves, short, if from the axils of stipules. Staminate flowers pedi- cellate, small, pendent at maturity, caducous soon after shedding pollen. Tepals greenish, rarely purplish, quincuncial in bud and widespread at an- thesis. Anther walls thin, bearing glandular hairs at the junctures of anther lobes; pollen sacs 4 (bilocular at anthesis) ; filaments flaccid. Car- pellate inflorescences congested series of axillary false spikes; the main axis, leaves, petioles and bracts associated with carpellate inflorescence often densely covered with resinous capitate or + sessile glandular hairs. Carpellate flowers sessile or nearly so, ensheathed by a glandular, beaked perigynal bracteole and subtended by a bract (stipule?). Perianth papery, continuous, closely appressed to the ovary. Fruit an ovoid, somewhat compressed achene with the perianth persistent, reticulate, and often mot- tled with brown; pericarp 5 layered, the inner palisade layer the thickest. Seed 1, ovoid; perisperm 1 cell-layer in thickness; endosperm fleshy, with a chalazal haustorium; embryo curved, U-shaped, flexed in the region of the epicotyl, with the radicle and cotyledon tips pointing toward the micropylar end of the seed (ovary apex); 27 = 18 + XX or XY. TyprE SPECIES: C. sativa L. (Ancient Greek and Latin name for hemp, said to come from Arabic, kinnab, or Persian, kannab.) — HEMP, MARIJUANA. A genus of one (or perhaps more) species, originally indigenous to tem- perate parts of Asia, probably to the desert region immediately to the south and east of the Caspian Sea, but now occurring as an adventive or naturalized weed throughout most of the world. Cannabis sativa, a de- cided nitrophile that grows well in close association with man and his wastes, is widespread in the United States and most frequently collected in Kansas, Nebraska, Iowa, and Illinois. It is known from only a few specimens and literature reports imply.* Cannabis ruderalis Janisch. (Uéen. Zap. Gos. Saratov. Univ. 2(2): 14. 1924), a possible second species oc- curring in Central Asia, southeastern Russia, and as a weed in eastern and Central Europe, is reported to differ from C. sativa in size and achene characters. Most authors accept only one species, and whether the diag- nostic characters of C. ruderalis are distinct and do not overlap the ex- tensive variability of C. sativa remains to be worked out carefully. Cannabis sativa has been much used as an experimental plant. Inter- est was focused early on monoecious hemp plants and the “reversion” of carpellate plants to staminate ones and vice versa. For example, seeds planted in summer at Columbus, Ohio, yielded about equal numbers of carpellate and staminate plants, but seeds from the same lot sown indoors * Haney and Bazzaz oar ag mae hs apparent rarity of Cannabis sativa in the PRE rah United St der of naturalized hemp in North America mostly from temperate es aod che fiber strains which are ill adapted to growing under environmental conditions present in the Southeastern States. 190 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 in December, gave a crop in which more than three-quarters of the car- pellate plants produced intersex or pure male flowers and a similar pro- portion of the staminate plants occurred with intersex or pure female flowers. After the discovery of photoperiodic responses in plants about 1920, it was realized that hemp was a short-day plant in which sex re- versal was correlated with exposure to ever shorter periods of light. Other factors, including temperature and available soil nutrients, ap- parently also exert some control. However, the genetic basis of sexuality in hemp is still not fully understood. Although environment was original- ly considered to have the major role and Mendelian inheritance to be in- consequential, different workers have demonstrated the presence of a hetero- morphic chromosome pair in staminate meiotic material from several dioecious strains. A heteromorphic pair has been less convincingly shown in certain monoecious strains. In dioecious hemp, sex determination is thought to be under an XX = @ and XY = 4@ system, and derived monoecism has been explained in two ways: monoecious plants are all XX, and variation in sex expression is due to heterozygous genes on the X-chromosomes and autosomes; or XX, XY, and YY monoecious plants can occur, but only autosomes determine sex. Unfortunately, somatic karyotypes have been imperfectly investigated. The morphology of the spicate carpellate inflorescence of hemp has been interpreted in several ways. Buds are reported to occur in the axils of both stipules and leaves, and, according to Camp (1932), stipular buds (two at each node) give rise to short determinate branches bearing single flowers, whereas the true axillary bud produces an indeterminate branch bearing leaves and stipular flowers but no secondary axillary branches. Bud primordia are visible in the axils of reduced leaves borne on the axillary branch, but suppression is broken only under extraordinary conditions. In an alternate interpretation (see Hayward, 1938) a pair of flowers, each subtended by a stipule, is claimed to arise in a leaf axil. A secondary branch, also issuing from the leaf axil, bears another leaf with stipules and flowers, and this, in turn, produces an axillary branch of the third order. The staminate inflorescence is composed of a main indeterminate axis bearing secondary branches from both the axillary and stipular buds near the base, but only from the stipular buds toward the apex (Camp, 1932). The staminate inflorescence of hemp is called a panicle or raceme by most authors, but the side branches are clearly cymose. Zander has studied the poorly differentiated, unbranched, and inarticu- late laticifers that occur in the stem of hemp. He concludes that “the anatomical behavior and developmental history of hemp laticifers are different in comparison with the rest of Moraceae”’ (transl.). Three economically important products are obtained from Cannabis sativa, Perhaps least significant (except in Russia) is a drying oil pressed from the fruits. The oil is used as a substitute for linseed oil in paints and varnishes and occasionally in soap making. After expressing, the “seed”- cake may be fed to cattle. In parts of eastern Europe and Russia roasted hemp “seeds” are eaten by humans, and in the United States Cannabis 1970] MILLER, GENERA OF CANNABACEAE 191 achenes are sold in seed mixtures prepared for cage-birds and outdoor bird-feeders. At one time uneaten achenes from such mixtures were a major source of spontaneous hemp plants, but now the “seeds” are sold only after the embryos have been killed. In many parts of the world hemp is grown on a commercial basis for fibers which are made into ropes, twines, bags, and webbing. Clothing was once made from the finer grades, and the strong, durable fibers of hemp were used extensively for sail cloth. A tarred caulking material, oakum, is produced from shorter fibers and tow. Hemp is most success- fully grown for fibers on rich, humified loam in a mild, humid climate. Individual fibers are about a centimeter long and occur in two zones in the stem. The outer, primary fibers are longer and thicker walled than the inner, secondary fibers, and both series are separated by several layers of parenchyma, a feature that enables the isolation of the primary fiber- bundles by retting. In general, internode and fiber-bundle length are in direct proportion, and tall cultivars with fewer nodes are sought. Staminate plants produce the finest grade of fibers. Because staminate plants die soon after shedding pollen, while carpellate plants live until the fruits mature, breeding programs have been directed toward producing either monoecious strains with the tall growth-habit of the staminate plant or uniform-ripening dioecious races. Colchicine-induced tetraploid strains, which give a greater yield of higher quality fiber than normal diploids, are grown to some extent. Hemp agriculture in the United States first started on a limited basis in New England, in 1632, with seeds brought to the Colonies from England. In the 1800’s, particularly during the midcentury years, hemp was extensively grown in the Bluegrass Region of Kentucky from seeds of Chinese origin. During the 1850’s, 75,000 tons were har- vested annually in the United States, but production had dropped to 5000 tons or less by the early 1900’s. Acreage in hemp dramatically in- creased during both World Wars. In 1943, as a response to the inac- cessibility of Manila hemp, nearly 63,000 tons were produced. All hemp fiber used in the United States in recent years has been imported, gen- erally from European sources. The hemp plant is the source of an important narcotic drug. Its effect on humans, although at present mostly lacking scientific documentation, seems to be to induce a general state of euphoria, frequently broken with periods of depression, and often accompanied by vivid mental imagery. The drug is apparently widely used throughout the World, in spite of severe penalties. There is some evidence that the use of hemp drugs in the United States started about 1910 in New Orleans. The narcotic principle occurs in the resinous material that accumulates most abundantly on tissues of the carpellate inflorescence. Among the numerous compounds isolated from the resin in recent years, the dibenz-a-pyrans, A!- and Al- transtetrahydrocannabinol (THC), have been found to be the major agents of intoxication in man. At present, tetrahydrocannabinols are reported from no other plant. Methods of taking the drug vary somewhat from culture to culture. 192 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 In the United States, marijuana is a mixture of the resin and various plant fragments, generally from the carpellate inflorescence, while Aashish is a term restricted to the pure resin. Both forms are most often smoked. In India the resin may be eaten or drunk in various carriers, especially milk. The word assassin, taken directly from the Arabic hashshdshin, hemp eaters, entered the English language at the time of the Crusades. The hashshashins were a group of murderers who were supposed, perhaps erroneously, to have carried out their missions, while intoxicated by hashish. The largest amount of resin is produced by flowering carpellate plants in hot, dry climates. An increase in humidity lowers production, and stems, lower leaves, and seeds contain the least amount of resin. Plants grown in India have been reported to contain 20 per cent resin, those from Mexico 15 per cent, and samples from Kentucky and Wisconsin 8 and 6 per cent, respectively. There are conflicting statements in the literature about the occurrence of THC in carpellate and staminate plants of dif- ferent geographic origin and in strains grown for fibers and oil. Selection of high-yield fiber or oil cultivars apparently may result in a loss of THC, in spite of the retention of high resin-production. Staminate plants from certain strains lack THC, but the intoxicating principle may occur in male plants of other cultivars. To date, a thorough analysis of the resin from wild self-seeding types and cultivated dioecious and monoecious strains grown in various parts of the World for fiber, oil, or the drug has not been made. Gas chromatographic methods would be useful in such a study. Forensic identification of marijuana is based on microscopic examina- tion of the sample, in conjunction with one or several chemical tests ap- plied to a petroleum-ether extract of the resin. The glandular and non- glandular hairs occurring on the upper and lower leaf surfaces, petioles, and bracts of the inflorescences of both sexes are considered distinctive, especially when compared with an authenticated specimen. Since the cystoliths of Cannabis sativa are composed of calcium carbonate, addi- tion of dilute hydrochloric acid will cause a slight effervescence, which is visible through a low-power microscope. REFERENCES: The references have been selected from an extensive bibliography dealing with various aspects of hemp. For additional literature on various topics refer to the cited bibliographies and review articles. Under family references see BECHTEL, Hitt, Martin, SCHREIBER (pp. 290-295), Srnord, WALTER (pp. 875-909), and WESTERGAARD, ApaMs, R. Marihuana. Science 92: 115-119. 1940. [Economic botany, effect of drug, behavior of marijuana users, and resin chemistry. | ANDREWS, G., & S. VinKENooG, eds. The book of grass, an anthology on In- dian hemp. xiv + 242 pp. New York. 1967. [Reviewed in Econ. Bot. 23: 82, 83. 1969.] Arnoux, M. Influence des facteurs du milieu sur l’expression de la sexualité du chanvre monoique (Cannabis sativa L.). I. Action du cycle photopério- 1970] MILLER, GENERA OF CANNABACEAE 193 dique. Ann. Amél. Pl. 13: 27-49. 1963; II. Action de la nutrition azotée. Ibid. 16: 123-134. 1966; III. Note yt gaa entre le cycle photo- périodique et la nutrition azotée. /bid. 62. Aen, A. L. Hemp — production and pany Econ. Bot, 2: 158-169. 1948, Hemp-growing and -usage in the U.S.] Bessey, E. A. Sex problems in hemp. Quart. Rev. Biol. 8: 194-200. 1933. Buiatt, A. H. A critical survey of the literature dealing with the chemical con- stituents of Cannabis sativa. Jour. Wash. Acad. Sci. 28: 465-477. 1938 BortHwick, H. A., & N. J. Scutty. Photoperiodic responses of hemp. Bot. Gaz. 116: 14-29. 1954. Bouquet, R. J. Cannabis. Bull. Narcotics 2(4): 14-30. 1950; 3(1): 22-45. 1951. [Literature review with emphasis on the drug BurKILL, I. H. A dictionary of the economic products of the Malay Peninsula. ed. 2. 2 vols. Kuala Lumpur. 1966. [Cannabis 1: 442-446. ] Camp, W. H. The floral anatomy of hemp (Cannabis sativa L.). Ohio State Univ. Abstr. Doct. Diss. 9: 50-59. . The antiquity of hemp as an economic plant. Jour. N.Y. Bot. Gard. 37: 110-114. 1936. CuaAren, S. Facts about marihuana, a survey of the literature. Am. Jour. Pharm. 117: 422-430. 1945. [Mainly effects of marihuana usage on man. | CueEvuvart, C. Expériences sur le développement de Cannabis sativa L. (sex- ualité et pigments foliaires) a température constante et sous différents ré- gimes de photopériodisme. Acad. Belg. Bull. Sci. V. 40: 1152-1168. 1954. COMMISSION ON Narcotic Drucs. The question of cannabis: cannabis bibliog- raphy. 250 pp. United Nations Economic and Social Council E/CN/479. 1965.* CoUNCIL OF SCIENTIFIC AND INDUSTRIAL RESEARCH. The wealth of India. Raw Materials. Vol. 2. xx + 427 pp. Delhi. 1950. [Cannabis, 58-64.] Dewey, L. H. The hemp industry in the ail States. Yearb. U.S. Dep. Agr. 1901: anne 3 pls. Washington. mp. Jbid. 1913: 283-346. 7 iy 1914, [ Cultivation. ] Exoratas, G. Pollen morphology and plant taxonomy. Angiosperms. Corrected rint and new addendum. xiv + 553 pp. New York & London. 1966. FARNSworTH, N. R. Hallucinogenic plants. Science 162: 1086-1092. 1968. [ Cannabis, 1086-1088. | Pharmacognosy and chemistry of Cannabis sativa. Jour. Am. Pharm. Assoc. II. 9: 410-414, 440. 1969. Gamace, J. R., & E. L. ZerKin. A comprehensive guide to the English-language literature ‘on Cannabis (Marihuana). STASH Bibliographic Series 1. xii + 265 pp. STASH Press. Beloit, yong 1969. [Author and subject indices; includes references through early 1969. ] H. The ancient cultivation of hemp. Antiquity 41: 42-49, 137, 138. 1967. [Europe. . Pollen-analytic evidence for the cultivation of Cannabis in England. Rey. Palaeobot. Palynol. 4: 71-80. 1967. [Hemp grown in England since early Anglo-Saxon time, ca. 400 a Goong, E., Ep. Marijuana. xvi + 197 pp. New York. 1969. [ Sociological, psychological, legal, and medical aspects Grinspoon, L. Marihuana. Sci. Am 1(6): 17-25. 1969. Haarer, A. E. Hemp (Cannabis Cae World Crops 5: 445-448. 1953. [Cul- tivation for fibers. | 194 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Haney, A., & F. A. Bazzaz. Some ecological implications of the distribution of hemp "(Cannabis sativa L.) in United States. Proc. 1st Int. Conf. Bot. Chem. Cannabis. (In press. Ciba Foundation. London.) Haywarp, H. E. The structure of economic plants. x + 674 pp. New York. 1938. [Morphology and anatomy of C. sativa, 214-245.] HIRATA, = Sex determination in hemp (Cannabis sativa L.). Jour. Genet. 19: 65-79. 1928. [Results of breeding experiments. | ——. ‘Cytological basis of sex determination in Cannabis sativa L. Jap. Jour. Genet. 4: 198-201. 2 pls. 1929.* [Heteromorphic pair of chromosomes ob- served in microsporocytes. | Horrcans W. Die Vererbung der Geschlectsformen des Hanfes (Cannabis sa- L.). I. Ziichter 17/18: 257-277. 1947; II. Ibid. 22: 147-158, 1952. teenies and chromosome ae aps , Cannabis sativa L. In: Roemer, T., & W. Ruporr. Handbuch der Pflanzenziichtung. ed. 2. 5: 204-264. Berlin. 1961. [Review of breed- ing methods and results Hopkins, J. F. A history of the hemp industry in Kentucky. xii + 239 pp. Lexington, Kentucky. 1951, [Hemp-growing for fibers. | Kincssury, J. M. Poisonous plants of the United States and Canada. xiv + 626 pp. Englewood Cliffs, N.J. 1964. [ Cannabis, 222-225.] Kouter, D. Die Entwicklung von Cannabis sativa unter dem Einfluss verschie- dener Tageslingen. Physiol. Pl. 11: 249-259. 1958. [Photoperiodic ef- fects . Homozygous males in hemp. Nature 195: 625, 626. 1962. [Breeding results. Kunpbvu, B. C. The anatomy of two Indian fibre plants, Cannabis and Cor- chorus, with special reference to fibre distribution and development. Jour. Indian Bot. Soc. 21: 93-128. 1 pl. 1942 LERNER, M. Marihuana: tetrahydrocannabinol and related compounds. Science 140: 175, 176. 1963. [Gas chromatography of marijuana seep Mackay, E. L. Sex chromosomes of Cannabis sativa. Am. Jour. Bot. 26: 707, 708. 1939. [Heteromorphic chromosomes visible in aoe division meta- anaphases from staminate buds. McPuee, H. C. The genetics of sex in hemp. Jour. Agr. Res. 31: 935-943. 1925 [1926]. [Selfed, normally pistillate plants produce ‘only pistillate progeny. | MEcHOoULAM, R., & Y. Gaont. Recent advances in the chemistry of hashish. Fortschr. Chem. Organ. Naturstoffe 25: 175-213. 1967.* MENZEL, M. Y. Meiotic chromosomes of monoecious Kentucky hemp (Canna- bis sativa). Bull. Torrey Bot. Club 91: 193-205. 1964. [Confirmation of heteromorphic pair of chromosomes in males of dioecious strains, but no heteromorphic pair found in monoecious plants. Monan Ram, H. Y., & R. Natu. The morphology and embryology of Cannabis sativa Linn. Phytomorphology 14: 414-429. 1964. Moors, L. A., Jr. Marijuana (Cannabis) bibliography, 1960-1968. Unpaged fi + 55 ped Bruin Humanist Forum. Los Angeles. 1969. Murpny, H. B. M. The cannabis habit: a review of seca psychiatric litera- ture. Bull. tae 15(1): 15-23. 1963. NAKAMURA, G. R. Forensic aspects of cystolith hairs of Cannabis and other plants. Jour. Assoc. Official Agr. Chem. 52: 5-16. 1969 Postma, W. P. Mitosis, meiosis en alloploidie bij Cannabis ‘sativa en Spinacia 1970] MILLER, GENERA OF CANNABACEAE 195 oleracea. (In Dutch; English summary.) x + 83 pp. Haarlem. 1946. | Nu- clear cytology. | Ram, M. Occurrence of endosperm haustorium in Cannabis sativa L. Ann. Bot. II. 24: 79-82. 1960. Roprinson, B. B. Hem mp. U. S. Dep. Agr. Farmers’ 1935. 16 ah [Hemp growing instructions issued in the U. S. during W. W. II a re- sponse to the ee of Manila hemp. | RosEvear, J. Pot, andbook of marihuana. 160 pp. New Hyde Park, New York. 1967. ene aspects. | Roy e, J. F. Fibrous plants of India. xiv + 403 pp. London. 1855. [Canna- bis, 314-340.] SAVELLI, R. Poliembrionia in Cannabis sativa L. Arch. Bot. Forli 4: 128-137. 1928.* SCHAFFNER, J. H. The fluctuation curve of sex reversal in staminate hemp plants induced by photoperiodicity. Am. Jour. Bot. 18: 424-430. 1931. [Sex “reversals” in individuals from seeds planted bimonthly for one year. SCHILLING, E. Zur Morphologie, Physiologie und diagnostischen Bewertung der Bastfasern von Cannabis sativa. Ber. Deutsch. Bot. Ges. 41: 121-127. 1923. [ Fibers. | SCHULTES, R. E. Hallucinogens of plant origin. Science 163: 245-254. 1969. [ Cannabis, 247, 248.] ScHULTz, O. E. Der Gegenwirtige Stand der Cannabis- eae Planta Med. 12: 371-383. 1964. [Chemistry of resin constituen Stmmons, J. L., ed. Marihuana, myths and realities. ee pp. North Holly- wood, Cultfoendu: 1967. [Sociological and legal aspects. | SINGH, S. P. Floral anatomy of Cannabis sativa L. Agra Univ. Jour. Res. Sci. 5(1): 155-161. 1956 SoLomon, D., ed. The marihuana papers. xxvi + 448 pp. Indianapolis. 1966. Reprinted as Signet Book W3442. 509 pp. New York. 1968. [An anthology of selected historical, psychological, and sociological writings on marijuana; includes a partial reprint of the LaGuardia Report (1944) on the mari- juana problem in New York City. ] Topp, A. R. The chemistry of hashish. London Roy. Coll. Sci. Sci. Jour. 12: 37-45. 1942.* . The hemp drugs. Endeavour 2: 68-72. 1943. [Chemistry of resin con- stituents. ] Torrott, F., U. Avico, & E. S. Crrannt. Methods of distinguishing biologically active cannabis and fibre cannabis, Bull. Narcotics 20(1): 55-59. 1968. [Possible to distinguish Indian hemp from Cannabis grown for fiber by using gas chromatography of the resins produced by each. | Watton, R. P. Marihuana, America’s new drug problem. x + 223 pp. Phila- delphia. 1938. [Broad review of the drug properties of hemp and the so- ciology and history of marijuana usage; written about the time the U. S Federal Marijuana Tax Act was passed. | Watt, G. A dictionary of the economic products of India. 6 vols. + index. Calcutta. 1889-1896. [Cannabis, 2: 103-126. 1889.] Wart, J. M., & M. G. Breyer-BrANDWIJK. The medicinal and poisonous plants of southern and eastern Africa. ed. 2. xii + 1457 pp. Edinburgh & Lon- don. 1962. [ Cannabis, 759-772.] Wayne, E. Chairman. Advisory Committee on Drug Dependence. Cannabis. 196 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 viii + 79 pp. London. 1968. [Bibliography of clinical literature, pharma- cology of marijuana, and data on marijuana usage in the U.K. Wen, A. T., N. E. Zrnzerc, & J. M. Netson. Clinical and psychological effects of marihuana in man. Science 162: 1234-1242. 1968. ee G. E. W., & J. Knicut, eds, Hashish: its chemistry and phar- cology. viii + 96 pp. Ciba Foundation Study Group No. 21. London. 1965 YOUNG, W. M. Chairman. Report of the Indian Hemp Drugs Commission, 1893-94. 7 vols. Simla. 1894. [Probably the most lengthy published ac- count of the effect of marijuana on man. Vol. 1: General information on 2, 3: Appendices; vols. 4-7: Evidence presented by witnesses. | ZANDER, A. Uber Verlauf und Entstehung der Milchréhren des Hanfes (Canna- bis sativa). Flora 123: 191-218. 1928. [Laticifers.] 2. Humulus Linnaeus, Sp. Pl. 2: 1028. 1753; Gen. Pl. ed. 5. 453. 1754. Rough, scandent and dextrorse-twining annual or perennial herbs of river and stream bottoms, thickets, hedgerows, roadsides, and other dis- turbed sites; with long taproots, or, if perennial, with a massive rhizome and abundant adventitious roots; aérial stems weakly ridged, often hexag- onal in cross section, rigid two-armed climbing hairs frequent near the apex and often on multicellular emergences; axillary branches mostly from upper nodes. Leaves generally decussate, or alternate when asso- ciated with an inflorescence, sometimes simple and cordate but more often palmately lobed, serrate; venation strictly palmate; petioles with two-armed trichomes; lower leaf surfaces with clear or yellowish resinous dots and widely spaced hairs on lamina and major veins, upper surfaces densely covered with cystoliths and short, rigid conic hairs; stipules lan- ceolate, two per petiole or coalesced laterally into one. Plants dioecious, rarely monoecious when growing in nature. Staminate inflorescences axillary, sometimes terminal, erect or + pendent, many flowered, loose cymose panicles. Staminate flowers pedicellate, small. Tepals greenish, bearing on the abaxial surface glandular dots and occasionally long, rigid trichomes. Glandular dots at junctures of the anther lobes either pres- ent or absent. Carpellate inflorescence spicate, consisting of a series of ovate or acuminate stipular bracts and bracteoles subtending solitary or paired flowers; bracts and bracteoles pubescent or ciliate, with sessile glandular dots or short-stalked cupulate lupulin glands on abaxial sur- faces. Carpellate flowers with short pedicels, borne in the axil of a brac- teole which at maturity is large and either brown and membranaceous or ae foliar and green, sometimes reddish pigmented. Perianth continuous, thin, appressed to the ovary, either covered with lupulin glands or not. Fruit a lenticular or + round achene, tightly or loosely ensheathed by the brownish or sometimes mottled persistent perianth; pericarp crus- taceous. Seed 1; embryo coiled into a tight spiral, cotyledons long, rad- icle pointing toward the micropyle. Type me ei _— L. (Late Latin name for this plant, from Old Slavic.) — 1970| MILLER, GENERA OF CANNABACEAE 197 A genus of two species native to portions of the North Temperate Zone but now widely cultivated, escaped, or naturalized throughout temperate areas of both hemispheres. Originally indigenous to eastern Asia (China, Taiwan, Manchuria, Korea, and Japan), Humulus japonicus Sieb. & Zucc.* is naturalized in eastern North America southward to North Caro- lina, South Carolina, and Tennessee. It differs from H. Lupulus L. in many characters, among which the occurrence of ciliate hairs on the margins of the bracteoles and stipular bracts, the absence of lupulin glands, and the presence of from 5 to 7 leaf lobes (vs. 0-3 [5] in H. Lupulus) are the most readily observed. Wild H. Lupulus is disjunct in the Northern Hemisphere. It occurs in Japan, a part of North America, and from central Asia to southern Europe, a range that appears to fit one of the Arcto-Tertiary distribution patterns. The populations in each re- gion have been considered distinct species, although recent work indicates that only subspecific differences exist between them. Humulus Lupulus presently is found throughout most of the United States and southern Canada, but the actual range of the native North American hop, which Nuttall, in 1848, named H. americanus (based on a Gambel collection Ww ' 8: spread have made it difficult to decide whether a particular plant is native or introduced. However, pollen of H. Lupulus has been found at several sites in the upper Midwest in lake sediments deposited long before the presence of European man in the New World. Future detailed analysis of postglacial sediments from elsewhere in North America could ultimately solve the problem of the pre-settlement distribution of the native hop on this continent. The spicate carpellate inflorescence of Humulus consists of a condensed primary axis bearing reduced secondary branches (Ehara, 1955; Hama- guchi, 1955). Internodes, although shortened, are clearly discernible. In H. Lupulus two flower pairs normally occur per node, each pair being subtended by a bract and each individual flower by a bracteole. Pe- duncles and pedicels are short, and the flowers are subsessile. The nodal flower clusters are interpreted as cymes in which the third (first opening) flower of each pair and the flower or branch between the pair are sup- pressed. Infrequently a reduced leaf is found at certain nodes between the two enlarged bracts, indicating their homology with stipules. Stipular bracts generally have pointed apices, while bracteoles are obtuse. The “Merrill (Trans. Am. Philos. Soc. 24: 138. 1935) contends that Antidesma scan- dens Lour. (Fl. Cochinch. 2: 617. 1790) is unmistakably the species widely known as Humulus japonicus Sieb. & Zucc. (Abh. d. Wiss. Miinchen 4(3): 213. 1846; also i io . Fam. Nat. 2: 89. 1846). If Merrill is correct, his combination, Humulus jcolien (Lour.) Merr., should be adopted. Un- fortunately, Loureiro’s reaper based on only a staminate plant, is eid ie and mentions glabrous leaves and unarmed stems, two characters that are difficult to ner to H. ri rie bi Further, since the specimen upon which Loureiro based is description apparently has not been preserved (see Merrill, loc. cit.), the appli- cability of the name H. scandens seems open to question 198 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 S * Gly tf ils sk dH ad acc 2 "Wie aged Ke Ke i 7s Humulus. H. japonicus: a-j. a, terminal staminate inflorescence, x /4; b, staminate flower, X 10; c, carpellate flower wi artly developed ; : ing ovule, style branches rminal carpellate infructescence, X 1/2; f, fruiting . . : x g, nearly mature achene in axil of bracteole, X 5; h, mature achene with tightly appressed perianth, X 6; i, achene with embryo in diagrammatic longitudinal 1970] MILLER, GENERA OF CANNABACEAE 199 carpellate spikes of H. japonicus and H. Lupulus are morphologically sim- ilar except that in the former only one flower (with its associated brac- teole) occurs in the axil of a stipular bract. Sterile bracts are commonly present at the base of the inflorescence. The diploid chromosome number in Humulus Lupulus is 20, and Winge in 1923 first demonstrated a heteromorphic pair of sex chromosomes (XY) in staminate plants of European origin (see Ono & Suzuki, 1962). Although carpellate plants of this species apparently uniformly have a homomorphic pair of sex chromosomes (XX), more recent research on the chromosomal makeup of staminate hop has shown considerable varia- bility which seems to be correlated with the geographic origin of the plants. Working with the wild hop of Japan, Ono (1955, 1962) found that the sex chromosomes almost invariably took multiple forms con- sisting of four or six chromosomes. Both types, the more frequent tetra- valent and the rarer hexavalent, presumably have arisen by reciprocal translocations between the original XY chromosome pair and certain autosomes. At meiosis, components of the sex chromosome complex con- jugate end to end. Cytological studies on staminate European and American hops have been less extensive, but, in general, the simple XY type seems to prevail in plants from Europe, while in American samples homomorphic XY males, heteromorphic XY males (X length: Y length = 10:8 instead of 10:5 as in the European hop) and tetravalent complexes have been observed. Sex chromosomes have been found in H. japonicus also, but instead of an XY mechanism, the staminate plant has a 2n chromosomal complement of 14 autosomes and 3 sex chromosomes, Y,X Yo. The Y chromosomes are heterochromatic for most of their length. Dur- ing anaphase I of meiosis in staminate plants, 8 chromosomes go to one pole and 9 go to the other. Carpellate plants have a 2” number of 14 + XX Bracteoles, perianths, and, to a lesser degree, stipular bracts of the mature fruiting spike of Humulus Lupulus are covered with large cup- shaped glands filled with a yellow resinous substance, lupulin, that is used to impart flavor and aroma to beer. Bacteriostatic agents in lupulin also inhibit the growth of spoilage bacteria, while permitting yeast to develop freely and ferment the wort, a mixture of sugar (obtained by enzymatic conversion of starch from various grains) and water. Hops may be added to the wort while it is being boiled prior to the onset of fermentation or to the final beer in the storage tank. In the former case, the bittering sub- stances are formed from the resin constituents during boiling, while in the latter the flavoring is derived from essential oils in the resin Lupulin is composed of soft and hard resins, but only donipanietits of the soft resin apparently contribute to a beer’s flavor. The aroma of beer section, area aoe hatched, * 6; j , rigid 2-armed trichome from petiole, « 25. B,. pul k, mature Pel in axil of bracteole, * 4; 1, mature achene with pe ee = inflated perianth and lupulin — >< 10; m, rigid 2-armed tricheae: frank petiole on multicellular pedestal, x 25 200 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 derives solely from essential oils, a distinct group of compounds. Soft resin can be separated into a-acid, B-acid, and uncharacterized fractions, each category containing a number of distinct organic compounds. Isom- erization of the a-acids during boiling yields water-soluble, bitter- tasting isohumulone and its analogues, which strongly inhibit the growth of Gram-positive bacteria. Lupulone in the f-acid fraction is also bac- teriostatic. The percentage of cohumulone in the a-acid component has been shown to vary in amounts which correspond to groupings or morpho- logical features recognized as characteristic of hops either native or long- cultivated in England, the European continent, and America (Davis & Burns, 1962). The cultivated hop was introduced into North America in 1629, and by 1650 commercial crops were being harvested in Virginia and New Netherlands. During the 1800’s and early 1900’s New York was an impor- tant region of hop cultivation, although following the Civil War, the in- dustry began to spread westward through Pennsylvania to Wisconsin. In the United States today the leading areas of hop production are Idaho, Washington, Oregon, and California, where three principal varieties are grown. The first, the English hop, was obtained by crossing a wild car- pellate plant from Manitoba with an English staminate plant, and the second, the Fuggle hop, was selected from seedlings raised in England. The origin of the third, Cluster hops, is less certain. The Late Cluster type originated during Colonial times on the eastern seaboard, perhaps as a seedling from an English variety then in cultivation, while the Early Cluster type was derived as a bud sport directly from the Late Cluster hop. At the present time the most widely grown hops in the United States are the Cluster varieties. Hops are also cultivated in Canada, Europe, northern India, China, Manchuria, Japan, South Africa, southeastern Australia and Tasmania, New Zealand, and southern South America. Pollination is not necessary for the development of the spike and lupulin glands, so staminate plants are rarely raised in hop yards, except for breeding purposes. Rhizome cuttings are used for propagation. Hops require a well-drained, nitrogen-rich soil and will not tolerate sharp temperature fluctuations or excessive rainfall during the growing season. Late summer dryness is necessary for proper maturation of the spikes, which must be picked within a few days of full ripeness, otherwise they are prone to shattering, and the lupulin quality is inferior. After harvesting, hops are dried in warm air kilns for several hours, cooled or cured in another building for about a week to equalize moisture content in the various pickings, baled, and marketed. In former times hops have been used medicinally as multipurpose tonics and to treat certain types of epidermal sores and irritations, usages per- haps related to the bacteriostatic quality of the resin. In Scandinavia phloem fibers from the aérial stems of H. Lupulus are made into twine and cloth to a limited extent. Humulus japonicus is occasionally used as an ornamental plant. 1970| MILLER, GENERA OF CANNABACEAE 201 REFERENCES: The references have been selected from a lengthy bibliography of publica- tions dealing with this genus. Only reviews or references of specific interest are included. Under family references see BECHTEL, HILL, MARTIN, SCHREIBER (pp. 283-290), StNoT6é, WALTER (pp. 822-874), and WESTERGAARD. BearD, F. H. Hops: their varieties and cultivation. Jour. Inst. Brewing 39: hiS—125;, 1943." Bent, PF. Twining of the hop (Humulus Lupulus L.). Nature 181: 1009, 1010. : BisHop, L. R. The resins of hops as antibiotics. Soc. Exper. Biol. Symp. 3: 101-104. 1949, Brooks, S. N., C. E. Horner, & S. T. Likens. Hop production. U.S. Dep. Agr. Agr. Inf. Bull. 240. 46 pp. 1961. [Cultivation in the U. S.] & S. T. Likens. Variability of morphological and chemical quality characters in flowers of male hops. Crop Sci. Madison 2: 189-192. 1962.* [See also Diss. Abstr. 22: 978. ] Burcess, A. H. Hops — botany, era and utilization. Jn: N. PoLuNrn, en. World Crop Books. xx + 300 pp. London & New York. 1964. CusHIne, E. J. Late-Wisconsin ona ieee and the glacial sequence in Minnesota. Pp. 59-88. 1 pl. In: E. J. CusHine & H. E. WricHr, Jr., eds., Quaternary Paleoecology. viii i 433 pp. New Haven, Connecticut, & Lon- don. 1967. [Pre-settlement Humulus pollen in Minnesota. | Dark, S. O. S. A survey of the present te a hop genetics. Wye Coll. Dep. Hop Res. Annual Rep. 1950: 58- OF. Darwin, C. The movements and habits of aiabiee one ed. 2, revised. viii + 208 pp. New York. 1891. [Humulus Lupulus mentioned in chapter on twining plants. | Davis, E. L. Morphological complexes in hops (Humulus Lupulus “" with special reference to the American race. Ann. Missouri Bot. Gard. 44: 271- 294. 1957. [See also Diss. Abstr. 16: 1566. 1956.] & R. L. Burns. The use of vapor fractometry in the analysis of some New England hops. Rhodora 64: 243-251. 1962. [Alpha acid components in three New England collections of wild hops suggest that two are de- rived from English sources and one is of American origin. Epwarpson, J. R. Hops—their botany, history, production and _ utilization. Econ. Bot. 6: 160-175. 1952. Enara, K. Comparative morphological studies on the hop (Humulus Lupulus L.) and the Japanese hop (H. japonicus Sieb. et Zucc.). I. Jour. Kyushu Univ. 10: 209-232. 5 pls. 1955; II. Zbid. 10: 307-324. 5 pls. 1956. Fanc, T. K. Observations on the morphology and anatomy in the rhizome of hop (Humulus Lupulus L.). (In Chinese; English summary.) Acta Bot. Sinica 6: 297-310. 1957.* FREEMAN, O. M. Notes on the flora of Polk County, North Carolina. Cas- tanea 20: 37-57. 1955. [H. japonicus, 44.] Hamacucui, T. Studies in hop. II. Growth and differentiation of the hop plant. (In Japanese; English summary.) Bull. Brewing Sci. 2: 67-120. Henpricks, H. V. Torsion studies in twining plants. II. Bot. Gaz. 75: 282- 297. ee eo of the aérial stem twining in H. Lupulus. | Howarp, G. . E. TarcHELL. Development of resins during the ripen- ing of ce. Bich Inst. Brewing 62: 251-256. 1956.* 202 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 JacopsEN, P. The sex chromosomes in Humulus. Hereditas 43: 357-370. 1957. [H. japonicus & H. Lupulus.| Kruara, H. The sex-chromosomes of Humulus japonicus. Jap. Jour. Genet, 4: 55-63. 1929.* [In staminate plants 27 = 14 + Y.XY, Lepreton, P., & G. Meneret. Eléments de chimiotaxinomie botanique. I. Généralités; notion d’espéce biochimique; cas des flavonoides chez Hu- mulus Lupulus L. Bull. Soc. Bot. France 111: 69-80. 1964. Miter, R. H. Morphology of Humulus Lupulus. 1. Developmental anatomy of the primary root. Am. Jour. Bot. 45: 418-431. 1958; II. Secondary growth in the root and seedling vascularization. Ibid. 46: 269-277. 1959. Muapentseva, M. S. Embryonal study of common hop. (In Russian.) Dokl. Moskov. Sel’skokh. Akad. Timiriazeva 59: 171-177. 1960.* Morecr, T. Some observations on sex chromosomes and sex expression in Humulus seis Sci. Rep. Tohoku Univ. Biol. 31: 7-16. s Muroca, T. Process of flower bud differentiation in the hop Sige (In Japa- nese; English Aaah Sci. Bull. Fac. Agr. Kyushu Univ. 13: 20-25. 1951. Myrick, H. The hop. viii + 300 pp. New York. 1914.* [Economic botany.] Neve, R. A. The place of polyploidy in hop breeding methods. Wye Coll. Dep. Hop Res. Annual Rep. 1955: 124-129. 1956.* Sex chromosomes in the hop Humulus Lupulus. Nature 181: 1084, 1085. 1958. [Heteromorphic chromosome pair in meiotic preparations from staminate plants. | Ono, T. Studies in hop. I. Chromosomes of common hop and its relatives. (In Japanese; English summary.) Bull. Brewing Sci. 2: 1-65. 1955. [See also HAMAGUCHI (1955).] . Review of research. Pp. 1 : T. Ono, The wild hop native to Japan. iv + 110 rei 10 pls. ae ite ite ecology, and cytology of H. Lupulus in Jap —— & H. Svuzukt, ‘Grteloenal studies. Ibid. 71-110. 5 pls. [H. Lupulus.] & S. Wapa. Ecology and morphology. Jbid. 12-70. 5 pls. [ H. Lu- pulus. Runner, D. K., & F. H. Smirn. The structure and development of the storage root of Humulus Lupulus L. (Abstr.) Proc. Oregon Acad. Sci. 2: 80. 1948- 51 [1951]. Satie, A. J., G. J. Jann, & M. Orpanixk. Lupulon—an antibiotic extracted from the strobiles of Humulus Lupulus. Proc. Soc. Exper. Biol. Med. 70: 409-411. 1949. [Antibiotic action on im vitro Gram-positive bacteria. | SALMON, E, S., & H. WorMatp. Humulus americanus Nuttall. Jour. Bot. 53: 132-135. 1915. |H. americanus and H. Lupulus considered distinct. | SCHAFFNER, J. H. Sex reversal in the Japanese hop. Bull. Torrey Bot. Club 50: 73-79. 1 pl. 1923. [Short days induce intersex flowers and carpellate and staminate plants to develop staminate and carpellate flowers, re- spectively. ] ScHerY, R. W. Plants for man. viii + 564 pp. Englewood Cliffs, N.J. 1952. [ Beer, 521-524.] SkovsTeD, A. Some colchicine experiments with hops (Humulus). Hereditas 39: 156-160. sg [Induced polyploidy in H. Lupulus and H. japonicus.] TouRNOIs, - Pp s sur la sexualité du houblon. Ann. Sci. Nat. Bot. IX. 19: 49-191. 5 pls. “sta, [Floral morphology, monoecious plants, carpellate to nei reversions and vice versa, fertilization, embryology, morphology 1970] MILLER, GENERA OF CANNABACEAE 203 of the fruiting spike, and Cannabis sativa * H. Lupulus and H. Lupulus x H. japonicus crosses (fruits matured but contained aborted embryos). | WEsTon, E. W. Changes in sex in the hop caused by plant growth substances. Nature 188: 81, 82. 1960. [H. Lupulus. | WEtTTSsTEIN, R. Fakultative Parthenogenesis beim Hopfen (Humulus Lupulus). Flora. 118/119: 600-604. 1925. [Occurrence of occasional somatic parthe- nogenesis. | Zus, L. Studies on growth and development of hop (Humulus Lupulus L.). (In Polish; English summary.) Hodowla Rosl. 2: 697-724. 1958.* ARNOLD ARBORETUM HARVARD UNIVERSITY 204 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 NOTES RELATING TO THE FLORAS OF NORFOLK AND LORD HOWE ISLANDS, I. P. S. GREEN In OctoseR AND NoveMBER 1963 I had the good fortune to visit first Norfolk Island and then Lord Howe Island in the southwestern Pacific, and to spend about a week on each, making botanical collections and observations upon the flora. The visits were made possible through funds from the Arnold Arboretum (where, at the time, I had the honor of being a staff member) and from the National Science Foundation (Grant No. GB1545), and grateful acknowledgement is made for their generous support. The immediate reason for the visits to these islands was to see certain members of the family Oleaceae, but the opportunity was taken to examine the general flora. Like most oceanic islands, Norfolk and Lord Howe are noted for the high degree of endemism in both their flora and fauna. In fact, they have been referred to as “two of the most interesting oceanic islands in the world” (Good, 1964, p. 144) and as such are of particular interest to biogeographers. But biogeography must be based on accurate taxonomy and when naming the collections made on the islands it soon became clear that the existing floristic lists are quite out of date (Laing, 1915; Oliver, 1917) and that a considerable amount of work upon these floras still re- mains to be done, especially when the plants are considered in relation to those of the various adjacent land-masses and islands. Because of this I am gradually working over the whole vascular flora of the two islands, examining it carefully, comparing specimens with related material from Australia, New Zealand, New Caledonia, and elsewhere, and reassessing their taxonomy. Due to many other commitments this work has not progressed as rapidly as I had hoped but I have been urged to publish at least the conclusions so far reached. LEGUMINOSAE Sophora howinsula (W. R. B. Oliver) P. S. Green, comb. nov. Edwardsia chrysophylla sensu C. Moore, Sketch of the Vegetation of Lord owe Island 3.1869 & in E. S. Hill, Lord Howe Island: Official Visit by the Water Police Magistrate and the Director of the Botanic Garden, Syd- ney. 28. 1870, non Salisb. apt oi tetraptera sensu F. Muell. Fragm. Phytogr. Austral. 7: 26. 1869 & . 1875; sensu C. Moore & Betche, Handb. Fl. New S. Wales 519. 1893; sensu Tate in Fletcher, Macleay Mem. Vol., Linn. Soc. New S. Wales 217. 1893; sensu Hemsley, Ann. Bot. 10: 235. 1896: sensu Maiden, Proc. Linn. Soc. New S. Wales 23: 128. 1898; non J. Miller. 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 205 S. tetraptera J. Mill. var. howinsula W. R. B. Oliver, Trans. & Proc. New Zealand Inst. 49: 139. 1917. Type: Lord Howe Island, Transit Hill, 1917, W. R. B. Oliver (? weELt, not seen). S. tetraptera J. Mill. subsp. howinsula (W. R. B. Oliver) Yakovlev, Proc. Leningrad Chem.-Pharm. Inst. 21 (Voprosy Farmakognozii 4): 57. 19 Yakovlev has recently published a synopsis of the genus Sophora (Ya- kovlev, 1967) in which he includes the Lord Howe representative as a sub- species of the New Zealand S. tetraptera, in sect. Epwarpsia (Salisb.) Taub." He recognizes ten species in this section, together exhibiting a most interesting distribution: S. denudata Bory from Réunion; S. chrysophylla (Salisb.) Seem. from Hawaii; S. macrocarpa Sm. from Chile; S. fernan- deziana (Phil.) Skottsb., S. masafuerana (Phil.) Skottsb., and S. reediana (Phil.) Yakovlev from the Juan Fernandez Islands; S. toromiro (Phil.) Skottsb. from Easter Island; and S. microphylla Ait. and S. tetraptera J. Mill. from New Zealand, except for subsp. macnabiana (R. Graham) Yakolev, a Chilean plant which he includes under S. microphylla, and the Lord Howe Island plant which he places as a subspecies of S. tetraptera. Careful examination of the Lord Howe material at Kew leads me to believe that specific recognition for this plant is justified. Certainly the number and type of differences from the New Zealand S. tetraptera are of a comparable order with those from S. microphylla and S. chryso- phylla. On the other hand the other subspecies which Yakovlev recog- nizes in this section, subsp. macnabiana, seems closer to the New Zealand material of S. microphylla, under which he places it, than do the Lord Howe Island specimens to S. tetraptera. The differences noted are as follows. Leaves with fewer leaflets (9—) 13— 17(—19) as opposed to (17—)19-33(-39) in S. tetraptera, slightly larger (1.3-)1.7-2.5(-3.5) by (0.6—-)0.8-1(-1.3) cm. in contrast to (0.6—)1- 2(-3) by (0.3—)0.5—0.7(—1) cm., and glabrous above even in the slightly sunken midrib. Flowers smaller 2.8—-3.5 cm. long (4-5 cm. long in S. tetraptera); calyx smaller 0.7—-0.8 cm. long, not 1—1.5 cm., lightly, not densely hairy; standard a little darter (2 cm. long) than the keel (2.5 cm. long) in contrast to 3-3.5 cm. and 4-5 cm. in S, tetraptera. Fruit with the surface of each segment minutely warty or uneven in contrast to smooth in the New Zealand plant. Lord Howe Island. Without locality, ex Phytologic Museum of Victoria (x), Milne (x), C. Moore 10 (kK) & Oct. 1936, McComish 19 (x); Transit Hill, low dryish forest, 12 Nov. 1963, Green 1616 (A, K); Transit Hill, in dense shrubbery, common, approx. 125 m. alt., 17 July 1965, van Balgooy 1014 (xk). ? Yakovlev, like other authors, attributes this section to Seeman (FI. Vit. 66. 1865). However, although Seeman sunk the genus Edwardsia Salisb. under Sophora L. he ; i (in Hook. fil. Fl. Brit. Ind. 2: 251. 1878, where in most cases s he carefully “designates “Subgen.” and “Sect.”) uses an unspecified infrageneric category denoted by “§”. 206 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 MALVACEAE Sida zahlbruckneri Rech. In 1907 Rechinger published the name Sida zahlbruckneri, attributing the species to Lord Howe Island and basing it upon a MacGillivray collec- tion at Vienna. There has been no other record of this species and, through the kindness of the Director of the Naturhistorisches Museum, to whom grateful thanks are expressed, I have been enabled to examine the holo- type on loan. It is clear that it has unfortunately been mislabelled at some time and that it was collected on the Isle of Pines, New Caledonia, and not Lord Howe Island. The particular MacGillivray collections of which it forms a part, were made, along with others by W. G. Milne, on the voyage of H.M.S. Herald, 1853-4. They were sent by MacGillivray and Milne to Kew and examination of the herbarium there reveals nothing resembling this species from Lord Howe Island, but both collectors made specimens of an identical Sida from the Isle of Pines, the next island visited by the Herald after leaving Lord Howe (Observatory Island, Isle of Pines, banks by the shore, Sept. 1853, MacGillivray 824, & Ob- servatory Island, Isle of Pines, on beach, abundant, Oct. 1853, Milne 85). On comparing the specimens it is clear that the type of S. zahlbruckneri agrees exactly with these collections and must have been part of them before becoming separated and mislabelled. In 1892 E. G. Baker in his “Synopsis of Malveae” described Sida num- mularia, based on the Milne collection cited above, and, therefore, not only is S. zahlbruckneri no longer to be classed as one of the Lord Howe Island endemics, but in fact falls into synonymy. Sida nummularia Bak. f. Jour. Bot. 30: 290. 1892; Daniker, Viert. Na- turf. Ges. Ziirich 78 (Beibl. 19): 261. 1933; Guillaumin, Fl. Nouv.- Caléd. 209, 1948. Type: Isle of Pines, New Caledonia, Milne 85 (x). S. zahlbruckneri Rech, Repert. Sp. Nov. 4: 1907. Type: “Lord Howe Island” [Isle of Pines, New Caledonia], ae (w). Sida nummularia, which is endemic to New Caledonia, is very similar to S. parvifolia DC. (Prodr. 1: 461. 1824) based on a specimen from Réunion and widespread on the Mascarene Islands. In fact they may not be specifically distinct; however, although the 34 collections I have examined from Mauritius, the Seychelles, and the Cosmoledo and Chagos Archipelagoes (I have not actually seen a specimen from Réunion) show considerable variation, the 11 New Caledonian collections seen are very consistent in leaf shape and size. Moreover, in each case where ripe fruit was examined, the seed of the Pacific plant was found to be puberu- lent at the apex, near the slit in the mericarp wall, but glabrous in the plants from the western Indian Ocean. The exact status of these two species (as well as S. discolor Bak. f., non Hook., described from “Timor” [Sumba]) must await careful biosystematic work on the group. Van Borssum Waalkes (in Blumea 14: 192, 193. 1966) equates Sida dis- 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 207 color Bak. f. with S. parvifolia DC. and says that “it has been found on the Pacific Islands.” No material is cited but it seems possible that these New Caledonian collections, together with others from Fiji, Tonga, and Samoa are involved. The specimens from these three latter groups of islands belong to S. samoensis Rech. (S. microphylla sensu Benth. in Hook. London Jour. Bot. 2: 211. 1843, & sensu Seem., Fl. Vitiensis 15. 1865, non Cav.) which is superficially very similar and also recorded from coral beaches, but with mericarps which are clearly biaristate, as pointed out by Daniker (Viert. Naturf. Ges. Ziirich 78 (Beibl. 19): 261. 1933). OXALIDACEAE Oxalis corniculata L. = Pl. 435. 1753; Oliver, Trans. & Proc. New Zealand Inst. 49: 140. 1917. Var. corniculata. O. reptans Sol. ex Forst. f. Prodr. 90. 1786, nom. nud.; Endlicher, Prodr. Fl. Norf. 72. 1833, nom. nud. O. corniculata L. var. reptans Laing, Trans. & Proc. New Zealand Inst. 47: 28. 1915. Type: Norfolk Island, on rocks, particularly near sea shore, 1912, Laing (CANTY) Norfolk Island. Anson Bay, 6 Jan. 1939, McComish 122 (x). Lord Howe Island. Near cultivated ground, Sept. 1853, MacGillivray 719 (x). Southeastern slopes of Malabar, dry open rocky and grassy patches in low forest, 11 Nov. 1963, Green 1559 (A, ia behind beach, Ned’s Beach, 5 m. alt., 20 July 1965, van Balgooy 1054 (kK Var. microphylla Hook. f. Fl. Nov.-Zeland. 1: 42. 1852 & Fl. Tasm. 1: 59. 1855; Young, Watsonia 4: 56. 1958; Allan, Fl. N. Zeal. 1: 239. 1961; Hoogland in Turner ef al. Austral. Conservation Foundation Spec. Publ. 1 (The Conservation of Norfolk Island) : 34. 1968. Types: New Zealand, Whaapu, Bay of Islands, 1833, R. Cunningham 234 (A, Cunningham 587) (K); Tasmania, A. Cunningham 1837/94 & Glen Leith, 14 Sept. 1840, J. D. Hooker (x). Norfolk Island. Near Kingston, alongside creek, 28 Jan. 1939, McComish 122A (kK). Kingston, ruins of old prison, 30 Oct. 1963, Green 1434 (a). Near Rocky Point, earthy banks and ea on sea cliffs, 30 Oct. 1963, Green 1461 (A). Phillip Is., eroded and rabbit infested hillside, 31 Oct. 1963, Green 1502 (a). The world-wide weed var. corniculata has been introduced to both Lord Howe and Norfolk Islands, but var. microphylla is most probably native on the latter. Captain Cook, when describing the discovery of the island in 1774, wrote that he found “cabbage-palm, wood-sorrel, sow thistle and samphire abounding in some places on the shores’ (“A voyage towards the South Pole and round the world” 2: 148. 1777 & in “The three voy- ages of Captain James Cook round the world” 4: 137. 1821). It is probably this species that he refers to as wood-sorrel (although it is not 208 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the Oxalis he might have known in Britain). In reference to the island, he also stated (op. cit. 147 & 136) that “we found it uninhabited, and were undoubtedly the first that ever set foot on it.” Yet, whether his party were, in fact, the first to land on the island may be open to ques- tion, even if it was uninhabited when they discovered it. Sow thistle, Sonchus oleraceus, now a widespread weed, is generally considered na- tive in the European, W. Asian and Mediterranean regions and must, therefore, have been introduced to Norfolk Island before Capt. Cook’s visit. Governor King, then Lieutenant (in Hunter, “‘A Historical Jour- nal of the Transactions at Port Jackson and Norfolk Island” 306. 1793), stated that on 5th April, 1788, in “a fine valley . . . . a number of plan- tain or banana tree” were found and later (p. 396) that ‘those planted .... have already yielded good fruit.” Although Endlicher (/. c. : 35) thought that Musa was indigenous he was probably mistaken. The island was previously probably inhabited, with the bananas planted and the sow thistle introduced (and the wood-sorrel?). (See also Thorpe in Jour. Polynesian Soc. 150: 123-126. 1929 and McCarthy, op. cit. 172: 267-270. 1934, for further evidence of previous occupation of the island.) Although var. microphylla is stated by Young (1958, p. 56) to be na- tive of the “mountainous parts of New Zealand and Tasmania” the very sparse field notes on the material in the herbarium at Kew give little or no indication that this may be its habitat. Allan (1961, p. 239) says that the distribution is uncertain and that the variety occurs in North Auck- land and I have certainly seen both varieties almost at sea-level on Nor- folk Island. Although Laing (1915, p. 28) gives the first valid description to go with Solander’s epithet reptans it is clear that he bases it on the plant from Norfolk Island which, in a footnote, he says may be endemic. Be- fore examining his type material, kindly sent on loan, I had expected it to be the small leaved var. microphylla; but I have been surprised to find that it proves to be var. corniculata although a semiprostrate, small- leaved form. Similarly, the specimen at the British Museum of the plant collected in New Zealand by Banks and Solander is a prostrate form of var. corniculata. Var. microphylla is recorded from both New Zealand and Tasmania and, from the material I have seen, is somewhat variable in pubescence. However, it seems to maintain its decumbent habit, very small leaves (less than 5 mm. long), short capsule (less that 12 mm. long), and small seeds (1 mm. or less in length). I have examined ma- terial of Oxalis corniculata from most parts of the world but have found var. microphylla from Norfolk Island, New Zealand, and Tasmania only, except for some specimens from Britain, where it is recorded as “a fre- quent garden escape’ (Young 1958). RUTACEAE Acronychia J. R. & G. Forst. A critical examination of the Norfolk and Lord Howe Island plants 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 209 that have passed under the names Acronychia baueri and A, endlicheri, together with the related plants from the mainland of Australia and islands in the Pacific, has led to a reassessment of the whole group. It is con- cluded that two species are involved, one consisting of three ital Soa subspecies and the other endemic to New Caledonia. They ma identified by the key below. It is possible that the plant described as Acronychia acidula F. Muell. (Fragm. Phytogr. Austral. 4: 154. 1864) is another subspecies, but no flowering material has been seen. It occurs in tropical parts of Queens- land (type locality: Seaview Range, Rockingham Bay) and has much larger leaves and fruit. Similarly, A. vestita F. Muell. (7. c. 155. 1864), of which I have seen only fruiting material may be another subspecies. It was also described from the Rockingham Bay area and the two taxa were combined by F. M. Bailey (Queensland Fl. 1: 210. 1899), but the isotypes and other material I have examined at Kew differ so much in indumentum that they may prove to be distinct. Key To ACRONYCHIA SIMPLICIFOLIA AND A. LEIOCARPA 1. Leaves usually more than 8 cm, long, (4.5—)7-13(-16) cm. se petioles (1-)2-4(-5) cm. long; calyx 1.5-2.5 cm. long; ovary and t hairy or velutinous; fruit apex at least slightly pointed. .......... .: setitesoles. 2. Apex of leaf rounded or obtuse, retuse or subretuse; fruit apex slightly beaked or distinctly pointed. 3. Petiole glabrous or minutely pubescent when young, eventually gla- brate; leaves ovate-elliptic; fruit only slightly hairy and slightly e subsp. simplicifolia. 3. Petiole pubescent, sometimes glabrous with age; leaves elliptic; fruit subvelutinous, distinctly pointed. .............. subsp. eo-scotica. 2. Apex of leaf acute, sometimes rounded subacute; fruit apex slightly pointed, more or less pinched; petiole minutely puberule nt when young. REG AD Rei | & Ub USN gg fies rE ea subsp. petiolaris. 1M eens less than 8 cm. long, (2—)3—7(—8) cm. long; petioles (0.8—)1-1.7(-2) m. long; calyx 1-1.5 mm. long; ovary and fruit glabrous; fruit apex not ied nll yy 5 oe oR wk e Ae Be A, leiocarpa. Acronychia simplicifolia (Endl.) McGillivray & Green, comb. nov. Vepris simplicifolia Endl. Prodr. Fl. Norf. 89. 1833. Type: Norfolk Island, . 1804, Bauer (w, not seen Acronychia simplicifolia (Endl.) Steud. Nom. ed. 2. 1: 21. 1840, nomen in synon.; Hoogland in Turner e¢ al. Austral. Conservation oe Spec. Publ. 1 (The Conservation of Norfolk Is.): 34. 1968, no Subsp. simplicifolia. FicureE la. Acronychia endlicheri Schott, Rutaceae. Fragm. Bot. 3. ¢. 2. 1834, nom. illegit.; KR. ragm. Phytogr. Austral. 9: 103. 1875; Moore & Betche, Handb. Fl. New S. Wales 518. Tas: Tate in Fletcher, Macias Mem. Vol., Linn. 210 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Soc. New S. Wales, 217. 1893; Maiden, Proc. Linn. Soc. New S. Wales 28: 699. 1903; Laing, Trans. & Proc. New Zealand Inst. 47: 29. 1915. Type as for Vepris simplicifolia Endl. A. baueri Schott, op. cit. 5. t. 3. 1834; Benth. Fi. Austral. 1: 366. 1863; F. Muell. Fragm. Phytogr. Austral. 9: 77 & 103. 1875; F. M. Bailey, Syn. Queensl. Fl. 54. 1883; Moore & Betche, Handb. Fl. New S. Wales 49. 1893; Tate in Fletcher, Macleay Mem. Vol., Linn. Soc. New S. Wales 217, 1893; Hemsl. Ann. Bot. 10: 233. 1896; Maiden, Proc. Linn. Soc. New S. Wales 23: 124. 1898; F. M. Bailey, Queensl. Fl. 208. 1899 & Compreh. Cat. Queensl. PI. 81. 1913; Oliver, Trans. & Proc. New Zealand Inst. 49: 141. 1917; Francis, Austral. Rain Forest Trees 167. 1929 & ed. 2. 191. 1951; Anderson, Trees New S. Wales ed. 2. 231. 1947 & ed. 4. 210. 1968; Beadle et al. Handb. Vasc. Pl. Sydney xi a 1963. Type: Locality unknown, ? Norfolk Island, Bauer (? w, not s A. ovata Endl. ex. Heynh. Nom. 2: e 46, m nud. A, hillii F. Muell. Fragm. Phytogr. Austral. C %, "1858, Type: Queensland, Moreton Bay, Hill & F. Mueller (MEL, not seen). Jambolifera endlicheri (Schott) Kuntze, Rev. Gri. Fi. 162. 1891, J. baueri (Schott) Kuntze, Joc. cit. 1891. Bauerella australiana Borzi, Bol. Orto Bot. Palermo 1: 155. 1898, nom. illegit. Type: as for Acronychia baueri Schott. Acronychia baueri Schott f. majoriflora Domin, Bibl. Bot. 22 (89) (Beitr. Fl. Pflanzengeogr. pita 848): 294. 1927. Type: Queensland, Brisbane River, Dietrich (not seen). Bauerella baueri chat) Daniker, Viert. Naturf. Ges. Ziirich 77 (Beibl. 19): 202. 1932, quoad ty Norfolk Island. Apr. 1835, Backhouse 616 (K); margins of woods, June 1830, A. Cunningham 29 & 148 (K); above Red Stone, 25 Aug. 1968 & 8 Mar. 1969, Owen Evans (kK). Lord Howe Island. Fullager (kK); 1869, C. Moore 17, 32 & 48(K); low dry forest, ani lower slope of Malabar, i Nov. 1963, Green 1574 (aA, K). Queensland. North Kennedy district: Strathdickie, Michael 1356 (GH); Rock- ingham Bay, Dallachy (x). South Kennedy district: Forest reserve 652 Cauley, approx. 30 miles S. of Proserpine, 22 May 1967, Mackay 67/144 (x); Dal- rymple Heights, rain forest, Aug—Oct. 1947, Clemens (GH, K). Port Curtis dis- trict: Middle Percy Island, Jan. 1906, Tryon (A); Northumberland Islands, R. Brown “5328” (kK). Wide Bay ree Mt. Perry, Aug. 1912, Boorman in NSW 16272 (kK); Nikenbah, Tryon (A); Kin Kin, Jan. 1916, White (x). Moreton district: Blackall Range, Dec. tae "White (A); Forest reserve, Yarraman, 1400 ft. alt., Aug. 1944, pes (A); Mt. Glorious, ridge forest, 2000 ft. it. Jan. 1945, Clemens (GH, K), & common in rain forest, 28 June 1937, White 11078 (Kk); Mt. Mistake, 3000 ft. alt., 2 Feb. 1944, Clemens 43591 (a); Bris- bane R., A. Cunn ningham (cH), 1829, Fraser 157, 183 & 213 (K) ee & Aug. 1855, F, Mueller (xk) & dark, dry woods, July, « Cunningham 3 ; Tam- bourine Mt., Oct. 1909, Simmonds (A); Moreton Bay, Oct. 1824, A. 7 eon ham 18 (k New South Wales. Upper Eungella, Tweed River, June 1923, White 13268 (A); Tweed Heads, Sept. 1910, Simmonds (a); Richmond R., C. Moore (x); Lismore, Aug. 1891, Maiden (a); Carrai Carrai State Forest, 30 miles approx. W. of Kempsey, frequent on limestone ridge, 810 m. alt., 11 June 1958, Con- 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 211 stable in NSW 46231 (kK); Port Macquarie, May 1819, A. Cunningham 49 (x); W. of Coneac (Manning R., approx. 18 miles NW. of Gloucester), small rain forest pocket by creek, 15 Oct. 1953, Johnson in NSW 26274 (xk): 16 miles ENE. of Singleton, patch of broad-leafed forest, 27 Mar. 1960, Storey 7203 (xk); Ash Islands, Newcastle, 1802-5, Brown “5329” (x); Wyong, Dec. 1898, Boorman (a); Illawarra, Nov. 18 A. Cunningham 169 & Aug. 1824, A. Cunningham 4 (Kk); 1 mile W. of Fe Tilba, very common in a dry t of rainforest on rocky hillside, 9 Sept. 1960, Johnson & Constable in NSW 52308 (K); slopes of Mt. Drom edary, Central Tilba (approx. SW. of Narooma), occasional in rain forest, 90 m. alt., 12 Sept. 1953, Constable in NSW 26537 (kK); Milton, on igneous rock, 5 Dec. 1905, Cambage 1420 in NSW 26312 (A, K). A careful examination of the Norfolk and Lord Howe Island plants shows that they do not differ taxonomically from that of the eastern main- land of Australia, from Queensland and New South Wales. When taken in the context of the Tribe Toddalieae the characters used by Borzi to define the genus Bauerella are insufficiently distinct to justify generic separation. Unfortunately Borzi failed to take up the epithet baueri, although citing it in synonymy, and thus rendered his binomial illegitimate. The correct combination B. baueri is sometimes attributed to Engler (in Engler & Prantl, Nat. Pflanzenfam. Nachtr, II-III. 4: 35. 1900) but it is not actually made by him at this reference. The earliest valid use that has been traced is that of Daniker referred to above. My friend and colleague D. J. McGillivray quite independently, and in connection with his work on the plants of New South Wales, came to the realization that the combination Acronychia simplicifolia had never been validly made and so I have pleasure in sharing with him in the com- bination above. Subsp. neo-scotica P. S. Green, subsp. nov., a subsp. simplicifolia petiolis pubescentibus, foliis constanter ellipticis et fructibus subvelutinis mani- feste acutis differt. Ficure 1b. Acronychia eriocar pa ry ex. Guillaum. Not. Syst. Paris 2: 98. 1911, nom. s. Col. Marseille II. 9 (Cat. Pl. Phan. Nouv. -Caléd. 1: 39): 111. 1911, nom. ayes A. baueri sensu Guillaum. JI. cc. (1911), non Schot Bauerella australiana sensu Guillaum. Il. cc. 40 & Nie 1911 & Jour. Arnold Arb. 12: 235. 1931, non Borzi. B. baueri sensu Daniker in Viert. Naturf. Ges. Ziirich 77(Beibl. 19): 202. 1932, pro parte, excl. spec. typ.; sensu Guillaum. Fl. Nouv.-Caléd. 168 1948. New Caledonia. Montaignes de Wailiou, 1861-67, Vieillard 2704 (K); Nouméa, 1868-70, Balansa 376 (kK); Nouméa (Ouen Toro), 3 Aug. 1930, Franc 2454 (a, K); dry stony slopes on Ouen Toro, 0-50 m. alt., 26 Feb. 1955, McKee 2163 (x); Anse Vata, low calcareous hill, 10 July 1955, ‘McKee 2731 (A); Mt. Dore, 1855-60, Viel 859 (kK); “auf den Hiigeln bei Yaouhé,” 17 Oct. 1902, Schlech- ter 15071 (xk); Prony, Franc (xk); Port-Boisé, 1861-67, Deplanche 511 (k, holotype) & Deplanche 510 (a). New Hebrides. Eromanga, Cook’s Bay, 22 July 1896, A. Morrison (x). Anei- 212 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 tyum: Anelgauhat, 5 June 1896, A. Morrison (x); Anelgauhat Bay, rain forest, common, 305 m. alt., 14 Feb. 1929, Kajewski 761 (A, K). This subspecies is most easily distinguished from the type by its fruit, which is fine velvety and relatively long pointed and rounded in section below the apex. In subsp. simplicifolium, as well as the fruit being only scattered hairy or glabrate, it is much less prominently pointed and in transverse section below the apex shows four distinct indentations (Fic- ures la & b). Also the petiole in this subspecies is more pubescent than in the others. The epithet chosen alludes to the fact that it occurs in both New Caledonia and the New Hebrides. a b c IGURE 1. Acronychia fruits, all ss ae 9 ' simplicifolia subsp. se dong ag (from Johnson & Constable NSW 5 2308) ; subsp. eéo-scotica jewski 761); c, subsp. petiolaris (from ae 114): d, A. leiocarpa (ivom McKee 211 15). Subsp. petiolaris (A. Gray) P. S. Green, comb. nov. FIGURE Ic. Acronychia petiolaris A. Gray, Bot. U.S. Expl. Exped. 1: 335. 1854 & t. 33A. 1857; Seem. Fl. Vitiensis 31. 1865; Gibbs, Jour. Linn. Soc. Bot. 39: 142. ati Por Jour. Arnold Arb. 32: 243. 1951; Parham, Pl. Fiji Is. 164. : Fiji, Mathuata, U.S. Exploring Expedition a holotype). ppecetiae: proscar (A. Gray) Kuntze, Rev. Gen. Pl. 102. 1891. Fiji. VaANuA Levu. Mbua: southern portion of Seatovo Range, 100-350 m. alt., dry forest, 20 Apr—2 May 1934, Smith 1702 (a, k); Bua, forest, not com- mon, Sept. 1878, Horne 1114 (GH, kK). Mathuata: without locality, U.S. Ex- ploring Expedition (GH, holotype of subsp. petiolaris); Dreketi, forest, Parham, Qoro & Kuruvoli in Dept. Agric. 13451 (kK); Dreketi district, near village Ba- sakalave, 32 km. W. of Seaqaqa, 40 m. alt., lateritic hill, rather dense forest, 26 Mar. 1964, Stauffer & Kuruvoli 5844 (x); Wainunu-Ndreketi divide, 200- 300 m. alt., dense forest, 17 May 1934, Smith 1841 (a, K); Seaqaqa, coffee plan- tation, 27 Feb. 1964, Parham & Kuruvoli in Dept. Agric. 13908 (kK); Seanggangga Plateau, in drainage of Korovuli R., vicinity of Natua, 100-200 m. alt., patches of forest in open rolling country, 25 Nov.—8 Dec _ 1947, Smith 6711 & ‘6728 (A, kK); southern base of Mathuata Range, N. of Natua, 100-250 m. alt., dense forest, 1-4 Dec. 1947, Smith 6779 (a, K); southern slopes of Mt. Numbuiloa, E. of Lambasa, 100-350 m. alt., open forest, 27 Oct-10 Nov. 1947, Smith 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 213 6407 (A, K); Lambasa, approx. 200 ft. alt., 17 Dec. 1922, Greenwood 566 (K); Nasuaca, forest, 22 July 1963, Qoro & Kuruvoli in Dept. Agric. 13486 (RK). Vitt Levu. Mba: Nandarivatu, 2600 ft. alt., 11 Mar. 1927, Mead in SFN 1987 (kK) & 3000 ft. alt., in copse, June 1927, Tothill 54 (x); Vicinity of Nan- darivatu, 850 m. alt., 22 Nov. 1927, Gillespie 3984 (cH), & 900 m. alt., 14 Dec. 1927, Gillespie 4311 (K); Sovutawambu, 5 miles from Nandarivatu, 750-800 m. alt., 27 Feb—4 Mar. 1941, Degener 14652 (a, K); southern slopes of Mt. Ndelainathovu, on the escarpment W. of Nandarivatu, 870-970 m. alt., dense forest, 26 June 1947, Smith 4934 (a, K); hills between Nandala & Nukunuku Creeks, along trail from Nandarivatu toward Lewa, 750-850 m. alt., thickets on fairly open slopes, 22 Sept. 1947, Smith 6150 (a, K). KANDAvu: understory in rainforest, Watkins 750 (kK). Without locality: Horne 599 (cu, «). The Fijian plant resembles subsp. meo-scotica in its fruit, which is not quite so pointed and is less completely rounded in section below the apex. But the leaf shape, however, is consistently different with an acute apex, rarely only subacute, never retuse or subretuse, and usually with more primary veins on each side of the midrib: (7—)9-10(-12) per side in contrast to (6—)7—9(—11) in the other subspecies. Acronychia leiocarpa P. S. Green, sp. nov., a A. simplicifolia foliis mi- noribus, (2~)3—7(—8) cm. longis, ovariis glabris, et praesertim fructi- bus minoribus glabris differt. Ficure 1d. A. baueri sensu Pancher in Sebert & Pancher, Not. Bois Nouv. Caléd. 245. [1874], non Schott. ; A. ligustroides Panch. ex Guillaum. Not. Syst. Paris 2: 98. 1911, nom. in synon. & Ann. Mus. Col. Marseilles 19 (Cat. Pl. Phan. Nouv.Caléd. 1: 40): 112. 1911, nom. nud. A, ovalifolia Panch. ex Guillaum. JJ. cc. 1911, nom. nud. Small tree to 5 m. tall, branchlets puberulent when young becoming glabrous. Leaves simple; petioles (0.8—)1-1.7(-2) cm. long, minutely puberulous; lamina elliptic or slightly elliptic-ovate, (2-)3-7(-8) cm. long, by (1.5—)2-3(-3.5) cm. broad, glabrous; margin entire; apex rounded, retuse; base obtuse and more or less rounded or acute; venation finely and clearly raised reticulate above and below, 7—9(—10) primary nerves on each side of the midrib. Inflorescence axillary, condensed panic- uloid, (0.5—)1—2(-2.5) cm. long, (1-)3-15-flowered, puberulent; flow- ers hermaphrodite, green or yellowish green; pedicels 0-1 mm. long. Calyx minutely tomentose externally, lobes 4, 1-15 mm. long, triangular ovate. Corolla valvate, minutely tomentose externally, lobes 4, 2.5~3.5 mm. long. Stamens 8, antesepalous 1.5—2 mm. long, antepetalous 1—1.5 mm. long; filaments flat, ciliate 0.7 mm. broad, abruptly tapered at the apex; anthers 0.5 mm. long. Ovary glabrous, 4-loculate, with several axile ovules; style arising from a slight depression on the top, 0.8 mm. long with somewhat flattened terminal stigmatic area 0.7 mm. in diameter. Fruit glabrous, rounded but more or less cuboid, apex not pointed, with 4 indentations (Ficure 1d), red (fide McKee 2115), 0.6-1 cm. long and 214 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 broad, endocarp hard boney, ? indehiscent, with 2-3 black seeds per loculus. ew Caledonia. Wailou, 1861-67, Vieillard 200 (x); Nouméa, 1868-70, Balansa 375 & 375a (K); Bois de Port-Despointes (Nouméa), “bois secs des collines littorales, schistes nummulitiques,” 14 Nov. 1942, Virot 810 (A), 812 (a) & 822(a); foot of Ouen Toro, 10 m. alt., 13 Feb. 1955, McKee 2115 (A, K); NE. slope of Ouen Toro, Nouméa, dry scrub, 26 Sept. 1963, Green 1211 (A, isotype; K, holotype); Isle des Pins, “ad rivulum ad pedem Pic Nga,” 100 m. alt., 18 July 1965, Bernardi 10067 (kK); without locality, 1862, Pancher s.n. & Deplanche 491 & 510 (K); “pieds de Ma . . . ,” 1900-1910, Le Rat 2273 (a). Acronychia leiocarpa has been confused with A. simplicifolia. Both species occur together, e.g. McKee 2115 and 2163 were both collected on Quen Toro near Nouméa, and it would be interesting to investigate their biology. However, A. leiocarpa is immediately separable by characters of the leaf, its smaller size and finer reticulation, and, above all, by the glabrous ovary and subsequent fruit. Other characters for separation are found in the smaller size of the flower parts, the relatively broad staminal filaments, which are usually more abruptly contracted below the anther, and the smaller, unpointed fruit (FicurE 1D). It appears that at one time Pancher recognized that this species was distinct and proposed the names Acronychia ligustroides and A. ovalifolia which he attached to some of his herbarium specimens, but in Sebert & Pancher “Notice sur les Bois de la Nouvelle Calédonie” 245 [1874], although describing the characteristic fruit, he equated this species with the Australian A. simplicifolia (as A. baueri). Guillaumin (in Not. Syst. Paris 2: 98. 1911) discussed Pancher’s proposed species and mentioned the glabrous ovary and abruptly narrowed staminal filaments (by in- ference from his description of the filament of A. simplicifolia) but he nevertheless seems to have concluded that only one species was involved. Zanthoxylum L. The plants that have passed under the names Blackburnia pinnata, Zanthoxylum blackburnia, Fagara pinnata, etc., also need reassessment, with the recognition of a new species from New Caledonia. Zanthoxylum pinnatum (J. R. & G. Forst.) W. R. B. Oliver, Trans. & Proc. New Zealand Inst. 49: 140. 6 July 1917; Druce, Rep. Bot. Exch. Club Brit. Is. 4: 653 [? Sept.] 1917; A. C. Smith, Bull. Bishop Mus. 141: 76. 1936; Hoogland in Turner et al. Austral. Conservation Foun- dation Spec. Publ. 1 (The Conservation of Norfolk Island): 34. 1968. Type: Norfolk Island, J. R. & G. Forster (pM, x). Blackburnia pinnata J. R. & G. Forst. Gen. Char. 12. ¢. 6. 1776; Forst. f. Prodr. 10. 1786; DC. Prodr. 2: 83. 1825; Endl. Prodr. Fl. Norfolk. 88. 1833; A. Cunn. in Heward, Hook. Lond. Jour. Bot. 1: 113. 1842. Ptelea pinnata (J. R. & G. Forst.) Linn. f. Suppl. Pl. 126. 1781. 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 215 Samara blackburnia Spreng. Syst. Veg. 1: 441. 1824, nom. illegit. Zanthoxylum blackburnia Benth. Fl. Austr. 1: 363. 1863; Hemsl. Ann. Bot. 10: tee 1896; Burkill, Jour. Linn. Soc. Bot. 35: 30. 1901; Laing, Trans. oc. New Zealand Inst. 47: 29. 1915; nom. illegit. X le blackburnia C. Moore, Sketch of the vegetation of Lord Howe Island 3.1869 & in E. S. Hill, Lord Howe Island: Official Visit by the Water Police Magistrate & the Director of the Botanic Gardens, Sydney 8. 1870; C. Moore & Betche, Handb. Fl. New S. Wales 518. 1893; Tate in Fletcher, Macleay Mem. Vol., Linn. Soc. New S. Wales 215. 1893: Maiden, Proc. Linn. Soc. New S. Wales 28: 699. 1903; nomen. Xanthoxylon howeanum F. Muell. Fragm. Phytogr. Austral. 9: 77. 1875, nom. nud.; Tate in Fletcher, Macleay Mem. Vol., Linn. Soc. New S. Wales 215. 1893, nom. nud. pice pinnata (J. R. & G. Forst.) ein in Engl. & Prantl, Pflanzenfam. III. 4: 1896, & ed. 2 19a: 224. 1931; A. C. the Jou Arnold Arb, 32: me ot goa Bull. Bishop Mus. 2202, 451, F. ellespieana A. m. Jour. Arnold Arb. 32: ie aa Type: Fiji, Viti yak ae Be (A, holotype; K, isotype). Lord Howe Island. “Section,’’ Sept. 1853, Milne (kK); Transit Hill, iy dryish forest, 13 Nov. 1963, P. S. Green 1615 (a, K) & 1617 (A, K); Erskine Valley, on lower part of ascent of Mt. Gower, dry forest on very rocky eaucd. Me Nov. 1963, P. S. Green 1669 (A, K). Without eae locality, C. Moore 9 & 36 (XK). Nopkalic Island. [1773], Forster (BM, isotypes); shady woods, 1830, A. Cunningham 4 & 147 (kK); Apr. 1835, Bickhouse 659 (kK); Nov. 1898, Robinson (BM); May 1904, Robinson (x). ew Hebrides. North central Eromanga, tableland covered with bracken, Saleacte ash, 500 ft. [132 m.] alt., 5 Aug. 1930, L. E. Cockayne 53 Fiji. Viti Levu: Nandronga & Navosa, northern portion of Rairaimatuku Plaieae: between Nandrau and Nanga, alt. 725-825 m., in dense forest, 4-7 Aug. 1947, A. C. Smith 5578 (a, holotype of Fagara pillespieana: K, isotype); sum- mit ridge of Mt. Loma Langa, 21 Nov. 1927, Gillespie 3943 (cH); Naitasiri (?), Colo-i-Suva, alt. 180 m., 19 May 1961, 7. Bola 35 (k). Fulanga: in forest on limestone formation, alt. 0-80 m., 22-26 Feb. 1934, A. C. Smith 1150 (GH, k). Tonga. Vava’u and Lifuka, Aug. & Oct. 1855, Harvey (kK); Vava’u, Dec. 1891, Crosby (k, male & female); Vava’u, in forest near Leimata village, NW. of Neiafu, 19 May 1953, Yuncker 16045 (GH). More material of this species, from throughout its range, is needed for examination. It is apparent that variation occurs between the popu- lations on the island groups, or at least between those on Lord Howe and Norfolk Islands, and perhaps the New Hebrides, and between these and Fiji and Tonga. The New Caledonian plant which has been treated as this species by some is described below as a separate, but closely related, species. No material has been seen from Tahiti but it is suspected that Zan- thoxylum nadeaudii Drake del Castillo, Ill. Fl. Ins. Mar. Pacif. 130. 1890 & FI. Polynésie Franc. 24. 1893 (Blackburnia pinnata var. tahitensis Nadeaud, Enum. PI. Indig. Ile Tahiti 75. 1873; Fagara nadeaudii (Drake) J. W. Moore, Bull. Bishop Mus. 226: 7. 1963) is part of this species, too. Of the main differentiating characters cited by Drake del Castillo, the 216 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 less deeply divided calyx could certainly apply to the specimens seen from Fiji and Tonga, and the leaf shape probably falls into the range of variation for this species. For each of the specimens cited above, and those which had been de- termined as this species from New Caledonia, I scored the lengths of the leaf rachis and leaflet, the texture and number of leaflets per leaf, the bud shape, the petal, filament and anther length, and the fruit di- mensions. After analysis of the results I was tempted to separate three subspecies together with the new species from New Caledonia, but in view of the range of variation on each of the other islands additional and better material is required to assess more accurately the following trends and differences which were noted. Nor¥roix Istanp (type locality). Here there is a tendency for fewer leaflets (4-6 per leaf, 5-8 in Lord Howe Island specimens). Lorp Howe Istanp. Two specimens, Green 1617 & 1669 (both male) appear very different from any other except the New Hebridean specimen by virtue of a sinuate-serrate upper margin to the leaflets, an articulation in the middle of individual pedicels (not more or less at the base), a small calyx, slim filaments more or less uniformly broad from base to apex, not broadest at the base and plant and, together with Moore 39 and Green 1615 bears foliage resembling more closely the Norfolk Island plant. New Hepripes. Unfortunately the only collection seen from these islands is a male specimen and therefore without fruit for comparison with the related New Caledonian species, but the leaves, buds, and flowers resemble the Lord Howe Island plants represented by Green 1617 & 1669, although slightly smaller in all parts. Fiy1 & ToncaA. Here there is a tendency for the texture of the leaf to be subcoriaceous, the leaflet number to increase, (6—)8-10(—15) per leaf, the calyx to be less deeply divided, and the petals to be longer (3.5-4.2 mm. long in con- trast to 2.7-3.0 mm. elsewhere). Fagara gillespieana was described by A. C. Smith from Fiji. I have examined the type, but the leaflet size and texture, upon which, to judge from the protologue, its differentiation seems mainly to have been based, can be matched elsewhere, and appear to me to fall within the range of variation for Zanthoxylum pinnatum. Zanthoxylum pancheri P. S. Green, sp. nov., affine Z. pinnato, a quo foliis plerumque brevioribus magis coriaceis, alabastris conicis acumi- natis, et praesertim fructibus minoribus differt. Blackburnia pinnata sensu Pancher in Sebert & Pancher, Not. Bois. Nouv. Caléd. 244. [1874], non J. R. & G. Forst. Zanthoxylum blackburnia sensu Guillaumin, Ann. Mus. Coll. Marseilles IT. 9 (Cat. Pl. Phan. Nouv.-Caléd. 1: 39): 111. 1911, non Benth. Fagara pinnata sensu Guillaumin, Fl. Nouv.-Caléd. 164. 1948, non Black- burnia pinnata J. R. & G. Forst. 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 217 Blackburnia pulchella Vieill. ex Guillaum. Ann. Mus. Col. Marseilles II. 9 (Cat. Pl. Phan. Nouv.-Caléd. 1: 39): 111. 1911, nom. nud. Type: New Caledonia, 1861-67, Vieillard 2452 (cH, K, isotype es). Zanthoxylum pinnatum sensu D§niker, Viert. Naturf. ie abdlit 77 (Beibl. 19): 195. 1932, non Blackburnia pinnata J. R. & G. F Evergreen shrub or tree to 10 m. high; branchlets glabrous or very young, (2.5—-)4-7(-9) cm. long; petiolules canaliculate 2—4(-5) mm. long; leaflets (4—) 6-8 (—14) in opposite pairs, subcoriaceous, asymmetrical- ly elliptic to ovate, the part on the adaxial side of the midrib much broader than that on the abaxial, (1.4~)1.8-4.5(-5.5) cm. long by (0.9—)1.2-2.4(-2.8) cm. broad; margin entire, slightly recurved; apex obtuse or bluntly and shortly acuminate; base obtuse or acute, adaxial part usually rounded, the abaxial narrowly acute; venation raised and more or less reticulate above and below. Jnflorescence axillary, cymose, ae (3-)4-8(-9) cm. long, very minutely puberulous. Flowers unisexual, 4-merous. Calyx synsepalous, lobes shallow or broadly tri- Pe RG 0.5-0.75 mm. long. Corolla valvate, more or less conical and bluntly acuminate in bud; petals lanceolate, 2-3 mm. long. Stamens antesepaline, filaments 1.2-1.5 mm. long; anthers 0.5-0.7 mm. long. Ovary, in male flowers narrowly conical, 1 mm. long, abortive; in female flowers ellipsoid, 1.8-2 mm. long, 1.2 mm. broad; style subapical, curved, 0.5 mm. long. Fruit more or less spherical, 1-seeded, bivalved capsule, 7-8 mm. long, 5—7 mm. broad; seed black, shiny. New Caledonia. Without locality: ‘‘collines buisées,” 1879, Pancher s.n. (holo- type, K; isotype, Bm); Le Rat s.n. (A). “In parva insula Jandé,” Vieillard 2451 (BM). Pum. 1861-67, Deplanch 10 & 297 (A, GH, K). Mt. Poume, 1868-70, Balansa 3137 (pM, K). Poume, frequent, less than 1000 ft. [308 m.] alt., scrub, 12 Dec. 1934, Compton 2386 (BM). Montagne de Gomouen, prés Gatope, 1861-67, Vieillard 310 (GH, K). Collines de Gomouen, eae 1861-67, Vieil- lard 2452 (BM, GH, K). Oundjo, between Koné and Voh, alt. 0-100 m., 30 Mar. 1956, McKee 4211 (a, K). Bourail, 1868-70, Balansa on (eM, K) & 29 Mar. 1869, Balansa 1171 (A, BM, K). Tontouta Valley, steep rocky slope, N. side of valley at junction of Epieudiols alt. 50-300 m., 15 July 1956, McKee 4894 (K). Nouméa, 1868-70, Balansa 433 & 1687 (per, K). Nouméa (Quen Toro), 1 May 1917, Franc 2099 (A). Bois de Port-Despointes (Nouméa), “bois secs des collines littorales, schistes nummulitiques,” 14 Nov. 1942, Virot 820 (A). After concluding that the New Caledonian plants which have passed under the name Zanthoxylum pinnatum constitute a distinct and unde- scribed species, it was interesting to discover that von Mueller had come to the same conclusion about a century ago (see Fragm. Phytogr. Aus- tral. 7: 141. 1871). I have followed his suggestion in naming the species after the French collector and botanist J. Pancher. Zanthoxylum pancheri is closely related to Z. pinnatum; the leaves are more coriaceous, although occasional specimens of the latter species from Fiji approach it in this character, also the leaves are generally 218 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 shorter and smaller. Most clearly, however, it is separable by the size of its fruit, 7-8 5-7 mm., as opposed to 10-12 & 7-10 mm. in Z. pinnatum. Furthermore, a good ‘ ‘spot” character when the flowers are in bud is the conical, somewhat acuminate, bud, in contrast to an obtuse, rounded bud in the other species. VIOLACEAE The nomenclature of the shrubby members of this family on Norfolk and Lord Howe Islands has been remarkably confused, but a comparison of specimens from these islands with material from New Zealand, Aus- tralia, and other Pacific areas has shown that three separate taxa are worthy of recognition. Beuzenberg (1961) has shown that the genera Melicytus J. R. & G. Forst. and Hymenanthera R. Br. should be united and, following him, the three taxa are placed in the genus Melicytus. Melicytus latifolius (Endl.) P. S. Green, comb. nov. Hymenanthera latifolia Endl. Prodr. Fl. Norfolk. 70. 1833 & Ic. Gen. PI. xiii. t. 108. 1840; Tate in Fletcher, Macleay Mem. Vol., Linn. Soc. New S. Wales 215. 1893; Kirk, Trans. & Proc. New Zealand Enak. 28: 514. 1896; Maiden, Kew 1908: 95. 1908; Laing, Trans. & Proc. New Zealand Inst. 47: 32. 1915; Hoogland in Turner et al. Austral. aoe Foundation Spec. Publ. 1 (The Conservation of Norfolk Is.): Suttonia ? tenuifolia Hook. f, Fl. Antarct. 1: ion ies in Engler, Pflan- zenreich IV. 236 (Heft 9): 335. 1902. Norfolk Island. Without precise locality: Bauer (xk, isotype), Caley (A) & April 1835, Backhouse 628 (K); skirts of woods, July 1830, A. Cunningham 43 (holotype of Suttonia tenuifolia, kK). Upper slopes of Mt. Pitt, 1937 & 1939, McComish 45 (A, K The exact identity of Hooker’s Suttonia tenuifolia has been held in some doubt; Laing in his list of Norfolk Island plants (Laing, 1915) says he knows nothing of it and Mez in his monograph of the M yrsinaceae in Engler’s Pflanzenreich places it as a ‘species dubia.” However, the type specimen, with the name in Hooker’s hand, had, to judge from the pencil annotations upon it, been early recognized as not belonging to the Myrsinaceae and marked “cfr. Melicytus” and “cfr. Hymenanthera,” where it was found in the Kew herbarium. In November 1937, Capt. J. D. McComish noted that he had only found one tree of Melicytus latifolius and that he had “made a thorough search, but cannot find another.” However, in 1939 he said that he has ‘now found several other trees but none as large as the original.” Turner, Smithers & Hoogland in their excellent report on “The Conservation of Norfolk Island” p. 35, say “A small tree, fairly common in the forests of Mt. Pitt Reserve, in particular at the higher levels.’ However, there may be some confusion with M. ramiflorus subsp. oblongifolius, see below. 1970] GREEN, NORFOLK AND LORD HOWE ISLANDS 219 Melicytus novae-zelandiae (A. Cunn.) P. S. Green, comb. nov. Scaevola ? novae-zelandiae A, Cunn. Ann. Mag. Nat. Hist. 2: 52. 1839. Type: New Zealand, North Island, at Matauri, on the sea coast opposite the Cavallos Isles, 1834, R. Cunningham [429] (K). Hymenanthera novae- zelandiae (A. Cunn.) Hemsley, Bull. Misc. Inf. Kew 1908: 96. 1908; Allan, Fl. New Zeal. 1: 194. 1961. Subsp. centurionis P. S. Green, subsp. nov., a subsp. novae-zelandiae foliis chartaceis angustioribus, caulibus junioribus minute papilloso- puberulis differt. Hymenanthera latifolia sensu Hemsley, Ann. Bot. 10: 231. 1896, non Endl. H. novae-zelandiae sensu W. R. B. Oliver, Trans. & Proc. New Zealand Inst. 49: 143. 1917, non (A. Cunn.) Hemsl. Shrub to 2.5 m. high with many stems; branches spreading, glabrous or very minutely papillose-puberulent when young. Leaves chartaceous; petioles 2-5 mm. long; lamina oblanceolate-elliptic, (3.5—)4—7 cm. long, (1.2—)1.5-2.7 cm. broad; base narrowly cuneate and decurrent onto the petiole; apex acute or obtuse and rounded; margin crenate-serrate with (3—)5—6(-8) teeth per side, not thickened or recurved; veins raised and reticulate above and below, (4—)5—6(—7) primary veins per side. IJn- jlorescence axillary to old leaves or their petiole scars with 1 or 2 fascic- ulate dioecious flowers; pedicels 1.5—2 mm. long with, in the middle, 2 opposite bracts with erose rounded margins. Calyx of 5 imbricate, erose, rounded lobes about 1 mm. long and 1-1.2 mm. broad. Corolla of 5 lan- ceolate rounded lobes 2.5—4 mm. long (the smaller in the @ ) more or less reflexed at their tips. Stamens 5, connate by a dorsal membrane produced laterally and above the anthers into 5 more or less lanceolate lobes 2 mm. long, slightly curled at the apex and erose-fimbriate; anthers 1 mm. long with a more or less cuneiform nectariferous sac borne dorsally 0.6 mm. long. Ovary elongated ovoid 2.5 mm. long, contracted into a very short style with 2 somewhat recurved stigmatic lobes. Fruit globose, (?) a berry, about 5 mm. in diameter, with a persistent style. Lord Howe Island. aon scarce, as I have only seen 4 plants,” 15 Nov. 1938, McComish 166 (holotyp K); Dawson’s Point, Eclipse Expedition, C. Moore 64 («K); without precise ocaliey Fullager (x). Although previously treated as the New Zealand plant, this subspecies is quite distinct. Not only are the leaves much thinner in texture and narrower in proportion to their length, giving them a more narrowly cu- neate base, but the margins are not, or scarcely, thickened and recurved. In addition the young stems and bases of the petioles when examined through a strong lens are found to be papillose-puberulent and not gla- brous as in the New Zealand plant. Any floral differences that exist are small and it is clear that separation at the rank of species is scarcely justified. ; It is a pleasure to name this plant in honor of Captain J. D. McComish who, in a letter to the Director of the Royal Botanic Gardens, Kew, 220 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 wrote on 9th November 1946, “I am convinced that this plant is not iden- tical with the New Zealand plant and Dr. Oliver agrees with me in this. However, all the plants, so far as I know, bear only pistillate flowers, but I have a man searching the Island for plants with staminate flower rs.’ Whether any were found I do not know but we may hope that staminate plants exist and that this subspecies, which must be very rare, does not become extinct. The plant from which the Fullager collection cited above came, bore male flowers. Melicytus ramiflorus J. R. & G. For Subsp. oblongifolius (A. Cunn.) P. S. Cua stat. & comb. nov. Hymenanthera oblongifolia A. Cunn. Hook. Lond. Jour. Bot. 1: 124. 1842. Melicytus ramiflorus sensu Hook. f. Handb. New Zealand Fl. 17. 1867, pro parte; F. Muell. Jour. Bot. 23: 354. 1885; C. Moore & Betche, Handb. Fl. New S. Wales 518. 1893; Maiden, Proc. Linn. Soc. New S. Wales 28: 697. 1903 ; Sip Trans. & Proc. New Zealand Inst. 47: 32. 1915, non J. R. & caine dentata R. Br. var. oe (A. Cunn.) Kirk, Trans. & Proc. New Zealand Inst. 28: 511. Norfolk Island. June-July 1830, A. Cunningham 44 (x); “on the skirts of woods, at Long Ridge, at the junction of the old cross road leading to Cascade Road,” July 1830, A. Cunningham 42 (127*) (holotype, K); April 1835, Back- house 641 (K); woods, June 1855, Milne 4 (x). No recent collections of this plant have been seen but Laing (1915) says that it is “not uncommon in the bush.” It is to be hoped that it is protected from extinction. LITERATURE CITED ALLAN, H. H. 1961. Flora of New Zealand 1. Government Printer, Wellington. Be pening. E. J. 1961. Observations on sex differentiation and cytotaxonomy e New Zealand species of the Hymenantherinae (Violaceae). New polires Jour. Sci. 4: 337-349. Goop, R. 1964. The geography of the flowering plants, ed. 3. Longmans, Green & Co., London Laine, R. M. 1915. A revised list of the Norfolk Island flora with some notes on the species. Trans. & Proc. New Zealand Inst. 47: 1-39. Oxtver, W.R. B. 1917. The peed and flora of Lord Howe Island. Trans. & Proc. a Zealand Inst. 49: 94— YAKOVLEV, G. 1967. Zametki po saiaplictes i geografii roda Sophora L. i blizkikh pai Se Leningrad Chem.-Pharm. Inst. 21 (Voprosy Far- makognozii 4): 4 Youne, D. P. 1958. neice in the British Isles. Watsonia 4: 51-69. RoyaL Botanic GARDENS K EW, RICHMOND, SURREY ENGLAND 1970] NEVLING, ELFIN FOREST, 12 221 THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 12. A NEW SPECIES OF GONOCALYX (ERICACEAE). Lorin I. NEVLING, JR. DuRING THE CouRSE of an ecological study of an elfin forest situated on Pico del Oeste, Sierra de Luquillo, Puerto Rico, a similar but not identical area, Cerro La Santa in the Bosque Estatal de Guavate-Carite, was visited and observed for comparative purposes.1 The Cerro La Santa site is situated approximately 25 miles southwest of Pico del Oeste at a slightly lower altitude (903 m.) and has a lower annual rainfall. Surprising- ly, the component genera of the two forests are very similar: some species are common to both, but in some genera different species are found. Two most interesting ericaceous plants belonging to the latter category are Gonocalyx portoricensis (Urb.) A. C. Smith from the Sierra de Luquillo and an undescribed species of the genus from Cerro La Santa. The pur- pose of this note is to describe the new species and to comment on certain aspects of the typification and date of publication of the genus. With the addition of Gonocalyx concolor the genus comprises four casas 17184 represents an undescribed species, probably referable to Gonocalyx. Only one flower is available and the specimen does not seem ample enough for description.””’ The specimen bears the following data, “Colombia, Departamento del Valle, Costa del Pacifico; rio Cajambre; Barco, 5—80 met. alt. 21-30 abril 1944.”’ The plant is an epiphytic shrub with carmine flowers and narrowly elliptic leaves. Smith may be correct in his placement of this entity but over-fractionation of genera in the Thibaudieae (or Vaccinioideae) makes intercalation of new material very difficult. Another collection, our specimen is sterile, which may belong to Gonocalyx is Idrobo & Schultes 961. “Colombia: Meta: Cordillera La Macarena (extremo nordeste) macizo Renjifo, cumbre y alrededores. alt. 1.300-1.900 m. Enero 6-20, 1951.’ The plant is epiphytic with a white corolla and leaves somewhat reminiscent in size and shape of those of G. grea e field work which led to this note was eo ote a by a grant from the Na- aise Sitios phe (GB: 3975) to Dr. Richard A. Howard with whom I ent many instructive hours in the field. Special thanks are due Dr. Howard Pieifer for lotutitie. pe photographing certain critical specimens and notes while at K Thanks are due Mr. R. Desm ew re d, Librarian at Kew, for arranging the photo- op tical text r. L. Emberger of Mo ier for the gift of photograph of the type of Gonocalyx pulcher. The line ee Youn of the anthers is the work of Miss Pamela Bruns. I am also grateful to ; B. Martinson Rico. for providing facilities and generous hospitality in Puerto 222 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Considerable emphasis has been placed upon the length of the corolla tube and the anther tubules in delimiting the genus Gonocalyx (see A. C. Smith, The American Species of Thibaudieae. Contrib. U. S. Natl. Herb. 28: 311-547. 1932, and H. Sleumer, Vaccinioideen-Studien. Bot. Jahrb. 71: 375-510. 1941). As a result of this approach, the genus may be Pohang of species which are not necessarily closely related (except for G. portoricensis and G. concolor). It is possible, if not probable, that the increase in length of corolla tube and anther tubules is the result of se- lection for specific pollinators and that this selection has occurred inde- pendently in Colombia, Dominica, and Puerto Rico. Each corolla seg- ment is innervated by three main veins: a median and two laterals. The laterals depart from the main vein at the very base of the corolla tube. Further, the laterals do not fuse with those of the adjacent corolla lobes, a condition which I believe indicates “incomplete” fusion and one which may permit greater flexibility in modifying tube length. It does not appear that length of corolla tube and length of anther tubule are cor- related i.e., the longer the tube the longer the tubules. On the contrary, it seems that in Gonocalyx the increase in length of corolla tube is a secon- dary and derived development and should not be considered of primary taxonomic importance in generic delimitation. KEY TO THE SPECIES OF GONOCALYX A. Leaves elliptic, ovate, or orbicular, 0.5-3 cm. long, the apex obtuse, retuse, r acute and apiculate; Paty borne singly or in pairs in the leaf axils; anther tubules 8-11 m . Leaf margin Rereger or inconspicuously crenate; corolla tube 12-16 mm. long. Plants of Puerto Ric C. Leaf margin usually ey revolute; calyx light green; corolla tubular, pink or rose fading to white at the apex, the corolla lobes erect; stigma at most subexserted at anthesis, white. «baa Aes eA Ce an op tne See ecraee OR yeti G. portoricensis. Leaf margin plane; calyx red; corolla campanulate, aa red, the corolla lobes spreading; stigma exserted at anthesis, red......... By Ret aa perenne ing Go RR agatha NU alae een nel color. B. Leaf margin shallowly but conspicuously crenate at least aes the apex; corolla rose, fading to white at the apex, tube ca. 20 mm. long. Plants Of Colom as a ee is 8 G. pulcher. A. Leaves ovate, 6-10 cm. long, the apex acuminate; inflorescence loosely race- mose, axillary, 3—8-flowered; corolla red; anther tubules ca. 3.5 mm. long. Paints OF TOMmiInieas es i ev Geer re ae G. smilacifolius. ice] Gonocalyx concolor Nevl., sp. nov. Frutex sempervirens epiphyticus cate irregulariter ramosus. Ramuli velutini rhodochroi. Folia ee ovata elliptica vel orbiculata 1.5—3 cm. longa 1.5—2.25 cm. lata + webhaae coriacea rigida plana apice acuta vel rotundata brevissime anteilats basi rotundata superficie glabra infra pilis sparsim instructa, nervo medio supra impresso, subtus promi- nenti. Inflorescentia lateralis uniflora. Calyx obprismaticus 3—3.5 mm 1970] NEVLING, ELFIN FOREST, 12 223 longus ruber glaber. Corolla campanulata 13-15 mm. longa 6-7 mm. in diametro ruberis lobis patentibus 2-3 mm. longis 4 mm. latis. Stamina 10 aequilonga exserta antheris ca. 3 mm. longis apiculatis tubulis ca. 10-11 mm. longis, filamentis liguliformibus ca, 2 mm. longis. Ovarium 5-loculare, ovulis pluribus. Stylus filiformis ruber; stigma minutum. Bac- cae non visae. Holotypus: Howard & Nevling 16947 (a). Small epiphytic shrubs, evergreen, the young branches and leaves rose- colored, minutely velutinous and glabrescent. Leaves simple, alternate, coriaceous, the blade ovate, broadly elliptic, or nearly orbicular, 1.5—3 cm. long, 1.5-2.25 cm. broad, apiculate and obtuse to acute at the apex, rounded at the base, glabrous above, with a few large trichomes beneath, the venation 5-pli from the base, outer set marginal, inner set submarginal and better developed, the midvein immersed above, emersed beneath, the margin entire except for a few inconspicuous crenations toward the apex; petiole to 2.5 mm. long, velutinous, young leaves brilliantly rose-colored ecoming green. Flowers bisexual, 5-merous, regular, uniformly vivid red (Nickerson 5R 5/13), borne singly on axillary brachyblasts, semipendent; pedicel terete, 9-11 mm. long, red, fringed with trichomes at the summit, bibracteolate near the base, articulated with the calyx; calyx tube ob- prismatic, 3-3.5 mm. long, conspicuously 5-ribbed, ribs cape gaits red, glabrous, the lobes very broad and apiculate, the calyx limb margin nearly entire; corolla tube campanulate, 13-15 mm. long, 6-7 mm. in diameter, carnose, with a few multicellular As tei i below the middle, glabrous within, deciduous, the lobes 5, 2-3 mm. long, 4 mm. broad, spreading; stamens 10, equal, free, inserted on a fleshy 10-lobed disc, deciduous, the anther sacs granular, ca. 3 mm. long, apiculately appen- daged, pollen in tetrads, the tubules ca. 10-11 mm. long, opening by in- trorse elliptical pores 0.5-0.75 mm. long, apices exserted, the filaments liguliform, pubescent on margins, ca. 2 mm. long; ovary inferior, 5-car- pellate and -loculate, the placentation axile, ovules many, the style fili- form, vivid red, glabrous, persistent, the stigma truncate, exserted at an- thesis. Berry not seen. Specimens examined: Puerto Rico. Bosque Estatal de Guavate-Carite, Cerro La Santa, Howard & Nevling 16947 (a, holotype; isotypes to be distributed), Howard 17255 (a), Wagner & Wagner, April 27, 1968 (A), Bro. Alain Liogier 10405 (GH, NY). There is little question that Gonocalyx portoricensis and G. concolor are derived from common ancestral stock. There is great similarity in many details and in aspect, as well as aeeane proximity. The dif- ferences which have developed are due, in my opinion, to a divergence in the pollinators. The flower of G. portoricensis ‘ pendent with a light green calyx and a corolla tube which is pink or rose at the base fading to white at the orifice. The filaments have inwardly and laterally directed trichomes protecting a green, glandular base. It is most likely that it 224 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ficure 1. Photograph of hiienie soe ewes fim’ & Nevling 16947) in nature, approximately twice natural s is insect pollinated although it must be pointed out that this species was under observation on Pico del Oeste for a period of two years and no pollinators were observed. It is in flower sporadically throughout the year and seems to set fruit regularly. The flower buds are sometimes parasitized by an unidentified insect larva which eats the anther sacs and pollen. The newly described G. concolor differs, in part, by having the flowers semipendent (borne at about 45° from the vertical), with vivid red calyx, corolla, and stigma. In addition, the corolla tube is campanu- late with the lobes spreading (see Ficure 1). Slight differences are also noted in the details of the anthers (FicurE 2). It is possible and purely speculative, for no pollinator has been observed here either, that this species has diverged towards hummingbird pollination. It has been col- lected in flower in December, January, February, and April and doubtless has a much longer flowering period. A problem which came to my attention during the preparation of this note was an apparent discrepancy in the date of publication of Gonocalyx pulcher. There are several publications of 1856 which must be con- sidered in some detail to resolve the matter. These are presented in pre- sumed chronological order for discussion. 1. GARDENERS’ CHRONICLE 1856: 152. (March 8) 1856. Under the title “New Plants” is listed “164. GoNocALYx PULCHER Planchon and Linden Fi. Columb. ined. We have received from Mr. Linden a coloured representation of this pretty plant which he offers for sale at 25 fr., and of which we translate his description.” The descrip- 1970] NEVLING, ELFIN FOREST, 12 225 . Stamens of pire de portoricensis ve G. concolor. a-c, G. portoricensis; a, Aiea adaxial view, X 4; stam lateral view ne only one tubule show oe 4: c, fila ment and anther sac, pestis ew, X 6 ag ee concolor; d, mide adaxial vi iew, X 4; e, stamen, la teral view ’ with ity ‘one tubule shown, x 4; f, filament and anther sac, lateral view, X 6. tion follows and is ee by a black and white illustration of a single flowering branc This publication is ata in Index Kewensis (Vol. 2) as the place of publication of Gonocalyx, an opinion which was accepted by two special- ists in American Ericaceae, A. C. Smith and H. Sleumer. The Index Nominum Genericorum lists the generic authority as Planchon et Linden ex Lindley, while Supplement 13 of Jndex Kewensis has an entry attribut- ing the genus to Planchon & Linden ex A. C. Smith! 2. FLORICULTURAL CABINET 24: 57. (March) 1856 Two plants, Cuphea eminens and Gonocalyx pulcher, are described in English, each in a separate paragraph. Each description is chatty in na- ture and presents some of the information included in Gardeners’ Chronicle plus a few notes on horticulture. Some errors have been introduced in- cluding the notation that the Gonocalyx is from Ocana, Mexico. A col- ored plate illustrates a flowering branch of each species. 3. BELGIQUE HorTIcoLe 6: 231, 232. 1856. In a section entitled “Plantes nouvelles de serre froide et tempérée.” and subtitled, “Mises pour la premiére fois dans le commerce, par M. J Linden”’ five species are listed: Calyptraria haemantha, Cuphea eminens, Monochaetum ensiferum, Gonocalyx pulcher, and Scutellaria trianaei. Pre- ceding the descriptions is a notation “Extrait du catalogue de M. Linden, 226 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 pour 1856.” The text is nearly a faithful reproduction of the Catalogue presentation. There are no illustrations. 4. Revue Horticore IV. 5: 162-165. 1856. In an article by Ch. Martins entitled “Revue de l’Horticulture étrangére” is a description of Gonocalyx pulcher which was taken from the Gardeners’ Chronicle. The description is a French translation of the English trans- lation. It would appear that no one seriously considered the possibility of valid publication of Gonocalyx in Linden’s Catalogue of 1855. It is abso- Jutely clear that Linden supplied the text and illustration to Gardeners’ Chronicle with the intent of publicizing his new plant. The Floricultural Cabinet of the same month indicates that the plants are available through Mr. Linden’s London agent. We suspect that there was a bit of promotion on Mr. Linden’s part and that the source of information and of the il- lustration in the Cabinet was the Gardeners’ Chronicle. The two re- maining publications of 1856 are clearly repetitions or translations of previous publications. Corroborative evidence indicating that this interpre- tation is correct is to be found in the nature of the illustrations. The Catalogue contains a fine colored plate which includes three flowering branches and several vegetative ones. The plate in the Chronicle is the left-hand flowering branchlet from the Catalogue printed without color and in reverse. The illustration in the Cabinet is the central flowering branchlet from the Catalogue. The complete (and original) plate which appears in the Catalogue was, in my opinion, originally prepared for pub- lication in the Belgian periodical Flore des Serres, but never used there. All facts lead to the conclusion that Linden’s Catalogue of 1855 did pre- cede all the 1856 publications and should be considered the place of valid publication for Gonocalyx pulcher. 1 believe the description (Cata- logue (Prix-courant de l’établissement d’introduction pour les plantes nouvelles) p. 5, 6. 1855) should be interpreted as a combined generic- specific description and that priority of Gonocalyx was established at that time. Indirect evidence (i.e., the title page lists “Printemps, Eté et Automne de 1855) may point to publication late in the year. A parallel problem with that of publication is that of typification. A. C. Smith typified Gonocalyx pulcher, the type species, by a specimen grown and deposited at Kew (specimen collected in October 1879). This sheet bears the Gardeners’ Chronicle reference and it is possible, if not probable, that the specimen had its origin from Linden’s original sale offering. The important questions are when, where, and by whom the original gathering was made. On the basis of the information supplied by Linden, the original discovery was made by Schlim (Linden’s half-brother) “en com- pagnie du Calyptratia haemantha, etc., dans les provinces de Pamplona et d’Ocafia.” According to Linden, Calyptraria was discovered in 1844 by Schlim at the Paramo de Cachiri and was later found also at Ocafia. 1970] NEVLING, ELFIN FOREST, 12 227 It is of interest that other sources? place Schlim with Linden from the beginning of 1844 (exploring the Santa Marta) until 4 March when they sailed for Europe. Coincidentally, Planchon sets the date of discovery of C. haemantha as 1847 (Flore des Serres 9: 171, 172, pl. 924. 1854). In October of 1845 Funck and Schlim returned to South America and began exploration and plant collection on behalf of Linden. They collected around Pamplona, Chinacota (and Ocafia?) until Funck departed in 1847 with the plant shipment for Europe. Perhaps it is to this date of introduc- tion for Calyptraria and not to the date of actual discovery to which Planchon refers. Linden stated that Schlim collected seeds and plants of the Calyptraria from the Ocafia site and that these were the origin of the plants which he offered for sale. It is possible that it was at this time that the Gonocalyx specimens were collected. However, Schlim re- mained in Colombia following Funck’s departure and continued to col- lect in the vicinity of Pamplona (all specimens lost by shipwreck) and then near Ocafia. He returned to Europe in August of 1852. I believe it is clear that the Gonocalyx was not collected in 1844 but sometime between 1845 and 1852. If, indeed, Schlim was the sole collector the date is nar- rowed to the 1847-52 period. Perhaps a more significant point is the role played by Planchon in this instance. Planchon collaborated with Linden and it was he who did the technical botanical aspects of their joint efforts. Planchon was at Ghent from 1849 to 1851, at Nancy from 1851 to 1853, and arrived in Montpellier in 1853. Linden would have sought Planchon’s critical opin- ion prior to any decision to publish, particularly with such an interesting plant in a difficult family. A specimen, deposited at Montpellier, of Gono- calyx pulcher bears the printed label of J. Linden for the “Voyage de L. Schlim.” The geographic data (Ocafia) correspond to expectations. The date is given as 1851 and the collection number is 153; further, the nota- tion “(vivant)” is included on the label. In addition, there are critical notes on the flower morphology. It is apparent that the Schlim 153 ( MPU) is the holotype (photo a) of G. pulcher Planchon & Linden and that Smith’s typification was in error. ARNOLD ARBORETUM and GRAY HERBARIUM HARVARD UNIVERSITY E. P. Kuru. Report of the Killip-Smith botanical expedition to > pe 1926-1927. . N. ¥. Bot. Gard. 28: 205-220, see especially footnote 3. JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ANALYSIS OF THE COMPLEX VASCULARITY IN STEMS OF DIOSCOREA COMPOSITA Epwarp S. AYENSU PLANT ANATOMISTS HAVE, for many years, been presented with various diagrams designed to represent the course of vascular bundles in mono- cotyledons. In a comprehensive study Bary (1884) classified the vas- N a JN , EIR v Kd fg PT a4 A acd = Ay) aS \ ARS Lx COND FN te Ss OY \ J SAF N Y oan \ é Ficures 1 and 2. Habit of Dioscorea composita Hemsl., * 1/2. 1. Male inflorescence. 1970] AYENSU, DIOSCOREA COMPOSITA 229 cular systems of the monocotyledons into three main categories viz., the palm type, the type of the Commelinaceae, and that of the “anomalous monocotyledons,” i.e. those “which differ fundamentally from .. . the very great majority of Monocotyledons.” In spite of some variation in the vascular tissue in stems of the palm type, the salient features of this construction are similar. Leaf trace bundles are present in all stems together with main vascular stem bundles. The main vascular bundles have the tendency to run near the periphery from each leaf enter the stem separately and take a radially oblique course. rge plants such as the palms the number of bundles reaches several hundred. The bundles from different parts of the same leaf penetrate Kf mL sy) stn a4 Le) GA Aen) 4 (> my WW rp /]S AN ROPR> N\A SD oe, a ZN Ans Wy \ 2. Female inflorescence. 230 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the stem to different depths. The midvein penetrates deepest, while those of the margins extend the least distance. The stem type of the Commelinaceae was described as similar to the dicotyledonous arrangement of tissues. The stem bundles occur in a circle lying near the periphery. These are some distance from the leaf trace bundles which are near the center. The so-called anomalous type of stem, including that of the Diosco- reaceae, was described as nearest to that of the dicotyledons in its vas- cular system. The leaf traces are arranged in a single circle around the circumference of the central cylinder. The vascular bundles penetrate ra- dially into the stem center, extending to unequal depths In an early study of the stem of the Dioscoreaceae Mohl (1845) made a comparison with the stem of the Palmae. He concluded that the vas- cular architecture of these two families was similar although there were some peculiar histological details in the Dioscoreaceae. Zimmermann and Tomlinson, in a series of articles on the anatomy of the palm Rhapis excelsa and Prionium (Juncaceae) (Zimmermann & Tomlinson, 1965, 1967, 1968; Tomlinson & Zimmermann, 1966a, 1966b, 1968a, 1968b), have made some very revealing studies showing more ac- curately than had previously been done the pathway of the vascular bundles. The studies of Rhapis excelsa, inter alia, are intended to serve as a model to represent (a) the course of vascular bundles including (1) leaf trace; (2) vertical bundles; (3) bridge bundles and (4) satellite bundles; (b) the dynamic flexibility that enables some monocotyledons to produce variable numbers of vascular bundles; and (c) the fact that there is no continuity of metaxylem between the stem and the leaf, hence, the leaf is irrigated solely by protoxylem. In a recent publication (Ayensu, 1969), an attempt, based on the con- ventional technique, was made to summarize and explain how the vascular tissue of two successive internodes maintains continuity in the complex nodal structure in stems of the Dioscoreaceae. It became evi- dent that a more reliable understanding of the pathway in the vascular system would be gained by the use of the motion picture analysis tech- nique. Four species of the Dioscoreaceae (Dioscorea composita Hemsl., D polygonoides Humb. & Bonpl., D. friedrichsthalii R. Knuth and Tamus communis L.) have so far been analysed by the Optical Shuttle method. Dioscorea composita has been selected to illustrate here the pathways observed in the species listed, MATERIALS AND METHODS Dioscorea composita (Ficures 1, 2) is a vigorously growing climber with a glabrous, Spehigg woody stem. The vine has a tendency to become sulcate when dry. The leaves are alternate, simple, and with en- tire margins. The leat blade is coriaceous, broadly ovate, cordate at base, abruptly acute at apex, or sometimes with the apex obtuse or apiculate. 1970] AYENSU, DIOSCOREA COMPOSITA 231 The leaves are 10-25 cm. long and 5-20 cm. wide with 7-9 veins promi- nent on both surfaces. Petioles are 6-12 cm. long, sulcate and rather firm. The male inflorescence is prominent, 1-2 in the axils, compound, 20-30 cm. long, and with rachis slightly tomentose. Flowers lack pedicels, are 1—2—3-agglomerate, and occur about 2-3 mm. apart. Bracts are acute, and broadly lanceolate. The perianth is fleshy, tubular, slightly tomentose, and about 3 mm. long; its segments are ovate- orbicular, obtuse and erect, and almost the same length as the tube. The fertile stamens are 6, central, and erect; the filaments averaging 2 mm. long are unequal and fleshy. The style is rudimentary but conspicuous. The female inflorescence is elongate, simple or compound; the capsules are ovoid-oblong, 1.5—2.5 cm. long. The tubers are large, long, and penetrate deeply into the soil. From the seeds sent me by Dr. Franklin W. Martin, plant geneticist, U.S. Department of Agriculture, Mayaguez, Puerto Rico, I was able to obtain germinated seeds of Dioscorea composita in petri dishes at 80° F, in ten days. Between two and three months later the vines were har- vested for histological studies. Other material used for this study was collected by me during a visit to Puerto Rico. A herbarium specimen of D. composita (Baldwin, 14238), collected in Vera Cruz, was also revived (cf. Ayensu, 1967) and used for this study. Material previously stored in the fixative FAA was washed thoroughly in running water for one hour. The stem pieces were then removed from the running water and passed through a series of alcohol concentrations from 50 percent to absolute. From the absolute alcohol the stem pieces were processed through the following changes: 1) 2/3 abs. alcohol and 1/3 chloroform 5) 2/3 chloroform and 1/3 paraffin 2) 1/2 abs. alcohol and 1/2 chloroform 6) 1/2 chloroform and 1/2 paraffin 3) 1/3 abs. alcohol and 2/3 chloroform 7) 1/3 chloroform and 2/3 paraffin 4) 2 changes of pure chloroform 8) 2 changes of paraplast These alcohol-chloroform- paraffin/paraplast changes were conducted in an oven at intervals of 30 minutes. Paraplast blocks uc the stems were cut at 10u, on a rotary microtome. Each section was cut and mounted on a separate slide. Sec- tions were numbered sequentially to facilitate the cinematographic analysis. The sections were dewaxed, stained with a mixture of safranin (95 parts of 1 percent solution in 70 percent ethanol) and Delafield’s haematoxylin (5 parts), dehydrated, and mounted in Canada balsam. The serial analysis of the sections was accomplished with the aid of the optical shuttle system (cf. Zimmermann & Tomlinson, 1965, 1966, 1968) consisting of two identical microscopes, discussion and image over- lap tube (Zimmermann Adapter), and a 16 mm. movie camera. A Data Analyser Projector was used in the study of the films. OBSERVATIONS Anatomy of mature aérial stem. The paragraph which follows is a 232 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 summary of the description of the internode encountered in Dioscorea composita. Trichomes absent; glands and stomata present. Cuticle thick and ridged. Epidermis composed of thick-walled circular or dome-shaped cells with cyto- plasmic contents. Cortex of 6 to 10 layers of cells of variable size; 1-3 outer layers collenchymatous. Endodermoid layer separating cortex from 2 to 3 layers of fibers. Both common and cauline vascular bundles present. Common bundles V-shaped with one phloem unit at converging ends of the V, and two phloem units terminating the flanges of the V. Cauline bundles elliptical with one pair of large vessels each of average diameter of 430y, and with one large phloem unit on inner side of innermost pair of large vessels close to the center of the pith, and another phloem unit at the outer end. Average diameter of large sieve tubes 96y. Pith area constituting one fourth of central cylinder. Crystals: idioblast- ye aL raphide bundles with hae: crystals are also present. Tannin cells observed. Extra floral glands pre Course of the vascular bundles. Internodes from the a portion of the plant show the following progression: (1) Starting from the base upwards in an internode, it is observed that 8 common and 8 cauline vascular bundles main- tain their strict individuality, as well as their shape, until about 850 before a connection is made with the nodal region. At this point alteration occurs in the two, or in some special cases in the four, cauline vascular bundles fac- ing the direction of the leaf insertion. The large phloem units on the inner side of the innermost pairs of metaxylem vessels begin to break down and anasto- mose (Ficures 4, 5). (2) About 1550, from above the innermost protoxylem elements of the same cauline bundles begin to anastomose. (3) This is fol- lowed by the fusion of the phloem elements from the two or four cauline bundles into a full manifestation of a phloem glomerulus. (4) Next is the development of the xylem glomerulus (FicurE 6). (5) Following (3) and (4), the phloem units and the xylem elements of the common vascular bundles begin to anastomose into xylem and phloem glomeruli (FicurE 7). (6) The large metaxylem vessels, until now not affected by the structural changes in the cauline vascular bundles, begin to lose their identity by merging into the xylem glomerulus (FIGURES 8-10). (7) The more centrally located phloem glomerulus “moves” radially between the remnants of the two adjacent cauline bundles and joins the outer one towards the glomerulus of the common bundle organized in the same radial direction. (8) The glomeruli of the other common bundles “move” in the direction of the point of the petiole insertion during the move- ment of (7). (9) The xylem glomerulus formed within the cauline bundles “fol- lows” rapidly behind the phloem glomerulus described in (8), thereby completing the breakdown and the incorporation of the large metaxylem vessels into the plexus. (10) The phloem units at the outer end of the cauline vascular bundles are unaffected by the processes of anastomosis occurring in the node. Instead, the position of the phloem unit is altered only slightly by the stresses applied as a result of the outward “movement” of the xylem and phloem glomeruli. (11) At about the time that the outward “movement” of the glomeruli is com- pleted, the reorganization of the tissues of the inner side of the cauline bundles begins. This reorganization is rapidly followed by the return of the outer half of the cauline bundle into its original position. At this point the elliptical shape of the bundles observed in transverse sections at the internode and in the petiole is restored (Ficures 11, 12). (12) The reorganization of the common bundles follows the process described for (11). (13) At 12,480y above the node 1970] AYENSU, DIOSCOREA COMPOSITA 233 sere a ing : Bay fet “# ‘J Suc iy. he ; FIGURE 3, saetiicdias region where each vascular bundle maintains its indi- viduality, X 105. Center arrow indicates direction of leaf insertion. Distance between FicuREs 3 and 4 is 380 Ficures 4 and 5. Large sihiees units on inner side of the innermost pair of metaxylem vessels yee to break down and anastomose, * 105. Distance be- ph. tween FIGURES 4 and 5 is 150 234 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 F Ps ee ie Ps . Ficures 6 and 7. Large metaxylem vessel an is 80 ABBREVIATIONS (for these and glomerulus; xy. g., xylem glomerulus; com v.b., cauline vascular bundle; m. m. s., main stem; p., petiole; r. b., i ¢ s begin to lose their identity, X pi. figures which follow): ph. g., phloe v.b., common vascular bundle; cau. v., metaxylem vessel; ph. u., phloem unit; raphide bundles. 1970] AYENSU, DIOSCOREA COMPOSITA 2 Mie? ae agee Zee? Ficures 8-10. Sequence showing the “movement” of phloem om 10S glomeruli in the direction (arrow) of the point of the petiole insertion, x 105. Distance between FicuREs 8 and 9 is 20y and between FicuREs 9 and 10 30u. is w& Lea 236 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the reorganization is complete and internodal structure with usually 8 common and 8 cauline bundles is entirely restored. DISCUSSION Although the number of bundles varies between species, and the alter- nation of common and cauline bundles is an established fact, it has been observed that occasionally two common bundles “‘run” side by side. This irregularity may be explained as follows: At the node two cauline bundles originate in the plexus and take their positions in the spaces on either side of the middle common bundle and the lateral common bundles. This ra- tio of two cauline and three common bundles is corrected a little farther down the stem by the elimination in the plexus of one of the lateral com- mon bundles. Therefore, in the internode there are almost always one or more common bundles in excess of the one to one ratio of common and cauline bundles. This irregularity is detected only in the internode be- cause the common bundles never unite during their course in the inter- node. In a similar manner the cauline bundles of some specimens of Dio- scorea macroura, D. preussii, and D. sansibarensis sometimes split into two with all the vascular elements above the pair of innermost metaxylem vessels forming a separate unit radially above the metaxylem Comparison of Dioscorea with Rhapis. In the course of this study it became quite obvious that the vascular architecture of Dioscorea com- posita was strikingly different from that of Rhapis excelsa (Palmae) de- scribed as a model for the monocotyledons by Zimmermann and Tomlin- son (1965). Their detailed study of the vascular bundles revealed four types of branch systems. A leaf trace was found to originate by the gradual transformation of the vertical bundle which reaches its maxi- mum size towards the center of the stem and turns sharply towards the stem periphery where it breaks up into several branches. The metaxylem deficient bundle that enters the leaf as a leaf trace was found to be the direct continuation of the vertical bundle. Zimmermann and Tomlinson (1965) concluded that there was no continuity of metaxylem between stem and leaf. Hence the leaf was irrigated solely by protoxylem. Apart from the studies on Rhapis excelsa the origin of the leaf trace and the dis- continuity of the metaxylem into the leaf has also been demonstrated in other monocotyledonous genera including Washingtonia, Geonoma (Palmae), and Prionium (Juncaceae). The present study, however, has demonstrated that in Dioscorea composita (and in other species of Dio- scoreaceae) the leaf is irrigated not solely by the protoxylem but by the metaxylem as well (FicuRrEs 6-8). Zimmermann and Tomlinson (1965) also noted that in most cases a single vertical bundle arises from a trace plexus. However, they observed that in many cases two or more bundles may arise. The implication of this irregularity in the number of vertical bundles is that a certain measure of flexibility exists in the palms and in other monocotyledons which mani- AYENSU, DIOSCOREA COMPOSITA bho tr ~I FIcure 11. 0 petiole, & 105 th 12 FicureE 12. e vascular b is 40. Transverse section of the petiole showing the reorganization of undies in ‘the petiole, < 105. Distance between FIGURES 11 and 238 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 fests itself in the unequal, sometimes large number of bundles observed in stems The Dioscoreaceae exhibit remarkable constancy in the total number of bundles in the stem. With few exceptions, the number of cauline and common vascular bundles established early in ontogeny is maintained in the plant, regardless of the formation of xylem and phloem glomeruli that reduce all the vascular bundles entering the nodal region into a plexus. The bridge bundles described and demonstrated in cinematographic analysis as branches from each departing leaf trace in Rhapis and Prionium were not encountered in Dioscorea composita. Bridges are short (the _longest one measured in Rhapis is 4.5 mm.) and they make a leaf trace and a vertical bundle appear to fuse directly. If we were to assume that bridge bundles were indeed present in the Dioscoreaceae they must emerge from the leaf trace together with the vertical bundles in a single package. Satellite bundles which Zimmermann and Tomlinson (1965) described as vascular branches from larger leaf traces originating in the region of bridge production were not observed in Dioscorea composita. Satellite bundles are best observed in plants which have fully developed inflores- cences. Zimmermann and Tomlinson also stated that even in plants where inflorescences have aborted very early in their development and are not exomorphically visible, satellite bundles can still be detected These comparisons drawn between the Palmae and the Dioscoreaceae after using similar instrumentation and methods confirm the fact that the vascular systems of these two taxa are basically different. This study also contradicts the earlier belief (cf. Falkenberg, 1876; Mohl, 1845) that the Palmae and the Dioscoreaceae do share a common architectural framework. The use of the Palmae as a model for the representation of the course of vascular bundles in the stems of monocotyledons is being questioned by some plant anatomists in view of the uniqueness of the palms amongst the monocotyledons. Vascular pattern studies in such genera as Trades- cantia (Scott & Priestly, 1925), Alstroemeria (Priestly, Scott, & Gillett, 1935), Ananas (Krauss, 1948), Oryza (Majumdar & Saha, 1957), Zea (Kumazawa, 1961), Ripogonum (Simpson & Philipson, 1969) and my own unpublished data on the bamboo Shibutaea kumusaca and Smilax megalantha have shown conclusively that because of distinct variations which exist it is too early yet to consider the structure of any one taxon as a model for the Monocotyledoneae. SUMMARY The complex vasculature of the aérial stems of Dioscorea composita was traced through the use of the Optical Shuttle method and the Data Analyser Projector. This cinematographic analysis has shown that the course of vascular bundles in D. composita and the other species of Dioscoreaceae so far examined, differs markedly from that of the palm 1970] AYENSU, DIOSCOREA COMPOSITA 239 Rhapis excelsa. Furthermore, the commonly held view that the Dio- scoreaceae and the Palmae share a common vascular framework is not supported. In view of the variations that exist in the vascular patterns in a number of monocotyledons, it is suggested that the selection of any one taxon as a typical monocotyledon should await results of further studies. ACKNOWLEDGMENTS I wish to express my appreciation of the efforts of Miss Monica Brook- man-Amissah who prepared the serial sections for this study. The study was supported in part by a grant from the Smithsonian Research Foun- dation. LITERATURE CITED AYENSU, E. S. 1967. Aerosol OT solution — An effective softener of herbarium specimens for anatomical study. Stain Tech. 42: 155, 156. . 1969. Aspects of the rg ey nodal anatomy ge the Dioscoreaceae. Jour. Arnold Arb. 50: 124— Bary, A. DE. 1884. rnin ats anatomy of the vegetative organs of the anerogams and ferns. [English Translation.] Clarendon Press, Oxford. tionsorgane der Monocotyledonen. 202 pp., 3 pls. Ferdinand Enke, Stutt- gart. Krauss, B. H. 1948. Anatomy of the vegetative organs of the pineapple ~217. Kumazawa, M. 1961. Studies on the vascular course in maize plant. Phyto- Mayumpar, G. P., & B. fk 1957. Nodal anatomy and the vascular system of — har of rice plant (Oryza sativa). Proc. Natl. Inst. Sci. India, B. 22: Mont, a voN. 1845. Untersuchungen iiber den Mittelstock von Tamus ele- phontibes L. Jn: Vermischte Schriften. Tiibingen. pp. 1 Priestey, J. H., L. I. Scort, & E. C. GmtteTr. 1935. The Cle of the shoot in Alstroemeria and the unit of shoot growth in monocotyledons. Ann. Bot. 49: 179. Scorr, i. 1, & J. f ved ee 1925. Leaf and 1 anatomy of Tradescantia Huasinsnsts Vell. Jour. Linn. Soc. (Bot.) 47: Smpson, P. G., & W. R. Puitirson. 1969. eek anatomy in vegetative shoots of Ripogonum scandens Forst. (Smilacaceae). New Zealand Jour. Bot. 7: 3-2 eres B., & M. H. ZrmMERMANN. 1966a. Anatomy of the palm — senelte: Il. Rhizome. Jour. Arnold Arb. 47: 248-261. 1966b. Anatomy of the palm Rhapis excelsa III. Juvenile Bisa! Jour. pers Arb, 47: 301-312. _ 1968a. Anatomy of the palm Rhapis excelsa V. Inflorescence. Jour. Arnold Arb. 49: 291-306. & : 8b. Anatomy of the palm oa excelsa V1. Root and branch insertion. Jour. Arnold Arb. 49: 307- 240 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ZIMMERMANN, M. H., & P. B. Tomitnson. 1965. Anatomy of - eae Rhapis excelsa, I. Mature vegetative axis. Jour. Arnold Arb. 46: —— & Analysis x ona vascular systems in ei Optical Shuttle sae Science 152: 73; ae ap the Palm Rhapis excelsa, IV. Vascular development in apex of vegetative aérial axis and rhizome. Jour. Arnold Arb. 48: 122-142. Vascular construction and development in the aerial stem of Prionium (Juncaceae ). Am. Jour. Bot. 55: 1100-1109. DEPARTMENT OF BOTANY SMITHSONIAN INSTITUTION WasuHincTon, D.C. 20560 1970] GILLETT, STATUS OF PROTOCYRTANDRA 241 THE TAXONOMIC STATUS OF PROTOCYRTANDRA (GESNERIACEAE) GrorcE W. GILuett ! THE GENUs Protocyrtandra is based upon C yrtandra todaiensis Kane- hira, of which the type (Kanehira 1854) was collected on the island of Korror, Palau, Caroline Islands, and described in 1933. In 1934, Hoso- kawa gave generic rank to this taxon on the basis of “4 perfect stamens sitting freely with each other.” The purpose of the present paper is to present an amplified description and an illustration of C. todaiensis, to re- late its geographical location to the distribution of the genus, and to clarify its taxonomic status. It must be emphasized that Cyrtandra is a little studied assemblage of some 600 species in which there is a remarkable propensity for evolu- tionary change, this being most noticeable in the morphology of the flower. The center of diversity for the genus is in the region of New Guinea, the Celebes, and Borneo, where there are about 300 species. In the territory from the Solomon Islands and Queensland, extending west and north to the Nicobar Islands and Botel Tobago Island (map, Ficure 1) there are perhaps 400 species. The genus is distributed from this area north and east to islands of the Pacific on which there are an estimated 200 species, over one-half of which are endemic to the Hawaiian Islands. The extensive distribution of this genus and of a multitude of other insular Pacific genera defies explanation on other than speculative terms. Perhaps the most conservative speculation that can be advanced for Cyrtandra is that dispersal was accomplished by frugivorous birds adapted to the conspicuous, fleshy fruits and very small seeds (Guppy, 1906; Carlquist, 1967; Gillett, 1967). ; Cyrtandra is adapted to almost continually wet, well-shaded habitats in forested ravines and gullies. Dispersal, therefore, has been limited al- most exclusively to those “high” islands on which there is an abundance of rain forest vegetation. There are no cyrtandras on Yap, a “high island on which the rain forest, presumably, was long ago removed by cultivation. No Cyrtandra has been collected on Niihau in the Hawaiian Islands. Although Niihau rises to approximately 400 meters, it is situated in the rain shadow of the island of Kauai and therefore has no rain forest vegetation. No Cyrtandra species occur in the Tuamotu Archipelago, an assemblage of “low” islands; none are known in the Marshall Islands, also “low” islands. co : ; The genus Cyrtandra has a remarkable diversity in habit, occurring as herbs, shrubs, trees, or vines. The leaves are always simple, but may be 1 Supported by the National Science Foundation and the Committee on Research, Academic Senate, University of California. ¥ Pid ify “ ~ 30°. we (} Qs BOTEL ees cr Ss ; 20°44 : : GUAM Drs 1Oo°44 Truk alau . @Pono CARG@LINE ‘ts. i Gxussie 140° 160° 180° 160° 140° “ ie te t + O- SQ5 Yo MARQUESAS *D i SAN ISO , i hae ohne A ; Futuna , 7 EW eo SP . ! + a : { ; W HEBRIDES, Nive Soke : we Ory .) Qi cook, "20% NEW hy, EvaSonca 1s! ' oS, CALEDONIA” UST: IS v 30° GEOGRAPHICAL DISTRIBUTION OF CYRTANDRA PROTOCYRTANDRA * Ficure 1. Map showing distribution of Cyrtandra and Protocyrtandra. cre WOLINOTUY GIONUVY AHL 40 TVNANOL TS “TOA] 1970] GILLETT, STATUS OF PROTOCYRTANDRA 243 alternate, opposite, or whorled, with a diverse expression of marginal de- tail and indument. The flowers are extremely variable in form, most often with two stamens having coherent anthers and three staminodes, some- times with the stamens not united, and occasionally (St. John, 1966) with sterile anthers borne on the staminodes. Therefore, the presence of four or five fertile stamens in the flowers of a given species does not seem particularly notable. While zygomorphy is the rule for the genus as we now know it, the corollas of some Hawaiian species come remarkably close to actinomorphy. It seems reasonable to predict that valvate, actino- morphic corollas, now known only in Cyrtandra todaiensis, will be found in other species. It is appropriate to point out that several species of Cyrtandra are dis- tinguished by possession of unisexual flowers. These species occur in New Guinea, the Solomon Islands, Fiji, and the Hawaiian Islands. The first to be discovered was accorded generic rank, as Cyrtandropsis, by Lauterbach (1910), and other species were added by Schlechter (1923). The genus was reduced by Burtt (1936) on the grounds that species with unisexual flowers thus brought together were very different in other characters. The propensity for the evolution of unisexual flowers is so theme. If we were to give generic status to those populations of Pitto- sporum, Wikstroemia, Santalum, Geniostoma, Alectryon, Pisonia, Sty- phelia, Myrsine, and of other Pacific genera in which unisexual flowers have evolved, the proliferation of new genera would be ludicrous. It is my judgment that this same reasoning applies to evolutionary changes in the number of stamens in Cyrtandra. Generic status for Cyrtandra todaiensis is rejected at this time because I believe there is a strong possibility that a definitive study of Cyrtandra will reveal intermediate expressions and character combinations that would make Protocyrtandra untenable. In this era of proliferation of new genera, it is held that a new genus must be demonstrated to be distinct before, not after, it is proposed. Such proof can be provided in this in- stance only by a review of all of the 600-odd species of Cyrtandra, after which an enduring decision can be reached as to whether this species should be accorded sectional, subgeneric, or generic rank. It is a great pleasure to acknowledge the assistance of Mrs. Sheila A. Hardy, who collected in the Palau Islands during her recent tenure as a Peace Corps volunteer in the Caroline Islands. Mrs. Hardy was assisted by Mr. Demei O. Otobed, who unerringly guided her on several field trips. The field work of Mrs. Hardy and Mr. Otobed provided a mass collection, including spirit collections of flowers and fruits. The following description was prepared from the study of this extensive material. The illustration of Cyrtandra todaiensis was prepared by Mrs. Marion R. Sheehan. The manuscript was constructively reviewed by Mr. B. L. Burtt, Royal Botanic Garden, Edinburgh, to whom appreciation is extended. 244 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 URE 2. Cyrtandra todaiensis. All drawings om fas. é& Otobed s.n. a, puhawite branch, X 1/4; b, 4-stamen flower, X 2; c, same in plane view, 1970] GILLETT, STATUS OF PROTOCYRTANDRA 245 Cyrtandra todaiensis Kanehira, Bot. Mag. Tokyo 47: 679. 1933. FIGURE 2. Protocyrtandra todaiensis (Kanehira) Hosokawa, Trans. Nat. Hist. Soc. For- mosa 24(132): 202, 203. 1934. Erect shrub 2~3 m. high with pubescence of light brown, septate, uni- seriate, noncapitate hairs ca. 30 in diameter and 1 mm. long; mature stems up to 2 cm. in diameter, round in cross section, fistulose; young foliage densely lanate with light brown hairs; leaves decussate, plabrate at maturity, the petioles glabrate, up to 14 cm. long, connate at bases, the blades lanceolate to oblanceolate to elliptic, up to 40 cm. long and 20 cm. broad, unequal and acute to cuneate at base, acute to rounded at apex, undulate to entire at margins, the venation obscure above, conspicuous beneath, the primary veins 10-14 per side, curved apically and distally parallel to the margin, the secondary veins concealed above, reticulate below; inflorescence a congested, sessile, bracteate cyme of 15 to 20 pedi- cellate flowers, the outer bracts villose, lanceolate to oblanceolate, 2 cm. long and 0.5 cm. broad, the inner bracts of similar shape but smaller, deciduous; flowers perfect, borne on pedicels 0.5 to 1.5 cm. long, the calyx foliaceous, deciduous, up to 12 mm. long, cleft into 3, 4 or 5 un- equal or equal, valvate lobes ca. 7 mm. long, densely pubescent on outer surface with noncapitate hairs ca. 60-120» in diameter and up to 4 mm. long, glabrous on inner surface; corolla actinomorphic, up to 17 mm, long, cleft into 4, 5 or 6 equal valvate lobes ca. 5 mm. long, pilose on the outer surface with noncapitate hairs ca. 90u in diameter and 2 mm. long, with- in pubescent at middle of tube with multicellular, uniseriate, non-capitate hairs (below) and capitate hairs (above) ca. 60. in diameter and 1 mm. long; stamens 4, 5 or 6, the glabrous filaments 3 mm. long and adnate at the middle of the corolla tube opposite and slightly below the sinuses of the corolla lobes, all bearing fertile anthers ca. 3 mm. long, the anthers not coherent; cupulate annular disc conspicuous at base of ovary, c mm. high; ovary and style ca. 8 mm. long, the ovary glabrous, the base of the style with capitate glandular hairs, the style cleft horizontally at the dilated apex into 2 applanate stigmatic lobes ca. 2 mm. long, the style separating ca. 1-2 mm. above the summit of the mature fruit; fruit a white, slightly translucent, ovoid, succulent berry with thick, fleshy pericarp and thin, intrusive placentae, up to 1.5 cm. broad and 2 cm. long, the apex rounded: seeds numerous, 2/3 mm. long, ovoid, foveolate. —_ Syntype. Caroline Islands, Palau, Korror (Koror) Island, Todaiyama, 150 m. in dense, wet jungles, August 1932, Kanehira 1854 (Kk). ; r of corolla, % 2; e, 5-stamen flower, 2; £6 in Diane view ata Me pa ng interior of corolla, X 2; h, young oda: ce wots ruined ular a of outer surface of ‘corolla, 20: 3, multi- cellular colleters of inner surface of corolla, x 20; k, ovary Aas mature flow- tr, 4; 1, mature fruit, « 2 246 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Caroline Islands. Toirechuil, Palau group. Growing in a limestone “sink” under dense canopy of shrubs and trees approximately 10 meters above the sea, Hardy & Otobed, s.n. (13 specimens) (A, BISH, E, UC, US LITERATURE CITED Burtt, B. L. 1936. Melanesian plants: II. Kew Bull. 1936: 459-466. wail S. 1966. The biota of long-distance dispersal. IV. Genetic systems e floras of oceanic islands. Evolution 20: 433-455. 1967. The biota of ees dispersal. V. Plant dispersal to Pacific Islands. Bull. Torrey Club 94: 129-162. oe: * W. 1967. The genus es | in Fiji. Contr. U. S. Natl. Herb. 37: 107-159. GuPPY, os B. 1906. Observations of a naturalist in the aay between 1896 and 1899. Vol. II: Plant Dispersal. Macmillan, Lon paneer: C. A. G. 1910. Gesneriaceae. Jn: “Results “ The Netherlands scientific expedition to New Guinea in 1907 under the auspices of Dr. H. A Lorentz.”” Nova Guinea 8: 331. ¢. 66. SCHLECHTER, R. 1923. Gesneriaceae papuanae. Jn: Barney zur Flora von Papuasien. X.” By C. Lauterbach. Bot. Jahrb. 58: 367-378. Sr. Joun, H. 1966. Monograph of Cyrtandra (Gesnericee on Oahu, Ha- waiian Islands. B. P. Bishop Mus. Bull. 229 DEPARTMENT OF LIFE SCIENCES UNIVERSITY OF CALIFORNIA RIVERSIDE, CALIFORNIA 92502 1970] THOMAS, POLLEN IN HYPERICUM PATULUM 247 HAPLOID AND DIPLOID POLLEN IN HYPERICUM PATULUM Joas L. THomas * HYPERICUM PATULUM Thunb. is a spreading shrub, native to Japan and China, but frequently cultivated elsewhere. It is evergreen or semi- evergreen in the southeastern states, but is killed to the ground each year in the Boston area. Even in areas in which it is winter-killed it grows to be 3 to 4 feet high each year, with a rounded growth habit. Cymes of golden yellow flowers 1% to 2 inches across appear throughout July and August. Hypericum patulum var. Henryi Bean has larger flowers, 2 to 2% inches across, and from this variety several cultivars have been derived, including ‘Sungold’, ‘Hidcote’, and ‘Golden Cup’. The cultivars have even larger, usually darker colored flowers and are more profusely blooming Polyploidy was suspected and a study of mature pollen from herbarium specimens revealed both a high percentage of abortive grains and an un- usually large size variation in those appearing normal and staining nor- mally with acetocarmine (Ficures 1 and 3). Subsequent studies of pol- len from buds of fresh material just prior to anthesis showed that the percentage of abortive grains, as determined by staining reaction with either acetocarmine or cotton blue, varied from a low of 30 percent in typical H. patulum to a high of 69 percent in the cultivar ‘Golden Cup’. Size frequency plots for normally staining grains are shown in FIGURES 5 to 8. The size range is unusually high in this material, the diameters varying from 18 to 45 micra for grains taken from a single anther. In each case the size frequency curve is slightly bimodal, and although there were intermediates, there were two fairly obvious size categories. These are readily apparent in Ficures 5 to 8. Although each of the graphs shown was made using measurements taken from a single flower, more than one anther was used in order to obtain a larger number of pollen grains. The pattern is essentially the same, however, when pollen from a single anther is measured. Samples from Hypericum patulum and H. patulum ‘Hidcote’ were examined using a single anther. Dividing the pollen grains into three groups, large, small, and abortive, the following counts were made; H. patulum: 165 large, 378 small, 547 abortive; H. patulum ‘Hid- cote’: 111 large, 203 small, 679 abortive. Germination tests, using a weak sugar solution, revealed that both size groups germinate, apparently with equal frequency (Ficurrs 2 and 4). Moreover, pollen tube growth for the first three hours following ger- mination seemed to proceed at approximately the same rate in the dif- ferent size categories, although measurements of tube length were made on only a small number of cells. wish to lags the help of Dr. Otto Solbrig who read the manuscript and ae ceal suggestions 248 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 oe Fic 1-4. Pollen grains from living material of a il ce iso col- cia. approximately 24 hours before a a Fic. 1, Hyperic oom ‘Hidcote’. Fic Hypericum patulum, Fic. 3, H ie hones. Fic. 4, Hypericum soit ‘Sungold’. Photographed - an initial magnification of ap- proximately 430 <. Further explanation in text. 1970] THOMAS, POLLEN IN HYPERICUM PATULUM 249 60 ~ 60 40 ~ PATULUM 40 GOLDEN CUP ~ ¢ re 2 20 4 20 \ ' t T t i SEG Bees | 24 32 40 48 24 32 40 46 DIAMETER IN MICRA DIAMETER IN MICRA €0 €0 4 40 4 HIDCOTE 40- SUNGOLD : : ES = 2 . 2 ath zo: + Li ‘2 Li 7 , 7 7 24 32 40 ae 24 32 40 a8 DIAMETER IN MICRA DIAMETER IN MICRA Ficures 5-8. Size frequency plots for re grains which showed a normal Staining reaction with acetocarmine. Fic. 5, Hypericum patulum. Fic. 7 2 bericum patulum ‘Golden Cup’. Fic. 7, Hypericum patulum ‘Hidcote’. Fic. 8. Hypericum patulum ‘Sungold’. A chromosome number of 2” = 36 was reported by Sugiura (1936) for Hypericum patulum, from material cultivated in a botanical garden. No other counts for this species were reported by Robson and Adams (1968) in a recent survey of chromosome numbers in Hypericum and related genera. In the present study chromosome counts and pollen measurements were made on each of the varieties and cultivars listed in Taste I. All counts were made from plants in the living collection of either the Arnold Arboretum, Harvard University (Aa), or the University of Alabama Arboretum (va). Column 4 in the table indicates the source from which the material was obtained by the respective Arboretum. Un- fortunately, no material from native populations was available for study. 18 AA 149-49 Kohankie Nursery Kew Gardens, England 18 AA 747-49 — Glasnevin, Ireland 18 UA 73-67 Neosho Nursery AA 263-51 Wayside Nursery 18 UA 41-67 Interstate Nursery Hypericum patulum Hoa on > > a : H. patulum ‘Golden Cup’ H. patulum ‘Hidcote’ H. patulum ‘Sungold’ ll a oo n i H. patulum var. ? n n n n 250 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 All counts and measurements were made from acetocarmine squashes of anthers. Meiosis in the pollen mother cells showed irregularities, how- ever, and there was some variation in chromosome number from one cell to the next. Bridge formation was of frequent occurrence in most of the material examined. In addition, the small and very sticky chromosomes in this species make this rather difficult cytological material. Pairin was irregular and multivalent formation was common. Both trivalents and quadrivalents were seen, with up to four multivalents occurring in a single nucleus. Univalents, too, were observed rather frequently. The univalents lag noticeably at anaphase and usually lead to the formation of micronuclei. This apparently accounts for the large number of abortive grains. Certain other pollen mother cells fail to undergo the second division of meiosis and form dyads rather than tetrads. These cells, there- fore, retain the diploid chromosome complement which probably accounts for the very large grains. Preliminary observations indicate that dyad a given individual, although the number of dyads counted represented a rather small sample. The remaining pollen apparently develops from tetrads following a more normal meiotic division. These occur with the greatest frequency, and with the various meiotic disturbances observed it is not surprising that there should be considerable variation in the size of the grains in this category also. Additional work is needed on these interesting plants, particularly studies of megasporogenesis, and controlled pollination experiments. It should be particularly interesting to make cytological studies of this ma- terial from native populations. Although 2” pollen seldom competes successfully with 1m pollen (Burnham 1962) it is apparent that func- tional 2n pollen is produced by these plants in significant quantities. BIBLIOGRAPHY BuRNHAM, C. R. 1962. Discussions in Cytogenetics. Burgess Publ. Co., Min- Rosson, N. K _& P. ApaMs. 1968. oo numbers in Hypericum and related es Brittonia 20: 95-1 Sbeteuk T. 1936. Studies on the tronniels numbers in higher plants, with special reference to cytokinesis. I. Cytologia 7: 544-595. COLLEGE OF ARTS AND SCIENCES UNIVERSITY OF ALABAMA UNIVERSITY, ALABAMA 35486 1970] THOTHATHRI, DERRIS 251 STUDIES IN THE LEGUMINOSAE, 11. A NEW SPECIES OF DERRIS FROM THE SOLOMON ISLANDS K. THOTHATHRI DvrING a stay at the Herbarium of the Komorov Botanical Institute, Academy of Sciences of the USSR, Leningrad, I came across an interest- ing specimen of Derris from the Solomon Islands (Kajewski 2290) which did not agree with material of any other species of this genus. After critical study, this specimen was found to represent a quite distinct un- described species. Information was then requested from the Arnold Arboretum of Harvard University regarding the duplicates of the Ka- jewski collection. It was learned that these had been incorrectly identi- fied as Derris elegans Benth. The characters of Kajewski’s plant are so well marked that it is being proposed here as a new species. Derris salomonensis Thoth., spec. nov. Prates I, IT. Pertinet ad sect. Derris (§ Euderris Benth.) affinisque est D. multi- florae Benth., a qua tamen differt foliolis maioribus, say desire glabra semper foliis ‘brevicre pedicellis longioribus et ovario puberu Planta volubilis in silva pluviali; ramuli glabri, ieutibaltat, striati. Folia imparipinnata, alterna, 32-40 cm. longa; foliola 5, ovato-oblonga vel oblonga, 14-22 6.5-9.5 cm., integra vel undulata ad margines, ad basin angustata, obtusa vel acuta, interdum breviter acuminata, sub- coriacea, glabra, petiolulata, stipulata; nervi laterales 7—9-jugi, reticulati pulchre, rachide glabra; petiolulis 6-9 mm. longis; stipulis deciduis et stipellis linearibus. Inflorescentia axillaris racemosa, 9-12 cm. longa, multo foliis breviore rachide glabra. Flores 2-3 brevibus stipitibus in- sidentes e nodis emergentibus, 1—1.2 cm. longi; pedicelli tenues, 8-10 mm. longi, glabri vel minute puberuli; bracteae lineares, ad basin pedicelli; bracteolae binae, lineares, quarum una pedicello sub calyce ipso et alterna basi calycis insident. Calyx campanulatus, 1.5—-2 > 2—4 mm., brunneus puberulus extus, integer vel molliter dentatus. Corolla alba, vexillum tusum vel emarginatum ad apicem; alae anguste oblongae, distincte un- guiculatae, 0.9—-1 cm. longae, auriculatae infra; carina navicularis, connata infra, 0.9-1 cm. longa, distincte inguiculata- stamina 10, monadelpha, columna 0.9—1.1 cm. longa, divisa supra; filamentis liberis in triente su- periore, antheris versatilibus. Pistillum 1—1.1 cm. longum, ovarium non stipitatum, elongatum, puberulum, stylo curvato, stigmate obliquo, ovulis 3—4. Legumen ignotum. Tyre. Solomon Islands. Bougainville Island, Karngu, 50 m., 22. 10. 1930, S. F. Kajewski 2290 (Lx, holotype; a, isotype). 252 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 The new species Derris salomonensis, a large climber in the rain forest, has handsome white flowers in pendent racemes. It is a member of sect. Derris (§ Euderris Benth.) and is related to D. multiflora Benth., from which it differs in its larger leaflets, glabrous inflorescence always shorter than the leaves, its longer pedicels and puberulous ovary. The stamens of Derris salomonensis are basically monadelphous, but the vexillary filament is always free at the base and in its upper portion; it is united with the column only in the middle portion. The staminal column is split open dorsally and the other filaments are free for their upper thi Additional material of this interesting new species, particularly in fruit- ing condition, would be most desirable. ACKNOWLEDGMENTS I wish to express my gratitude to Dr. J. T. Vassilcezenko, Curator of the Herbarium, Komorov Botanical Institute, Leningrad, for according me facilities to pursue this work; to Dr. Father H. Santapau, Director, Biology Department, St. Xavier’s College, Bombay, for the Latin trans- lation of the description of the new taxon; to Dr. K. Subramanyam, Di- rector, Botanical Survey of India, Calcutta, for his kind encouragement; and to Dr. Lorin I. Nevling, Jr., of the Arnold Arboretum and Gray Her- barium of Harvard University, Cambridge, Massachusetts, for kindly supplying information about the duplicates of the collection studied. CENTRAL NATIONAL HERBARIUM BOTANICAL SURVEY OF INDIA INDIAN BoTaNIc GARDEN Howrah, INDIA EXPLANATION OF PLATES PLATE I Derris salomonensis Thoth. Photograph of holotype, Kajewski 2290 (LE). PLATE II Derris salomonensis Thoth. Drawings of flower parts. Ficure 1. Flower, in- cluding pedicel and bracteoles; Ficure 2. Standard; Ficure 3. Wing; FIGURE 4. Keel petal; Ficure 5. Staminal column; Ficure 6. Ovary with style and stigma. Enlargements as indicated on each figure. Jour. ARNOLD Ars. VoL. 51 PLATE I THOTHATHRI, DERRIS Jour. ArNoLp Ars. VoL. 51 Pirate II THOTHATHRI, DERRIS The purpose, function, and nonprofit status of this or- Have not chonged oO Hove changed during Lseseattgt and the exempt status for Federal income tox sdlard preceding 12 preceding 12 months Purpote: JOURNAL OF THE ARNOLD ARBORETUM VoL. 51 Juty 1970 NUMBER 3 THE GENERA OF ACANTHACEAE IN THE SOUTHEASTERN UNITED STATES + Rosert W. Lonc ACANTHACEAE A. L. de Jussieu, Gen. Pl. 102. 1789, “Acanthi,” nom. cons. (ACANTHUS FAMILY) Herbs [shrubs] or vines [rarely small trees]. Leaves simple, opposite exstipulate; pubescence of stems and leaves of simple hairs; cystoliths of various forms often present in parenchyma or epidermis. Inflorescences basically cymose (dichasial), often condensed in leaf-axils, modified into racemes or panicles, or flowers sometimes solitary. Flowers perfect, nearly actinomorphic to zygomorphic; bracts and bracteoles usually present and often colored, sometimes large and enclosing the flower. Calyx segments 5, occasionally 4, not infrequently fused, persistent. Corolla sympetalous, the limb 5-lobed or bilabiate, rarely 1-lipped, lobes convolute or im- bricate in bud. Nectariferous disk nearly always present, below ovary. 1Prepared for a generic flora of the southeastern United States, a project of the Arnold Arboretum and the Gray Herbarium of Harvard University made possible through the support of the National oral Foundation (Grant GB-6459X, principal investigator Carroll E. Wood, Jr.). e treatment in this paper, which is the 52nd published in the series, follows ine ese established in the first paper pen Arnold Arb. ot Britain! 1958). The area covered includes North and South Caro- lina, Georgia, Florida, Tennessee, ae bama, Mississippi, Arkansas, and Louisiana. The deactictines are based primarily on the plants of this area, with additional in- formation in ana? Retnnes which have not been seen and verified are marked by a asterisk: The Pasa for this paper was prepared while I held a Mercer Research Fel- lowship at the Arnold Arboretum in the latter half - ies I wan cae express my thanks for the facilities extended to me during my s in Ca bale and to the several people who have aided pula iaeat with this uae of ie heeatiaaibie. Dr. Carroll Wood has spent a great deal of time in editing and checking the manu- script, in making numerous additions and sugg' gestions, in preparing the dissections for and supervising the illustrations, and in aga the legends for them. The illustrations are the work of the set nego Marsh ip ites 5, i-n) and of Diane C. Johnson and Virginia Savage, whose initials will be found on the appro- priate Mary C. Dillon Sacttty checked most of ‘ie references and the abbreviations of books and journals, and Mrs. Nancy Dunkly verifie d most of the remainder and prepared the typescript. My appreciation is extended to them all. 258 JOURNAL OF THE ARNOLD ARBORETUM [vor. 51 Stamens 4 (5), didynamous, or 2 (staminodia often present in 2-stamened flowers, 1, 2, or seldom 3), epipetalous; anthers 1- or 2-loculate, longi- tudinally dehiscent, the connective of various widths; pollen very variable, of many kinds, mostly tricolporate. Gynoecium of 2 united carpels; stig- mas simple or 2-cleft or -lobed, the posterior lobe often aborted or wanting; style filiform, simple; ovary 2-loculate, superior, placentation axile (parie- tal in Nelsonioideae), ovules 2—10 in each locule, anatropous, amphitro- pous, or campylotropous, 1-integumented, tenuinucellate. Fruit typically a 2-valved loculicidal capsule [drupaceous in Mendoncioideae|. Seeds 4(2)—20, usually flat, without endosperm |{endosperm present in Nelson- ioideae| borne on hooklike projections, the modified funiculi of the ovules, termed retinacula or jaculators, these papilliform or apparently lacking in a few genera; seeds usually dispersed through sudden opening of fruit; seed coat smooth or roughened, often mucilaginous when moistened; em- bryo straight or bent; cotyledons broad and flat. Embryo-sac develop- ment normal, monosporic (Polygonum type), embryogeny generally con- forming to Onagrad type (Nelsonioideae) or Solanad type (Ruellioideae). Chromosome numbers x = 7-18, 20, 21. (Including Thunbergiaceae Bremek., Mendonciaceae Bremek.) TypE GENUS: Acanthus L. A pantropical family of about 250 genera and 2600 species with four main centers of distribution: Indo-Malaya, Africa, Brazil, and Central America. About 12 genera, including the two largest, Justicia (about 300 species) and Ruellia (about 225 species), are pantropical. Of the remain- ing genera about 40 per cent are confined to the Western Hemisphere. Approximately 120 genera are monotypic and restricted in range. Rela- tively few taxa are found in the United States; only three genera occur in the Northeastern States, seven additional ones are indigenous to the Southeastern States, and a few more are represented by species in the southwestern United States. The family has long been noted for its taxonomic difficulty. Some authorities (e.g., Lindau, Grant, Melchior) consider the Acanthaceae to a natural family, presumably monophyletic in origin, while others (e.g., Van Tieghem, Bremekamp) have viewed the original composition of the group as being artificial, presumably polyphyletic. For the great majority of genera the family may be recognized by the presence of the peculiar structure of the funiculus that becomes hook-shaped in the mature fruit, and is termed the retinaculum or jaculator. This structure, however, is not typical of three of the subfamilies that contain a few genera of uncertain relationshi The family also is characterized by the presence of cystoliths in the leaves and stems of most (but not all) genera and by the explosive de- hiscence of the mature capsule. Cystoliths are visible as slight protuber- ances or, more commonly, as streaks in the epidermis. Bracts and brac- teoles are generally present and are often showy. Classification within the family is based primarily on minor characters of the androecium, particu- larly the anthers, and on pollen sculpture. Such bases for classification 1970] LONG, GENERA OF ACANTHACEAE 259 often make difficult the accurate identification of genera and species. A wide variety of life forms is found, including many hygrophiles, xero- morphic species with spiny or acicular leaves, lianas, and even mangrove plants. Arborescent forms are rare, and most species are shrubs or herbs with opposite, usually decussate, entire, exstipulate leaves. In the tropics the Acanthaceae, particularly the “Strobilanthes” group of the Old World, are conspicuous, showy elements of the local vegetation. Santapau, for example, listed 42 genera found in the immediate vicinity of Bombay, India. Various genera have been the subject of anatomical and morphological investigations, Detailed accounts and references have been given by Sol- ereder, who also made special reference to the occurrence and morphology of cystoliths. These appear in the form of silicified bodies with or without a cellulose skeleton. Linsbauer earlier had shown that their nature and distribution are valuable in the recognition of genera, especially in the Ruellieae and Odontonemateae. Slightly lignified cystoliths occur in the pith of certain species of Ruellia, and pigmented cystoliths are reported for Ruellia, Sanchezia, and Strobilanthes. In addition to cystoliths, cal- cium oxalate crystals may be secreted in the form of prisms, needles, or other shapes. Solereder reported that petiolar anatomy exhibited either a single arc- shaped vascular strand, or a concentric group of separate bundles with the ends of the crescent very much incurved in some species, or a circle of more or less separate bundles (e.g., Thunbergia), or, rarely, a cylindrical vascular strand (e.g., Crossandra). The epidermis is occasionally papillose (e.g., Ruellia). Stomata are present on both leaf surfaces in some species but are confined to the lower surface in others, and they are nearly always caryophyllaceous, although in at least Lepidagathis they are rubiaceous. A wide assortment of both glandular and nonglandular hairs is reported for various genera. The family is characterized anatomically by the presence of anomalous secondary thickening that occurs in a number of genera. This usually appears as inconspicuous groups of intraxylary and interxylary phloem and as bundles of bodies (described as acicular fibers) that resemble large raphides. The vessels typically are very small (less than 50) or extremely small (less than 254). Wood is semi-ring-porous, and the vessel elements have fine, spiral thickenings, simple perforations, and scanty parenchyma. The vascular system of Justicia americana has been found to consist of seven distinct steles, one central and six arranged around the circumference. The meristeles and the central medullary genera is monostelic, but in some respects the stem anatomy resembles certain Campanulaceae because of the presence of a medullary bundle. The roots generally contain less well-developed xylem. Unusually long root hairs, intraxylary phloem, and pitted lignified pith are also recorded for certain genera. The anatomy of several Ruellieae, including Blechum, Dyschoriste, 260 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Hemigraphis, Hygrophila, and Ruellia, has been described in detail by Tchouproff, who also attempted, rather unsuccessfully, to show relation- ships graphically by means of anatomical comparisons. Danert analyzed leaf, bud, and floral arrangements in selected genera, including Blechum, Justicia, Ruellia, and Thunbergia. He concluded that Acanthaceae exhibit “plagiotropic” growth owing to the presence of a “Sprossendodermis,” a character he attributed to phylogenetically older stocks. Vesque suggested an anatomical characterization of the family, but only in connection with the leaf anatomy. He also gave a classification of cystoliths. The Acanthaceae have long been notorious for their very great varia- tion in pollen morphology, useful in taxonomic diagnosis. Probably no other family has such a wide range of pollen morphological features as the Acanthaceae. Early detailed studies of acanthaceous pollen were made by Bischoff, Fritzsche, and Von Mohl, but the taxonomic value of pollen characters in the classification of the family was first pointed out by Radlkofer. It was on the basis of his work that Lindau elaborated the use of pollen morphology as the chief basis for his classification of the family in Engler & Prantl’s Die Natiirlichen Pflanzenfamilien. Hallier, who reported both three-porous and sparsiporous grains in Pseuderanthe- mum, was among the first to question seriously whether pollen is any more valid than floral morphology as a tool for determining relationships. Later students of the family have continued to emphasize the use of pollen, however. Raj, after an exhaustive survey of pollen types in 260 species of 103 genera, concluded there are considerable grounds for doubting the con- vincing use of pollen as a basis for classification, although he apparently is in general agreement with Bremekamp’s use of pollen in his revisionary studies. The Thunbergioideae have the largest pollen, varying from 60 to 100 p, anomatreme, spheroidal, eight-colpate; Nelsonioideae show much less variation and typically have three-colpate, prolate pollen; the Acan- thoideae sensu Lindau have a very wide range of pollen types, including two-porate bilateral, pantoporate, polyporate, and the most common form, three-colporate, found in most of the tribes. Although many tribes form homogeneous groups, others, such as the Ruellieae and Odontonemateae, have various kinds of pollen sculpturing. Recent studies of pollen and its use as a taxonomic tool in the family have not been altogether reassuring. Bhaduri concluded that the pollen of Nelsonioideae closely resembles that of Andrographis (Ruellioideae), although Raj strongly disagreed. Sharma & Vishnumittre found four widely different pollen types in Eranthemum and Pseuderanthemum. Bremekamp’s rearrangement of tribes and genera is based primarily on his interpretation of pollen relationships. This has led him to recognize a number of segregate genera, especially in the Ruelliinae, based on minor pollen differences. It has yet to be established whether or not these “splinter” taxa are worthy of recognition. The Acanthaceae also have a great range of chromosome numbers. Twenty different haploid numbers have been determined, with n = 7, 9, 1970] LONG, GENERA OF ACANTHACEAE 261 10, 13-22, 25, 26, 28, 30, 31, 34, 40, and 66. Basic numbers of (5)°6, 7; 8, 9, 10, and 11 are known, most species generally falling into one of three series, x = 7, 8, or 9, with x = 7 represented by the greatest number of genera. The lowest haploid number (” = 7) is found in Thunbergia; the chromosomes of Elytraria are among the smallest in the family, about I» in length chiefly with median centromeres. Pizzolongo & D’Arienzo examined Acanthus cytologically and reported the genus as dibasic, with * = 7 or 8; A. spinosus is a sterile polyploid with 2n = 112, but the closely related octoploid A. mollis, 2n = 56, and decaploid A. montanus, 2n = 80, are fertile. Grant, who studied the family cytologically, largely to investigate generic relationships, concluded that polyploidy, especially aneuploidy, has undoubtedly occurred and that the wide range of numbers represents an ascending and descending aneuploid series. Aneuploidy is probably responsible for the large number of genera and species. Grant found that all but three of the 18 genera, where more than one species was examined, had euploids, aneuploids, or both types of polyploids present. Narayanan used cytological studies to demonstrate the close relationships of some genera and intrageneric groups. Grant also based some of his speculations regarding relationships within the Justicieae on chromosome numbers. The considerable embryological literature in Acanthaceae has been re- viewed by Schnarf, Mauritzon, and, most recently, by Maheshwari & Negi. Detailed references are also given by Davis. The linear tetrad develops into a Polygonum-type embryo sac; the embryogeny follows either the Onagrad (Elytraria) or Solanad type. Mauritzon pointed out that the endosperm shows a mode of development peculiar to the T ubiflorae ; this is a cellular type of endosperm that results from free nuclear divisions followed by wall formation in the central chamber. The first two divisions result in a row of three cells which form the endosperm proper, while the micropylar and chalazal cells develop into haustorial organs. The chalazal haustorium is lacking in Thunbergia. Bhaduri concluded that Acanthus and Crossandra ought to be considered the basic type for the family be- cause they lack the free nuclear stage. The advanced type would then be represented by Ruellia and Adhatoda (Justicia) where free nuclear divisions occur. Supernumeraries and persistent antipodals are found rarely in the family, although Hartman reported them in A phelandra. Embryological data have been adduced by Johri & Singh as evidence that the Nelsonioideae properly belong in the Acanthaceae, and not in the Scrophulariaceae, as proposed by Bremekamp, who discounted the im- portance of comparative embryological similarities in Acanthaceae. The family continues to provide considerable interest for the plant embryol- ogist, however. A recent investigation by Sahi presented the details of the distinctive embryo sac and embryogeny of Andrographis that throw some light on the taxonomic relationships of this genus. Unfortunately, the floral biology of the family is not as well understood as the embryology. Characteristically, the flowers are proterogynous, nectar-yielding, and brightly colored and are often arranged in large in- 262 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 florescences. Usually, insects hit the downward directed tips of the pollen sacs and are dusted with pollen on the back. The insect transfers the pollen to projecting stigmas of another flower. Premature dispersal of pollen is apparently prevented by the hairs that frequently surround the anthers. Highly specialized pollinating systems involving extrafloral nec- taries have been reported in Thunbergia. Ornithophily (hummingbirds) and possibly myrmecophily may be factors in the pollinating systems of red-flowered rain-forest species of Aphelandra. Uphof reported cleistogamy in six genera of Acanthaceae. Tharp & Barkley found that all paniculate taxa of Ruellia in Texas are cleistoga- mous, and Long & Uttal discovered that all species of Rwellia of the eastern United States are cleistogamous during part of the reproductive cycle. The general occurrence and importance of cleistogamy in Acanthaceae have yet to be evaluated, but they deserve further attention. Monocarpy ap- parently is common in the Old World genus Strodilanthes; after a number of years of vegetative growth, the plants flower, then die off. Santapau has observed, however, that renewed vegetative growth may occur after flowering in S. callosus Nees if there has been an abundance of ra Seeds are dispersed for short distances by the explosive ones of the ripe capsule. Bremekamp attributed this mechanism to the progressive loss of water from the maturing fruit. Many genera have seeds covered with hairs or scales that become erect with moistening; possibly this is an adaptation that helps to anchor the seed in a suitable substratum. Breeding studies and hybridization experiments are few in number. Meeuse & DeWet have reported natural and artificial intergeneric hybrids between the South African Ruttya, 2n = 36, and Ruspolia, 2n = 42. They suggested that the vigorous but completely sterile hybrid (2n = 39) may have horticultural value and that other intergeneric hybrids may occur. Lindau did not place these two genera in the same tribe. Long has made numerous intrasectional and a few intersectional hybrids in Ruellia. In- vestigations reported thus far suggest that large scale cross-breeding work in Acanthaceae would be of considerable value both taxonomically and horticulturally. Perhaps the difficulty of obtaining materials of this largely tropical group accounts for the dearth of genetic experiments in the family. Little is known about the phytochemistry of the family, perhaps be- cause no important poisonous, medicinal, or useful species are known. Pharmacological investigations of drugs from certain genera have been made, and diuretic and antiasthmatic effects have been credited to mem- bers of the family. Chemical constituents include polyphenols, essential oils, non-volatile isoprenoid compounds, heterosides, and alkaloids. Arndt, Eggers, & Jordaan have determined by spectrographic analysis the struc- ture of five new alkaloids obtained from the South African genus Anisotes and some flavenoid studies have recently been reported by Nair. Ob- viously, much remains to be learned concerning the chemotaxonomy of genera in the famil The Acanthaceae have had a long, tortuous history, taxonomically, since they were first recognized as “‘Acanthi” by B. de Jussieu in 1759. The 1970] LONG, GENERA OF ACANTHACEAE 263 name was adopted by A. L. de Jussieu, who placed eight genera in the group. A general discussion of the family and its affinities was given by Lindley, who included a list of genera supplied by Nees and references to the early literature. Nees’s account in De Candolle’s Prodromus was the first comprehensive treatment of the family, and it established limits for many genera that have been followed up to the present. Endlicher, follow- ing Nees, divided the “order” Acanthaceae into three tribes, Thunbergieae, Nelsonieae, and Echmatacanthi, the last further divided into seven sub- tribes. On the basis of floral morphology Bentham & Hooker recognized 120 genera in five tribes: Thunbergieae, Nelsonieae, Ruellieae, Acantheae, and Justicieae, with the Ruellieae further divided into five subtribes, the Justicieae into six. RadIkofer demonstrated the value of pollen characters in the classifica- tion of Acanthaceae and, as mentioned above, Lindau greatly expanded the critical use of pollen characters in his treatment of the family. Lindau distinguished four subfamilies, Nelsonioideae, Mendoncioideae, Thunber- gioideae and Acanthoideae. Van Tieghem proposed that the ‘“Thunber- giacées” be a separate family to consist of three tribes, ““Thunbergiées,” “Mendonciées,” and ‘“Nelsoniées.” He proposed dividing the remaining Acanthaceae into two subfamilies, the “Acanthoidées” (without cysto- liths), and the “Justicioidées” (with cystoliths). Wettstein was in genera agreement with this change because he concluded there was close affinity between Lindau’s first three subfamilies. However, instead of regarding them as a distinct family, he placed them all in Thunbergioideae, thereby reducing Lindau’s subfamilies to tribes. Interestingly, this was the orig- inal position taken by Nees, whose Anechmatacantheae (without retinac- ula) and Echmatacantheae (with retinacula) correspond to Wettstein’s Thunbergioideae and Acanthoideae. More recently, Bremekamp has made a revision of the family based largely, but not entirely, on detailed pollen characters. He divided the Acanthoideae sensu Nees into Acanthoideae and Ruellioideae, while aban- doning Lindau’s familiar ‘“(Contortae” and ‘“Imbricatae.” Lindau’s 16 tribes were reduced to seven, largely through transfers and expansion of tribal circumscriptions. Bremekamp changed his mind about an earlier reduction of Nelsonioideae to a mere subtribe of Acantheae and removed them from the family, recommending they be placed in the vicinity of the Rhinantheae (Scrophulariaceae). He raised Lindau’s Mendoncioideae and Thunbergioideae to family status and made numerous changes in generic limits, especially in Old World tropical taxa. On the basis of his palynological studies, Raj concurred with Bremekamp’s revisions, but Grant said there is no “cytological criterion” to suggest that the sub- families are sufficiently different to be considered separate families. Grant maintained that the evidence from chromosome studies suggested that a classification based on pollen morphology may be somewhat artificial. Wernham suggested that the Thunbergieae are closely related to the Convolvulaceae, but that they ought to be classified with Acanthaceae because of their “primitive” retinacula. Melchior maintained the same 264 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 general classification of the family as Lindau, while admitting that the subfamilies are indeed strongly marked. Leonard also followed Lindau’s circumscriptions in his treatment of tropical American groups hanges in and additions to the general classification of Acanthaceae continue to the present, with the general tendency of one school (repre- sented by Bremekamp and Raj) to emphasize minor differences in pollen morphology, anther, and other floral characters in recognizing numerous segregate taxa; and the other school (represented by Melchior and Leonard) to follow the broad generic interpretations of Lindau. Melchior stated the situation accurately when he said, ‘“‘Auch die natiirliche Glieder- ung der Acanthaceae . . . ist noch in Fluss.” The phylogenetic relationships of Acanthaceae are also controversial. Baillon thought the family was related to both the Scrophulariaceae and the Labiatae on the basis of stamen morphology and to the Bignoniaceae on the basis of calyx and fruit characters. He thought Acanthaceae were also related to Boraginaceae and to Convolvulaceae through their nearly actinomorphic flowers. Bessey considered Acanthaceae as descended from Scrophulariaceae or groups ancestral to them. Engler and Diels placed the family between Globulariaceae and Myoporaceae in the vicinity of the large and complex Gesneriaceae. Hegnauer, on the basis of phytochemical evidence, suggested that there is close relationship with the Verbenaceae- Labiatae complex, but he may also have been influenced by Wettstein, who had placed Acanthaceae in the Tubiflorae between Bignoniaceae and Verbenaceae. Apparently, students generally agree that Thunbergioideae is the most primitive subfamily on the basis of anatomical, embryological, cytological, and palynological grounds. Melchior considered the family to be an old, highly developed and very natural group distinguished by the “sling mechanism,” pollen mor- phology, and the presence of cystoliths and thought that certain parallel developments have occurred in the closely related Scrophulariaceae. Bremekamp stated a diametrically opposing view that Acanthaceae sensu Lindau cannot be regarded. as a natural entity. He attempted to resolve this problem by excluding what he considers are extraneous elements, as the Thunbergiaceae, the Mendonciaceae, and the Nelsonieae. Bremekamp’s “natural” Acanthaceae is closely similar, then, to Lindau’s subfamily Acanthoideae. Raj concluded from his pollen studies, particularly of the sporoderm, that Acanthaceae sensu Bremekamp is an assemblage of primitive and advanced tribes. He concluded that the Strobilantheae, Barlerieae, and Andrographideae are primitive, while Justicieae, Aphel- andreae, and Acantheae are the most advanced As in most angiospermous groups, there is a sketchy fossil record. Velenovsky & Viniklar believe Acanthaceae date back to Cretaceous times because of their identification of fossils as Strobilanthes. Raj, from photo- graphs by Kuyl, Muller, & Waterbolk, identified fossil pollen taken from Miocene deposits as resembling that of Ruellia and Sanchezia. Because there are so few tree-forms in the family and most taxa are tropical, it is unlikely that fossil woods will be discovered. Investigations of fossil pollen 1970] LONG, GENERA OF ACANTHACEAE 265 appear to be the most promising avenue for historical documentation of Acanthaceae. The family is of little economic importance. A few plants have been re- ported to have medicinal properties, but none is included in modern works on materia medica. Bremekamp pointed out that the leaves of two new species of the Indonesian genus Filetia contain inulin, a carbohydrate not previously found in the family. Burkart listed a number of species, no- tably Justicia campestris Griseb., that can be used for emergency forage. Metcalfe & Chalk reported that blue dyes have been obtained from Strobilanthes species and yellow dyes from Justicia species, but they ap- parently have no economic value. The family is best known horticulturally for its many unusual and interesting ornamental plants that occur in the tropics. Because of their tropical nature, it is not surprising that the Acanthaceae of the southeastern United States reach their best development in Florida. Some genera only reach southern Florida, and others, such as Justicia and Ruellia, are best represented in the United States within the state bound- aries. Other genera represented in Florida, such as Barleria and Hemi- graphis, are apparently not well established outside of cultivation, and still others are found only in gardens. Andrographis has been reported as occurring on chrome ore piles in Virginia but probably is not established in our flora. These three genera are included in the key unnumbered and in brackets. Cultivated members of the Acanthaceae in the southeastern United States, particularly in Florida, include Acanthus montanus T. Anders. (leaves spiny); Aphkelandra tetragona (Vahl) Nees (scarlet corollas, hooded upper lobes); Crossandra infundibuliformis (L.) Nees (bright scarlet-orange corollas); Eranthemum pulchellum Andr. (E. nervosum (Vahl) R. Br. ex Roem. & Schult.) (blue curved corollas) ; Fittonia Verschaffeltii E. Coem. (creeping herb; leaves white or red veined) ; Graptophyllum pictum Griff. (shrubs, leaves variously marked, corollas purplish or red, two lipped); various species of Jacodinia that resemble Justicia (brightly colored two-lipped corollas, inflorescences various) ; Odontonema strictum Kuntze (corollas bright red, flowers in close ra- cemes); Pachystachys coccinea (Aubl.) Nees (Jacobinia coccinea (Aubl.) Hiern) (shrub with dense heads of scarlet flowers); Peristrophe speciosa (Roxb.) Nees (herb, flowers violet, subtended by two large bracts) ; Pseuderanthemum atropurpureum (Bull ex De Vos) Radlk. (leaves purplish or yellow blotched); Sanchezia nobilis Hook. f. (corollas yellow, with yellow revolute rims); and Strobilanthes Dyeriana Mast. (Perilepta Dyeriana (Mast.) Bremek.) (distinctive rose-purplish foliage). REFERENCES: ANperson, T. An enumeration of the species of Acanthaceae from the con- tinent of Africa and the adjacent islands. Jour. Linn. Soc. Bot. 7: 13-54. 1864a. 266 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 On the identification of the Acanthaceae of the Linnean Herbarium, in the usons of the Linnean Society of London. /bid. 111-118. 1864b. _ An enumeration of the Indian species of Acanthaceae. /bid. 9: 425, 426, 3 Arnot, R. R., S. H. Eccers, & A. arent The alkaloids of Anisotes sessili- florus C. B. Cl. (Acanthaceae) —5 new 4-quinazolone gaan Tetrahe- dron 23: 3521, 3532. 1967. Perea ey spectrographic analysi Asrupitto, R. V. A study of the vegetative characters of the eect species of the family Acanthaceae in Mount Makiling and their taxonomic impor- tance. Philip. Jour. Forestry 19: 15-40. 1963 BarLtton, H. Mono ee ane oninesati Hist. Pl. 10: 403-466. 1891. [ Divides family into s BECKNER, J. Becca nen (Acanthaceae), a new Florida weed. Sida oe a | 67. BENOIST, R. Recherches sur la structure et la classification des Acanthacées de la tribu des Barlériées. Théses Fac. Sci. Univ. Paris 1465(Sér. A:706): 5-105. 1912. [Includes references to earlier literature. ] Contribution 4 la flore des Acanthacées de |’Afrique francaise. Not. Syst. Paris 2: 284-288. 1912; 289, 290. 1913. [Notes and comments on 6 genera; includes key to Thunbergia s spp. | Acanthacées de Madagascar. Bull. Mus. Hist. Nat. Paris 31: 386- 388. 1925. [Includes key to Mendoncia and new spp. in genus. | BentHaM, G., & J. D. Hooker. Acanthaceae. Gen. Pl. 2: es ee? 1886. [ Treatment prepared by Bentham BuapurI, S. A contribution to the morphology of pollen grains of agin and its bearing on taxonomy. Calcutta Univ. Jour. Dep. Sci. II. 1: 25-3 1944. [Acanthus, Crossandra, Adhatoda, cite Ruellia.) BreMeKAmp, C. E. B. On the opening mechanism of the acanthaceous fruit. S. Afr. Jour. Sci. 23: 488-491. 1926. (CE. “ig Centralbl. 159: 218. 1930.) [Role of water-loss in sudden opening of fruit. | Notes on the Acanthaceae of Surinam. Rec. Trav. Bot. Néerl. 35: ae 1938. [Includes key to genera and spp. based on pollen charac- . Materials for a monograph of the Strobilanthinae (Acanthaceae). Verh, Nederl. Akad. Wet. Afd. Natuurk. 2. 41(1): 1-306. pls. 1-6. 1944. [Rejects or as artificial, refers emended Strobilanthinae to broadened Ruellie . Notes on he “Aatineete of Java. Ibid. 45(2): 1-78. 1948. [Dis- cussions of generic problems in Justicia, Thunbergia, Hygrophila, Asystasia, Dicliptera. | The delimitation of the Acanthaceae. Proc. Nederl. Akad. Wet. C. 56: 533- 546. 1953. . The Acantheae of the Malesian Area. I. General considerations. bid. 58: 162-171. 1955a. . Notes on some acanthaceous genera and species of controversial ie tion. Acta Bot. Neerl. 4: 644-655. 1955b. [Emphasizes use of morphology in classification of genera and higher categories. ] . A revision of the Malaysian Nelsonieae (Scrophulariaceae). Reinward- tia 3: 157-261. 1955c. [Staurogyne Wall., Nelsonia R. Br., and Elytraria Vahl; Ophiorrhiziphyllon Kurz and Gynocraterium Bremek. also belong to Nelsonieae. | 1970] LONG, GENERA OF ACANTHACEAE 267 New Bornean Acanthaceae. Blumea 10: 151-175. 1960. [New genus and new tribe, Borneacanthus, Borneacantheae. . Remarks on the position of some Australian Acanthaceae. Acta Bot. Neerl. 11: 195-200. 1962. [Realignments in Ruelliinae and Justiciinae. | . On the systematic position of the Australian Nelsonias and Thunbergias and of the Rwellia species which by Domin were referred to Aporuellia Clarke. Proc. Nederl. Akad. Wet. C. 67: 301-306. 1964. [Includes Bru- noniella, nov. gen. | . Studies in the flora of Thaila nd. 32. sere array Nelsonieae; Thunbergiaceae; Acanthaceae. (Second list.) Dansk. Bot. Ark. 23: 195- 224. 1965a. [Includes descriptions of new genera and spp. ———— .. Delimitation and subdivision of the Acanthaceae. Bull. Bot. Surv. India 7: 21-30. & N. E. NANNENGA- BREMEKAMP. A preliminary survey of the Ruel- liinae (Acanthaceae) of the Malay Archipelago and New Guinea. Verh. Nederl. Akad. Wet. 2. 45(1): 1-39. 1948 BurkartT, A. Sree ee como forrajeras de emergencia. Darwiniana 6: 192-202. 1 pl. sie ae ee ie - se Acanthaceae. Jn: W. T. THISELTON-DYER, , Fl. Trop. Afr. 5: 1-192. 1899; 193-262. 1900 Cia H. Anatomia de la madera de dos géneros de las Acantaceas. Re- vista Forest. Venez. 4: 7-15. 1961. [Vessel elements larger in Bravaisia than in Trichanthera.| Crarke, C. B. Acanthaceae. Ja: J. D. Hooxer, ed., Fl. Brit. India 4: 387- 512. 1884; 513-558. 1885 DANERT, - oe die Symmetrieverhaltnisse der Acanthaceen. Flora 140: 307- 325: Davis, G. Ms Sy stematic embryology of the angiosperms, viii + 528 pp. New York. 1966. [Acanthaceae, 31, 32; extensive bibliographic references. | De, A. Cytological investigations in the family Acanthaceae. Sci. Cult. 32: 1966. 198, 199, . Cytological, ear ag and palynological studies as an aid in tracing affinity and phylog in the family Acanthaceae. I. Cytological studies. Trans. Bose Reseach. Inst. 29: 139-175. 1966; II. Floral anatomy. /bid. 30: 27-43. 1967; III. General ere Ibid. 51-65; IV. Palynology and final conclusion. /bid, 31: 17-2 De Bary, A. Comparative oy of ‘iS ety organs of the phanerogams and ferns. (Transl. and annot. F. O. Bow . Scott.) xvi + 659 Oxford. 1884. bptecuini 137-14 Devens G. Des Miao médicinales. These. Ecole Supér. Pharm. Paris. vii + 186 pp. 1896. [Anatomical and pharmacological information. | ELtts, J. L. Chromosome numbers in some members of the Acanthaceae. Sci. Cult. 28: 191, 192. 1962. [Barleria, Crossandra, Ecbolium, Justicia, Petali- dium, Strobilanthes.} ENDLICHER, S. Acanthaceae. Gen. Pl. 696-708. 1839. ERDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. xii + so pp. frontisp. Stockholm; Waltham, Mass. 1952. [Acanthaceae, 30- An introduction to a Scandinavian 1961. [Revised pollen terminology. | Ital. II. Pp BERGLAND, & J, PRAGLOWSKI. pollen flora. Grana Palynol. 2: 1-92. oy GicanTEe, R. Embriologia dell’Acanthus mollis L. Nuovo Giorn. Bot. 36: 5-33. pls. 1-10. 268 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Goop, R. D’O. Anomacanthus, a new genus of Acanthaceae. Jour. Bot. London 61: 161-164. 1923. [Vine related to Mendoncia with unusually large fruit up to 10 cm. in diam. Grant, W. F. Cytogenetic and cytotaxonomic studies in Eupatorium, Celosia, and Acanthaceae. PH.D. diss. Univ. Virginia, Charlottesville. 1953 . A cytogenetic study in the Acanthaceae. Brittonia 8: 121- 149. 1955. Gray, A. Acanthaceae. Synop. Fl. N. Am. 2: 321-331. 1878. Hatter, H. enue Acanthaceen. Nova Acta Acad. Leop.-Carol. 70: 195-240. pls. 9-16. ber Sear oc metallicum sp. n. und das System der Acan- thaceen. Ann. Jard. Bot. Buitenzorg 15: 26-36. pl. 9. 1898. [Raises ques- tions regarding value of pollen characters in determining phylogenetic re- lationships. | HarTMANN, A. Zur Entwicklungsgeschichte und Biologie der Acanthaceen. Flora 116: 216-258. 1923. olseeiehaiu Ruellia, Hemigraphis, Thunbergia, Blechum, Barleria, Belopero HEGNAUER, ae der Pflanzen. Band 3. Dicotyledoneae Acanthaceae- -Cyrillaceae. 743 pp. Basel & Stuttgart. 1964. [Acanthaceae, 41-49. eo R. W. Identification of New World timbers. Trop. Woods 86: 14-25. 1946. [Acanthaceae, 15-20; includes key to ame based on microscopic anatomy; Bravaisia and Trichanthera attain tree siz Hosein, M. Uber den systematischen Wert der Cpuarihen bei den Acantha- ceen. Bot. Jahrb. 5: 422-440. 1884. Huarp, D. Origine de quelques anomalies observées chez des pollens d’Acan- thacées. Pollen Spores 7: 19-26. 1965. [Meiotic abnormalities produce aborted pollen in Jacobinia, Crossandra. | JosepH, J. Chromosome numbers and abnormalities observed in a few mem- bers of Acanthaceae. Curr. Sci. Bangalore 33: 56, 57. 1964. [Nelsonia, sata inion Cardanthera. Kaur, J. omosome numbers in Acanthaceae. II. Sci. Cult. 31: 531, 532. 1965; ir Ibid, 32: 142, 143. 1966. KnutuH, P. Handbook of flower pollination. Vol. 3. (Transl. J. R. A. Davis.) iv + 644 pp. Oxford. 1909. [Acanthaceae, 237-240. In German ed. cf. 3t2): aa ee Kuyt, O., J. MULLER, & H. WaTERBOLK. The application of palynology to - geology with ner to western Venezuela. Geol. Mijnbouw 3: 49-7 1955. LAKELA, O., & F. C. CRAiGHEAD. Annotated checklist of the vascular plants of Collier, Dade, and Monroe counties, Florida. pp. agar Nia Gard. & Univ. Miami Press, Coral Gables. 1965. [Acanthaceae, 82, 83.] & R. W. Lone. Plants of the Tampa Bay area. 109 pp. Univ. South Florida Bookstore, Tampa. 1970. [Acanthaceae, 83, 84. LEONARD, E. C. Botany of the Maya area. Miscellaneous Papers. X. The Acanthaceae of the Yucatan Peninsula. Carnegie Inst. Publ. 461: 191- 238. 1936. : e Acanthaceae of Colombia. Contr. U.S. Natl. Herb. 31: 1-117. 1951; 19-322, ee 323-781. 1958. —. enus of Acanthaceae from Mexico. Wrightia 2: 1-3. 1959a. (Lande, cated to Aphelandra. | ————. Five new species and one new variety of Acanthaceae recently col- 1970] LONG, GENERA OF ACANTHACEAE 269 lected in Venezuela. Bol. Soc. Venez. Ci. Nat. 21: 7-15, 1959b. [ New spp. in Justicia, Ruellia. & . SMITH. Sanchezia and related American Acanthaceae. Rho- dora 66: 313-343. 1964. [Includes new spp. ] Liepau, O. Beitrage zur Anatomie und Morphologie der Mangrove-Pflanzen, insbesondere ihres Wurzelsystems. Beitr. Biol. Pflanzen 12: 182—213. 1913. | Acanthus ilicifolius, a mangrove plant. ] Litto, M. Catalogo de las Acantaceas Argentinas. Lilloa 1: 21-66. 1937. LInpav, Beitrage zur Systematik der Acanthaceen. Bot. Jahrb. 18: 36-64. pls. 1894. ; . anthaceae. Nat. Pflanzenfam. IV. 3b: 274-354. 1895. Linley, J. Acanthaceae. Nat. Syst. Bot. ed. 2. 284, 285. 1836. [Early ref- erences to family, together with revised classification. ] a K. Uber die kalkfreien Cystolithen der Acanthaceae. Ber. Deutsch. Bot. Ges. 39: 41-49. 1921. [Details of development of cystoliths. Lusgock, J. A contribution to our knowledge of seedlings. Vol. 2. viii + 608 pp. London; 646 pp. New York. 1892. iasaaain 348-365. ] MAHESHWARI, P., & V. Nect. The embryology of Dipteracanthus patulus (Jacq. ) Nees. Phytomorphology 5: 456-472. 1955. [Includes review of embryological research in family. Maiti, P. C. Andrographolide: the active principle of Kalmegh. Bull. Bot. Surv. India 6: 63-65. 1964. [Kalmegh is Andrographis paniculata, pre- scribed for liver ailments; includes bibliography. | MartTIN, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513-660. 1946. [Acanthaceae, 627, 628, 629, 646 Mavritzon, J. Die Endosperm und Embryoentwicklung einiger Acanthaceen. Lunds Univ. Arsskr. II. Sect. 2. 30(5): 1-41. 1934. MEEusE, A. D. J., & J. M. J. DE WET. Ruttyruspolia, a natural intergeneric hybrid in Acantharene. Bothalia 7: 439-441. 1961. Metcuior, H. Acanthaceae. Engler’s Syllabus der Pflanzenfamilien. ed. 12. 2: 456-460. 1964. pa C. R., & L. CHatx. Acanthaceae. Anat. Dicot. 2: 1014-1023. 1950. Mever, P. G. Bei eitrag zur Kenntnis der Acanthaceen ——— Mitt. Bot. Staatssam. Miinchen 2: 300-306, 368-385. Die 7 und Euphorbiaceae og Inaugural-Diss. Naturw. Fak. Ludwig-Maximilians Univ. Munchen. 1961. [Includes keys to spp. MoHAN ea H. Y. Endosperm and embryo development in some Acantha- ceae. Proc. 43rd Indian Sci. Congr. 1956(3): 235. 1956. [Peristrophe, Adhatoda, and Barleria. Embryological characters in the classification of the Acanthaceae. Proc. IX Int. Bot. Congr. 2: 267. 1959 . Endosperm in Eranthemum nervosum. Curr. Sci. Bangalore 28: 169, 170. 1959.* . The development of the seed in Andrographis oe Am. Jour. Bot. 47: 215-219. 1960. [Endosperm cellular from beginning. Masanp, Endosperm and seed development in Andrographis echioides Nees. Curr. Sci. Bangalore 31: 7, 8. ; Embryology of Nelsonia campestris R. Br. Phytomorphology W Endosperm in Acanthaceae. /bid. 14: 388-413. 1964. [Systematic review of studies in various genera; includes bibliography. | 270 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Nair, A. G. R. Luteolin as a characteristic flavone of Acanthaceae. Curr. Sci. hina 34: 79, 80. 1965.* NARAYANAN, C. R. Somatic aS aeCipia in the Acanthaceae. Jour. Madras Univ. B. 21: 220-231. 1951la _ Nucleolar conditions in meiosis of Barleria Prionitis. Ibid. 232-235. 51 1b. . Note on chromosome fragmentation in meiosis of Barleria Prionitis. Ibid. 236, 237.\195ic.* _ Studies in the Acanthaceae; microsporogenesis in Justicia Betonica, Barleria Prionitis and Ruellia tuberosa. Ibid. 26: 189-207. 1956 Natarajan, A. T. Studies in the morphology of pollen—Tubiflorae. Phyton 58 genera, in 10 families, including Thunbergia, Ruellia, Asystasia, Hygro- phila, Andrographis, Justicia, and Barleria NEES VON ESENBECK, C. G. Acanthaceae Tadd Orientalis, Jn: N. WALLICH, Pl. Asiat. Rar. 3: 70-117. 1832. ———. Characters of new species of Indian Acanthaceae. Comp. Bot. Mag. 2: 310, 311,: 1837, . Acanthaceae. DC. Prodr. 11: 46-519. 1847. . Acanthaceae. Jn: K. F. P. von Martius, Fl. Brasil. 9: 6-163. pls. 1847. 1-31. Pat, M. Chromosome numbers in some Indian angiosperms I. Proc. Indian Acad. Sci. B. 60: 347-351. 1964. [Aphelandra, Barleria, Dianthera, Jus- ticia. | Pax, F. See L. RADLKOFER. PuHatak, W. G., & K. B. AmBEGAOKAR. Embryological studies in Acanthaceae — the female gametophyte. Jour. Univ. Baroda 4: 87-97. 1955.* [Female gametophyte of Elytraria acaulis, Barleria Prionitis, and Acanthus ilici- folius. Cf. Proc. Indian Sci. Congr. Assoc. 43(3, abstr.): 227. 1955. ee mbryological studies in Acanthaceae — endosperm and em- ee development in Barleria Prionitis L. Jour. Univ. Baroda 5: 73-87. 1956 & pgid tetas bageant: in the Acanthaceae V. Development of embryo sac and endosperm in Blepharis maderaspatensis (L.) Roth. Proc. Indian Acad. Sci. B. 57: 38-95. 1963. PizzoLonco, P., & M. D’Artenzo. Contributo alla cariologia del genere Acan- thus. Ann, Fac. Sci. Agr. Portici Univ. Stud. Napoli III. 30: 175-183. Rasicer, F. H. Untersuchungen an einigen Acanthaceae und Urticaceae zur Funktion der Cystolithen. Planta 40: 121-144, 1951. [Includes discussion of possible role of cystoliths in ogee acer Raprorp, A. E., H. E. Anzes, & C. R. BELL. Manual of the vascular flora of the Carolinas. Ixi + 1183 pp. Univ. of North Carolina Press, Chapel Hill. 1968. [Acanthaceae, 971-974. RADLKOFER, L. Ueber den systematischen Werth der Pollenbeschaffenheit sig den Acanthaceen. Sitz-ber. Math.-Phys. Akad. Wiss. Miinchen II 256-314. 1883. [First called attention to value of pollen characters in systematics of family. See F. Pax, Bot. Jahrb. 5(Lit-ber.): 19, 20. 1884, r a critical review. | Raj, B. Pollen morphological studies in the Acanthaceae. Grana Palynol. 3: 3-108. 1961. 1970] LONG, GENERA OF ACANTHACEAE 271 Rao, V. S. The floral anatomy of some Bicarpellatae. I. Acanthaceae. Jour. Univ. Bombay II. 21(5,B): 7-34. 1953. Recorb, S. J. Classification of various anatomical features of dicotyledonous woods. Trop. Woods 47: 12-27. 1936. [Includes references to Acantha- ceae. REED, C. F. Andrographis, a genus of Acanthaceae new to eastern United States. Castanea 26: 128. 1961. [A. echinoides Nees on chrome ore piles near Newport News, Va. | Rickett, H. W. Wild Flowers of the United States. Vol. 2. The southeastern States. Part 2, pp. 323-688. pls. 117-241. New York. 1968. [Acanthaceae, 444-449, pls. 164, 165. Species of Ruellia, Dyschoriste, Stenandrium, Di- cliptera, and Justicia illustrated in color. | Russow, E. Mittheilungen iiber secretfiihrende Intercellularginge und Cysto- lithen der Acanthaceen, so wie iiber eine merkwiirdige bisher nicht beobach- tete Erscheinung in einzelnen Weichhaftzellen mehrer eee der genannten ae Sitz-ber. Naturf-ges. Univ. Dorpat 5: 308-316. SAHI, B. Embryo sac endosperm in Andrographis Ssinccisaee Nees (Acan- aa Curr. Sci. Bangalore 35: 368, 369. 1966. [Distinctive embryo sac and mature embryo. | SANTAPAU, H. The Acanthaceae of Bombay. Univ. Bombay Bot. Mem. 2: 1- 104. 1951. [Keys to genera together with descriptions. ] . Notes on the Acanthaceae of Bombay. Jour. Bombay Nat. Hist. Soc. 51: 349-368. 1954. [Corrections and appendix to 1951 paper The flowering of Strobilanthes. Ibid. 56: 677. 1959. [Absence of typi SAYEEDUD-D1Nn, M. Some common Indian herbs with notes on their anatomical characters. III. Andrographis echioides Nees. Jour. Bombay Nat. Hist. Soc. 41: 548-550. 1940. [General anatomy. ] ScHArFFnit, E. Beitrage zur Anatomie der Acanthaceen-Samen. Beih. Bot. Centralbl. 19(1): 453-521. 1906. [Detailed anatomical study of the 4 ypes of seeds described for family. | ScHNARF, K. Vergleichende Embryologie der Angiospermen. vi Berlin. 1931. [Contains review of embryology of Acanthaceae, especially ScHREMMER, F, Acanthus mollis, eine europaische Holzbienenblume. Osterr. Bot. Zeitschr. 107: 84-105. 1960. [Includes pollination by Xylocopa. | Scott, F. M. Cystoliths and plasmodesmata in Beloperone, Ficus, and Boehm- eria. Bot. Gaz. 107: 372-378. 1946. SELL, Y. Lee complexes inflorescentiels de quelques Acanthacées. Etude p eee des phénoménes de condensation, de racémisation, akira ae n et de troncature. Ann. Sci. Nat. Bot. XII. 10: 225-300. 1969. Seieue B. D., & VisHNnu-Mitrre. Contribution to the pollen morphology of the genera Eranthemum L. and Pseuderanthemum Radlkof. (Acanthaceae). Proc. Natl. Inst. Sci. hence B. 29: 520-526. 1963. [Four pollen types oc- cur in 15 spp. of 2 g SOLEREDER, H. Sisllaunehe Anatomie seg Dicotyledonen xvi + 984 pp. Stuttgart. 1899. [Acanthaceae, 692—705.] SREEMADHAVAN, C. P. Bremekampia Casanthaceae’ a new generic name. Bull. Bot. Surv. = 6: 323, 324. 1964. [New name for Haplanthus.| STEENIS, C. G. G. J. v. Gregarious flowering of Strobilanthes oo in Ma laysia. Ann. Bet Gard. Calcutta 150 Anniv. Vol.: 91-97. Takizawa, Y. Die Struktur der Pachytanchromosomen einiger Ee hcela 272 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 sowie eine Reihe neu bestimmter Chromosomenzahlen. Cytologia 22: 118- 126. 1957. [Identification of heterochromatic regions in meiotic chrom somes in Thunbergia, Strobilanthes, Hemigraphis, Ruellia, A elinden Peristrophe, Hypoéstes, Thyrsacanthus, Fittonia, Justicia, Beloperone.| TcHovuprorr, O. Quelques notes sur l’anatomie systématique des Re iacees. Bull. Herb. Boiss. 3: 550-560. 1895. TrecHeEM, P. van. Structure du pistil et = Vovule du fruit et de la gra des Acanthacées. Ann. Sci. Nat. Bot . 7: 1-24. 1908. 'Thunbergiacées proposed; not validly published. | Upnor, J. C. Cleistogamic flowers. Bot. Rev. 4: 21-49. 1938. [Cleistogamy Vesgue, J. Sur quelques formations cellulosiennes locales. 1. Eléments nou- veaux “i le liber des Acanthacées. Ann. Sci. Nat. Bot. VI. 11: 181-185. pl. 9 : ao ractéres des principales familles Gamopétales, tirés de l’anatomie “- la she Ibid. VII. 1: 183-360. pls. 9-15. 1885. [Acanthaceae, 326- 335; some anatomical characterizations of family, includes classification of cystoliths. WeErRNHAM, H. F. Floral evolution with particular reference to the sympetalous dicotyledons. New Phytol. 11: 145-166. 1912. WETTSTEIN, R. von. Handbuch der systematischen Botanik. ed. 4. x + 1152 pp. Leipzig, Eres 1933-1935. [Acanthaceae, 909-911.] Witus, J. C. A dictionary of the flowering plants and ferns. ed. 7. (Revised by H. K. Atry SHAW.) xxii + 1214 pp. + Key to the families of flowering plants (liii pp.). Cambridge, England. 1966. [Includes Mendonciaceae Bremek., Thunbergiaceae Bremek. | KEY TO THE GENERA OF ACANTHACEAE IN THE SOUTHEASTERN UNITED STATES A. Retinacula papilliform or absent; herbs [shrubs] or vines op oe anthers 2-loculate; fruit a beaked or cuspidate capsule; seeds 4 B. Vines; ovules 2 in each locule; calyx [annular or] iat: ea 4{or 2], basal lobes of anther spurred or blunt; capsule beaked; pollen spheroidal, spirotreme (subfam. Thunbergioideae). .... 1. Thunbergia. Erect herbs; ovules numerous in each locule; calyx more or ties equally 5-parted; cna 2, basal lobes of anther blunt; capsule beaked or cuspidate; pollen 3-colpate, syncolpate, prolate (subfam. Nelsonioideae). Foie in abet eae mu ae ee Ee a 2. Elytraria. . Retinacula hooklike, subulate, truncate or more or less fimbriate at tip; herbs with or without cystoliths: anthers 1- or 2-loculate; fruit not beaked; seeds 2 to man C. Shoots not articulated; cystoliths absent; stamens 4, anthers 1-loculate; ae). w > pollen colpate (subfam. Acanthoidese). ..... 0.2.0... 3. Stenandrium. C. Shoots articulated; cystoliths cas stamens 2 or 4, if 4 then at least 2 provided with 2-loculate anthers; pollen colporate or porate (subfam. Ruellioideae D. Aestivation of corolla contorted; stamens usually 4; corolla actino- morphic, subactinomorphic or zygomorphic. E. Calyx actinomorphic or nearly so; stamens didynamous; in- Ce various. 1970] LONG, GENERA OF ACANTHACEAE 273 F. Corolla limb actinomorphic or nearly so; capsule stipitate. Inflorescence spiciform; bracts large, closely imbricate; lechum, Q . Inflorescence axillary, or axillary and terminal, cymose, racemose, or paniculate, rarely in heads: bracts usually small, rarely imbricate; corollas projecting well beyond the bracts. H. Calyx lobes lanceolate or linear; anther sacs blunt; corolla limb actinomorphic or subactinomorphic. eet ee eg eee eS 5. Ruellia, H. Calyx lobes bristle-like; anther sacs awned or pointed at the base; corolla limb subactinomorphic. ee ee ON eM eS Fe SP Dyschoriste, F. are zygomorphic, bilabiate; capsule cylindrical, not C ies racemose; stamens united in a short tube; 2 fascicles or rows of hairs present on posterior of the faa. AD Ae Wee a CED Si Hemigraphis |. iG Trifloréscesite axillary, clustered, or flowers apparently solitary; stamens didynamous; corolla without hairs. . . sical Eat spe igh aaa ar Mee ee UND Vs pmlc taaenea aes 7. Hygrophila, E. Calyx zygomorphic, the segments very dissimilar, ete 0 one i D. Aestivation of corolla cochlear; stamens 2 or 4; corolla zygomorphic, hee! strongly bilabiate. Ovules six or more in each locule; capsule not stipitate. 0h Sy Pee eS ed ee ieee ee rographis |. J. Ovules two in each locule; capsule more or less distinctly stipitate Ki. Starnens Aah Gisele ogee. See 8. Asystasia. K. Stamens 2 L. Stems 6-angled; calyx subtended by two partially united bracts; placenta separating from the mature capsule WHIVOR Cer nu a Gere tor arte wes 9. Dicliptera. . Stems subterete or slightly quadrangular; bracts distinct; placenta remaining attached to the valves of the capsule. 10. Justicia, . Subfam. THUNBERGIOIDEAE Lindau 1. Thunbergia Retzius, Physiogr. Sillsk. Handl. 1(3): 163. 1776, nom. cons Climbing, often twining herbaceous [or woody] vines [or shrubs] with articulated stems. Leaves prevailingly sagittate, hastate, or cordate at the base. Flowers large, pedunculate, solitary in the axils [or in racemes, rarely cymose]; subtending bracts 2, spathaceous, large. Calyx persistent | variously reduced or shortly cupuliform, truncate] or 5—16-lobed. Corolla subactinomorphic to slightly zygomorphic, large, funnelform or salverform, tube slender, short, curved or straight, enlarged into a spreading 5- lobed 274 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 limb. Stamens didynamous, attached near the base of the tube; anthers with an apiculate connective, appendaged at base; pollen large, mostly 85-100, in diameter, spirotreme, spheroidal. Disc present, fleshy. Stigma infundibular or peculiarly 2-lobed (see Fic. 1f); ovary fleshy, ovules 2 in each locule, funiculus pulvinate. Capsule rounded, coriaceous, abruptly flat-beaked, subtended by persistent bracts, loculicidally dehiscent; seeds 2 in each valve. Seeds semiglobose, with a conspicuous excavation on the inner side; retinacula none. (Not Thunbergia Montin, Vet. Akad. Hand]. Stockholm 34: 288. pl. 11. 1773, nom. rejic. = Gardenia Ellis | Rubia- ceae|.) Typr species: T. capensis Retz. (Named for Carl Pehr Thun- berg, 1743-1822, Swedish student and successor to Linnaeus, who traveled in Japan and South Africa and wrote floras of these regions, later professor of botany at Uppsala.) — CLocK-VINE. About 200 species of the Old World tropics, with 12 or more occurring in the American tropics and subtropics in cultivation or more or less naturalized, two of these in peninsular Florida. Thunbergia alata Bojer ex Sims, 2n = 18, a slender, herbaceous vine with winged petioles and pale yellow to orange corollas, usually with a dark-purple ‘“‘eye,”’ is locally abundant along roadsides in moist thickets and near abandoned homesites southward from Brevard and Sarasota counties, Florida. It is a native of East Africa but is now thoroughly naturalized in many tropical areas. None of the Florida specimens has the rusty-villous pubescence found on younger stems and leaves of many specimens from the West Indies. Bre- mekamp (1948) formally described several new varieties in T. alata in Java Roxb. (T. volubilis Pers.), also a smooth, herbaceous vine with usually hastate leaves but with white corollas, has been collected in hammocks along the Indian River, Brevard County, Florida. A number of va- rieties, again based on pubescence and on leaf shape, have been recog- nized in this species by both Nees and Clarke. Thunbergia grandiflora Roxb., 27 = 28, 56, an herbaceous or somewhat woody twining vine with large, bright-blue to white, more or less campanulate corollas, a native of India, is cultivated in southern Florida and may persist after cultivation, but apparently has not become established as an escape. All three species belong to sect. THUNBERGIA (§ Euthunbergia Benth. fa. I. Thunbergia. a—m, 7. alata: a, portion of vine with nearly mature fruits, X 4; b, flower, x 114: c. same in partial section, corolla lobes not one one bract removed, calyx and disc intact, to show relative seep of stamens ium and gynoec , mm 05-4, Oller stamen, X 12; e, inner stamen, < 12; f, aiema: 12; g, ovary, disc, ean calyx in section cut to show placenation of two of four ovules, h, mature fruit before pina X 3; i, inner surface +4 valve of capsule after genie , X 3; j, seed, abaxial surface <6 a adaxial aiigees ie ie ge Aue ‘abaxial pete oaks ote radicle and position oi cotyledons, < 6; m, same, in vertical section, adaxial surface to right — radicle, act a ‘and folding of Speedin, x LONG, GENERA OF ACANTHACEAE 1970] 276 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 & Hook.) on account of their two-lobed stigmas and solitary axillary flowers. The remarkable anatomy of Thunbergia has occasioned studies by Chodat, Roulet, and Mullenders. Interxylary phloem is produced directly from the inner surface of an otherwise normal cambium in T. mysorensis T. Anders. (sect. HEXACENTRIS (Nees) Benth. & Hook.) and in T. grandi- flora. This “centrifugal” phloem is produced independently of the other secondary tissue. Also, xylem bridges that differentiate in a tangential fashion occur in normal phloem tissue. The presence of medullary xylem in a number of species is similar to that found in Mendoncia. Roulet also investigated certain intrageneric relationships from an anatomical comparison of the stem and the morphology of pollen. He concluded that T. fragrans and T. alata are not closely related, but rather that T. fragrans is more closely allied with the African T. Aastata, T. capensis, and T. hirta; T. alata is related to T. reticulata and T. annua, of East Africa. Some interesting observations have been made regarding the repro- ductive biology of Thunbergia. Cammerloher (see also Faegri & Van der Pijl) reported that the extra-floral nectaries of T. grandiflora function as part of the pollination system. Certain carpenter bees, such as X ylo- copa aestuans and X. latipes, have been observed as pollinators of this species. The bee must crawl into the corolla tube and in so doing pushes against the stigma. The stigma and anthers are in an expanded outer chamber of the tube, while a constriction in the tube separates this from a small inner chamber that incloses the ovary and nectary. The thick wall of the tube at the constriction prevents tearing of the corolla and blocks passage of the insect. A narrow channel runs down to the nectary alongside the style, and the insect must extend its tongue along this to reach the nectar. Hairs attached to the anthers are touched by the insect from the outside will chew a hole through the base of the corolla thereby obtaining access to the nectar. This “illegitimate” circumvention of the pollinating mechanism is prevented, however, by ants attracted to the extrafloral nectaries on the bracts, pedicel, and calyx. If another insect, such as a Xylocopa, is similarly attracted by the extrafloral nectar, the ants aggressively chase it away. Flowers lacking these nectaries almost invariably have corollas with holes in their bases. Various other floral adaptations to insect pollination have been reported for T. alata and T. erecta (Benth.) T. Anders. by Knuth. Cytological studies in Thunbergia point to the widespread occurrence of polyploidy. Mangenot & Mangenot have reported 2m = 32, 48, and 64 for three species, which suggests a base number of x = 8. Other numbers reported for the genus (2m = 18, 28) may have resulted through aneuploidy. Additional counts would be important in Thunbergia, but the available evidence already suggests that variation in chromosome number has been important in speciation within the genus. 1970} LONG, GENERA OF ACANTHACEAE 277 The classification of plants related to Thunbergia has been a matter of long-standing controversy. Nees (1847) maintained Thunbergia, Meyenia Nees, and Hexacentris Nees as separate genera. Anderson, how- ever, did not, aud later Bentham & Hooker treated the three as sections of an inclusive Thunbergia. Radlkofer suggested on the basis of pollen and anatomical characters restoration of Meyenia to generic rank. He was followed by Lindau (1893), who treated Thunbergia, Meyenia, and Pseudocalyx as constituting subfam. Thunbergioideae, while placing the tropical American Mendoncia in a separate subfamily, Mendoncioideae, on the basis of its drupaceous fruit, absence of retinacula, and pollen morphology. (Lindau’s is the most recent review of Thunbergia in its entirety. In it he established four sections based on the morphology of the stigma and the inflorescence.) Roulet (1894), after a detailed anatomical study, concluded that the three genera could not be maintained separately. Van Tieghem, however, disagreed and, stressing pollen, anther dehiscence, and petiolar anatomy, restored Hexacentris to generic rank and concurred with Radlkofer and Lindau in maintaining Meyenia as distinct from Thunbergia. Bremekamp initially accepted the inclusion of Thundergia in the Acan- thaceae but later (1953) decided to follow Van Tieghem (1908; see under Family References), who had proposed the establishment of a separate family, Thunbergiacées, on the basis of the rostrate capsule and distinctive excavation on the ventral side of the seeds. Bremekamp thought Thunbergia to be more closely related to Mendoncioideae, Big- noniaceae, and Pedaliaceae than to Acanthaceae. Both Lindau and Bremekamp attached great taxonomic significance to pollen types in their classification of Thunbergia, and Raj concurred with Bremekamp that Thunbergioideae ought to be raised to family status on the basis of pollen characters. He also recommended that Meyenia be excluded from the family and transferred to the Pedaliaceae, again on the basis of pollen characters. It appears obvious that the uncertainty of the rela- tionships of Thunbergia cannot be removed by conventional taxonomic approaches. Additional information derived from cytogenetical and chemo- taxonomic techniques is very desirable for this genus. REFERENCES: Under family references see ANDERSON (1864a), BAILLON (p. 423), BENTHAM & Hooker, BREMEKAMP (1938, 1948, 1965), CLARKE, HARTMANN, KNUTH 239), Lrnpau (1895), NEEs (1847), RAJ, SOLEREDER, and VAN TIEGHEM. Benorst, R. Contribution 4 la connaissance des Acanthacées africaines et mal- gaches. Not. Syst. Paris 11: 137-151. 1944. [ Thunbergia, 144-149. | Bor, N. L., & M. B. Rarzapa. Some beautiful Indian climbers and shrubs VIII. Thunbergia. Jour. Bombay Nat. Hist. Soc. 42: 685-697. 1941. [Includes key to 9 cae i species. | BREMEKAMP, C. E. The Thunbergia species of the Malesian area. Verh. Nederl. Maa. tae Afd. Natuurk. 2. 50(4): 1-90. 1955. BuRKILL, I. H. Thunbergia. Bull. Natl. Bot. Gard. Lucknow 22: 1-9. 1958. 278 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 CAMMERLOHER, H. Zur Kenntnis von Bau und Funktion extra- otal Nektarien. Biol. General. 5: 281-302. 1929. [Floral biology of T. grandiflo Cuopar, R. Sur l’origine des tubes criblés dans le es seat seedy Arch. Sci. Phys. Nat. Genéve III. 27: 229-239. pl. 1 Curtis, S., & W. J. Hooxer. Thunbergia alata: ‘albitora). Bot. Mag. 62: Faecri, K., & L. vAN DER PijL. The principles of pollination ecology. ix + 248 pp. Pergamon Press, Toronto, London, etc. 1966. [7. grandiflora, 162-164, with drawings. | Herxke tots, G. A. C. Thunbergia. Occas. Notes Hong Kong Hort. oe 1: 14—- 21. 1931. [Notes on cultivated spp., including T. alata, T. fragrans. | — G. Ubersicht tiber die bisher eer Arten der Gattung Piaiccss Bot. Jahrb. 17(Beibl. 41): 31-43. oe S., & G. MANGENOT. Enquéte sur sre nombres chromosomique dans collection des espéces tropicales. mks Cytol. Biol. Vég. 25: 411-447. 962. [T. erecta, 2n = 64; T. laevis, n = 24; T. togoensis, 2n = 32. ives W. Lori e du phloeme rapespicniee dans les genres Thun- bergia et Stylidium. Cellule 51: 1-48. 1947. [Includes T. fragrans and T. Rapin Narr, A. G. R., S. NacarajANn, & S. S. SUBRAMANIAN. Chemical ea, sa nectar in Thunbergia grandiflora. Curr. Sci. Bangalore 33(13): 1964.* naa L. vAN DER. Houtbij bloemen bij Costus, Bauhinia, Centrosema en Thun- bergia. Trop. Nat. 30: 5-14. 1941.* [Pollination of T. grandiflora. | RADLKoFER, L. Ein Beitrag zur afrikanischen Flora. Abh. Naturw. Ver. Bremen 8: 369-442. 1884. [Includes discussion of systematics of Thunbergia based on anatomy and pollen morphology. | Routet, C. Résumé d’un travail d’anatomie er Nes systematique du genre Thunbergia. Bull. Herb. Boiss. 1: 370-386. 1893. . Recherches sur oom comparée du genre Thunbergia Lin. fil. [bid. 2 259-326, 342-383. SHIBATA, K. Estudios solo de plantas colombianas silvestres y sein Jour. Agr. Sci. Tokyo Agr. Coll. 8: 49-62. 1962. [Chromosome number in Thunbergia. | Stms, J. Thunbergia fragrans. Bot. Mag. 44: pl. 1881. 1817; Sar grandi- flora. ey 50: pl. 2366. 1822; Thunbergia alata. Ibid. 51: pl. 2591. 1825. TIEGHEM, P, vAN. Restauration es genre Hexacentre dans la Puce nouvelle des inbred Ann. Sci. Nat. Bot. IX. 7: 111-116. 1908. Subfam. NELSONIOIDEAE Lindau 2. Elytraria Michaux, Fl. Bor.-Am. 1: 8. 1803, nom. cons. Acaulescent [or caulescent] rigid herbs with basal rosettes. Leaves alternate or sometimes subopposite, basal [or crowded at the ends of branches]. Flowers borne in dense, scapose, terminal spikes, peduncles and inflorescence bearing imbricate, coriaceous bracts; bracteoles sub- tending calyx somewhat shorter than the calyx. Calyx 4-parted, scarious, segments narrow, dentate or entire at the apex. Corolla blue or white, tube cylindrical, limb slightly bilabiate, lower lip 3-lobed. Stamens 2, barely 1970] LONG, GENERA OF ACANTHACEAE 279 exserted, anthers sometimes awn-tipped at the base, 2-loculate, locules equal, parallel; staminodia usually absent; pollen 3-colpate, syncolpate, prolate, about 30 X 40 » in diameter, shed at the 2-celled stage. Ovary 2-loculate, placentation parietal, ovules 6-10 in each locule, amphitro- pous, funicular obturator and retinaculum absent. Capsule narrow, acute at apex, contracted at base, thick-walled, with explosive dehiscence. Seeds more or less flattened; embryo spatulate, slightly bent; endosperm pres- ent, ruminate. Embryo sac with persistent antipodal cells, often be- coming binucleate; endosperm cellular, conforming to the Pedicularis type; embryogeny conforming to Lythrum-variation of Onagrad type. (Tubi- flora J. F. Gmel., nom. rejic.) Type species: E. virgata Michx., nom. illeg. = E. caroliniensis (J. F. Gmel.) Pers. (Name derived from Greek elytron, sheath, referring to the coriaceous bracts that cover the scapes and subtend the flowers.) A small genus of perhaps 15 species in the tropics and subtropics of both Old and New worlds. The American species were treated by Leonard, who recognized twelve species in the Western Hemisphere. In our flora the genus is represented by Elytraria caroliniensis (J. F. Gmel.) Pers. var. caroliniensis (E. virgata Michx., Tubiflora caroliniensis [Walt.| J. F. Gmel.), which occurs on the Coastal Plain from southern South Carolina to southern peninsular Florida (Lee County), where it is lo- cally abundant in bottomlands, wet pine barrens, and in calcareous gravel. It is a well-marked species with large, basal leaves in a rosette, and a stout, virgate scape with one or more spikes. In southern Florida (Collier and Dade counties) a narrow-leaved form, £. caroliniensis var. angustifolia (Fern.) Blake (Tubiflora angustifolia (Fern.) Small) is some- times frequent locally, also in wet or rocky soil. Beyond our range, E. bromoides Oerst., a somewhat dwarf form that may prove to be a geo- graphic subspecies of the E. caroliniensis complex, occurs in Texas The taxonomic position of Elytraria and that of the Nelsonioideae have been controversial since St. Hilaire first included the genus in the Acan- thaceae (1805). Sprengel, for one, did not accept this decision and trans- ferred the genus to the Scrophulariaceae (1817), a course also followed by Link (1821). Lindley (1847) placed Elytraria in his segregate family Nelsoniaceae. Lindau established the subfamily Nelsonioideae because, on the basis of the general habit of the plants, the presence of papilliform retinacula in a number of genera, and the presence of explosively dehiscent capsules, he believed the genera properly belong in the Acanthaceae. The Nelsonioideae, including five genera, three of them monotypic, is the largest of his first three subfamilies. After a detailed study of various groups of genera, Bremekamp (1953, 1965) proposed that the Nelson- ioideae be transferred to the Scrophulariaceae “in the vicinity of the Rhinantheae.”’ Earlier, however, Bhaduri had expressed the view that possibly the Nelsonioideae did not even deserve subfamily rank, since the pollen resembles that of Acanthus and Crossandra, of the Acanthoideae, and is not otherwise distinctive. 280 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Johri & Singh challenged Bremekamp’s views and after a detailed study of microsporogenesis, megagametogenesis, embryogeny, and endosperm formation concluded that Elytraria is in general conformity with other members of the Acanthaceae. The only feature of special interest is the formation of peripheral endosperm cells that form prominent ridges owing to unequal growth. This imparts an uneven or ruminate appearance to the structure, a feature uncommon in Acanthaceae. In an elaborate de- fense, Bremekamp (1965) emphasized the alternate leaves, parietal placentation, scapose habit, and type of endosperm in Nelsonioideae, all characters that tend to associate the group with Scrophulariaceae, particu- larly tribe Rhinantheae. He concluded that Johri & Singh did not prove that the Nelsonioideae belong with the Acanthaceae, although he not only ignored the embryological data, but also the pollen characters and the hemiparasitism of Rhinantheae, a characteristic unknown in Nelsonioideae. Morton has reported 2x = 38 for three African species of Elytraria, sug- gesting a base number similar to that for other Acanthaceae. Martin de- scribed what he termed a “‘spatulate,” slightly bent embryo in E/ytraria, although the Acanthaceae generally produce non-spatulate, strongly bent embryos. Certainly the total evidence is equivocal, and one must conclude that the position of Elytraria and other Nelsonieae is uncertain at the present time. Probably no special advantages would accrue from a trans- fer to the Scrophulariaceae since other problems would then arise. Addi- tional cytological information, as well as genetic and chemotaxonomic evidence, ought to shed light on the systematic position of the group. REFERENCES: Under family references see BAILLON (p. 425), BHADURI, BREMEKAMP (1938, 1942, 1953, 1965), Leonarp (1951), Linpavu (1895), Martin, and Rag. Jouri, B. M., & H. Stncu. The morphology, embryology, and systematic posi- tion of Elytraria acaulis (L. f.) Lindau. Bot. Not. 112: 227-251. 1959. Leonarp, E. C. The American species of Elytraria. Jour. Wash. Acad. Sci. 24: 443-447. 1934. [Six spp.; taxonomic synopsis only, with keys. | . New species of Elytraria from the West Indies and Peru. Jbid. 28: 308- 313. 1938. [E. prolifera, E. crenata, E. planifolia, E. Klugii, spp. nov. Genus limited to Cuba and Hispaniola in W. Indies. ] . Two new tropical American species of Acanthaceae. Proc. Biol. Soc. Wash. 52: 17, 18. 1939, [E. macrophylla, from San Luis Potosi, Mexico, and a Mendoncia from Venezuela. Morton, J. K. The West African species of Elytraria (Acanthaceae), a taxo- nomic and cytological study. Revista Biol. Lisbon 1: 49-58. 1956. SMALL, J. K. Tubiflora acuminata. Addisonia 18: 55, 56. pl. 604. 1934. [E. bromoides Oerst. | Warp, D. B. The genus Anonymos and its nomenclatural survivors. Rhodora 64: 87-92. 1962. [E. caroliniensis (J. F. Gmel.) Pers.] Wirsur, R. L. The identity of Walter’s species of Anonymos. Jour. Elisha Mitchell Sci. Soc. 78: 125-132. 1962. [E. caroliniensis (7. F. Gmel.) Pers. ] 1970] LONG, GENERA OF ACANTHACEAE 281 Subfam. ACANTHOIDEAE Lindau emend. Bremek. Tribe APHELANDREAE Nees emend. Bremek. Subtribe Aphelandrinae Bremek. 3. Stenandrium Nees in Lindley, Nat. Syst. Bot. ed. 2. 444. 1836, nom. cons. Perennial acaulescent [caulescent or suffrutescent] herbs. Leaves entire, often radical. Flowers borne in scapose spikes [or subsessile or sessile, peduncles axillary, leafless], simple [or branched]; bracts herbaceous, imbricate, lanceolate to ovate, entire [sometimes dentate]; bracteoles Fic. 2. Stenandrium. a—j, S. dulce var. floridanum: a, plonering plant, X 1; b, coun | in vertical section. A show position of stamens and stamino ium, flower with bract and two bracteoles, corolla removed, X 4; d, Milvoren of coro tube in vertical section showing two of four stamens and staminodium, < i - f, ovary in vertical section to show septum and two ovules, seeds retained — note retinacula se, i i i le spine-like 4; h, seed with echinate seed coat, after moistening, X 5; i, sing hygroscopic hair from a ee after moistening, < 25: htouns moistened, oriented as in seed (h), X 282 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 subtending calyx short, setaceous. Calyx lobes 5, subequal, narrow, acute, striate-nerved. Corolla infundibuliform, pinkish, purple [or white], tube cylindrical, slender, somewhat oblique and slightly enlarged at the throat, limb [4- or] 5-lobed, spreading, lobes rounded, subequal, retuse or obo- vate, imbricate. Stamens didynamous, adnate in the throat of the tube, included; filaments very short; anthers 1-loculate, linear, connivent or almost cohering in pairs, muticous at base; pollen variable, spheroidal, with indistinct apertures [or 3-colpate, prolate, reticulate or anomotreme, pantoporate]. Disc inconspicuous, truncate. Style filiform, stigma sub- clavate, obconical, simple or obscurely 2-lobed; ovules 2 in each locule. Capsule subfusiform, oblong to subterete; seeds 4, or sometimes fewer by abortion, orbicular, plano-compressed, minutely muricate, echinate [to hispid |, each subtended by a long retinaculum. (Gerardia r. nom. rejic.) LECTOTYPE SPECIES: S. mandioccanum Nees. (Name from Greek stenos, narrow, and andros, male, referring to the narrow anthers characteristic of the Aphelandreae. ) A genus of remarkable distribution with perhaps 30 species, all in tropical and subtropical regions of the Western Hemisphere, except for one that is endemic to Madagascar. Our representative is S. dulce (Cav.) Nees var. floridanum Gray (S. floridanum (Gray) Small, Gerardia flori- dana (Gray) Small), an inconspicuous, uncommon, low herb of seasonal- ly wet pinelands, moist sandy soil, and grassy roadsides of southern peninsular Florida, from Citrus and Osceola counties south into the Florida Keys. Our plants are very similar in appearance to the highly variable var. dulce, which ranges from southern Chile to Mexico; they differ chiefly in usually having upper bracts and bracteoles glabrous or spar- ingly hirsute-ciliate, rather than hirsute. They are highly variable in pubescence and in leaf morphology, the blades varying from narrowly spatulate to suborbicular. Individual plants may be from 4-6 cm. to 15 cm. tall. Superficially, S, dulce var. floridanum also appears to be re- lated to the Mexican S. subcordatum Standl., of Yucatan, and to S. mexi- canum Leonard. Other Mexican plants, such as S. fasciculare (Benth.) Wasshausen may also be elements of the S. dulce complex. Our plants do not, however, appear to be closely related to S. barbatum Torr. & Gray, a very hirsute, many-stemmed plant with numerous, crowded leaves, which occurs in Texas and New Mexico. The relationships of the genus are apparently with other tropical American Aphelandrinae. Nees (1847) thought Stenandrium to be near- est Crossandra. He divided the 16 species into two mtorenal sae ‘“Acaulia vel subacaulia ...” and ‘“Caulescentes .. . Strangely, Bentham & Hooker placed the genus in the tribe Justicieae, subtribe Asystasiinae, among genera to which it bears no relation. Lindau fol- lowed Nees in placing Stenandrium in Aphelandreae; his classification suggested intergeneric relationships with the monotypic Central American Strobilacanthus Griseb. and with the large tropical American genus A phel- andra R. Br. Lindau recognized two unequal sections, sect. StENANDRIUM 1970] LONG, GENERA OF ACANTHACEAE 283 ($ Schizostenandrium Lindau) and sect. SPHAEROSTENANDRIUM Lindau, based entirely on differences in pollen morphology. Our species would be placed in the latter. The present subgeneric disposition of species must be considered unsatisfactory, since one species, the Brazilian S. trinerve Nees, has one variety in one section, another variety in the other! Breme- kamp initially placed the genus in ‘Acantheae (1948) but later returned it to the Aphelandreae (1965). Stenandrium is one of several genera of Acanthaceae that exhibit a wide range of pollen types. Raj found that the pollen of S. barbatum was 3- colpate, goniotreme, prolate; that of S. trinerve anomotreme, spheroidal; that of S. Pohlii Nees (Brazil) 6—7-porate, spheroidal; and that of S. droseroides Nees 3-colpate, prolate, with finely reticulate sculpturing. Pol- len-grain size, however, was fairly uniform. With this much variation in pollen morphology it appears that pollen characters are not a sound basis for future monographic revision, Chromosome counts are highly desirable, since not a single species of Stenandrium has been reported on cytologically. REFERENCES: Under family references see BAILLON (p. 461), BREMEKAMP (1948, 1953, 1965), Gray, HALLIER ee Lrnpau (1895), Leonarp (1953), LINDLEY (1836), NEEs (1847), andR LEONARD, E. C. reas pee novae vel criticae. Wrightia 2: 75-82. 1960. [Includes description and figure of S. Carolinae Leonard & Proctor and formal publication of S. bracteosum (Britt. & Millsp.) Britton ex Leonard, comb. nov., a close relative of the new sp. | A new Stenondrium from the State of Durango, Mexico. /bid. 83-85. [S. pelorium Leonard, an unusually large sp. | STANDLEY, P. C. Three new plants from Yucatan. Jour. Arnold Arb. 11: 47, 48. 1930. [Includes S. subcordatum Standl. } THIERET, J. W. Proposal for the conservation of the generic name Stenandrium Nees versus Gerardia L. (Acanthaceae). Taxon 5: 58. 1956. Subfam. RUELLIOIDEAE Bremek. Tribe RUELLIEAE Nees emend. Bremek. The Ruellieae as redefined by Bremekamp (1944) and Bremekamp & Nannenga-Bremekamp (1948) is a large tribe composed of several sub- tribes, none of which is precisely delimited. They all, however, are characterized by usually contorted aestivation of the corolla lobes, by spheroidal or ellipsoidal, 3- or more-porate pollen grains, and by reticulate, spinulose, or banded pollen surfaces. Lindau’s tribes Hygrophileae, Pet- alidieae, Strobilantheae, and Barlerieae, are retained as subtribes with considerable emendation by Bremekamp; they are distinguished largely on the basis of differences in pollen morphology. The descriptive anatomy of several genera of Ruellieae showing comparative relationships was re- ported by Tchouproff. 284 JOURNAL OF THE ARNOLD ARBORETUM [von. 51 Subtribe Blechinae Bremek. 4. Blechum P. Browne, Civ. Nat. Hist. Jamaica 261. 1756. Perennial herbs. Leaves petiolate, entire, repand, dentate or crenate. Flowers small, sessile or on very short pedicels in the axils of bracts; in- florescence a more or less dense, terminal |or axillary] spike or spiciform raceme; bracts imbricate, ciliate, broad, suborbicular [or elongate]. Calyx deeply 5-parted, segments linear-subulate, nearly equal [or un- equal, almost bilabiate|. Corolla infundibuliform, white or purplish; tube elongate, slender; limb short, nearly equally 5-lobed, contorted in aestiva- tion, barely or not extending beyond the subtending bracts of the flower. Stamens didynamous, included, adnate about the middle of the tube; anthers 2-loculate, muticous, locules parallel; pollen 3-colporate, syncol- porate, spheroidal, about 47 » in diameter. Style filiform; stigma subulate, obscurely lobed, the posterior lobe minute; ovules few to 6 in each locule. Capsule broadly oblong [ovoid or orbicular], with a short, narrow base. Seeds suborbicular, plano-compressed, surface with mucilaginous hairs when moistened; supporting retinacula acute; septum breaking away in the mature capsule. Type species: Ruellia Blechum L. = Blechum Brownei Juss. (Apparently a Greek name for wild pennyroyal, Mentha Pulegium L.) Blechum is a small genus of ten or possibly fewer species of herbs in- digenous to the American tropics; one species has been introduced into the Old World tropics as a weed and is now rather common in warm cli- mates of the world. In general, the genus is recognized by its dense, sometimes 4-sided spike, very reduced flowers, and the mature capsules in which the septum breaks away from the capsule wall. Our single repre- sentative is B. Brownei (B. pyramidatum (Lam.) Urban; cf. Bremekamp, 1938, p. 149), 2% = 34, which occurs as an occasional naturalized weed in extreme southern Florida, usually in damp soil and in the margins of mangrove areas (Lakela 26876, GH, USF). The species can be easily recognized by its dense, conspicuously bracteate, 4-sided spikes and small flowers. It is widely distributed throughout the American tropical regions in open cultivated or waste places, in both dry and moist soil, from eastern and southern Mexico through Central America and the West Indies to northern South America. It has been introduced as a weed in the Old World tropics. The intergeneric relationships of Blechum are uncertain. Nees (1847) placed the genus in his subtribe Dicliptereae, believing it to be allied to Dicliptera and Rungia on the basis of the detachable septa in mature capsules of all three genera. Nees’s reference to the occurrence of Blechum in Madagascar and the East Indies is unclear, since he does not record any of the eight species described as having been found there. Bentham & Hooker placed the genus in the subtribe Petalidiinae, while recognizing that its three to many ovules per locule made it dissimilar to other genera 1970] LONG, GENERA OF ACANTHACEAE 285 in the group. Lindau classified Blechum in his tribe Petalidieae together with four other genera, but Bremekamp & Nannenga-Bremekamp (1948) removed it to a subtribe of its own because of its racemose inflorescence and peculiar pollen. It should also be pointed out that they stated that the separation of Petalidiinae, Blechinae, and Ruelliinae is based only on differences in the morphology of pollen. Melchior has gone a step farther and recognized the tribe Blecheae (including only Blechum) and has placed it near the Barlerieae. Grant reported that Blechum Brownei has 34 somatic chromosomes, as does Ruellia, and said that karyology and chromosome number indicated affinity of the two genera and provided little evidence for their separation. Satellite chromosomes were observed in both genera, with the satellites attached by a slender thread to the short arm of a chromosome with a subterminal centromere. He suggested that Blechum may have been de- rived RY pti sees of plants with basic chromosome numbers of « = 8 and x = 9. Leonard (1951) also emphasized the similarity of Blechum and Rueliia, pointing out that the only conclusive difference between the two is the breaking away of the septum from the mature capsule in Blechum, while it remains intact in Ruellia. The type species of Blechum was first thought to be a Rwellia because of its close morphological similar- ity to that genus Oersted recognized in Blechum two subgenera based on morphology of the calyx and bracts, BLecHum (subg. Eublechum Oerst.) and CHILO- BLECHUM, which were treated as sections by Lindau (1895) with the species about equally divided between the two. In his treatment of Acan- thaceae of the West Indies Lindau (1900) recognized three species of Blechum, with the others restricted to Central America and Mexico, but both sections are represented over the range of the genus. REFERENCES: Under eorest apie se see BAILLON (p. 433), BENTHAM & HOOKER, BREME- NANNENGA-BREMEKAMP (1948), GRANT, LEONARD (1951), LINDAU AMP (1895), Metecnee Nees (1847), and Raj. Jussieu, A. L. pe. Mémoire sur le Dicliptera et le Blechum, genres nouveaux de plantes, composés de plusieurs a auparavant réunies au Justicia. Ann. Mus. Hist. Nat. Paris 9: 251-271. 1807. LINDAU, G. Ppt ince uae S nc Symb. Antill. 2: 170-250. 1900. [ Blechum, 185~ 188. ] OERSTED, icos og Centralamerikas acanthaceer. Vidensk. Medd. A. S. Mex Naturl. For. Kisbechon 6: 113-181. 1854. [Subg. Chiloblechum. | Subtribe Ruelliinae Bremek. 5. Ruellia Linnaeus, Sp. Pl. 2: 634. 1753; Gen. Pl. ed. 5. 238. 1754. Perennial caulescent herbs [or shrubs], usually pubescent. Leaves ses- sile or petioled, mainly entire, undulate [rarely dentate]. Flowers large 286 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 and showy (usually), or small and cleistogamous, solitary, or more often clustered in the leaf axils, or borne in terminal or axillary cymes or cy- mose panicles; bracteoles sessile or subsessile, narrow, small [or sometimes oblong to lanceolate, herbaceous, imbricate, and conspicuous]. Calyx deeply 5-parted or cleft, the segments linear or narrow, acute, subequal {or posterior lobe larger, or 2-lipped]. Corolla infundibuliform, purple, lavender, white [red, rose, or yellow], sometimes saccate; tube short to long (sometimes quite long), narrow below, straight or oblique; limb of 5 obtuse, spreading lobes, nearly regular |or 2 posterior lobes connate at the base forming a lip], aestivation convolute; cleistogamous flowers with corolla small, tubular, closed. Stamens 4, rarel 5, didynamous, adnate below the throat, included | or exserted | : anthers parallel, 2-loculate, oblong-sagittate, muticous; filaments dilated at the base, anterior filaments larger; pollen 3-porate, spheroidal, from 47 to 99 » in diameter. Disc in- conspicuous |or frequently conspicuous]. Style slender, usually slightly recurved at apex; stigma simple, subulate or 2-lobed, the lobes equal or posterior lobe short or obsolete; ovules 2-10 in each locule, anatropous. Capsules linear-oblong, narrow, or clavate, somewhat flattened, base very short or contracted into a solid, elongate stipe. Seeds 4-20, plano-com- pressed, suborbicular, with a mucilaginous coat that exhibits, after moisten- ing, tapered, short hairs with walls marked internally by rings or spirals; retinacula elongate, acicular, hooked. 2” = 34. (Including Aphragmia Nees in Lindl., Arrhostoxylum Nees in Mart., Cryphiacanthus Nees, Dip- teracanthus Nees.) LecrotyPr species: Ruellia tuberosa L.; see Britton & Brown, Illus. Fl. No. U.S. ed. 2. 3: 241. 1913. (Named in honor of Jean de la Ruelle, 1474-1537, a French physician and herbalist. ) The second largest genus of Acanthaceae with about 250 species dis- tributed throughout the tropics and subtropics ; relatively few species in the temperate regions of North America; six indigenous species and three introductions in the southeastern United States. The genus has an inter- esting quadricentric distribution that corresponds in general to that for the family, with centers of variation in Brazil, Indo-Malaya, Central America, and Africa. The circumscription of Ruellia adopted here is es- sentially that of Lindau (1895). The genus has long been recognized as taxonomically very difficult, and Lindau freely admitted that his intrage- neric classification was tentative Our most important species. is the polymorphic Ruellia caroliniensis (J. F. Gmel.) Steud. subsp. caroliniensis, sensu stricto (R. hybrida Pursh, R. parviflora sensu Small), which occurs throughout the Southeastern States in a wide variety of habitats, such as bottomlands, sandy woods, limestone soils, old fields, disturbed sites, scrub vegetation, etc. Fernald described six varieties, in addition to the typical one, and five forms, pri- marily on the basis of slight differences in leaf shape and on pubescence patterns. These appear to be of doubtful taxonomic value, since they describe minor variations induced by environmental modification or by small genetic differences. In southern peninsular Florida occurs an en- 1970] LONG, GENERA OF ACANTHACEAE 287 fee, oS a Veo i WwW Riess Chay, UAAY iv p ili f} i fl — i ai° Va ss Fic ellia. a~j, R. caroliniensis var. heteromor pha: a, flowering and fruiting ‘ranch, % ey ~ portion of leaf, showing trichomes and cystoliths, x 12; c, flow bud, ; d, flower with five stamens, corolla partly laid open, ae Repti fron a " flower with four stamens and st taminodium, < 3; f, g, two views of anther, < 8; h, stigmata, X 8; i, ovary in ecco section to show engin and ovules te one locule, X 12; j, n ve two n — mature fruits, X 2. : k, open capsule iiondite retinacula, one aborted seed remaining in right. hand valve, oF with covering of hygroscopic hairs, after moistening, x 6 Il hair’ (ab out half as long as on from seed coat, showing ers spiral sae annular thickenings, < 200: n, embryo from moistened seed, x 288 JOURNAL OF THE ARNOLD ARBORETUM [von. 51 demic variety, R. caroliniensis subsp. caroliniensis var. succulenta (Small) . W. Long? (R. succulenta Small), a distinctive ecotype generally re- stricted to the Everglades region. Ruellia caroliniensis subsp. ciliosa (Pursh) R. W. Long® is found in dry pinelands and scrub vegetation from South Carolina to central Florida, thence west to southeastern Louisi- ana. Hybrid populations of the two subspecies are frequent in northern and central Florida in disturbed sites. A second endemic variety, R. caro- liniensis subsp. ciliosa var. heteromorpha (Fern.) R. W. Long,* occurs in sandy pinelands and in disturbed sites from Volusia and Lee counties, Florida, south into the Keys. Hybrid populations of the two endemic varieties are not uncommon in southern Florida. Fernald also described a number of new but doubtiully important varieties in R. humilis Nutt., a highly variable species that is found only in the western part of our area, in open fields, dry prairies, and rocky soil from Iowa and Kansas south to Texas, extending into western Virginia, Tennessee, Alabama, and western Florida. Some introgression between R. humilis and R. caroliniensis occurs in their area of overlapping dis- tribution. Ruellia noctiflora (Nees) Gray is endemic to wet pinelands and savannahs from southwestern Louisiana to northern Florida and eastern Georgia; it is one of our most distinctive species because of its elongate corolla tube and capsule. Genetic evidence (Long, unpublished) indicates that it is related to the R. humilis complex. The Ozarkian R. pedunculata Torr. ex Gray subsp. pedunculata extends from southern Illinois south into western Louisiana, eastern Texas, and eastern Okla- homa in calcareous or rocky soils. A disjunct subspecies, R. pedunculata subsp. pinetorum (Fern.) R. W. Long,® a smaller and smoother form but with the characteristic elongate peduncles of subsp. pedunculata, oc- curs in low pinelands of South Carolina, northern Florida, and Louisiana. Ruellia strepens L. (including R. strepens f. cleistantha (Gray) McCoy) reaches its greatest development in the Mississippi basin and adjacent areas and extends into the Carolinas and Virginia, principally in bottom- lands and on calcareous soils. A related taxon, R. Purshiana Fern. (in- cluding R. Purshiana f. claustroflora Fern.) , is endemic to the mountainous vs nig ee subsp. caroliniensis var. succulenta (Small) R. W. Lon R. pee a Small, Bull. N.Y. Bot. Gard. 3: 437. V* Ruellia caroliniensis subsp. ciliosa ng, R. W. ee stat. nov. R. ciliosa Pursh, Fl. Am. Sept. 2: 420 Dipteracanthus ciliosus (Pursh) Nees, peat: 16: = a sensu Small, Fl. Southeast. U.S. 1084. oa re Man. Southeast. FI., 933, non Nutt. “Ruelli caroliniensis sgh ciliosa (Pursh) R. W. Long var. heteromorpha Long, stat = hetoromorpha & ernald, ‘Rhodo ra 47: 32. hybrida sensu Small, Fl. Southeast. us. 0 1903, non Pursh, Fl. Am. Sept. V® Ruellia pedunculata Torr. ex Gray subsp. pinetorum (Fern.) R. W. Long, stat. nov. Ruellia pinetorum Fernald, Rhodora 47: 24. 1945. 1970] LONG, GENERA OF ACANTHACEAE 289 region and Piedmont plateau from western Maryland south to Alabama, in woods and on calcareous or acid soils. The morphology of the plant sug- gests that it may be of hybrid origin involving R. strepens and R. caro- liniensis, although Uttal reported that the two species grow in adjacent areas and apparently do not hybridize. All these species represent sect. DipTreRACANTHUS (Nees) Lindau. A second group of species in our flora has been introduced into Florida from tropical America. The Mexican and Antillean R. Brittoniana Leonard emend. Fern. (R. malacosperma sensu Small, non Greenman), of sect. PHYsIRUELLIA Lindau, has escaped from cultivation and is nat- uralized in waste areas and around former homesites in southern F lorida, possibly in Louisiana, and also in Texas. A single collection of R. mala- cosperma Greenm., of sect. RuELLIA, from Key West, Florida, indicates that this plant is locally escaped. Apparently R. Lorenziana Griseb., an Argentinian species of sect. RUELLIA related to R. tuberosa L., is a new addition to our flora that is locally naturalized near Homestead, Florida. Although Britton & Millspaugh in their Bahama Flora state that R. tuberosa occurs in the southern United States, no specimens of this com- mon tropical weed have been seen from our area. Ruellia is the only acanthaceous genus for which there is some infor- mation about interspecific relations as determined by hybridization tests. Probably one of the first interspecific hybrids was made by Miiller, who successfully crossed two Brazilian species, R. formosa and R. silvaccola. Morphologically intermediate hybrids were produced by almost 50 per cent of the seeds formed from crossing experiments. Bowden reported a sterile hybrid, R. ciliosa (= R. humilis Nutt.2?) * R. tuberosa. Long success- fully crossed R. caroliniensis with R. humilis and suggested that natural hybridization may be a factor causing the observed intergradation of these species in their area of overlapping distribution. From a study of breed- ing systems in populations of southern Florida, he concluded that the four taxonomic species involved are ecological races of a single polytypic species, R. caroliniensis. An investigation of interspecific hybrids resulted in the production of 22 different F,; hybrids involving 12 North American taxa. Intersectional crosses, such as R. brittoniana R. occidentalis, pro- duced partially or completely sterile but sometimes vigorous hybrids. Ab- normal meiosis in these hybrids may account for the high sterility. The breeding systems in both the eastern species and the Texan “panicu- late’ species centered on R. nudiflora involve showy, chasmogamous flowers and small, tubular, cleistogamous flowers. Chasmogamous flowers favor outcrossing both structurally and by regular proterogyny. Generally, cleistogamous flowers are produced more abundantly towards the end of the growing season in the eastern species (at the beginning, in the panicu- late species) producing numerous fruits with good seeds. The total evi- dence suggests that local populations are predominantly autogamous, par- ticularly in the northern portion of our range, but that outbreeding oc- curs frequently in ulations from Florida. Long fa oa no revision of the genus ought to be attempted 290 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 until a thorough investigation of the reproductive biology of Ruellia spe- cies has been completed. Genetic barriers to interspecific crossing are lacking between many species, although these same species have been placed in different genera by various authors (especially Bremekamp). If it can be assumed that crossability is correlated with evolutionary rela- tionship, the broad generic interpretation of Lindau may be a more ac- curate reflection of the natural limits of Rwellia than the numerous segre- gate genera advocated by Bremekamp. Embryo sac development and microsporogenesis have been studied by Mauritzon, Maheshwari & Negi, Narayanan, Mohan Ram, and Wall & Long. No walls form after the first reduction-division during pollen forma- tion in Dipteracanthus patulus; quadripartition occurs by furrowing with tetrads decussate or isobilateral. The single-layered nucellus is completely absorbed by the time the four-nucleate stage of the embryo sac has been reached. Development of the embryo sac is of the Polygonum type, with the tip of the sac extending into the micropyle. Both synergids and antip- odals are ephemeral, but pollen tubes persist, and remnants can be recognized even in mature seeds. The endosperm is of the cellular type and is composed of a two-nucleate micropylar haustorium, a four-nucleate chalazal haustorium, and a central chamber that later separates into a cellular upper portion and a free-nuclear lower portion. A notable feature is the occasional formation of twin embryos. Additional embryos may arise by the proliferation of some of the suspensor cells. In an investigation to determine whether there are embryological dif- ferences between Ruellia tuberosa and Dipteracanthus patulus, Mohan Ram concluded that, in general, both show the same type of embryology, although twin embryos were not found in R. tuberosa. The only notable difference was the presence of a cellular basal apparatus, a structure ap- parently absent in D. patulus. Chromosome numbers may be different for the two genera, however. Kaur and De both report 2n = 44 for D. pro- stratus; the somatic number for R. tuberosa is 2n = 34. In any event, the investigations thus far in the genus clearly demonstrate the usefulness of experimental approaches to the definition of both genera and species. Hybridization tests may yet prove to be a more satisfactory means of defining natural groups than pollen characters or floral morphology within Ruelliinae. The genus has little economic importance. A number of tropical species are popular cultivars, mostly in greenhouses. Ruwuellia Devosiana Morr. and R. Makoyana Hort. both have purplish leaves, usually with attractive white venation; the former has white corollas marked with purple or lilac, the latter bright reddish corollas. The bright-blue-flowered R. squarrosa Hort. is grown in southern Florida as a ground cover and rock garden lant. Members of the subtribe Ruelliinae have pollen with three equatorial pores, or rarely four to seven evenly distributed pores, or “sparsiporous,” alveolate, echinulate or granulate exines. In his key to the genera of Ruelliinae of the Malay Archipelago and New Guinea, Bremekamp in- cluded eight genera, four of which were included as synonyms by Lindau 1970] LONG, GENERA OF ACANTHACEAE 291 in his treatment of Ruellia, and two of which are new. The recognition of segregates from Ruellia is based largely on inflorescence patterns, corolla morphology, and pollen sculpturing. For instance, Ruellia is said to differ from Arrhostoxylum in having pollen with sinailer, shallower, more regular alveolae. Ruellia was emended by Bremekamp and restricted to the four species malacosperma Greenm., a circumscription coinciding with the original delimitation of Cryphiacanthus Nees. In contrast, Lindau (1895) used a broad, inclusive definition of Rwellia, including 19 segregate genera. He recognized nine sections (four monotypic) based on morphology of the capsule and the flower. His sect. RUELLIA is entirely American and cor- responds with Ruellia L. emend. Bremek. His sect. DripreERACANTHUS (Nees) Lindau corresponds in part to Dipteracanthus Nees emend. Bre- mek., although Bremekamp restricted the genus to paleotropic species with sparsiporous pollen and subringent corollas. It is not clear from Breme- kamp’s key in what genus our species would fall, although he stated that the closest allies of Ruellia L. emend. Bremek. are Aphragmia (based on Ruellia inundata HBK.) and Arrhostoxyvlum (based on A. glabrum Nees), both restricted to tropical and subtropical America. Most of our species possibly would fall into Aphragmia, resembling Ruellia in pollen, inflores- cence, and seed characters, but differing in the clavate capsules with fewer seeds per capsule. Bremekamp also suggested that perhaps our species ought to be transferred to a new genus based on R. strepens L. REFERENCES: Under family references see BAILLON (p. 426), BENTHAM & HOOKER, BREME- KAMP (1965), BREMEKAMP & NANNENGA-BREMEKAMP, DE, GRANT, KAUR (1966), Lronarp (1951; 1959), Linpau (1895), inane ‘Mercatre & CHALK, Nees (1847), RAJ, SANTAPAU, SOLEREDER, and TCHOUP Diers, L. Der Anteil an Polyploiden in den ar der West-kor- dillere Perus, Zeitschr. Bot. 49: 437-488. 1961. [ R. floribunda Hook., ca. 28 = 34, Everett, T. H. Ruellia amoena. Addisonia 22: 55, FERNALD, M. L. Ruellia in the eastern United States. ie 47: 1-38; 47-63; 69-90. 1945. Forrste, A. F. Notes on structures adapted to cross-fertilization. Bot. Gaz. 13: 151-156. 1888. [Includes observations on a sp. of Ruellia. Horm, T. The root-structure of Spigelia marilandica L., Phlox ovata L. and Ruellia ciliosa Pursh. Am. Jour. Pharm. 78: 553-559. 1906. . Ruellia and Dianthera: an anatomical study. Bot. Gaz. 43: 308-329. pls. 11, 12. 1907. [General description of vegetative structures. ] LeConte, J. E. Observations on the North American species of the gen Ruellia. Ann. Lyc. Nat. Hist. N. Y. 1: 140-142. 1824. [Habitat ie on R. ciliosa and R. stre epens a Leonarp, E. C. Ruellia tuberosa and a few of its close relatives. Jour. Wash. Acad. Sci. 17: 509-520. 1927. Lonc, R. W. Convergent patterns of variation in Rwuellia caroliniensis and R. humilis (Acanthaceae). Bull. Torrey Bot. Club 88: 387-396. 1961. 292 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 . Biosystematic investigations in South scarp populations of Ruellia (Acanthaceae). Am. Jour. Bot. 51: 842-852. Artificial interspecific hybridization in oe (Acanthaceae). bid. 53: 917-927. 1966a. —. The artificial hybridization of the tropical species Ruellia Brittoniana < R. occidentalis and its taxonomic significance. Bull. Torrey Bot. Club 93: 181-187. 1966b. . Observations regarding the occurrence and relationships of pies Lorentziana (Acanthaceae) in southern Florida. Rhodora 68: 432- 1966c. —. Origin of the “dwarf Rwellia humilis” (Acanthaceae) populations of central Florida. Bull. Torrey Bot. Club 95: 16-27. 1968. [Apparently R. caroliniensis * R. ciliosa introgressants. | . Urrat. Some observations on flowering in Ruellia (Acanthaceae). Rhodora 64: eh Ties 1962. [Predominance of cleistogamous reproduction in natural populatio McCoy, S. A ran dings Ruellia. Am. Bot. 43: 22-24. 1937. [R. strepens var. cleistantha Gray, an autumnal form of the sp. MauesHwart, P., & V. Nect. The embryology of Dipteracanthus patulus (Jacq.) Nees. Phytomorphology 5: 456-472. 1955. [Includes extensive bibliography. | Mouan Ram, H. Y. Postfertilization studies in the ovule of Ruellia tuberosa Linn. Lloydia 23: 21-27. 1960 Miter, F. von. Mischlinge von Rwellia formosa und silvaccola. Abh. Naturw. Ver. Bremen 12: 379-387. 1892. Prain, D. Ruellia Devosiana. Bot. Mag. 137: pl. 8406. 1911 STANDLEY, P. C. Two additions to the ia of Louisiana. Torreya 14: 21-24. 1914. [R. spectabilis = R. Brittoniana Leonard TuHarp, B. C., & F. A. BARKLEY. The genus Rvellia in Texas. Am. Midl. Nat. Urtat, L. J. Observations on Rwellia Purshiana (Acanthaceae) in Virginia. Castanea 30: 228-230. 1965. . A plant-insect relationship i in Ruellia L. (Acanthaceae) coincidental to Cobubatha, favoring seed production by later cleistogamous flowers. | . Seed differentiation in east North American Rwellia. Castanea 33: 254, 255. 1968. WaLL, W. E., & R. W. Lonc. Megasporogenesis and embryo sac development in Ruellia caroliniensis (Acanthaceae). Bull. Torrey Bot. Club. 92: 372- 377-1965. Warp, D. B, The genus Anonymos and its nomenclatural survivors. Rhodora 64: 87-92. 1962. [R. caroliniensis (J. F. Gmel.) Steud., 90. Wirzeur, R. L. The identity of Walter’s species of Anony Jour. Elisha Mitchell Sci. Soc. 78: 125-132. 1962. [R. cpaenaae vat F. Gmel.) Steud. ] Subtribe Petalidiinae Lindau emend. Bremek. 6. Dyschoriste C. G. D. Nees in Wallich, Pl. Asiat. Rar. 3: 75, 81. 1832. Perennial caulescent herbs [sometimes suffrutescent at base] with 1970] LONG, GENERA OF ACANTHACEAE 293 glabrous or (more often) pubescent, erect, ascending [or prostrate] stems. Leaves sessile, subsessile [or petioled], usually entire [frequently fas- cicled|. Flowers in axillary clusters or cymes, sometimes solitary [rarely racemose], purplish, blue, or less commonly white, subtended by linear to oblong foliaceous bracts and bracteoles. Calyx deeply 5-cleft, lobes usually linear or subulate-setaceous, ciliate. Corolla-tube incurved or straight, cylindrical, often slightly ampliate at base, limb oblique, spread- ing, nearly regular to distinctly bilabiate. Stamens didynamous, a short and a long filament connate below and adnate to corolla tube, pubescent; anthers 2-loculate, more or less distinctly mucronate or aristate at base, locules equal, oblong, parallel to slightly divergent; pollen 3-colporate, prolate or occasionally subprolate, 40-50 « 47 28 y. Stigma linear, anterior lobe somewhat flattened, oblique, posterior lobe rudimentary or absent; style filiform; ovary 2-locular, glabrous, ovules 2 or occasionally 1 in each locule. Capsule included in the persistent calyx, oblong-linear, 2—4-seeded, glabrous, separating with difficulty at maturity into 2 valves with 1 or 2 seeds within each valve held in position by the retinacula; capsule base contracted into a solid stipe; septum adnate. Seeds flat, orbicular or suborbicular, mucilaginous when wetted, hairs with faint rings or spiral thickenings within. (Including Calophanes D. Don, Apassalus Kobuski.) Lectotype species: D. depressa (Wall.) Nees; see Britton & Brown, Illus. Fl. No. U. S. ed. 2. 3: 240. 1913. (Name from Greek dys, difficult, and choristos, separable, in reference to the tenaciously coherent valves of the capsule. About 100 species widely distributed in the tropics and subtropics, with a few taxa extending into the warm-temperate regions of the world. About 40 supposed species are American, but it is probable that a monographic revision will reduce this number substantially. Two species and two varieties occur in our area. Dyschoriste humistrata (Michx.) Kuntze (Apassalus humistratus (Michx.) Kobuski) is locally abundant in rich, sandy loam, limestone materials, and low river terraces from north- central Georgia to central peninsular Florida. D. oblongifolia (Michx.) Kuntze var. oblongifolia, 2n = 30, is found on the Coastal Plain from South Carolina to Sarasota and Brevard counties, Florida, westward to Louisiana, in disturbed areas, sandy soil, pine barrens, and sandhills. It is quite variable both in leaf size and in pubescence of the stems and sepals. An endemic variety, D. oblongifolia var. angusta (Gray) R. W. Long,® is locally abundant in extreme southern Florida and the Florida Keys in moist pinelands, disturbed sites, especially burned-over areas, rocky roadsides, limestone rocks, and similar places. Nees (1832) established the genus by the transfer of species of ome authors from Ruellia and Justicia. The key characters used were “. . antherae bilocellatae, locellis seeoie aequalibus basi mucronatis. Cap- sula . . . prope a basi tetrasperma. . Dissepimentum adnatum.” Only fe pi tericgl oblongifolia var. angusta (A. Gray) . W. Long, comb. nov. alophanes angusta Gray, Synop. Fl. N. Am. 2: 324. 1 Posse angusta (Gray) Small, Fl. Miami 168. iE f Florida Keys 135. 1913. 294 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 a year later David Don described his genus Calophanes, based on Ruellia oblongifolia Michx., apparently collected in Georgia. Both genera were used in later botanical literature, Dyschoriste representing Old World and Calophanes, New World taxa. Nees accepted this arrangement, but Bentham & Hooker, and later Clarke, placed Dyschoriste in the synonymy of Calophanes. Kuntze revived the earlier name; he was followed by Lindau, who placed the genus in his tribe Strobilantheae. The complex problems in the typification of Dyschoriste have been dis- cussed by Kobuski, who monographed the American species. His choice yh hy ay by, SS VS Fic. 4, Dyschoriste. a—g, D. oblongifolia var. oblongifolia: a, flowering stem, x %; b node with two flowers, one bractlet removed from pedicel of flower at left, X 2: c, flower in longitudinal section, x 3; d, anther after shedding of pollen, yt ‘8; e, stigma, X 10; f, ovary in vertical section to show placentation, x capsule, is see ygroscopic hairs, after moistening, ate oe embryo from mes seed. Mae 1970] LONG, GENERA OF ACANTHACEAE 295 of Ruellia erecta Burm. as the type is based on Kuntze’s transfer of Ruellia erecta to Dyschoriste thereby making it the oldest known epithet in the genus. The choice presumes the accuracy of the identity and synonymy of the several taxa, including D. depressa Nees, that Clarke referred to Calophanes Nagchana Nees. Ruellia erecta is typified by a reference of Burmann to a plate and description by Van Rheede. Un- fortunately, the plant illustrated and described has alternate, lance-ovate leaves, annual roots, and an apparently one-lipped corolla. The calyx is described as four-parted, with two segments larger than the others. It is apparently not acanthaceous, but its true identity is uncertain. For the present the lectotype chosen by Britton & — seems to be the best available until the Old World plants can be examine Bremekamp (1955) transferred Dyschoriste pe Strobilantheae to his Petalidiinae, a subtribe characterized by axillary cymes that rarely com- bine in a terminal panicle, by a subactinomorphic corolla never provided with a row of hairs functional in retaining the style, and, most importantly, by pollen grains that are prolate and banded, with three of the bands in the equatorial plane, dilated, and containing a pore. Most of the genera of the subtribe are tropical African in distribution, but Dyschoriste is pantropical. Kobuski decided not to place the American species in sub- genera or sections because of the relative uniformity of morphological characters in the genus. Instead, he identified three distinct geographical areas where the species appear to be related: the southeastern United States (2 taxa); Texas and southern Arizona to the Isthmus of Tehuan- tepec (21 taxa); and South America (17 taxa). Kobuski segregated his West Indian genus Apassalus from Dyschoriste on the basis of the much smaller leaves, smaller flowers, fruit characters, and muticous anther appendages. He considered it intermediate between the hypothetical ancestors of Apassalus and Dyschoriste proper in his interpretation. Apassalus was described as having only two seeds per capsule, but only one species (of three) has two seeds. The absence of anther appendages appears to be the only consistent character for the genus, and this hardly warrants recognition of a new genus in the Acan- thaceae. The phylogeny and relationships of Dyschoriste are obscure. The large number of closely allied species was considered by Kobuski probably to have evolved from common ancestral stock. He proposed that the taxa of the southeastern United States possibly are off-shoots of the widespread and morphologically generalized complex of D. decumbens (Gray) Kuntze, of Texas, Arizona, and Mexico, that developed parallel to the Mexican D. Greenmanii Kob. and D. jaliscensis Kob. Another morphological rela- tionship is with the largely Texan D. linearis (Torr. & Gray) Kuntze. On the other hand, Kobuski thought that our southeastern taxa possibly are most closely related to his “Apassalus group” in the West Indies and have migrated into the United States from that direction. This theory assumes evolution of plants with large leaves, large flowers, anthers with appen- dages, and four-seeded capsules from plants with small flowers, muticous 296 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 anthers, and two-seeded capsules, a proposition that is difficult to accept. The absence of Dyschoriste in the Mississippi Valley does not rule out the possibility that the range of the genus prior to the Oligocene extended continuously across the southern United States. Possibly the distribution was then split into two parts by the encroachment of inland seas during later epochs of the Cenozoic. Such an interpretation would explain, at least in part, the relationship of our species with those of Texas and Mexico. REFERENCES: Under family references see BAILLON (p. 428), BENTHAM & HOOKER, BREME- KAMP (1955b, 1965), GRANT, Linpau (1895), NEES (1832, 1847), and Ray. ALEXANDER, E. J. Dyschoriste humistrata. Addisonia 18: 25. pl, 589. 1933. BREMEKAMP, C. E. B. List of the Acanthaceae collected in Celebes by Dr. W. Kaudern and Dr. G. Kjellberg. Sv. Bot. Tidskr. 42: 372-403. 1948. [In- cludes D. celebica Bremek., sp. nov., and notes on pollen differences in Dyschoriste and Hygrophila. ] KogpuskI, C. E. A new genus of the ae Ann. Missouri Bot. Gard. 15: 1-8. 1928. [Apassalus Kob., gen monograph of the American ied of the genus Dyschoriste. Ibid. 9-90. [Includes 12 new species. | Ropertson, C. Florida flowers and insects. Trans. Acad. Sci. St. Louis 25: 277- 324. 1927. [D. oblongifolia (as Calophanes), 285, 286; visited by long- and short-tongued bees, Diptera, and butterflies. J STERNON, F. Etude d’une plante médicinale mexicaine: Dyschoriste (Calo- phones) linearis Gray. Bull. Soc. Bot. Belg. 65: 14-21. 1932. [Anatomical study demonstrating relationship with Ruellieae. | Subtribe Hygrophilinae (Nees) Bremek. 7. Hygrophila R. Brown, Prodr. Fl. Nov. Holland. 479. 1810. Perennial [or annual] erect, ascending [or diffuse] caulescent herbs; stems glabrous, sparsely pilose jor villous], unarmed Jor with axillary spines]. Leaves entire, lanceolate to obovate. Flowers borne in axillary fascicles [or terminal heads, rarely solitary]; bracts linear [elliptic or lanceolate|, ciliate; bracteoles lanceolate [or oblong], shorter than the calyx. Calyx tubular, segments 5 [4], equal to subequal, more or less fused to above the middle. Corolla tube cylindrical, ventricose at the apex, slightly ampliate; limb bilabiate, posterior lobe erect, concave, 2- cleft or 2-toothed, anterior segment 3-lobed, white [or pale purple]. Stamens didynamous |or 2 fertile stamens and 2 staminodia], not exserted, adnate to the tube above the middle; filaments glabrous, each pair connate at the base by a membrane, barely dilated; anthers oblong, muticous or submucronulate, 2-loculate, anther sacs parallel, divergent-sagittate. Pol- len grains 3(4)-colporate, prolate spheroidal to subprolate spheroidal, varying in size from 35 X 33 » to 70 X 66 pw. Disc inconspicuous. Style long, pubescent; stigma subulate, incurved, posterior lobe abortive; ovules usually 6-8 in each locule [or rarely 2, up to 18]. Capsule narrow, oblong, 1970] LONG, GENERA OF ACANTHACEAE 297 nonstipitate, the partition adnate. Seeds 12-18 [rarely 4, up to 36], or- bicular-compressed; retinacula short, obtuse, acute. Typr species: Ruel- lia ringens L. = H. salicifolia (Vahl) Nees (H. angustifolia R. Br.); cf. T. Anderson, Jour. Linn. Soc. Bot. 7: 113. 1864. (Name from Greek hygros, wet, and philos, loving, a reference to the usual habitat of the plants. A pantropical genus of about 80 species primarily of the Old World tropics but with a number of species in tropical America. Nees’s treat- ment (1847) is the only account of the entire genus. He informally divided the 24 species into two groups based on the morphology of the inflorescence. Bentham and Hooker interpreted the genus broadly, re- duced a number of Nees’s segregate genera to synonymy, and did not recognize any subgeneric categories. Clarke divided Hygrophila into two subgenera, Hycropuita (§ Euhygrophila Clarke) and AsTERACANTHA (Nees) Clarke, based on the presence or absence of spines, on calyx morphology, and on the number of seeds per capsule. He stated that Hygrophila may have 40-100 minute seeds per capsule (surely a mistake! ) but cited no species with more than 32. Lindau (1895) recognized five sections, but Asteracantha Nees was treated as a separate genus. Our single representative belongs to the largest and taxonomically most dif- ficult section, HycropHita (§ Euhygrophila Clarke emend. Lindau), which is distributed from tropical Asia to tropical America. Hygrophila lacustris (Schlecht.) Nees, 2n = 32, occurs from south- western Georgia and western Florida to eastern Texas, with its greatest development apparently in the lower Mississippi River valley and delta region, It is also found in the West Indies. The plant is a weak-stemmed perennial herb with narrow, sessile leaves and sessile axillary clusters of small white flowers. Hygrophila lacustris appears to be related to the widespread tropical H. conferta Nees, of Mexico and Central America. Nees observed that H. lacustris differs from H. conferta only in having sessile, attenuate leaves and somewhat smaller corollas. Other relation- ships, also in tropical America, may include H. portoricensis Nees and H. brasiliensis (Spreng.) Lindau. Hygrophila polysperma (Roxb.) T. Anders., a native of the East Indies, has been reported by Reams as quickly estab- lishing itself in lakes at Richmond, Virginia, when introduced, but it is doubtfully naturalized. Although Hygrophila is pantropical, its relationships are apparently with Old World genera, particularly with the African Brillantaisia Beauv. and the largely East Indian Synnema Benth. Lindau suggested interge- neric relationships also with the African genera Eremomastax Lindau and Mellera S. Moore, but Bremekamp believed these genera properly belong in the subtribe Petalidiinae. The Hygrophilinae resemble the Petalidiinae in inflorescence characters, but differ in the bilabiate corolla and distinc- tive pollen sculpturing. Typically, Hygrophila produces pollen with three, rarely four, germ pores that have four grooves between two of the pores. Reams, in an examination of environmentally induced variations in the 298 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 morphology and ontogeny of two types of hydathodes (water stomata and water-secretory trichomes), found that submerged leaves have water- stomata but nonfunctional, recessed trichome-hydathodes, while aérial or emersed leaves produce functional, bulbous trichome-hydathodes and water-stomata. Hydathodes are replaced by simple hairs on the bracts. The differentiation of the three kinds of trichomes begins immediately after the second cell-division of the primordium. Other evidences of the essentially aquatic nature of this genus were noted by Solereder who described the typical lacunar, hydrophytic cortex and the peculiar appear- ance of the interlocking parenchyma cells around the air chambers. REFERENCES: Under family references see BAILLON (p. 430), BENTHAM & HOOKER, BREME- KAMP (1938, 1948, 1955b, 1965), CLARKE, Lrnpau (1895), Nees (1847), Raj, and SOLEREDER; under Dyschoriste see Bremekamp. BERTHOLDT, Fhe seins temple plant (Hygrophila sp.). Aquarium Jour. 29: 386, 38 Kunpvu, B. c rh morphology of the spines of Hygrophila spinosa T. Anders. Jour. Bombay Nat. Hist. Soc. 43: 678-680. 1943. [Spines are modified branches. | LANGAWANKAR, J. D., N. L. PHALNiKar, & B. V. BHIpE. Chemical investigation of Hygrophylia (sic) ried (N. 0. Acanthaceae). Jour. Univ. Bombay II. 13(5A): 15-1 MrEceE, J. a tide a nombres chromosomiques d’espéces d’Afrique o cidentale. Revue Cytol. Biol. Vég. 24: 149-164. 1962. [H. ai lekse (Nees) T. Anders. and H. spinosa T. Anders., 2n = 32.] RANGASWAMY, K. Cytomorphological studies in Asteracantha longifolia Nees (Hygrophila spinosa A. And.). Proc. Indian Acad. Sci. B. 14: 149-165. Reams, W. M., Jr. The occurrence and ontogeny “7 hydathodes in Hygrophila polysperma T. Anders. New Phytol. 52: 8-13. THORNE, R. F. Vascular plants Sahelian A A Susant from Georgia. Castanea 16: 29-48, 1951. [Includes H. lacustris. | StopoLa, J. Encyclopedia of water arly 368 pp. T.F.H. Publications, Jersey City, N.J. 1967. [H. polysperma, H. salicifolia, 269, 271, 272, 275.] Tribe JustiIctiEAE Nees emend. Bremek. Subtribe Odontonematinae Lindau emend. Bremek. 8. Asystasia Blume, Bijdr. Fl. Nederl. Indié 796. 1825 [1826]. Perennial herbs [or shrubs], [erect,] procumbent or clambering, pubes- cent [or glabrous]. Leaves [narrowly lanceolate or] ovate, entire or somewhat dentate. Inflorescences terminal simple or branched racemes {or spikes], often unilateral; flowers opposite or alternate, solitary or glomerate in axils or bracts; bracts linear to narrowly deltoid, bracteoles minute. Calyx lobes 5, linear-setaceous [or lanceolate], subequal. Corol- la purplish, blue, yellow, or white, infundibuliform, the limb 5-lobed, the tube straight or incurved, long, narrow or ventricose, the throat ampliate 1970] LONG, GENERA OF ACANTHACEAE 299 or campanulate; aestivation imbricate. Stamens didynamous, or occasion- ally the posterior pair reduced to filiform staminodia, included; anthers 2-loculate, muticous at base or mucronate, locules parallel, equal, approxi- mate, or unequal; pollen 3-porate (pleurotreme), prolate. Disc cupulate or annular. Style subulate; stigma obtuse or minutely 2-parted (or subcapitate); ovary with 2 ayeles per locule. Capsule stipitate, pubescent [or glabrous], elliptic, the base contracted and solid, adnate; seeds 2—4 or fewer by abortion. Seeds ovate or compressed orbicular, irregularly angular, rugose or tuberculate, glabrous, retinacula elongate, subulate. TYPE SPECIES: A. intrusa (Forssk.) Blume. (Name possibly from Greek asystatos, not holding together, in reference to the wide-spreading valves of the dehisced capsules. ) A genus of possibly 40 species (although about 70 have been described) native to the Old World tropics, chiefly in the East Indies, India, and Africa, and now thoroughly naturalized in the New World tropics. Asytasia is represented in our flora by A. gangetica (L.) T. Anders. (A. coromandeliana Nees), 2n = 50, 52, a native of India. The plant is apparently a rather recent introduction into southern Florida where it has pear to be identical or closely similar to those from the West Indies and other areas of tropical America where the species is used as a ground cover and has escaped from cultivation. The corolla color varies from pale bluish lavender to yellow or white with purplish darker lines on the limb. The genus is in critical need of monographic revision. Nees described 15 species in his treatment of Asystasia for DeCandolle’s Prodromus, and he included 7 varieties in the most variable species, A. gangetica. Bentham & Hooker enlarged the genus to include two groups of shrubs, Dicentran- thera T. Anders., from the African tropics, and Mackaya Harv., from South Africa. Lindau (1895) placed Asystasia in its own tribe, Asystasi- eae, on the basis of its distinctive pollen and reduced number of seeds per capsule, but Bremekamp, in his realignment of the Justicieae (1965), transferred the genus to his emended Odontonematinae. REFERENCES: Under family references see BAILLON (p. 459), BENTHAM & Hooker, BREME- KAMP (1948, 1965), Kaur, Leonarp (1953), Linpau (1895), NEES (1847), and Raj. AKAMINE, E. K. Germination of Asystasia gangetica L. seed with special refer- ence to effect of age . nig temperature requirement for germination. PI. Physiol. 22: 603-607. 1 De, A. Cytological investigations i in the wer! Le coups Sci. Cult. 32: 198, 199. 1966. [A. gangetica, n = 25, = Hooker, W. J. Asystasia coromandelion see Mag. 72: pl. 4248. 1846. MANceNor, S., & G. Mancenor. Nombres chromosomiques nouveaux chez diverse dicotylédones et sale one d’Afrique occidentale. Bull. Jard. 300 JOURNAL OF THE ARNOLD ARBORETUM [vox. 51 Bot. oe 27: 639-654. 1957. [22 = 26, a new number for 4. coro- mandeliana (= A. gangetica) a the Acanthaceae; see also KAuR. | a "C. R. Nuclear behavior and chromosomal aberrations in mitosis of Acanthus ilicifolius and Asystasia coromandeliana. Indian Jour. Genet. Pl. Breed. 11: 205-210. 1951 Subtribe Justiciinae Bremek. 9. Dicliptera Jussieu, Ann. Mus. Hist. Nat. Paris 9: 267. 1807, nom. cons Perennial herbs [or shrubs], erect, ascending, or diffuse, pilose, hirtel- lous [or variously pubescent], with branching, often lax stems more or less hexagonal in cross-section. Leaves petioled, blades lanceolate to ovate, entire (or undulate). Flowers sessile, solitary or clustered in contracted cymes, these forming erect, secund, en lax ves or panicles; bracts 2, frequently small, subulate; bracteoles 2 or 4, opposite in pairs and Calyx deeply 5-parted, the lobes linear-setaceous, equal or subequal, hyaline. Corolla red, violet, blue, pink, or white; tube narrow, resupinate, slightly ampliate; limb nearly regular or more often bilabiate, the upper lip entire, emarginate, or 2-toothed, flat or concave, the lower lip spread- ing, slightly 3-lobed; aestivation imbricate. Stamens 2, exserted but shorter than the upper lip, staminodia absent; anthers 2-loculate, anther sacs often unequal, muticous or the longer ones sometimes spurred; pollen 3-colporate, prolate or rarely perprolate, mostly 60 > 25 » in diameter. Disc cupular, oblique-truncate, or sinuate-dentate. Style filiform; stigma entire or minutely 2-toothed; ovary with 2 ovules per locule. Capsule ovoid to suborbicular, subsessile, contracted into a short, solid stipe; placentae separating elastically from the walls and rupturing on dehiscence. Seeds 4 [or 2] or fewer by abortion, lenticular, with smooth, muricate, or squamose surface; retinacula uncinate, broadened at the base, acute rs (Including Diapedium Konig, nom. rejic., Yeatsia Small, Gatesia Type species: D. chinensis (L.) Juss., typ. cons. (Name from Greck diklis, folding two ways, a double or folding door (7.e., having two valves), and pteron, wing, in reference to the two recurved, apparently winged valves of the dehisced capsule.) A large, taxonomically complex genus of possibly 150 species (although according to Leonard nearly 300 have been described) in the tropics, subtropics, and warm-temperate regions of the world, best developed in the East Indies, South Africa, and tropical South America. Three species in three sections occur in our Section DiciipTERA (§ Platystegiae Nees) is characterized by orbicular, ovate or oe bracteoles, the bases not cuneate. Dicliptera viridiflora (Nees) R. W. Long,’ an endemic species of southwestern V7 Dicliptera viridiflora (Nees) R. W. Long, comb. n Rhytiglossa viridiflora Nees in DC. Prodr. 11: 346; ore 1970] LONG, GENERA OF ACANTHACEAE 301 Georgia and adjacent Florida in rich, wooded bottomlands of the Flint and Apalachicola rivers, where it may be locally abundant, has white, pale blue, or pink, nearly regular corollas in compacted spikes with conspicuous, ovate, acuminate bracts. The species appears to be morphologically close to the West Indian D. martinicensis Juss. Section SPHENOSTEGIAE Nees, the largest subgeneric group, is char- acterized by floral bracts more or less cuneate at the base and is repre- sented with us by D. brachiata (Pursh) Spreng. var. brachiata (Diapedi- um brachiatum (Pursh) Kuntze, Diapedium attenuatum (Gray) Heller), 2n = ca. 80. Recognizable by its green, rather broad, wedge-shaped bracts, usually large, ovate-lanceolate to elliptic, long-petioled leaves, and strongly bilabiate, pink or reddish corollas, this erect herb is widely dis- tributed on the Coastal Plain from Virginia to Texas, inland to Arkansas, Missouri, and Kansas in river bottoms, hammocks, and low ground. It is variable in morphology of leaf, bract, and pubescence; differences in the latter caused Fernald to describe two minor varieties from Texas Section DAcTYLOSTEGIAE Nees is a small group of tropical American species characterized by the presence of two floral bracts larger than others in the involucel. Our representative is D. assurgens (L.) Juss. (Diapedium assurgens (L.) Kuntze), 2n = 80, a West Indian weedy species char- acterized by stiffly erect stems, and conspicuous bright red or scarlet bilabiate corollas in narrow, terminal or axillary lax panicled spikes. Two varieties, both occurring in peninsular Florida, have been distinguished. coral soil, and in hammocks. Less frequently, var. assurgens, characterized by subulate- attenuate bracts, can be found in extreme southern Florida, from Sanibel Island, Lee County, to the Florida Keys. here is some evidence that polyploidy has been an important cause of variation in Dicliptera. Grant pointed out that both D. assurgens and brachiata are tetraploids in comparison with the Central American D. resupinata, which Sigiura reported as having a gametic chromosome number of 20. Because of their morphological similarities, it is possible that D. brachiata may have been derived either as an autopolyploid or allopolyploid derivative of D. resupinata. Kaur has reported the chromo- some number of D. elegans as 2n = 20 and that of D. Leonotis as 2n = 48. Additional chromosome counts for Dicliptera species are very desir- able. In the only treatment for the entire genus, Nees (1847) established five sections (based on the morphology of the floral bracts), which were accepted, in general, by Lindau (1895). Initially, Bremekamp (1943) accepted Lindau’s placement of Dicliptera in the tribe Odontonemateae, Dicliptera Halei Riddell, New agen ee Hive Jour. 8: 756. “eo Gatesia laete-virens Gr. od Pro . Arts Sci. 13: 365. Justicia laetevirens Buckl hae oa an ie 176. 1843, not J. cee Vahl. Yeatsia laete-virens (Buckl.) Small, Bull. Torrey Bot. Club 23: 405-410. 1896. 302 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 subtribe Diclipterinae but later transferred the genus to his emended Justicieae. Intergeneric relationships appear to be closest with the Old World tropical genera Hypoéstes Soland. ex R. Br. and Peristrophe Nees, and possibly with Rungia Nees (although Bremekamp suggested that pollen and capsule characters indicate that Rungia ought to be in a differ- ent subtribe from Dicliptera). Pollen sculpturing apparently is not as useful a taxonomic tool in the Justicieae as in some other tribes of Acan- thaceae. Eight different pollen types have been reported from a single anther of D. javanica (cf. Davis)! Dicliptera needs a critical taxonomic review because the proposed inter- generic relationships and intrageneric alignments appear to be highly artificial, if not simply arbitrary. The present construction of subgeneric categories based on minor differences in the morphology of floral bracteoles does not seem to reflect any natural relationships of species groups within the genus. This emphasis on small differences has carried over into the description of species. Leonard included 29 species, 26 of them new, in his account of Dicliptera in Colombia. Morphological criteria for the separa- tion of many of these new species are minimal, and it is probable that future study will prove them unworthy of taxonomic recognition. REFERENCES: Under family references see BAILLON (p. 461), BREMEKAMP (1948, 1965), Davis, GRANT, LEONARD (1958), Linpau (1895), Nees (1847), and Ray. BREMEKAMP, C. E. B. L’identité du Jacobinia suberecta Andr. et la délimita- tion des Diclipterinae Lindau. Boissiera 7: 182-201. 1943 Evers, R. A. Illinois flora: notes on Limnosciadium, Dicliptera, and Iresine. Rhodora 64: 282, 283. 1962. [Second locality for D. brachiata in s. Minois.] FERNALD, M. L. Two varieties of Dicliptera brachiata. Rhodora 43: 287, 288. 1941. [Var. glandulosa (Scheele) Fern., Tex. to Vera Cruz, Mex., and var. Ruthii Fern., Tex. ] Gomez, D. M. Catalogo de las periantiadas cubanas, espontaneas y cultivados. Anal. Hist. Nat. 23: 267-302. 1894. [D. assurgens var. Vahliana. Kaur, J. Chromosome nueerat in Acanthaceae I. Curr. Sci. Bangalore 34: 295. [D. elegans Dalz., 2n = 20; D. Leonotis Dalz. ex Clarke, 2n = 48. SicturA, T. Studies i in the hens numbers of higher plants III. Cytologia 10: 205-212. 1939. [Includes chromosome numbers for Dicliptera. | 10. Justicia Linnaeus, Sp. Pl. 1: 15. 1753; Gen. Pl. ed. 5. 10. 1754. Perennial herbs [rarely shrubs] with erect or ascending, smooth or variously pubescent stems. Leaves petiolate, linear, lanceolate, or more often ovate to oblong, entire. Flowers small or moderately large, fascicled, or in lax terminal or axillary cymes, spikes or panicles [or solitary]; bracts various, often small, linear or subulate, distant, or sometimes both bracts and bracteoles conspicuous, large, imbricate. Calyx deeply 5 [or some- times 4]-parted, cleft nearly to the base, at aa subulate or linear, acute, nearly equal. Corolla white, red, pink, purple, sometimes with purple or white markings in the throat: a short [rarely long], straight 1970] LONG, GENERA OF ACANTHACEAE 303 or incurved, rather narrow, slightly ampliate above; limb bilabiate the upper lip erect or concave, incurved, spreading, frequently 2-dentate, external in bud, lower lip 3-lobed. Stamens 2, staminodia none, usually slightly exserted but not exceeding the corolla lips, adnate to the throat of the corolla; anther 2-loculate, anther sacs separated, more or less super- posed, one or both sacs apiculate or produced into a spur or tail, lobes parallel or obliquely affixed, connective broad or narrow: pollen grains 2-porate, bilateral, ca. 66 & 47 * 40 pn, 3-colporate (pleurotreme), sub- prolate, ca. 28 « 22 ». Disc cupulate or cyathiform, entire or sinuately 2- or 3- or S-lobed. Stigma entire, capitate, or minutely 2-dentate; style filiform; ovules 2 in each locule. Capsule ovate, oblong, clavate [or a lar], contracted at the base into a short, solid stalk. Seeds 4 [or 2] o fewer by abortion, plano-compressed, suborbicular, smooth, we as rugose, tuberculate or muricate; retinacula arcuate, acute, truncate below. (Including Dianthera L., Beloperone Nees, Rhytiglossa Nees, Drejerella Lindau.) Lecrorypr species: J. hyssopifolia L.; see Hitchcock & Green, Int. Bot. Congr. Cambridge 1930. Nomencl. Propos. Brit. Bot. 116. 1929. (Named in honor of James Justice, a Scottish botanist and horticulturist of the 18th sarin author of The British Gardener’s Calendar, 1754.) — WATER-WILLOW The largest genus in the family with about 300 species distributed throughout the tropics and subtropics of both hemispheres and extending into the temperate region of North America. The most widespread Justicia of our area is J. americana (L.) Vahl (Dianthera americana L.; including J. mortuifluminis Fern.), 2n = 26, the common water-willow of margins of ponds, lakes, rivers, and occasion- ally swamps throughout most of the eastern United States and probably even into Mexico. Justicia ovata (Walt.) Lindau var. ovata (J. humilis Michx.) occurs on the Coastal Plain in swampy woods, shallow water, and bottomlands from southern Virginia to northern Florida; var. lanceo- lata (Chapm.) R. W. Long,’ 2n = 14, 14 + 1, is found in wet soil and bottomlands from central Florida and Tennessee, west to Louisiana and Texas; var. angusta (Chapm.) R. W. Long,’ a narrower-leaved, smaller- flowered plant occurs as an endemic in central Florida, also in marshy places. Two other well-marked, endemic species also occur in Florida: J. crassifolia (Chapm.) Small is found in wet pinelands and glades in Gulf County at the mouth of the Apalachicola River, and J. Cooleyi Leonard & Monachino has been found in hammocks on limestones in Her- BN co ct ovata (Walt.) Lindau var. lanceolata (Chapm.) R. W. Long, comb. Pesach: ovata var. lanceolata Chapman, FI. So. U.S. 304. 1860. J. lanceolata (Chapm.) Small, Man. Southeast. Fl. 1231. 1933. : . hu Michx. var. lanceolata (Chapm.) Gleason, a erg 4: 24. 1952. ve 1 usticia ovata (Walt.) Lindau var. angusta (Chapm.) R. W. Long, comb. nov. nthera ovata var. angusta Chapman, Fl. So. U.S. 304. 1860. 7 pats (Chapm.) Small, Man. ‘Southeast. FI. 1231. 1933. 304 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 nando County. A Mexican species, J. Brandegeana Wasshausen & Smith 1” (Beloperone guttata T. S. Brandeg., Drejerella guttata (T. S. Brandeg.) Bremek.), has escaped and apparently become naturalized in peninsular Florida, especially in waste places, citrus groves, and around former habita- tions. This is the common “shrimp plant” that is grown widely for its attractive spikes of reddish-brown or greenish-yellow, ovate bracts and white flowers. Our species need careful revision, for it is apparent that some of the forms intergrade and may, therefore, be conspecific. Experimental hy- bridizations should be especially useful in determining interspecific rela- tionships, particularly in the J. ovata complex. It appears that our species are not closely related to tropical American taxa, although possibly J. americana may be allied to certain Mexican and Central American groups. Justicia ovata and J. crassifolia are possibly related to West Indian and Central American taxa, such as J. secunda Vahl and J. filibracteolata Lindau. Monachino and Leonard thought that /. Cooleyi is related to J. Pringlei Robins., of Mexico, and to J. galapagana Lindau, of the Galapagos Islands. The taxonomic history is so complex that only a general review, and that only with special reference to taxa of our area, is possible here. Nees described the North American species under his Rhytiglossa, a confused assemblage of dissimilar elements supposedly from South Africa and America. He placed the genus in his tribe Aphelandreae, but later Ben- tham & Hooker transferred the group to their Justicieae and included it under Dianthera. For the most part, Bentham & Hooker followed Ander- son in his general circumscription of Justicia, although he also included Adhatoda Medic. Bentham & Hooker recognized 11 sections in Justicia, but Beloperone and Dianthera were considered to be separate genera. It is not altogether clear from their descriptions where our species would be placed, but presumably they would be under Dianthera. Lindau (1895) divided Justicia into three subgenera, Justicta (subg. Eujusticia Lindau), GENDARUSSA (Nees) Lindau, and DiantHERA (L.) Lindau based on morphology of the inflorescence and anther sac and to some extent on pollen sculpturing. Subgen. Jusrict, the largest division, was further divided into seven sections, three of which contained American taxa. Our species all belong to subgen. D1antTHERA which supposedly is characterized by blunt or short-stalked anther sacs. Bremekamp (1948) was of the opinion that the generic circumscriptions of Anderson, Bentham, and Lindau were too vague and that their defini- tions of Justicia could not be held together any longer. He recommended that Justicia be confined to J. Ayssopifolia L., of the Old World, and - Pe Brandegeana Wasshausen & L. B. Smith, Fl. Illus. Catarinense 1(Acan): . 1969. ohare guttata T. ie fag rar Vers Calif. prs Bot. 4: 278. 1912; non Justicia guttata Wall. Pl. Asiat. Rar. 1: 24. pl. Calliaspidia guttata (T. 2 Brandes.) pale ae ght Akad. Wet. Afd. Natuurk. 2. 45(2): 54.1 Drejerella guttata (T. S. aes Bremek. Philip. Jour. Sci. 80: 14. 1952. Fic. 5. Justicia. a-g, J. ovata var. angusta: a, small plant with flower, x 14; b, flower and flower buds, x 3; c, anther, i : isc and ovary with one side removed to show four ovules, 4 eae e, oe fruit, one seed removed — -_ Eb a Oo La | = Qa ao c a eee pe¥] co ae p — | Oo 3S aa bt co 5 ee 3 ao Ee i=) 2g oO i=] g*] Q. = x ~_ ou , anther, from below, X 10; 1, p aially mature fruit and flower from which corolla has fallen, & 2. m,n, J. amer cana: m, corolla with stamens, from above, x 3; n, anther from below, m1 306 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Dianthera (based on D. americana L.) be restricted to the New World. There is no evidence from pollen morphology to suggest that Dianthera ought to be retained as a genus, although Grant thought there is some cytological basis for recognizing the genus. The slight morphological dif- ference for their separation is based on the position of the anther sacs, which are stalked obliquely at the tip of a branched connective in Dian- thera, while in Justicia one anther sac is borne horizontally or obliquely at the tip of the connective and one is borne perpendicularly, or nearly so, below it. In both instances the anther sacs are blunt, not pointed at the base, a character Lindau considered of sufficient weight to place all species of Justicia of the United States in his subgen. DIANTHERA. Leonard (1958) united both Dianthera and Beloperone under Justicia after point- ing out that continuous variation in anther characters made it impractical to maintain the separate genera. Bremekamp transferred Beloperone guttata to still another segregrate genus, Drejerella. Until additional evi- dence is available, a say interpretation of the genus appears to be more in keeping with the Inter- and wae relationships in Justicia have been the subject of a wide variety of investigations. Mohan Ram & Sehgal studied pollen- grain formation, embryo-sac development, and embryogeny in ]. simplex D. Don, a native of tropical Africa and Asia. They reported that chalazal haustoria generally are two-nucleate in Justicia, as in Beloperone and Adhatoda. No significant embryological differences were found between the three genera. Dicliptera and the African genus Barleria also resemble Justicia in the morphology of the chalazal haustoria, but Blechum, Ruel- lia, and Hemigraphis produce four-nucleate haustoria. The broad basal apparatus characteristic of the three latter genera is represented by a rudimentary, few-nucleate structure in Justicia Palynological and cytological investigations ‘of taxonomic relationships have generally been inconclusive. Raj was not able to find any differences between the pollen grains of Justicia and Dianthera. However, Bhaduri concluded that Lindau subordinated Adhatoda as a section in Justicia be- cause he misconstrued the morphology of the pollen of A. vasica (J. Adhatoda) and further concluded that the two genera should be separate. Bentham & Hooker maintained Adhatoda as a separate genus on the basis of other morphological grounds, but Leonard (1958), apparently on simi- lar bases, submerged Adhatoda in Justicia. Grant concluded there is suf- ficient evidence to retain Adhatoda as a separate genus because the somatic chromosome number typically is 2n = 54. Justicia species are known with 2m = 28, 32, and 36, suggesting three basic numbers: x = 7, 8, and 6 or 9. The same numbers are also found in Beloperone and Jacobinia. Grant suggested that aneuploidy is a probable cause for the diversity in chromo- some number in Justicia and that possibly related groups, such as Adha- toda, arose through allopolyploidy. The discovery of “B” chromosomes in J. ovata var. lanceolata (cf. Lewis et al.) may be evidence of the occur- rence of aneuploidy that perhaps was caused by fragmentation of chromo- somes or is a vestige of reduction from higher ploidy levels. Additional 1970] LONG, GENERA OF ACANTHACEAE 307 cytological studies should be useful in any future mene studies, but crossing experiments might be more helpful at this poi Unlike many Rvellia species, Justicia populations fay be largely if not entirely allogamous. Although no hybridization studies have been made, there is some evidence that outbreeding occurs regularly in J. simplex. Jain & Joshi conducted a genetic analysis of gene frequencies at a petal- color locus in local populations and concluded that the tendency for markedly local differentiation in these polymorphic plants could best be explained by considering them as partially isolated, panmictic populations. Additional investigations of causes of variation in the genus ought to in- clude artificial hybridization experiments to ascertain whether barriers to crossing occur between species. It is possible that some of the observed patterns of variation in our species are caused by natural hybridization. Future work on Justicia may result in additional evidence that would provide a basis for splitting up the genus into two, three, or more segregate genera. At the present time the evidence for this course of action is equivocal, but it seems clear that most of our southeastern species are more closely related to one another than they are to other taxa outside of the range of the Flora. If segregate genera are to be recognized they would be Dianthera L. (based on D, americana L.), Adhatoda Nees (based on Justicia Adhatoda L. = A. vasica Nees), and Justicia (based Justicia species have also been used in various other biological and chemical investigations. Anatomical and morphological studies have been reported by Sterling, Jones, e¢ al.; studies of toxins, alkaloids, and other chemical constituents have been carried on by Munakata e¢ a/. and Bhat- nagar & Popli; and pollination studies by Buxbaum. Many of the attrac- tive cultivated plants that pass under the name Justicia Cage belong to the tropical American genus Jacobinia Nees ex Moric REFERENCES: acs family references 8 BAILLON (pp. ti hae 445), BENTHAM & Hooker, BREMEKAMP (1948, 1965), ELLIs, HARTMANN, JOSEPH, LEONARD (1958, “t959), LINDAU Gai, Nees (1847), PAL, “ei and SOLEREDER Benorst, R. Contribution a la connaissance des Justicia de seandiae Syst. Paris 13: 118-131. 1947. [Describes 13 new spp. BHATNaGaR, A. K., & S. P. Porpzt. Mass fragmentation of the alkaloids of Adhatoda vasica Nees. Indian Jour. Chem. 4: 291, 292. 1966. [Spectro- scopic characteristics of vasicine and vasicinol. | BREMEKAMp, C, B. E. Philippine Acanthaceae, I. Philip. Jour. Sci. 80: 11-21. 1952. [Drejereila vs. Beloperone, 13, 14 Buxsaum, F. Die Biene als Honigdieb bei Justicia Adhatoda L. Biol. General. ee 104-106. 1926.* FERNALD, M. L. Justicia umbratilis, sp. nov., J. humilis Michx., J. americana var. subcoriacea, var. nov. In: Another century of additions to the flora of Virginia. Rhodora 43: 639-642, pl. 693. 1941. [See also Rhodora 44: 92. 942: J. umbratilis Fern. = J. mortuifluminis Fern., nom. nov., not J. umbratilis S. Moore, 1913.] Not. 308 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 & B. G. Scuupert. Studies of American types in British Herbaria. Part IV. Some species of Thomas Walter. Rhodora 50: 190-208, 217-229. pls. 1103-1115. 1948. [Dianthera ovata Walt., 224-226; J. ovata (Walt.) Lindau replaces J. humilis Michx. Hom, T. Ruellia and Dianthera: an anatomical study. Bot. Gaz. 43: 308-329. pls. 11, 12. 1907. [General description of vegetative structures; Ruellia has monostelic structure, Dianthera polystelic. | Jan, S. K., & B. C. Josut. Local differentiation in some natural populations of Justicio swniples: Genetics 47: 789-791. 1962. Jones, W. R. The development of the vascular structure of Dianthera ameri- cana, Bot. Gaz. 54: 1-30. pls. 1-4. 1912. [Seedlings are monostelic at first, later peripheral meristeles develop. | Lammers, W. T. Accumulation of radioisotopes of Justicia americana L. in the Clinch River. (Abstr.) ASB Bull. 8: 23. 196 Lewis, I. M. A black knot disease of Dianthera americana L. Mycologia 4: 66-70. 1912. [Bagniesiella Diantherae, sp. nov., an Ascomycete. | Lewis, W. H., H. L. Strrptinc, & R. G. Ross. Chromosome numbers for some angiosperms of the southern United States and were Rhodora 64: 147- 161. 1962. [J. ovata var. lanceolata,n = 14; n = 14+ 1, 2n = 29.] Mowan Raa, H. Y., & P. P. SEHGAL. The Biehistary st Justicia simplex Don. Phytomorphology 8: 124-136. 1958. MonacuHino, J., & E. C. LEonaArD. A new emma of Justicia from Florida. Rhodora 61: 183-187. 1959. [J. eng Munakata, K., S. Marumo, K. Onta, & Y. CHEN. Justicidin A. and B., fish-killing ‘onntiponents of pies Hayatai var. decumbens. eta, Lett. 47: 4167-4170. 1965.* Nac, N. C., & K. N. Bose. Chemical examination of some Indian medicinal plants. Trans. Bose Res. Inst. Calcutta 6: 201-211. 1931. [Includes J. PENFOUND, W. T. The biology of Dianthera americana L. Am. Midl. Nat. 24: 242-247. 1940. [In connection with mosquito control. Raj, B. Pollen morphology of two species of Acanthaceae. Curr. Sci. Bangalore 319, 320. 1965. [J. prostrata (Clarke) Gamble and Rungia repens Nees. Ropertson, C. Flowers and insects. VII. Bot. Gaz. 17: 65-71. 1892. [J. americana (as Dianthera), 65, 66; pollinated by Apidae and Andrenidae, but also visited by syrphid flies and by butterflies. ] ScHULTES, R. E., & B. Hotmstepr. De plantis toxicariis e Mundo Novo trop- icale commentationes II. Rhodora 70: 113-160. 1968. [J. pectoralis Jacq. var. stenophylla Leonard used in preparation of a narcotic snuff from Virola theiodora. | SEN, P. K., T. Bosr, & R. N. Bose, Effects of boron and molybdenum on root- ing of semi- -hardwood cuttings of Justicia Gendarussa L. Indian Jour. Pl. Physiol. 2: 21-28. 1959 oe L, H. Dianthera sila L. var. subcoriacea (Fern.) Shinners, mb. nov. Field Lab. 25: 79, 80. 1957. [Accepts Bremekamp’s circum- atise of Justicia. | Sms, J. Justicia americana. Bot. Mag. 50: pl. 2367. 1822. SMALL, J. K. Dianthera casi Addisonia 3: 79, 80. pl. 120, 1918. [Plant illustrated is J. ovata var. angus . Justicia Runyonii. "Tbid. 15: 29, 30. 1930. [From Texas. | 1970] LONG, GENERA OF ACANTHACEAE 309 SMITH, E. B. I.0.P.B. Chromosome number reports VII. Taxon 15: 155-163. 1966. [J. americana, 2n = 26. STERLING, C. The primary body of the shoot of Dianthera americana. Jour. Bot. 36: 184-193. 1949. [Developmental anatomy with reference pei stelar concept; includes references. | TURNER, B. L. Justicia Warnockii (Acanthaceae), a new species from Trans- Pecos Texas. Field Lab. 19: 100, 101. 1951. [Shrubby perennial, vegeta- tively unlike any other sp. of Justicia in U.S.] DEPARTMENT OF BoTANy & BACTERIOLOGY UNIVERSITY OF SOUTH FLORIDA TAMPA, FLORIDA 33620 310 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 FLORAL MORPHOLOGY AND VASCULAR ANATOMY THE HAMAMELIDACEAE: THE APETALOUS GENERA OF HAMAMELIDOIDEAE A. LINN BoGLE THe HAMAMELIDACEAE ARE A FAMILY of twenty-seven genera and pos- sibly 100 species distributed in the tropical, subtropical, and temperate regions of both the New World and the Old. The genera are small, four- teen are monotypic, six contain only two to three species, while seven comprise from five to fourteen species. Most of the genera are narrow endemics or very restricted in their distribution, only two (Hamamelis, Liquidambar) being present in both Asia and North America. The genera are morphologically diverse, ranging from huge evergreen trees to low deciduous, rhizomatous shrubs. Their leaves vary from palmately lobed and veined to elliptic-oblong and pinnately veined and may be spirally arranged, two-ranked, or opposite. They are stipulate in all genera except one (Rhodoleia). Inflorescences vary from simple racemes to panicles, spikes, heads, or densely congested and fused, complex, capitate flowering branch-systems. The flowers may be showy or inconspicuous, petaliferous, apetalous or naked, bisexual or unisexual, with few to numerous stamens, many ovules to only one, and few winged seeds per carpel to mostly a single wingless seed. Perhaps the one unifying morphological feature of the family is in its characteristic 2(—3)-carpelled, septicidal (and some- times partially loculicidal as well) woody, capsular fruits. The morphological, systematic, and phyletic relationships of the Ham- amelidaceae were last reviewed in detail forty years ago (Harms 1930). Many of the problems encountered then concerning morphological inter- pretations and systematic or phylogenetic affinities have remained unre- solved up to the present time. Furthermore, in the intervening years, the concept of the family has been expanded through the discovery or recogni- tion of new genera which have increased its diversity, rather than bridged its internal gaps. Indeed, the diversity of the family is such that some consider two or more families to be represented, rather than one. The limits of some tribes, or the tribal affinities of a few genera are uncertain, while the morphological nature of certain floral organs has evaded satis- factory interpretation in some cases, or has been widely misinterpreted in others. These problems have prompted several workers, in recent years, to point out the need for more detailed study of the group. Foremost among the areas in need of examination is that of floral vascular anatomy, which has only recently come under scrutiny (Bogle 1967, 1968, 1969; Endress 1967; Horne 1914). Evidence from such studies in other groups has been very valuable in the interpretation of difficult problems of floral morphology, which is among the principal bases on which systematic and phylogenetic relationships may be established. 1970] BOGLE, HAMAMELIDACEAE 311 This paper is the first in a series which will provide a detailed compara- tive account of the floral vascular anatomy, coupled with other observa- tions on floral and inflorescence morphology, for each of the 27 known genera of the family. The ultimate purpose of this work is the clarification of the morphological homologies of the organs of the hamamelidaceous “flower,” and of the phylogenetic trends among the divergent floral types within the family. Such information, along with evidence from other lines of research, should then permit a re-evaluation of the interfamilial and phylogentic relationships of the Hamamelidaceae. The opinions expressed in the literature concerning this family differ widely. It has been thought to be 1), related to several extant magnolialian families, or directly derived from a magnolialian or proto-magnolialian stock (Solereder 1899; Hallier 1903, 1912; Tippo 1938; Tang 1943; Chang 1961; Cronquist 1968; Takhtajan 1969); or 2), derived from a rosalian or proto-rosalian stock, with close affinity to the extant Rosaceae, Saxifragaceae, or more especially the Cunoniaceae (Bentham & Hooker 1865; Baillon 1871, 1874; Eichler 1875; Reinsch 1889; Niedenzu 1891; Bessey 1915; Hutchinson 1926, 1959; Tong 1930; Harms 1930; Melchoir 1964; Endress 1967; Jay 1968); or 3), to be derived from an amentiferous ancestral stock (Wett- stein 1935; Croizat 1960, 1964). On the other hand, many of the authors cited above derive from a proto-hamamelidalian stock the highly special- ized and advanced group of families included in the “Amentiferae” (par- ticularly the Betulaceae, cf. Endress 1967) which are characterized by greatly condensed inflorescences and reduced, typically unisexual, mono- chlamydeous, wind-pollinated flowers. The authors mentioned above who view the ‘“‘Amentiferae” as primitive, however, consider the Hamamelid- aceae and its allies transitional to the more advanced Polycarpicae, Ro- sales, or Cornales, with dichlamydeous, bisexual flowers, but this view receives little support from current workers. MATERIALS AND METHODS All of the specimens collected in the field in Southeast Asia and Mexico for anatomical study were preserved in fluid. Fresh specimens were pre- served in formalin-acetic acid-alcohol (F.A.A.) for the most part, or in 70 per cent ethyl alcohol. With collections in Hong Kong, however, a reasonable preservation was achieved with a makeshift solution consisting of equal parts of Rectified Spirits of Wine B.P., and distilled white vingar, giving concentrations of about 50 per cent ethyl alcohol and 2.5 per cent acetic acid. These specimens were later transferred to F.A.A. Flowers from herbarium specimens have been used for a few genera, but were often found to be unsatisfactory for anatomical purposes due to distortion and rupturing of the tissues caused by pressing, drying, and re-expansion. Flowers at both anthesis and post-anthesis stage were sectioned when available, since the ovary and its vasculature are poorly developed in some genera at flowering time. : : Cleared and uncleared whole flowers, hand sections, and serial microtome 312 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 sections were utilized according to the nature of the material to be ex- amined. -Flowers from herbarium specimens were softened and re-ex- panded, using the modified Juel method described by Tillson & Bamford (1938), and then treated as fresh material. Specimens to be cleared were first soaked in water to remove excess F.A.A., then bleached to near-whiteness at room temperature in a solution consisting of nine parts 2.5 per cent sodium hydroxide to one part 5 per cent sodium hypochlorite or commercial Clorox. The bleaching solution was rinsed out thoroughly with water before gradual dehydration followed by clearing in xylene, chloral hydrate, or paraffin oil. Where necessary and possible, pubescence was removed from floral surfaces after clearing in xylene, when the brittle epidermis lifts off easily. Flowers to be sectioned were of necessity bleached and softened. Un- treated flowers proved generally unsatisfactory for sectioning and staining due to 1), tearing of the sections caused by the hardness of the lignified, often dense epidermal pubescence, or hard internal tissues such as the developing bony endocarp, or 2), the presence of densely staining or- ganelles or intracellular substances of undetermined nature with a particu- lar affinity for safranin, which often obscure the vasculature. A distinct disadvantage of this harsh process is the collapse of cells that it may cause in delicate parenchymatous epidermal surfaces, as in young ovules, or the transmitting tissues of the stylar canal, etc. After bleaching, the flowers were dehydrated in a normal-butyl or tertiary-butyl alcohol series, em- bedded in “Paraplast,” and sectioned at 10, 15, or 20 micra, depending on the size of the flower. Various stain combinations were tried, including erythrosin and crystal violet, safranin and anilin blue, safranin and fast green, and Foster’s tannic acid-iron chloride (Johansen 1940). The last two methods gave the most satisfactory results for tracing vasculature, and were used almost exclusively. Polarized light was frequently used to aid in tracing small vascular bundles. rawings were made with the aid of the drawing attachments designed for the Wild M-—20 and M-—5 microscopes. The floral pubescence so ob- scures the outlines of the floral organs in many genera that it has been omitted in most of the figures for the sake of clarity. ACKNOWLEDGMENTS This work is based in large part on a doctoral dissertation presented to the Graduate Faculty of the University of Minnesota in November, 1967. Additional material has since been studied as it became available from collectors or institutions. I am deeply grateful for the financial support given this study by the Graduate School and the Department of Botany of the University of Minnesota, The National Sicence Foundation, and the Directors of the Gray Herbarium and the Arnold Arboretum of Harvard University. I wish to express my profound appreciation to Dr. Ernst C. Abbe, who suggested the investigation, made possible the collec- 1970] BOGLE, HAMAMELIDACEAE 313 tion and study of critical Old World genera in Southeast Asia, and took an active interest in the progress and results of the work. It is impossible to thank my wife, Margaret, enough for her help during the course of this project. Generous support has been given the work through the donation or loan of specimens from many sources. I would like to thank Dr. E. C. Abbe, Dr. A. O. Dahl, Dr. Delzie Demaree, Dr. Lawrence C. W. Jensen, Dr. Robert B. Kaul, Dr. A. R. Kruckeberg, Dr. M. Mizushima, and Mr. H. C. Tang, for collections made in my behalf. Thanks are also due to the directors of the following arboreta, botanic gardens, and organizations, who have made important contributions of specimens, or have permitted me to make collections from their gardens: the Arnold Arboretum; Long- wood Gardens; the Morris Arboretum; the United States Plant Introduc- tion Station at Glenn Dale, Maryland; the University of Washington Arboretum; the Royal Botanic Gardens, Kew, Great Britain; the Botanic Gardens of Tokyo University, Japan; the East African Herbarium, Kenya; the Botanical Research Institute of the Republic of South Africa; the Botanical Survey of Swaziland; and the Botanic Garden of the Taiwan Forestry Research Institute. The curators of the following herbaria have allowed me to borrow specimens or to examine their collections; to all of them I express my appreciation: the Arnold Arboretum and the Gray Herbarium of Harvard University, the Chicago Natural History Museum; the University of Michigan Herbarium; the University of Minnesota Herbarium; the Herbarium of the Department of Botany of the United States National Museum; the Herbarium of the Urban Services Department, Hong Kong; the Herbario Nacional del Instituto de Biologia, Universidad Nacional Auténoma de México; the Muséum National d’Histoire Naturelle, Paris; the Philippine National Herbarium, Manila; the Herbarium of the Botanic Gardens, Singapore; and the Herbarium of the Taiwan Forestry Research Institute. : 5 Particular efforts were put forth in my behalf by Dr. Wei-Fang Lin, Mr. Tao Kiang, and Dr. T. Liu, of the Forestry Research Institute of Taiwan, who arranged several weeks of collecting in the mountains of Taiwan. Similarly, Dr. Eizi Matuda, of the Universidad Nacional Auton- oma de México, Dr. S. M. Martinez, Director, and Mr. I. L. Ostria, Sub- director, of the Department of Natural Resources, State of Mexico, ar- ranged a collecting trip for me to remote areas of the State of Mexico. Through their efforts I was able to obtain critical specimens for this study, and I am extremely grateful to them. GENERAL CONSIDERATIONS For the purposes of this work the genera of the family will be considered as comprising three fairly natural groups. The apetalous tribes and genera of the Hamamelidoideae (Distylieae: Distylium, Sycopsts, M atudaea, Molinadendron,; Fothergilleae: Parrotia, Parrotiopsis, Fothergilla) will be 314 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 considered here, followed later by the petaliferous Hamamelidoideae (tribes Hamamelideae, Eustigmateae, Corylopsideae), and finally the four small and distinct tribes Liquidambaroideae, Rhodoleioideae, Disanthoideae, and Symingtonioideae. The order of consideration of the genera here differs somewhat from that given in the system of Harms (1930, as revised by Melchior, 1964, and with four tentatively placed genera included), which is the most comprehensive available. The tribe Distylieae Hallier (1903) originally contained only Distylium Sieb. & Zucc. and Sycopsis Oliv. and was characterized by Hallier (op. cit., p. 255) as follows: Leaves leathery, entire or toothed, evergreen, with small lanceolate stipules, spicular cells, two-layered palisade tissues, and small single crystals. Inflorescences pedunculate, solitary, of axillary heads or racemes; bracts small; flowers monoecious or andromonoecious, lacking petals; anthers oblong, narrowed below to a short filament and above into a short point, dehiscing by two lateral, longitudinal, simple slits. The Central American genus Matudaea Lundell (Lundell 1940) was considered by Walker (1944) to conform in most respects with Hallier’s description of the tribe, and has been tentatively included in it. Walker also maintained two Central American species of Distylium which, with a third species since described, were recently and rightfully removed as the segregate genus Molinadendron Endress (Endress 1969). Endress con- siders Molinadendron misplaced in the Distylieae, and closer to Fothergilla and Parrotiopsis (Fothergilleae) or to Fortunearia and Sinowilsonia (Corylopsideae). The genus Sinowilsonia Hemsl. (Hemsley 1907) was added to the Distylieae (Harms 1930) but later removed to the Corylop- sideae (Schulze-Menz 1964). I shall describe it with the petaliferous Ham- amelidoideae. Leaf venation in the Distylieae is pinnate. In Molinadendron the veins terminate in the distal, marginal teeth, but in the other three genera the veins are, for the most part, curved-anastomosing (camptodromous type) well within the margins, or in a few species basally triplinerved (Matu- daea, Sycopsis laurifolia) and approaching a camptodromous condition. The tribe Fothergilleae DC. (1830) contained only Fothergilla L. Niedenzu (1891), however, recognized a tribe Parrotieae which combined the apetalous genera Distylium, Parrotia C. A. Mey., and Fothergilla (including Parrotiopsis) with the petaliferous Corylopsis (Sycopsis was placed in the tribe Hamamelideae between Eustigma and Hamamelis). The tribe Fothergilleae in Harms’s system (1930) contains Fothergilla, Parrotia, and Parrotiopsis (Niedenzu) Schneid., and is characterized in his key to the family (op. cit., p. 315) as having flowers perfect, in con- gested spikes or small heads. Petals lacking. Stigma punctiform or capi- tate. Number of floral parts irregular. Stamens often more than five. Anthers dehiscing by longitudinal slits or valves. The leaves of the three genera of Fothergilleae are membranous, stipu- late and deciduous, with ascending, often more or less parallel pinnate veins which terminate in the distal lobes of the sinuate-dentate to coarsely toothed margins (craspedodromous type). 1970] BOGLE, HAMAMELIDACEAE 315 The structure of the ovary is rather uniform within the Hamamelidoi- deae. Therefore, a general description is provided at this point in order to avoid repetition in the generic descriptions below. The trends of vas- cular specialization will be discussed later for the entire subfamily. € gynoecium consists of two, or not infrequently three, involute carpels. The carpel margins are fused below, and the carpels united adaxially through the length of the ovary, forming an incompletely septate, essentially bilocular capsule. The two locules become confluent above, just below the level of ovule insertion near the apex of the locules, as the opposing fused adjacent carpel margins incorporated in the septum separ- ate to form the parietal placentae. Above the ovules the ovarian cavity continues upward as a narrow, elongated canal which opens to the outside through the style bases. Above the ovary the carpels are free, in the Distylieae and Fothergilleae terminating in exserted, linear-subulate styles which are often recurved, and sometimes slightly expanded, bearing ex- tensive exposed or included papillate stigmatic surfaces along their ven- tral faces. One or occasionally two, pendent, anatropous ovules may occur in each carpel, inserted on both margins, but usually only one of these (occasionally two) is functional. Each carpel is supplied with five bundles: the dorsal midvein, two submarginal lateral bundles, and two marginal ventral bundles. In the syncarpous ovary the lateral bundles or adjacent fused carpel walls are incorporated in discrete or diffuse compound lateral bundles (the synlater- als of Endress, 1967; lateral bundles of Horne, 1914) which ascend at each edge of the septum. The compound laterals may dichotomize to form individual lateral bundles well below the level of ovule insertion, or re- main united almost to the top of the ovary, where they fuse with the ventral bundles. With few exceptions the ventral bundles arise in the septum from a narrow, largely procambial remnant of the receptacular cylinder (the inner series of Horne, 1914), which enters the base of the septum before consolidating as a single axial bundle (the synventralmedian of Endress, 1967) and dividing to form two compound ventral bundles (the synventrals of Endress, the septal or placental bundles of Horne). The ascending compound ventrals may also divide to form distinct ventral bundles well below the level of ovule insertion, or remain united to the level of fusion with the lateral bundles near the top of the ovary. After fusing with the lateral bundles the ventrals continue upward into the margins of the style. Each functional ovule receives a single trace from the adjacent ventral or compound ventral bundle. No traces have been observed to supply abortive ovules. Within the ovary walls the dorsal and lateral bundles supply lateral branches to the vascular network of the carpels. At anthesis this network is poorly developed and mainly pro- cambial, but in the fruit it forms a coarse reticulum of thickened bundles over the surface of the bony endocarp. Above the locules the dorsal bundles pass upward as the major bundle to the style bases, where they may remain distinct, or unite with the ventrals as a single broad vascular arc, 316 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 In the following descriptions the general morphology of the inflores- cence and flowers of each genus is discussed, followed by a description of the vascular anatomy. Abbreviations for various organs and anatomical features illustrated in the figures are introduced as necessary in the text. In the absence of petals, and due to the uncertain morphological origin of the perianth in some genera, the terms perigon and tepal are used, rather than calyx and sepal. An appendicular hypanthium consisting of fused sepal, stamen, and petal bases is present in the petaliferous genera of Hamamelidoideae, but is absent in certain apetalous genera (Distylium, Molinadendron, Matudaea), or is replaced by an analogous structure, possibly consisting of foliar, axial, and floral tissues united in a cup around an ovary. This structure, which has not previously been investigated, is here referred to as a kypanthium for the sake of simplicity. But, as seen in Sycopsis dunnii, staminate flowers may occur at various levels on its outer wall or on its rim, in which case the term “hypanthodium” * in the sense of Emerson & Emerson (1960) might be more appropriate. The shallow floral cups in the flowers of Parrotia, Fothergilla, and Parrotiopsis may be of similar origin, although this is not known for certain, and they are also termed hypanthia. The term stele is used here in a general sense with reference to the vascular system of the peduncle, pedicel, and re- ceptacle, and may include both primary and secondary tissues. The terms bundle, trace, and strand are used interchangeably. The term trunk bundle is applied to those vascular bundles which traverse the hypanthium longi- tudinally, and branch distally to supply traces to the perianth and androe- cium. OBSERVATIONS AND INTERPRETATIONS 1. Distylium Siebold & Zuccarini, Fl. Jap. 1: 178. ¢. 94. 1841 FIGURES 1 —3, 4A—D. The widespread but poorly understood Asian genus Distylium contains about 11 species of evergreen shrubs and trees distributed in temperate, subtropical, and tropical regions from Japan and southern Korea south- ward to Taiwan, eastern and southeastern China, westward in the Hima- laya to Assam, and southward through Burma, Indo-China, and Malaya to Java and Sumatra (Walker 1944; Airy-Shaw 1963; distribution map in Balgooy, 1966). Three Central American species originally ascribed Fs this genus were recently segregated as the genus Molinadendron Endress 1969 The complexities associated with the variation and reduction of flower and inflorescence in Distylium are such that it is difficult to consider one without the other, for it is often difficult to distinguish between flower (euanthium) and inflorescence (pseudanthium), or between vegetative *Eames (1961, p. 249) notes that the term hypanthium has been loosely used to n as “an inflorescence with concave capitulum on whose walls the flowers are arranged.” 1970] BOGLE, HAMAMELIDACEAE 317 — @ € ee? feed as oN Seas (ean A GS ee on B= B28 O% G1 St OS) OS FS 2 e Ni eg es 4 23, ea =$3° geo ey c \S ‘@ ® @ (93) ‘ eo D kK 7 “~~ Stipulate leaf ~ Stamen ie Reduced stipulate leaf ° Vestigial ovary on —. Bract (7 2°or 3°). re) Lateral male flower i | male flower with ee hiae tower ® Lgestigial terminal ovary FicuRE 1. Distylium racemosum: A-C, structure of a complex inflorescence, Each unit is secieneey by the same number in a ree A, inflores escence at anthesis, some bracts = procidh removed — note ain ig oe n of func- tional ovary in each unit; B, matic representation nee A; C, inflorescence diagram of 4; p, anther — solid pile eis path of dehiscenc 318 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 axis and inflorescence axis. The inflorescence of Distylium racemosum is, therefore, discussed in some detail. The following descriptions and inter- pretations of the infloresence and floral morphology are based primarily on material of Distylium racemosum Sieb. & Zucc., D. gracile Nakai, and D. stellare O. Ktze. in my collections, but I have also examined available herbarium specimens of other species of the genus, and find no morpholog- ical evidence to contradict the interpretation. Morpuotrocy: The inflorescences of Distylium are small, bracteate, axil- lary or terminal spikes, racemes, or panicles, produced from a bud of the previous or current growth season (Ficures 1, 4). The flowers are naked (Vink 1957) and unisexual, but through aggregation of staminate and pistillate flowers often appear bisexual, with a functional or abortive ovary. The stamens at maturity are strongly exserted, with oblong to ovate, dark- red anthers in which the connective is produced as a short apiculus, and dehiscence is slightly dorsi-lateral by means of longitudinal slits which are simple in some species and basally subvalvate in others (F1icuRE ID). The functional ovaries are terminal on the inflorescence axes and are ver small at anthesis, with exserted, straight to recurved styles. A hypanthium such as that found in most genera of the family is never present A compound inflorescence of Distylium racemosum (n = 12, Sugiura 1936), containing several axillary inflorescence units arranged along a short lateral axis, is illustrated in Ficure 1a, and diagrammatically in Ficure 1B and c. Examination of each of the inflorescence units reveals a variety of vegetative and floral modifications and reductions such as those noted by Hemsley (1907) and by Vink (1957), as follows: inflores- cence unit number | is axillary to a large bract (br, excised in FicuRE 1A, but scar area hatched); units 2, 3, and 4 are axillary to well developed stipulate leaves, such as that figured with unit 4 (excised at 2 and 3, scar areas hatched); units 5, 6, 7, and 8 are axillary to trimerous bract groups which continue the two-ranked phyllotactic pattern of the lower nodes and represent reduced stipulate leaves; unit 9 is represented only by the terminal ovary. Within unit 1, in ascending order: the proximal node bears a primary bract which encloses two secondary bracts inserted oppositely in the trans- verse plane, flanking a male flower containing six stamens (one of them sterile) and a tiny abortive ovary; the second node bears a trimerous bract group to which is axillary a small, transversely inserted secondary bract, a flower consisting of five stamens, and an abortive ovary; the third node bears a trimerous bract group with three stamens in the axil, but no secondary bracts and no ovary; the inflorescence appears to terminate with a set of trimerous bracts unilaterally inserted at the base of a well developed, functional ovary which is surrounded basally by a cycle of six irregularly inserted stamens, two of which are more or less axillary to the bracts, while the other four are inserted on the receptacle around the naked side of the ovary. In the systematic literature this terminal group of bracts, stamens, and ovary would be interpreted as a bisexual flower with 1970] BOGLE, HAMAMELIDACEAE 319 three sepals, but morphological and anatomical evidence indicate that the bract group and its associated stamens represent a fourth node, that the remainder of the stamens represent a fifth node at which the bracts have failed to develop, and that the ovary represents a terminal, naked pistillate flower. Bracts such as those which I interpret as abortive at node five (indicated by dashed symbols in Ficure 18-1, c-1) are ac tually present at the tips of inflorescence units 2, 3, and 4. In the material available, all stages of reduction of such bracts can be found, down to three abortive primordial protrusions on the receptacle on the faked side of the ovary. The interpretation of the ovary as a terminal, naked, pistillate flower is based on vestigial vasculature in its base (desdtied below), which sug- gests the former presence of a hypanthium or floral envelope of some sort in the recent phylogenetic history of the genus Each higher inflorescence unit is progressively more reduced than unit 1, with fewer bracts and flowers. Unit 5, for example, is axillary to a trimerous bract group and consists of a viceectia ovary surrounded by five hypogynous stamens and four bracts of varying size. But one of the bracts is obviously larger, is inserted opposite and morphologically lower than the other three in the transverse plane, and bears two fertile stamens in its axil, while three stamens are associated with the opposing bract group. This inflorescence would also be described as a bisexual flower, but actually consists of two lateral, naked staminate flowers and a terminal, naked pistillate flower. Unit 6 consists of single primary bract and three fertile stamens, representing a naked staminate flower associated with an abortive terminal ovary. Units 7, 8, and 9 are aggregated as a terminal group of six bracts, six “hypogynous” stamens, and a terminal functional ovary. But three of the bracts, with four associated stamens, morpholog- ically lower and opposite the others on the axis, represent an inflorescence (unit 7) reduced to a naked staminate flower axillary to a bracteate stip- ulate leaf. Unit 8 is similarly reduced to two stamens axillary to the three higher bracts, and the ovary, designated as unit 9, represents a terminal, naked pistillate flower. Units 7, 8, and 9 thus duplicate the condition apparent at the tips of units 2, 3, and 4, with six bracts, five or six stamens, and a functional ovary forming what has generally been considered an incomplete, bisexual flower, a condition brought about by the reduction of leaves and distal internodes of the inflorescence axes, resulting in the aggregation of naked unisexual flowers and their associated subtending bracts to form a bisexual pseudanthium. ; More complicated flowering branch systems have been seen in which the lateral axis illustrated in FicurE 1A would represent only one of numerous progressively reduced units arranged along an axis which terminates in a similar pseudanthium. In such a system branches to the fourth order are involved, each terminating in a Spices hanes each pseudanthium in- corporating elements of two orders of branching The terminal position of the functional ovaries and the reduction of subterminal units are more clearly demonstrated in FIGURE RES 4A-D, of Distylium racemosum. Ficure 4A illustrates an axillary, paniculate in- 320 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 florescence bearing seven maturing terminal ovaries subtended by trimer- ous bract groups. The stamens have fallen away from all the lateral sta- minate flowers, in some cases, revealing abortive ovaries. A subterminal ovary reduced to one carpel and an abortive bicarpellary ovary (ab ov) are shown in FicurE 48 and c, while a tricarpellary ovary with an abortive organ protruding from its side is illustrated in FicurE 4p. The protruding organ is probably a stamen which has become adnate to the ovary and developed a small stigma-like structure terminally. I have found, in my material, several isolated examples of such monstrous stamens consisting of part anther and part stigma, associated with terminal ovaries. In inflorescences from flowering specimens of Distylium buxifolium (Hance) Merr. (FE. H. Wilson 115, 3537, |A|) the inflorescence units were found to be spirally arranged rather than two-ranked as in D. racemosum, and much reduced from the condition seen in the latter, with shorter, more congested primary axes, fewer nodes and lateral flowers, considerable modification and fusion of the primary and secondary bracts, and pseudanthial complexes containing from 4 to 10 bracts (for the most part in obvious groups of three, and representing the reduced stipulate leaves of possibly 2 to 4 subterminal nodes) surrounding the subterminal staminate and terminal pistillate flowers. An abortive subterminal ovary may be present among the stamens axillary to the morphologically lowest group of bracts in the terminal complex toMy: The vascular supply to the petiole of the foliage leaf and to the iiticreseeate bracts of Distylium racemosum and D. gracile consists of three traces derived from three gaps in the stele, with a single trace provided to each stipule from the adjacent lateral leaf trace (FIGURE 24). } OF y--O A Ficure 2. Steps in the aa of the — — to the inflorescence bracts of Distylium racemosum and Sycopsis dunn 1 race, trilacunar node of the stipulate leaf; B, lateral traces = pate vestigial; c, latera i traces absent, only the stipules supplied from the lateral gaps; D, stipulate leaf reduced to a single bract receiving one trace from one gap Within the flowering branch system reduction of the leaf blade is accom- panied by a corresponding reduction in the vascular supply to the pet- iole. Reduction of the lamina results first in the failure of the lateral traces to the petiole to develop fully, as indicated in Figure 2B, where the lateral traces supply traces to the stipules, then end blindly as vestigial stubs in the cortex near the petiole base. The petiole then receives only a single median trace. Further reduction of the lamina results in the loss of the vestigial stubs, so that only the stipules receive traces from the 1970] BOGLE, HAMAMELIDACEAE 321 lateral gaps in the stele (FicuRE 2c). The bracteate blade may fuse with the stipules to form a single broad, trifid bract, or the leaf blade may be lost and the persistent stipules may fuse into a single bifid bract. The penultimate stage of reduction is a single bract supplied by one trace from one gap. The latter condition is encountered most frequently at the base of an axis (FIGURE 2p). In view of the great variation in the flowers of Distylium I have based the following anatomical description on a selected specimen which I think exhibits the general morphological and anatomical features of the species which were available for anatomical study. Ficure 3a illustrates the tip of an inflorescence of D. racemosum which is similar to those described above except that here the leaf blades at the subterminal nodes are fully developed rather than bracteate. Group 1 consists of a naked staminate flower of four stamens axillary to a leaf. Group 2 looks like a terminal bi- sexual flower, but consists of two or three stamens axillary to the distal leaf, while the remaining stamens inserted on the naked side of the re- ceptacle (one with a partially laminar anther) are interpreted as re- ferable to a third node (not numbered), the bracts of which have failed to develop. The axis terminates in the terminal ovary. At the base of group 1, in the wake of the departing leaf traces (If tr), a small axillary stele (ax 2) is pinched off from the main axis to supply the base of the sessile staminate flower. At anthesis this stele is largely procambial, with scattered protoxylem elements apparent only in po- larized light. As the stele ascends in the receptacle four large collateral bundles (st) organize (FIGURE 3c, D, E). Scattered protoxylem strands lying between these bundles in the stele gradually die out, leaving only procambial arcs (stippled) which die out as the four stamen traces depart. At the base of the distal complex, after the departure of the leaf traces, the stele consists of a ring of numerous collateral bundles. Five more or less equally spaced stamen traces (st) differentiate, then arch outward to enter the bases of the filaments (FicurE 3r-H). The gaps left by the traces are shortly repaired as the stele consolidates in the receptacle. Although the stamens are inserted at the base of the ovary morphologically, their traces originate well down in the axis anatomically, as much as 500 micra below the ovary base. In longitudinal sections and cleared specimens the ovary appears to be subtended by a relatively long vascular cylinder or stipe (stp in FicurE 30, stamen traces and scars not shown in the receptacle). : ; In the base of the ovary, at a level slightly above the insertion of the filaments, the tips of numerous small, recurved, procambial strands (peri) are encountered in the cortex. When followed upward these can be seen to branch or anastomose to a slight extent before connecting with con- spicuous large reflexed bundles which originate from the receptacular stele at the base of the gynoecial supply (F1curE 3c-1). Although most of this peripheral system appears in the base of the ovary below the lo- cules, some branches of it continue upward and end blindly in the ovary walls above the level of the locule bases (FIcURE 3J-t, 0). et) 322 JOURNAL OF THE ARNOLD ARBORETUM [voL, 51 Ly. o 2 w aor @ oa 7 ®O Ov G ase lium ra a subterminal s minate flowe o o ip anthium — no atic re — ue ne omg ea subte ane C-E, transverse gia atc ‘through group i (slightly oblique) c, depa leat and eHpale traces — secondary axis developing; p, stamen traces aioe in and departing from he ae Aes ‘axis; E, stamen filaments in axil of leaf; 1970] BOGLE, HAMAMELIDACEAE 323 In the base of the ovary the receptacular stele is slightly elliptic in transverse sections, with its long axis in the plane of the dorsal bundles. From the median sides of the ellipse broad dorsal carpel bundles (d) depart, along with adjacent bundles of the peripheral system (FIGURE 3H, 1). At a slightly higher level two compound lateral bundles (11) organize on opposite sides of the stele in the transverse plane and di- verge slightly toward the periphery (Ficurr 31, J). The stele then con- stricts to form a narrow procambial cylinder at the base of the locules, which divides in the median plane to form two procambial compound ventral bundles (vv) in the base of the septum. The dorsal bundles ascend in the ovary as broad vascular arcs. The compound laterals dichotomize transversely at about the level of ovule insertion near the top of the locules. The compound ventral bundles are not entirely developed at anthesis, and those portions which are present are entirely procambial. They appear to die out as they ascend in the base of the septum, but reappear at a slightly higher level as four indi- vidual ventral bundles (v, Ficure 3j-L). This discontinuity in their development suggests that their differentiation proceeds both acropetally and basipetally in the developing septum. Near the top of the locules the ventrals approach and fuse with their adjacent lateral bundles (1), but the ventral bundle of each fertile margin branches just prior to fusion to provide a trace to the solitary ovule (ov). The most striking features in the floral apparatus of Distylium are the extreme variation in the external morphology of the inflorescences and flowers, the foliar nature of the inflorescence bracts, the development of bi- sexual pseudanthia, and the curious system of vestigial vascular bundles, here referred to as the peripheral system, in the base of the ovary. The flowers of the genus have not previously been studied anatomically, but have been variously interpreted by systematic workers as polygamous, or as unisexual and monoecious, or as andromonoecious, with a variable number (0-6) of free and unequal sepals (or bracts, or bracteoles) in- serted at different levels, an irregular number (2-8) of hypogynous stamens, and a superior ovary (Bentham & Hooker 1865; Baillon 1874; Niedenzu 1891; Harms 1930; Walker 1944; Vink 1957). The gynoe- cium of Distylium has long been considered a prime example of the su- perior ovary in a family generally characterized by half-inferior ovaries. Hemsley (1907) described each of several flowers from two inflores- cences of Distylium chinense Hemsl. in detail, and could find no uni- formity in their organization, nor any way to differentiate between the various “bracteoles” and “sepals.” He concluded that the flowers of F-N, transverse sections (slightly oblique) through group 2; F, stamen traces departin le; G, stamens separating — a rag foe “the pe ies H, ca aa reac bundle and adjacent peripheral bundle le; 1, several peripheral bundles and second dorsal bundle departing from the stele; J, base of locules — some peripheral bundles still evident; x, ovary — individual ventral bundles appearing; L-M, insertion of ovules; N, style bases; 0, gynoecial vasculature as seen In a cleared ovary (semi- atic). diagrammatic 324 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 he genus are functionally either male or female, but never hermaphro- dite. Walker (1944) states that ‘the variations in the floral parts between one species and another are so little, or so inconsistently variable that they are of scant use in differentiating species .. .” “It is often difficult to determine whether flowers are unisexual or bisexual, because there may be found stamens and pistils of all degrees of development even in the same inflorescence.’ Vink (1957) noted that the floral bracts of Distylium stellare most often occurred in threes and sixes, and that the middle bract of a group of three sometimes was leaf-like in appearance, while the two lateral bracts were always more or less lanceolate in shape. Since the leaves of the genus are stipulate, he interpreted the bracts as reduced stipulate leaves, and concluded that the flowers of Distylium are asepalous, as well as apetalous. His is the only interpretation of the inflorescence bracts which is supported by the anatomical evidence. He also observed that distal flowers are often merged in twos, forming ‘‘twin flowers” at the tips of the inflorescences. Croizat (1960, 1964) rejects any conventional explanation for the in- florescence bracts and floral organization of Distylium, considering the flowers of the genus exemplary of his proposed mieetaiae stage of “flower making” between a unisexual amentiferous inflorescence and a bisexual angiospermous flower. He considers the unisexual flower and inflorescence primitive, and derives the bisexual flower through the ‘“sexualization into maleness”’ of pistillate flowers by the de novo origin of stamens on scales (the “sepals,” “bracts,” “bracteoles” of Distylium) subtending the pis- tils. These stamens supposedly assume the functions of the male inflo- rescences which are then suppressed. However, I find no evidence in my material of a unisexual staminate inflorescence of the type figured by Croizat (1960, p. 296, fig. 36A). The “twin flowers” mentioned by Vink are interpreted by Croizat (1960) as “Scales (‘‘carpels”) originally ar- ranged to establish two separate floral structures . . . here rearranged to establish one only, thus factually marking the final transformation of an inflorescence into a single flower.” Morphologically and anatomically, “twin flowers” are nothing more than two bisexual pseudanthia, one ter- minal the other subterminal, closely associated at the tip of an inflores- cence. Each is subtended by a trimerous bract group, stamens of both are inserted between the two pistils, and vestigial peripheral bundles are present in both ovaries. Although Croizat was correct in his interpreta- tion of the pseudanthial nature of some “flowers” in Distylium, the evi- dence now available contradicts his views concerning the method of their derivation. 2. Sycopsis D. Oliver.? Trans. Linn. Soc. London 23: 83. 1860. FicureEs 4&-F, 5, 6. An Asian genus of evergreen shrubs or trees containing about nine species ranging in distribution from Assam through Southeast Asia and *The generic name is derived from the Greek sykos, for fig, and opsis, meaning 1970] BOGLE, HAMAMELIDACEAE 325 the Malay Archipelago to New Guinea. Sycopsis sinensis Oliv. has the most northerly distribution, occupying a broad east-west band across seven provinces of central China, while Sycopsis dunnii Hemsl. has a very broad southern distribution from southern China southward to Su- matra and eastward to the Philippines and New Guinea. The other seven species have relatively restricted distributions. Prominent among them are S, griffithiana Oliv. in Assam and possibly Yunnan, S. laurifolia Hemsl. in Yunnan, S. tutcheri Hemsl. in Kwangtung (Hong ‘Kong) and Hainan, and S. salicifolia H.-L. Li in Walker, in Hainan Only Sycopsis sinensis is well bow in cultivation, and it has recently been shown to be interfertile with Parrotia persica, although the hybrid, x Sycoparrotia semidecidua, is apparently sterile (Endress & Anliker 1968) The following descriptions are based primarily on flowers and inflo- rescences from cultivated plants of Sycopsis sinensis, and from herbarium specimens of S. dunnii and S. laurifolia, but available specimens of the several other species were also examined morphologically. Morpuotocy: The inflorescences of Sycopsis are short, axillary or terminal panicles, spikes or heads. The congested spikes or heads of Sycopsis sinensis (FicuRE 6N) and S. griffithiana, with each flower en- closed by a broad, densely dark-brown-hirsute primary bract, differ sig- nificantly from the short open spikes or racemes of S. dunnii and related species, with flowers subtended by unilaterally inserted trimerous groups of small bracts (see FiGURE 4e—-H). Secondary bracts are generally ab- sent in the inflorescences, but one or two may occur flanking a basal flower, either free or partially adnate to the hypanthium The axillary or terminal flowers are sessile to subsessile, asepalous (Vink 1957) and apetalous, and appear to be andromonoecious (I have found no evidence of strictly pistillate flowers that would substantiate the interpretation of polygamous inflorescences encountered in the sys- tematic literature). Staminate flowers contain 3 to 7 stamens in Sycopsis dunnii, and (5—)7 to 10 in S. sinensis, inserted perigynously in a single cycle on the rim of a rudimentary, hirsute, naked (S. dunnii) or irreg- ularly toothed and lobed (S. sinensis, Ficure 7A) hypanthium, around a minute abortive ovary. At anthesis the stamens become strongly eXx- serted, with elliptic to ovate (S. dunnii, S. laurifolia) to distinctly oblong (S. sinensis), 2-celled anthers containing four pollen-sacs, and dehiscing by means of lateral longitudinal slits which are simple above, but ven- trally subvalvate below. The anther connective is produced as a short, blunt point. Bisexual flowers contain a 2(-—3)-carpellary, superior to slightly semi-inferior ovary, surrounded by an urceolate subglobose hypanthium which, in Sycopsis dunnii and related species, bears irregular bracts or bract groups (the bracteoles and sepals of most authors) in- likeness (Rehder nears Oliver (1860, p. 83), in his ce description, tage that “the young subobov fruits, narrowed and (from drying?) more or sulcate below, together with ei re ‘SESS of the plant, fiat some of hea narrower- leaved species of East Indian F 326 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ae 4, Distylium racemosum and Sycopsis dunnii: a-c, Distylium racemos- ; A, maturing inflorescence, stamens absent — note te rminal functional ovaries of a few axillary abortive ovaries; B, pistil of subterminal pseudanthium re- 1970] BOGLE, HAMAMELIDACEAE 327 serted unilaterally at its base, at various levels on its outer surface, and on its often oblique rim. Bu t i in S. sinensis and S. griffithiana there are no unilaterally inserted bract groups, and the level to oblique hypanthium rim bears a single cycle of (4—)5 to 7 irregularly lobed to incised tepals which are basally connate into a tube above the rim (cf. FiGuREs 5A and 6A). Three to seven stamens are present in the bisexual flowers of S. dunnii, while 8 to 9 stamen scars could be counted on the hypanthium in the post-anthesis flowers of S. sinensis which I examined. The long styles are spreading (S. dunnii) to spirally curved (S. sinensis), A single ovule or seed occupies each locule. In inflorescences of Sycopsis dunnii and S. laurifolia it appears to me that only the terminal ovary (i.e., in the terminal bisexual flower) on each inflorescence axis is func- tional. The lateral and subterminal flowers on each axis are staminate through reduction of the ovary. A staminate flower, or an abortive sterile flower, is usually axillary to the bract group at the base of the terminal flower, and stamens may also be axillary to the bract ~ in- serted at mid-height on the hypanthium (Ficures 4£-H, 5a, C). sinensis the ovary of each bisexual flower in the spike may be Salen The morphological differences between Sycopsis dunnii and S. sinensis are sufficient to place their congeneric relationship in question. Conse- quently, the vascular anatomy of both species is considered here. ANATOMY OF SycoPsIS DUNNII: Terminal flowers with functional ovaries, associated bracts, and in some cases abortive subterminal flowers, taken from herbarium specimens (Steward & Cheo, 165, 170, 199|a]; also S. laurifolia, A. Henry, 11365[A]), were sectioned or cleared. The flowers are variable in their organization with regard to the level of in- sertion of the bracts, and in the number of bracts (0-2) and stamens on the rim. Relatively few flowers of either species were available for sec- tioning, and the vascular organization of individual flowers differs with their morphological organization. The gk of S. laurifolia appear to agree in general with those of S. dunn The vascular supply to the aneeae bract groups subtending the la- teral staminate flowers and the terminal bisexual flowers is similar to the nodal anatomy typical of the foliage leaf, i.e., three traces derived from three gaps, supplying the central reduced leaf blade, with a trace sup- plied to each stipule by the adjacent lateral leaf trace. The vascular supply to the three bracts mounted on the side of the hypanthium may be similar to the typical pattern or represent a reduced modification of it, ultimately consisting of three separate traces, each of which supplies only one bract. The modifications of the vascular supply to the bracts are similar to those described above (FicurE 2) for Distylium. The duced to a single carpel; c, subterminal pistil abortive; sghooed -carpelled ay a istil — note soar: adnate organ. E-H, p. unnit; E, ma i ‘ cihactaue note trimerous ‘bract groups on fla inks of hypanthia ; “ subter doin ower almost fully developed, aio functionally ee subte wer abortive and sterile; H, subterminal flower reduced t bristle-like pees fi (ab fl) adnate to the hypanthium 328 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 st FicurE 5. Sycopsis dunnii: a, view of a ape ie bisexual — and subterm- nal abortive sterile flower (post- anthesis); 8B, flor agram; C, view of an in- Resice tip at anthesis, with two subterminal eatate fis beneath the 1970] BOGLE, HAMAMELIDACEAE 329 traces supplying bract groups inserted on the side of the hypanthium originate from the peduncular stele well below the departure of the main vascular supply to the hypanthium, and ascend in the parenchyma- tous tissue near the outer surface of the hypanthium, entirely apart from the remaining vasculature (FIGURE 51-0). In one instance the trace to a solitary distal bract was discontinuous for a considerable distance in the upper portion of the hypanthium, possibly having been disrupted by elongation of the latter (a phenomenon noted also in Fortunearia). The vascular supply of a single trace to each of the bracts inserted on the rim of the hypanthium is derived within the hypanthium from narrow secondary vascular cylinders or trunk bundles which also supply the stamens and constitute the main vascular supply to the hypanthium. The major supply to the hypanthium develops from the peduncular stele well below the base of the ovary (about 600 to 700 micra below the base of the locules in the sectioned flower of Ficurr 5), where as many as seven large collateral bundles diverge on as many radii into the cortex (FIGURE 5J-L). Immediately behind each of these, and from a slightly higher level as seen in transverse sections, one to three small branches from ad- jacent bundles bend outward (FicuRE 5J, arrows) and take up positions on the adaxial side of the larger bundle, where they remain distinct through almost the entire length of the hypanthium, uniting with the larger abaxial portion only as the rim is approached and the strands con- the phloem accompanying the small adaxial xylem strands is oriented laterally to adaxially, so that the aggregate of vascular strands in each group then varies in appearance from a narrow, more or less continuous to discontinuous ring of xylem strands with associated external phloem, as in a narrow stele, to two opposing tangential narrow bands of xylem with phloem on the abaxial face of the outer band, and on the adaxial face of the inner band. In some flowers a relatively substantial vascular cylinder occurs in the hypanthium, originating from the peduncular stele approximately in the transverse plane of the ovary (and in the plane of the two-ranked bracts), and ascending to supply traces to bracts and stamens on the rim (FIGURE 5M-R). From the bases of these trunk bundles and secondary steles lateral terminal bisexual flower; pD, longitudinal median section (semidiagrammatic) of a maturing terminal flower (bracts are out of the plane of section) ; E; ene F-H, sections through the abortive subterminal flower; F, leaf rhe - iv ing secondary stele; G, bundles of abortive hypanthium and carpel se “462 vind H, fragment of abortive ovary apparent; I-R, transverse sections t ng a ep tacle and terminal flower; 1, three leaf traces departing the peduncular stele; J, hag! secondary nee q undles forming on right side; M, ' sgn and peripheral "bundles on left side; N, about middle of ovary —note s 3 trace, distinct ventral bundles on left; 0, ovule insertion — note sees hig ing from the hypanthium; p, larger secondary axis in the hypanthium a ivide into three smaller units; Q, R, upper levels of hypanthium and sty a S, Tr matic representation of a terminal bisexual “flower” interpreted as a pseud- anthium. 330 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 branches are given off to either side near the base of the hypanthium. The branches move laterally and upward in the hypanthium, often branch- ing and anastomosing with strands from adjacent bundles. In transverse sections these branches are situated randomly between the major bundles (FicurE 5M—0). Much of this system dies away as it ascends (at least in the post-anthesis flowers examined), but in some cases a single bundle may persist to the rim of the hypanthium between and slightly adaxial to each pair of major bundles (arrow in FicuRE 5R), suggesting a vestigial supply to abortive stamens. I could find no anatomical evidence in the hypanthium suggesting the former presence of sepals abaxial to the sta- mens on the rim. Above the departure of the major bundles to the hypanthium the re- maining bundles of the peduncular stele shortly consolidate to form two dorsal carpellary bundles (d), a narrow receptacular stele which enters the base of the septum to form the compound ventral bundles (vv), and two diffuse groups of traces on either side of the receptacle in the trans- verse plane from which are derived the compound lateral carpellary bundles (ll) centripetally, and centrifugally a series of weak traces (peri) which diverge almost horizontally into the base of the hypanthium just beneath its separation from the ovary, and about 100 to 200 micra high- er than the divergence of the major hypanthial bundles (Ficure 51, M, Ss). Within the base of the hypanthium these traces, in association with some minor strands derived from branches of the main hypanthial system, form an irregular ascending, branching, and anastomosing system of bundles in the parenchyma just within the adaxial face of the hypanthium, to either side of the ovary in the transverse plane (FIGURE 5M-P, S). This system also gradually dies away as it ascends. The vascular supply to the hypanthium, therefore, consists of two independent sets of traces derived from different levels in the receptacular stele. The ovary often appears to be fused with the hypanthium to a slightly higher level on the transverse side which bears the distal bracts (cf. F1c- URE 5M). Within the pistil the compound lateral and compound ventral bundles divide in the upper half of the ovary to form distinct laterals and ventrals which fuse at the level of ovule insertion. A single ovular trace departs from the ventral of each fertile margin just prior to its fusion with the lateral (Ficure 50). ANATOMY OF SYCOPSIS SINENSIS: Bisexual flowers. Post-anthesis flowers from which the stamens had fallen were taken for sectioning from the lower half of inflorescences where there is less crowding of the flowers (cf. FicurE 6N) and less likelihood of deformation of the receptacular vasculature. A cylindrical pedicellar stele subtends the flower. In the receptacle 8 to 13 large bundles diverge sharply from the stele to enter the base of the hypanthium, in which they ascend more or less directly to the rim. Some of the bundles function only as traces to the shallow perigon tube and its irregular lobes, while others are trunk bundles supplying traces BOGLE, HAMAMELIDACEAE FIGURE 6. Sycopsis sinensis (bisexual flower): A, post anthesis flower — note irene lobing of the isipeer or one green stamen shown; B, floral diagram; c, drawing from cleared porti of the m e hypanthium showing perianth lobes, ta eae scars, trunk rein sacar akan of the lobes; p-m, transverse sections of post-anthesis flower; D, pedicellar stele and base of primary bract; E, receptacle — note departing hypanthial trunk bundles and et ee re- on = pound ventral bundles; 3, mid-height in ovary; K, insertion of | si ovules; tos formation of tepal and stamen traces in hypanthium; M, insertion of stam on the hypanthium rim; n, fruiting inflorescence, only terminal iewse t in fruiting condition — note splitting hypanthium $32 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 to both stamens on the rim and to the tepals. At various levels in the hypanthium the trunk bundles may produce single lateral branches which function as dichotomizing traces to the perigon (cf. FIGURE 6c), but branching is sparse within the hypanthium. I could find no regular patterns in either the lobation of the perigon, or in its vascularization. In transverse sections some of the hypanthial bundles are collateral, while others, particularly the trunk bundles, are somewhat variable, often appearing as narrow vascular cylinders as in the hypanthium of Sycopsis dunnii. There is no evidence, however, of the two systems of numerous minor bundles derived from branches of the trunk bundles and from the receptacular stele in the omen of the developing componnd lateral car- pellary bundles, as in S. dun Occasionally a sdinusens fn bundle, such as those indicated e instance the larger bundle was about 500 micra long, while its smaller companion was only about 80 micra sate Both ended blindly above and below Slightly below the level of stamen insertion many, but not all, of the perigon tube, where it dichotomizes irregularly, forming 2 to 3 smaller bundles which supply the tepal lobes (FicuRE 6c, K—M) Above the departing hypanthial bundles a prominent receptacular stele continues upward for a short distance (about 200 to 350 micra in the post- anthesis material examined), becoming broadly elliptic in the median plane in transverse sections as the dorsal carpellary bundles (d) organize and depart. As each dorsal moves away from the stele one or two median or lateral traces from it bend sharply inward and return almost hori- zontally toward the stele (arrow in FicURE 6F), where they are in- corporated into a smaller vascular cylinder remaining at the center of the receptacle. From this cylinder two to four bundles depart on either side in the transverse plane to take up positions in the bulging walls of the exocarp and on the edges of the septum as lateral bundles (1) or dif- fuse compound lateral bundles (ll, Ficure 6c-1). Their departure leaves two small opposing vascular arcs in the median plane which consolidate to form a narrow, largely procambial receptacular cylinder containing several strands of protoxylem in the base of the septum (FIGURE 6G-H). At the base of the locules the hypanthium is essentially free from the su- perior ovary, and the major bundles of both are apparent in transverse ections Within the ovary the dorsal bundles are relatively massive and occupy prominent dorsal bulges in the carpel walls. In the bundle groups at either edge of the septum the central bundles function as the compound laterals, while the lateral bundles move outward into lateral bulges and even- tually ramify and anastomose into the vascular networks of the carpel walls. The narrow receptacular cylinder in the base of the septum shortly divides to form two compound ventral bundles which ascend unbranched 1970] BOGLE, HAMAMELIDACEAE 333 to the base of ovule insertion where each dichotomizes. The distinct ven- trals thus formed then fuse with adjacent distinct lateral bundles. The ovule traces are supplied from the ventrals just prior to fusion (FicuRE 6J, K). In some ovaries it appears that the ventral bundle of the fertile margin goes entirely to the ovular funiculus, free of any connection to the adjacent lateral bundle, but such connections may form later in the ontogeny of the ovary. The marginal ventral bundles in each style soon become associated with the dorsal bundle to form a vascular arc, rather than three distinct bundles (Ficure 61, m). Staminate flowers. The hypanthium is strongly reduced, and the gyn- oecium rudimentary in the sessile staminate flowers (FIGURE 7A). The shallow, membranaceous perigon tube which projects above the rim of the hypanthium is irregular in its lobing and vascularization. The lobes bear long dark-brown stellate hairs on their distal margins. The tepals in- dicated in the floral diagram (FicureE 7B) correspond to the lobes of the perigon tube as they appeared in the sectioned flower illustrated. A well organized pedicellar stele was present in some, but not all of the specimens examined. A reduced pedicellar stele may be present only as a lobe of the peduncular stele on the lower side of the flower, while on the upper side the bundles to the receptacle originate from the margins of a gap in the peduncular stele. In the receptacle of the flower the entire stele is incorporated into 9 to 12 large bundles, most of which enter the hypanthium (FicurE 7c-k). In the median plane two dorsal carpel bundles (d) are derived either from a single bundle of the stele, or by the fusion of 2 to 3 traces from adjacent hypanthial bundles (FicuRE 7£, F). No compound ventral or lateral car- pel bundles are formed, but there is frequently the appearance of vestigial traces to such bundles in the form of protoxylem strands which bend in- ward from the bundles in the receptacle at the level of dorsal bundle for- mation. : ; The ascending bundles in the shallow hypanthium may remain entire, or give rise to one or two lateral branches which eventually supply the perigon tube (FicurE 7H). The trunk bundles, situated on the radii of the 7 to 10 stamens, branch just below the insertion of the stamens to produce a large adaxial stamen trace (st), and a smaller abaxial trace (te) which enters the perigon tube, where it may divide irregularly and supply branches to a tepal. A few (1 to 3) bundles in the hypanthium divide at about the level of stamen insertion to provide traces to the perigon only. There was no correlation between the number of bundles in the hypanthium and the number of stamens on its rim in the flowers examined. In Sycopsis, as in Distylium, it becomes difficult to distinguish between flower and inflorescence. The flowers, especially, present problems of in- terpretation and nomenclature, particularly the tubular structure surround- ing the ovary and bearing the stamens. This has been termed an urceolate receptacle (Harms 1930; Tong 1930; Walker 1944; Vink 1957), a hy- 334 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 AY ie se A: A: a s : ¥ ib of stamens on hypanthium; j, through the perianth tube, filaments, and styles; K, separation of some tepals and upper portion of the primary bract. panthium bearing calyx lobes (Endress & Anliker 1968, ref. S. sinensis), or a calyx tube with short lobes (Oliver 1860; Niedenzu 1891). The nature of the perianth has also proved problematical. Hemsley (1907), in his original descriptions and illustrations of Sycopsis dunnii, 1970] BOGLE, HAMAMELIDACEAE 335 S. tutcheri, and S. philippinensis (= S. dunnii), clearly indicates the uni- lateral insertion, variability in number, and morphological similarity of the “bracteoles” of the inflorescence and the “sepals,” counting as the latter the 1 to 3 bracts occurring on the hypanthium rim. The variability in the numbers of sepals is also evident in the species descriptions in Walker’s (1944) revision of the genus. Vink (1957), in his comparative notes on the flowers and bracts of Distylium stellare and Sycopsis dunnii, concluded that the flowers of Sycopsis are also asepalous, and that the usually trimerous bract groups at the base, sides, and rim of the urceolate hypanthium are actually reduced stipulate leaves. The vascular supply to the bract groups, described above, clearly indicates that his interpreta- tion is correct, at least for S. dunnii and S. laurifolia. The foliar nature of the tepals of Sycopsis sinensis and S. griffithiana is not so clear cut, however. In these species the 5 to 7 tepals are inserted in a single cycle on the rim of the hypanthium, projecting beyond the rim as an irregularly lobed and fused perigon tube. Oliver (1861), in his original description of the genus, based on S. griffithiana, states of the perianth “it is doubtful how far the lobation and degree of obliquity of the calyx limb is con- stant.” He later (1890) described the perianth of S. sinensis as “calycis tubo irregulariter fisso.”’ Vink’s interpretation of the bracts of S. dunnii does not appear directly applicable to the markedly different perianths of these two species. Interpretations of the sexual condition of the flowers have varied. Oliver described the flowers of S. griffithiana as unisexual through abortion, and questioned whether they were ever hermaphrodite. Walker found the flowers of the genus to be “unisexual and monoecious or andromonoe- cious,” while Vink considered them “polygamous monoecious.” In cul- tivated specimens of Sycopsis sinensis Endress (Endress & Anliker 1968) found only bisexual and staminate flowers (i.e., andromonoecious). He observed that the buds of all flowers are bisexual, and that the stamens develop in all flowers, but that the ovary is abortive in many. The pistil- late flowers reported in the literature actually represent bisexual ones from which the stamens have fallen. He also found that inflorescences terminal on short or long shoots contained mostly bisexual flowers, with only one or two basal staminate flowers, while inflorescences from lateral buds were in many cases entirely staminate and soon deciduous. Stamen development is often irregular in bisexual flowers of Sycopsis sinensis with several stamens abortive. This condition has also been noted in S. griffithiana by Oliver (1860), and in the generic description by Vink (1957, presumably for S. dunnii). Walker noted that the hypanthium does not develop and enlarge with the fruiting ovary, as do the hypanthia of other genera of the family, but splits irregularly almost or quite to the base, forming a shallow 2- to 3-valved cup around the base of the mature fruit (FicuRE 6N, terminal fruiting flower). In S. laurifolia, however, as in Sinowilsonia, the hypanthium does appear to develop with the fruit, forming a thin papery urceolate covering which ruptures only as the woody capsule within springs open. 336 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 The morphology and anatomy of Sycopsis dunnii and S. laurifolia in- dicate that their floral apparatus is rather specialized. Such morphological features as the open spicate to paniculate inflorescences; the terminal posi- tion of the functional ovaries; the reduction of lateral flowers to a stam- inate condition; the form and dehiscence of the anthers; the absence of tepals; the presence of reduced or modified stipulate leaves as the bracts of the inflorescence; and the crowding out of subterminal lateral flowers through the failure of internode elongation, all suggest a relatively close relationship to the genus Distylium. The complex structure of the hypanthium in Sycopsis dunnii and S. laurifolia is indicated by such anatomical features as the presence of leaf traces, independently derived from the peduncular stele, supplying the bracts on the side of the hypanthium; the presence in some flowers of secondary vascular cylinders supplying bracts and stamens on its rim; and the two complex systems of minor peripheral bundles or trunk bundle branches in its base. The accumulated evidence suggests that the hypanthium in S. dunnit and S. /aurifolia may consist of foliar, axial, and floral tissues attributable to bracts and staminate flowers of subterminal nodes, which, through con- densation of the inflorescence axis, have developed upwards and fused into a tube around the naked ovary of a terminal pistillate flower. If this interpretation is correct it would appear that, as in Distylium, only naked unisexual flowers are present in the inflorescence, with the staminate flowers incorporated into the hypanthium around the pistillate flower, forming a bisexual pseudanthium. The hypanthium is thus neither “ap- pendicular” nor “‘receptacular” (cf. discussion of these terms in Eames, 1961, p. 249, 250). It is suggestive of the syconium of Ficus, but differs both morphologically and anatomically (cf. Condit 1932). The term hy- panthodium may be appropriate for it (see footnote 1). The ultimate fate of the hypanthium as an outgrown, ruptured structure at the base of the fruit further suggests that it is not an intrinsic part of a bisexual, perigyn- ous flower. The exceptional behavior of S. /aurifolia in this respect should be investigated. The vestigial peripheral bundles in the receptacles and ovaries of S. dunnii and S. laurifolia, although suggestive of the peripheral bundles in the ovary of Distylium, are less numerous and more localized in their origin from the receptacular stele and do not persist in the base of the ovary. Their direct contribution to the hypanthium suggests that they may represent the vestigial supply to subterminal lateral organs squeezed out in the condensation of the axis and no longer apparent morphologically, but possibly incorporated into the hypanthium. I have attempted to express this interpretation of the pseudanthium of Sycopsis dunnii diagrammatically in Ficure 5s. Sycopsis sinensis and S. griffithiana differ strongly from the other species of the genus in the morphology of their inflorescences and flowers, partic- ularly in the absence of obvious reduced stipulate leaves or bract groups associated with the flowers, and the disposition of the tepals in a single cycle on the hypanthium rim. However, the two groups of species show 1970] BOGLE, HAMAMELIDACEAE 337 some similarity in their leaf form, and in the possession of urceolate hypanthia which are ruptured by the developing fruit. Anatomically, S. sinensis differs from S. dunnii in the absence of prominent secondary vas- cular cylinders and of the two minor bundle systems in the hypanthium. Vink’s interpretation of the inflorescence bracts of S. dunnii would apply to S. sinensis only if one hypothesizes that, through a process of con- tinued inflorescence reduction and further modification of the floral ap- paratus, terminal bisexual flowers such as those seen in S. dunnii become the axillary, sessile, apparently bisexual flowers of S. sinensis, with the inflorescence bracts localized in a cycle on the hypanthium rim in each flower. The irregular number and form of the tepals, the irregular number, insertion, and development of the stamens, and the irregular vascular supply to the hypanthium, perianth, and androecium might tend to support such an interpretation. But a better alternative lies in the fact that Sycopsis sinensis is closer morphologically to Parrotia persica, even though the two species differ strikingly in their leaf form, and in the degree of development of their hypanthia. The existence of morpholog- ically intermediate first generation hybrids between them, in conjunction with the morphological and anatomical differences separating S. sinensis and S. dunnii, strongly suggests that the two groups of species in Sycopsis are not congeneric. 3. Parrotia C. A. Meyer, Verz. Pfl. Caucasus 46. 1831 Ficures 8, 9. A monotypic genus, containing only Parrotia persica (DC.) C. A. Mey., a species of small (ca. 5 m.) shrubby trees forming the characteristic arborescent component of the moist deciduous forest on the south shore of the Caspian Sea in northern Iran, where it is endemic. Its leaves are rather similar in appearance to those of Parrotiopsis, Fothergilla, and Hamamelis, often becoming brightly colored in the fall. MorpuHo.ocy: The inflorescence of Parrotia (n = 12, Pizzolongo 1958) consists of 2 to 8 small, inconspicuous, perfect or staminate flowers clus- tered in a terminal or axillary, involucrate, bracteate head. Staminate flowers, when present, are usually basal in the head, and result from reduction of the ovary. The flowers are sessile in the axils of large, broad, primary bracts (B, in Ficure 8A) which are similar to the primary bracts of Sycopsis sinensis in their dark-brown, stellately pubescent abaxial sur- faces and glabrous adaxial surfaces. The primary bracts become sharply reduced upward in the inflorescence, and secondary bracts are generally absent. The perianth consists of a perigon of 5 to 9 (to 10, according to Endress, 1968) green tepals which are irregular in width and length, and bear long, dark-brown, silky tufted hairs at their tips. The tepals are inserted with the stamens on the rim of a shallow hypanthium. Above the hypanthium rim the tepals may be free or irregularly connate and lobed. Ten to four- teen (8 to 15) stamens are inserted irregularly on the hypanthium rim in no particular order with regard to the tepals. The stamens are similar 338 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ficure 8. Parrotia persica: A, view res sonia from a cleared segment a a hypanthium — note stamen Jails, rink bundles rregular easier al supply to the tepal stamen; E-K, transverse sec- tions (sight y obliqu a of a post-anthesis flow B, through receptacle — note ane formed recep ieee st o and partial ‘conor of the hypanthium ; F, a fy ace note * epart ompound lateral bundles from the Abbreviated receptacula stele; G, Sere gk aca bases — note insertion of stamens and parenchymatous collar on hypanthium; H, mid-ovary — note perigon tube and 1970] BOGLE, HAMAMELIDACEAE 339 to those of Sycopsis sinensis in their strongly exserted, dark- red, linear- oblong anthers, each of which contains four pollen sacs and dehisces laterally by means of simple longitudinal slits. The anther connective is produced as a short apiculus (Ficure 8p). On the inner edge of the hy- anthium rim a number of small, vascularized, parenchymatous lobes, or a pe shallow collar, may alternate with or enclose the bases of ‘the stamen filaments (col, Ficure 8F, G). In the mature, non-fruiting flower the collar appears as a thickened and irregularly lobed ring of tissue on the rim, the protuberances of which suggest the abortive development of floral organs. (FIGURE 9A). The hypanthium closely surrounds the base of a superior or nearly Superior 2(—3)-carpellary ovary which is rudimentary at anthesis. The styles are long and recurved, with extensive papillate stigmatic surfaces along the length of their ventral faces. In isolated instances tepals of Parrotia become quite leaf-like in appear- ance, with a petiole and short narrow blade (Ficure 9A), and are similar in this respect to reduced stipulate leaves in the inflorescences of Distylium racemosum or Sycopsis dunnii. Furthermore, in a considerable number of flowers examined, particularly from the large tree in the Arnold Arbo- retum, but in other collections as well, aberrant carpels or bicarpellary ovaries have been observed in different positions around the functional ovary. These are inserted not on the receptacle, but on the inner face or rim of the hypanthium (FicurE 9p, c). From 1 to 3 small ovaries of this type have been found on the hypanthium in single flowers, and in two or three flowers of the same inflorescence, occasionally developed to the point of containing rudimentary ovules. Sections of a flower containing two such ovaries are illustrated in Ficure 9p-F, and their anatomy is briefly described below Anatomy: A pedicellar stele may subtend some flowers, but is scarcely discernible beneath others in the material examined. In the latter case many of the bundles which function in supplying the hypanthium and gynoecium appear to develop almost directly from the margins of a large gap in the peduncular stele, or from a very short and longitudinally dis- torted cylinder of loosely arranged bundles which diverge from the gap. In the receptacle, well below the base of the locules, 10 to 13 prominent bundles diverge from the pedicellar stele to enter the base of the hypan- thium. Within the hypanthium the trunk bundles vary somewhat in the kind and numbers of organs they supply. In general, each trunk bundle divides just below the rim to supply one abaxial trace (te) to a tepal and another larger trace (st) to the base of a stamen (FicurE 8£-G). Varia- tions from this pattern include bundles which supply only a stamen, only a perianth lobe, or two closely set stamens and a perianth lobe. In some young flowers stamens have been found on the rim with no apparent vas- free tepals; 1, os through level of ovule insertion; J, through the style bases; K, throu gh styles L, median longitudinal section semidiagrammatic) throug h Ower; M, transverse section through a tricarpellary ov 340 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 cular connection through the hypanthium between the base of the filament bundle at the level of insertion and the receptacular stele. The number of traces to the base of a particular perianth lobe varies with its breadth and with the number of stamens included within the width of its base. The trunk bundles may appear collateral in transverse sections, or oc- casionally as narrow vascular cylinders. In addition to the bundles described above, lateral branches from the trunk bundles, largely procambial at anthesis, were observed to move adaxially within the hypanthium to end blindly at the base of the lobes of the shallow parenchymatous collar (col) on the inner edge of the rim. The significance of these traces is uncertain, but they could indicate the abortive development of stamens in these positions. n the ovary at anthesis only the dorsal carpel bundles are apparent, usually as protoxylem strands. The septum and ovules are scarcely, if at all, developed. In the post-anthesis ovary the dorsal bundles are prom- inent, and develop in close association with the hypanthial bundles (FicurE 8k, left side). The long internode apparent between the separa- tion of the hypanthial bundles and the carpel dorsal bundles from the receptacular stele in Sycopsis sinensis is absent in the receptacle of Par- rotia persica. The compound lateral bundles (ll) originate from the receptacular stele in the transverse plane well above the departing dorsal bundles, and ascend as multi-stranded vascular bands at each edge of the septum. The central bundles in each band consolidate into a broad vascular arc in the upper half of the ovary. In polarized light two distinct protoxylem strands are apparent in each arc, indicating two adjacent lateral bundles. The in- dividual lateral bundles separate at about the level of ovule insertion (FIGURE 8F-1). Above the departure of the compound laterals the remaining portions of the receptacular stele consolidate to form a narrow and largely pro- cambial cylinder. At about mid-height in the septum the cylinder divides in the median plane to form two largely procambial compound ventral bundles (vv, Ficure 8H, M). The latter divide transversely just below the level of ovule insertion, and the individual ventral bundles shortly supply traces to the ovules, then fuse with the adjacent lateral bundles and pass upward as the marginal bundles of the styles (FicurE 8H-K). In the base of some ovaries examined a few small procambial bundles occur which are similar to the peripheral bundles in the base of the ovary of Distylium, They originate from the receptacular stele well above the departure of the hypanthial bundles and mainly in the vicinity or even from the base of the departing compound laterals. They are deflexed, and appear to end blindly near the periphery of the ovary base, near or just below the level of separation of the hypanthium from the ovary. Occasion- ally branches from such a bundle can be traced into and upward in the hypanthium, where they end blindly in or near the parenchymatous collar, and upward in the ovary wall above the level of their separation from the stele. 1970] BOGLE, HAMAMELIDACEAE 341 E 9. Parrotia persica: A, leaf-like tepal on hypanthium rim; 8B, whole Fic tec 4 near tepals a the single cota ime ge a hypanthium, with suture directed outward; c, section of hyp: bearing the carpel; D-F, transverse epic through a flower es on asia ap Gates on the ince (slightly oblique); p, through base of functional ovary and hypanthium (above) — note the two steles in the hypanthium; £, ovaries and abaxial stamens and tepals forming on ‘the nthium; F, areas of ‘perianth and androecium supplied by each stele siieatodl as zones 1 & 2. In those flowers containing anomalous ovaries on the hypanthium rim, anatomical study shows that a substantial vascular cylinder is pinched 342 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 off from the pedicellar stele and enters the hypanthium beneath each ovary. Most of the bundles in each cylinder are used up in supplying the ovary, but tepals and stamens adjacent and abaxial to the ovary on the rim also receive their traces from the cylinder. In FicuRE 9p-F the sectors (1 and 2) of the perigon and androecium supplied by two such steles are indicated. However, no regular pattern of insertion and vascular supply of tepals and stamens relative to the anomalous ovaries was apparent. The morphological resemblances between Parrotia persica and Sycopsis sinensis are striking, and the description of the intergeneric hybrid Sycoparrotia confirms their close affinity. They are very similar in their capitate inflorescences; in the form and pubescence of the primary inflores- cence bracts; in the pubescence, variable number, and extremely irregular form of their tepals; in the irregular number and insertion of the stamens, and the similar form, dehiscence, and reddish color of their anthers; in their essentially superior ovaries; and in the irregular vascular supply to the hypanthium and its attendant organs. In Parrotia persica the number of tepals (5 to 10) and stamens (8 to 15) is usually greater, and the tepals considerably longer and broader, than in Sycopsis sinensis, which has 4 to 7 tepals and 5 to 10 stamens. In S. sinensis the hypanthium is longer, and the basal connation of its tepals into a perigon tube above the hypanthium rim is more pronounced than in Parrotia persica. In fruit the shallow hypanthium of Parrotia is not ruptured, as in Sycopsis sinensis. But Parrotia exhibits certain morphological and anatomical features in common with Distylium, and to a limited extent with Sycopsis dunnii. The tepals of Parrotia, in particular, suggest varying manifestations of the reduced stipulate leaves which form the inflorescence bracts and blade among the tepals of Parrotia enhances this interpretation of their morphological nature. Also striking in Parrotia is the presence in the receptacles of some flowers of peripheral bundles which supply branches both to the base of the ovary and the base of the hypanthium, as in Sycopsis dunnii. These are similar in origin and appearance to those of Distylium, but not as numerous or as strong On the basis of the available evidence it appears possible that the flower of Parrotia persica may represent a pseudanthium of a more ad- vanced type than that seen in Distylium, developed through the complete reduction of an inflorescence axis, and resulting in the aggregation of reduced stipulate leaves (bracts) of subterminal nodes, and modified naked staminate flowers of suppressed lateral axes, around the functional ovary of the naked terminal pistillate flower. I have observed terminal and lateral pseudanthia approaching this level of reduction in some rela- tively reduced inflorescences of D. gracile and D. buxifolium. The union and elevation of the bracts and modified staminate flowers on the shallow 1970] BOGLE, HAMAMELIDACEAE 343 hypanthium may explain the variability in the number of bundles supply- ing the base of the hypanthium, and the variability in their form. The peripheral bundles of the receptacle may be attributable to suppressed lateral organs of the condensed inflorescence axis, and the appearance of abortive carpels or ovaries on the hypanthium in otherwise normal flowers may be due to the aberrant development of pistils attributable to the terminal ovaries of lateral axes which are usually suppressed in the pseu- danthium. The occurrence of these anomalous ovaries, and leaf-like bracts, on the hypanthium is certainly teratological, and of only secondary im- portance as evidence. But such organs have not been observed on hypan- thia in any other genus of the family, and should not be ignored. The interfertility of Parrotia persica and Sycopsis sinensis indicates that they are relatively recent derivatives of a common ancestral stock. If the above interpretation of the Parrotia flower is correct, then the longer hypanthium of Sycopsis sinensis may represent only an elaboration in length of a shorter ancestral hypanthium, perhaps as in Parrotia, with concomitant reduction in the number of bracts and stamens inserted on the rim of its narrower throat, and in increased basal connation of the tepals above the rim. 4. Fothergilla J. A. Murray, Syst. Veg. ed. 13. 418. 1774. Ficure 10. The genus Fothergilla consists of two species of low deciduous shrubs which usually reach a height of 1 to 3 meters and frequently form dense clumps through rhizomatous proliferation (Ernst 1963; Weaver 1969). The species are limited in distribution to the coastal plains and uplands of the southeastern United States. MorpuHorocy: The inflorescence of Fothergilla contains numerous ses- sile flowers solitary in the axils of broad primary bracts spirally arranged in a terminal spike (FicurE 10A). The flowers are incomplete, perigynous, and generally bisexual, although a few basal flowers in the inflorescence may be functionally staminate, containing only a rudimentary ovary. The perigon and androecium are inserted on a shallow, campanulate hypanth- ium which surrounds the ovary and is fused with it for its lower third. The perianth consists of 5 to 7 (—8) small, irregular tepal lobes (te) of variable size and position (Ficure 10a, 1-K). I found 18 to 24 stamens in flowers of Fothergilla gardenii (n = 24, Weaver 1969), and up to 31 stamens in those of F. major (n = 36, Weaver 1969; Weaver found 12 to 24 stamens in F, gardenii, and 22 to 32 in F. major), inserted in a single cycle on the rim of the hypanthium with their bases enclosed externally by the lobes of the perigon and internally by a shallow, much lobed inner parenchy- matous lip (col) of the hypanthium (FicureE 101-K). The distally clavate, unequal filaments are several times longer than the hypanthium, and bear small, basifixed anthers which contain four pollen sacs. Anther dehiscence is subvalvate and lateral by means of longitudinal I-shaped slits (FicurE 10c). There are no staminodia. The ovary is up to one-fourth semi- inferior. The two carpels terminate in filiform styles with inconspicuous 344 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ficure 10. Fothergilla gardenii: A, a flower at anthesis, stamens and styles cut away; B, floral diagram; c, stamen; D-J, transverse sections through a flower at anthesis; D, through receptacle — note dorsal bundles and receptacular stele; £, bases of the locules — note compound lateral bundles and residual receptacular stele; ¥, partial separation of the hypanthium, compound ventral bundles formed; 1970} BOGLE, HAMAMELIDACEAE 345 papillate stigmas within the folded margins of the distal half. Each locule contains one or occasionally two ovules, the lower of which is fertile. ANATOMY: The vascular supply to the base of each flower varies from a short but cylindrical pedicellar stele to a reduced condition in which a variable number of large bundles enter the receptacle directly from the margins of an elongate gap in the peduncular stele. In the receptacle, well below the base of the locules, several large trunk bundles (tb) diverge from the pedicellar stele to enter the base of the hypanthium, eventually to supply the perianth and androecium. An inter- node is not present in the receptacular stele between the trunk bundles and the carpel dorsal bundles, and the latter are formed at either side of the receptacle in the median plane by the fusion of 2 to 3 lateral branches from single or adjacent trunk bundles (d in Ficure 10p, ©). Above the departure of the associated trunk and dorsal bundles the remaining bundles of the receptacular stele converge on the center of the receptacle to form a narrower receptacular cylinder (Ficure 10p) which then shortly gives rise to two compound lateral bundles (Il) to either side in the transverse plane. The remaining traces in the stele consolidate to form a still nar- rower and largely procambial cylinder which enters the base of the septum. In polarized light at least two strands of protoxylem appear in this cyl- inder, situated on opposite sides in the tranverse plane (FicuRE 10E). The number of bundles entering the base of the hypanthium approxi- mates half the number of stamens eventually supplied, but there is no exact correlation in this respect. Within the hypanthium the number of trunk bundles is increased by branching. Rarely, lateral branches from adjacent bundles fuse to form a single trunk bundle. More often, minor lateral branches of the trunk bundles end blindly in the hypanthium slight- ly below the level of stamen insertion. Each stamen receives a single trace. A single trunk bundle may supply only one stamen, but many of the trunk bundles branch immediately below the insertion of the stamens to form 2 to 3 stamen traces. In addi- tion, almost every trunk bundle produces a single, relatively minor trace (te) at this level, which enters the base of a tepal. Occasionally a rela- tively weak hypanthial bundle will supply only a tepal. There was no apparent regularity in the vascular supply to the tepals. Within the tepal the traces may branch, and anastomoses may form between adjacent branches. In the ovary the division of the compound ventral (vv) and compound lateral (ll) bundles to form distinct ventrals and laterals usually occurs slightly below the level of ovule insertion, as figured by Horne (1914), but occasionally the compound laterals were observed to divide in the lower half of the ovary, or to remain entire to a level slightly above ovule in- sertion. The free ventral and lateral bundles of each margin then fuse, G, ovary — note fusion of compound lateral and compound ventral bundles; 4, ovule insertion; 1, insertion of the stamens on the anthium; J, thr ough the various organs of the flower; K, —— median saaediiial hand section (semi- diagrammatic) of a flower at anthesis 346 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 51 and immediately above the point of fusion an ovular trace separates from the ventral bundle of each fertile margin and enters the funiculus of the ovule (FicurE 10H). At anthesis the upper portions of the ventral bundles are frequently in only a procambial condition, and in some carpels the ventral bundle to the sterile margin appears to die out soon after separating from the adjacent ventral which supplies the fertile margin. Fothergilla, one of the better known genera of Hamamelidoideae, is widespread in cultivation and easily accessible for study. Baillon (1871) described the organogeny of Fothergilla alnifolia L. (= F. gardenii Murr.). According to his observations 5 to 7 short sepal primordia develop succes- sively on the rim of a shallow, cupulate receptacle. The first and second primordia appear on either side, then the others develop in the intervals between the first two. Later, one stamen primordium develops before each small sepal, then 2, 3, or 4 stamen primordia appear secondarily on the sides of each initial stamen. Baillon considered the smaller, secondary stamen primordia opposite each calyx lobe to be exterior to the primary stamen initial, and interpreted the groups of stamens as having arisen through the divisions of a primitive stamen opposite each sepal. He did not interpret his observations in terms of cycles of stamens. Tong (1930), however, states that the normal number of stamens is 14 in Fothergilla major, arranged in two whorls, the outer of which al- ternates with the calyx lobes. He considered the outer whorl of stamens derived through the transformation of petals to stamens, a view which he attributes justifiably also to DeCandolle (1830), and Griffith (1838). Excess stamens were interpreted as being derived through splitting of stamen primordia. Tong’s floral diagram accordingly shows the stamens arranged in two cycles. Shoemaker (1905) observed some stages of the organogeny and em- bryology of Fothergilla gardenii. His observations on the development of the stamens agree with those of Baillon. Flint (1957) has provided a detailed description of pipe accra and megagametogenesis in both Fothergilla gardenii and F.m Horne (1914) illustrated his ‘brief description of the floral vasculature of Fothergilla major with three drawings of transverse sections at critical levels. My observations on Fothergilla are in general agreement with those of Horne. There is a slight difference in the relative positioning of the stamens on the rim of the hypanthium, so that the stamen opposite the mid-line of each tepal appears slightly adaxial to its neighbors. This is especially apparent in those flowers having more numerous stamens (Fothergilla major), but it is not sufficiently pronounced either morphologically or anatomically to provide a basis for recognizing more than one cycle of stamens on the hypanthium The lobed collar on the inner edge of the hypanthium rim of Fothergilla is similar to that seen in Parrotia persica, but there is no vascular supply 1970] BOGLE, HAMAMELIDACEAE 347 / ase loé % @) Figure 11. Parrotiopsis jacquemontiana: A, flower at anthesis, me ger cut away; B, floral diagram; c, stamen; D-K, transverse sections through a mend ge dice , through the receptacle — note sepiee bundles differentiating; r, bases of the locules; G, ovary — note diffuse Sy hae lateral bundle traces; H, partial separation of the hypanthium, compound later: ty) J, ovule insertion; x, through the style bases; L, median longitudinal hand-section (semidiagrammatic) of a flower at anthesis. 348 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 evident to its base. Nor is there any evidence in the receptacle of peri- pheral bundles or vestigial vasculature of any kind. The loss of an inter- node in the receptacular stele, resulting in the close association or partial fusion of the carpel dorsal bundles with the hypanthial trunk bundles, is also seen in Parrotia, but in both genera a segment of the receptacular stele intervenes below the departure of the compound lateral bundles, and continues upward into the base of the septum. The origin of the ovular traces above the fusion of the lateral and ventral bundles is a deviation from their departure prior to fusion in Distylium, Sycopsis, and Parrotia. 5. Parrotiopsis (Niedenzu) Schneider, Handb. Laubholzk. 1: 429. 1905. FIGureE 11. This monotypic Asian genus of deciduous shrubs or small trees is known only through the species Perrotiopsis jacquemontiana (Decne.) Rehd., which has a restricted distribution in the Himalayan mountains of northern Kashmir, West Pakistan, and Afghanistan (see Kammeyer 1957, p. 73, for distribution map). Morpuo.ocy: The inflorescence of Parrotiopsis (n = 12, Anderson & Sax 1935) contains about a dozen small, inconspicuous flowers arranged in a terminal subcapitate spike. The blades and stipules of the leaves at the two or three lowest nodes of the inflorescence are conspicuously ex- panded into spreading white bracts which form a showy involucre around the flowers. A single sessile flower, usually staminate through reduction of the ovary, is axillary to each set of involucral bracts. Primary bracts may subtend a few flowers just above the involucre, but are absent at the distal nodes of the inflorescence, and secondary bracts were not present in the material examined. The flowers are regular, apetalous, perigynous, and usually bisexual, with a semi-inferior ovary (Ficure 11H, L). The perianth and androecium are inserted on the rim of a shallow, cupulate hypanthium which is fused basally with approximately the lower third of the ovary, while the free portion of the hypanthium forms a shallow cup around the middle of the ovary. The limb of the perianth is reduced to irregular lobes on the margin of the hypanthium. Occasionally one or two tiny, pubescent tepals, less than 1 mm. long, develop. About 21 to 25 stamens with unequal, terete filaments, are inserted in a single cycle on the rim of the hypan- thium. The small, oblong, truncate anthers contain four pollen sacs and dehisce laterally by means of four valves (FicuRE 11c). The anther con- nective may be produced as a very short apiculus on the adaxial edge of the truncate tip. A very shallow, thin, parenchymatous collar on the inner edge of the hypanthium encloses the stamen filament bases. In the ovary one or occasionally two ovules are found in each locule. The styles are linear, with papillate stigmas running nearly the entire length of the ventral margins. ANATomy: A cylindrical pedicellar stele is present beneath each flower. In the receptacle the stele expands as 13 to 19 trunk bundles differentiate 1970] BOGLE, HAMAMELIDACEAE 349 and diverge toward the periphery to enter the base of the hypanthium (Ficure 11p-H). Adnation between the bundles of the hypanthium and gynoecium in Parrotiopsis is relatively advanced over that in Parrotia and Fothergilla, as evidenced by the fact that there is no indication of a vascular cylinder in the receptacle. The 13 to 19 trunk bundles (tb) which enter the base of the hypanth- ium eventually supply 21 to 25 stamens, as well as the tepals. As in Fothergilla, each stamen receives a single trace (st) from a trunk bundle, and some trunk bundles divide within the hypanthium to form additional bundles, or divide just below the insertion of the stamens to form two or three stamen traces (FIGURE 11F-1). The limb of the perianth is so reduced that few of the lobes were vas- cularized in the material which I examined. The few traces (te) produced by the trunk bundles to the tepal lobes are very inconspicuous at anthesis. This is in contrast with Fothergilla, in which almost every one of the numerous trunk bundles produces a single abaxial branch to the perianth. Because of this reduced condition I am unable to confirm Saunder’s ob- servation (1939) that the stamens are supplied from both the midveins and lateral veins of the calyx lobes. In the receptacle the gynoecial bundles appear to have independent origins in the fusions of branches from the bundles of the expanding pedicellar stele, rather than arising from a cylindrical receptacular stele. At either side of the receptacle in the median plane 3 to 4 small branches from diverging trunk bundles converge and fuse to form a carpel dorsal bundle (d, Ficurr 118, F). In the transverse plane 4 to 6 traces converge centripetally to function in the formation of the compound ventral (vv) and compound lateral bundles (ll). The compound ventrals are often the first to differentiate, in some flowers appearing as a single bundle on each side of the receptacle (FicurE 11c, E), while the comopund laterals form at a slightly higher level as an aggregation of 4 to 5 weak bundles arranged in a loose band at each edge of the septum (Ficure 11F, G). In the ovary the several bundles of each compound lateral consolidate distally to form a narrower band, or a single broad bundle which dicho- tomizes at about the level of ovule insertion to form the independent lateral bundles (1), The compound ventrals divide above the middle of the ovary, forming four ventral bundles (v) which fuse with the adjacent laterals at the level of ovule insertion. Ovular traces (ov) depart from the ventrals of the fertile margins just prior to this fusion (Ficure 11H- ‘F). The genus Parrotiopsis, like Fothergilla, is also widespread in cultiva- tion, and has been studied by several workers. Horne (1914) included two figures of transverse sections illustrating the basic vascular pattern of the ovary (as Parrotia jacquemontiana), and in one of these two ovules are conspicuous in each locule. Tong (1930) interpreted the stamens of Parrotiopsis as forming three alternating septenate cycles within a cycle of seven calyx lobes. Saunders (1939) described the origin of the stamen traces as being “detached from the sepal lateral veins as well as from 350 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Ficure 12. Molinadendron sinaloense: a, view e a flower at eee B, nearly ot fruit— note shallow hypanthium; c, flora gra’ nsverse sections of a flower at anthesis 5 (alightly ’ oblique) ; D, Gisanh she receptacle, se seo and stamen traces forming; E, receptacle, gynoecial bundles forming (d, 11); F, bases of the tae in insertion of some te pals and stamens; G, stamens separating from base of the ovary; H, about mid-height in the ovary; 1, base of ovule insertion; J, top of ovule insertion; K, through the style be [Fe oo” ia & | 1970] BOGLE, HAMAMELIDACEAE 351 the midrib bundles and [standing] in a single ring.” Kaul (1969) found that the endosperm is ad initio cellular in Parrotiopsis The systematic affinities of the genus are indicated by the fact that at one time or another, Parrotiopsis jacquemontiana has been included in both Fothergilla (Niedenzu 1891) and Parrotia (Decaisne 1844), as well as by Rehder’s statement (1920) that Parrotiopsis differs from Fothergilla “only in the capitate inflorescence supported at the base by large bracts, hea in the less numerous stamens with linear not club-shaped filaments .”, a view expressed earlier by Hooker (1896). However, Parrotiopsis is more advanced than Fothergilla in its fewer-flowered, more congested inflorescences, in the extreme reduction of its perianth, and in the slightly more pyendduend adnation between hypanthium and ovary. It is also more advanced anatomically in the absence of a residual receptacular cylinder, resulting in basal adnation between the hypanthial and gynoecial vascular systems. The diffuse origin of the compound lateral bundles, and the individuality of the compound ventrals in the receptacle, are apparently artifacts of the reduction of the receptacular stele and the greater adnation between hypanthium and ovary. I could find no morphological or ana- tomical basis for assigning stamens to more than one cycle on the hypan- thium. 6. Molinadendron Endress, Bot. Jahrb. 89: 353. 1969. Ficure 12. The genus Molinadendron is a small Central American group recently segregated from Distylium Sieb. & Zucc. (cf. Harms 1933; Walker 1944; Gentry 1948; Balgooy 1966). The genus contains three as yet little known species of large evergreen trees. Molinadendron guatemalense (Radlk. ex Harms) Endress is apparently restricted to the vicinity of Coban, Guatemala. Molinadendron hondurense (Standl. in Walker) En- dress occurs widely in Honduras, while M. sinaloense (Standl. & Gentry) Endress is restricted to the mountains of the State of Sinaloa, Mexico. The following descriptions are based primarily on material of M. sinaloense which I collected in 1964. In addition, flowers from herbarium sheets of the other two species were also examined anatomically, and showed no significant difference from those of M. sinaloense. Morpuotocy: The inflorescences of Molinadendron sinaloense are axillary or terminal on short lateral leaf-bearing shoots of the current season’s growth. Each consists of about a dozen small inconspicuous flowers spirally arranged in a short, congested spike. The proximal flower may be axillary to a much reduced stipulate leaf, the blade of which may be little more than a narrowly triangular bract. Intermediate flowers in the inflorescence are subtended by obvious primary bracts which are more or less hooded and cover the flower in bud; but at distal nodes the primary bracts are reduced and adnate to the base of the flower, often intergrading with the tepals of the perigon. Similarly, a pair of subopposite secondary longitudinal section through a post-anthesis ovary, rather oblique — note slightly perigynous insertion of stamens 352 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 bracts may subtend the lowest flower or two, but at higher nodes these either fail to develop, or are adnate to the floral receptacle and indistin- guishable from the tepals. It is often easier to fix the presence and posi- tion of these bracts in the elongated, more open fruiting inflorescence than ut anthesis. The flowers at anthesis are sessile or nearly so, incomplete, subperigyn- ous, and bisexual. The perianth consists of 2 to 8 linear to broadly cune- ate-dentate and sometimes cucullate tepals (or fewer if some of these are secondary bracts). Petals are absent. Two to eight (usually 5 to 7) bright-red stamens are arranged in a single cycle around the base of the sessile, broad-based ovary. The anthers, which are short-oblong with a truncate apex and a short adaxial apiculus, are borne on short terete filaments which elongate after anthesis. Each anther contains four pollen sacs, and dehisces laterally by means of four narrow, longitudinal valves. The two- (occasionally three-) carpellary ovary appears superior but is slightly sunken in the broad receptacle, and hence slightly inferior. It is incompletely bilocular, and contains one or sometimes two (then one on each margin) ovules in each locule. The placentation is parietal in my material, rather than axial as described by Endress (1969) (cf. FIGURE 121). The styles are exserted, recurved, and somewhat expanded distally, with extensive papillate stigmatic surfaces. he basal flower(s) and sometimes the terminal flower in an inflores- cence may be staminate through reduction of the ovary. Abortive stamens, reduced in varying degrees to subulate sterile filaments (sto in FIGURE 12A), and abortive linear, bristle- or scale-like, pubescent perianth bracts frequently appear in any part of the inflorescence. A striking feature of the flowers is the irregularity in number and insertion of the tepals and stamens. There appears to be a rough correlation between the number of stamens and the number of tepals in the perianth. They are often equal, with a stamen opposite each tepal, but frequently the number of stamens is larger, and their insertion irregular. And in some flowers the stamens alternate with the tepals. In fruit the marcescent tepals and stamens, or their basal portions, are prominent on the rim of the very shallow hypan- thium around the base of the nearly superior bony capsule (FIGURE 12B). Anatomy: In the pedicel a short cylindrical stele assumes an elliptic configuration as its vascular strands become organized into traces to the floral organs. Each tepal (te) receives a single trace, as does each stamen (st). Opposing tepals and stamens frequently receive their traces from a common trunk bundle in the receptacle, but this is not a constant feature, since, in some cases, each may be supplied by a trace derived independent- ly from the stele. Alternating tepals and stamens each receive independent traces from the stele. When two stamens oppose a single tepal each stamen is supplied by an independent stamen trace, rather than by branches of a single stamen or tepal trunk bundle. In transverse sections the adnation of the perianth and androecium to the base of the ovary is evident (FIGURE 12F,G b 1970] BOGLE, HAMAMELIDACEAE 353 In the receptacle, the carpel dorsal bundles may be fused basally for a short distance with some of the traces supplying the perianth and androe- cium as a result of the loss of an intervening internode in the receptacu- lar stele. However, a short segment of the stele persists above the de- parture of these traces, giving rise to the compound lateral and compound ventral bundles. At the median sides of the expanding receptacular stele, 2 to 3 small vascular strands from diverging tepal or stamen bundles move centripetal- ly to associate or fuse into a dorsal bundle (d in Ficure 128, F). In the base of the ovary a dorsal bundle may be entire (Ficure 12k, right side) or consist of two large collateral bundles which then fuse at a higher level into a single broad bundle (FicureE 12«-¢, left side). Above the departure of the sepal, stamen, and dorsal bundle traces the residual bundles of the receptacular stele move toward the center of the receptacle. In the plane of the septum 1 to 3 of these on each side cease their centripetal movement and assume a position about midway between the periphery and the center. These form the compound lateral bundles (Il) at each edge of the septum. The remaining residual bundles converge and form a narrow cylinder in the base of the septum (Ficure 12k, F). A compound lateral may appear as a single broad bundle in the base of the ovary, or as a group of 2 to 3 vascular strands which fuse above into a common compound bundle (Ficure 12E-H). The compound laterals usually dichotomize transversely at the level of ovule insertion or slightly above. The narrow residual receptacular cylinder divides about midway up the septum to form two compound ventral bundles (vv), each containing 1 to 3 strands of protoxylem, which diverge toward the edges of the septum. At anthesis the compound ventrals and laterals may vary in form in the upper part of the septum and parietal placentae. Usually each dicho- tomizes in the transverse plane at about the level of ovule insertion to form individual ventral and lateral bundles which then fuse in pairs and pass into the styles as marginal bundles. In this case the ovular trace originates from an individual ventral bundle prior to its fusion with the adjacent lateral. Sometimes, however, the compound ventral and lateral bundles fail to divide, but fuse in the placenta, and the bundle thus formed then dichotomizes at a slightly higher level to form the marginal bundles of the styles. In this case the ovular trace may originate from the com- pound ventral either below or above its fusion with the compound lateral. Exceptionally a compound ventral bundle will pass into the funiculus of the ovule without dividing, with no discernible connection to the adjacent compound lateral bundle. The latter condition may be due to the im- mature state of the ovary and its vasculature at anthesis, or it may repre- sent a form of reduction in the vascular pattern of the ovary concomitant with reduced size, It has been noted in other genera of the family as well. The flowers of Molinadendron differ from those of Distylium in their spicate arrangement; in the presence of a perianth; in their bisexual condition (or occasionally staminate through reduction of the ovary); 354 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 in the valvate dehiscence of the anthers; in the slightly inferior condi- tion of the ovary, and in the persistence of the sepals and stamens on the rim of a very shallow hypanthium around the base of the fruit (FicurE 128. See also Gentry 1948, figure 1). Anatomically, the vestigial peripheral bundles of the Distylium ovary are absent, the carpel dorsal bundles are fused basally with the tepal and stamen traces, and fusion of adjacent lateral and ventral bundles in the septum and placentae tends to be more advanced. Perhaps the most striking feature of the flowers is the irregularity in the numbers and insertion of the tepals and stamens, and the intergrada- tion of the primary and secondary bracts with the tepals. One is fre- quently at a loss to know which is which. Endress (1969) concluded that ““Molinadendron does not even belong to the tribe Distylieae (Disty- lium, Sycopsis, Matudaea), but shows more relationship to Fothergilla/ Parrotiopsis and Fortunearia/Sinowilsonia.”’ However, the pollen morph- ology of Molinadendron differs markedly in the form of its exine reticu- lum, and in the shape and sculpturing of its apertures (Chang 1960, Bo- gle, unpublished scanning electron micrographs), from that of Fothergilla and Parrotiopsis, and is closer to that of Sinowilsonia and Distylium. The distinct one-trace tepals of Molinadendron also differ from the irregularly vascularized tepals of Fothergilla. e genus Fortunearia differs from Molinadendron most strikingly in its regular, complete, usually pentamerous flowers, as well as in its numerous-flowered racemose inflorescences, subvalvate anther dehiscence, rescences are well developed, and the flowers are racemosely arranged, regular, more or less complete, and usually pentamerous, with an urceolate hypanthium surrounding the superior ovary in the pistillate flowers. Sta- men dehiscence in the staminate flowers is subvalvate, by means of I- shaped slits, as in Fortunearia. These morphological differences, in conjunction with its New World distribution, suggest that Molinadendron occupies a relatively isolated position among the apetalous Hamamelidoideae. 7. Matudaea Lundell, Lloydia 3: 209. 1940 FicurE 13. The genus Matudaea consists of only two poorly defined species of large, possibly evergreen trees distributed in Mexico and Central Ameri- ca. Matudaea trinervia is known from a few scattered locations in Hon- duras, Guatemala (Standley & Steyermark 1946), and the State of Chiapas, Mexico. Matudaea hirsuta Lundell (1961) is known only from a small area in the southern part of the State of Mexico, and differs from M. trinervia principally in its densely hirsute twigs. Specimens more or less intermediate between these two species have been collected in the State of Jalisco, Mexico, by McVaugh ef al. (personal communication). The following descriptions are based primarily on flowers of Matudaea BOGLE, HAMAMELIDACEAE URE 13. Matudaea hirsuta: A, flower at perk B, floral diagram; c, post- canoes flower — note wrap scars crowded beneath the — ova D, stipulate primary bract; ©, deciduous syntepalous ora F-M, transverse sec- tions through a flower e anthesis (slightly Ea & F, ea stele and departing perianil bundles; G, through receptacle, sepal traces departing to right, stamen traces to the left; ee receptacle, part of gynoecial vasculature formed igi peri e wall; J, about mid-height in ry; K, ovule insertion; L, through fused carpels above the ] ; ugh style ; ectio through ovary just above the ae of I, cepa slightly to show vestigial vas- culature; 0, ug ty) of M. trinervia—note vestigial ection through bas vasculature; P, near-median enaieadinal section through an ovary. 356 JOURNAL OF THE ARNOLD ARBORETUM [VoL 51 hirsuta which I collected in Mexico in 1964. Flowers of M. trinervia from herbarium specimens were also examined anatomically, and appear to agree with those of M. hirsuta Morpuorocy: The inflorescences of Matudaea are small, congested, and either spicate or paniculate. They are borne in the axils of the two- ranked leaves. Both the leaves and their associated inflorescences become abruptly reduced toward the tip of the shoot axis. The distal leaves may take the form of a much reduced lamina, no larger than its flanking stipules, or of a bract either free from, or variously fused with the two bracteate stipules. At the distal nodes the inflorescences become reduced to a single, short pedicellate, or sessile flower, and the tip of the shoot axis may become similar in appearance to the lateral inflorescences along its length. Larger inflorescences in my collections attain a length of about 2 cm., and bear one or two short secondary axes at the proximal nodes. The secondary axes may bear one or more flowers Flowers are situated in the axils of bract-groups which are arranged along the primary and secondary axes in the same one-half phyllotactic sequence as that of the leaves on the shoot axis. The bract-groups repre- sent a reduced leaf and its stipules, an interpretation which is supported both by the obvious reduction of the leaf along the shoot and inflorescence axes, and by nodal anatomy. There has been much reduction in the inflorescence. Failure of the primary axis to elongate sufficiently at its base results in congestion of proximal secondary axes in the leaf axil, so that two or more axes may appear to have distinct origins in the axil. Failure of internode elonga- tion at the distal end of an axis results in congestion and eventual abor- tion of subterminal flowers or axes. The compound nature of some few- flowered spikes is frequently indicated by the presence of reduced bracts o The flowers are regular, incomplete, and eed The perianth con- sists of an irregular, syntepalous, circumscissile perigon. The androecium contains 18 to 30 hypogynous stamens, inserted in a congested, shallow ring around the base of the ovary (Ficure 13c, Pp). The oblong anthers are subsessile in bud, but strongly exserted at maturity. Each contains four pollen sacs, and dehisces laterally by means of four longitudinal valves. The anther connective is produced as a short apiculus. The two- carpelled superior ovary contains one or occasionally two ovules in each locule. Each carpel terminates in a free, elongate, recurved style, with expanded papillate stigmatic margins In bud the perigon forms a cap which tightly encloses the stamens and ovary. At anthesis the cap splits irregularly and longitudinally from an apical pore, expanding to form an irregularly lobed cupulate or reflexed perianth tube (Ficure 134, c, £E). The margins of the lobes are incised 1970] BOGLE, HAMAMELIDACEAE 357 with many often capitate fimbriae. The tube is shed by means of a circumscissile abscission layer which forms at its base around the rim of the slightly expanded receptacle. There is no morphological indica- tion of the number of tepals incorporated into the tube. In some flowers the stamens appear to be inserted in a more or less spiral fashion, but in others, particularly those with fewer stamens, there is a faint tendency toward an irregularly whorled arrangement approaching three alternat- ing cycles (cf. floral diagram, Ficure 138). There is no external evi- dence of staminodia between the stamens and the ovary, ANATOMY: The cylindrical pedicellar stele subtending each flower ex- pands rapidly as it enters the receptacle, with some dichotomizing of its bundles. There is little or no regularity or pattern in the vascular sup- ply to the various floral organs. A particular bundle in the receptacle may supply a single organ, or through divisions and anastomoses con- tribute to more than one A number of traces (9 to 12) diverge from the pedicellar stele to supply the base of the perianth (te in FicurE 13F, G). Some of these undergo branching, and the branches of adjacent bundles frequently anastomose, while still within the receptacle. Although usually irregular and complex, the patterns of origin, branching, and anastomosis of the perianth traces in some flowers approach the pattern seen in the nodal anatomy of the foliage leaf (as described in Distylium), and suggest the connation of modified stipulate bract groups to form an irregular perigon. In this respect it should be noted that the primary and secondary bracts sub- tending the flowers have incised margins with capitate fimbriae like those of the perianth lobes (FicurE 13p, E). Above the departing perianth traces the supply to the androecium forms. In many cases the stamen trace is formed by the fusion of two bundles in the receptacle, and sometimes of three. Occasionally a single bundle will branch into three, the central trace of which will supply the outer (or lower) of two stamens lying on a given radius, while the two lateral bundles fuse to form the trace to the inner, and morphologically higher, stamen. bs As the numerous traces to the androecium diverge, the remaining bundles of the receptacular stele converge centripetally to form the vas- cular supply to the gynoecium. In transverse section at this level this short internode of the receptacular stele appears as a complex vascular plexus, as seen toward the right side of FicurE 13H, with part of the gynoecial system evident (dorsal bundle, compound lateral bundle) to- ward the left side. In the base of the ovary two dorsal bundles (d), two compound lateral bundles (ll), and a narrow residual receptacular cylinder are apparent. Each of these has a complex and independent origin in the anastomosis of several vascular strands. Also at the base of the ovary one encounters a series of vestigial bundles, suggestive of the peripheral bundles in the ovary of Distylium, of which the morphological origins are not known 358 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 (peri in Ficure 130, of Matudaea trinervia, Standley 25570). These may be entirely procambial at anthesis, or contain small strands of pro- toxylem, and appear to branch and anastomose to a certain extent. In flowers of Matudaea hirsuta in my collections, similar vestigial bundles were found originating from the bases of the dorsal and compound lateral bundles (FicurE 13N). These are similar in this respect to certain hy- panthial bundles in several genera of petaliferous Hamamelidoideae (Ha- mamelideae) which are adnate basally to the gynoecial system, but eventually separate to supply organs on the hypanthium In the ovary, the residual receptacular stele dichotomizes in the septum just above the base of the locules. The two compound ventral bundles formed ascend to the placentae where each fuses with the adjacent com- pound lateral bundle. Just prior to this fusion each compound ventral usually provides a single ovule trace to one of the solitary ovules. But ovular traces have also been observed to depart above the point of fusion (FicurE 13k). Slightly above the level of fusion the compound bundle formed dichotomizes to provide marginal bundles to the style bases. The inflorescences of Matudaea are similar to those of Distylium race- mosum in their progressive reduction upwards along the shoot, resulting in the merger of inflorescence and shoot distally, and in the abortion of subterminal flowers beneath the terminal flower, due to congestion caused by the failure of the distal internode to elongate. It often appears that in the last stages of its reduction, the subterminal flower develops as a small knot of capitate fimbriae at the base of the terminal flower. But the flowers of Matudaea are apparently bisexual, with a definite perianth, rather than unisexual or obviously pseudanthial and atepalous, as in Distylium, and there is no direct morphological evidence that the stamens of Matudaea are attributable to subterminal naked staminate flowers aggregated about the base of a terminal naked pistillate flower, as in Distylium. Matudaea thus stands apart from the other apetalous Hama- melidoideae by virtue of its calyptrate perianth and numerous hypogynous stamens, the latter being approached only by the fewer and subperigynous stamens of Molinadendron. Its pollen is also distinct, and has apparently not been described in the literature. Anatomically there is no parallel among the other apetalous genera to the complex vasculature of the perianth. The vasculature of the ovary, like that of Molinadendron, is relatively advanced in the fusion of com- pound ventral and lateral bundles in the placentae, rather than of indi- presence of a perianth and numerous hypogynous stamens would suggest that these bundles may represent traces to a number of abortive stamens. A specific example of an apparently vestigial stamen, adnate to the base of the ovary and receiving such a trace, is illustrated (v st, in FIGURE 131, J, N). If this interpretation should prove to be correct, the flower 1970] BOGLE, HAMAMELIDACEAE 359 of Matudaea with undifferentiated perianth, numerous hypogynous sta- mens, and superior ovary, could represent a relatively primitive type, morphologically, among the apetalous Hamamelidoideae. However, the vestigial vasculature is similar to that of Distylium, and considered in conjunction with the irregular vascular supply to the perianth, and the similarities of inflorescence morphology, suggests the need for further study and for more information from other areas of investigation before the idea of a pseudanthial origin of the flowers can be discarded entirely. DISCUSSION The morphological and anatomical observations recorded above reveal a striking diversity of floral forms and structures which share no common basic pattern. Several parallel lines of specialization appear to be rep- resented, some of which transgress currently recognized generic and tribal limits. A perianth is absent in Distylium, and in those species of Sycopsis re- lated to S. dunnii. The morphology and vascular supply of the inflo- rescence and “floral” bracts in these two groups clearly indicate that they represent reduced stipulate leaves. In Sycopsis sinensis, S. griffithiana, and Parrotia persica an irregular perigon of 4 to 10 tepals is inserted on the rim of a shallow to deep hypanthium.* The tepals of Parrotia are typically more numerous (6 to 10), and more variable in width and length, than those of Sycopsis sinensis (4 to 7; cf. Endress & Anliker 1968, figs. 3, 4), usually appearing as distinct bracts rather than as irregular lobes to form the perigon tube beyond the rim of its longer hypanthium. The 5 to 8 tepals of Fothergilla, although smaller in size, are similar to those of Parrotia and Sycopsis sinensis in their variable width and irregular vascular supply, and may represent a modification from a perigon similar to that of Parrotia. Such reduction may be observed also in the minute, irregular tepals of Parrotiopsis, which are so poorly developed that little can be said about their vascular supply. In the androecia of all seven genera the stamens are variable in number and insertion relative to the members of the perianth, and are strongly exserted. The 18 to 30 hypogynous stamens of Matudaea would appear to mark this genus as rather unspecialized among the apetalous taxa. It is not clear whether the large numbers of perigynous stamens on the hypanthia of Fothergilla (12 to 32) and Parrotiopsis ( 21 to 25) are de- rived from a many-stamened, hypogynous ancestor similar to Matudaea, or whether they have developed secondarily from an ancestral type, per- " i i the term hypanthium is used here only in a de- es Hoagie ry ig poneegt the sled Ga General Considerations, above, for a brief discussion of the morphological nature of the hypanthium in the Hamameli- doideae. 360 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 haps similar to Parrotia (8 to 15). The seven to ten stamens inserted around the narrower mouth of the hypanthium in Sycopsis sinensis probably represent a slight reduction from a larger ancestral number on a more extensive hypanthium rim, such as that seen in the closely related Parrotia persica. Three to seven perigynous stamens have been counted in Sycopsis dunnii, while two to six stamens occur in the subperigynous flowers of Molinadendron, and in the staminate flowers of Distylium. Anthers vary in size and shape among the genera from small and slightly oblong (Fothergila epee to large and linear-oblong (Par- rotia, Sycopsis sinensis). anther connective is barely, if at all, pro- duced as an apiculus in exces and Parrotiopsis, and slightly to moderately produced in the other genera, becoming longest in Matudaea, Parrotia, and in the male flowers of Sycopsis sinensis. Dehiscence is by means of four more or less distinct valves in Fothergilla, Parrotiopsis, Ma- tudaea, and Molinadendron, by slits which tend toward a subvalvate con- dition in Distylium and several species of Sycopsis, and by simple slits in Sycopsis sinensis and Parrotia. Although the flowers and stamens of these apetalous genera are presumably modified for wind pollination, the conspicuous white involucral bracts surrounding the inflorescence of Parrotiopsis, and the dense clusters of long, white, clavate stamen fila- ments in Fothergilla, suggest adaptations toward visual attraction of in- sect pollinators. Unfortunately, observations regarding such character- istics as floral fragrance and pollinators are lacking in the literature. A hypanthium is absent in Matudaea and Distylium, and only weakly developed in Molinadendron, where it is more obvious around the base of the mature fruit. The shallow hypanthium of Parrotia is free of any fusion to the ovary wall, but the hypanthia of Fothergilla and Parrotiopsis are partially adnate to the base of the ovary. The tubular hypanthium of Sycopsis sinensis probably represents an elaboration of a shallower ancestral type, such as that of Parrotia, but the morphological nature of the more or less urceolate hypanthium in Sycopsis dunnii and its related species is not entirely clear. It may represent the fused tissues of re- duced stipulate leaves, and associated axillary axial and floral tissues into a tube around an ovary, and may not be homologous at all with the hy- panthium of Sycopsis sinensis. The general characteristics of the gynoecium in the subfamily Hamame- lidoideae have been described above (see General Considerations). Among the apetalous genera the position of the ovary ranges from superior to partially inferior. The pistil of Distylium is apparently superior morph- ologically, but the presence of a vestigial peripheral bundle system in its base, in conjunction with the absence of a perianth and androecium, suggests the possibility that it may be only secondarily superior. A ves- tigial system of peripheral bundles is also present in the base of the su- perior ovary of Matudaea, but in the presence of a perianth and androe- cium this vasculature may represent vestigial traces to abortive stamens. The superior ovaries of Sycopsis sinensis and Parrotia, although lacking well developed vestigial bundle systems, are associated with other morph- 1970] BOGLE, HAMAMELIDACEAE 361 ological and anatomical peculiarities which suggest a relationship to Distylium, and the possibility that their ovaries may also be secondarily superior. The ovary of Fothergilla varies from more or less hypogynous to one third inferior (cf. Weaver 1969), while in the closely related Parrotiopsis the lower fourth or third of the ovary is adnate with the proximal portion of the hypanthium. The pistil of Molinadendron is sessile and slightly inferior in an expanded receptacle. Considering only the evidence of floral morphology a general trend of floral development among the apetalous Hamamelidoideae could be hy- pothesized to have progressed from a relatively primitive, bisexual flower, containing a perianth consisting of a variable number of undifferentiated tepals, numerous hypogynous stamens, and a superior bi- or tricarpellary ovary, essentially as in Matudaea (which differs in having the tepals fused in a tubular perigon). From this basic plan could be derived forms in which the tepals and stamens have become fused basally into a shallow, hypogynous hypanthium (Parrotia, Fothergilla), possibly accompanied by reduction in stamen number (Molinadendron, Parrotia, Sycopsis sinen- sis), and partial adnation of the hypanthium to the ovary (Fothergilla, Parrotiopsis). Elaboration of the hypogynous hypanthium into an ur- ceolate tube, possibly accompanied by reduction and loss of the tepal lobes morphologically (Sycopsis dunnii and related species), and sup- pression of the development of the gynoecium in lateral flowers of the inflorescence axes (Sycopsis dunnii), or in some whole inflorescences (Sycopsis sinensis), would lead to more advanced andromonoecious floral types (Sycopsis). Finally, suppression of the hypanthium and its asso- ciated perianth lobes (if present) would lead to terminal naked pistillate flowers and lateral naked staminate flowers (Distylium). The vestigial vasculature in the base of the pistil of Distyliwm would then be attribu- table to the lost hypanthium, rather than to lost stamens as hypothesized for Matudaea. This sequence appears reasonable with regard to the pro- gressive steps of modification necessary to derive the various floral forms involved. It differs from that proposed by Tong (1930) and Harms (1930) in that they derived the polygamous, apetalous genera of the tribes Fothergilleae and Distylieae from a petaliferous ancestral type through the transformation of petals into stamens, rather than from an apetalous ancestral type such as Matudaea, which was unknown at that time In such a series the flowers would represent euanthia, and the morph- ological nature of the perigon, lacking only in Distylium and those species of Sycopsis related to S. dunnii, and of the hypanthium, would be homo- logous. If correct, the apetalous genera of Hamamelidoideae may rep- resent the remnants of a line of floral specialization quite apart from that which gave rise to the petaliferous genera, for Matudaea has no close morphological counterpart among the petal bearing members of the subfamily. Among the latter group, on the basis of accumulated evidence of wood anatomy, pollen morphology, floral morphology , and floral vascular anatomy (personal observations, in manuscript), the most 362 JOURNAL OF THE ARNOLD ARBORETUM [vot. 51 primitive genera appear to be Maingaya Oliv. and Dicoryphe Thou. Their pentamerous (or tetramerous in some species of Dicoryphe) flowers con- tain alternating cycles of sepals, petals, 5(-—4) stamens, 5(—4) large staminodia which approach the stamens in shape, and an inner cycle of 10(—8) scales or lobes, all inserted perigynously on the rim of a tubular hypanthium above a half-inferior or essentially inferior ovary. However, in view of the unusual features of floral morphology and vascular anatomy in Distylium, Sycopsis, and Parrotia, it may be worth considering another, more speculative, interpretation of the course of floral development among at least some of the apetalous genera, leading from a hypothetical unisexual, monoecious, ancestral type, to bisexual pseudanthia through the condensation of inflorescences and the aggrega- tion of staminate and pistillate flowers within a cycle of bracts. Among extant genera the species of Distylium could represent an early stage in the development of such pseudanthia. Its naked unisexual flowers have probably been derived from those of a bisexual ancestor through the sup- pression of the perianth and gynoecium in the case of the lateral staminate flowers, or of the perianth and androecium in the terminal pistillate flow- ers on each inflorescence axis. Such reduction must be assumed regard- densation of inflorescence axes and the aggregation of bracts and staminate flowers around a terminal pistillate flower have already produced a condition in which the external identity, and even the anatomical identity, of the individual flower is lost, and the inflorescence merges into a false flower. From a common ancestor with unisexual flowers more or less similar to those of the present day distyliums, the apparently bisexual flowers of Sycopsis dunnii and its related species may have developed through a more pronounced reduction of inflorescences axes, resulting in the aggre- gation and fusion of lateral trimerous primary bract groups with their associated axillary axial tissues and staminate flowers into a tubular false hypanthium around a naked terminal pistil. From a similar hypothetical ancestor, the capitate inflorescences and bisexual flowers of Parrotia could have evolved through the condensation of entire lateral inflorescence axes, resulting in the aggregation and fusion of a variable number of primary bract groups and highly modified lateral staminate flowers in a shallow hypanthium around the persistent naked terminal pistil. From a condition such as that seen in Parrotia it is only a short step, in one direction, to Sycopsis sinensis through a lengthening of the hypan- morphological origins of the hypanthium of S. sinensis would then = pear to differ somewhat from those of the similar structure in S. dunn and the two species would represent an interesting case of Pe lonenetie 1970] BOGLE, HAMAMELIDACEAE 363 convergence, resulting in urceolate hypanthia that are ruptured by the developing fruit. In another direction could be derived Fothergilla and Parrotiopsis. In these genera the increased number of stamens may rep- resent either the retention of an ancestral condition or a secondary de- velopment. In the context of a pseudanthial interpretation, the flowers of Matudaea might represent a less advanced New World counterpart of Parrotia, its aggregated connate bracts not yet having become fused in a hypanthium with the stamens. The vestigial vasculature in the base of the ovary, like that in Distylium, might then be attributable to a phylogenetically lost floral envelope which once surrounded the ovary. In Molinadendron, on the other hand, the irregular relationships and variable numbers of the stamens and one-trace tepals are suggestive of an aggregation of bracts and stamens similar to those seen in some highly reduced inflo- rescences of Distylium buxifolium. LITERATURE CITED Atry-SHAw, H. K. Distylium racemosum. Curtis’s Bot. Mag. 160: pl. 9501. 1 ANDERSON, E., & K. Sax. Chromosome numbers in the Hamamelidaceae and their phylogenetic significance. Jour. Arnold Arb. 16: 210-215. 1935. BaILLon, H. Nouvelles notes sur les Hamamélidées. Adansonia 10: 120-137. 1871 ———. Saxifragacées, Hist. Pl. 3: 325-464. 1871. [Hamamelis series, 389- 397; 414-415; Hamamelideae, 456-461. Liguidambar series, 397-400; 415; Liquidambareae, 461, 462.] BaLcooy, M. M. J. van. Distylium. In: Pacific Plant Areas 2: 132, 133. Blumea Suppl. vol. 5. [Distribution map. BENTHAM, G., . D. Hooxer. Hamamelideae. Gen. Pl. 1: 664-669. 1865. Bessey, C. E. The phylogenetic taxonomy of flowering plants. Ann. Missouri Bot. Gard. 2: 109~164. 1915. [Hamamelidaceae, 151.] Bocte, A. L. Floral vascular anatomy and the nature of the hamamelidaceous flower. Ph.D. thesis, University of Minnesota. 1967 [1968]. . Major features of morphology and vascular anatomy of the hamameli- daceous gynoecium. Am. Jour. Bot. 55: 715. 1968. [Abstr. ] Flower and pseudanthium in Déstylium. XI International Botanical Congress Abstracts, 17. 1969. CANDOLLE, pe. Hamamelideae. Prodr. 4: 267-270. 1830 [Tribe Hama- meleae, 268, 269; Tribe Fothergilleae, 269, 270. Carr, S. G. M., & D. J. Carr. The functional significance of syncarpy. Phy- tomorphology 11: 249-256. 1961. ie, Goatees K Ai "The pollen morphology of Liquidambar L. and Altingia Nor. Bot. Zhur. 44: 1375-1380. pls. 1-5. 1959. [In Russian with English sum- mary. gee pollen morphology of the families Hamamelidaceae and Altin- giaceae. Acta Inst. Bot. Acad. Sci. URSS. 1 Fl. cSyst: PL: Vase... 13: 173-232. pls. 1-21. 1964. eos aa Conprr, I. J. The structure and development of flowers in Ficus carica L. Hilgardia 6: 443-481. 1932 364 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 CROIZAT, fe Principia botanica, la. xiii + 632 pp. Caracas. 1960. (Chapt. 4b, 257-300. ] : eae on high systematics, phylogeny, and floral morphogeny, with a note on the origin of Angiospermae. Candollea 19: 17-96. 1964. Cronoutst, A. The evolution i classification of flowering plants. x + 396 pp. Eames, A. J. Morphology of the angiosperms. xiii + 518 pp. New York. 961 EICHLER, rt W. Bliithendiagramme. 2. xx + 575 pp. Leipzig. 1878. [Hama- melideae, 436-438 Enpress, P. Systematische Studie iiber die verwandtschaftlichen Beziehungen zwischen den Hamamelidaceen und Betulaceen. Bot. Jahrb. 87: 431-525. pls. La 1967. —— & J. ANLIKER. X Sycoparrotia semidecidua hybr. nov. (= Par persica C. A. Mey. X Sycopsis sinensis Oliv.). Schweiz. Heir. aL 16-18: 6-28. 1968. Ernst, W. R. The genera of Hamamelidaceae and Platanaceae in the south- eastern United States. Jour. Arnold Arb. 44: 193-210. 1963. FLINT, F. F. Megasporogenesis and megagametogenesis in Fothergilla Sabet Murr. and Fothergilla major Lodd. Trans. Am. Microscop. Soc. 76: $Ll GENTRY, H. S, Additions to the flora of Sinaloa and Nuevo Leon. Brittonia 6: 309-331. 1948. GRIFFITH, W. Description of two genera of the family of Hamamelideae, two species of Podostemon and one species of Kaulfussia, Asiat. Res. 19: 94- 114. pls. 13-19. Hatiier, H. Ueber den Unitas. die Gliederung und die Verwandschaft der Familie der Hamamelidaceen. Beih. Bot. Centralbl. 14: 247-260. ———. L’origine et le systéme phylétique des Angiospermes exposés a Paide de leur arbre généalogique. Arch. Néerl. Sci. Nat., Ser. 3B. 1: 146-234. pls. 1-6. 1912. Harms, cae Hamamelidaceae. Jn: A. ENGLER & K. PRANTL, Nat. Pflanzenfam. ed. 2. 18a: 330-343. 1930. eber eine neue Art der oe Distylium Sieb. et Zucc. aus Mit- tel-Amerika, Notizbl. Bot. Gart. Berlin 11: 714-718. 1933. Hemstey, W. B. Distylium chinense "Teal Hooker’s Te. i ee, 2855. 1907; Sycopsis tutcheri Hemsl., S. sinensis Oliv. Ibid. pl. 2834; Sycopsis dunnii Hemsl. Ibid. pl. 2836 TS. laurifolia Hemsl. and S. philippinense Hemsl. discussed but not illustrated in pl. 2836.] Henpverson, I. F., & W. D. HENDERSON. A dictionary of scientific terms. ed. 7. xiv + 595 pp. New York ; Hooker, J. D. Parrotia jacquemontiana. Curtis’s Bot. Mag. 122: pl. 7501. Horne, A. L. A contribution to the study of the evolution of the flower, with special reference to the Hamamelidaceae, Caprifoliaceae, and Cornaceae. Trans. Linn. Soc. Lond. II. Bot. 8: 239-309. pls. 28-30. HUTCHINSON, re The families of flowering plants. I. Dicotyledons. xiv + 328 pp. London. 1926. [Hamamelidaceae, 213, 214.]; ed. 2. xv + 510 pp. Oxford. 1959. [Hamamelidaceae, 180, 181. camo B. D. A glossary of botanic terms. ed. 4. xii + 481 pp. London. 1928 1970 | BOGLE, HAMAMELIDACEAE 365 Jay, M. Distribution des penn chez les Hamamélidacées et familles af- fines. Taxon 17: 136-1 1968. JOHANSEN, D. A. Plant Berar xi + 523 pp. New York. 194 KamMEyeR, H. F. Die Schénen Zaubernusse. 92 pp. Wittenberg. oe. Kau, U. Endosperm in Parrotiopsis jacquemontiana. Phytomorphology 19: 197-199. 1969. LuNpDELL, C. L. Three noteworthy new ore from southern Mexico. Lloydia 3: 209-212. 1940. [Matudaea, 209-21 . Plantae Mayanae—IV. New w species, nomenclatural changes, and new records for trees and shrubs of Mexico and Central America. Wrightia 3: 1-20. 1961. |Matudaea hirsuta sp. nov., 1-3. oo F. Hamamelidaceae. 7m: A. ENGLER & K. Prantt, Nat. Pflanzen- III. 2a: 115-130. a _Nachtrige II zum II-IV. Teil: 29. 1900; bid. (Erganzungshefte II): OLIVER, 7 On Sycopsis. Trans. = ee Lond, 23: 83-89. pl. 8. 1860. Sycopsis sinensis Oliv. Hooker’s Ic. Pl. 20: pl. 1931. 1890. P1zzoLonco, P. Ricerche cario- lepiaie ong su alcune Hamamelidales. Ann. Bot. Roma 26: 1-18 ae : 2. 1958. [In Italian, English summary. Par- rotia ers pl. 1, figs. REHDER, A. New s ecies, Cae and combinations from the herbarium and the collections of the Arnold Arboretum, Jour. Arnold Arb. 1: 254— 263. 1920. price of cultivated trees and shrubs hardy in North oe ed. xx + 996 pp. New York. 1954. [Hamamelidaceae, 311-3 ices: va Ueber die anatomischen Verhaltnisse der oe mit Riicksicht auf ihre systematische Gruppierung. Bot. Jahrb. 11: 347-395. pl. 8. 1889. SAUNDERS, E. R. Floral morphology, 2. xiv + 133-609 + vii pp. New York. 1940. [Hamamelidaceae, 162-164; Corylopsis, Hamamelis, Parrotia, Par- rotiop SCHULZE- cael G. K. Hamamelidaceae. Jn: A. ENGLER’s Syllabus der Pflan- zenfamilien. ed. 12. 2: 196-198. 1964 SHOEMAKER, D. N. On the development of Hamamelis virginiana. Bot 39: 248-266. pls. 6, 7. 1905. [Observations on organogeny of Fothergilla ardeni SoLsKaDex, Zur Morphologie und Systematik der Gattung Cercidiphyllum Si t Zucc., mit wee ay Sar iy der Gattung Eucommia Oliv. Ber. sch. Bot. Ges. 17: 38 pl. 28. 1899. SoANDLE, P; ie.” Celie ne of Guatemala. Fieldiana, Bot. 4 (4): 426-430. 1946. [Distylium = Molinadendron, Matudaea, Liqui- cae | Sucrura, T. A list of chromosome numbers in angiospermous plants. II. Proc. Imp. Acad. Japan 12: 144-146. TAKHTAJAN, A. Flowering plants — origin and dispersal. x + 310 pp. Wash- 1969. Tanc, Y ‘Sy stematic anatomy of the woods of ee Hamamelidaceae. Bull. Fan Memorial Inst. Biol. 1: 8-63. tables 1-3. Trttson, A. H., & R. BAmrorp. The floral Le - the Aurantioideae. Am. Jour. Bot. 25: 780-793. 1938. ; Trero, O. Comparative anatomy of the Moraceae and their presumed allies. Bot. Gaz. 100: 1-99. 1938. [Hamamelidaceae, 2, 3, ; Tone, K. Studien iiber die Familie der Hamamelidaceae, mit besonderer Beriick- 366 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 sichtigung der Systematik und eo von Corylopsis. ee Dept. Biol. Coll. Sci. Sun Yatsen Univ. 2: 1-72, + 1-14 [in Chinese]. 30. — W. Hamamelidaceae. Jn: C. G. G. J. vAN STEENIS, Flora Malesiana I. rs 363-379. 1957. Waker, E. H. A revision of Distylium and Sycopsis (Hamamelidaceae). Jour. Arnold Arb. 25: 319-341. 1944. Weaver, R. E. Jr. Studies in the North American genus Fothergilla (Hama- melidaceae). Jour. Arnold Arb. 50: 599-619. 1969. WETTSsTEIN, R. Handbuch der Systematischen Botanik. ed. 4. x + 1149 pp. 1935. [Hamamelidaceae, 684-686. | ARNOLD ARBORETUM HARVARD UNIVERSITY 1970] KAZMI, BORAGINACEAE 367 A REVISION OF THE BORAGINACEAE OF WEST PAKISTAN AND KASHMIR * S. M. A. Kazi 8. Pseudomertensia Riedl in Rechinger, Fl. Iranica 48: 58. 1967. Oreocharis (Decne.) Lindl. Veg. Kingd. 656. 1848, non Benth. 1876, pro parte. Lithospermum L. ais . Oreocharis Decne. in Jacquem, Voy. Inde Bot. 122. , pro par a Schrad. ie Oreocharis (Decne.) DC. Prodr. 10: 123. 1846, pro Hee: auct. fl. asiatica pro parte — non Roth, Catalecta 1: 34. 1797. Type species: Lithospermum elongatum Decne. Perennial, strigose or softly hairy herbs. Leaves usually alternate. Racemes terminal, scorpioid, usually simple, dense or lax, ebracteate. Flowers blue to purplish- -blue, pedicellate. Sepals narrow, sometimes en- larged in fruit. Corolla campanulate, cylindrical-campanulate, or hypocra- teriform; tube as long as, or much longer than the calyx; throat with or without appendages; lobes 5, imbricate in bud, short, spreading or erect. Stamens 5, included or exserted from the corolla tube; filaments very short to very long; anthers oblong, obtuse, or rarely acute. Ovary 4-lobed, style filiform, stigma capitate, small. Nutlets 4, erect, ovoid-oblong, dor- sally slightly convex, ventrally carinate, slightly angular, margins often acute; areola distinct, sub-basal, oblique, affixed to the short pyramidal gynobase. Species 11 to 12, distributed in Western Himalayas from Afghanistan to Nepal KEY TO THE SPECIES a. Anthers exserted from the corolla tube, filaments 2-5 mm. long. b. Corolla more or less niga the calyx; corolla lobes up to 1.5 mm. long, feucal anonnioes BOM oes oe aes sin 1, P. parviflora. b. Corolla aie oe calyx; corolla lobes 2.5—3 mm. lone eee appen prese P. trollii. a. Anthers sscladcd or ee exserted from the corolla tube; steel tee "hat 2 mm. long. c. Faucal appendages absent. ; Corolla not constricted at the throat; internally with minute pubes- cence instead of faucal appendages; filaments 2 mm re ener es hate UMC Se in Gey ee er Ne ea oe 3. efornicata. d. Corolla constricted at the throat, internally glabrous; lament hardly 4. P. echioides. ong. c. Faucal appendages present, poorly to well developed. * Continued from volume 51, p. 184. 368 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 or ere me A thd Banks 5. P. elongata. e. Corolla not daanpamulate: distinctly rs lindrical or infundibuliform. f. Faucal appendages longer than g. Faucal appendages 2-3 mm. (eae 1s mm. broad at the base, triangular, ciliate at the margins. ......... . P. drummondii. g. Faucal appendages 0.8-1.2 mm. long, 0.5 mm. broad at the base, lingulate, not ciliate at the margins. .. 7. P. chitralensis. f. Faucal appendages broader than long. h. Filaments usually longer, rarely equalling the anthers in length. SERS gsc icey MIR Aiss a) A ee eS a 8. P. anjumiae. h. Filaments always shorter than the length of anthers. i. Leaves, including petioles, 40-160 mm. long, 10-45 mm. broad; corolla tube twice as long as the oe anthers up Wer eek et eee nemorosa. i. Leaves, ace petioles, 15-80 mm. long, z 101 mm. broad; corolla tube three to five times as long as the calyx; an- oy Ys See steed | 10. P. moltkioides. 1. P. parviflora (Decne.) Riedl in Rechinger, Fl. Iranica 48: 60. 1967. Ls Lori aie parviflorum Decne. in Jacquem. Voy. Inde Bot. 1: 126. ¢. 130. 1844. Moltkia parviflora (Decne.) C. B. Clarke in Hook. f. Fl. Brit. India 4: 171. 1883. Mertensia exserta I, M. Johnston, Jour. Arnold Arb. 37: 305. 1956; R. R. Stewart, Biologia 13: 93. 1967. Type: In summis pascuis 4 Hlahabad ad Hyderabad trans jugum Pir- penjal, 2681 m., Jacquemont s.n. (P). Icon.: Jacquemont, l. c. ¢. 130. 1844. Perennial with stout woody underground rhizomes; emerging shoots many, suberect or sometimes trailing, 6-15(—25) cm. long, clothed with short appressed trichomes. Basal leaves short petiolate, linear-oblong, entire, obtuse or subacute, 40-50 mm. long, 4-6 mm. broad, covered on both surfaces with short, appressed trichomes; cauline leaves mostly sessile, shorter and narrower, acutish. Inflorescence usually forked, some- times simple, short and scorpioid when young, later elongated, curved, 2-6 cm. long. Calyx divided to the base, lobes narrow oblong, 2.5-3 mm. long, densely hairy on the lower parts and margins, obtusish, in fruits slightly enlarged; pedicels 1-4 mm. long. Corolla 2.5—3 mm. long, more or less equalling the calyx, cylindrical, glabrous, tube 2—2.5 mm. long, lobes 0.5—1(—1.5) mm. long, 0.5—0.8 mm. broad at base, erect. Stamens much exserted from the corolla tube; anthers obtusish, elliptic, 0.5—0.6 mm. long, versatile; filaments up to 3 mm. long, inserted about 2.5 mm. above the corolla base. Style 6 mm. long, stigma set tae Nutlets ovoid-oblong, acute, smooth, shining, ca. 2 mm. DISTRIBUTION: West Pakistan, Kashmir. West Pakistan: CHITRAL STATE: Ziarat, Lowari Pass, 2600 m., Stainton 2337 (pm). Hazara Dist.: Pipe line, Changlagali, 2500-3000 m., Dickason 83 1970] KAZMI, BORAGINACEAE 369 (MICH); between Dungagali and Changlagali, 2500-3000 m., Kazmi 746a (PES); Nathiagali, 2700 m., Kazmi 84b (PES), KurRAM AGENCY: Kurram Valley, Afan- di 404 (PEs); Ziran, Kazmi 207a (pes): Shalozan, Kazmi s.n. (pes), Afandi sm, (PES); Kaiwas & Shendtoi, 2700-3000 m., Aitchison 397 (GH). Rawat- PINDI Disr.: Murree Hills, 2300 m., R. R. Stewart 12559 (GH), Asghar Ali, s.m. (MIcH); Patriata Forest, Kazmi ‘4c (PES); Murree, Kashmir Point, Kazmi 2480 (PES); Upper Topa, Kazmi 2487 (prs). Kashmir: Poonch, Trarkhel, 2000 m., R. R. Stewart 12084 (cH); Gulmarg, Aitchison 74 (x), Keshawanond 1162 (cH), Trotter 110 (pm); Basin “ Chenab, Thomson (4. 5. 1848) s.n. (K); near Avantipura, Thomson (4. 5. 1848) s.n, (kK); Lidder Valley, Kinimola Nulla, Inayat 25705 (Kk). Miscellaneous: 5000— 7000 ped., Herb. Ind. Orient. Hook. f. & Thoms., Thomson s.n. (cH). 2. P. trollii (Melch.) Stewart & Kazmi, comb. nov. Mertensia coventryana S. Clay, The Present Rock Garden 20: 379. ¢. 27a. 1937, nomen nudum Moltkia trollii Melch. Notizbl. Bot. Gart. Berlin 15(1): 115. 1940 Mertensia trollit (Melch.) I. M. Johnston, Jour. Arnold Arb. 37: 303. 1956; Riedl in Kgie & Rechinger, Biol. Skr. 13(4): 227. 1963 Type: Kashmir: Tragbal Pass, in der Nadelwaldregionen mit Pinus, Picea und Abies, 3000 m., Bliiten hellblau, C. Troll 7028 (s). Icon.: S, Clay, l. c. ¢. 27a. 1937, under Mertensia coventryana. Perennial with underground branched rhizomes to 20 cm, long; emerg- ing shoots usually curved, sometimes erect, to 10 cm. tall, clothed with short, thick, white, appressed trichomes. Basal leaves including petioles 4—6 cm. long, 4-18 mm. broad, lamina elliptic or elliptic-oblong, entire, apices obtuse, glandular, nerves prominent below, at the base gradually attenuated into long petioles, more or less ciliate at margins, broad at the base; both surfaces covered with short, white, appressed trichomes, cauline leaves oblong to oblong-lanceolate, shorter, narrower, sessile to subsessile. Inflorescence terminal, usually solitary, contracted, scorpioid, 1-3 cm. long. Calyx shorter or equal to the corolla tube, divided to the base, lobes linear-oblong, acute, densely hairy on the margins, few scattered trichomes externally, marginal trichomes 0.5—0.8 mm. long and antrorsely spreading; pedicels 1-3 mm. long. Corolla sky-blue to dark purplish-blue, 7-8 mm. long, glabrous, tube cylindrical ca. 5 mm. long, lobes oblong, 2.5— 3 mm. long, ca. 1.8 mm. broad at the base, subpatent; faucal appendages 0.5 mm. long and broad. Stamens exserted from the corolla tube to exceed- ing the corolla lobes; anthers versatile, elliptic, obtuse, 1-1.5 mm. long; filaments 2.5—5 mm. long, slightly narrowed at the apices, inserted at the level of the faucal appendages, about 4 mm. above the corolla base; style filiform, equalling or shorter or little exceeding the stamens; stigma minute, subcapitate. This species is closely related to Pseudomertensia parviflora in having stamens conspicuously exserted from the corolla tube, but differs from it in having a much longer corolla with well developed lobes and five in- 370 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 vaginated hemispheric appendages in the throat. The corolla lobes in P. trollii are oblong or elliptic, 2.5-3 mm. long and clearly exceeding the calyx lobes, while those of P. parviflora are more or less triangular and usually 1-1.5 mm. long. The length of calyx, corolla, filaments, and anthers is very variable in Pseudomertensia trollii and the three following varieties may easily be distinguished. KEY TO THE VARIETIES a. Calyx about 2/3 the length of corolla tube; corolla 9-11(- Z mm. long. SI oad eae eC aM cr Re eager 2a ene san eae rere Ons var. pies a. Calyx usually equal to the corolla tube; corolla up to 8 mm. long. ai rs 1 mm. long; filaments 4-5 mm. long; faucal appendages well de- 2a. BE irik, a eaten Oba ie RS Vaud bay id oe koe Ss . troll, b. he ee 1-1.5 mm. long; filaments ca. 2 mm. long; faucal Fees Be EE hee io oe ie hs BHR DR ae ® eee 20 2b. var. harrissi. 2a. Var. trollii Basal leaves narrower and with short petioles compared to var. edel- bergii; calyx more or less equal to the corolla tube; corolla up to 8 mm. long, lobes acutish; anthers 1 mm. long; filaments 4-5 mm. long. DISTRIBUTION OF VAR.: West Pakistan, Kashmir. West Pakistan: G1Lcir AGENCY: On Indus river near Rondu, E-W. of Skardu, © about 40 miles, Thomson (18.3.1848) s.. (kK); Gurais, Mrs. Earl in R. R Stewart 1258 (GH). Hazara Dist.: Kaghan Valley, Bhimbal, 2300 m., Duthie 19486/a (kK); Batakundi, 2700 m., Champion (13.5.1927) sn. (GH); Kazmi 5.n, (PES). Kashmir: Tragbal Pass, 3000 m., Troll 7028 (B-type); Kajnag range, Duthie 11073 (£); Banihal Pass, 3000 m., Coventry 1448 (pm); Pahlgam, 3000 m., Miss Ward 116 (BM). 2b. Var. harrissii Kazmi, var. nov. Type: West Pakistan: Dir, Harriss 16397 (Gu-holotype). Folia basalia ad var. trollium similia; tubum calycis corollae plus min- usve aequans; corolla usque ad 8 mm. longa, lobi aliquot obtusi; antherae 1—1.5 mm. longae; filamenta 2.5 mm. longa. DISTRIBUTION OF VAR.: West Pakistan. West Pakistan: Dir State: Dir, Harriss 16397 (GH-type). Hazara DISsT.: Naran, near the Ganji Pahari top, 3000 m., Kazmi 2218b (pEs); Saiful Maluk Sar, D. McVean (June 1960) s.m. (E); between Dadar and Kaghan (probably at Shahid Pani ?), Jnayat 22006 (x). 2c. Var. edelbergii (Rech. f. & Riedl) Kazmi, comb. nov. nhac composed Rech. f. & Ried] in Kgie & Rechinger, Biol. Skr. 13(4): 170.0177. 1963. 1970] KAZMI, BORAGINACEAE S71 Pseudomertensia edelbergii (Rech. f. & Riedl) Riedl in Rechinger, Fl. Iran- ica 48: 60. 1967. Type: Afghanistan: Nuristan: Pashki, 2300 m., Edelberg 630 (w-holo- type, C-isotype). Icon.: Riedl 1. c. ¢. 176, 177. 1963. Basal leaves broader with longer petioles as compared to the other two varieties; calyx about 2/3 the length of the corolla tube; corolla 9-11 (—12) mm. long; anthers ca. 1 mm. long; filament 2.5 mm. long DISTRIBUTION OF VAR.: West Pakistan, Afghanistan, West Pakistan; Swat State: Kalam, 3000 m., Kazmi 2468 (PES); Indus Kohistan, beyond Ushu, 3200 m., Kazmi s.n. (pes); Kalam mountain, S-E. of village, upper section of hill above ‘alp’ rocks, near ridge, flowers deep blue fading to deep pink, 3000-3200 m., Lamond 1841 (£); Lamond in Rechinger 30854 (w), Rechinger and Riedl (/. c. 1963) in their original description of Pseu- domertensia edelbergii gave the length of the corolla as 9-11(-12) mm.: later Riedl (/. c. 1967) cited the length of the corolla as 7-8 mm. which is the length of the corollas in varieties trollii and harrissii. In all the specimens of var. edelbergii cited above, the length of the corolla is def- initely more than 8 mm. Also, the original description of P. edelbergii does not indicate the length of the filaments, which was measured by me to be more or less 2.5 mm., much shorter than the length of the filaments of P. trollii, but almost the same as in var. harrissii, from which var. edel- bergiu differs in having a shorter calyx in comparison with the length of the corolla tube, as well as shorter anthers and larger leaves. Both var. edelbergiit and var. harrissii differ from the typical variety in the length of the filaments which varies between 4 and 5 mm. Because the length of the filaments in the P. trollii complex is not very constant, and since var. harrissii represents an intermediate form between P. edelbergii and P. trollii, I consider all three taxa as only varieties of P. trollii. 3. P. efornicata (Rech. f. & Riedl) Riedl in Rechinger, Fl. Iranica 48: 60. 1967 Mertensia efornicata Rech. f. & Riedl in Kgie & Rechinger, Biol. Skr. 13(4): 231, 1903: Type: Chitral, Drosh, 11500 ft., Bowes Lyon 186 (BM). Icon.: Riedl, 1. c. 232. fig. 178. 1963, under Mertensia efornicata. Perennial; rhizomes stout, long, more or less 1 cm. thick, branched, branches short, underground; emerging shoots many, 18-25 cm. long, erect, simple, or in the upper parts, bifurcate, covered loosely with pa- tent to subappressed trichomes. Basal leaves lanceolate to oblong lan- ceolate, apices apiculate, attenuated at the base into petioles, including the petioles 5-12 cm. long, 7-18 mm. broad, on the lower surface thickly, 372 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 on the upper surface loosely covered with short trichomes; upper leaves few, attenuated towards the base, sessile, 2-4 cm. long, 3-8 mm. broad. Inflorescence solitary, short when young, later elongated up to 10 cm., bearing many distant pedicellate flowers; pedicels 3-5 mm. long. Calyx divided to the base, lobes linear, acute, + 5 mm. long, 0.5—0.7 mm. broad, loosely covered with subappressed short trichomes. Corolla 10-11 mm. long, cylindrical-campanulate, glabrous externally, minutely hairy in- ternally in the throat in place of faucal appendages, throat not con- stricted, lobes and tube not distinct, lobes ovate to roundish, 2—2.5 mm. broad. Anthers + 1.5 mm. long, sagittate, acute, slightly exceeding the sinuses between the corolla lobes; filaments filiform, 2 mm. long, at- tached 5-6 mm. above the corolla base. Style filiform, equalling or slightly exceeding the corolla; stigma capitate. DISTRIBUTION: West Pakistan. West Pakistan: CHITRAL STATE: Drosh, Painogh, 3500 m. (13. 7. 1958), Bowes Lyon 186 (3M) The absence of faucal appendages brings P. efornicata very close to P. echioides but the presence of pubescence in place of them distin- guishes this species from the latter. In P. echioides the anthers are sub- sessile or the filaments never exceed the length of 0.5 mm., whereas in P. efornicata filaments are about 2 mm. and always longer than the anthers. 4. P. echioides (Benth.) Riedl in Rechinger, Fl. Iranica 48: 62. 1967. Lithospermum Lesion es? in Royle, Illustr. Bot. Himal. Mount. 1: 305. r. 10; 63. Lithospermum pinay Pinay Rae in Jacquem. Voy. Inde a 120. 1844. Eritrichium secundiflorum (Decne.) A. DC. Prodr. 10: 124. 1846. Mertensia echioides (Benth.) Benth. & Hook. f. Gen. Pl. 2: Ae) ta70; . B. Clarke in Hook. f. Fl. Brit. India 4: 170. 1883; Riedl in Kgie & Rechinger, Biol. Skr. 13(4): 229. 1963. Mertensia secundiflora (Decne.) Brand, Pflanzenr. IV. 252(Heft 97): 200. 1931. Mertensia strigosa Melch. Notizbl. Bot. Gart. Berlin 15: 113. 1940, syn. nov. Type: Soongnum in Kunawar-Kherang Pass, R. /nglis s.n. (kK). Icon.: Riedl, 1. c. fig. 175. 1963, under Mertensia echioides. Perennial with long underground rhizome. Emerging shoots usually erect, sometimes decumbent or curved, 5—20 cm. long, clothed with short usually spreading, rarely appressed, thin trichomes. Basal leaves includ- ing long petioles (25—)40-90(—150) mm. long, (5—)10-15(-—25) mm. broad; lamina ovate, elliptic-lanceolate or lanceolate, usually acute, some- ips leaves few, shorter and narrower, sessile. Inflorescence short and scor- 1970] KAZMI, BORAGINACEAE 373 pioid when young, later elongated up to 13 cm., lax, bearing 10-20 sub- sessile pedicellate flowers to 10 mm. apart; pedicels up to 3 mm. long. Calyx divided to the base, lobes linear, + obtuse, erect, 4-5 mm. long, 0.5 mm. broad at base. Corolla blue to deep purple-blue, subcylindrical to funnel-shaped, glabrous, bearing few trichomes at the sinuses between the lobes; corolla tube 4.5-6 mm. long, slightly dilated above: lobes usually 34.5 mm., sometimes even longer, 1.5 mm. broad: faucal ap- pendages absent. Anthers oblong, acute or rarely emarginate at the apices, usually emarginate or roundish at base, 1.5-1.9 mm. long exserted from the corolla tube, sometimes anther tips reaching half the length of the corolla lobes; filaments not more than 0.5 mm. long, thick, attached 4.5—5.5 mm. above the corolla base. Style 10-12(-15) mm. long, fili- form; stigma capitate. Nutlets ovoid-trigonal, acute, pubescent on ridges, 1.8—2.5 mm. long, 1—1.5 mm. broad. DIsTRIBUTION: West Pakistan, Kashmir, and India. Most authors have placed Pseudomertensia echioides with the species having faucal appendages and have distinguished it from those species because the appendages are less developed and the anthers are ex- serted from the corolla tube. With these characters it becomes some- times quite difficult to distinguish P. echioides from the allied species, which have faucal appendages at various stages of development and anthers usually exserted from the corolla tube. In the specimens, includ- ing the type, which I have examined, I found that P. echioides has no faucal appendages, and if very rarely these are present, they are in in- 0.5 mm. in length; filaments are thick and dilated at the base. These characters are well correlated with the long (1.5-1.9 mm.) anthers, usually acute at their apices, with usually slightly produced connectives and with the presence of trichomes at the sinuses between the corolla lobes. The presence of trichomes at the sinuses is not found in any other species of our area. The size and form of the leaves of Pseudomertensia echioides are very variable and two varieties may be distinguished easily. 4a. Var. echioides. Shoots few, not profusely leafy, leaves thick, usually oblong-lanceolate, up to 1.5 cm. broad, densely covered with trichomes; inflorescence more compact and less hairy in comparison to var. /ahulensis. DISTRIBUTION OF VAR.: West Pakistan, Kashmir. West Pakistan: Gitcir AcENcy: Astor valley, Kalapani, Kamrinala, Jnayat 25708 (kK); Shankargarh above Astor village, Giles 66 (kK); Astor to Degru, Conway 334 (Kk); Upper Shingu valley, Deosai region, 3000 m., R. R. Stewart 22231 (cH); Baltistan, Satpara Nullah, 3000 m., Webster & Nasir 6356 (cH), , R. R. Stewart 20244 (GH, MICH, US), 3000 m., Siddiqui, Nasir & Zaffar Ali 4231 (BM, RAW), Duthie (12. 7. 1892) sm. (BM); Nathar waterfall, 374 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 R. R. Stewart 26343 (raw); near Skardu, 2300-2500 m., Schlagintweit 794 (pm); Marpu Nullah, above Dras, 4000 m., Duthie (3. 7. 1892) sn. (BM); Burja La, 4000 m., C. B. Clarke 29930 (BM, x). Kashmir: Shakkar, Rangdum, 4000 m., Koelz 5969 (GH, MICH, US); Barnaj Nullah, near Sapphire Mines, Kishtwar, 3500 m., Ludlow & sind 9181 (GH); Suru valley, N. of Srinagar, 4000 m., Osmaston 212 (kK); LapAKH: Hanu Yegma via Handamir up to Chorbat La Pass (Pass between the Indus valley and the sgl valley), Schlagintweit 6511 (us). Miscellaneous: Hab. Himal. Bor. Occ., 5000-6000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (BM, GH, je Falconer s.n. (GH). Mertensia strigosa Melch. agrees in all its details with Pseudomertensia echioides var. echioides and represents only a more densely pubescent form of the variety. 4b. Var. lahulensis (Brand) Kazmi, comb. nov. Lindelophia lahulensis Brand, Repert. Sp. Nov. 19: 70. 1923. Type: Lahul: Kyelang, an Felsen des Nyimephed, 4200 bis 4400 m. U.M., im Juli, 1880, Heyde s.n. (z1— Herb. Bernhard Lorenz). Shoots many, profusely leafy, leaves thin, usually ovate to ovate-lan- ceolate, more than 1.5 cm. broad, loosely covered with trichomes; in- florescence lax; fruits more hairy than in var. echioides. DISTRIBUTION OF VAR.: INDIA (Lahul): Kyelang, Koelz 470, 5183, 8408 (GH, MICH, US). 5. P. elongata (Decne.) Riedl in Rechinger, Fl. Iranica 48: 61. 1967. Lithospermum elongatum Decne. in Jacquem. Voy. Inde Bot. 121. 1844. Mertensia elongata (Decne.) Benth. & poke t: am Pie ois: C. B. Clarke in Hook. f. Fl. Brit. India 4: 173. Eritrichium elongatum (Decne.) A. DC. in DC. ee 10: 124. 1846. Type: KAsHMiIr: Supra Pendjegram, 2600 m., Aug. 21, 1831. Jacque- mont s.n. (GH, P). Icon.: Jacquemont, l. c. ¢. 126. 1844. Perennial with underground rhizomes 10-15 cm. long and 0.5 cm. thick. Emerging shoots simple, greenish-white, erect, 10-25(—30) cm. tall, cov- ered densely with antrorse, appressed to subappressed, white, trichomes up .7 mm. long, rarely longer. Basal leaves petiolate; petioles up to 30 mm. long, margins ciliate, base broad, clasping the shoots; lamina oblong to oblong-lanceolate, narrowed at both ends, entire, acute, 15—50 mm. long, 6-8 mm. broad, covered on both surfaces, below ‘comparatively densely, with thin appressed trichomes, usually arising from minute tu- berculate bases; middle cauline leaves sessile or subsessile, to 70 mm. long, and 12 mm. broad; upper cauline leaves sessile and shorter than the middle ones. Inflorescence solitary, short and scorpioid when young, later elongated, 2—6(—9) cm. long, bearing 5—15 closely set (to 6 mm. 1970] KAZMI, BORAGINACEAE 375 distant) flowers; pedicels of the lower fruits up to 6 mm. long, gradually decreasing in size upwards, hairy. Calyx 3-5(-5. 5) mm. long, divided to the base, lobes linear, acute, erect, up to 0.5 mm. broad, covered sparsely dorsally and densely at the margins with thin, white, sub- appressed or rarely patent trichomes. Corolla blue, bluish- -purple, or deep blue, campanulate, (6—)7—9(-9.5) mm. long, glabrous, tube equal or slightly longer than the calyx, gradually extended from the base; lobes roundish, patent, 2.5-3.5 mm. long and 2-3 mm. broad at the base; faucal appendages poorly to well developed, 3-3.5 mm. above the corolla base, the area above the appendages usually puberulous. Anthers 1.5—-2 mm Hs oblong, acute, emarginate at bases, 1.5-2 mm. long; filaments 0. §: 1.5 mm. long, thick, attached 2.5~3.5 mm. above the corolla base. Nut- lets 4, glabrous, smooth, shining, acute, ovoid-trigonal, pallid, 2.5 mm. long, 1.5-1.6 mm. broad DISTRIBUTION: West Pakistan, Kashmir. West Pakistan: CuirraL State: Shishi Kuh, 2700 m., Harriss s.n. (pm). Gitcir AcEency: Babusar village, 4200 m., Siddiqui 27274 (MICH). Kashmir: East of Pirpanjal, above Tilpatra forests near ets Jhelum poms 3500-4000 m., Ludlow & Sherriff 7754 (pM, GH); Rajdhiangan Pass, 3500-4000 m., R. R. Stewart 17974 (GH, MICH, US), 225566 ean). 19292 (GH); Tragbal, 3300 st R. R. Stewart 4800 (micH), Koelz 9167 (GH); Poonch, near Bantara Gali, 2700 m., R. R. Stewart & Nasir 24084 (pm); above Gulmarg, Ru k: Stewart 10419, 8752 (GH); Haramukh, Ludlow & Sherriff 7870 (cH). Miscellaneous: Herb. East Ind. Co., Falconer s.n. (kK); Kohli 21(x). 6. P. drummondii Kazmi, sp. nov. Type: Kashmir: Pir Panjal (south side), 11,000 ft., June 27, 1902, Mr. J. R. Drummond 13919 (x-holotype). Herba perennis, rhizomatibus horizontalibus subterraneis. Surculi emergentes simplices, erecti vel suberecti, ad 20 cm. alti, dense vel laxe pilis albis, plus minusve crispatis, patentibus, ad 1 mm. longis vestiti. Folia basalia et inferiora petiolata, petioli 10-20 mm. longi, alati, hirsuti, ad basim dilatati, lamina ad 45 mm. longa et 15 mm. lata, late lanceolata, obtusa vel subacuta, supra infraque pilis albis ad 1 mm. longis, plerumque patentibus vel subappressis vestita; folia caulina mediana et superiora ad folia inferiora similia, sessilia, sursum decrescentia. Inflorescentia ter- minalis, simplex vel bifurcata, 5—9-florae, in juventute brevis et scorpioi- dea, demum eee pedicelli desunt vel ~ 1.5 mm. longi. Calyx ad basim divisus, lobi ad 5.5 mm. longi et 3 mm. lati, ovato-lanceolati, ex- terne nervo primario et marginibus dense biveitis cetera laxe hirsuti; in- terne paene glaber, erectus. Corolla atrocaerulea ad purpurea, glabra, cylindrica, tubus 7~10 mm. longus, calycem excedens vel paene duplo longior; lobi ad 4 mm. longi et 3 mm. lati, ovati vel subrotundati, integri, interne subpuberuli, erecti vel A GE appendices faucales triangulari- lingulatae, 2-3 mm. longae, 1-1.5 mm. latae ad basim, margines ciliati. 376 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Antherae 1.5—2 mm. longae, oblongae, subsessiles vel filamenta ad 0.5 mm. longa, 3-6 mm. supra basim corollae inserta. Stylus 10 mm. longus, stigma capitatum. Fructus non visus. DISTRIBUTION: West Pakistan, Kashmir. West Pakistan: Gitcir AGENcY: Naltar, Sept. 1960, McVean s.n. (&). Kashmir: Pir Panjal, 3600 m., Drummond 13919 (E-type). Only three species of the genus Pseudomertensia have faucal appen- dages, which are longer than broad, viz. P. lindelofioides (Rech. f. & Riedl) Riedl (in Rechinger, Fl. Iranica 48: 61. 1967), P. chitralensis and P. drummondii. P. lindelofioides, which is distributed in eastern and northeastern Afghanistan and which can be expected in West Pakistan, is distinguished from the other two by having large calyces, 7-8 mm long, corollas 14-15 mm. long, faucal appendages 3-4 mm. long and fila- ments 2—3 mm. long. In the other two species, along with the differences in form, the calyces do not exceed 6 mm., the corollas 12 mm., the faucal appendages 3 mm., and the filaments 2 mm. in length. The new species Pseudomertensia drummondii is more closely related to P. chitralensis than P. lindelofioides. It is distinguished from P. chitra- lensis by having denser and longer, up to 1 mm. long, spreading trichomes on the stems and leaves. The trichomes in P. chitralensis are appressed and do not exceed 0.3 mm. in length, except for a few scattered ones on the stem and petioles. The leaves in P. chitralensis have much longer petioles, and ovate laminae which are roundish or obtuse at the apices, whereas the leaves of P. drummondii have shorter petioles and lanceolate laminae, which are obtuse or acutish at the apices. In texture, the leaves of P. chitralensis are thin, while those of P. drummondii are thick. Flow- ers of P. drummondii are subsessile or short pedicellate and the pedicels attain a maximum length of 1.5 mm., while the pedicels of P. chitralensis ordinarily reach a length of 4 mm. The calyx lobes of the former are ovate to ovate-lanceolate, while those of the latter are linear. Faucal appendages of P. drummondii are 2-3 mm. long, 1-1.5 mm. broad at base, triangular and ciliate at the margins, while those of P. chitralensis are 0.8-1.2 mm. long, 0.5 mm. broad at base, oblong-lingulate, narrowed in the middle and entire at the margins. The former has longer anthers and shorter filaments compared to those of P. chitralensis. The specimen collected by McVean from Gilgit is a variation of P. drummondii. It agrees with the type specimen in all respects, except that it has slightly narrower calyx lobes, smaller corolla and the filaments are inserted only 3 mm. above the corolla base. 7. P. chitralensis (Riedl) Riedl in Rechinger, Fl. Iranica 48: 62. 1967. Mertensia chitralensis Riedl, in Kgie & Rechinger, Biol. Skr. 13(4): 228. 1963. Type: Chitral: Runbur, 3200 m., Bowes Lyon 664 (BM-holotype). Icon.: Riedl, l. c., fig. 174. 1963, under Mertensia chitralensis. 1970] KAZMI, BORAGINACEAE 377 Perennial with underground rhizomes. Emerging shoots simple or branched, + 15 cm. tall, tender, clothed with thin, short, appressed tri- chomes. Basal leaves petiolate; petioles 1-8 cm. long, winged, membranous, ciliate below, clasping the shoots at the bases; lamina shorter than the pet- ioles, elliptic, oblong-ovate or ovate-lanceolate, gradually attenuated to- wards the base, apex obtuse or roundish, rarely acute, margins entire, 2—5 cm. long, 0.8—1.7 cm. broad, uniformly covered on both surfaces with more or less loose, short, appressed trichomes: upper leaves few, smaller, and with shorter petioles. Inflorescence solitary, short, 2-3 cm. long when young, later elongated, 7-8 cm. long, bearing 7-12 flowers; pedicels 1—4 mm. long. Calyx divided to the base, lobes 4~4.5 mm. long, up to 0.7 mm. broad, in fruit elongated to 6 mm., linear, obtuse, erect, margins densely pubescent, trichomes up to 1 mm. long, few scattered trichomes on the upper surface. Corolla blue to bluish purple, infundibuliform, glabrous, 10-12 mm. long, tube 7-9 mm. long, slightly dilated upwards, lobes roundish, obliquely patent, ca. 2 mm. long and broad; faucal ap- pendages 0.8-1.2 mm. long, 0.5 mm. broad, narrowed in the middle, slightly emarginate above. Anthers oblong, more or less rounded at both ends, ca. 1.2 mm, long, 0.4 mm. broad, tips approaching the sinuses be- tween the corolla lobes; filaments 0.7 mm. long, slightly dilated at base, attached below the faucal appendages. Style filiform, ca. 11 mm. long; stigma capitate. Nutlets oblong or ovate, trigonous, 2.5—3 mm. long, ca. 1 mm. broad, slightly puberulous externally, pubescent inside, pallid. DISTRIBUTION: West Pakistan. West Pakistan: CHITRAL STATE: Rumbur, 3200 m., Bowes Lyon 664 (BM); Pattisun, Toppin 191 (x). 8. P. anjumiae Kazmi, sp. nov.t Type: West Pakistan: Kurram, up the Shend Toi valley, near water at an elevation of 11,000 ft., Aitchison 192 (pm-holotype, GH-isotype). Surculi tenues subterranei, radices fibrosas emittentes fibroso-radicantes et squamis parvis obsiti, Caules floriferi ad 20 cm. longi, arcuati vel erecti tenues herbacei, pilis patulis sparse obsiti, in cincinnum singulum terminalem aphyllum abeuntes. Folia basalia plura, lamina oblongo-ovata vel oblanceolata, (1—)2—4(-—5) cm. longa, 5-10 mm. lata, basi longe at- tenuata, apice breviter attenuata vel subrotundata, plana, integra, tenuis- sime molliterque herbacea, costa mediana tenuis, nervis secundariis vix visibilibus, supra et subtus pilis sparsis brevissimis appressis tecta, longe tenuiter petiolata, petiolo 2—5 cm. longo; folia caulina pauca breviter petiolata, petiolis 4 cm. longis, 0.5 mm. latis. Inflorescentiae solitariae, primo densae, scorpioideae demum ad 8 cm. elongatae, 5~20-florae. Pedi- celli 1-10 mm. longi. Calyx 3-4 mm. longus ad basin usque in laciniis anguste linearibus obtusis fissus, pilis arcuatis patulis ad 1 mm. longis obsitus. Corolla quam calyx duplo longior, in sicco caeruleo-violacea, 8 *I named this species after the name of my wife, Anjum. 378 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 mm. longa, subcylindrica vel infundibuliformis, utrinque glabra, lobis erectis vel oblique patulis, ca. 1.5 mm. longis et latis, rotundatis. For- nices tenuiter evoluti. Filamenta 1—-1.2 mm. longa, 7 mm. supra basin tubi me Antherae ca. 1 mm. longae versatiles, atrae, oblongae, ad apicem t basin rotundatae. Stylus corollam saepe superans, tenuiter filiformis, a. Stigma minutum subcapitatum. Nuculae ignotae. DISTRIBUTION: West Pakistan. West Pakistan: KurrAM: Shend Toi valley, Aitchison 192 (BM, GH). The specimen cited here, collected by Aitchison from Kurram valley, has been cited by Ried] (in Rechinger, Fl. Iranica 48: 62. 1967) under portant distinguishing characters of P. echioides are the very short (0.5 mm.) filaments, shorter than the anthers (1.5-1.9 mm.) which are acute at their apices, and the absence of faucal appendages. These characters are well correlated with the long (3—4.5 mm.) and narrow (1.5 mm corolla lobes, which more or less equal or sometimes even exceed the corolla tube in length. Pseudomertensia anjumiae differs from P. echioides in having longer (1—1.2 mm.) filaments, equalling or slightly exceeding the length (1 mm.) of the anthers which are roundish at apices. The faucal appendages are quite developed. Corolla lobes are much reduced, only 1.5 mm. long and broad, about 1/5 the length of the corolla tube, and not oblong-ovate but roundish. Pseudomertensia anjumiae is dis- tinguished from the related species, P. moltkioides, in having shorter corolla lobes and longer filaments. In P. moltkioides corolla lobes are 2.5-4.5 mm. long, 2-3 mm. broad, and oblong-ovate in shape; the fila- ments, 0—-0.5 mm. long, are dilated at the base. 9. P. nemorosa (DC.) Stewart & Kazmi, comb. nov. Lithospermum ovalifolium Decne. in Jacquem. als pred ieee 121. 1844. Eritrichium nemorosum A. DC. in DC. Prodr. 10: 846. Mertensia nemorosa (DC.) I. M. Johnston, Jour. pare Arb. 37: 305. 1956. Mertensia moltkioides (Benth.) C. B. possi e var. thomsoni C. B. Clarke in ook. f. Fl. Brit. India 4: 170. 1883, syn. Type: Kashmir: In humidis nemorum ad Ouri, May 4, 1831, Jacque- mont s.n. (GH, P). Icon.: Jacquemont, 1. c. ¢. 124. 1844, under Lithospermum ovalifolium; Coventry, Wild Fl. Kashmir 3: ¢. 44. 1930, under Mertensia moltkioides. Perennial with stout, little branched, underground rhizomes, up to 20 cm. long, sometimes to 10 mm. thick. Emerging shoots weak, curved or erect, 6-15(—20) cm. tall, clothed mostly with spreading, thin, white trichomes up to 1 mm. long. Basal leaves petiolate; petioles 2—8 cm. long, hairy, broad and clasping the shoot at their bases; lamina ovate, elliptic, or ovate-lanceolate, margins entire, mostly acute at apex, round or ob- 1970] KAZMI, BORAGINACEAE 379 tuse at base, rarely truncate, never cordate, (2—)3-7(-8) cm. long, 1-4.5 cm. broad, upper surface slightly hairy, lower surface comparatively densely hairy especially on nerves, trichomes thin, not arising from tu- berculate bases, appressed to suberect, rarely patent; upper cauline leaves shorter, narrower, with short petioles, subsessile or sessile. Inflorescence at flowering 4~9 cm. long, bearing 6~14 flowers, in fruit much elongated, with fruits sometimes up to 10 mm. distant; pedicels 1-9 mm, long, hairy. Calyx divided to the base, lobes linear-oblong, obtusish, erect, up to 7 mm. long, covered externally with subappressed or spreading trichomes. Corolla blue, bluish-purple to deep blue, glabrous, corolla tube cylindrical, (8-)9-11 mm. long, lobes ovate, suberect to patent, 3.5-4 mm. long, 3- 3.5 mm. broad at the base; faucal appendages small 0.7-0.8 mm. broad, 0.5-0.6 mm. long, anthers 1 mm. long, 0.5 mm. broad, oblong, rounded at both ends. Filaments very short, nearly triangular, 0.5 mm. high, 0.7 mm. broad at the dilated base, attached in between and slightly below the bases of the faucal appendages. Style filiform, ca. 8 mm. long; stigma small, subcapitate. Nutlets 3 mm. long, 1.5 mm. broad, acute at apices, slightly verruculose on the ridges, dark brown. DISTRIBUTION: West Pakistan, Kashmir, northwest India. West Pakistan: Hazara Dist.: Kaghan valley, between Naran and Batakundi, Kazmi 795a (PEs); Swat State: Mt. Ilam, R. R. Stewart 24373 (Raw); Mis- cellaneous: locality unknown (probably from Swat) Kazmi s.n. (PEs). Kashmir: Poonch: Serimang, 2000-2700 m., R. R. Stewart & Nasir 25540 (sm); Lidder valley: Kolhoi, Znayat 25711 (kK); 3300 m., Ludlow 8 (BM); Chashma Shai, 2300 m., R. R. Stewart 10926 (GH); above Uri on Poonch road, 2000 m., R. R. Stewart 12954 (GH); ad Ouri (Uri ca. 20 miles from Baramula and 40 miles west of Srinagar), Jacguemont s.n. (GH, P); Lidderwat, 3000 m., Coventry 575 (kK), Evershad s.n. (Bm); Sind valley, Gangangir, 2500 m., Rich 1075 (K); Kishanganga valley: near Nilam, Kazmi s.n. (pes); Islampur, Kazmi 295b (pes); Kel, Kazmi s.n. (pes); between Shardi and Kel, 1950-2100 m., Schmid 1744 (pm, MicH); Sind valley, Mehanmarg, 2500 m., Ludlow & Sherriff 8147 (pm); Chenab-Ravi watershed, Badawar to Padri Pass, June 3, 1848, Thomson s.n. (K); south of Chenab valley between Baleta and Katti, Herb. Ind. Or. Hook. f. & Thoms., May 14, 1948, Thomson s.n. (K). Miscellaneous: Hab. Himal. Occ., 6000-7000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson sm. (GH); Herb. East India Co., Falconer s.n. (GH); Palmer s.n, (BM). Pseudomertensia nemorosa is very Closely related to Mertensia race- mosa (Royle) C. B. Clarke (in Hook. f. Fl. Brit. India 4: 171. 1883) of Kumaon, Bashahar and Kulu, which also agrees more closely in its characters to Pseudomertensia than to Mertensia and is here recombined as Pseudomertensia racemosa (Royle) Kazmi.? Pseudomertensia nemo- rosa differs from P. racemosa in its blue-purple rather than white corollas, its more pubescent leaves, not subcordate at the bases, and its coarser * Pseudomertensia racemosa (Royle) Kazmi, comb. nov. : Myosotis racemosa Benth. in Royle, Ilustr. Bot. Himal. Mount. 1: 305. 1839. Mertensia racemosa (Royle) C. B. Clarke in Hook. f. Fl. Brit. India 4: 171. 1883. 380 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 habit. The next most closely related species is P. moltkioides from which it differs mainly in the form and size of leaves. In P. moltkioides the leaves are oblong, oblong-lanceolate, lanceolate, or rarely narrowly lan- ceolate and are up to 10(—12) mm. broad, their bases always attenuated into short or long petioles; while in P. nemorosa the leaves are ovate, elliptic, or ovate-lanceolate, and are usually 10-45 mm. broad, their bases always round, obtuse or rarely truncate, and not gradually attenuated into petioles as in P. moltkioides. Dr. R. R. Stewart has compared the specimens, /nayat 25711 and Rich 1075 at the Kew Herbarium with Thomson’s collection from Islam- abad, Kashmir, on which Mertensia moltkioides var. thomsoni C. B. Clarke (in Hook. f. Fl. Brit. India 4: 170. 1883) is based, and found them to be identical. These specimens (cited above) and Clarke’s description “Radical leaves 3 & 11/2 in. (75 mm. long, 40 mm. broad), base sub- obtuse” agree with P. nemorosa. 10. P. moltkioides (Royle ex Benth.) Kazmi, comb. nov. Myosotis moltkioides Royle ex mene ig qt Illustr. Bot. Himal. Mount. 1: 305. 1836; ¢. 73. fig. 1. [Dec.j] 1 Anchusa moltkioides Royle ex Benth. ie .; DC. Prodr. 10: 51. 1846. sep nanategese pacar (Royle ex Benth.) Decne. in et Voy. Inde Bot. 122. Mertensia ene C. B. Clarke in Hook. f. Fl. Brit. India 4: 171. 1883 Mertensia nuristanica Rech. f. Ann. Naturh. Mus. Wien 58: 56. 1951 Type: Kashmir: Pirpenjal, Royle (?) Icon.: Royle, /. c. 1835, under Anchusa moltkioides. Perennial with long, slender, branched, underground rhizomes, up to 5 mm. thick. Emerging shoots profusely leafy at the bases, naked or with few distant leaves above, 5-15 cm. tall, clothed usually with short and appressed or sometimes with long and patent trichomes. Basal leaves petiolate, petioles shorter or longer than lamina, lamina oblong to oblong- lanceolate to lanceolate, rarely linear-lanceolate, entire, acute to obtuse, (including petioles) 15—70(-80) mm. long, 2—10(—12) mm. broad, cov- ered on both surfaces with short appressed to subappressed trichomes, rarely glabrous below, cauline leaves few, sessile, shorter and narrower than the basal ones. Inflorescence 2—3 cm. long with sessile or subses- sile closely set flowers when young, later elongated up to 7 cm. long, with distant (up to 10 mm, apart), short pedicellate flowers; pedicels up to 3 mm. long. Calyx divided to the base, lobes linear to linear-lanceolate, obtuse to acute, erect, 3-4(—5) mm. long, 0.5 mm. broad, covered with short, patent to subpatent trichomes. Corolla blue, bluish-purple to deep bluish-purple, infundibuliform, externally glabrous, internally glabrous to puberulous or minutely pubescent above the throat, tube 8-15 mm. long, lobes oblong-ovate, patent, 2.5-4.5 mm. long, 2-3 mm, broad; faucal ap- pendages well developed, broader than long, slightly emarginate and puberulous. Anthers oblong, roundish at both ends, 1—1.75 mm. long; 1970] KAZMI, BORAGINACEAE 381 filaments more or less broad at the base, 0-0.5 mm. long. Style filiform; stigma subcapitate. Nutlets ovate, trigonous, light to dark brown, 3 mm. long. DISTRIBUTION OF SPECIES: Afghanistan, West Pakistan, Kashmir, northwest India, and Tibet. This species, Pseudomertensia moltkioides, is very variable in the length of petioles and lamina, number of leaves on the shoots, length of calyx lobes and corolla tube, indument of minute trichomes on the corolla throat, and the insertion of the anthers on the corolla. The four varieties which follow may easily be distinguished from one another. KEY TO THE VARIETIES a. Anthers distinctly below the bases of the faucal appendages............... PE TN Ee er a ee ee ja. var. moltkioides. a. Anthers distinctly exceeding or surpassing the bases of the faucal appendages. b. Leaves: linearspathulate, 06° 0 go avin ae 10d. var. tanneri. b. Leaves oblong to oblong-lanceolate. c. Leaves short petiolate, scapes naked, corolla internally, above the b tHYOSE, TRNGUE i ee eo . var. primuloides. c. Leaves with long petioles, scapes leafy, corolla internally, above the throat, densely minutely pubescent. ............ 10c. var. leichtlinit. 10a. Var. moltkioides. Basal leaves usually with long, rarely with short petioles; corolla tube mostly 3—5 times longer than the calyx; corolla usually glabrous within, rarely puberulous; anther tips just below to a few millimeters below the faucal appendages. West Pakistan: CHITRAL STATE: Golen, Shishikuh, 4700 m., Bowes Lyon 106 (pM). Gitcir AGENcy: Rama valley, southwest of Astore, 4000 m., Lankester Kamri top, 4000 m., R. R. Stewart 18675 (GH); Mekerum, Hispar glacier, 3500 m., Russell 1450 (cH); Haramosh La, Culbert 86 (pm); Agres Camp, below Hispar Pass, Karakorum, 4100 m., Conway 209 (Bm); near Hispar, Shukurri, 4400 m., Conway 166 (kK); Nagar, Gharesa glacier, slope facing north, 12 miles east of Nagar, Polunin 6127 (pm); Minapin glacier, 4100 m., Lloyd & Megan 105 (pm). Hazara Dist.: Kaghan valley, between Naran and Saiful Maluk, ca. 2700 m., Kazmi 200b (pes); Ganji Pahari, near Naran, half way to the top, ca. 2500 m., Kazmi 852c (PES); 1.5 miles north of Naran, Burtt & Arshad Ali B857 (£); Basal, Schmid 363 (£); between Shahid Pani and Musa ka Musalah, on rocky slope facing Kaghan valley, ca. 3000 m., Kazmi 2514 (pes); Siran valley, near the foot of Musa ka Musalah, ca. 2800 m., Kazmi 2491 (PEs). Kashmir: Apharwat above Gulmarg, 4000 m., Rich 1230 (xk); Apharwat, 00 m., Polunin 56/196 (£); above Gulmarg, Fuller 153 (x); Pirpanjal, Inayat 25707 (K); top of Masjid Gali, 4500 m., R. R. Stewart 18363 (Gu, 382 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 MIcH); Kishtwar, Dudhari Pass, 4000 m., Ludlow & Sherriff 9216 (GH); Kish- anganga valley, above Nilam, ca. 3200 m., Kazmi 302c (PEs); Chinari, Zakia Bandi, on rocky slopes, Kazmi 258a (PES) The binomial Mertensia moltkioides (Benth. in Royle) C. B. Clarke (in Hook. f. Fl. Brit. India 4: 170. 1883) is based on the illustration of Anchusa moltkioides (Royle ex Benth. in Royle, 1. c. ¢. 73. fig. 1. 1835), described by Royle ex Bentham (in Royle, /. c. 1: 305. 1836) as Myoso- tis moltkioides, is a misidentification which was further confused by C. B. Clarke’s description (/. c.) and, therefore, ignored by many authors. The illustration of Anchusa moltkioides gives all the necessary details of the form of the leaves and the structure of flowers and should be ac- cepted as type. This illustration shows clearly the scapose peduncle, oblong leaves with long petioles, and the anthers distinctly below the faucal appendages. On the basis of these distinguishing characters two of the three specimens cited by C. B. Clarke (/. c.) (Kashmir, Falconer, and near Islamabad, Thomson) do not agree with M. moltkioides, but are, rather, Pseudo- mertensia nemorosa. The collection of Dr. Watt from Pangee, which I was unable to examine, may also not belong to Mertensia moltkioides. Actually C. B. Clarke’s description of Mertensia tibetica represents the type of Pseudomertensia moltkioides. He cited the following details ‘“— leaves small subradical long-petioled elliptic, peduncles long subscapose — anthers entirely below the scales — nutlets as of M. moltkioides.” Pseudo- mertensia nuristanica Rech. f. represents only a form of P. moltkioides in which the anthers are attached to the corolla tube a few millimeters below the faucal appendages. Both these binomials are therefore synonyms of P. moltkioides. Ried] (in Rechinger, Fl. Iranica /. c.) recognized the specific name Pseudomertensia primuloides for the taxon containing the forms of Mer- tensia tibetica and M. nuristanica. As the dates of publication of Royle’s Tllustr. (/. c. 1835, 1836) precede the date of publication of Jacquemont’s Voyage (l. c. 1844), P. moltkioides has priority. 10b. Var. primuloides (Decne.) Kazmi, comb. nov. Eritrichium primuloides Decne. in Jacquem. Voy. Inde Bot. 123. 1844; DC. Pro 25. 1846. getie ct hesiaraet (Decne.) C. B. Clarke in Hook. f. Fl. Brit. India 4: 171. var. Pil Seabee rimuloide (Decne.) Riedl in Rechinger, Fl. Iranica 48: pa Type: Kashmir: rhea Jacquemont s.n. (P). Icon.: Jacquemont, I. c. ¢. 128. 1843. Basal leaves usually with short, sometimes with long petioles; corolla tube usually 2 to 3 times longer than the calyx; corolla internally, above the throat, usually puberulous, sometimes minutely pubescent; anthers distinctly exceeding or surpassing the bases of faucal appendages. 1970] KAZMI, BORAGINACEAE 383 West Pakistan: CurtrRaL State: Drosh Nullah, 3800 m., Toppin 757 (x); Shera Shing Pass, 4800 m., Stainton 3208 (pm); Ziarat (Seveiik Pass), 3000 m., Stainton 3208 (BM), 2330 (E). Hazara Dist.: Kaghan valley, Champion $M. (GH); Makra, 4500 m., Duthie s.n. (kK); between Shogran and Bhunja, Kazmi 732b (PEs); near Sari, between Shogran and Makra, ca. 3000 m., Kazmi ncaa (PES); Chapri, 28. 7. 1899, Inayat s.n. (kK); Naran, stony ground, 3000 , Jafri & Ali 3389, 3332 (gE); Naran, June 1960, McVean sn. (E). Miscel- pati northwest India (marked b), J. L. Stewart s.n. (£). Kashmir: Apharwat, above Gulmarg, 3400 m., Pinfeld 193 (pm), Rich 1257 (kK), R. R. Stewart 15486, 10327, 8604 (cH), Steone 43 (E); above Gulmarg, Aitchison 73 (K); Pirpanjal, 10. 8. 1901, Znayat s.n. (K); Bringhi valley, Mantnar Nullah, 4300 m., Ludlow & Sherriff 8050 (Bm, GH); Mantnar valley, near Desu, 4100 m., Ludlow 115 (GH); Kishenganga valley, near Nilam, Kazmi 316b (PES); Muzafarabad, Musa, 18. 7. 1899, Imayat s.n. (K); Kamri Pass, 4500 m., Duthie 12576 (sm); Kamri Pass top, 4300-4600 m., R. R. Stewart 22690 (GH); Ablau valley, Bringhi river, 4000-4300 m., Ludlow 21 (pm). Among the specimens ee above, Ludlow 21 differs have the others in having filaments up to 0.5 mm. lon 10c. Var. leichtlinii Kazmi, var. nov. Type: Hazara, Max Leichtlin 4 (BM). Folia basalia longipetiolata, petiolis saepe laminis aequantibus; corol- la intra, supra appendices fauces, minute densique pubescens; scapi ple- rumque foliati. West Pakistan: HAzArA Dist.: Duthie s.n. (kK); Leichtlin 4 (pm-holotype of var.); between Balaket and Babusar Pass, Abel 132 (Bm); Kaghan valley, Naran, Shaukat Ali 125 (BM). 10d. Var. tanneri (C. B. Clarke) Stewart & Kazmi, comb. nov. Mertensia primuloides (Decne.) C. B. Clarke var. tanneri C. B. Clarke in Hook. f. Fl. Brit. India 4: 171. 1883. Type: Daskin, Astore, Tanner 10 (kK). Icon.: Riedl, in Kgie & Rechinger, Biol. Skr. 13(4): fig. 125. 1963. Leaves linear-spathulate with longer petioles, calyx lobes 4-5 mm. long; corolla violet, corolla tube ca. 8 mm. long. West Pakistan: Grtcir AGENcY: Daskin, Astore, Tanner 10 (xk); Sai, Gilgit, Tanner s.n. (K); Rattu et Goriket, Schmid 1774 (pm); Rama valley, southwest of Astore, 3500 m., Lankester & Pearson 2030 (pM). Hazara Dist.: Kaghan valley, Naran, Barrett s.n, (kK 9. Anoplocaryum Ledeb. FI. Rossica 3: 154. 1847. Echinospermum sect. 3. Anoplocaryum Turcz. in Bull. Soc. Nat. Moscou 23(1): 522. 1850. Type species: Anoplocaryum compressum (Turcz.) Ledeb. 384 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Annual (or perennial ?) herbs. Basal leaves usually petiolate. Inflo- rescence bracteate, pedicellate, pedicels usually longer than the calyx and reflexed in fruit. Calyx divided to the base, not much enlarged in fruit. Corolla hypocrateriform to subrotate; faucal appendages distinct. Gyno- base shortly pyramidal with 4 triangular depressions. Nutlets ovoid, rough or slightly tuberculate, margins irregularly setulose, carinate in- ternally from the carunculate scar to the top. Species about 5, distributed from China to the Western Himalayas. A. brandisii Brand, Repert. Sp. Nov. 22: 100. 1925, Pflanzenr. IV. 252(Heft 97): 116. 1931. PE: Chamba State: Sach Pangi, Brandis 3166 (pp); Lahul, Brandis 3236 (pp); Haelle (?), Brandis 3996 (pp), holotype not indicated. Perennial or annual herb. Stems many, slender, thinly covered with short, thin, appressed trichomes, 15-30 cm. tall. Basal leaves 15-25 mm long, 2-4 mm. broad, entire, obtuse, gradually narrowed at the base into a short petiole, covered with thin, short, appressed trichomes on both surfaces; cauline leaves subsessile to sessile, shorter and narrower. In- florescence terminal, lax, lower flowers bracteate; pedicels hairy, 1-2 mm. long, elongated in fruit, to 12 mm., recurved. Calyx divided to the base, lobes lanceolate, slightly hairy, in flower 1.5 mm. long, in fruit usually subreflexed, to 2.5 mm. long, 0.5 mm. broad. Corolla subrotate, 4-5 mm. in diameter; faucal appendages semilunar. Gynobase triangular, distinct from the equally long style. Nutlets 3-4, ovoid, 2 mm. long, margins densely minutely setulose, granulate and slightly rugose on both surfaces, internally distinctly and externally slightly carinate, attached obliquely to the gynobase by a large carunculate scar, DISTRIBUTION: West Pakistan, Kashmir, northwest India. Pakistan: GILcIr AGENCY: Niltar valley, north of Gilgit, 3500-3700 m., Duthie 12425 (GH, K). Reported from near Gilgit, Roberts (Brand, 1. c.). 10. Eritrichium Schrad. Comment. 4: 186. 1820. Eritrichium Schrad. sect. Eu-eritrichium DC. Prodr. 10: 124. 1846. Type species: FE. nanum (Vill.) Schrad. Annual or perennial, strigose or villous herbs with alternate leaves. Racemes simple or branched, usually terminal or sometimes axillary. Calyx divided to the base, lobes 5, sometimes enlarged and reflexed in fruit. Corolla subrotate to infundibuliform, lobes spreading; faucal ap- pendages distinct. Stamens inserted on the corolla tube, included; anthers ovate. Ovary 4-lobed, style filiform, stigma capitate. Gynobase sub- prismatic-pyramidal, shorter than the style. Nutlets 4, surpassing the gynobase, attached obliquely to the depressions in the gynobase, at margins usually appendiculate; appendages various. Species about 25, centering in Asia, distributed to Europe and America. 1970] KAZMI, BORAGINACEAE 385 KEY TO THE SPECIES = ey leaves in dest heaina covered on both surfaces with A ai 1-2 m. long, margins ciliate: (26.423 2.405.-. Se ee 1. E. nanum. Basal leaves not in Rms 5 petiolate, covered on both surfaces ee trichomes p to 1 mm. long, margins usually not ciliate, rarely with a few scattered long trichomes b. Cauline lewves gradually narrowed towards the bases, seis to short POUCINE. ee eG Cae eee 2. E. fruticulosum. b. Cauline leaves broad at the bases, always sessile. Cauline leaves ovate to oblong- ovate, roundish to obtuse at apices, usually covered with unequal trichomes. Pane a 3. E. spathulatum, Cauline leaves oblong-lanceolate to linear-filiform, apices obtuse to acute, usually covered with equal trichomes. d. Nutlets triangular-oblong (excluding appendages), 1 . long, 1 mm. broad, outer surface convex or slightly eid a raised in sis middle, marginal appendages 0.2 mm. long, distant, not con- fluent at their DASPSs: on die ah Nee eas pean, 4. E. patens. Nutlets ovate (excluding appendages), 2 mm. long, 1.5 mm. broad, outer surface plane or slightly concave, marginal appendages 1 mm. long, contiuent At their haseg? = 0 ee 5. E. canum » ee ea 1. E, nanum (Vill.) Schrad. Comment. Soc. Regiae Sci. Gott. 4: 186. . Pop. in Fl. URSS. 19: 514. 1953; Riedl in Rechinger, Fl. Tranica 48: 64. Myosotis nana Vill. Hist. Pl. Dauph. 2: 459. 1878, Prosp. Pl. Dauph. 21. 879. Type: “Sur les sommets des Alpes au dessus de Brande et d’Allemont .. without citation of collector’s name (?). Icon.: Vill. 1. c. ¢. 13. 1878; Hegi, Ill. Fl. Mitt.-Eur. 5(3): ¢. 219, fig. 2a-c, & fig. 3095. 1927. Perennial, densely caespitose; roots much branched near the surface of the ground, each branch bearing a thick rosette of leaves and producing a single, usually simple or rarely branched stem. Stems very short, to 12 cm. long, covered thinly or densely with thin, silky, usually spreading trichomes. Basal leaves numerous, sessile, oblong-lanceolate, entire, roundish or obtuse at apices, margins thickly ciliate, 5-10 mm. 2-3(—5) mm. broad, both surfaces covered with thin, silky, trichomes up to 2 mm. long; cauline leaves few, usually narrower. Inflorescence short, 3—6-flowered, lower flowers bracteate, bracts leaflike, 1-2 mm. long; upper flowers subsessile, lower flowers pedicellate, pedicels up to 1.5 mm. long, hairy. Calyx divided nearly to the base, lobes oblong-lanceolate, in flower 2.5-3 mm. long, in fruit elongated to 4 mm., densely hairy, tri- chomes long, sometimes yellowish. Corolla usually blue, rarely white, with a yellow eye, campanulate-rotate, tube equalling or a little longer than the calyx, slightly narrowed at the throat, lobes ovate, spreading, long, 386 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 0.5-3.5 mm. long, 1-3 mm. broad; faucal appendages linear-horizontal. Nutlets ca. 2.5 mm. long, at margins dentate, glabrous. DISTRIBUTION OF SPECIES: European Alps, Caucasus, Altai, Siberia, western China, Tibet, Afghanistan, West Pakistan, Kashmir, northwestern India. la. Subsp. nanum. Basal leaves many; stems short; corolla 6-7 mm. long. DISTRIBUTION OF SUBSP.: Represents the species in the European and western Asian portions of its range. 1b. Subsp. villosum (Ledeb.) Brand, Pflanzenr. IV. 252(Heft 97) : 189 1931; Riedl in Rechinger, Fl. Iranica 48: 64. 1967 Myosotis villosa Ledeb. Mem. Acad. St.-Pétersb. 5: 516. 1815; Flora Altaica 91. 1829. Eritrichium villosum (Ledeb.) Bunge, Verzeichn. Altaigebirg. Pflanzen 14. 1836; DC. Prodr. 10: 126. 1846. Echinospermum sericeum Benth. in Royle, Ilustr. Bot. Himal. Mount. 306. Eritrichium sericeum (Benth. in Royle) DC. Prodr. 10: 126. 1846. Eritrichium basifixum C. B. Clarke in Hook. f. Fl. Brit. sag rs 165. 1883. E. nanum subsp. villosum var. eu-villosum Brand, 1. c. 190. 1931. Type: “Habitat in Sibiriae alpibus,”’ without citation of collector’s name (LE). Icon.: Ledeb. Icon. Fl. Rossica 2: ¢. 215. 1830. Basal leaves few, stems longer than in the typical subsp.; corolla 3—5.5 mm. long. DISTRIBUTION OF suUBSP.: Altai, Siberia, Tibet, Afghanistan, West Pak- istan, Kashmir, and northwest India. Pakistan: GiLcir AGENCY: Rimochagma valley 4700 m., 20. 7. 1892, Duthie s.n. (BM, E); Baltistan, Hunter-Weston 10253 (MIcH); snow field on the south slope of Burja La, 4700-5000 m., Webster & Nazir 6546 (GH); Talala, 4300 m., Koelz 9747 (cH), R. R. Stewart 20770 (GH); Jutial Nullah, R. R. Stewart 26326 (BM); Ramna valley, southwest of Astore, 4000 m., Lankester & Pearson 1345, 1346 (pm); Chota Deosai, 4000-4500 m., R. R. Stewart 19934 (GH, MICH, US). Hazara Dist.: Kaghan valley, between Paya and Makra, above Shogran, ca. 3000 m., Kazmi 2144b (PEs); Gitidas, 3700 m., Metz s.n. (micH); Kaghan valley, /. : Chaudhri s.n. (RAW); Siran valley, near Shahid Pani, ca. 3100 m., Kazmi 193c (PEs). shmir: Kamri Pass top, 4500 m., R. R. & I. D. Stewart 18708, 22677 (GH); Burzil Pass, 4300 m., Duthie 14038 (pm); Zaskar, Pansi La, 5500 m., Koelz 5938 (GH, MICH, us); Kargia, Lagong, 4500 m., Koelz 5392 (cH, MICH, US); Ladak, Hanu Yogma vi a Handamir up the Che rbat La Pass (Pass between Indus valley and the Ghapek oalniors Schlagintweit 6513 (GH); Barnaj Nullah, Sap- 1970] KAZMI, BORAGINACEAE 387 phire mines, Kishtwar, Ludlow & Sherriff 9151 (GH); Masjid Gali top, 4000 m., R. & I. D, Stewart 18363B (cx). 2. E. fruticulosum Klotzsch, Reis. Prinz Wald. Bot. 96. 1862. E. strictum Decne. in Jacquem. ae Pines oa (Klotzsch) C. B. Clarke in Hook. f. Fl. Brit. India 4: 164. 1 Type: Himalaya, Dr. coil s.n, (HEID). Icon.: Klotzsch, 1. c. t. 62. 1862. Perennial herb with woody tap root up to 15 cm. long. Stems many, erect to decumbent, slender, up to 30 cm. long, simple or sometimes branched in the upper part; branches short, stem and branches green, covered with thin, white, antrorsely appressed or subappressed soft tri- chomes not exceeding 0.5 mm. in length. Basal leaves few, petiolate, petioles slender, hairy, up to 20 mm. long; lamina pale green, ovate-lan- ceolate, ca. 25 mm. long, 5 mm. broad, obtuse, attenuate towards the base, covered uniformly and loosely on both surfaces with thin, appressed trichomes up to 0.5 mm. long; cauline leaves subsessile to short petiolate, petioles sometimes up to 3 mm. long, lamina as in the basal leaves, acutish, shorter and narrower. Inflorescence simple or rarely geminate, racemes loose, 10—15-flowered, bracteate; pedicels slender, hairy, more or less re- flexed in fruit, up to 6 mm. long. Calyx divided to the base, lobes 1-1.5 mm. long, 0.2—0.3 mm. broad in flower, lanceolate, narrower towards the base, pubescent on both faces, enlarged in fruit to 2.5 mm. long and 0.7 mm. broad, spreading or slightly reflexed. Corolla light blue with a yel- low eye, rotate, tube more or less equalling the calyx, lobes roundish, 1.5 mm. long and broad. Nutlets 4, excluding glochidiate appendages, 1.7- 2 mm. long, 0.8-1 mm. broad, roundish at the base, acutish at apex, outer face convex, margined with glochidiate non-confluent appendages, 0.5 mm long, 0.2 mm. broad at base, pubescent on both faces with short, stiff, erect trichomes. DisTRIBUTION: West Pakistan, Kashmir, northwest India. West Pakistan: Gitctr AcENcy: Gilgit proper, Manugah Nala, river bank, 3500 m., J. W. Thornley 27 (sm); Baltistan, E agobe: valley, 3000-3500 m., Duthie 12144 (pm); Hunza, near Baltit, 3000 m., R. R. Stewart 26327 (om). Quetta Dist.: Ziarat, 2600 m., 10. 9. 1887, Hiss s.n. (E). mir: La , Gya, Koelz 6412 ’ (Mic, us); Nubra, Tsanglung Pass to idseveas 3 Peas ae 1856, Schlagintweit sn. (GH); Rupshu, 4700 m., Koelz 6654 (MICH, US). Miscellaneous: Himal. Bor. Occ., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (BM, GH). C. B. Clarke (J. c.) recognizes Eritrichium fruticulosum as a variety of E. strictum and distinguishes it from £. strictum and its variety thomsoni by “weaker, greener, stems diffuse sometimes with divaricate branches.” Eritrichium strictum and its variety thomsoni are considered in this work as synonyms of E. canum from which E. fruticulosum differs not only in 388 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 being more herbaceous with slender and more or less decumbent stems, but also in the form and pubescence of the leaves, and the shape and appendages of the nutlets. The figure given by Klotzsch (/. c.), which should be accepted as the type, shows very clearly the sessile or short petiolate cauline leaves nar- rowed towards the base, generally thinly and uniformly covered with very short appressed trichomes. These characters together with the ob- long form of the nutlets, which are roundish at the base and acutish at apex, margined with non-confluent appendages 0.5 mm. long, 0.2 mm. broad at the base and with both surfaces Esl with short, stiff, erect trichomes, distinguish this species from its close relatives. Th E. canum and its varieties the leaves are densely pubescent and are not gradually attenuated towards the base, but are broad and sessile. The nutlets are larger, with longer appendages, confluent at base and forming narrow or broad marginal wings. : as = Fe ser sar Fa rae in Royle) C. B. Clarke in Hook. f. Fl. Brit. India 4: 164. perenne spathulatum ee in Royle Illustr. Bot. Himal. Mount. 306. 1836; DC. Prodr. 10: 142. 1846. Brivichiam jacquemontit ohare in Jacquem. Voy. Inde Bot. 122. ¢. 127. E. aa (Pall.) Bunge var. pectinatum (Pall.) Brand subvar. spathulatum (Benth. in Royle) Brand, Pflanzenr. IV. 252(Heft 97): 193. 1931. Type: Chango in Kunawur, with citation of collector’s name (K, P). Icon.: Jacquemont, |. c. ¢. 127. 1843, as Eritrichium jacquemontii. Perennial, weak, decumbent herb. Stems many, to 20 cm. long, simple or branched, greenish, covered thinly or densely with short, appressed, white trichomes, Basal leaves petiolate, petioles 10-20 mm. long, slender, hairy; lamina, ovate-lanceolate, 20-30 mm. long, 3-7 mm. broad, gradu- ally attenuated towards the petiole, obtuse to roundish at apex, margins entire, both surfaces covered thinly or densely, usually with unequal, subpatent white trichomes sometimes arising from tuberculate bases, up to 0.5 mm. long or even longer at the margins; cauline leaves sessile, ob- long-ovate to ovate, gradually becoming reduced in size upwards, indu- ment similar to that of basal leaves. Inflorescence usually branched, up to 20 mm. long, hairy, many flowered, bracteate, bracts ae pedicels slender, hairy, in flower up to 2 mm. long, in fruit to 6 mm. long, ee Calyx divided to the base, lobes oblong-ovate, 1.5 mm. long, 0.5 m broad, usually densely hairy with longer trichomes at the margins, ee in fruit slightly enlarged and sometimes reflexed. Corolla sky blue with a broad yellowish eye, 3-4 mm. long, rotate-campanulate, tube more or less equalling the calyx, lobes 2—2.5 mm. long, roundish, patent. Nut- lets 2—3.5 mm, long, 1.3-2.5 mm. broad, glabrous or minutely hairy on both surfaces, margins appendiculate, appendages up to 0.6 mm. long, 1970] KAZMI, BORAGINACEAE 389 0.2 mm. broad at base, weakly glochidiate, not at all to slightly confluent at base. DISTRIBUTION OF SPECIES: West Pakistan, Kashmir, Tibet (?). 3a. Var. spathulatum. Nutlets 2~2.5 mm. long, 1.3—1.5 mm. broad, sieaicgd hairy on both sur- faces, appendages usually not confluent at bas West Pakistan: Gitcrr AGENcy: Baltistan, Lamchau Nala, above Dras, 3700- 4000 m., Duthie 13786, 13767 (K). s Poonch, near Bedori, rocks, 3300 m., Rashid, Nasir & Stewart 23942 (BM), R. R. Stewart & Nasir 24038 (BM), 23024 (x): LaDAK: Tog, on rocky slopes, 5000 m., Koelz 2641 (cu); Ke La, 6000 m., in wet gravel, Koelz 2508 (GH, K, ay Miscellaneous: Tibet Occ., Herb, Ind. Or. Hook. f. & Thoms. Thomson 11 (Gu 3b. Var. thomsonii (C. B. Clarke) Kazmi, comb. nov. Omphalodes thomsoni C. B. Clarke in Hook. f. Fl. Brit. India 4: 155. 1883 Eritrichium thomsoni (C. B. Clarke) I. M. Johnston, ous Arnold Arb. 21: 3. 1940, non E. strictum var. thomsoni C. B. Clarke I. c. 164. Type: Western Tibet: Nubra, 13,000 ft., Thomson 8 a). Nutlets 3-3.5 mm. long, 2-2.5 mm. broad, usually glabrous, appendages more or less confluent at their bases. Kashmir: Lapak: Tsullak, by side of rocks on dry banks along streams, 5300 m., Koelz 2479 (GH); Rupshu, Tso Morari Korzok, on sandy slopes from side of "rocks, 5000 m., Koelz 2198 (cH, us); Korzok, 5000 m., Koelz 2206a (cu, US Eritrichium spathulatum may be distinguished from the closely related E. fruticulosum by its broader, ovate, sessile cauline leaves, covered with unequal trichomes and its larger nutlets; and from E. canum by its more herbaceous, decumbent habit, its broader cauline leaves, and the much shorter appendages on the margins of the nutlets. 4. E. patens Decne. in Jacquem. Voy. Inde Bot. 125. 1844; DC. Prodr. 10: 128. 1846. Type: D’Regui ad Tchini, Jul. 14. 1830, Jacquemont 1316 (P-holo- type, GH-isotype). Erect perennial herb to 35 cm, tall. Stems woody at the base, many simple or branched above, brownish, densely covered with white, stiff, subappressed trichomes. Basal leaves petiolate, petioles up to 20 mm. long, slender, hairy; lamina linear, up to 1-1.5 mm, broad, en- tire, acutish, covered densely on both surfaces with short, stiff, appressed white trichomes, sometimes arising from tiny white tubercles, nerves in- conspicuous; cauline leaves similar to the basal leaves, sessile, up to 390 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 15(-—20) mm. long, 1-1.5 mm. broad, sometimes with comparatively longer trichomes below and at the margins. Inflorescence terminal or sometimes axillary, simple to germinate, bracteate, many flowered; pedi- cels up to 6 mm. long, slender, pubescent, recurved in fruit. Calyx divided nearly to the base, lobes lanceolate, pubescent, acute, 1-1.5 mm. long, 0.2-0.3 mm. broad, slightly enlarged in fruit. Corolla blue, campanulate, tube equalling the calyx, lobes 2 mm. long, 1.5 mm. broad, ovate, patent. Nutlets 4, excluding appendages 1.5 mm. long, 1 mm. broad, roundish at the base, acutish above, outer surface convex or slightly raised longitu- dinally in the middle, usually tuberculate, sometimes minutely hairy, trichomes arising from minute tuberculate bases, inner surface glabrous, margined with distant, ee appendages, 0.2 mm. long, 0.1 mm. broad at base and not conflue DISTRIBUTION: West Pakistan, Kashmir, northwest India, Tibet. West Pakistan: Gitcir AcENCcy: Baltistan, Hushe to Brumi on the way from Hushe up to the Sospor glacier, Schlagintweit 6006 (BM Kashmir: Hab. Tibet Occ., 13,000-14,000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (pm); without locality, 1835, Royle (kK, mounted with Clarke’s No. 30370c of Aug. 10, 1876). Brand (Pflanzenr. IV. 252(Heft 97): 199. 1931) has placed E. patens in the synonymy of Lappula barbata var. cariensis but examination of the type showed it is an Eritrichium, Eritrichium patens closely resembles E. fruticulosum, from which it differs in its stout, densely pubescent, erect stems, very narrow, pubescent, basal leaves, and distinctly sessile cauline leaves. The nutlets of E. patens are comparatively smaller than those of E. fruticulosum. They are tuberculate on the outer face and glabrous on the inner side, while those of E. fruticulosum are not tubercu- late but are pubescent all around. Eritrichium patens also has much shorter appendages at the margins of nutlets compared to E. fruticulosum. 5. E. canum (Benth, in Royle) Kitamura, Act. Phytotax. Geobot. 19: 103. 1963; FI. Pl. vee Pakistan 121. 1964; Riedl in Rechinger, Fl. Tranica 48: 64. 1967. oe canum Benth. in Royle, Illustr. Bot. Himal. Mount. 1: 306. pier me strictum Decne. in Jacquem. Voy. Inde Bot. 125. 1844; DC. Prodr. 10: 128. 1846; C. B. Clarke in Hook. f. Fl. Brit. India 4: 164. 1883. Eritrichium strictum var. thomsoni C. B. Clarke I. c. aeons longifolium Decne. in Jacquem. /. c. 124. t. 129. DC. Prodr. 10: 846. Woodie longifolia Decne. in Jacquem. I. c. Eritrichium rupestre (Pall.) Bunge « genuinum Herder, Acta Horti Petrop. 1: 539. 1872. Eritrichium rupestre (Pall.) Bunge var. pectinatum (Pall.) Brand, Pflanzenr. IV. nig agit 97): 191. 1931, pro parte non E. pectinatum (Pall.) DC. Prodr. 10: : Ssapthe sericeum Aitch. Jour. Linn. Soc. Bot. 19: 178, 1882, non DC. I. c. 1970] KAZMI, BORAGINACEAE 391 Type: Lippa and Pangee in Kunawur, Kherang Pass, R. Inglis s.n. K). Icon.: Jacquem. Voy. Inde Bot. ¢. 126. 1844. Perennial, usually erect, sometimes decumbent, up to 40 cm. tall. Stems woody at base, thick, grayish or white, simple or branched in the upper part, densely covered with appressed white trichomes. Basal leaves pet- iolate, petioles up to 50 mm. long, hairy, more or less broad at the base; lamina narrowly lanceolate, acutish, gradually attenuated towards the petioles, to 80 mm, long and 6 mm. broad, covered densely on both surfaces with short, white, appressed trichomes: middle cauline leaves many, sessile, lanceolate, up to 30 mm. long and 3 mm. broad, obtuse or sometimes roundish at apex, gradually becoming reduced in size above; upper cauline leaves ovate, acute. Inflorescence simple, geminate or ter- nate, up to 15 cm. long, many flowered, bracteate, bracts linear, those of the upper flowers very short, pedicels in flower short, in fruit up to 10 mm. long, erect, usually straight, pubescent. Calyx divided nearly to the base, lobes ovate, 1 mm. long, 0.6 mm. broad, in fruit enlarged to nearly twice their size, densely pubescent, obtuse or sometimes roundish at apex. Corolla bright blue with a yellow or orange eye, rotate-campanu- late, tube slightly shorter than the calyx, lobes + 3 mm. long and broad. Nutlets 4, (excluding appendages) 2 mm. long, 1.5 mm. broad, outer face plane or slightly concave, covered loosely to densely with thin short trichomes, rarely glabrous, inner side glabrous to pubescent or sometimes slightly tuberculate, margins appendiculate, appendages irregular, up to 1 mm. long, confluent at base, glochidiate at apex. DIstriBuTIOoN: Afghanistan, West Pakistan, Kashmir, northwest India, and S.W. Tibet West Pakistan: CHITRAL STATE: Haute vallée de Yarkhun, Hautes gine 4000 m., Schmid 2283 (pm); upper Yarkhun valley, Pamir, Schmid 30 (BM). Gmerr Acency: Chhantir gah, Schmid 2163 (K). Hazara Dist.: Kaghan valley, between Batakundi and Burawai, Burtt & Reshan Din 922 (£). KURRAM AcEeNcy: Kurram valley, Kaiwas, Mt. Sikaran, 3300 m., Aitchison s.n. (GH); Kurram valley, Harsukh 15408 (x); Safed Keh, north of Kurram valley, Col- lett 110, 84 (kK) Shalozan, ca. 2700 m., Kazmi 204b (pes); Ziran, Kazmi s.n. : nmin R. R. Stewart 25366 (BM, MICH). Se N. W. F. P. (det. Jafri) sn. (£). Re- ported from Baluchistan, Lac Kashmir: Sonamarg, C. B. Clarke 30873 (K), 30873H (BM, type of E. strictum var. thomsoni), 3000 m., Rich 1119 (kK), R. R. Stewart aetigee (kK), 9291, 12422a (cH); Gagangu to Sonamarg, Young s.n. (BM); vicinity of Sonamarg. on Sind river, 50 road-miles east-northeast of Srinagar, Dickason 102 (MIcH); vicinity of Pahlgam on east of Lidder river, 27 road-miles north of Islamabad, Dickason 103, 104 (micH); vicinity of Baltal, on Sind river, 59 road-miles east- northeast of Srinagar, 3100 m., Dickason 105 (MicH); Pangtaran to Thani Young s.n, (BM); Pir Panjal, sip m., Drummond 14019 A upper Shieshinlg valley, 3700 m., Drummond 14186 (n): near Aliabad, m., Drummon 13924 (E); above Kainmal, DIcKISON PLATE IV Jour. ARNoLD Ars. VoL. 51 ; ACEAE VI KISON, DILLENI Dic 1970] HARTLEY, NOTES ON ZANTHOXYLUM 423 ADDITIONAL NOTES ON THE MALESIAN SPECIES OF ZANTHOXYLUM (RUTACEAE) Tuomas G. HARTLEY SINCE THE PUBLICATION of my revision of the Malesian species of Zanthoxylum (Jour. Arnold Arb. 47: 171-221. 1966) I have examined a considerable number of additional Malesian collections of that genus. As a result, the descriptions of some of the species should be amplified and several range extensions may be reported. Also, it has come to my attention that one of the names used, Z. limonella (Dennst.) Alston, is invalid, and, accordingly, the necessary correction is made. The species are numbered here in the same order as in the revision. Only those collections pertinent to the comments are cited in full. How- ever, a complete list of all additional numbered collections examined is included as a supplement to the original Index to Exsiccatae Grateful acknowledgment is made to the directors and curators of the Herbarium Bogoriense, Djalan Raya, Bogor, Indonesia (Bo) and the Rijksherbarium, Leiden (1), for the loan of herbarium specimens. Ben- jamin C. Stone, of the University of Malaya, Kuala Lumpur, is also thanked for sending a duplicate of his recent collection of Zanthoxylum acanthopodium from Malaya. 2. Zanthoxylum nitidum (Roxb.) DC. The range of this species may be extended to include the Philippine island of Mindanao. Philippines. Minpanao: Lake Lanao, Clemens, May 1907 2 (Bo). 5. Zanthoxylum avicennae (Lam.) DC. The range of this species may be extended to include the Celebes. Also, it is now confirmed that the collection cited for the provisional name Fagara anisata Back. ex Bakh. f. (Backer 11431) is Zanthoxylum avicennae Java. East Java. Kediri, G. Wilis, Backer 11431 2 sheets,é and ? (so). Celebes. Res. Manado, Paloe, NIFS bb 3 2 (Bo), Pesik NIFS bb 9 (xo). 6. Zanthoxylum megistophyllum (Burtt) Hartley With the addition of the recent Bougainville collections of R. Schodde and L. Craven the description of this species may be amplified somewhat. Also, the range may be extended slightly to include Santa Ysabel Island in the Solomons. Small to medium trees to 17 m. Inflorescences to 44 cm. long. Anthers 424 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 of staminate flowers 0.5-1 mm. long. In primary and secondary rain forests at altitudes up to 690 meters. Solomon Islands. BOUGAINVILLE: Near Koniguru No. 1, ca. 10 miles N of Buin, Craven & Schodde 107 8 (tL); Lake Laloru Crater, ca. 14 miles N of Buin, Craven & Schodde 260 & (1); vicinity of Aku, ca. 9 miles W of Buin, Schodde & Craven 4110 & (1). SANTA YsaBeL: Moloforu Bay, Beer’s Col- lectors BSIP 7376 2 (L). 7. Zanthoxylum forbesii Hartley The original description of this species was based on a single fruiting collection from Sumatra. With the addition of a staminate flowering specimen from Sumatra and a fruiting specimen from Java, the descrip- tion and range may be amplified as follows: Small (4 m.) to large trees; dioecious. Leaves paripinnate or imparipin- nate, glabrous, 40-70 cm. long; petiolules of lateral leaflets 0.6—-1 cm. long; leaflets with 12-19 veins on each side of the midrib, 15-27 cm. long, 5-10 cm. wide, obtuse to acutely oblique at base, short to rather long acuminate at apex, the acumen to 1.7 cm. long. Inflorescence terminal, paniculate, 24 cm. long and about half as broad, the axes and branches glabrous. Staminate flowers (in bud) 3 mm. long; pedicels 1 mm. long; sepals 4, green, broadly triangular, 1 mm. long, connate at about half their length; petals 4, white, elliptic, 2.5 mm. long; stamens 4, filaments 1 mm. long, anthers 2 mm. long; disc pulvinate, about 0.5 mm. high; rudimentary carpels 2, about 0.5 mm. high. tra. Padangsidimpoean, Kostermans 22042 8 (1). Java. West JAVA. Tokoka, Koorders 6983 2 (Bo). 9. Zanthoxylum retroflexum Hartley A second collection of this species has leaves ascending rather than sharply reflexed as in the type collection. This necessitates deleting each of the first statements in the 7th couplet of the key (p. 176) and altering the description to read, with regard to the leaves: “...ascendentibus vel acute reflexis,...” Field studies are desirable to determine what factors are involved with the reflex movement of the leaves in this species. Sumatra. Palembang District, Tjabang Forest Res., Kostermans § 42 2 (L). 10. Zanthoxylum rhetsa (Roxb.) DC. The name used in the revision, Zanthoxylum limonella (Dennst.) Alston, is invalid since the basionym, Tipalia limonella Dennst., represents a com- bined new generic name and specific epithet listed with only a reference to a plate and description in Hortus Indicus Malabaricus. Rickett and Stafleu (Taxon 10: 80. 1961) have pointed out that such names in Denn- stedt’s publication are nomina nuda because new generic and specific combinations cannot be validated by reference to a previous publication. Details of Dennstedt’s Schlussel zum Hortus Indicus Malabaricus with 1970] HARTLEY, NOTES ON ZANTHOXYLUM 425 lists of the valid and invalid names were given recently in a paper by H. Manitz (Taxon 17: 496-501. 1968). The choice of the name Zanthoxylum rhetsa over Z. budrunga (Roxb.) DC., the basionyms of which were published on the same date, follows Hooker f., Fl. Brit. India 1: 495. 1875. The range of this species may be extended to Nei Madoera Island, just north of Java, and the islands of Bali and c Java. Madoera Island, N. van Soemenep, Backer 20669 (Bo). Lesser Sunda Islands. Batt: Becking 49 (Bo), 70 (Bo). Trmor: N Midden Timor, N/FS bd 7223 (BO). 14. Zanthoxylum pluviatile Hartley A recent collection by J. S. Womersley from near the type locality is the first carpellate flowering material I have seen of this species. The description of the flowers follows. Also, the range may be extended to include New Britain, apparently the first Zanthoxylum ever collected from that large island, and to another of the Solomon Islands, Malaita Island. Carpellate flowers 4-5 mm. long; pedicels, sepals and petals as in staminate flowers; rudimentary stamens 0.2 mm. long; disc subpulvinate, about as thick as the base of the ovary, 0.75 mm. high; gynoecium 1- carpellate, 2 mm. high, style excentric, stigma flattened. Territory of New Guinea. Morose District: Markham River logging area, Ades NGF 19198 2 (L). New Britain. Kandrian Subdistrict, Pulie River, y & Frodin NGF 27243 & (1). Solomon Islands. Malaita Island, Dala, See BSIP 4477 (1). 16. Zanthoxylum integrifoliolum (Merr.) Merr. The following collection documents the report of this species from Taiwan: Taiwan. Lanyu Island (Botel Tobago), Chang 3136 2 (1). 17. Zanthoxylum ovalifolium Wight The range of this species may be extended to include Borneo. Indonesian Borneo (Kalimantan). Martapoera, Daoen Limauan NIFS bb 568 2 (BO). 18. Zanthoxylum acanthopodium DC. This species was recently reported from Malaya by B. C. Stone (Malay- an Nat. Jour. 23: 31, 32. 1 fig. 1969). This new station considerably lessens the gap in the previously cited distribution, between Thailand and umatra Malay 1 Peninsula. PAHANG. Cameron Highlands near Brinchang Village, Stone 5700 & (a). 426 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 SUPPLEMENT TO THE INDEX TO EXSICCATAE The numbers in age refer to the a species be iho following list: 1. Zanthoxylum scandens; 6. Zanthoxylum meriophilm: flexum; 10. rsa eat rhets Spersipunctatum; 16. 18. Zanthoxylum pasion Backer 5753 (1); 11431 (5); 20669 (10); 22526, 25365 (1); 27132, 29457 (10); 32842, 34281, 34282 (2) Bakhuizen oy (2) Becking 49, 70 (10) Beer’s Collectors BSIP 7376 (6) Beumée 1930 Borssum agree 324, 563 (10) Bruggeman 448 Carr 11583 (14); ena 11953, 14730 (17) 17 Chang 3136 (16) Chew & Corner RSNB 4895 (5) Docters van Leeuwen-Reynvaan 1003 Ms Evans 55 (14) Flenley ANU 2329 (15) Hallier 361 (1); 361a (2) Henty & Frodin NGF 27243 (14) Hildebrand E 397 (1); NIFS Ja 4676 17) ensen 8 (5 Kjellberg 1345 (2) Koorders 6982 (10); 6983 (7); 6984, 6985, 6986, 6987, 6988, 6989, 6990, 6991, 6992, 6993, 6994, 6996, 7036, 7038, 11583, 11584, 11585, 11586, 13123, 13683, 18774, 20321, pore 25575 (10); 25093, 26257 (1 arsed, F086: C1Oy> F195 7, 31501 (1); 33408 (2); 33743 (1); 34378 ARNOLD ARBORETUM HARVARD UNIVERSITY Zanthoxylum niti 2 pe ate paren . Zanthoxylum pluviatile ; nthosylam integrifoliolum ; xylum avicennae ; 9. grew oxylum retro- 15. Zanthoxylum con- 17. Zanthoxylum ovalifolium ; (2); 36500, 36766 (10); 36908 (2); & Koorders-Schumacher 43376, 47694 (2) Kostermans S 42 (9); 22042 (7) Lam 245 (17) Lorzing 3712 (2); 6221, 7841, 9482 18 Koorders 18076 (17); Netherlands Indies Forest Service (NIFS), the following by anony- mous collectors: bb 3 (5); Ja 2251, bb 7223 (10) Ouwehand 304 (18) Pesik NIFS bb 9 (5) Pigl 246 Rensh 303 (5); = > Royen NGF 1637 Sayers & Millar : cr 19851 (15) Scheffer HB 14634 (1) Schodde 2967 am Schodde & Craven 4110 (6) Slooten 279 (1) Smith, J. J., & Rant 253 (1) Snyder BSIP 4477 (14) Steenis 4214, 4917, 5181, 6861, 12261 1 Stone 5700 (18) Teysmann HB 14625 (10) Thorenaar 336 (10 Winckel 2 (2) Wirawan 417 (10) Womersley NGF 19198 (14); NGF I5 Zwickey 1092 (16) 1970 | ELIAS, NOTES ON GALIPEA (RUTACEAE) 427 NOTES ON THE GENUS GALIPEA (RUTACEAE) IN CENTRAL AMERICA ? Tuomas S. ELIAs IN THE SMALL GENUS Galipea, consisting of eight to ten species with a center of distribution in northern South America, are two species which reach eastern Panama. Aublet (1775) first described Galipea with a single species, G. trifoliata, from French Guiana. Additional species of Galipea were soon described, mainly from Brazil. In 1912, Pittier collected a single fruiting collection of G. jasminiflora (St. Hil.) Engler in eastern Panama (Darien). This locality represents a disjunction from southern Brazil and Guyana. The second species, G. panamensis, described here for the first time, is endemic to Panama, although it is expected to occur in bordering areas of Colombia. Two other species described from Central America, G. guatemalensis Standl. & Steyerm. and G. nicaraguensis Standl. & Williams do not belong in Galipea, but should properly be placed in Angostura (Cusparia). Nomenclatural problems associated with the valid generic name for Cusparia have been treated by Elias (1970), and the taxonomy of the Central American species of Angostura will be the subject of a subsequent note The genus Galipea belongs to the tribe Cusparieae, which comprises approximately 22 to 24 genera confined mainly to tropical South America. The tribe is characterized by palmately compound 1-5-foliolate leaves, rarely simple; flowers regular to zygomorphic; corolla often fused and tubular; stamens usually 5, staminodia often present; carpels 4 to 5, ovules 2, collateral with one slightly superposed on the other; embryo curved to uncinate with little or no albumen, radicle retracted between the cotyledons. The relationships between the two closely related genera Angostura and Galipea have often been obscured by the indiscriminate practice of de- scribing new species when generic limits are not clearly understood. The taxonomy of the two genera as well as of most of the tribe Cusparieae is poorly known, due largely to the paucity of collections. Engler (1874) helped to clarify the limits of these two genera by transferring eight species, those with individual mericarps rather than capsules, from Galipea to Cusparia (= Angostura). He observed that the gynoecium in species of Angostura was composed of five closely packed but free carpels. In species of Galipea, however, the five carpels have undergone partial fusion and form a 5-lobed ovary. Consequently, in fruiting condition the individual *I am grateful to Dr. Robert L. Dressler for providing adequate material of the new species, including pre rar flowers for ction. Dr. Thomas G. Hartley has kindly read the manuscri pt and offered several useful comments. Dr. Lily M. Perry has graciously proofread jah corrected the Latin description. 428 JOURNAL OF THE ARNOLD ARBORETUM [vor. 51 carpels of Angostura develop into separate mericarps, while in Galipea a 5- parted capsule is formed. Although the structure of the fruit, apocarpous versus syncarpous, has been very useful in distinguishing the two genera, it may not prove to be absolutely reliable. Examples of rutaceous genera which may have capsular as well as apocarpous or at least subapocarpous fruits include Esenbeckia, Evodia, Boninia, Melicope, Tetractomia, Pelea, and M yrtopsis (Hartley, 1969). Additional characters that can be used to separate the two genera may be found in the flowers. In Galipea the petals are united into a distinct, often curved tube, while in Angostura the petals are free or fused only at the base. The stamens of Galipea are inserted near the apex of the corolla tube as opposed to stamens inserted at the base of the disc in Angostura. Also the anthers of Galipea usually have a small basal ap- pendage while those of Angostura lack an appendage. In an attempt to find other characters which would be useful at the generic level, pollen of the two genera was examined and found to be of value. Although ma- terial was not available for all species the grains of the species of Galipea sampled were large, suboblate to subspherical, coarsely reticulate, and 4-5-porate. In contrast are the medium large, prolate, reticulate, and 3- colporate pollen grains found in species of Angostura. KEY TO THE CENTRAL AMERICAN SPECIES OF GALIPEA & . Leaves unifoliolate; corolla tube stout, 1.1-1.4 cm. long, the lobes narrowly obovate, acute at the ACE 23-16 COM tone ss G. panamensis. Leaves trifoliolate; corolla tube slender, 9-11 mm. long, the lobes oblong to narrowly obovate, ‘obtuse at the apex, 4-5 mm. long. ...... G. jasminiflora. S Galipea panamensis Elias, sp. nov. A Galipea simplicifolia (Nees & Mart.) Engler inflorescentiis paniculatis, 0.5-2 c ere corolla tubulosa gracilis, lobis 5 mm. longis differt. ne que ca. 7 m. altae; rami teretes, leviter sulcati, murini vel bruneoli, seb lenticellati, lenticellae + elevatae. Folia 1- foliolata, petio- lata, petioli superne a sha expansi et indurati ad basin, articulati prope basin laminae, 9-16 m. longi: lamina coriacea, elliptica vel anguste elliptica, basi eae. apice acuminata, pellucido-punctata, glabra, 11- 21 cm. longa, 3—9.2 cm. lata, nervis lateralibus utroque 5—11-jugis subtus prominentibus. Pi naira terminales et axillares; corymbi congesti, ca. 12~—22-flori; pedunculi erecti vel semi-erecti, conspicue lenticellati, validi, 1.2-3.5 cm. longi. Flores albi; calyx campanulatus, pellucido- punctatus, puberulus, 3-4 mm. longus, lobis transverse ovatibus, apice acutis minute ciliatis, 0.5-1 mm. longis; corolla puberula, conspicue pel- lucido-punctata, 2.3-3 cm. longa, tubo cylindrico, arcuato in alabastro, 1.1—1.4 cm. longo, lobis in aestivatione ee anguste obovatis, apice acutis, dilatatis vel leviter recurvatis, 1.3-1.6 cm. longis, ad 3.5 mm. latis; stamina 6—7, inclusa, apicem tubi versus ao Alas enta brevissima, haud 1 mm. longa, antherae fertiles binae, basifixae, longitudinaliter dehiscentae, 1970] ELIAS, NOTES ON GALIPEA (RUTACEAE) 429 Figure 1. Galipea agecoosinona a, branchlet wena inflorescence, Lb; atu ¥%, b, er nia or x 1%; e bud showing superposed ovules, longitudinal section, X 3; d, cross section of ovary, greatly peti aor e, mature capsule a be sion of dehiscence, x 114; f, endocarp with seed, < 24. a-f from Oy eid 3682 (holotype a). 4-6 mm. longae; staminodia libera, complanata, 6-8 mm. longa; stylus cylindricus, 10-12 mm. longus, stigmata 5, oblonga, 1-2 mm. longa; ovarium 5-carpellatum; loculi biovulati, ovulis plus minusve superimpositis, placentatio axilis. Capsulae 5-loculares, subglobosae, 1-1.5 cm. longae, 430 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 cocci basi apiceque connati, ventraliter ac dorsaliter carinati, dehiscentes a basi fere usque ad medium; endocarpium cartilagineum bivalvatum, 1- spermum, basi integrum, alae apice obtusae; semen + orbiculatum vel obovoideum, lateraliter parum compressum, pubescens. Panama. CoLon: Santa Rita ridge lumber road, ae of Colon, Correa ‘2 Dressler 766 {fis., frts.} (110), Dressler 3659 [fls., frts.] (A, Mo). PANAM Cerro Jefe, in coffee plantation, ca. 700 m. altitude, Dressler 3682 | fls., fats] (holotype A, isotype MO). While most species of Galipea have trifoliolate leaves, G. panamensis differs in having them unifoliolate. Also, its corymbose inflorescence the inflorescence is narrowly paniculate. Both G. panamensis and G. jasminiflora, known only from eastern Panama, presently represent the northernmost occurrence of the genus LITERATURE CITED AuBLeT, M. F. Pl. Guiane Francaise 2: 662. t. 269. Exias, T. S. The correct name for the genus Cleiur: (Rutaceae). Taxon ENGLER, A. Rutaceae. Jn: Martius, Flora Bras. 12(2): 78-195. pls. 14-28. 1874. [Galipea 95-99, pls. 17-19. HartLey, T. G. A revision of the genus Flindersia (Rutaceae). Jour. Arnold Arb. 50: 481-526. 1969. ARNOLD ARBORETUM HARVARD UNIVERSITY i VWs 3 AN WZ A he Lar Hid ee Sa SA il) Zs \\ Rahs = AY YA Nf lee Missoun Fors y-nabL IOCT 2 61970 vanced meget | Volume 51 Number 4 JOURNAL OF THE ARNOLD ARBORETUM HARVARD UNIVERSITY JOURNAL OF THE ARNOLD ARBORETUM VoL. 51 OcTOBER 1970 NUMBER 4 THE GENERA OF MOLLUGINACEAE AND AIZOACEAE IN THE SOUTHEASTERN UNITED STATES! A. LINN BOGLE MOLLUGINACEAE Hutchinson, Fam. FI. Pl. 1: 128. 1926, nom. cons. (CARPET-WEED FAMILY) Scarcely or non-succulent, annual [or perennial], glabrous or stellately pubescent herbs [subshrubs, or shrubs], with mostly prostrate to erect, often much-branched stems from a short, narrow taproot. Leaves simple, narrow to broad, glabrous or pubescent, alternate, opposite, or falsely whorled, exstipulate [or stipulate]. Inflorescence basically cymose, axil- lary or terminal, diffuse or congested. Flowers sessile or pedicellate, regu- lar, perfect [or imperfect]. Perianth uniseriate, inconspicuous. Sepals (tepals) usually 5 [4], free [or slightly united basally], imbricate or quin- cuncial, persistent. Petals absent [or small, staminodial in origin]. Sta- mens 3—10[~—20], alternating with the carpels when 3 or with the sepals when 5, hypogynous [or perigynous]; anthers 2-loculate, dehiscing by longitudinal slits; filaments free, united basally [or fascicled when stamens numerous]; pollen 3-colpate, or 3—5-zonicolporoidate. Gynoecium of 3-5 ‘Prepared for a generic flora of the southeastern United States, a project of the Arnold Arboretum and the Gray Herbarium of Harvard University made possible in the first paper i in the series (Jour. Arnold Arb. 39: 296-346. 1958) and continued through those in volumes 40-50 (1959-1969). The area covered includes North and lorida, Tennessee, Alabama, Mississippi, Arkansas, and mentary info a8 in brackets. References which the author has not seen are marked by an risk. I wish to nae Dr. Jack Fisher for rt specimens of Sesuvium Portulacastrum, and Dr. John Thieret for specimens of Trianthema Portulacastrum. due to Miss Rachel A. Wheeler, who ne the illustrations of Mollugo, to Miss Virginia Savage, who drew the remaining vege enceara nes o Mrs. Nancy Dunkly, who checked the bibliography and typed the manuscri I am particularly grateful to Dr. Wood for his ae during the course of this work, and for his careful editing of the manuscript. an John Thieret very kindly read the iecign and made several helpful commen 432 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 free or united carpels, when united the ovary superior, 3—5-loculate; styles or sessile stigmas as many as the carpels; ovules one per carpel and basal, or several to many on axile placentae, anatropous to anacampylotropous. Fruit an achene or 3—5-valved loculicidal capsule. Seeds reniform to roundish; seed-coat smooth, punctate, tuberculate, or longitudinally ridged; saccate caruncle or hilar peg present or absent. Embryo periph- eral, curved; cotyledons fleshy; endosperm starchy. Type Genus: Mol- lugo L. A small family of 14 more or less weedy genera containing about 95 species distributed primarily in the tropics and subtropics of both hemi- spheres. The family is best developed in southern Africa, where twelve genera occur. Of these, seven are confined to Africa (Adenogramma Reichenb., Coelanthum E. Mey. ex Fenzl, Hypertelis E. Mey. ex Fenzl, Pharnaceum L., Polpoda Presi, Psammotropha Eckl. & Zeyh., Suessengu- thiella Friedr.), three extend to southwestern Asia (Corbichonia Scop., Limeum L.) or southeastern Asia (Gisekia L.), and two are pantropical weeds (Glinus L., Mollugo L.). Glischrothamnus Pilg. is known only from Brazil, and Macarthuria Hueg. is restricted to Australia. Mollugo and Gisekia are each represented in our area by a single species, Glinus by two. Gisekia has only recently been found in Florida, and is thus not included in the older floristic works. The genera included here have appeared in the literature under several family names. Bentham & Hooker considered them a tribe, Mollugineae, in their order Ficoideae (family Ficoidaceae). Baillon recognized a se- ries Mollugineae in his family Portulacaceae. In the two editions of Eng- ler & Prantl’s Die Natiirlichen Pflanzenfamilien they appear in the Aizoaceae, and in both Small’s Manual of the Southeastern Flora and the North American Flora under Tetragoniaceae (= Aizoaceae). However, taxonomic opinion (Hutchinson, 1926, 1959; Friedrich, 1955; Takhtajan, 1959; Adamson, 1960; Eckardt, 1964; Cronquist, 1968) is now heavily in favor of segregating this group from the Aizoaceae. In this case the family name Molluginaceae Hutchinson is conserved against Glinaceae Link. Of the four tribes now recognized in the family (Eckardt), only the Mollugineae (including Mollugo L., and Glinus L., with carpels united and several to many seeds per locule) and Gisekieae (containing only Gisekia, with apocarpous gynoecium and solitary, basal ovules) reach our area, while the tribes Corbichonieae (formerly Orygieae) and Limeeae are confined to the Old World. From the related Aizoaceae the Molluginaceae differ in having a scarcely (if at all) succulent habit; in possessing red pigments of the anthocyanin rather than the betacyanin type; in lacking anomalous secondary growth in the stems and roots; in having floral parts few, usually free (except in Coelanthium), hypogynous, and cyclically arranged; and in lacking a calyx tube. The family is considered to be closely allied with the Phyto- laccaceae through the apocarpous gynoecium of Gisekia and the solitary, 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 433 basal ovules of the tribe Limeeae, but differs in having cymose-dichasial, rather than racemose or spicate, inflorescences. There are also relation- ships with the Portulacaceae and Caryophyllaceae. Available chromosome counts indicate that x — 9 for the Molluginaceae, with several levels of polyploidy represented in the common genera and species. From a study of the floral morphology and vascular anatomy of seven species in four genera (Gisekia, Glinus, Mollugo, Semonvillea | Limeum)) Sharma (1963a) concluded that progressive condensation of the floral axis has resulted in a change from acyclic to cyclic origin of the tepals and tepal traces and from independent origin of the tepal and stamen traces to various stages of cohesion and adnation. In the androecium he suggests reduction from three pentamerous cycles to one (the innermost) ; in the gynoecium reduction from three bundles in each carpel to two in Limeum pterocarpum (Gay) Heimerl (as Semonvillea pterocarpa Gay) through the loss of the dorsal bundle; and in the placentation reduction from axile to basal (Gisekia) or partially parietal. The embryology of a few genera (Gisekia, Glinus, Limeum, Mollugo, and Corbichonia) has been studied, although not intensively. Ovules vary from anatropous to anacampylotropous. The inner of two integu- ments forms the micropyle. An aril may be present around the funiculus (Glinus, Corbichonia, Macarthuria). A normal Polygonum-type embryo sac develops within the crassinucellar ovule, followed by either a caryo- phyllad or solanad type of embryogeny. Anthers are tetrasporangiate. The pollen grains are 3—5-colpate or zonicolporoidate (i.e., distributed around the equator of the grain; Sharma) and are usually 3-celled when shed. The family is of no great significance either economically or horticul- turally, but individual species may be utilized regionally as vegetables or in folk medicine or serve as grazing plants for livestock. REFERENCES: Apamson, R. S. The phytogeography of Molluginaceae with reference to Southern Africa. Jour. S. Afr. Bot. 26: 17-35. 1960. [Includes maps. | BAItton, H. Portulacées. Hist. Pl. 9: 54-80. 1886. [Series Mollugineae, 61-64, 65, 76-80; Mollugo, 76, 77, figs. 92-98.] : Beck, E., H. Merxmitter, & H. WacNner. Kurze Mitteilung iiber die Art der Anthocyane bei Plumbaginaceen, Alsinoideen und Molluginaceen. Planta 58: 220-224. 1962. [Anthocyanins in two species of Hypertelis.] BENTHAM, G., & J. D. Hooker. Ficoideae. Gen. Pl. 1: 851-859. 1867. [Tribe Mollugineae, 850-859; Mollugo (incl. Glinus), 357; Gisekia, 859. | CANDOLLE, A. P. pe. Ficoideae. Prodr. 3: 415-456. 1828. [Glinus, 455.]; Phy- tolaccaceae (by Mogutn-Tanpon). /bid. 13: 2-40. 1849. [Tribe Giesekieae, 26-35; Giesekia, 26-28, 459.]; Caryophylleae. Jbid. 1: 351-422. 1824. Mollugo, 391-393. eles The pote and classification of flowering plants. x + 396 pp. Boston. 1968, [Molluginaceae, 182. Detailed discussion of phylogenetic relationships among families of subclass Caryophyllidae. ] 434 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Davis, G. L. Systematic embryology of the angiosperms. ix + 528 pp. New York. 1966. [Molluginaceae, 177, 178. DuPont, S. Sur la morphologie des plantules de Mesembryanthémacées et d’Aizoacées. Bull. Soc. Hist. Nat. Toulouse 95: 356-360. 1960. [Seedling morphology; Mollugo, Glinus. | Ecxarpt, T. Molluginaceae. H. Metcuror. Engler’s Syllabus der Pflanzen- familien, ed. 12. 2: 86, 8 EICHLER, A. W. = ccs eae Pt. 2. Leipzig. 1878 [Aizoaceae, 119-124; Mollugo, Glinus.| ENDLICHER, S. Portulaceae. Gen. Pl. 946-955. 1840. [Tribe Mollugineae, 952- 954; Mollugo, Glinus.| Fenzi, E. Monographie der Mollugineen und Steudelieen. Ann. Wiener Mus. Naturgesch. 1: 337-384. 1836. [Portulacaceae, tribe Talineae, sectio Mollugineae, 352.]; Supplement. Verbesserungen und Zusatze zur ersten Abtheilung der Monographie. /bid. 2: 279-307. 1839. [Portulacaceae, tribe Mollugineae, 299.]; Monographie der Mollugineen (Zweiter artikel). Ibid. 2: 245-279. 1839. FriepricH, H. C. Beitrage zur Kenntnis einiger Familien der Centrospermae. Mitt. Bot. Staatssamml. Miinchen 12: 56-66. 1955. [Molluginaceae, 58. | HermerL, A. Phytolaccaceae. Nat. Pflanzenfam. III. 1b: 1-14. 1889. [Gzsekzia, LO, 17. Hutcuinson, J. The families of flowering plants. I. Dicotyledons. London 1926. [ Molluginaceae, 128, 129.] Ed. 2, Oxford. 1959. [Molluginaceae, 428, 429. ae C. Aizoaceae. In: C. E. Hupsarp & E. Mitne-RepHEap, eds., FI. rop. E. Afr. 37 pp. London. 1961. [Gisekia, 2; Glinus, 11; Mollugo, 16; well illustrated. | Miter, K. Beitrage zur Systematik der Aizoaceen. Bot. Jahrb. 42(Beibl. 97): 54-94. 1908. [Including poe pag Mollugo. Pax, F., & K. HorrMann. Aizoaceae. Nat. he presse ed. 2. 16c: 179-233. 1934, {Including Gisekia, Choke: Moll RowepsacH, P. Molluginaceae. Jn: KF. von Martius, Fl. Brasil. 14(2): 229-244, 315. pl. 55.1872. [Mollugo, aE SHarMaA, H. P. Studies in the order Centrospermales. II. Vascular anatomy of the flower of certain species of Molluginaceae. Jour. Indian Bot. Soc. 42: 19-32. 1963; IV. Pollen morphology of some species of families Ficoi- daceae, Molluginaceae, Nyctaginaceae, and Portulacaceae. Ibid. 637-645. TaxuHtTajANn, A. Die Evolution der Angiospermen. viii + 344 pp. Jena. 1959. Soa rns Wight; Gisekia, 196. THIERET, J. W. Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2: 352-360. 1966. [Aizoaceae, including Mollugo verticillata, M. Cerviana, 355, fig. 2; Glinus lotoides, 357, fig. 3; excellent illustrations and descriptions of the seeds. Urnor, J. C. T. Dictionary of Economic Plants. ed. 2. 591 pp. Lehre. 1968. | Mollugo, 348; Gisekia, 244.] Witson, P. Tetragoniaceae. N. Am. Fl. 21(4): 267-277. 1932. [Mollugo Glinus. | KEY TO THE GENERA OF MOLLUGINACEAE IN THE SOUTHEASTERN UNITED STATES General characters: low herbs with simple, alternate, opposite or falsely whorled, often distinctly unequal, exstipulate leaves; stems mostly prostrate 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 435 (to ascending) ; inflorescence an axillary or terminal, diffuse or congested apne; ; flowers incomplete, small and inconspicuous, regular, hypogynous; sepals 5 petals none; stamens mostly 3-5, alternate with the sepals (or with the carpal when 3 ), free or slightly united basally in a shallow ring; carpels 3-5, free or united in a several-loculate superior ovary; ovules 1 to many ; ae ae or axile; fruit an achene or many-seeded ‘oculicidal capsule; seeds + reniform, the seed-coat smooth, punctate, tuberculate, or longitudinally ridged. A. Plants glabrous; seeds lacking caruncle and filiform appenda B. Carpels 5, free; ovule asal; fruit a thin-walled achene; petiole aks often decurrent on the stem, the ribs often bearing scale- like 1 TSA ar eo SUT RAR hors om I aia hie hp LCL COR 1. Gisekia. ne 3(-5), united; ovary 3(—5)-loculate; ovules numerous in each locule on axile placentae; fruit a 3(-5) -valved loculicidal wees petiole TATE: Tee CCE nh ey Se eee eee ollugo. Plants thinly to densely stellate-pubescent; seeds numerous, ba seats a long filiform appendage and a whitish, saccate caruncle around the p sistent funiculus; sepals mucronate ees ovary compound, 3(—5)- joculate; stigmas 3(-5), sessile OF SORES GA. oc. ie ct a tere doa tar 3 > Tribe GISEKIEAE Endl. 1. Gisekia Linnaeus, Mant. Pl. Alt. 554, 562. 1771. Scarcely succulent annual herbs with prostrate to ascending branches radiating from a central axis. Stems glabrous, reddish, subequal, with opposite branching; nodes swollen. Leaves exstipulate, opposite to fasci- culate, the fascicles separated by distinct internodes; blades narrowly lanceolate to linear-oblong, unequal, entire, the margins sometimes revo- lute at maturity; midvein prominent below, depressed above, secondary veins inconspicuous; margins of the petiole or leaf-base decurrent, form- ing ribs on the terete stems. Inflorescence an axillary or terminal several- flowered compound dichasium [varying to umbellate]. Flowers small, per- fect [or imperfect], about 2 mm. long. Sepals 5, free, boat-shaped, trans- lucent margins whitish below, pinkish above. Stamens 5{—15], alternate with the sepals; filaments basally dilated; anther basifixed, 2-loculate at anthesis, dehiscing by lateral longitudinal slits, white. Gynoecium apo- Carpous, carpels 5 [3 or 10], laterally compressed, wall translucent, cov- ered with soft, whitish papillae, at maturity the carpel margins weakly erose-winged; style short, inserted on inner side of summit of carpel, erect in bud, curving outward at maturity; stigma + terminal; ovule soli- tary in each carpel, basal. Fruit a thin-walled achene. Seed nearly cir- cular in outline, laterally compressed + lenticular in section; seed coat black, lustrous, finely punctate at maturity. Embryo nearly annular, peripheral; cotyledons oblong, fleshy; endosperm starchy. TYPE SPECIES: G. pharnacioides L. (Named for Paul Dietrich Giseke, 1741-1796, pro- fessor at Hamburg, Germany.) A small genus of perhaps five species distributed primarily in the warm- er, drier parts of tropical and subtropical Africa and Asia. The most 436 JOURNAL OF THE ARNOLD ARBORETUM (von. 54 widespread of these, G. pharnacioides, 2n = 36, is the only species oc- curring in our area, apparently as a very recent adventive. Godfrey de- scribed and illustrated plants of this species from orange groves in Orange and Polk counties in central peninsular Florida, where they flourish, some- times in large numbers, in loose cultivated sands. The species may be spreading from this area. This habitat is similar to that of the Indian representatives of this species (which the plants of our area closely ap- proach) as described in detail by Joshi & Kambhoj, who rarely found the species in other than sandy habitats, with the plants appearing after the first few rain showers and the life-cycle being completed in five to eight weeks. This species, with its leaves with large, thick-walled epidermal cells bearing thick cuticle and wax layers and the palisade cell layers internal, is well adapted to xerophytic conditions. The stems, leaves, and all floral organs are conspicuously white-flecked by bundles of raphides produced in the parenchymatous tissues. There is disagreement concerning the systematic position of the genus. It has been referred to the Phytolaccaceae (DeCandolle, Baillon, Heimer) but was excluded from that family by Walter; to the Portulacaceae (Gagnepain), a position that has found no support from other workers; to the Ficoidaceae or Aizoaceae (Bentham & Hooker, Pax & Hoffmann) ; and, most recently, to the Molluginaceae (Hutchinson, Friedrich, Eckardt, Takhtajan, Cronquist). In the Aizoaceae, Molluginaceae, and Phytolac- caceae Gisekia comprises a monotypic tribe, Gisekieae Endl. With the Phytolaccaceae Gisekia shares free carpels with solitary, basal ovules. Among the genera of Molluginaceae Gisekia is perhaps closest (but only distantly related) to Limeum. In its anacampylotropous ovules with few-layered nucellar epidermal caps Gisekia compares favorably with other Molluginaceae, rather than Phytolaccaceae. It differs from those species of Aizoaceae so far investigated in lacking radially stretched epi- dermal cells in the nucellar caps and in the absence of starch grains in the embryo sac. The pollen is tricolpate and three nucleate when shed (Shar- ma, 1963b; Joshi & Rao). Secondary growth is normal, rather than anomalous. The floral vascular anatomy of G. pharnacioides has been described by Sharma (1963a). The nature of the red pigments, whether antho- or betacyanous, has not yet been determined. Orthographic variations of the generic name include Giesekia, Gieseckia, Giseckia, Gisechia, and Giesechia, while the specific name may appear as “pharnaceoides”, rather than as the original “pharnacioides” (Adam- son). The genus is of little economic significance. Gisekia pharnacioides is an important grazing plant for cows, goats, sheep, and camels in India, and is used in folk or Ayurvedic medicine against a broad range of complaints, including scabies, rhinitis, bronchitis, leprosy, leucoderma, urinary diseases, and as a powerful antihelminthic in cases of Taenia (tapeworm) infection. The leaves are used in the preparation of dal (the split seeds of various le- gumes), and as a potherb in times of famine (Uphof). 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 437 REFERENCES: Under family references see BAILLON, BENTHAM & HooKER, CANDOLLE, CRONQUIST, ECKARDT, FRIE RICH, HEIMERL, HUTCHINSON, JEFFREY, Pax & HOFFMANN, SHARMA, and Tee. ApaMson, R. S. The South African species of Aizoaceae. X. Gisekia L. Jour. S. Afr. Bot. 27: 131-137. 1961. GAGNEPAIN, F. Place de pa genres soi-disent de la famille des Ficoides. Bull. Soc. Bot. France 65: 7-10. 1918. Goprrey, R. K. Gisekia pharnacioides, a new weed. Rhodora 63: 226-228. 1961. Josut, M. C., & O. P. Kampuoj. Studies on the autecology of Gisekia pharna- ceoides Linn. Jour. Indian Bot. Soc. 38: 8-34. 1959 & V. R. Rao. The embryology of Gisekia pharnaceoides Linn. Proc. Indian Acad. Sci. B. 3: 71-92. 1936. [Includes review of previous embry- ological studies in Aizoaceae and Phytolaccaceae. | Wa ter, H. Phytolaccaceae. Pflanzenr. IV. 83(Heft 39): 1-154. 1909. [Gisekia, 1, 24, 25.] Tribe MoLiucIneae™ [ Fenzl ] 2. Mollugo Linnaeus, Sp. Pl. 1: 89. 1753; Gen. Pl. ed. 5. 39. 1754. Rather variable glabrous annual herbs [or subshrubs] of dry, open, disturbed habitats. Stems prostrate to decumbent, radiating from a short, slender, tapering taproot, dichotomously much-branched, slender, wiry, with slightly swollen nodes. Leaves simple, unequal, exstipulate [or stipules rare, obsolete or early deciduous], scarcely fleshy, forming basal and cauline whorls of 3—8, the whorls often separated by long internodes, basal leaves usually largest, up to 4 cm. long, cauline leaves gradually reduced upwards; blades linear lanceolate to oblong, elliptic, spatulate or obovate, the apex acute to obtuse, the base gradually tapering to a short, membranaceous-margined petiole. Inflorescence an axillary [or terminal] umbellate cluster of 2-5 inconspicuous, short-pedicellate flowers, the pedi- cels strongly reflexed after flowering. Sepals 5, free, imbricate (or quin- cuncial), green without, whitish within, margins hyaline, spreading at anthesis. Petals none.2 Stamens usually 3 and alternate with the car- pels, sometimes 4 or 5 [rarely more] and alternate with the sepals, united basally by a very shallow ring of filament tissue; anthers versatile, the 2 lobes separated below. Carpels 3[-8], syncarpous; styles 3 [-5], free, very short, erect, recurved; stigma linear, papillate; ovary superior, 3-loculate; ovules numerous in each locule, the placentation axile. Fruit a thin-walled 3[—5]-loculate, ellipsoidal capsule, longer than and included within the persistent sepals, dehiscing loculicidally by 3{—5] persistent valves, the partitions separating from the persistent central axis bearing the funiculi of the ovules. Seeds numerous, somewhat compressed, reni- d find no evidence in the many North oe specimens of Mollugo ] verticillata in the Harvard University Herbaria (A, GH, NEBC) of the five white, petaloid staminodia described for this species by Ahles ix "hice Ahles, & Bell, 1968). 438 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 FicurE 1. MOLLUGINACEAE. a-h, Mollugo. M. verticillata: a, part of a flower- ing plant, from above X %; b, flowering and fruiting branch enlarged, pa, eon flower, X 15; d, flower with forward a depressed to show enlarging superior ovary ‘and hypogynous stamens, X 15; e, cross section of ovary showing placen- tation, X 15; f, mature fruit before dehiscence, — persistent sepals and sta- oe xX 8; g, fruit after dehiscence, < 8; h, seed with peglike hilum, * 25. Glinus. G. lotoides: seed — note long coiled ange and inflated aril around Sviccalis, A ook form to roundish, reddish- to blackish-brown; seed-coat bearing 3-7 prominent to obscure longitudinal ridges or smooth; a distinct hilar peg present, but caruncle and filiform appendage lacking. Embryo periph- eral, curved around a small amount of hard, semi-transparent endosperm; cotyledons oblong, fleshy. Lectotype species: M. verticillata L.; see Britton & Brown, Illus. Fl. No. U.S. ed. 2. 2: 35. 1913. (An ancient generic name derived from Latin “Mollitie plantae”, pliant or delicate 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 439 plants; see Linnaeus, Crit. Bot. 109. 1737; Philos, Bot. 157. 1751) ,— CARPET-WEED, INDIAN CHICK-WEED, DEVIL’s GRIP. A genus of about 35 species native to the tropics and subtropics of both hemispheres, but now widespread in temperate regions in North America and Europe as a weed of disturbed areas. Wilson lists nine species in the North American Flora, but these are mainly distributed in the West Indies and Central and South America. Only Mollugo verticillata L., m — 64, occurs in our area, ranging from Florida northward through New England to Nova Scotia, westward to Ontario and North Dakota, southward to Texas, New Mexico, and Mexico, while on the Pacific coast it ranges from southern California northward to Oregon, Washington, and Idaho. A second species, M. Cerviana (L.) Seringe, 2m = 18, ranging from Arizona to Lower California and Mexico, is distinguished from the rela- tively large, coarse, mat forming plants of M. verticillata by its small, ascending, diffuse habit, filiform stems, small linear leaves, long-pedicellate flowers usually with five stamens, and seeds with a reticulate seed-coat. This species has been reported to occur on ballast in the vicinity of New York City. In our area Mollugo verticillata may occur in almost any disturbed sit- uation, including cultivated ground, roadsides, railroad beds, sandy river banks, exposed lake-bottom muds, beaches, sand-dunes, and gardens. It flowers from spring to fall. Similar open or disturbed habitats are report- ed in autecological studies in India by Bakshi & Kapil on M. Cerviana, M. nudicaulis Lam., 2n = 54, and “M. hirta Thunb.” (= Glinus lotoides L.). The species are adapted to xerophytic conditions, with short life cycles that progress from germination after a few seasonal showers to flowering and fruiting within a few days, to death within 30 to 45 days. Self and insect pollination and wind dispersal of seeds are reported for these species. The genera Coelanthum, Glinus, Hypertelis, and Pharnaceum are closely allied to Mollugo, which is possibly the most primitive of the group, being least specialized in habitat, habit, and floral structure (Adamson). The species of Glinus were treated by Bentham & Hooker and in parts of Index Kewensis as synonyms of Mollugo, an unfortunate taxonomic judgment that created much nomenclatural confusion in the literature. a5 Scattered chromosome counts for seven species of Mollugo indicate a polyploid series with diploid numbers of 18, 36, 54, and 64. The genus has attracted relatively little interest. Howell described a complex of endemic evolution among eight closely related species in the Galapagos Islands in which two divergent lines have developed from a single variable glabrous annual prototype that had flowers with three or five stamens, a condition found in M. verticillata, to which the Gala- pagean species are “perhaps most closely related.” Differentiation has re- sulted in “one line developing a subshrubby perennial habit and large flowers, the other line developing a low woody caudex and strikingly 440 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 glandular herbage. In both lines there is an increase in the number of stamens, approaching or equalling eight, the sum of the five stamens op- posite the sepals and the three stamens alternate with the cells of the capsule, two staminal arrangements which occur in the M. verticillata type. The geographic separation of the several forms on the different islands has undoubtedly been an important factor in the segregation and maintenance of the specialized developments.” Payne described several interesting anatomical features exhibited by Mollugo verticillata, including an increase in thickness of the cortical tissues to produce swollen nodes, pseudoannulation in the xylem of the stems and roots, and the presence of rosettes of cells in the leaf epidermis representing vestigial trichome bases. Anomalous secondary thickening, reported by Metcalfe & Chalk (after Solereder) for “M. radiata R. & P.” (= Glinus radiata (Ruiz & Pav.) Rohrb.) pertains to Glinus and has not been found in species of Mollugo. The floral vascular anatomy of three species of the genus has been described by Sharma. Excellent illustrations of the seeds of M. verticillata and M. Cerviana are provided by Thieret. Stomata of the ranunculaceous type occur on both surfaces of the leaves, while clustered and single crystals of calcium oxalate are found in tissues of the leaves and stems. Tartaric acid, tannins, and bitter glucosides, but no alkaloids, have been found in M, nudicaulis. Pollen grains with four germ pores and a two-celled condition at shedding are recorded for M. verticillata (Payne), in contrast to the three-celled condi- tion reported elsewhere in the family, and Sharma reports pollen grains three-colporoidate in M. nudicaulis, and three- to five-zonicolporoidate in M. pentaphylla L Embryological studies by Payne (Mollugo verticillata), Bhargava (M. nudicaulis), and Kshirsagar (M. stricta L.), indicate that ovules are anacampylotropous at fertilization. Embryo-sac development is normal (Polygonum type), while development of the embryo is probably of the solanad type. Endosperm formation is initially free-nuclear, later becom- ing cellular. The genus is of no economic importance. Plants of Mollugo Cerviana and M. pentaphylla are used in the Old World as a vegetable and in folk medicine against a wide variety of complaints, including skin disorders and diarrhoea, or as a mild laxative. REFERENCES: Under family references see ADAMSON, BAILLON, BENTHAM & Hooker, Dvu- Pont, EcKARDT, EICHLER, FENZL, Fxtepnicy, JEFFREY, PAYER, Raprorp et dl., ROHRBACH, SHARMA, THIERET, and Up Apamson, R. S. The South African species of Aizoaceae. IV. Mollugo, Phar- naceum, Coelanthum, and Hypertelis. Jour. S. Afr. Bot. 24: 11-65. 1958. BAKSHI, T. S., & R. N. Kapi. The autecology of Mollugo nudicaulis Lam. Bull. Bot. Soc. Bengal 6: 45-48. 1954. e morphology and ecology of Mollugo Cerviana Ser. Jour. Indian Bot. Soc. 33: 309-328. 1954. 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 441 Buarcava, H. R. Contribution to the eee of Mollugo nudicaulis Lamk. Proc. Indian Acad. Sci. B. 1: 271-278. 1935. CoriLLion, R. Sur la présence du ie verticillata L. oe a état spontané dans le Val de Loire. Mém. Soc. Bot. France 105: 51, 1958. Hom, T. Mollugo verticillata L. Am. Jour. Sci. IV. 31: 525-532. cre HowELL, J. T. The genus Mollugo in the Galapagos Islands. Proc. Calif. Acad. Sci. IV. 21: 13-23. 1933. Hsu, C. Preliminary chromosome studies on the vascular plants of Taiwan (1). Taiwania 13: 117-129. 1967. [M. pentaphylla, 2n = 18, 36.] Ksurrsacar, B. G. Contribution to the embryology of Polycarpaea corymbosa Lamk. and Mollugo stricta Linn. (syn. Mollugo pentaphylla Linn.). Proc. 47th Indian Sci. Congr. 3: 357. 1960 Kumart, K. A remarkable case of abnormality in the flowers of Mollugo dis- ticha Ser. Proc. 41st Indian Sci. Congr. 4: 30, 31. 1954. [Abnormal ovaries with ovules replaced by leaflike structures bearing ovules at their tips, and axillary condensed shoots bearing leaves and flowers; leaflike carpels with axillary shoots; normal stamens; leaflike sepals. Martin, C. A. The comparative internal anatomy of seeds. Am. Midl. Nat. 36: 513-660. 1946. [M. verticillata, 557.] Payer, J. B. Traité d’organogénie comparée de la fleur. vii + 749 pp. 145 pls. Paris. 1857. [{Mollugineae, 331, 334, pl. 70; under Portulacaceae. | Payne, M. A. Morphology and anatomy of Mollugo verticillata L. Univ. Kansas Sci. Bull. 21: 399-419. 1933 . The seni and seed of “Mollugo verticillata. Ibid, 22: 5-25. 1935. Pitot, A. A tir des espéces ouest-africaines poe bieiniend sur les genres “Moll ugo” . “Glinus.”” Webbia 19: 751-808. Rickett, H. W. Wild Flowers of the United ire Yat Zs : Southeastern States, Part 1. New York. 1967. [M. verticillata, 142, pl. Sosa, A. Sur ene saponosides cristallisés de Mollugo . Lamk. tAjaodobes). Compt. Rend. Acad. Sci. Paris. 248: 2243-2245. 1959. STEYERMARK, J. A. eva of Missouri. Ixxxiii + 1725 pp. Ames, Iowa. 1963. [M. verticillata, 632.] 3. Glinus Linnaeus, Sp. Pl. 1: 463. 1753; Gen. Pl. ed. 5. 208. 1754. Annual herbs, with radiating, much-branched, procumbent to ascending stems, forming mats up to 0.5 m. in diameter. Foliage and axes covered with a dense, short, whitish, stellate pubescence, particularly when young [or glabrous]. Leaves alternate to whorled, exstipulate, unequal, entire, + fleshy, elliptic to ovate, spatulate, obovate or orbicular, gradually nar- towed below to a slender petiole, apex rounded to abruptly acute, Inflorescence a dense, sessile to short-pedunculate axillary dichasium. Flowers few to numerous, sessile to short pedicellate. Sepals 5, free, externally stellate-tomentose, margins hyaline, tip mucronate, aestivation quincuncial, persistent. Stamens (3—)5[—20], alternate with and shorter than the sepals [or fascicled when numerous], hypogynous; filament filiform, anther versatile, 2-loculate at anthesis, the 2 lobes separated above and below the point of insertion, dehiscence dorsilateral by longi- tudinal slits. Carpels 3(—5), united; styles wanting; stigmas 3(—5), sessile, of hyaline papillae, persistent [or styles 3-5, short, erect]; ovules numer- 442 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ous, anacampylotropous, the placentation axile. Fruit an ovoid, loculicidal capsule, the 3(—5) valves separating from a central axis bearing the funiculi and sterile ovules. Seeds numerous, reniform, with a short funic- ulus enclosed within a bladder-like caruncle and a long filiform appendage coiled round the seed; seed-coat tuberculate to nearly or quite smooth and shiny, reddish- to blackish-brown. Embryo curved; cotyledons oblong, fleshy, endosperm starchy. Type species: G. lotoides L. (Name from Greek glinos, sweet juice, the significance not clear.) A genus of about thirteen species with pantropical distribution, reaching temperate areas as an introduced weed. Two species are found in our area. Glinus lotoides L., with obovate to orbicular leaves and usually tuberculate seeds, has been found from Louisiana (Thieret) to Oklahoma, Arkansas, and Missouri, in moist situations on alluvial muds and sands on the margins of rivers and lakes, as well as on moist forest soils. Glinus radiatus (Ruiz & Pav.) Rohrb., with narrower leaves and smooth seeds, is reported from Louisiana, Oklahoma, and California, south- ward in Mexico to Central America, and in the West Indies. These two species are distinguished primarily on the texture of the seed coat (cf. Wilson), but the limits of their variation and distribution in the southern United States appear rather indefinite, suggesting the need for further collection and study. Although considered distinct by Linnaeus, Glinus was later reduced to synonymy under Mollugo L. by Bentham & Hooker and in the earlier volumes of the Index Kewensis. The extensive and very confusing syn- onymy which resulted is discussed in detail by Pitot. Glinus lotoides frequently appears under the synonym Mollugo hirta Thunb. The genera are now generally accepted as distinct, with Glinus differing from Mollugo principally in the presence of a conspicuous caruncle and a filiform ap- pendage (frequently mistaken for the funiculus) on the seed, and in the dense stellate pubescence of Glinus. Adamson (1960) suggests that Glinus was derived from the same stock as Mollugo, but along a distinct line, and that the Brazilian Glischrothamnus Pilg. is derived from Glinus. Chromosome counts of 2% = 36 have been reported for both Glinus lotoides and G. oppositifolius (L.) A. DC. The report by Solereder (re- peated in Metcalfe & Chalk) of anomalous secondary thickening in stems and roots of G. radiatus (as Mollugo radiata Ruiz & Pav.) should perhaps be reconfirmed. Both anomocytic and paracytic stomatal types occur in the leaf epidermis (Inamdar). The floral vascular anatomy of G. lotoides and G. oppositifolius is described by Sharma. The embryology of only G. lotoides is known. The walls of the tetrasporangiate anther are four cell-layers thick. The cells of the single tapetal layer each contain two to three many-nucleolated nuclei. Mature pollen grains are tricolpate (Sharma), and distinctly three-celled when shed. Ovules are anacampy- lotropous and crassinucellate, with two integuments, The caruncle (or “aril”) may finally enclose as much as the lower third of the ovule in some instances, but its size varies in different specimens, and seeds of 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 443 G. lotoides lacking the caruncle are illustrated by Pitot. Embryo-sac de- velopment is of the Polygonum type, and the mature embryo sac contains abundant starch grains. The genus is of no commercial value. In India the leaves of Glinus lotoides and G. oppositifolia are used as a vegetable or as an appetizer. As an herb the plants are used as a purgative or as a remedy for diarrhoea, itches, and skin diseases. REFERENCES: Under family references see ADAMSON (1960), BAILLON, BENTHAM & HOOKER, DuPont, FENZL, FRIEDRICH, JEFFREY, ROHRBACH, SHARMA, THIERET, and WILSon. ApAmson, R. S. The South oii species of Aizoaceae. IX. Glinus. Jour. S. Afr. Bot. 27: 125- og: E.-Hamior, A., S. Necm, & A SHARAF, An re ama of Glinus lotoides. Qualitas Plant. sv epped Veg. 14: 147-155. 1967.* Inampar, J. A. Epidermal structure and stomatal ontogeny in some Polygonales and Ceatrosierinas. Ann. Bot. II. 33: 541-552. pl. 1. 1969. [G. lotoides.] Mirra, K., & N. Darra. Jn: 1.0.P.B. Chromosome number reports XIII. Taxon 16: 445-461. 1967. [G. lotoides, G. oppositifolius. | Narayana, H. S., & B. C. Lops. Contribution to the development of male and female gametophytes of Glinus lotoides. Proc. 48th Indian Sci. Congr. 3: 274. 1961. Piror, A. A partir des espéces ape africaines considerations sur les genres “Mo llugo” et “Glinus”. Webbia 19: 751-808. 1965. SHAH, C. S., & M. SUKKAWALA. goouminaae Naar study of Mollugo hirta Thunb. (Glinus lotoides Linné). Indian Jour. Pharm. 21: 64-68. 1959.* SHARMA, A. K., & N. K. BuatracHaryaA. Cytogenetics of some members of Portulacaceae and related families. Caryologia 8: 257-274. 1956. [Mollugo THIrRET, J. W. Thirty additions to the Louisiana flora. Sida 3: 123-127. 1967. [G. lotoides, 124.] AIZOACEAE Rudolphi, Syst. Orb. Veg. 53. 1830, tea a Sprengel, Anleit. ed. 2. 2(2): 842. 1818, “Aizoiden’’|, nom. (MESEMBRYANTHEMUM FAMILY) Slightly fleshy to markedly succulent annual or perennial herbs and low subshrubs_ [or shrubs], with glabrous, papillose, or pubescent, prostrate to erect and usually much-branched stems spreading from a small, narrow taproot, often forming mats. Leaves + succulent, cauline, alternate or opposite, stipulate or exstipulate, sessile or petiolate, the petioles often dilated, the membranaceous margins sometimes decurrent, or connate in a Sheath about the node; blades simple, entire, flattened to + cylindrical, glabrous, papillate or pubescent. Flowers mostly inconspicuous, regular, perfect; axillary or terminal (but appearing axillary), sessile to pedicel- 444 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 late, solitary, paired, or few in a cymose or spike-like cluster [or inflores- cence a dichasium], often flanked by a pair of small, sessile, marcescent bracts. Sepals (tepals) (3—)5, equal or unequal, united below in a usually shallow tube, the tube free from or united with the ovary, the lobes imbricate or quincuncial, erect, hooded or incurved above, + fleshy, sometimes keeled and bearing a subapical dorsal appendage, greenish without, white or pink to purplish within, persistent or falling as a unit. Petals none [or few to numerous, staminodial, often colorful]. Stamens (1-)3, 5, (7-)10(-13), to numerous, solitary or in pairs or groups al- ternate with and opposite the calyx lobes or in a continuous cycle, free or united basally, inserted on the floral tube, persistent or falling with the sepals; anthers 4-sporangiate, 2-loculate at anthesis; pollen mostly 3-colpate or colporoidate. Carpels rarely 1 or 2, usually 3—5[—20], united; styles as many as the carpels, free or united below; ovary superior or half-inferior |to infer rior], 1—5-locu late or incompletely 2-chambered by to smal wrinkled or tuberculate-striate, brown to black; embryo + curved to annular; nena starchy, hard and semitransparent or soft and white. TYPE GENUS: Aizo6n L., A family of about eleven more or less weedy genera (including Tetra- gonia and Mesembryanthemum, sensu lato, but excluding the fourteen genera now segregated in the Molluginaceae) and approximately 2500 species, with a primary center of distribution in South Africa, but also well developed in the Mediterranean region and Australia, and distributed in dry regions of the tropics and subtropics of both hemispheres. Five genera (Cypselea, Galenia, Sesuvium, Tetragonia, Trianthema), and six species occur in our area, mainly as strand plants or weeds of waste places. Only two of the six subfamilies now recognized in the family ae gen Eckardt) reach our area: the monotypic Tetragonioideae Lindl. apetalous flowers, half-inferior ovaries, and stony indehiscent rn and the Aizodideae, containing ten apetalous genera with superior ovaries and capsular fruits. The subfamilies Aptenioideae, Hymenogynoideae, Caryotophoroideae, and Ruschioideae are either largely or entirely confined to Africa. The family appears to reach an evolutionary climax in the genus Mesembryanthemum, sensu lato, with its many curious, strongly succulent plants and striking flowers with many colorful petaloid stamino- dia. Mesembryanthemum has been split into about 120 segregate genera * The monotypic genus Geocarpon Mackenzie, disjunctly oasis in south- western Missouri (Greene, Jasper, Polk, and St. Clair counties) a ee cn Arkansas (Drew and Bradley counties), and cited under Precis nhac various authors (Fernald, Gleason, Pax & Hoffmann, Wilson), has been transferred to Caryophyllaceae by Palmer & Steyermark (1950) as the type of a monotypic tribe Geocarpeae (in subfam. Alsinoideae 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 445 and elevated to family rank (see Ihlenfeldt, Schwantes & Straka, 1962; Eckardt, 1964; Pax & Hoffmann, 1934; for literature citations and ac- counts of the systematics of this diffcult group). In the older, more conservative sense the Aizoaceae consisted of a broad and ill-defined alliance of about 25 genera (including those now placed in Molluginaceae) that has appeared in the literature under several family names, including Aizoaceae (Pax, Pax & Hoffmann), Ficoidaceae (Bentham & Hooker, De Candolle, Jussieu), Sesuviaceae (Horaninow), and Tetragoniaceae (Small, Wilson). The question of the correct family name has been problematical (cf. Sprague, Rowley, Friedrich), but Aizo- aceae is now conserved. In the older literature various genera have been attributed to such related families as Caryophyllaceae, Phytolaccaceae, and Portulacaceae, or more recently placed in segregate families such as Molluginaceae Hutchinson, nom. cons. (Rohrbach), Mesembryan- themaceae Fenzl, nom. cons. (Lindley; Endlicher; Ihlenfeldt, Schwantes, & Straka; Schwantes), or Tetragoniaceae Nakai, sensu stricto, nom. cons. Two families, Molluginaceae and Aizoaceae (or Ficoidaceae, including Mesembryanthemum and Tetragonia), are most often recognized in the recent literature (Cronquist, Eckardt, Adamson, Hutchinson, Friedrich). The Aizoaceae are distinguished from the Molluginaceae by their characteristically succulent habit, sepals united basally to form a cal tube, stamens inserted perigynously, and ovary often partly or entirely inferior; by the presence of betacyanin pigments (betalains, reported thus far for 13 genera and 36 species) rather than anthocyanins; and by the frequent occurrence of anomalous secondary growth in stems and roots. The family has long been considered to have been derived from a phytolaccaceous stock, and it is closely related to the Portulacaceae through Sesuvium and Trianthema. Relationships to the Cactaceae have also been proposed (Buxbaum). Available chromosome counts indicate a base number of x = 8 (or 9) for the “semi-succulent” group (including Sesuvium, Tetragonia, and Trianthema), with diploid numbers of 2” = 16, 26, 28, 32, 36, 48, and 96; while x = 9 for the “succulent” group (largely confined to Africa), with numerous counts of 2m = 18, 27, 36, 54, 72, and 108 (Darlington & Wylie). The embryology of the family has not been intensively studied. Avail- able details include tetrasporangiate anthers, with development of the Basic type and containing Ubisch granules (Sesuvium Portulacastrum), and pollen grains three colpate (Sharma) and three-celled when shed. Ovules are anacampylotropous to campylotropous and crassinucellar, with the micropyle formed by the inner of two integuments. A third integu- ment or aril, is reported in Galenia, Sesuvium, and Trianthema. Embryo- sac development is of the Polygonum type (but the Adoxa type has been described for Mesembryanthemum pseudotruncatellum). Starch grains are present in the mature embryo sac. Endosperm formation is initially nuclear, becoming cellular (but remaining nuclear up to eleven days after pollination in Trianthema Portulacastrum). Embryo development is of 446 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the solanad type, and polyembryony has been reported in Trianthema Portulacastrum (Davis). In a study of the floral vascular anatomy of species of Galenia, Sesu- vium, and Trianthema, Sharma (1962) found progressive fusion of the “tepals,” and increasing adnation between the perianth tube and the stamen filaments. These trends are accompanied by progressive stelar condensation in the receptacle, resulting in a transition from independent origins of the tepal and stamen traces, to their adnation and origin as common bundles. He concluded that specialization in the androecium among these species has resulted in a reduction from numerous to few stamens, rather than in an increase in number via “dédoublement.” In all species the carpel walls were supplied with five or more bundles (as opposed to three in the Molluginaceae), a condition which he considered derived. He considered the ‘“Ficoidaceae’ advanced over the Mollugin- aceae in the possession of fused tepals whose traces arise at the same level from the stele and divide within the receptacular cortex; perigynous stamens; semi-inferior to inferior ovaries whose carpellary traces divide before leaving the cortex; and fewer ovules. The organogeny of the flower in Tetragonia and Trianthema has been described in detail by Payer. The family is of economic significance primarily for the many species of Mesembryanthemum and related genera cultivated as ornamentals, and for the various species of Sesuvium, Tetragonia, and Trianthema cultivated for use as vegetables or in folk medicine. REFERENCES: BarLton, H. Mésembryanthémacées. Hist. Pl. 9: 46-53. 1886. [Série Tetra- goniées, 48, 49, 52, 53; figs. 63-69; Tetragonia expansa, T. fruticosa.| Portulacacées. Ibid. 54-80. [ Série Aizoidées, 65, 73-76; Cypselea, Galenia, BENTHAM, G., a D. Hooxer. Ficoideae. Gen. Pl. 1: 851-859. 1867. r Klarung der phylogenetischen Stellung der Aizoaceae und Cactaceae a Pflanzenreich. Sukkulentenkunde 2: 3-16. 1948. [Floral morphology, anatomy, embryology. } CaNDOLLE, A. P. bE. Ficoideae. Prodr. 3: 415-456. 1828. [Tetragonia, 451, 453; Sesuvium, 453-455.] Portulaceae. Ibid. 351-364. [Cypselea, 353; Trian- thema, 351, 352 mt Cronquist, A. The evolution and classification of flowering plants. x + 396 oston. 1968. [Aizoaceae, 181. Detailed discussion of phylogenetic rela- RET) among families of subclass Caryophyllidae, 172~180.] Davis, G. L. Systematic embryology of the angiosperms. 528 pp. New York. 1966. [Ficoidaceae, 121, 122.] D’Husert, E. Recherches sur Je sac embryonnaire des plantes grasses. Ann. Sci. Nat. Bot. VIII. 2: 37-128. 1896. [Tetragoniées, Aizoacées, 111; presence of starch granules in the embryo sac. DuPont, S. Sur la morphologie des plantules de Mesembryanthémacées et d’Aizoacées. Bull. Soc. Hist. Nat. Toulouse 95: 356-360, 1960. [Seedling morphology; distinguishes three distinct forms; Sesuvium, Trianthema, Tetragonia similar. | ; 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 447 Eckarpt, T. Aizoaceae. Jn: H. Metcutor. Engler’s Syllabus der Pflanzen- familien. ed. 12. 2: 87-90. 1964. EIcHLer, A. W. Bliithendiagramme. Pt. 2. Leipzig. 1878. [Aizoaceae, 119-124; Sesuvium, Trianthema, Tetragonia. | FENZL, E. Monographie der Mollugineen und Steudelieen. Ann. Wiener Mus. Naturgesch. 1: 337-384. 1836. [Portulaceae, tribe Portulacellae, 351.] Supplement. Verbesserungen und Zusitze zur ersten Abtheilung der Mon- ographie. /bid. 2: 279-307. 1839. [Tribe Tetragonieae, 287; tribe Aizoideae, incl. Galenia, 288; Sesuvieae, incl. Trianthema, 289, 290, Sesuvium, 292, Cypselea, 293, 294.] FriepricH, H. C. Beitrige zur Kenntnis einiger Familien der Centrospermae. Mitt. Bot. Staatssamm. Miinchen 12: 56-66. 1955. [Ficoidaceae, 58, 59; Tetragoniaceae, 59, 60. Horaninow, P, Systema Naturae. xiii + 142 + xxiv pp. St. Petersburg. 1834. [Sesuviacene, 83, incl. Sesuvium, Tetragonia, Trianthema, Mesembryan- themum, Aizoon. Hurcutnson, The families of flowering plants. I. Dicotyledons. London. 1926. [Ficoidaceae (Aizoaceae), 129.] Ed. 2. Oxford. 1959. [Ficoidaceae, IHLENFELDT, H. D., G. Scuwantes, & H. Straka. Die hoheren Taxa der esembryanthemaceae. Taxon 11: 52-56. 1962. [Mesembryanthemaceae Fenzl emend. Herre & Volk; subfamilies and tribes defined. | Jerrrey, C. Aizoaceae. Jn: C. E. Hupparp & E. MILNE-REDHEAD, eds., FI. Trop. E. Afr. 37 pp. London. 1961. [Sesuvium, 20; Trianthema, 22; Zaleya, LinDLey, J. The vegetable kingdom. ed. 3. lxviii + 908 pp. London. 1853. [Tetragoniaceae, 209; incl. Cypselea, Sie: Sesuvium, Tetragonia, Tri- anthema. | Lussock, J. A contribution to our knowledge of seedlings. Vol. 2. 646 p London n. 1892. [Ficoideae, 13-18; Tetragonia expansa, 17, 18, figs. 403, Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513-660. 1946. [Aizoaceae, 557.] Metcatre, C. R., & L. CHALK. Anatomy of = dicotyledons. Vol. 1. Ixiv + 724 pp. Oxford. 1950. [Ficoidaceae, 706—712.] Mutter, K. Beitrige zur Systematik der Aizoaceen. Bot. Jahrb. 42(Beibl. 97): 54-04, 1908 Nakat, T. Neinlic ad plantas Asiae Orientalis (18). Jour. Jap. Bot. 18: 91-103. 1 [Aizoaceae, 105, 106; Tetragoniaceae, 103, 104; Demidovia (= Tetragonia), Tetragonia, 91-93.] Pax, F. serene (Ficoideae, Mesembrianthemaceae). Nat. Pflanzenfam. IT. 1b: 33-51. & K. rasa. Aizoaceae. Ibid. ed. 2. 16c: 179-233. 1934. Payer, J. B. Traité d’organogénie comparée de la fleur. Texte, viii + 749 pp.; Atlas, viii + 154 pls. Paris. 1857. [Tetragoniaceae, 349-355; Trianthema, bl. 76, Tetragonia, pl. 77.] RicKett, H. W. Wild flowers of the United States. Vol. 2. The oo Slates: Part 1. New York. 1967. [Sesuvium Portulacastrum, 153, pl. S. verrucosum; Pt. 2: 152, 643, We yt Roureacn, P. Ficoidaceae, In: EK P. N Martius. Fl. Brasil. 14(2): 307-314, 316. pls. 70, 71. 1872. poeta Tomueswe ia. | 448 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Row ey, G. D. ee in Ficoidaceae, I. The family name. Natl. Cact. Succul. Jour. 6: 27, 2 SCHWANTES, G. ce ae (Juss.) em. Hutchinson. Kakteen Sukkul. 8: 167— 169. 1957. SHarma, H. P. Studies in the order Centrospermales. III. Vascular anatomy of the flower of certain species of the Ficoidaceae. Proc. Indian Acad. Sci. B. 56: 269-285. 1962. [Galenia, Sesuvium, Trianthema.| IV. Pollen morphology of some species of the families Ficoidaceae, Molluginaceae, Nyctaginaceae, and Portulacaceae. Jour. Indian Bot. Soc. 42: 637-645. 1963. [Galenia, Sesuvium, Trianthema, Tetragonia. | Spracuge, T. A. The nomenclature of plant families. Jour. Bot. London 60: 69- 73. 1922. [Ficoidaceae, 71.| TuieErET, J. W. Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2: 352-360. 1966. [Cypselea humifusa, 357, fig. 3; si ane Portula- castrum, 357, fig. 4; Sesuvium Portulacastrum, 357, fig. Upnor, J. C. T. Dictissacy of economic eos ed. 2. 591 pp. Ashes 1968. WIiison, P. Tetragoniaceae. N. Am. Fl. 21: 267-277. 1932. Woutpart, A., & T. J. Masry. The pelts and phylogenetic significance of the betalains with respect to the Centrospermae. Taxon 17: 148-152. 1968. KEY TO THE GENERA OF AIZOACEAE IN THE SOUTHEASTERN UNITED STATES General characters: fleshy to strongly succulent, glabrous or pubescent, pros- d and forming mats; leaves below into a shallow floral tube, the lobes green, white, pink or purplish within; petals none; stamens mostly 3-10 or numerous, solitary, paired, or in groups, alternate with or opposite the sepals; ovary superior or semi-inferior, 1-5 locu- late ; carpels 1-5, united; ovules and seeds 1—many; placentation ake basal, or parietal; fruit a loculicidal or circumscissile capsule or bony and indehiscent ; seeds reniform to pyriform; seedcoat smooth, wrinkled or tuberculate A. Ovary superior; fruit a loculicidal or circumscissile capsule; leaves alternate or opposite. B. Capsule loculicidal by 3-5 valves; ovules and seeds solitary; stamens 8-10, in pairs alternate with the sepals; stems and alternate leaves densely white-pubescent; stipules absent. ................ 1. Galenia. B. Capsule circumscissile; ovules and seeds few to numerous; stamens 1 many; leaves ORpOsHe C. Ovary 3-5-loculate; seeds numerous; stamens 5 to said sepals appendaged, persistent; leaves + equal: stipules absen vis # 5 Wile EEE Ley ea TA epi ese) See dae A ee ra " Sesuvium. C. Ovary 1- or 2-loculate; leaves unequal; stipules present. D. Stamens 5-10; sepals appendaged; ovules and seeds few; cap of the ovary fleshy and crested; stipules scarious nea S aA sahes CAGES bat eae ban enne ores 3. Trianthema. Stamens 1-3(—5); sepals unappendaged; seeds numerous; cap of the ovary thin-walled and rounded; stipules —— Tuo ELE TRUER Sale RR ee ee Cypselea. o 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 449 A. Ovary half-inferior; fruit horned, indehiscent, bony; leaves patie eX- stipulate, the blades flat, + ovate-triangular, papillos ae . Tetragonia. Subfamily AIZOOIDEAE [Schwantes] 1. Galenia Linnaeus, Sp. Pl. 1: 359. 1753; Gen. Pl. ed. 5. 169. 1754. Perennial, much-branched, grayish [or yellowish to green], prostrate [or erect| herbs or subshrubs [or shrubs]; branches woody at the base, forming dense mats, lateral branches alternate, + secund, spreading ; stems and leaves initially densely coarsely villous with loosely appressed, whitish, flattened hairs, becoming + glabrescent. Leaves alternate [or opposite], secund, entire, fleshy, sessile exstipulate, the epidermis coarsely papillose; leaves variable in size, those of the main stems largest, with blades broadly obovate, tapering below to a broad base, the apex obtuse, younger leaves + elliptic to obovate and acute. Flowers small and in- conspicuous, on short lateral branches, sessile, axillary, solitary or in groups of 2—5 [or inflorescence a terminal, paniculate cyme]. Sepals (4) 5, united below in a shallow floral tube, the lobes erect, the tips curved inward and downward, pubescent without, glabrous and whitish within. Stamens (8—)10, of unequal length, inserted on the floral tube in pairs alternate with and shorter than the calyx lobes; anthers small, yellow; filaments flat, tapering upwards [or filiform]; pollen 3-colpate. Gynoe- cium of [2- ig. 5 united carpels; styles [2—|3—5, free or united below, stigmatic along the inner surface; ovary superior, truncate and [2—|3-5- angled above, the center depressed, 3—5-loculate [or 2- or 1-loculate by abortion]; placentation axile; ovule solitary in each cell, pendulous on an arching funiculus, campylotropous. Fruit a small, leathery, [2—]3—5- angled capsule with depressed apex, dehiscing loculicidally from the top [or indehiscent]. Typr sprcies: G. africana L. (Named for Claudius Galenus, ca. 130-ca. 200 a.v., noted Greek naturalist and physician of Pergamos in Asia Minor.) A genus of about 27 species with a natural distribution confined to southern Africa. The genus is represented in our area by one adventive species, Galenia secunda (L. f.) Sond, (subg. KoLLertA), which can be recognized by its prostrate and secund habit, small axillary flowers, paired stamens, superior 5-angled ovary, and one-seeded carpels. The species is represented in the Harvard University Herbaria by only two sheets of a collection by A. H. Curtis (no. 6869, in 1901) from Pensacola, Florida. In a penned note accompanying one of these Curtis states that “of my Pensa- cola plants none but #6865 grew on strictly ballast ground, but the seeds of most of them doubtless came in ballast — No. 6869 forms dense mats probably 4 ft. across. Leaves thickish & with cinereous color.” These specimens bear flowers but no fruits. Small described the ovary of this species as partly inferior, but in the specimens at hand, and as described by Adamson, the ovary is superior. Seeds from a South African specimen of the species are small, compressed, and semilunate in outline, with a 450 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 coriaceous, iridescent blackish-brown seed coat with longitudinal, tuber- culate striations. The embryo is curved around a copious starchy endo- sperm (but is not nearly annular), and has oblong, fleshy cotyledons about as long as the radicle. The continued presence of the genus in our area needs confirmation. A chromosome count for the genus is lacking. Two subgenera (Fenzl, Sonder, Adamson) or sections (Pax, Pax & Hoffmann) have been consistently recognized in the genus. The composi- tion of the subgenera (GALENIA and Korterta Fenzl) has been emended by Adamson in the only recent treatment of the genus. Galenia is closely related to the South African genus Plinthus Fenzl, the two forming a group within the tribe Aizoéae (Adamson). The pubescence in our specimens is simple, but two-armed hairs occur in some species of the genus. Large bladder-like water storage cells are present in the leaf epidermis. In the dried specimens these collapse, giving the leaf surface a pitted appearance. Delicate spiral thickenings of the vessel walls and anomalous secondary thickening are reported for the genus (Metcalfe & Chalk). The genus is of no economic significance. Galenia spathulata Fenzl may be used as fodder for sheep in southern Africa. REFERENCES: Under family references see BAILLON, besgiog & Hooker, FENZL, METCALFE & CHALK, MULLER, Pax, Pax & Horrm Apamson, R. S. The South ord species of Aizoaceae. III. Galenia L. Jour. _ Afr. Bot. 22: 87-127. . [Revision of the genus. | Scott, J. D. Preliminary ee on the root system of inure nem on the Worcester Veld Reserve. S. Afr. Jour. Sci. 34: shcisgytge SONDER, O. W. Mesembryaceae. Jn: W. H. Harvey & a SonperR, Fl. Capensis 2: 386-479. 1862. [Galenia, 473-479, 2. Sesuvium Linnaeus, Syst. Nat. ed. 10. 1052, 1058, 1371. 1759. Annual or perennial, prostrate or ascending, succulent, glabrous herbs or subshrubs, Stems several from a short tap-root, sometimes rooting at the membranaceous margins clasping [or connate], exstipulate. Flowers perfect, terminal but appearing axillary, solitary or clustered, sessile to long pedicellate, a pair of marcescent bracteoles at the base of the pedicel. Sepals 5, united below, the lobes imbricate, margins scarious, hooded above and bearing a short subapical dorsal mucro, pink to lavender within, spreading at anthesis, persistent. Stamens 5, free and alternisepalous or numerous in groups alternate with the sepals or in a continuous cycle, in- serted on the floral tube, shorter than the sepals, persistent; filaments filiform or subulate; anthers small, often reddish; pollen tricolpate [or polymorphous? |. Styles 2-5, linear, erect; stigma longitudinal, papillose; 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 451 ovary superior, 2-5 loculate; ovules numerous, anacampylotropous, pla- centation axile. Capsule membranaceous, enclosed within the persistent perianth, circumscissile near the middle. Seeds numerous, small, round- reniform, smooth, lustrous, black, arillate, the aril membranaceous, tightly investing the seed; embryo annular or slightly coiled; cotyledons oblong, fleshy; endosperm soft, starchy. (Including Halimum Loefl., Halimus Kuntze, Squibbia Raf.) Type species: S. Portulacastrum (L.) L. (Name Latin, presumably after Sesuvium, land of the Sesuvii, a Gallic tribe, the significance here not explained.) — SEA PURSLANE. A genus of about eight species distributed in the tropics and subtropics of both hemispheres, inhabiting saline beach-dunes, marshes, hammocks, and disturbed situations in coastal areas. Two rather variable species occur in our area. Sesuvium maritimum (Walt.) BSP. (S. pentandrum Ell.) with flowers small and sessile or nearly so and five alternisepalous stamens, ranges from Long Island southward to Florida and the West Indies, westward to northern Louisiana and Texas. Sesuvium Portulacas- trum (S. sessile Pers.) 2n = 16, 36, 48, with larger and distinctly pedicel- late flowers and numerous stamens, appears to occur sporadically in the Carolinas, and from Georgia southward through Florida to the West Indies, westward to Texas, and southward to Mexico, Central America, and South America. Sesuvium crithmoides Welw., a native of Angola, with long, narrowly linear leaves and sessile flowers, was collected once in 1902 on the waterfront in Brunswick, Georgia, but has not been reported since. In the southwestern United States Sesuvium verrucosum Raf., with an often more erect habit, numerous stamens, and vegetative surfaces densely papillate with large epidermal water-storage cells, ap- proaching our area in Oklahoma and Texas, ranges northward and west- ward through Kansas, Colorado, Utah, Nevada, and central California, and southward to Mexico. Sesuvium is closely related to Cypselea Turp. and Trianthema L. It has been recognized as distinct by all authors except Degener & Degener, who reduce it to synonymy under Trianthema, stating that the difference between the two genera is mainly “the number of cells in the ovary, a character . . . of trivial importance.” These three genera have been 2 omnled allied, and the group is thought to link the Aizoaceae to the Portulaca Poneidane counts, available only for Sesuvium Portulacastrum, in- dicate the presence of polyploidy and aneuploidy, with base numbers of eight (and nine), and 2n = 16, 32, 36, 48. The presence of betacyanin has been demonstrated in S. Portulacastrum. Species of Sesuvium exhibit structural adaptations to xerophytic condi- tions. The succulent leaves of S. Portulacastrum have stomata on both surfaces and contain subepidermal palisade tissues that are four to five cell-layers thick and enclose a central zone of loose, water-storing paren- chyma. In contrast, the leaves of S. maritimum contain a homogeneous chlorenchyma interrupted only by large intercellular spaces that underlie 452 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 t mbra eed in s sheesh curved embryo sh ee asdnecaee pon ir coanient Xa large water-storage cells in the epidermis. The epidermal cells have a thickened outer wall with a smooth cuticle. Stomata are of the ranun- culaceous type. Crystal clusters occur in cells of the pith, cortex, and leaves. Anomalous secondary thickening is reported in the stems and roots of S. Portulacastrum, The ovules of Sesuvium Portulacastrum have two integuments (the inner of which forms the micropyle) and an aril (or third integument). The inner integument is two cell-layers thick, the second integument is initially two layers thick but becomes three layered, while the aril is three cell-layers thick. A Polygonum-type embryo sac is formed, and embryo development probably conforms to the Linum variation of the solanad type. The nucellus immediately above the embryo sac consists of two to three layers of radially elongate cells, while five to six cell- layers are present along the sides of the sac. The endosperm is initially free nuclear, but later becomes cellular. Anthers are tetrasporangiate, with walls five cell-layers thick, the innermost layer forming a glandular, binucleate tapetum. Ubisch bodies are present in the anthers. Pollen grains are three celled when shed. Sesuvium Portulacastrum is cultivated as a vegetable in some Asian countries. 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 453 REFERENCES: Under family references see BAILLON, BENTHAM & Hooker, Davis, DuPont, FENZL, MaArtTIN, METCALFE & CHALK, MULLER, Pax, Pax & HorrMann, RICKETT, UpHoF, WILSON, and WouHLPART & Mapry. ADAMSON, R. S. The South African species of Aizoaceae. XII. Sesuvium, Tri- anthema, and Zaleya, Jour. S. Afr. Bot. 28: 243-253. 1962. DEGENER, O., & I. DEGENER. Trianthema L. Flora Hawaii. Fam. 116. Aizoaceae. 1960 [ Trianthema and Sesuvium united. | Futvio, T. E. pt. Recuento cromosomicos en angiospermas argentinas. Kurt- ziana 4: 87-90. 1967. [S. Portulacastrum, n = 8. HARSHBERGER, J. W. The comparative leaf structure of the sand dune plants of Bermuda. Proc. Am. Philos. Soc. 47: 97-110. pls. 1-3. 1908. [S. Portula- castrum, 106. JEFFREY, C. Notes on tropical African Aizoaceae. Kew Bull. 14: 235-238. 1960. [Segregates the intermediate genus Zaleya Burm. f. to clarify con- fused limits between Sesuvium and Trianthema. | Kearney, T. H., Jr. The plant covering of Ocracoke Island; a study in the ecology of the North Carolina strand vegetation. Contrib. U. S. Natl. Herb. KEINHOLz, R. An ecological anatomical study of beach vegetation in the Philippines. Proc. Am. Philos. Soc. 65(Suppl.): 58-100. 1926. [S. Portula- castrum, leaf anatomy, 81. | RAGHAVAN, T. S., & V. K. Srinivasan. Studies in the Indian Aizoaceae, Ann. Bot. IL. 4; 651-661. 1940. [S. Portulacastrum; embryology, 653; » = 24, 659. ] SHarMA, A. K., & N. K. BuarracHaryya. Cytogenetics of some members of Portulacacese and related families. Caryologia 8: 257-274. 1956. [S. Portulacastrum, 2n = 36.]| 3. Trianthema Linnaeus, Sp. Pl. 1: 223. 1753; Gen. Pl. ed. 5. 105. 1754. Succulent, glabrous [or pubescent], herbaceous or suffrutescent, annual or perennial plants of waste places. Stems prostrate or erect, from a short tapering taproot, often much branched, the branches alternate or opposite and usually bearing decurrent lines of hairs from the petiolar sheath to the node below, glabrescent. Leaves opposite, the members of a pair conspicuously unequal; blades glabrous, fleshy, elliptic-lanceolate to spatu- late, obovate, or suborbicular, the apex acute or obtuse to truncate-acumi- nate or retuse, the base acutely tapering to rounded; petiole margins expanded, membranaceous, connate into a sheath around the node, stipu- late. Flowers axillary, solitary or cymosely clustered |or inflorescence a dichasium], sessile or short-pedicellate, perfect. Sepals 5, united below and falling as a unit [or persistent], the tube fused with two bracteoles, the petiolar sheath, and even the stem, the lobes imbricate, cucullate, with a sometimes-barbed dorsal mucro just below the apex [or mucro absent], inner surface colored bright- to purplish-pink [or white]. Sta- mens 5-10, inserted on the floral tube in two cycles alternate and opposite the sepals [or stamens 1-3 or numerous in fascicles alternate with the sepals], shorter than and falling with the sepals; anthers small, rotund 454 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 j £ ¥ E i a FIGURE 3. ianthema, a-k, Portulacastrum: a, flowering branch, the ely flowers Tidden by the taal sheathing leaf bases and secondary bracts, xX 1;.D,-an axillary flower with connivent calyx lobes — note sharply. pointed secondary Baer in rece d gps petioles, < 3; c, flower in semi idiagram- matic vertical section to show connate sepa 5, peri gynous aaa of “fo ral parts, semi-inferior ovary, pavietit placentation, ae: horizontal septation of pig X 7; d, stamen, X 20; e, lateral view of two piles note aril develop- ing from funiculus, x "30; f, ‘nearly mature fruit inclosed by tube consisting of fused leaf bases, secondary ‘bract cts, and perianth— note fleshy cap protruding, X 3; g, same, floral tube removed — note horizontal line of dehiscence between fleshy cap pr nd | membranaceous poo of pyxis, X 7; h, fruit in semidiagrammatic vertical section, X ection of ovary, showing parietal plncentation, x pap wit vith tight ly fete aril, x “ k, seed in section, showing curve par ies inclosing endosperm (stippled), ~~ to oblong; filament filiform; pollen tricolpate. Style 1[—2], short (in ours), excentric; stigma longitudinal, papillate; ovary superior, one or incompletely 2-chambered by a horizontal partition, + truncate above and bearing an excentric, often bilobed marginal crest [or peripheral corona]; ovules few, campylotropous, arillate, placentation basal or parietal. Capsule superior, + turbinate, closely surrounded below by the floral tube, walls membranaceous below but fleshy above, apex truncate, 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 455 circumscissile near the middle, the fleshy cap exserted and bearing a bilobed marginal crest and including the upper chamber of the ovary. Seeds [1—]several, round-reniform, arillate, wrinkled, reddish-brown to black, the upper 1 or 2 often enclosed in the upper chamber of the ovary and falling with the indehiscent operculum, 2 or more remaining in the lower chamber. Embryo annular to slightly coiled, surrounding the starchy endosperm; cotyledons oblong, fleshy. (Including Portulacastrum Juss.). Type species: T. Portulacastrum L., 2n = 26. (Name derived from Greek treis, three, and anthemon, flower.) — SEA PURSLANE, HORSE PURSLANE. A small pantropical genus of perhaps 25 species inhabiting sandy sea- beaches, hammocks, alkaline or clay soils, and waste places where in- troduced. The genus has centers of development in Africa and Australia and is represented in our area only by Trianthema Portulacastrum, which ranges from New Jersey to Florida and the West Indies, westward to Texas and southern California, northward to Oklahoma and Missouri, and southward to Mexico and Central and South America. The species is quite variable in habit, pubescence, development of the inflorescence, length of the bracteoles and sepals, number and length of stamens, and form of the crest of the operculum. Two sections were recognized in the genus in De Candolle’s Prodromus (sect. ZALEYA Burm. f. and sect. Rocama Forsk., which included T7i- anthema Portulacastrum). More recently Jeffrey (followed by Adamson) has recognized two subgenera: TRIANTHEMA, with flowers usually solitary and ovaries containing four to numerous ovules, and PAPULARIA (Forsk.) Jeffrey with flowers usually clustered and ovaries containing two super- posed ovules. Jeffrey resolved the confused generic limits between Tri- anthema and Sesuvium in Africa by reestablishing the genus Zaleya Burm. f. to contain the digynous species T. decandra L. (locules and stigmas two, operculum two valved). Trianthema is also closely related to Cypselea Turp. The few chromosome counts available suggest the development of polyploidy and aneuploidy within the genus, since 2n = 16 in Trianthema argentina Hunziker & Cocucci, T. decandra, T. pentandra L., and T. polysperma Hochst., while 2n = 26 and 36 in T. Portulacastrum (as T. monogyna), and 2n = 28 and 32 in T. crystallina Vahl. A relationship between level of ploidy and ecological characteristics in several African species is suggested by Hagerup. Morphological adaptations to xerophytic conditions are apparent in the leaf structure of Trianthema Portulacastrum, in which a central, compact, palisade-like chlorenchyma tissue is bounded above and below by one or two layers of thin-walled water-storage cells and the epidermal layers are uncutinized. Stomata (both anomocytic and paracytic types: Inamdar) occur on both surfaces of the leaf. The upper epidermis consists of large, angular, thin-walled cells, and bladder-like epidermal cells are recorded in T. crystallina Vahl. The leaf veins are surrounded by a sheath of 456 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 parenchyma cells that lack chloroplasts but contain abundant aici grains. Drusiform clusters of calcium oxalate crystals are abundant in the parenchymatous tissues of the leaf and stem of T. soon Anomalous secondary thickening has been reported in Trianthema, but its occurrence should perhaps be confirmed. Embryological studies of Trianthema argentina, T. decandra, and T. Portulacastrum (as T. monogyna L.) indicate that ovules are either campy- lotropous or amphitropous and have a bulky nucellus surrounded by two integuments (the inner of which forms the micropyle) and an aril or “third integument.” In 7. argentina a characteristic air space is reported between the inner and outer integuments at the chalazal end of the ovule, and the cells of the nucellar epidermis are reported to elongate and undergo periclinal divisions to form the nucellar cap (Cocucci), while in 7. Portula- castrum the epidermal cells are said merely to stretch out radially (Bhar- gava). Embryo-sac development is of the Polygonum type, and starch grains accumulate in the embryo sac. The endosperm is initially nuclear, but later becomes cellular from the micropylar end. Development of the embryo is of the solanad type (probably the Linum variation). Poly- embryony has been reported in T. Portulacastrum. The anther walls develop normally and contain five cell-layers at maturity, including a binucleate cellular tapetum. Pollen grains are three celled when shed. The alkaloid trianthemin (of unknown structure) has been isolated from the Indian drug “punarnava,” which is derived from Trianthema Portula- castrum (T. monogyna) and is used in the Ayurvedic medicine of India in cases of heart disease, anemia, and other disorders. The presence of betacyanin has also been demonstrated in this species. The genus is of no commercial value, but some species are cultivated as vegetables in eastern Asia. Leaves of Trianthema Portulacastrum are used as a vegetable in India and Africa, and a decoction of its roots serves as an emenagogue in the Philippines and may be abortive in large doses. Its root is also used as an amenorrheic or as a cathartic in powdered form with ginger. Ashes of the herbaceous T. salsoloides Fenzl are mixed with simson oil and lime in Sudan to make a soap. Plants of T. pentandra L. are also used in Sudan as an astringent in abdominal diseases, while the leaves are eaten as a potherb in times of famine, and ashes of the plant are a source of salt for the natives (Uphof). Several species of the genus, including 7. Portulacastrum, are reputed to be poisonous to livestock because of their content of alkaloids. ove saponins, or nitrates. REFERENCES: Under family references see BAILLON, BENTHAM & Hooker, Davis, DE CAN- DOLLE, ECKARDT, EICHLER, FENZL, MARTIN, eee Pax & HorrMAnn, PAYER, RICKETT, UpHor, WILson, and Woutpart & MAsr Apamson, R. S. The South African species of Aizoaceae. XII. Sesuvium, Tri- anthema, Zaleya. Jour. S. Afr. Bot. 28: 243-253. 1962 Buarcava, H. The life-history of Trianthema monogyna Linn. Proc. Indian Acad. Sci. B. 2: 49-58. 1935. [Embryology. | 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 457 Cocucct, A. E. Embriologia de Trianthema argentina (Aizoaceae). (English summary.) Kurtziana 1; 105-122. 1961. Everist, S. L. A review of the poisonous plants of Queensland. Proc. Ro oy. Soc. Queensl. 74: 1-20. 1964. [T. galericulata, T. Portulacastrum, T. Hacervp, O. Uber Polyploidie in Beziehung zu Klima, Okologie und Phylogenie. Chromosomenzahlen aus Tumbuktu. Hereditas 16: 19-40. 1932. [T. crystal- lina, fig. 18; T. ge peor fig. 20; T. pentandra, fig. 19; pp. 24, 26, 30.] Hunziker, A. T.., . E. Cocucct. El género Trianthema (Ateoaens) en la Reptblica peed (English summary.) Bol. Acad. Nac. Ci. Cérdoba 41: 17-28. 1959. [Includes map, annotated bibliography of recent taxonomic literature. | Inampar, J. A. Epidermal structure and stomatal ontogeny in some Polygonales and Centrospermae. Ann. Bot. II. 33: 541-552, pl. 1. 1969. [T. Portula- castrum. | Jerrrey, C. Notes on tropical African Aizoaceae. Kew Bull. 14: 235-238. 1960. [Revision of Trianthema, two subgenera recognized, and genus Zaleya KEINHOLz, R. An ecological-anatomical study of beach vegetation in the Philippines. Proc. Am. Philos. Soc. 65(Suppl.): 58-100. 6 pls. 1926. [Leaf anatomy of T. Portulacastrum. i] Racuavay, T. S., . K. Srinivasan, Studies in the Indian Aizoaceae. Ann. Bot. II. 4: 651- 661. 1940. [Embryology of T. decandra L.; chromosome counts for T. decandra and T. Portulacastrum. | SHARMA, A. K., & N. K. BuatrracHaryyA. Cytogenetics of some members of Portulacaceae and related families. Caryologia 8: 257-274. 1956. [T. mon- ogyna (= T. Portulacastrum), 2n = 26.| Wess, L. J. Guide to the medicinal and poisonous plants of Queensland. Counc. Sci. Industr. Res. Bull. 232. 1948. [T. crystallina, T. decandra, T. Portula- castrum, T. pentandra, 14.]| 4. Cypselea Turpin, Ann. Mus. Hist. Nat. Paris 7: 219, pl. 12, figs. 0- 11. 1806 Small, prostrate, glabrous to papillose, fleshy, herbaceous or suffrutes- cent, densely branched annuals forming small, leafy mats; branches slender, from the top of a short, slender taproot. Branching alternate, sympodial (at least in part). Leaves small, opposite, entire, fleshy, the members of a pair distinctly unequal; blades elliptic to elliptic-oblong or Ovate, apex acute or obtuse, base acute to rounded; petiole often as long as the blade, dilated below, the margins scarious, clasping, those of a leaf pair sometimes connate and sheathing the node; stipules scarious, laciniate. Flowers appearing axillary, solitary, small, accompanied in the axil by two small, scarious, laciniate bracteoles. Sepals 5, unequal, erect, united below for about one third their length, the margins scarious, the lobes thicker and slightly keeled above, lacking a dorsal appendage, green or purple-tinged without, white to reddish within, quincuncial, persistent. Stamens 1—5, alternate with and shorter than the calyx lobes, inserted at the sinuses, persistent; anthers small, subglobose, whitish; filaments fili- form. Styles 2[-3], very short, erect, slightly united below; ovary 458 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 superior, 2[—3]-loculate at anthesis; ovules numerous, campylotropous, exarillate; placentation axile. Fruit a subglobose, 1-loculate, thin-walled pyxis circumscissile below the middle, remnants of the septum remaining; seeds numerous, small, on long funiculi from the persistent central axis, subreniform. Seed coat smooth, brown; embryo arcuate around a scanty, starchy endosperm; cotyledons fleshy. (Including Radiana Raf., Mil- legrana Juss. ex Turpin.) Type species: C. humifusa Turpin. (Name from Greek, kypsele, a hollow vessel or a beehive, probably because of the shape of the pyxis.) A very small New World genus of possibly three species, inhabiting drying swamp land and other moist, disturbed sites on shore lines, ham- mocks, and sandy pinelands. The genus is represented in our area by Cypselea humifusa in peninsular Florida and in Louisiana; this species is also known from central California. It may have been introduced into all three of these areas from the West Indies (Cuba, Hispaniola, St. Thomas, Guadeloupe, and others). Cypselea humifusa, usually with green calyx lobes and with stamens mostly one to three, was first described from Haiti (Saint Domingue). Cypselea rubriflora Urb., with longer pedicels, reddish calyx lobes and five stamens, has been described from Cuba. C. Meziana K. Miill., of Paraguay, is said to differ from both of these species in having three styles and a three-loculate ovar The extent and taxonomic significance of color variation in the calyx lobes and other plant parts is not known, since collectors’ notes on color are rare. Specimens of a recent collection by Thieret of Cypselea hu- mifusa from St. Landry Parish, Louisiana, are lightly tinged with purple in the stems, leaf tips, calyx tube and lobes, and darkly so in the fruits, but such coloration is lost in older herbarium specimens, which become rather brown Although plants of the genus are glabrous, the nodes often appear pilose due to the protruding segments of the laciniate bracteoles and stipules. The solitary flowers appear to be axillary but are actually ter- minal. Growth of the vegetative axis beyond each flower is sympodial, the axillary bud of the larger leaf continuing the axial growth while the axillary bud of the smaller leaf produces only a short fascicle of small leaves. At anthesis the ovary is distinctly septate, with axile placentation, but the septum is ruptured during fruit development, giving the ap- pearance of being unilocular with the free-central placentation that is so often cited in taxonomic accounts. Anomalous secondary thickening and solitary crystals are reported for the genus. Shea counts are lack- ing, and the identity of the red pigments is not know e genus is considered to be closely related to Seaton L. and Tri- anthema L. (Miiller). REFERENCES: Under family references see BAILLON, BENTHAM & HooKeER, De CANDOLLE, FENZL, METCALFE & CHALK, MULLER, Pax, and Pax & HorrMANN 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 459 Urpan, I. Plantae Haitienses et Domingenses novae vel rariores VII. a cl. . L. Ekman, 1924-1928 lactae. Ark. Bot. 22A(17): 1-115. 1929. re. rubriflora, 15.| Subfamily TETRAGONIOIDEAE Lindl. 5. Tetragonia Linnaeus, Sp. Pl. 1: 480, 1753; Gen. Pl. ed. 5. 215. 1754. Annual or perennial herbs or subshrubs with somewhat succulent, dense- ly papillate [or pubescent], procumbent or ascending, spreading branches. Leaves cauline, mostly alternate, petiolate, exstipulate; blades entire, fleshy, ovate-deltoid to ovate-rhomboid, the apex acute to obtuse, the base mostly truncate to cuneate, papillate [or pubescent]. Flowers axillary [or terminal], solitary or paired [or few in a cymose or spike-like cluster ], nearly sessile or short pedicellate, perfect [or polygamous through reduc- tion of the ovary]. Sepals 4 (3-5), fleshy, united in a tube and adnate to the ovary, color varying in ours from greenish to yellowish within | to reddish]. Stamens 7-13 [1 to many, or in fascicles alternate with the calyx lobes], inserted on the floral tube, shorter than the calyx lobes; of a fruit — note seed of single fertile carpel, sterile locules crowded out, X 4; f. indehiscent horned ruit, X 3; g, seed, X 10; h, seed in section, to show curved embryo partly inclosing endosperm (stippled), X 12. 460 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 anthers subglobose; pollen yellow. Carpels 3-9 [rarely 1 or 2], united; styles 3 or more, equaling the locules, short, erect; ovary half-inferior [to inferior], 3—9-loculate; placentation axile; ovules solitary, pendent, campylotropous (?). Fruit obconic, nutlike, bony, indehiscent, with usual- ly 4 or 5 short, sharp horns [or fruit compound, variously angled to winged or thorny, the thorns often producing axillary buds or sessile flowers in T. dimorphantha Pax! |. Seeds solitary, + pyriform to subreniform; seed coat smooth, membranaceous, light brown; embryo hook or horse-shoe shaped {to arcuate] around an abundant, white, starchy endosperm; cotyledons linear-oblong, fleshy. (Including Demidovia Pall.) LECTOTYPE SPECIES: T. fruticosa L.; see Hitchcock & Green, Int. Bot. Congr. Cambridge, 1930. Nomencl. Prop. Brit. Bot. 159. 1929. (Name from Greek fetra, four, and gonia, angle, in reference to the fruit; a shortened form of the pre-Linnaean name Tetragonocarpos.) — NEW ZEALAND SPINACH. A genus of about 50-75 species distributed chiefly in the Southern Hemisphere. More than half the species are restricted to southern Africa, while a large number are concentrated in southern South America, par- ticularly in Chile. A few species occur in Australia, New Zealand, Poly- nesia, southeastern Asia, and North Africa. The genus is represented in our area by Tetragonia expansa Murr. (T. tetragonoides (Pall.) O. Kuntze), 2% = 32, of subgenus TeTRAGoNoIpEs. Although native to the coastal regions of the southern and western Pacific, T. expansa has been widely cultivated as a leafy vegetable and has become naturalized in many areas after escaping from cultivation. The species is reported from old fields and waste places in North Carolina and Florida, Bermuda, Con- necticut and Massachusetts, and California. The number of specimens available is small, however, and it is not clear whether the species is truly naturalized in these areas or occurs only as a waif after escaping cultiva- tion. None of the available specimens is complete; the lower parts are missing from all of them. The broad distribution of T. expansa around the shores of the Pacific basin is probably explained by Hashima’s finding that the fruits can stay afloat in sea water for more than a month, with the enclosed seeds retaining their viability, thus allowing for long-distance dispersal by ocean currents. From other genera of Aizoaceae Tetragonia is distinguished by its half- inferior to inferior ovaries with a single pendent ovule in each locule, by the absence of staminodial bodies (Adamson), and by its indehiscent, cartilaginous or stony (and sometimes compound) fruits. The genus has been subdivided into two or three sections (Pax & Hoffmann, Post & Kuntze), or two or four subgenera (Fenzl, Adamson). The most recent revision is that of Adamson who recognizes four subgenera: TETRAGONIA (containing seven series), TETRAGONOIDES (DC.) Adamson, ANISOSTIGMA (Schinz) Adamson, and TripuLtocarpus (Moore) Adamson. Only sub- genus TETRAGONOIDES ranges beyond southern Africa. The systematic position and relationships of Tetragonia are not clear. The genus was once considered close to Mesembryanthemum (Baillon, 1970] BOGLE, MOLLUGINACEAE & AIZOACEAE 461 Bentham & Hooker, De Candolle, Pax) on the basis of floral structure (presence of a floral tube, numerous stamens, half-inferior to inferior ovaries), but more often it has been segregated as the type, if not the sole member, of a tribe Tetragonieae (Fenzl, Miiller, Pax & Hoffmann), sub- family Tetragonioideae (Eckardt, Schwantes), or family Tetragoniaceae (Friedrich, Lindley, Nakai). The chromosome complement of the genus is based on eight, with numbers of 2” = 32 reported for Tetragonia expansa, T. crystallina L’Hérit., and T. echinata Ait., and 2n = 96 for T. trigyna Banks & Sol. ex Hook. f. Payer described and illustrated the floral organogeny of Tetragonia expansa in detail, but the morphology and anatomy of the genus have received little attention. Secondary thickening in stems and roots is anomalous, producing xylem and phloem in more or less complete con- centric rings. Large bladder-like water-storage cells occur in the epidermis among smaller unspecialized cells. In dried specimens these become col- lapsed and flattened, giving the plant a hoary appearance. Stomata of the ranunculaceous type are reported. Columnar crystals occur in the pith of T. spicata L. f., and clustered crystals in the cortex of T. expansa Apart from a statement of D’Hubert on the presence of starch grains in the embryo sac of the “Tetragoniées” the embryology of the genus is apparently unknown. Coutinho et al. investigated the structure of chloro- plasts in T. expansa and found them intermediate in form between the Euglena and Aspidistra types, in that they contain few, small, irregular grana and a relatively large amount of stroma lamellae extending through- out the length of the chloroplast. Red pigments of the betacyanin type are reported in Tetragonia crystal- lina. Toxic alkaloids and saponins are also reported for T. expansa (as T. tetragonoides). The economic significance of Tetragonia expansa appears to rest solely on its cultivation as a leafy vegetable, a substitute for spinach. Analysis of the dried plant, however, has shown that the content of anhydrous oxalic acid may attain levels of 10 per cent or greater, thus posing the danger of sharply reducing the absorption of calcium from a diet contain- ing adequate amounts of that element. REFERENCES: Under family references see BAILLON, BENTHAM & Hooxer, D’HUBERT, Ecxarpt, EICHLER, FENZL, FRIEDRICH, LINDLEY, Martin, Metcatre & CHALK, MUtter, Pax, Pax & HorrMANN, PAYER, SCHWANTES, Upnor, Witson, WOHL- PART & Mapry. Apamson, R. S. The South African species of Aizoaceae. II. Tetragonia. Jour. S. Afr. Bot, 21: 109-154. 1955. [37 species in 4 subgenera. | Autes, H. E., C. R. BELL, & A. E. Raprorp. Species new to the flora of North or South Carolina. Rhodora 60: 10-32. 1958. [T. expansa. ] CANDOLLE, A. DE. Origine des plantes cultivées. Paris. 1883; Engl. transl. ed. 2. 462 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 1886; reprinted by Stechert-Hafner. New York. 1959. |T. expansa, 89; notes on discovery and distribution. | Coutinuo, L. M., H. Ruska, M. R. Epwarps, & G. A. Epwarps. The fine structure of chloroplasts of Tetragonia expansa Thunb. Anais Acad. Brasil. Ci. 27: 505-517. 1955. Everist, S. L. A review of the poisonous plants of Queensland. Proc. Roy. Soc. Queensland 74: 1-20. 1964. [T. expansa, as T. tetragonioides. | Hamasuima, S. On the dispersion of seed of Tetragonia expansa Murr. (In ooreel English summary.) Hokuriku Jour. Bot. 6(3): 72-74. 1957. [In- cludes m KINGSBURY, I M. Poisonous plants of the United States and Canada. xiii + 626 pp. Englewood Cliffs. 1964. [T. expansa, Meunier, E, La tétragone. Revue Hort. 89: 282, 283. 1917. [T. expansa Murr., T. implexicoma J. Hook. Moore, S. Le M. Alabastra diversa. Pt. XXXIV. 1. Plantae Rogersianae — VI. Jour, Bot. London 59: a Brig ay [T. dimorphantha segregated as Tribulocarpus Moore, gen. nov., 228. ] Post, T. von, & O. Kuntze es Bae Phan. xlvii + 714 pp. Stuttgart. 1904. [Tetragonia, p. 554. WALKER, J. Tetragonia expansa. Bot. Mag. 50: 2362, 1823. [Color plate. | ARNOLD ARBORETUM Present address: HARVARD UNIVERSITY DEPARTMENT OF BOTANY UNIVERSITY OF NEw HAMPSHIRE DurHAM, NEw HAMPSHIRE 03857 1970] CONDE & STONE, JUGLANDACEAE 463 SEEDLING MORPHOLOGY IN THE JUGLANDACEAE, THE COTYLEDONARY NODE Louis F. CoNpDE AND DoNa_p E. STONE ! DESPITE NUMEROUS sTuUDIEs in the Juglandaceae, the number of genera and their phylogenetic positions are still unsettled. Alfaroa, Annamocarya, Carya, Engelhardia, Juglans, Oreomunnea, Platycarya and Pterocarya were recognized by Leroy (1955), while Manning (Manning & Hjelm- qvist, 1951; Manning, 1961) has continued to emphasize the similarities of Oreomunnea (Engelhardia sect. OREOMUNNEA sensu Manning) with Engethardia and those of Annamocarya (Carya sect. RHAMPHOCARYA sensu Manning) with Carya, thus arguing for six genera. The walnut family is generally conceded to be a very natural assemblage (Lawrence, 1951; Cronquist, 1968), so it is not surprising that generic boundaries are debatable. This is not to say that the taxa are difficult to characterize: all representatives are, in fact, large forest trees which may be recognized by combinations of features that range from differences in bark and wood (Heimsch & Wetmore, 1939) to phyllotaxy (Standley, 1927), flowers (Manning, 1940, 1948), pollen (Whitehead, 1965), inflorescences (Man- ning, 1938), and particularly fruits (Leroy, 1955). It is precisely this comparative wealth of information that makes the Juglandaceae so tempt- ing to the phylogenist. Because of the remarkable diversity of the flowers and fruits and as a result of some preliminary studies on seedling germination, we had reason to believe that seedling morphology would be useful in assessing generic relationships. An earlier study by Davey (1916) on the Amentiferae covered the descriptive aspects of two species of Carya, four species of Juglans, one of Pterocarya, and the monotypic genus Platycarya, A wide variety of cotyledonary nodal types is reported in the literature. Trila- cunar and multilacunar nodes are noted for Carya (Davey, 1916; Lang- don, 1934), and Juglans (Davey, 1916, 1935; Nast, 1941; Gravis, 1943) and unilacunar two-trace nodes for Platycarya and Pterocarya (Davey, 1916). This diversity of nodal types, representing both the most primitive and advanced forms as proposed by Marsden and Bailey (1955) in their study of nodal evolution, as well as the absence of information on the primitive members of the family, namely Alfaroa and Engelhardia, offered additional promise for the current survey. An implicit consideration of our study was an examination of the evidence giving rise to the general belief * Adapted from the M.A. thesis of the first author. The collection of field samples and the laboratory research were supported by National Science Foundation grant GB-5233X. Special thanks are due also to Dr. Delzie Demaree, who provided several important collections, and to Dr. Wayne E. Manning, who offered helpful comments on the manuscript of the thesis. The line drawings are the artistry of Susan Carlton Smith and were sponsored by the Duke University Council on Research, 464 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 that dicotyledonous seedlings have conservative vascular anatomy (Sin- nott, 1918; Foster & Gifford, 1959; Carlquist, 1961; Eames, 1961). MATERIALS AND METHODS Of the six genera and 60 species recognized by Manning (1961), 12 species of six genera were sampled (Taste 1). Seedlings were obtained from field- and greenhouse-grown collections. Field specimens were pre- served in formalin-acetic acid-alcohol (FAA) at the same time herbarium specimens were prepared. Most of the seeds were stratified in a standard greenhouse mixture of sand, peat, and loam and stored at 40°F for 85 to 167 days prior to removal to the greenhouse. Fruits of Engelhardia rox- burghiana, however, were planted in the greenhouse without cold treat- ment. Seedlings in various stages were killed, fixed, and stored in FAA. Material for sectioning was dehydrated in a tertiary butyl alcohol series and embedded in 61°C Tissuemat. Serial transverse sections of most seed- lings were cut at 7 to 12 ». The older and woodier specimens of Carya cordiformis were sectioned at 25 » on a sliding microtome. Sections were stained in safranin or safranin-fast green. Photomicrographs were taken with a Leitz Ortholux microscope, plano objectives, and Panatomic-X sheet film. RESULTS TABLE 1 presents a summary of the cotyledonary nodal survey. This same sequence of species is adopted in the following account of the seedling characteristics. The monotypic genus Platycarya strobilacea Sieb. & Zucc. is a native of eastern Asia and possesses a combination of some of the most primitive and advanced characters in the family (Manning, 1961). The pistillate catkin resembles a spiny cone with the individual bracts remaining at- tached to the inflorescence at the time of fruit dispersal (Manning, 1940). The minute fruits depend on small lateral outgrowths of the bracteoles for wind dissemination. Seed germination is epigeal. The cotyledons are deeply bilobed and both of the primary lobes are again lobed, but in an unequal fashion (Lubbock, 1892; Hickel, 1914; Davey, 1916; FicurE 1). The cotyledonary node is unilacunar two-trace (FicuRE 2), with the traces separated by a very narrow band of parenchyma (FicureE 15). Davey (1916) notes the anatomical similarity of the seedling of this species to that of Pterocarya. The species of Pterocarya, a genus with six species and several varieties known from the Caucasus Mountains and eastern Asia (Manning, 1961), have elongate pistillate inflorescences bearing numerous 2-winged fruits that are well adapted to wind dispersal. Germination is epigeal in the three species investigated, including P. rhoifolia Sieb. & Zucc. (Davey, 1916), P. fraxinifolia (Lam.) Spach (Lubbock, 1892; Hickel, 1914), and P. stenoptera C. DC. (Ficure 3), The thin, foliaceous cotyledons are TABLE 1. Survey of cotyledonary nodal anatomy in the Juglandaceae * COTYLEDONARY TAXA COLLECTION DATA GERMINATION NoDE Platycarya strobilacea Fruit, Kokawa 9 epigeal 1-gap 2-trace Pterocarya stenoptera Fruit, Herbst Bros, 5270 . 1-gap 2-trace Juglans ailantifolia Fruit, Gerardi Nursery hypogeal 3-gap 3-trace Carya aquatica Fruit, Demaree 57353 - 6-gap 6-trace Carya cordiformis Fruit, Stone 1848 <4 4-gap 4-trace Carya laciniosa Fruit, Gerardi Nursery i 5—6-gap 5—6-trace Carya ovata Fruit, Demaree 57494 “i 3-gap 3-trace Carya tomentosa Fruit, Demaree 55007 * 3-gap 3-trace Engelhardia roxburghiana Fruit, Stone GH-112 epigeal i-gap 2-trace Engelhardia mexicana Seedlings, Stone 2143 hypogeal 1-gap 3-trace Engelhardia pterocarpa Seedlings, Stone 1016A - 1-gap 3-trace Alfaroa costaricensis Seedlings, Stone 2156 is 1-gap 3-trace * Voucher specimens have been deposited in the Herbarium of Duke University. [OL6T ‘“ANOLS ® AANOO aVaAOVONVIONLS Sor 466 [vor. 51 JOURNAL OF THE ARNOLD ARBORETUM ERC 4 - FO ter oni ue << SS ‘ ai } Mg so Se anaes FIG Seedlings and cotyledonary so of oa ees 1, seedling of acearoliel strobilacea with epigeous cotyledo 1.5 ap two-trace 1 3; Kat 4. 3 tenop us y : ears og me with epigeous cotyledons, X 0.7. 5, seedling of Alfaroa costaricensis ith hypogeous cotyledons, * 0.4. 6, one-gap ‘three- trace node characteristic of 1970] CONDE & STONE, JUGLANDACEAE 467 deeply bilobed, and, in contrast to Platycarya, each of the primary lobes is again deeply and essentially equally bifurcated (FicurE 3). The cotyle- donary node of P. stenoptera is unilacunar two-trace (Ficures 2, 18) as Davey (1916) reported for P. rhoifolia. In five of eight nodes examined a small central trace was situated between the main pair (Ficure 17) but in only one case did the third trace pass into the cotyledon as a separate strand. Generally it fused with the lateral traces, or with the stelar tissue adjoining the gap, or terminated in the cortical tissue. The genus Juglans is divided into three sections and twenty species with representatives in both the New and Old World (Manning, 1961). The genus is closely related to Pterocarya but in general has more advanced features. The fewer-flowered inflorescence of Juglans is associated with the development of heavy fruits with thick shells and an indehiscent husk. The large cotyledons serve as food storage organs and remain enclosed in the shell below ground at germination (Lubbock, 1892; Rowlee & Has- tings, 1898; Davey, 1916; Pammel & King, 1918; Henrotin, 1933; Nast, 1941; Gravis, 1943; Ficure 8). The cotyledonary node of J. ailantifolia Carr. (= J. sieboldiana, Davey, 1916) is basically trilacunar (FicurE 9), although bilacunar nodes do occur. Occasionally both types are found in the same seedling (FIcurEs 21, 22). Nast (1941) reports that J. regia L. has a four-gap four-trace cotyledonary node, similar to some seedlings of Carya. The genus Carya also has representatives in southeastern Asia and the eastern part of Canada, the United States, and Mexico. By all standards it is the most advanced member of the family. In the aspect of seeds and seedlings the species of Carya are similar to certain species of walnuts. The fruits are large, with a very hard shell and thin to thick dehiscent husk. As in Juglans the fruits are adapted to animal dissemination. The heavy seeds display hypogeal germination (Rowlee & Hastings, 1898; Pammel & King, 1918; FicurE 10) and the cotyledons remain in the shell. Considerable variation was found in the number of traces supply- ing the cotyledons. Carya aquatica (Michx. f.) Nutt. has multilacunar cotyledonary nodes with six gaps and six traces (Ficures 11, 23). Carya cordiformis (Wang.) K. Koch was found to have a four-gap four-trace cotyledonary node (FicurE 12), although Davey’s report (1916) of hexarch vascular poles in the hypocotyl indicates that three-gap three-trace nodes might also be common. Davey also reported octarch vascular poles in the hypocotyl of C. illinoensis (Wang.) K. Koch (= C. olivaeformis, Davey, 1916) so the seedlings probably have a four-gap four-trace node. Carya laciniosa (Michx. f.) Loud. has extremely large fruits with mul- tilacunar cotyledonary nodes supplied with five to six gaps and traces (Ficure 13). Carya ovata ( Mill.) K. Koch is a close relative of C. lacin- seedlings drawn in 5 and 7. 7, seedling of Engelhardia pterocarpa showing hypo- geal germination, x 0.4. 468 JOURNAL OF THE ARNOLD ARBORETUM [voL, 51 ) 5 - 8 a O O on & = i2 & oO ¢@ © oO O Oo a s o ry = # Oo Oo a. O 2 A z a ww O ww je Aah 7 M ds Ms I a 7 \ aaa ae J h ay 2 \ 4 Ny \ en We y a i} \g f a Dee | O O i] a = O Oo @ || ee ®, oO Wr are O O pe 2 4 Pf Sh} O O J \ | rt Oo ‘e) \ \ ? s 8-14. Seedlings and cotyledonary nodes of Scat aps see of Fuga ailantifolia showing hypogeal germination, trilacuna ary nodes of aa: ailantifolia. pecan with hypogeous cotyledons, x 0.4. aquatica. 12, four-gap four-trace node of Carya cordiformis. 13, pentalacunar and hexalacunar nodes of Carya laciniosa. 14, trilacunar nodes characteristic of C arya ovata and C. tomentosa. ling ‘ nar and Car 1970] CONDE & STONE, JUGLANDACEAE 469 iosa but has trilacunar cotyledonary nodes (FIGURE 14), as does the tetra- ploid Carya tomentosa Nutt. (Ficure 14). The genus Engelhardia, as recognized by Jacobs (1960) and Manning, (1966), has two Old World sections and a New World section distributed from southern Mexico to Costa Rica. The unifying and distinctive feature of this genus is the elongate pistillate inflorescence bearing dense clusters of 3-winged fruits. While the fruits vary greatly in size they all seem to have aerodynamic stability for wind dispersal. Germination is of two sorts. Both Old World sections have members with epigeal germination (sect. ENGELHARDIA, E. serrata Bl. [Jacobs, 1960]; E. apoensis Elmer ex Nagel; sect. PstLocarPEar, E. roxburghiana Wall., Ficure 4). Species of section OREOMUNNEA, on the other hand, display the hypogeal germina- tion pattern that characterizes all other New World taxa of Juglandaceae (FicuRE 7). The seedlings examined have nodal patterns correlated with the type of germination. Engelhardia roxburghiana (Ficures 2, 16) has epigeal germination and a one-gap two-trace cotyledonary node similar to Platycarya and Pterocarya. Both E. mexicana Standley (FicurE 19) and E. pterocarpa (Oersted) Standley (Ficures 6, 7) have hypogeal germina- tion with a one-gap three-trace node. The small genus A/faroa is confined to the premontane rain forests (Holdridge, 1964) of southern Mexico, Central America, and northern Colombia. It has many vegetative and reproductive characters in common with Engelhardia sect. OREOMUNNEA but differs markedly in those of the fruit. The well-developed wings of Engelhardia fruit appear only as small basal tabs on fruits of Alfaroa, while the floral tissue enlarges to form a thin to thick indehiscent husk. There is a striking superficial resemblance between the fruits of Juglans and Alfaroa. Alfaroa costaricensis Standley has hypogeal germination (FicurE 5) and a one-gap three-trace cotyledon- ary node (Ficurgs 6, 20). A common pattern of five vascular bundles is seen in the cotyledonary petioles in the Juglandaceae (Stone, in press). In Pterocarya the veins of the lamina form three double-stranded traces in the distal end of the petiole (Ficure 24). Near the middle of the petiole the central trace separates, one half merging with each lateral trace. The two resulting bundles in the proximal end of the petiole enter the stele at a single gap. In Platycarya the levels of branching and fusion are slightly different but the basic pattern is the same. Davey (1916) reports three double-stranded traces in the petioles of Carya cordiformis (= C. amara Nutt.), Juglans nigra, J. ailantifolia, J. cinerea, and J. regia. In these species the three traces do not anastomose in the proximal end of the petiole but enter the stele at three separate gaps. The four-gap four-trace node of J. regia reported by Nast (1941) is a variation of the same pattern. Again, three double- stranded traces are formed in the petiole but the central trace divides in two in the basal end of the petiole and four traces enter the stele at four 470 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 gaps. In a cleared cotyledon of Juglans olanchana Standley the double central trace fused with one of the lateral bundles entering the bilacunar node (FIcuRE 25). DISCUSSION In 1914, after a broad survey of the angiosperms, Sinnott proposed the trilacunar node as the primitive type from which unilacunar and multila- cunar nodes were derived. With this pattern as the basis, subsequent in- vestigators have used nodal anatomy as an aid in determining the sys- tematic position of families within orders and genera within families (Bailey & Howard, 1941; Money, Bailey & Swamy, 1950; Swamy & Bailey, 1950; Canright, 1955). By 1955 sufficient information had ac- cumulated to spark a reevaluation of Sinnott’s sequence of nodal types (Marsden & Bailey, 1955; Canright, 1955). The prevalence of the uni- lacunar two-trace node in lower vascular plants, primitive woody Ranales, and in the cotyledonary node of angiosperm seedlings has led to the recog- nition of this nodal pattern as the basic type. This revised concept of ifford, Ts , Rt RY . Y bi Ld s mk _ 2 2 aenenrninennliiinwiniais EXMma ee. Ficures 15-18. Cotyledonary nodes of epigeous J e. 15, one-gap two-trace node of Platycarya strobilacea, X 47 , one-gap two-trace node of Engelhardia roxburghiana, X 55. 17, one-gap two + one-tr e of Ptero- i stenoptera, * 55. 18, one-gap two-trace node of Pterocarya stenoptera, 1970] CONDE & STONE, JUGLANDACEAE 471 Recently the validity of the Marsden-Bailey model of nodal evolution has been questioned (Pant & Mehra, 1964; Benzing, 1967; and Cron- quist, 1968). In particular, Benzing’s studies on foliar nodes of the woody Ranales suggest that those families with unilacunar two-trace nodes are characterized by specialized floral features and derived decussate phyl- lotaxy (i.e. Chloranthaceae, Monimiaceae, Austrobaileyaceae). Also, as Carlquist (1961) has noted, it is curious to find the unilacunar two-trace condition in foliar nodes of such advanced families as the Labiatae (leaves opposite or whorled), Solanaceae (leaves alternate or becoming opposite at or near the inflorescence: Lawrence, 1951), and Verbenaceae (leaves opposite or whorled). “The possibility that two-trace unilacunar nodal anatomy evolved in association with opposite phyllotaxy from an ancestral condition characterized by alternate arrangement . . . is quite likely” (Benzing, 1967). Of course the Marsden and Bailey model of nodal evolution does not rely exclusively on the patterns of foliar nodes. One of the mainstays of the argument rests on the relatively conservative nature of the cotyledon- ary node. In a survey of the seedlings of 99 families of dicotyledons, Bailey (1956) found that 77 percent had an even number of vascular Strands at the cotyledonary node, and 60 percent were unilacunar two- trace. The trilacunar cotyledonary node was found in about 10 percent of the species but not a single multilacunar attachment was encountered; hence the conclusion that cotyledonary nodal anatomy is more conserva- tive than foliar and the unilacunar two-trace node is primitive. Benzing (1967) points out, and rightly so, that “The anatomy of cotyledonary nodes does not necessarily reflect ancestral conditions in the mature stem; the unique seedling morphology and decussate insertion of the cotyledons makes this unlikely.” If we are able to accept the available information at face value, it seems likely that evolution of the foliar node has progressed independently of the cotyledonary node. The usefulness of cotyledonary nodal anatomy in phylogenetic studies depends, of course, on its degree of conservativeness. Certainly Bailey’s survey (1956) suggests that cotyledonary attachment is conservative, but unfortunately few details are provided on families or species examined and no information on cotyledonary size, position, or activity. This informa- tion is relevant because there is circumstantial evidence that relates coty- ledonary development to physiological activity of the seedling (Compton, 1912). A rapidly growing seedling with a large cotyledonary food supply or one that has large cotyledons that are photosynthetically active might be expected to have the best developed vascular system (Hill & de Fraine, 1913). Boureau (1954) and Carlquist (1961) note that complexity of vascularization does not parallel size of seedlings, but it must be recognized that vascular supply and seedling demand are relative. Functional sig- nificance of vascular supply would lead to adaptive changes that involve both increases and decreases in vascular complexity. Fitchia, an excep- tional member of the Compositae with enormous epigeous cotyledons and pentalacunar nodes, seems to represent a good example of increased vas- 472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 cularization (Carlquist, 1957). Decreased vascularization associated with reduced functional demand might be the case in two epigeous members of the Fagaceae (Carlquist, 1961, after Gravis, 1943). The large foliaceous cotyledons of Fagus sylvatica have multilacunar nodes, whereas the smaller, fleshy green cotyledons of Carpinus betulus (Lubbock, 1892) are vas- cularized with a highly reduced unilacunar system e evidence at hand suggests that daacitieteation of seedlings is com- Bie to the situation in other parts of the plant. In those groups where selection pressure is intense for a particular characteristic, conservatism is shed. The Juglandaceae offer additional proof of the adaptability of ee morphology and anatomy. asic vascular supply to the cotyledons in the Juglandaceae is rela- Ve uniform. As we have been able to confirm for Pterocarya, Platy- carya, Juglans, and Carya (Davey, 1916; Nast, 1941), the veins from the aa" ozs we ee 5 - ¢ tak: SiS tee, eS! <%. = 2 Ficures 19-23. Cotyledonary nodes of (oe etc 19, one-gap three-trace node of Engelhardia mexicana, X 104. 20, one-gap three-trace node of ievege costaricensis, X 75. 21, two-gap two-trace node of Juglans ailanti- oul , three-gap three-trace node of he oes ailantifolia, * arrows 1970] CONDE & STONE, JUGLANDACEAE 473 four cotyledonary lobes merge to form five traces in the distal region of the petiole. In Platycarya and Pterocarya the central double strand fuses with the two laterals to enter the single gap as two compound traces (FicurRE 24). The bilacunar, trilacunar, and multilacunar nodes of Jug- lans and Carya probably result from varying degrees of separation and fusion of the double-stranded units (F1GURE 25). The presence of two to a maximum of six gaps supports this interpretation. The spatial separa- tion of the strands is certainly correlated with the larger size of the nodal region and petiole base. Sinnott (1918) has noted similar variations of the double-stranded central trace in other angiosperms. In the hypogeal seedlings of Carya and Juglans, and to a lesser extent the New World Engethardia and Alfaroa, the cross-sectional area of the vascular bundles is noticeably larger than in the epigeal seedlings. This may well be the result of extensive branching of the vascular system in the massive cotyle- dons of the hypogeal forms (Stone, in press). It is at the level of the cotyledonary node that the Juglandaceae show such remarkable diversity of vascular patterns, patterns which may be correlated with seed and seedling size. The small seedlings of Platycarya (Ficure 1), Pterocarya (Ficure 3), and Engelhardia (sect. ENGELHARDIA and sect. PstLocarPEAE; FicurE 4) have epigeous, foliaceous cotyledons with unilacunar two-trace nodes. The larger hypogeal seedlings of Engel- hardia sect. OREOMUNNEA (FIGURE 7) and Alfaroa (Ficure 5) are unila- cunar three-trace; those of Juglans (FicurE 8) and Carya (FicuRE 10) have bilacunar, trilacunar, and multilacunar nodes. These three levels of nodal complexity generally parallel proposed generic phylogenies derived from independent assessment of wood (Heimsch & Wetmore, 1939), inflorescences (Manning, 1938), flowers (Manning, 1940, 1948), pollen (Whitehead, 1965), and fruits (Leroy, 1955). The genus Engelhardia, for example, as viewed by Manning (1966), has three sections: sect. ENGELHARDIA and sect. PstLOcARPEAE are Old World; sect. OREOMUNNEA is New World. Engelhardia roxburghiana of sect. Pstto- CARPEAE, with its primitive pollen and characteristics of both sect. ENGEL- HARDIA and sect. OREOMUNNEA, has been suggested as the intermediate or connecting link (Manning, 1966; Stone, 1968). Information from cotyledonary nodal anatomy is compatible with such a sequence. The Old World sections of Engelhardia both have primitive epigeal germination with unilacunar two-trace nodes. The New World sect. OREOMUNNEA is a derived member with hypogeous cotyledons and unilacunar three-trace nodes. The end point in this series is the genus Alfaroa. By all standards it is closely related to Engelhardia sect. OREOMUNNEA but unlike the Engelhardias it has a large, wingless, nutlike fruit (Manning, 1949, 1959; Stone, 1968). In the few species we have examined, Alfaroa has hypogeal germination and unilacunar three-trace nodes similar to Engelhardia sect. OREOMUNNEA. The increase in nodal complexity is evident also in the Pterocarya- Juglans alliance. Pterocarya, which is generally acknowledged to be the more primitive of the two (Manning, 1938, 1940, 1948, 1949; Whitehead, 1970] CONDE & STONE, JUGLANDACEAE 475 1965), has small 2-winged fruits with epigeal germination and unilacunar two-trace nodes. /uglans, on the other hand, is noted for its large, wingless fruits, hypogeous cotyledons, and trilacunar three-trace cotyledonary nodes. The genera Platycarya and Carya are more difficult to place on the phylogenetic tree. Platycarya has primitive floral features (Manning, 1938, 1940, 1948) but advanced wood (Heimsch & Wetmore, 1939) and distinctive pollen with pseudocolpi (Whitehead, 1965). This monotypic genus is usually treated as an isolated element derived from Engelhardia stock. Seedling morphology is consistent with this view. The fruits are small with minute wings and the seedlings have epigeous cotyledons and unilacunar two-trace nodes similar to those of Engelhardia and Pterocarya. Without doubt Carya is the most advanced member of the family. The fruits are large, germination is hypogeal, and cotyledonary nodal anatomy ranges from three-gap three-trace to six-gap six-trace, The general similar- ity of fruit and seedling characteristics shared with Juglans suggest a close relationship, but the totality of evidence (Hjelmqvist, 1948; Manning, 1948, etc.) seems to favor the idea that Carya and the Pterocarya-Juglans alliance had separate evolutionary pathways stemming from the Engel- hardia plexus (Whitehead, 1965). The remarkable diversity of cotyledonary nodal anatomy may at first seem inconsistent with the view that the genera of Juglandaceae comprise a very natural plant family (Lawrence, 1951). On closer inspection, how- ever, it is apparent that the nodal attachments are part of the syndrome of characters associated with patterns of adaptive radiation. Vegetatively the family has been relatively conservative. All members are trees of temperate or cool tropical areas with pinnately compound, resin-dotted, aromatic leaves (Manning, 1961). Diversification of the genera has pro- ceeded through the elaboration and adaptation of flowering and fruiting structures. Modifications of the inflorescences, staminate flowers, and pistillate flowers appear to be refinements for wind pollination and seed dissemination. The primitive fruits were probably small and had minute wings like those of Platycarya (Manning, 1940) or Rhoiptelea (Manning, 1961). From this stage of development fruit dispersal by wind and animals evolved along several generic lines. In Engelhardia the development of larger wings led to larger fruits, and eventually a shift from epigeal to hypogeal germination with a concomitant increase in vascular complexity. Once hypogeous cotyledons and complex nodes were fixed the exploitation of new niches was pursued in Alfaroa by the loss of wings, development of a husk, and increase in size of fruit (Stone, 1968). Similar adoption of even more complex nodes is coupled with the advent of hypogeal germina- tion in Juglans and Carya. The general pattern of evolution of the jug- landaceous seedling shows a one to one correspondence between the adapta- tion to hypogeal germination and increased nodal complexity. The Ficures 24, 25. Drawings of cleared cotyledons of Juglandaceae. 24, vascular anatomy of a one-gap two-trace “cotyledon” of Pterocarya stenoptera, X 5, 25, vascular anatomy of a two-gap two-trace “cotyledon” of Juglans olanchana (Stone 2701), x 3. 476 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 enlarged nodal region of hypogeal seedlings may well have been advan- tageous in overcoming the “spatial and other difficulties in the development of multilacunar nodes where the leaves are opposite” (Bailey, 1956). Thus, the conservatism of seedling anatomy in the dicotyledons may well be related to the constraints placed on nodal development in most groups by the prevalence of primitive, epigeal germination. LITERATURE CITED BarLey, I. W. Nodal anatomy in retrospect. Jour. Arnold Arb. 37: 269-287. 1956. & R. A. Howarp. The comparative morphology of the Icacinaceae. I. Anatomy of the node and internode. Jour. Arnold Arb. 22: 125-132. 1941. Benzinc, D. H. Developmental patterns in stem primary xylem of woody Ranales. II. Species with trilacunar and multilacunar nodes. Am. Jour. Bot. 54: 813-820. 1967. BoureEAv, E. Anatomie végétale. Vol. 1. Presses Univ. de France, Paris. 1954. CANRIGHT, J. E. The comparative morphology and relationships of the Mag- noliaceae. IV. Wood and nodal anatomy. Jour. Arnold Arb. 36: 119-140. 1955. CARLQUIST, e The genus Fitchia (Compositae). Univ. Calif. Publ. Bot. 29: 1-143, 1957. gone plant anatomy. Holt, Rinehart, & Winston, New York. 961. Compton, R. H. An investigation of Ae seedling structure in the Leguminosae. Jour. Linn. Soc. Bot. 41: 1-122. 1 Cronguist, A. The evolution and Seem of plants. Houghton Mifflin Co., Boston. Davey, A. J. Seedling anatomy of certain Amentiferae. Ann. Bot. 30: 575-599 1916. . Note on the structure of the epicotyl in Juglans nigra. New Phytol. 34: 201-210. 1935. Eames, A. Morphology of the angiosperms. McGraw-Hill Book Co., New York. 1961. Foster, A. S., . M. Gtrrorp, Jr. Comparative morphology of vascular plants. W. H. pee & Co., San Francisco. Gravis, A. Observations anatomiques sur les embryons et les plantules. Lejeunia Mém. 3: 1- ree 1943. HErmscuH, C., Jr., . H. Wermore. The significance of wood anatomy in the taxonomy as ie Juglandacese. Am. Jour. Bot. 26: 651-660. 1939. Henrotin, M. Introduction a l'étude de bourgeons series: eens des tiges dans le genre Juglans. Bull. Soc. Bot. Belg. II. 66: 65-85. Hicket, R. Graines et plantules des Angiospermes. Chez l’auteur, ade 1914 rel Dendrologi Forestiére. Encyclopédie Economique Sylviculture 5: 1- 5, H eos a Studies on the floral er ae and phylogeny of the Amenti- ferae. Bot. Not. Suppl. 2: 5-171. 1948 HI.1, T. G., & E. DE FraIne. A consideration of the facts relating to the struc- ture of seedlings. Ann. Bot. 27: 257-272. 1913 Hotpripce, L. R. Life zone ecology. Tropical Science Center, San José, Costa Rica. 1964 1970] CONDE & STONE, JUGLANDACEAE 477 nae M. Juglandaceae. J Flora Malesiana I. 6: 143-154. 1960, Lancpon, L. M. Embryogeny of Carya and Juglans, a comparative study. Bot. en ‘06: 93-117. 1934. LawkENcE, G. H. M. Taxonomy of vascular plants. MacMillan Co., New York. 1951 Leroy, J. — as sur les Juglandaceae. Mém. Mus. Paris II. B (Bot.) 6: 1- 246. 19 Lusgock, J. contribution to our knowledge of seedlings. Vol. 2. Paul, Trench, Triibner, & Co., London. 1892 MANNING, W. E. The morphology of the flowers of the Juglandaceae, I. The inflorescence. Am. Jour. Bot. 25: 407-419. 1938; II. The pistillate flowers and fruit. Jbid. 27: 839-852. 1940; III. The staminate flowers, Ibid. 35: 606-621. 1948. . The genus Alfaroa. Bull. Torrey Bot. Club 76: 196-209. 1949. . Alfaroa and Engelhardia in the New World. Ibid. 86: 190-198. 1959, Juglandales. 7m Encyclopedia Biological Sciences, Reinhold Publ. Corp., New York. 1961. ———. New combinations and notes on te a (Juglandaceae) of the Old World. Bull. Torrey Bot. Club 93: 34-52. 1966. . HJELMQvist. Annamocarya, Rhamphocarya, and Carya sinensis. Bot. Not. 4: 319-330. 1951. Marspen, M. P. F., & I. W. Battey. A fourth type of nodal anatomy in dicoty- ledons, illustrated by Clerodendron trichotomum Thunb. Jour. Arnold Arb, 36: 1 1955. Money, z, i I. W. Battey, & B. G. L. Swamy. The morphology and relation- ships of the dee Jour. Arnold Arb. 31: 372-404. 1 Nast, C. G. The embryogeny and seedling morphology of Juglans regia L. Lilloa é 163-205. 1941. PAMMEL, L. H., & C. M. Kina. The germination of some trees and shrubs and their juvenile forms. Proc. Iowa Acad. Sci. 25: 291-340. 1918. Pant, D. D., & B. Menra. Nodal anatomy in retrospect. Phytomorphology 14: 384-387. 1964. Row eg, W. W., & G. T. Hastincs. The seeds and seedlings of some Amenti- ferae. Bot. Gaz. 26: 349-353. 1898. Stnnott, E. W. Investigations on the phylogeny of the angiosperms. I. The anatomy of the node as an aid in the classification of angiosperms. Am. Jour. Bot. 1: 303-322. 1914. . Conservatism and variability in the seedling of dicotyledons. Jbid. 5: 120-130. 1918. STANDLEY, P. C. The American species of Engelhardtia. Trop. Woods 12: 12-15. 1927 STONE, D. E. New World ehoupeciin F a new species of Alfaroa from Mexico. Am. Jour. Bot. 55: 477-484. 1968. . Evolution of cotyledonary and nodal vasculature in the Juglandaceae. Am. Jour. Bot. Swamy, B. G. L., & I. W. BAI LEY. Sarcandra, a vesselless genus of the Chlor- a ated, Tour: Arnold Arb. 31: 117-129. 1950. WuireHeap, D. R. Pollen morphology in the a II. Survey of the family. Jour. Arnold Arb. 46: 369-410. 1965 DEPARTMENT OF BOTANY DvuKE UNIvErRSI Duruam, NortH CaroLina 27706 478 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 THE VASCULAR SYSTEM IN THE AXIS OF DRACAENA FRAGRANS (AGAVACEAE) 2. DISTRIBUTION AND DEVELOPMENT OF SECONDARY VASCULAR TISSUE M. H. ZIMMERMANN AND P. B. TOMLINSON IN THE FIRST PAPER of this series, we described the distribution and origin of primary vascular bundles in Dracaena fragrans (L.) Ker-Gawl. (Zimmermann & Tomlinson, 1969). We showed that the principle of primary vascular structure and development in Dracaena is similar to that of other monocotyledons we had studied (Zimmermann & Tomlinson, 1965, 1967, 1968). However, like a few other woody monocotyledons, Dracaena differs from palms because it has a vascular cambium which produces secondary vascular tissue. In this way Dracaena resembles a woody dicoty- ledon although the secondary tissue produced from the monocotyledonous kind of cambium differs from that of most dicotyledons because it matures as a series of anastomosing vascular bundles within a matrix of secondary parenchyma (Tomlinson & Zimmermann, 1969). In this present paper we examine the distribution and development of secondary tissues and its relation to primary vascular development. The structural relation between primary and secondary vascular bundles in Dracaena was one of the earlier problems of monocotyledonous anatomy to be investigated. It was studied by von Mohl (1824, 1849) and his contemporaries, notably de Mirbel (1845). The observations and inter- pretations of these early botanists with regard to the development of the Dracaena type of stem were inconclusive and often contradictory. Con- tinuity between primary and secondary bundles was indeed described by a number of these early authors and in this sense our own observations can be said to do no more than confirm earlier work. However, we now know that a knowledge of secondary vascular development must be based upon an understanding of primary vascular development within the crown. This information had not been available until our own analysis of the course of procambial strands in the developing crown of Dracaena was presented. With this understanding we can now discuss in a meaningful way the distribution of secondary vascular tissues in monocotyledons. Most of the background information to this paper, including growth habit and general histology of Dracaena has been presented in the first paper of this series (Zimmermann & Tomlinson, 1969). MATERIALS AND METHODS Sources of materials, methods of sectioning and serial analysis by ciné microphotography have been dealt with in the first paper. The relation between primary and secondary tissues was investigated in the sapling 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 479 axis of a plant grown from seed to an overall height of 75 centimeters within three years. During this time it had not flowered, and remained unbranched. The basal 12 centimeters of the aérial stem of this sapling was fixed for investigation. Outgrowth of lateral buds below a terminal inflorescence and _ their method of establishing vascular continuity with a parent stem was studied, following a flowering period in December, 1968. Shoots were dissected at intervals after flowering, fixed in FAA, embedded in paraffin and sec- tioned serially. Detailed analyses of two successive stages were made by the cinematographic method. DISTRIBUTION OF SECONDARY TISSUE Establishment growth in seedlings. Most monocotyledons have no secondary growth, and in order to produce a stable axis thickening growth must precede growth in height (Schoute, 1903). In the majority of arborescent monocotyledons, represented by most palms, the leafy before growth in height begins with longer internodes. 1b. e found i danus an few palms such as Socratea: growth in height begins early, the Widening stem is supported by prop roots. 480 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ye iti. ( Pon i ee FIG. 2 Ne POST - SEEDLING 1 ai GROWTH ite, oie ~| es ie > —_—_> ——_—_ a b c d mas Lr i 7 «f FIG. 3 | on Tale a ae Om —~ ae SYMPODIAL (' GROWTH od ——-» ——— a 1b c d ican aes FIG. 4 geo Er fe ) GROWTH AFTER DECAPITATION a b l 1c Ficures 2—4. Distribution of secondary vascular tissue (black) in Dracaena fragrans. 2a—d, Establishment growth during the post-seedling stage. Growth in height begins early, the stem increases in width by secondary growth as it grows in height. Secondary vascular bundles are initially produced in close association with primary bundles (FicurE 2a, b). Eventually, secondary growth is “left 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 481 crown of seedlings remains at ground level and enlarges gradually. Growth in height does not begin until crown size and stem diameter have reached that of the adult tree. During this time of “establishment growth” height growth is minimal, internodes are very short, and an underground obconical axis develops (FicurE 1a). In a few palms (e.g. Socratea) and in Pan- danus establishment growth is associated with growth in height because internodes elongate, but the resulting axis is stabilized mechanically by supporting prop roots (FicurE 1b). During this period of establishment growth the number of axial vascular bundles increases by a process of growth which has been described for the palm Rhapis in one of our earlier papers (Tomlinson & Zimmermann, 1966). Conifers and dicotyledons do not depend upon primary establishment growth because secondary growth provides for increase in thickness as the stem grows in height. As the tree grows taller its stem thickens propor- tionally so that mechanical stability is achieved. The seedling growth of monocotyledons of the Dracaena-type is similar in this respect. In Dracaena, germination of seeds is hypogeal and is followed by the development of the seedling axis characteristic for monocotyledons. This involves an initial widening of the crown by establishment growth as is shown in Ficure la, and a multiplication of primary vascular bundles which we presume is similar to that described for Rhapis (Tomlinson & Zimmermann, 1966) although we have not yet examined this in detail. dicotyledons, i.e. by secondary stem thickening while the overall height is increasing. This is shown in Ficure 2a—d, Although this second phase of establishment growth is similar to that in dicotyledons, because it in- volves the production of secondary tissue, it differs from it in that the number of primary bundles in the shoot also increases (FicureEs 2a, b). We shall discuss later how this is possible. At the moment it is important to state that during the early post-seedling growth secondary vascular bundles are produced in close association with primary growth of the crown region (FIGURE 2a), but at a later stage of shoot growth (Ficure 2d) primary growth is very similar to that of palms and involves no closely associated secondary bundles. This latter mode of primary growth formed the substance of the first paper of this series (Zimmermann & Tomlinson, Establishment growth in older parts of the plant. Pandanus, behind” (Ficure 2c, d). 3a—d. Production of secondary vascular bundles follow- ing flowering (establishment growth in the sympodium). A terminal inflores- cence is produced (Ficure 3a), declining apical dominance enables a lateral bud to grow out (Ficure 3b). The outgrowing lateral “has access” to only few Primary vascular bundles but regenerates primary vascular tissue (white) by part (Ficure 3b-d). 4a-d. Establishment growth of lateral shoot after de- capitation of the parent axis. Regeneration of the primary vascular tissue by exogenous growth produces primary bundles which are secondary (black) in their lower part. 482 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Dracaena and numerous other woody monocotyledons grow sympodially. The primary axis produces a terminal inflorescence. Apical dominance thus declines and one or more of the lateral vegetative buds below the inflo- rescence grows out. The morphology of this process has been described by earlier botanists, notably by Schoute (1903, 1906), and is summarized in our introductory paper (Tomlinson & Zimmermann, 1969). If more than one lateral bud grows out the axis will fork. It is indeed largely by this mechanism that many monocotyledonous trees become branched. In order to simplify our description, we will consider here a situation in which only one vegetative bud grows out so that the sympodium remains single. Ficure 3 illustrates the sympodial growth of a Dracaena axis in a simplified manner. In F1cuRE 3a the shoot has reached the flowering state. In Ficure 3b the terminal inflorescence is beginning to age and a lateral bud begins to grow out. Now it is perfectly clear that the inflorescence is supplied by a large number of primary axial bundles but these are “‘lost” from the sympodium because growth of the inflorescence is determinate. This means that the newly developing vegetative shoot has to undergo a period of establishment growth in order to restore the approximate equiv- alent number of vascular bundles. In Dracaena this process again in- volves secondary vascular tissue (FicurE 3c). Hence, at the time the new vegetative crown is fully re-established, the number of primary vascular bundles is fully restored. An unbranched shoot which flowers repeatedly over a period of time acquires a segmented appearance (cf. Figure 3 in Zimmermann & Tomlinson, 1969). Two points will be important as we discuss secondary growth later in this paper. First, establishment growth of the new vegetative shoot of a sympodium involves secondary vascular tissue. Second, this secondary vascular tissue (shown black in Ficure 3b— d) is not necessarily continuous with the secondary vascular tissue towards the base of the tree. The kind of establishment growth found in flowering shoots also occurs in buds released from apical dominance by other ways, as in the decapita- tion of a shoot (Ficure 4) or on the upper side of a leaning branch. Photographic illustrations of this kind of shoot are given in Tomlinson & Zimmermann (1969, Figures 14 and 15). Bundle continuity in the post-seedling axis. FicureE 5 shows longitudinal and transverse sections of the basal portion of a young Dra- caena plant, drawn to scale. As a result of establishment growth the di- ameter of the primary vascular cylinder had increased from 2 mm. at ground level to 12.5 mm. at 11 cm. height. An overall conical stem shape was maintained by secondary growth although the primary vascular cylinder itself is an inverted cone. An 8 cm. long piece from between 3 cm. and 11 cm. height was sectioned and analyzed in detail. The results are shown in Ficure 6. The sections show that the number of primary vascu- lar bundles also increases with height during establishment growth. The diameter of the primary cylinder at the base of the piece shown in FIGURE 6 is about 7.5 mm. and contains 290 primary vascular bundles. At the top, the diameter of the primary vascular cylinder is 12.5 mm. and contains 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 483 11 cm 10 GROUND HEIGHT ABOVE Coen ANALYZED PIECE Ficure 5. Longitudinal and transverse sections (0, 2, 3, 5, and 11 cm. above ground) of the basal part of the analyzed post-seedling Bee showing establish- ment growth by production of secondary tissue, emt * scale. The prim ne stem diameter increases from 2 mm. at ground lev 12.5 mm. at 11 height. The 8 cm.-long piece between 3 cm. and 11 ¢c age ight was used for the analysis shown in Figure 6. Secondary vascular oa is indicated by stippling. 540 primary vascular bundles. Therefore, the question is: where do the 250 new primary vascular bundles come from? Before we can answer this question we may briefly recapitulate our answer to the same problem in the post-seedling stage of Rhapis. There we found that the number of vascular bundles increased because each lower leaf trace produced more than one vertical-(axial-)bundle branch. Thus the number of axial bundles is augmented above the departure of each leaf-trace. In Dracaena .—) _— + —— ¢ sm + —STEM | ee a ee PK ’ pm r 7) CORTEX nt L3 de cce h e 0 1 2 3 4 5 6 ? cm Ficure 6. Course of vascular bundles in an 8 cm.-long piece of a stem produced ee post-seedling og moi growth. nly the central part of the primary stem gre w endogenously as do palms. An example of an a axial-(vertical- )bundle branch resulting from this kind of growth is as wn at Y. Establishment growth ca in number of primary vas- cular bundles) is by exogenous growth. greet ples of these exogenous (outer) bundles are secondary in their ee part. Primary bundles are shown as solid black or, secondary bundles as dotted lines. The point of transition from primary to secondary is marked by an arrow at X. Note that all bundles are shown in a single radial plane. The axis is fore- shortened about twice. t8r WOLANOTUVY GIONUV AHL AO TVNUNOL TS “TOA | 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 485 it is probable that this type of primary establishment growth takes place during the very early phases of seedling growth. Morphological evidence suggests this, although we have not yet analyzed such early stages anatom- ically. Post-seedling establishment growth of the axis shown in FIGURE 6 is very different and involves secondary vascular bundles. It can be seen in FicurE 6 that some of the primary bundles of the upper part of the stem are continuous as secondary bundles in the lower part. This is why there are more primary bundles in the upper stem part. The develop- mental significance of this finding will be discussed later, after the an- atomical situation has been described in more detail. FicuRE 6 is based upon accurate measurements of bundle positions in a young post-seedling axis. As in previous publications radial distances from the stem center are shown regardless of the side of the stem on which they are located. In other words, the whole stem is represented in a single radial plane. One bundle is shown which is entirely primary (FIGURE 6, at Y), three bundles (and a short section of a fourth one) are shown which are primary in their upper part and secondary in their lower part (FicurE 6, at X). The bundles in Ficure 6 were plotted on graph paper while being followed individually through a series of transverse sections. Followed towards the base of the stem each approaches the periphery of the primary vascular cylinder and “becomes” a secondary bundle. When followed further down as a secondary bundle their identity was main- tained, but only in an arbitrary way, because secondary bundles anasto- mose extensively. hen the analytical film of the axis used to produce Ficure 6 is projected so that bundles are followed downwards, one observes that all peripheral primary bundles “sink” into the secondary vascular cylinder, thus “becoming” secondary vascular bundles. When projected in reverse (upwards), secondary bundles “emerge” as primary ones and “move” towards the stem center. Ficure 7 illustrates four positions of the same vascular bundle at the point of transition. At the highest level (F1cuRE 7a) the bundle is collateral (primary) in the distribution of its vascular tissues; at the lowest level (FIGURE 7d) it is amphivasal (secondary) and at intermediate levels (F1cURE 7b, c) an “intermediate” condition can be recognized. These indicate that part of the vascular differentiation within a procambial strand was carried out at a primary phase of growth and completed in a secondary phase. Bundle continuity in older stem parts. Establishment growth of an outgrowing lateral bud following decapitation of the parent stem (FIGURE 4a-d) is anatomically very similar to that of the post-seedling stem. The secondary bundles are continuous from the daughter into the parent axis. They extend down and around the parent axis in a basal direction. In this type of union it is important to realize that secondary bundles are not continuous with primary ones at their lower extremity. They anastomose extensively among themselves, but as one descends the secondary vascular cylinder gets thinner and finally disappears almost entirely. Vascular con- tinuity between this more recent secondary cylinder and the primary 486 JOURNAL OF THE ARNOLD ARBORETUM VOL. 51 ws fe tate! Shed Sane : e:* >: -** oa © &* Fi i ee “4@, ae a Th. ee 2 ee. 2 a , ae, x7 | pt * GuRE 7. A single exogenous vascular bundle (arrow) shown in the area of oot from primary to secondary tissue. In the highest position Rehabs 7a, m. above the bottom section, FIGURE 7d) the bundle is entirely primary. I the bation: section (Ficure 7d) it is entirely secondary, Int erinedinte positions are seen in FiGuRE 7b (2.8 mm. above 7d) and Ficure 7c (1.6 mm. above 7d). 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 487 cylinder below is given by radially-running leaf traces which connect to the secondary bundles via short upwardly-directed bridges. Leaf traces continue to elongate radially as long as leaves are still attached to the stem. We do not yet know if their elongation continues after the leaves have fallen off the stem. We assume that the attachment of lateral outgrowing vegetative buds on the upper side of leaning stems (illustrated in Figure 15, Tomlinson & Zimmermann, 1969) is similar to that after decapitation, but we have not yet investigated this situation in detail. The vascular anatomy of renewal shoots of a sympodial branch is some- what more complicated. Though we have found, without question, the same basipetal transition of primary into secondary vascular bundles, there are many primary bundles of the daughter shoot which are contin- uous with primary bundles of the mother shoot. This means, of course, that growth of the lateral shoot begins sufficiently early that it is possible for numerous primary bundles of the branch to develop in continuity with primary bundles of the parent shoot. For this reason, there is no sharp distinction between primary and secondary vascular tissues at the level of union when maturation is complete. This contrasts with the sharp boundary between primary and secondary vascular tissue in both the post- seedling and decapitated axis. LEAF AND BUD TRACE Foliage leaves in Dracaena fragrans are long lived, and can persist for two to three years as functional assimilating organs. All leaves subtend a bud, which remains minute and strongly suppressed, but retains its ability to grow as can be seen in decapitated or leaning shoots. Buds are first recognizable as a meristem in the axil of a leaf in position P15 and bud traces are differentiated at position P16. These traces may be satellites (branches from a major leaf trace) or axial bundles directed into the bud (Zimmermann & Tomlinson, 1969, p. 381). Bud traces, although few and small, are long persistent, Both foliage leaves and buds in older stems are connected across the secondary cylinder by a series of traces which owe their continuity with the central cylinder to their primary origin. Traces are not ruptured by the activity of the cambium which they cross and it is, therefore, of some interest to understand how they can retain a translocating function across a zone of tissue which increases in thickness continually by meristematic activity. The mechanism is simple and not essentially different from the development of leaf traces within the basal meristems of leaves in all large monocotyledons. Continuity of conducting tissues along the trace is main- tained by the continued differentiation of extensible elements of xylem and phloem. Traces are more or less horizontal within the zone of secondary tissue. Within the cambial zone undifferentiated elements persist around the trace. These cells are somewhat longer than the cambial elements proper which, 488 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 together with recent cambial derivatives, make up the cambial zone. gi in length of the trace, to accommodate for secondary increase in the stem diameter, is the result of occasional vertical divisions in these or Genie initials (i.e. divisions transverse to the long axis of the leaf trace). Other divisions are horizontal (the new cell wall is parallel to the long axis of the trace) so that the trace grows in thickness. Existing proto- xylem and protophloem elements are eventually ruptured by the extension RE 8. fa a: arg through a stem of Dracaena fragrans outside the seca vascular tissue. A median leaf trace (LT) is cut transversely, above it numerous racenetoaee Tanek (two 0 them marked with arrows), connecting a dormant axillary bud with the primary vascular tissue. 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 489 of the trace but new ones are continually generated to take their place. The mechanism whereby these new elements are differentiated has not been studied in complete detail. The trace always includes much more xylem than phloem, the latter is most easily recognized by the collapsed elements on the lower side of the trace which stain densely with haematoxylin. It is not known if new conducting elements are added entirely from the sur- rounding cells, or from the undifferentiated cells between xylem and phloem, or from both sources. These details have not been sought. A group of (still procambial) bud traces of a dormant bud and the median leaf trace of the supporting leaf are shown in a tangential section through the secondary cortical tissue in Ficure 8. DISCUSSION Before we discuss the origin of vascular bundles which are primary in their upper and secondary in their lower part, we must recapitulate the principle of development of the vascular pattern of those monocotyledon- ous plants which have only primary bundles. A description of this has appeared in print for three species, namely Rhapis excelsa (Zimmermann & Tomlinson, 1967, see Figure 2 therein), Prionium serratum (Zimmer- mann & Tomlinson, 1968, Figures 14 and 17) and Dracaena fragrans (Zimmermann & Tomlinson, 1969, Figure 13). In these papers it has been shown that monocotyledonous vascular bundles originate as branches of leaf traces and grow upwards into a meristematic region, immediately below the shoot apex proper, which we have called the cap. The upper end of a potential vascular bundle in the cap remains undetermined until it is connected to a newly differentiating leaf trace associated with a young leaf primordium. Leaf traces which connect to a potential bundle in the cap center become major bundles, leaf traces which appear during a later stage of leaf-primordium development link with a potential bundle at the cap periphery and thus give rise to a minor bundle with a shorter leaf- contact distance and a more limited radial penetration. This pattern of vascular development is typical and unique for monocotyledons, as far as we know, as it leads to what we call an “inner” (endogenous) vascular system. Additional new leaf traces may subsequently arise outside the Cap at a point where no free upper ends of vascular bundles are available. In Rhapis and Prionium these late-differentiating leaf traces become cor- tical bundles and end blindly below, sometimes (as in Rhapis) anastomos- ing among themselves. This suggests that there is no physiological mech- anism available to “propagate” these bundles. In Dracaena, on the other hand, which has a vascular cambium, such late-appearing (“outer” or exogenous) leaf traces can continue to grow below as secondary vascular traces since there is a vascular cambium available whereby they can be “propagated.” This pattern is more like dicotyledonous vascular development and fundamentally different from the “normal” monocotyledonous development of the “inner” vascular System. This developmental pattern in the young sympodial branch pro- 490 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 vides an easy and, indeed, the only seemingly possible explanation of primary bundles which are continuous below as secondary bundles. When this situation became clear it lead to the postulate that secondary bundles of Dracaena are homologous with the cortical bundles of Rhapis and Prionium. This, in turn, necessitated the prediction that monocotyledons with secondary growth have no cortical bundles. Examination of sections of a number of species representing genera in the Agavaceae (Agave, Beaucarnea, Cordyline, Dasylirion, Pleomele, Yucca; the Liliaceae (Aloé) ; and the Xanthorrhoeaceae (Lomandra, Xanthorrhoea) in the microslide collection of one of us (P.B.T.) showed that the prediction was indeed supported, The reverse is not necessarily true, however, because we do know of monocotyledons (e.g. certain Bromeliaceae) in which there is no cortical system, but neither is there a secondary vascular system. In summary, our investigations so far suggest that monocotyledons pos- sess two patterns of vascular development, an inner one (such as the central vascular system of palms) and an outer one which is represented by a system of cortical fibrous bundles in Rhapis and Prionium, but in Dracaena includes some of the primary bundles (the more peripheral ones in regions of establishment growth) and all the secondary bundles. This last statement needs qualification. We realize that secondary vascular bundles can grow in Dracaena without direct contact with the crown. Such a situation exists in the basal part of the stem as shown in FicureE 2c,d. Secondary tissue here “creeps” up on the stem by progressive acropetal development. We know no more about how this vascular development is regulated than we know how vessel development at the base of a ring- porous dicotyledonous tree is regulated. We realize that many questions are still unanswered. It will be neces- sary to find out how sharply definable the two patterns of development are in Dracaena. We have seen very clear-cut examples of both types, but we have the impression that a sharp line cannot always be drawn between the two. Ultimately, the problems of developmental physiology will have to be studied. Vascular strands must always develop between two physiological poles. The positional arrangement of these poles is much more complicated in monocotyledons than in dicotyledons. In spite of all these open questions, however, we feel that Dracaena has brought us considerably closer to the understanding of monocotyledonous growth. SUMMARY In the region of establishment growth some of the primary vascular bundles of Dracaena fragrans are continuous below with secondary vas- cular bundles. In terms of development this can be accounted for by late differentiating leaf traces which fail to make contact with the meristematic cap of the developing crown. Unlike the corresponding bundles in palms which form a feebly developed cortical system, these bundles in Dracaena continue their differentiation in association with the secondary vascular cambium as secondary vascular bundles. The cortical system of Rhapis 1970] ZIMMERMANN & TOMLINSON, DRACAENA, 2 491 and Prionium is thus homologous with the secondary vascular system of Dracaena. Further evidence in support of this comes from the observation that all monocotyledons with secondary vascular tissues which have been examined lack an independent cortical system. ACKNOWLEDGMENTS Support for continuing studies on the anatomy of monocotyledons is made possible for one of us (P.B.T.) by a grant from the National Science Foundation (GB—5762-X). We are indebted to G. W. Bailey for assistance in preparing serial sec- tions. LITERATURE CITED MireBEL, C.F.B. pe. 1845. Suites des recherches anatomiques et phan a sur quelques végétaux monocotylés. Ann. Sci. Nat. Bot. III. 3: 321-337. Mout, H. von. 1824. De palmarum structura. Jn: K.F.P. von mae His- toria ieee Palmarum 1: pp. I-LII. 16 pls. 849. On the structure of the palm stem. Rep. Ray Soc. 1849: 1-9 ner. J. C. 1903. Die Stammesbildung der Monokotylen. Flora aa 92: 32-48. ee Ueber die Verdstelung bei monokotylen cgi “ oo Verastel- on Pandanus. Ann. Jard. Bot. Buitenz. 20 (11.5): 5 ee P. B., & M. H. ZIMMERMANN. 1966. Anatomy of ce palm Rhapis excelsa, III. Juvenile phase. Jour. Arnold Arb. 47: 301-312. a 1969. Vascular anatomy of monocotyledons with secondary wth — an detrudnetion, Ibid. 50: 159-179, au ck. M. H., & P. B. Tomirnson. 1965. Anatomy of the palm Rhapis excelsa, I. Mature vegetative axis. Jour. Arnold Arb. 46: 160-180. . 1967. Anatomy of the palm Rhapis excelsa, IV. Vascular development in apex of vegetative aérial axis and rhizome. /bid. 48: 122- 142, —. 1968. Vascular construction and development in the aerial stem of Prionium (Juncaceae). Am. Jour. Bot. 55: 1100-1109. 969. The vascular arin in the axis of Dracaena fragrans (Agavaceae), i ib a and development of primary strands. Jour. Arnold Arb. 50: 370-38 [M.H.Z.] ied lie HARvarpD UNIVERSITY FAIRCHILD TROPICAL GARDEN Cazot FOUNDATION 10901 OL_p CuTLER RoapD PETERSHAM MIAMI MASSACHUSETTS 01366 FLORIDA 33156 492 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 RELATIONSHIPS IN THE PIRIQUETA CAROLINIANA- P. CISTOIDES COMPLEX (TURNERACEAE) ROBERT ORNDUFF IN ASSOCIATION WITH an investigation of the breeding systems of Pi- riqueta caroliniana (Walt.) Urban sensu lato (Ornduff & Perry, 1964) and P. cistoides (L.) Griseb. (Turneraceae), I have carried out a crossing program to determine whether the conspicuous morphological variation within these species is associated with (or maintained by) inter- or intra- specific barriers to crossing or with hybrid sterility. This paper presents the results of the hybridization program and briefly discusses some of the evolutionary and taxonomic implications of these results. Many of the variants in each species have been accorded taxonomic recognition by other authors. In the following paragraphs, however, the use of binomials or of varietal names is one of convenience in referring to the morphological variants and is not a reflection of taxonomic decisions that I have made in the group. Both species discussed in this paper are in need of a taxonomic revision that must utilize a wider range of techniques than those I have used here. I am indebted to W. R. Anderson, W. R. Ernst, F. W. Martin, and J. W. Purseglove for their generous assistance in providing seeds of Piri- queta cistoides and to J. D. Perry for providing the results of his pre- liminary investigations of the P. caroliniana group. Much of this work was carried out at the Department of Botany, Duke University, and was sup- ported in part by research grants from the National Science Foundation and Associates in Tropical Biogeography at the University of California. Piriqueta caroliniana is usually a perennial herb which ranges along the Coastal Plain from South Carolina southward through Florida to Cuba and Haiti (Urban, 1883; Small, 1933; Brizicky, 1961; Ornduff & Perry, 1964). A few collections referred to this species also have been made in scattered localities in Brazil, Venezuela, and Colombia (Urban, 1883). The species is absent from the Lesser Antilles. Piriqueta carolini- ana occurs in somewhat sandy soil of dunes, grassy areas, or open wood- lands, and, although it seems able to tolerate ecological disturbance such as fire, lumbering, or grazing, it is not an invasive weed. Urban (1883) recognized six varieties of Piriqueta caroliniana which differ largely in pubescence and foliar characters. Three of the varieties occur in the United States, although only one is restricted to this country. Small (1903, 1933) recognized four species of Piriqueta in the United States that are distinguished primarily by characters of the pubescence. Three of these species occur only in Florida. In contrast, Brizicky (1961) recognized only a single species (P. caroliniana) with five varieties in the United States and suggested that the “entire genus is much in need of a modern taxonomic revision, and field observations, culture experiments, 1970] ORNDUFF, PIRIQUETA (TURNERACEAE) 493 and cytogenetic studies are needed to determine the status (probably intraspecific) of the variants in our area.” A close relative of Piriqueta caroliniana is P. cistoides, a widely dis- tributed rather weedy plant that occurs in the Greater Antilles and the Lesser Antilles, and also ranges from Mexico through Central America into Brazil (Urban, 1883). Although described as an annual, some plants of this species will continue to flower for at least three years in cultivation. Piriqueta cistoides is variable throughout its range. Urban (1883) recognized eight varieties in this species. Piriqueta cistoides var. cistoides is the most widely distributed and occurs almost throughout the range of the species; in pubescence characters it resembles P. caroliniana var. caroliniana. Four of the varieties are restricted to mainland South America, one is endemic to Jamaica, and two occur in both the West Indies and on the South American continent. In general, however, subsequent authors (including Urban, 1920) have not recognized infraspecific taxa in P. cis- toides (e.g., Boldingh, 1913; Fawcett & Rendle, 1926; Standley, 1928; Stahl, 1936; and Alain, 1957). Piriqueta caroliniana and P. cistoides are diploid with » = 7 (TABLE 1; Lewis et al., 1962). The most consistent morphological differences between them are floral ones. Pirigueta caroliniana has large, distylous, and strongly self-incompatible flowers; P. cistoides has small, homostylous, and largely autogamous flowers. HYBRIDIZATION PROGRAM A program of artificial inter- and intraspecific hybridizations was car- ried out with Piriqueta caroliniana and P. cistoides. Plants of P. caro- liniana used in this program were referable to each of the morphological variants recognized by Small (1933; TABLE 1). Three of the collections of P. cistoides were referable to var. cistoides; the Trinidad plants were closest to var. latifolia Urb. (TABLE 1). Seeds from South American and Meso-American populations of these species have not been available, and, as a result, this study has been limited to representatives from the south- eastern United States and the West Indies. Artificial hybridizations were carried out in insect-proof cages in the greenhouse. The large, self-incompatible flowers of Piriqueta caroliniana were not emasculated prior to hybridization, but the anthers of the small, autogamous flowers of P. cistoides were removed before anthesis to prevent self-pollination. Intraspecific interpopulation hybridizations are easily accomplished in P. caroliniana and result in a full seed-set. In contrast, the seed-set following such hybridizations within P. cistoides is reduced, possibly because of damage to the flowers as a result of removal of the anthers. Interspecific hybridizations were possible only when f. cistoides served as the seed parent, indicating the presence of a unilateral incom- patibility system. When the hybrids flowered, pollen samples from them TABLE 1, Origin and chromosome numbers of collections of Piriqueta used in crossing program TAXON DESIGNATION been ORIGIN UMBER (P. cistoides var.) cistoides Dominica n=7 Dominica, West Indies, W. R. Ernst 2141, cistoides Jamaica Kingston, Jamaica, M. Crosby, H. Hespen- heide, & W. Anderson 1264. cistoides Puerto Rico ee Mayaguez, soc Rico, F. W. Martin latifolia Trinidad a= 7 St. icoelaa: Trinidad, J. W. Purseglove s.n. (P. caroliniana var.) glabrescens 1519 — Collier go 5 east of Monroe Sta- on, J. D. Perr caroliniana 64541 a= 7 South Carolina. sao County: west of antee. tomentosa 6743 Florida. Hillsborough County: Tampa. glabrescens 6746 n= 7 Florida. Collier County: east of Naples. tomentosa-viridis-glabrescens * 6750 n= 7 Florida. Dade County: southwest of Home- stead, caroliniana 6752 2n = 14 Florida. Dade County: ri of Perrine. tomentosa-caroliniana * 6754 m=7 Florida. Dade County: Sunilan viridis 6755 g=7 Florida, Glades County: puncrntee of Palm- dale. caroliniana 6758 n= 7 Florida. Highlands County: south of DeSoto City. caroliniana 6762 South Carolina, Jasper County: near Hardee- ville 1 Collection numbers are those of the author unless otherwise noted. * Population contains plants referable to each taxon or intermediate between the taxa. b6b WOLAINOAAVY GAIONUV AHL AO TYNUAOL IS “t0A} 1970] ORNDUFF, PIRIQUETA (TURNERACEAE) 495 were mounted in aniline blue- ae iinig and scored for viability on the basis of their staining reaction (TABLE RESULTS Fifteen hybrid progenies of Piriqueta caroliniana representing 14 inter- population combinations were obtained (Tastes 1, 2). The average TABLE 2. Results of artificial hybridizations of Piriqueta caroliniana and P, cistoides Cones AVERAGE POLLEN (RANGE OF POLLEN VIABILITIES} VIABILITY NUMBER OF PLANTS IN PROGENY) [Intraspecific hybridizations: P. caroliniana] 6454 X 67551 88 (64-99; 5) Reciprocal 76 (59-99; 7) 6454 X 6758 89 (72-100; 5) 6743 X 6746 90 (75-98; 8) 6743 X 6754 94 (84-97; 6) 6746 X 6750 93 (82-100; 12) 6746 X 6754 92 (76-99; 4) 6752 X 6746 70 (43-99; 4) 6754 X 6454 90 (79-97; 8) 6754 X 6750 92 (80-100; 15) 6754 X 6755 94 (80-98; 6) 6755 X 6743 86 (75, 98; 2) 6755 X 6746 95 (86-100; 5) 6762 X 6743 “ (93-100; 6) 6762 X 6755 (85-100; 11) [Intraspecific hybridizations: P. toutes Dominica < Jamaica 43 (39-46; 4) Reciprocal 48 (3 oa 0; °) poi ra Puerto Rico 46 ( Reciproc 57 (40-69: é) aR < Trinidad 34 (22, 46; 2) Reciprocal 19 (17,23; 2) Jamaica X Puerto Rico 96 (95, 98; 2) Reciprocal 95 (92-97; 3) Trinidad X Jamaica 61 (50-81; 7) oy 37 (1-63; 6) nidad < Puerto Rico 72 (68-78; 6) Garena: (aaa Trinidad x 1 74 (74, 75; 2) F, 48 (0-80; 11) Trinidad & 6454 68 (9-90; og F, 46 (2-81; Trinidad & 6746 75 (60-92; 2) Trinidad & 6755 75 (47-86; 9) F, 53 (32, 74; 2) *See Taste 1 for origin and further gala of these populations. In the eilecd | column the seed parent is liste 496 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 pollen viabilities of 13 of these progenies were above 85 percent. Two hybridizations produced progenies with average pollen viabilities below 85 percent. However, the reciprocal of one of these crosses produced progeny with a high average pollen viability. The other of the progenies involved parental taxa (vars. caroliniana and glabrescens) which pro- duced fertile progeny when individuals from other populations were hy- bridized. These results indicate there are no consistent internal barriers to hybridization among the morphological variants of P. caroliniana, nor is sterility prevalent in the hybrid progeny. Examination of the morphology of the F; hybrids in Piriqueta carolin- iana indicates a tendency for pubescence characteristics of the more pubes- cent parent to be expressed in the progeny. For example, hybrids between parents with the caroliniana (hirsute and tomentose herbage) and viridis (glabrous) phenotypes resemble the caroliniana parent; those between glabrescens (tomentose pedicel and calyx only) and viridis phenotypes resemble the glabrescens parent. Those between tomentosa (tomentose herbage) or caroliniana and glabrescens resemble either of the first two parents. Thus, the hybrids are not intermediate between the parents in the pubescence characters which have provided the chief morphological bases for making a taxonomic distinction between them. Ten hybrid progenies of Piriqueta cistoides representing six combina- tions of parents originating on different West Indian islands were grown (TABLE 2). Reciprocal hybrid progenies were grown for four of these hybrid combinations. With one exception, the hybrid progenies of P. cistotdes exhibited a reduced pollen viability. The average pollen viability of the single F, generation was lower than that of the F;. The pattern of low interpopulation pollen fertility in P. cistoides provides a sharp contrast with the pattern obtained in P. caroliniana. Four progenies were obtained as a result of artificial hybridizations be- tween plants of the Trinidad collection of P. cistoides and P. caroliniana; hybridizations using other collections of P. cistoides failed to produce seed. The average pollen viabilities of these interspecific hybrid progenies ranged from 68 per cent to 75 per cent (TABLE 2). A reduction in average pollen viability occurred in the three F, generations that were grown. DISCUSSION There are no consistent sterility barriers separating the morphologically distinctive variants of Piriqueta caroliniana in the southeastern United States that have been accorded taxonomic status by some authors. This indicates that these distinctive variants of P. caroliniana are very closely related genetically. The ease of making artificial hybrids among them, and the generally high fertility of the resultant F, hybrids and of subse- quent generations, may explain in part the occurrence of natural popula- tions which share characters of two or more of the segregate taxa, since in many areas of Florida populations of two or more of these taxa occur in close proximity. Nevertheless, the majority of herbarium specimens I 1970] ORNDUFF, PIRIQUETA (TURNERACEAE) 497 have examined at FLAs, Fsu, and usF are referable to these taxa and show no obvious indication of hybridization. Some years ago Dr. J. D. Perry mapped the distribution of these variants in Florida, based on his examina- tion of herbarium specimens at DUKE, GA, NCU, Nsc, and us. Examination of his unpublished map indicates that populations of P. caroliniana var. caroliniana and var. tomentosa are considerably more abundant than are hose of vars. glabrescens and viridis, and, furthermore, that the latter two taxa are confined to the southern half of peninsular Florida. It seems possible that field studies may reveal the existence of differences in ecolog- ical tolerances of these four taxa and that the differences may provide the basis for their continued genetic and morphological integrity over much of their ranges. The low interpopulation crossability and the associated hybrid sterility in Piriqueta cistoides are unexpected in view of the contrast these results provide with the behavior of the closely related P. caroliniana. The degree of morphological differentiation among the plants of P. cistoides used in the crossing program was considerably less than that in material of P. caroliniana. Nevertheless, in P. cistoides this low degree of differentiation is in general associated with a genetic variability that results in a reduced pollen fertility of intraspecific hybrids. The high genetic individuality of each population of P. cistoides may be related to the autogamous breeding System and weedy habit of the species. It is probable that many of its populations are built up from one or a few initial colonizers of disturbed land. These populations are consequently rather uniform genetically. Homozygosity for random genetic changes or alterations in chromosome structure can become rapidly established in autogamous annuals, particu- larly under conditions where there are fluctuations in population size. In an outcrossing non-weedy perennial of stable habitats such as P. carolin- jana, the establishment of homozygosity for such random cytogenetic changes would be a slower process, even when generation time is taken into consideration. In addition, the genetic individuality of the populations of P. cistoides undoubtedly is strongly reinforced by the spatial isolation of the populations on West Indian islands. This speculation suggests, therefore, that the observed differences in the fertility of interpopulation hybrids of P. cistoides compared with those of P. caroliniana is a conse- quence of the differences in the breeding systems, duration, population structure, and distribution patterns of these two morphologically similar species. cer The close relationship between Piriqueta cistoides and P. caroliniana is indicated by their strong morphological similarity and by the moderately high fertility of their interspecific hybrids. The floral differences which separate P. cistoides and P. caroliniana are those which are associated with their different breeding systems. The showy, heterostylous flowers of Piriqueta caroliniana are outcrossed by halictid bees (Ornduff & Perry, 1964). In contrast, the flowers of P. cistoides are homostylous and are usually self-pollinated before anthesis. The smaller perianth and the reduced length of the reproductive structures of the flowers of P. cistoides 498 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 represent a familiar syndrome of features by which an autogamous species differs from its allogamous relatives (see Ornduff, 1969, p. 128). The floral morphology of P. cistoides indicates that this species has been derived from a heterostylous ancestor. The flowers of P. cistoides are long- homostylous, that is, they fundamentally combine a long style with the stamen length and pollen size of short-styled flowers (Ornduff, unpub- lished). Where heterostyly and homostyly occur in closely related taxa, homostyly is considered to be a derivative condition. Evidence from morphology, geographical distribution, and artificial hybridizations sug- gests that P. caroliniana is the probable ancestor of P. cistoides. The re- sults of this study have been useful in assessing genetic relationships among populations of these two species and also in explaining some of the differences in their variation patterns, Although these results may be pertinent to the taxonomic decisions that ultimately must be made in this group, the decisions will have a stronger foundation if they include a con- sideration of careful field, herbarium, and nomenclatural studies. LITERATURE CITED ALAIN, Bro. (ALAIN H. Liocter). 1957. Flora de Cuba. Vol. 3. Havan BotprincH, I. 1913. Flora voor de Nederlandsch West- Indische a ae Ko- loniaal Instituut Amsterdam. Brizicky, G. K. 1961. The genera of Turneraceae and Passifloraceae in the southeastern United States. Jour. Arnold Arb. 42: 204-218 Fawcett, W., & A. B. RENDLE. 1926. Flora of Jamaica. Vol. 5. London party 1 W. 1953. Heterostyly in Pirigueta caroliniana. Castanea 18: 103- tweet ie H., H. L. Striptinc, & R. G. Ross. 1962. Chromosome numbers for some angiosperms of the southern United States and Mexico. Rhodora 64: 147-161. ORNDUFF, be he as Reproductive biology in relation to systematics. Taxon 18: 12 & 4 eo Perry. 1964. pre aay biology of Piriqgueta caroliniana (Turneraceae). Rhodora 66: 100-109. SMALL, J. K. 1903. Flora of the southeastern United States. New York. . 1933. Manual of the southeastern flora. Chapel Hill. STAHL, A. 1936, Estudios sobre la flora de Puerto Rico. o 2. San Juan: Federal Emergency Relief Administration. 3 vols. 911 STANDLEY, P. C. 1928. Flora of the Panama Canal Joe ioe U.S. Natl. Herb. 27: x + 416. Urpan, I. 1883. Monographie der Familie der Turneraceen. Jahrb. Bot. Gart. Berlin 2: 1-152. . 1920. Symbolae Antillanae seu Fundamenta Florae Indiae Occidentalis 8: 418- 860. Berlin: Borntraeger. DEPARTMENT OF BOTANY UNIVERSITY OF CALIFORNIA BERKELEY, CALIFORNIA 94720 1970} KAZMI, BORAGINACEAE 499 A REVISION OF THE BORAGINACEAE OF WEST PAKISTAN AND KASHMIR * S. M. A. Kazumi 15. Lappula Gilib. Fl. Lithuan. 25. 1781. Echinospermum Sw. ex Lehm. Asperif. I: 113. 1818, TYPE SPECIES: not indicated. Annuals, rarely biennials or perennials. Leaves alternate. Inflorescence bracteate. Calyx divided to the base, lobes after flowering not at all to slightly enlarged. Corolla blue or white, funnelform to salverform, Fau- cal appendages present. Anthers included. Gynobase narrowly pyramidal. Style short, filiform; stigma capitate. Nutlets 4, attached throughout their length to the gynobase, sometimes apices free on the inner face, appendic- ulate, appendages glochidiate to aculeate. About 45 species, usually distributed in temperate Europe and Asia, . few in Africa, one in America. KEY TO THE SPECIES a. Nutlets smooth, tuberculate all over their surface; tubercles large, thorn- like, neither prickly nor bearing glochidia at their apices. b. Nutlets pyramidal to ovate-pyramidal in outline, usually longer than broad, tubercles less unequal, basal ones not much enlarged........... sk Tyce gon bible 6 @ aime a dbo ooh 2g IRE aa dene gE aoe et eens 1. L. spinocarpos. b. Nutlets broadly pyramidal in outline, usually broader than long, tubercles very unequal, basal ones much enlarged. .........--- Os Dis ceratophora. a. Nutlets usually rough, appendiculate at the margins, appendages bearing glochidia at their apices, usually tuberculate, on the dorsal and ventral sur- faces, tubercles small, very rarely tuberculate only. Nutlets usually not, or only rarely, indistinctly margined with few glo- chidiate appendages; usually indistinctly verruculose, more or less smooth, 3. L. sinaica, Be Nutlets distinctly tmargined with appendages, appendages glochidiate. d. F d fruits sessile, fruits pyramidal, nutlets triangular, mar- ginal appendages usually dilated at base, bases joined to form a narrow marginal wing. ........----+----: eee 4. L. went Flowers and fruits with short to long pedicels, fruits ovate, nutlets ovate to oblong, marginal appendages not at all to slightly dilated at base, bases not joined to form a marginal wing. e. Nutlets with uniseriate marginal appendages. f. Inflorescence strongly elongated; nutlets small, ie snp iba marginal appendages always less than 1 mm. long; eaves linear; Gas oe fi iiay' ER co he SN eee 5. L. microcarpa. : Pp. * Continued from volume 51, p. 402. 500 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 f. Inflorescence elongate to more or less contracted; nutlets usually large, 3-4(-5) mm. long; marginal appendages always longer than 1 mm., usually 1.5-2 mm. long; leaves lanceolate; bracts usually large. g. Leaves on the lower surface subglabrous. ............... FE LENORE UO REE eR ERT . L. semiglabra. g. Leaves on the lower surface densely pubescent. .......... Pe ety eae a tao Oe bcc aunt SNe pak eee x aly 7. L. patula. e. Nutlets with multiseriate marginal appendages. h. Nutlets with three series of appendages. .. 8. L. consanguinea. h. Nutlets with two series of appendages. i. Corolla funnelform to salverform, 3.5-7 mm. in diameter; stem covered with soft, white, appressed trichomes. ...... Ree ae ee eg Toe at oo 9. L. barbata. i. Corolla salverform, 3-4 mm. in diameter; stem covered with stiff, spreading trichom j. Appendages of the inner series a longer than those OF Sie Outer GOTK nso os os ok ots QO. L. heteracantha. j. Appendages of the inner series more or less equalling those of the outer series. ............ 11. L. myosotis. 1. L. spinocarpos (Forssk.) Ascherson & O. Kuntze, Acta Horti Petrop. 10: 215. 1887; Riedl in Rechinger, Fl. Iranica 48: 69. 1967 Anchusa spinocarpos Forssk. Fl. Aegypt.-Arab. 41. 1775. Echinospermum vahlianum Lehm. Asperif. 1: 132. 1818. E. tuberculosum Ledeb. in Eichw. Pl. Casp.-Cauc. re 11, 1831. E. spinocarpos (Forssk.) Boiss. Fl. Orient. 4: 249. 1875. gs ee fa spinocarpos (Forssk.) Brand, oe hes IV. 252 (Heft 97): 98. 1931 TYPE: Aten Forsskal, s. n. (c). Icon.: Brand l.c. fig. 1 B. 1931, under Sclerocaryopsis spinocarpos. Annual herb, usually decumbent or sometimes erect, branched at the base, 5-20 cm. tall. Stem and branches covered densely with appressed to subappressed white trichomes, arising from tiny bulbous bases. Basal leaves usually linear-spathulate, entire, obtuse, 15-35 mm. long, 1-3 mm broad, covered densely on both surfaces with white subappressed tri- chomes; cauline leaves shorter and narrower, sometimes narrowly lan- ceolate, not much attenuated towards the base. Inflorescence lax, brac- teate; pedicels in flower about 1 mm. long, in fruit elongated to 4 mm. Calyx divided to the base, lobes in flower 1-2 mm. long, to 0.75 mm. broad, erect, covered with white appressed trichomes, in fruit enlarged, to 7 mm. long and 1.5 mm. broad, with subpatent trichomes arising from prominent tuberculate bases. Corolla infundibuliform, 3-4 mm. long, tube white, limb blue, more or less campanulate. Nutlets pyramidal to ovate- pyramidal, usually longer than broad, 3—5 mm. long, tuberculate, tubercles large, thorn-like, not prickly, very rarely glochidiate, less unequal, basal ones not much enlarged, surface smooth to verruculose or rarely minutely aculeate-tuberculate. 1970] KAZMI, BORAGINACEAE 501 DISTRIBUTION: North Africa, Sinai, Palestine, Irag, Iran, Afghanistan, central Asia, Tien Shan, Pamir Alaj, West Pakistan. West Pakistan: DERA GAzI KHAN Dist.: Fort Munro, 1800 m., Kazmi 1862a (PES); Rechinger 29727 (w). Quetta Dist.: 45 miles from Sibi to Quetta in Bolan Pass, Kazmi 1398b (PEs); ZHOB Dist.: between Zhob and Sambaza, 1500 m. Kazmi 1923b (pes); Rechinger 29845 (w); 30 km. N. Fort Sandeman, Rechinger 29858 (w). Reported from: Sibi, Banerji in Williams 55; Nai Ublo, Harsukh 20603. The leaves of Lappula spinocarpos vary from linear to linear-spathulate, to narrowly lanceolate, or sometimes to oblong-ovate. The surface of the nutlets and of the large tubercles varies from shining and smooth to rough, to minutely verruculose, or even to minutely aculeate. Similarly the tri- chomes on the leaves are very dense and appressed in some cases (Persia, May, 1869, Bunge (cH) ), while loose and erect in others (specimens from Kasakistan, Caucasus, Egypt, and Iraq). 2. L. ceratophora (M. Pop.) M. Pop. Fl. URSS 19: 417. 1953; Riedl in Rechinger, Fl. Iranica 48: 70. 1967. Echinospermum ceratophorum M. Pop. in Korov. Kultiasow Pop. Descr. rt. Nov. Turk. 67. ¢. 17. 1916. Tyre: In provincii bucharicis Karschi et Burdalyk Turkestaniae, J. J. Sprygin & M. V. Kultiasow s. n. (Lf). Icon.: M. Pop. 1. c. 67. ¢. 17. 1916; M. Pop. 1. c. #. 20. fig. 1. 1953. Annual, usually decumbent sometimes erect, to 15 cm. tall, much branched at the base. Stem and branches densely covered with white, short, depressed trichomes. Leaves linear to linear-spathulate, obtuse, entire, 20-40 mm. long, 1-3 mm. broad, both surfaces covered with white appressed to subappressed short trichomes, margins (especially towards the base) bearing some scattered long trichomes. Inflorescence lax, brac- teate; pedicels in flower up to 1 mm. long, in fruit elongated to 4 mm. broad, erect, in fruit enlarged to 3-6 mm. long and 1.5 mm. broad, erect or slightly curved towards the fruit, covered densely with subappressed trichomes, arising from prominent tuberculate bases. Corolla blue with a white eye, infundibuliform, slightly dilated at the throat, lobes erect or patent, Nutlet broadly pyramidal, usually broader than its length, tuberculate, tubercles large, thorn-like, usually smooth and shining at the surface, rarely rough, very unequal, basal ones much enlarged, to 2 mm. long. DistripuTion: Iran, Afghanistan, West Pakistan, Syr-Darya, Amu Darya, Kara Kum, Pamir Alaj. West Pakistan: QuETTA Dist.: Quetta to Chaman, 30 km. northeast of Quetta, Kazmi 1468b (pes); Lamond 958 (E); Rechinger 28866, 28908b (w). The only difference between Lappula spinocarpos and L. ceratophora 502 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 is in the form of the fruit and the length of the tubercles. In L. spino- carpos the nutlets are longer than broad with comparatively shorter tu- bercles, varying in length from the top to the base of the nutlet. Usually the middle ones are the longest, but this character is not very constant and sometimes the lower tubercles are longer than the middle ones, thus bringing this species very close to L. ceratophora in this respect. In L ceratophora the nutlets are broader than long with comparatively longer tubercles, increasing in their length, more or less gradually, from the top towards the base, the basal tubercles are the longest. This character is more stable in L. ceratophora. For distinguishing the two species from one another it is necessary to consider and correlate the ratio of the length and breadth of the nutlets and the length ratio of the tubercles from the top to the base of the nutlets. There is little difference in the form of leaves or floral parts in the two species. After examination of more material L. ceratophora may prove to be a subspecies or variety of L. spinocarpos. 3. L. sinaica (DC.) Ascherson ex Schweinf. Mem. Inst. Egypt 2: 111. 1887; Brand, Pflanzenr. IV. 252(Heft 97): 144. 1931; M. Pop. FI. URSS 19: 475. 1953; Riedl in Rechinger, Fl. Iranica 48: 76. 1967. Echinospermum sinaicum DC. Prodr. 10: 141. 1846; Boiss. Fl, Orient. 4: 251. 1875 E. kotschyi Boiss. Diagn. Pl. Orient. 7: 29. 1846. E. divaricatum Bunge, Relig. Lehm. 410. 1851 ex descr.; Mem. Acad. Sci. St. Pétersb. 7: 410. 1851. Lappula divaricata B. Fedtsch. Rast. Turk. 663. 1915. Type: Sinai, Aucher 118 (c). Annual erect to more or less decumbent, less branched herb, 6—50 cm. tall. Stem and branches flexuose, slender, covered with white, appressed to subpatent trichomes arising from minute white tuberculate bases. Basal leaves petiolate, oblong to elliptic-oblong, entire, obtuse, (includ- ing petioles) 15-50 mm. long, 5-15 mm. broad, covered on both surfaces loosely to densely with equal to unequal long, subappressed trichomes in the old leaves sometimes arising from white tuberculate bases; cauline leaves sessile, gradually decreasing in size upwards. Inflorescence in flower long, very much elongated in fruit; pedicels lacking to very short in flower, elongated to 5 mm. and reflexed in fruit. Calyx divided to the base, lobes linear-oblong in flower, 1—-1.5(—2) mm. long, hairy, erect, not much enlarged in fruit. Corolla bluish-purple with a white or yel- lowish eye, infundibuliform, + 3 mm. long, + 2.5 mm. in diameter; faucal appendages trapeziform. Nutlets oblong-ovoid, 2—2.5 mm. long, indistinctly margined, tuberculate to quite smooth, sometimes indistinctly verruculose or bearing few short glochidiate appendages, totally attached to the gynobase. DIsTRIBUTION: Sinai, Iraq, Iran, Caucasus, Afghanistan, Turkestan, Pamir Alaj, Tien Shan, West Pakistan, northwest India. 1970] KAZMI, BORAGINACEAE 503 West Pakistan: CHITRAL STATE: Chitral village, 2 miles south, on gravel slope, 1500 m., Stainton 2213 (BM). KALAT STATE: between Kalat and Mastung, 40 km. from Kalat, 2000 m., Kazmi 1278b (pes); Rechinger 28377 (Ww). QUETTA Dist.: Quetta, Lace s. n, (E); Spin Karez, 1900 m., Kazmi 1632c (PEs); Rech- inger 29235 (w); between Qila Abdullah and Sheila Bagh, 2300 m., Kazmi 1529 (GH, PES); Bolan Pass, 2000 m., Kazmi 1407b (pes); Rechinger 28423 (w); between Kolpur and Machh, 900 m., Kazmi 1401b (pes); Rechinger 28431 (w). Reported from Baluchistan: Khojak, 2300 m., Rechinger 29067 (w); Inter Bostan et Saran Tangai, 1800 m., Rechinger 29192 (w); Ziarat, 2100 m., Rech- inger 29361 (w); Tak, 30 km. S. Quetta, 2100 m., Rechinger 28395 (w); 10 miles S. Kalat, 1900-2200 m., Rechinger 28362 (Ww). 4. L. sessiliflora (Boiss.) Giirke in Engl. & Prantl, Nat. Pilanzenfam. IV. 3a: 107. 1893; Brand, Pflanzenr. IV. 252 (Heft 97): 144, 1931; M. Pop. Fl. URSS 19: 477. 1953; Riedl in Rechinger, Fl. Iranica 48: 78. 1967. Echinospermum sessiliforum Boiss. Diagn. Pl. Orient. 11: 124. 1849; FL Orient. 4: 253. 1875. Heterocaryum divaricatum Stocks ex Boiss. Fl. Orient. 4: 253. 1875. Cynoglossospermum sessiliflorum O. Kuntze, Rev. Gen. Pl. 2: 437, 1891. Type: Persia: Esfahan, Aucher 5017 (Gc). Icon.: Brand, 1. c. fig. 15. F—J. 1931. Annual, erect or decumbent herb, branched at the base, 5—20(-25) cm. tall. Branches divaricate, curved. Stem and branches densely covered with short, white to brownish, appressed trichomes. Basal leaves petiolate, linear-oblong to oblong-spathulate, entire, obtuse, (including petioles) 15- 25 mm. long, 1-2.5 mm. broad, covered on both surfaces uniformly with thin, patent, white trichomes to 0.5 mm. long, intermixed with scattered thick trichomes, to 1 mm. long, arising from white tuberculate bases; cauline leaves sessile, narrower and shorter, covered uniformly with tri- chomes 0.5-0.75 mm. long, arising from prominent, white, tuberculate bases. Inflorescence long, bearing distant sessile flowers or fruits, brac- teate, bracts leaflike, oblong to oblong-ovate. Calyx divided to the base, lobes oblong, hairy, erect, not enlarged in fruit, to 2 mm. long. Corolla blue, infundibuliform-campanulate, 2-2.5 mm. long; faucal appendages minute to inconspicuous. Nutlets broadly triangular, 2 5-3 mm. long, margined with uniseriate appendages, appendages glochidiate at their apex, free to confluent at base, sometimes forming quite broad scarious wings at margins, dorsal and ventral surfaces tuberculate to tuberculate-verruculose, rough, totally attached to the gynobase. Distripution: Iraq, Iran, Caucasus, Pamir Alaj, Tien shan, Turkestan, Afghanistan, West Pakistan. West Pakistan: KALaT STATE: Between Kalat and Mastung, 2000 m., Kazmi 1273 (PEs); Rechinger 28386 (w). Quetta Dist.: West of Ziarat, 2200 m., Kazmi 1669b (pes); Rechinger 29285 (w); Spin Karez 1900 m., Kazmi 16416 (pes); Rechinger 29234 (w); between Quetta and Ziarat, Kazmi 1574b (PEs); 504 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 near Kolepur, Kazmi 1400b (res). Reported from: Quetta, inter Qila Abdullah et Sheila Bagh, 1600-1900 m., Rechinger 29011 (w); inter Dulai et Kanak, 1600 m., Rechinger 29095 (w). Baluchistan, Stocks s. n. (GH-type of Hetero- caryum divaricatum Stocks ex Boiss.). The sessile fruits and triangular nutlets of Lappula sessiliflora distin- guish this species from allied species having nutlets with appendaged margins. The appendages on the margins of the nutlets of L. sessiliflora are very variable in length and breadth. Sometimes they are quite dis- tant to slightly confluent at their bases, while in other cases they are so strongly joined together that they form a broad scarious wing at the margins. These wings may be turned vertically to give the nutlets a cup-like appearance. Usually narrowly winged and broadly winged nut- lets are found on the same plant at different stages of development. 5. L. microcarpa (Ledeb.) Giirke in Engl. & Prantl, Nat. Pflanzenfam. IV. 3a: 107. 1893; Brand, Pflanzenr. IV. 252 (Heit 97): 142. 1931, excl. syn. E. rigidum DC. |ex Persia]; M. Pop. Fl. URSS 19: 462. 1953; Riedl in Rechinger, Fl. Iranica 48: 74. 1967 cocoa ea microcarpum Ledeb. Fl. Alt. 1: 202. 1829; Icon. Fl. Rossica 2: 830; DC. Prodr. 10: 140. 1846; Boiss. Fl. Orient. 4: 251. 1875. z. osm 208 & Kir. Bull. Soc. Nat. Moscou 14: 715. 1841; DC. Prodr. 10: 140. 1 E. oligacanthum Ledeb. Fl. Rossica 3: 161. 1846-1851 Heterocaryum inconstans Vatke, Zeitschr. Ges. Naturw. 11: 129. 1875. Type: Habitat in lapidosis montium Arkaul, versus cacumen, in mon- tibus ad rivulum Urmuchaika prope Buchtarminsk, Ledebour s.n. (LE ?). Icon.: Brand, l. c. fig. 15. A-E. 1931; Ledeb. 1. c. #. 183. 1830, under Echinospermum microcarpum. Annual or biennial, erect herb, 15-55 cm. tall. Stem solitary or many, simple or usually branched above, branches long, divaricate; stem and branches covered with white, appressed trichomes, 0.5—0.75(—1) mm. long on the lower part of the stem, usually intermixed with longer and stouter trichomes arising from prominent, white, tuberculate bases. Basal leaves oblong, obtuse, gradually attenuated towards the base, 30-45 mm. long, 2-3 mm. broad, forming a rosette, covered densely or loosely on both surfaces with spreading trichomes to 2 mm. long, arising from prominent white tuberculate bases; cauline leaves sessile, linear-oblong, narrower and shorter. Inflorescence short, bearing closely set, subsessile flowers, later much elongated, with pedicellate fruits up to 5 mm. apart; pedicels hairy, stout, erect, to 2 mm. long; bracts small. Calyx divided to the base, lobes 1-2 mm. long, erect, not much enlarged in sie Corolla bluish- white, infundibuliform, 2. = 3 mm. long, 2-3 mm. in diameter; faucal appendages 0.5—0.75 mm. long. Nutlets ovoid, 2. $3 mm. long, with a single series of short, not glochidiate appendages at the margin (not more than 1 mm. long), dorsal and ventral surface tuberculate or granulate. 1970] KAZMI, BORAGINACEAE 505 DISTRIBUTION: Iran, Caspian Arals, Turkestan, Tien shan, Pamir Alaj, Afghanistan, West Pakistan, Kashmir. West Pakistan: CHITRAL STATE: Agram, Arkari Gol. west of Tirich Mir, 3000 m., Stainton 2616 (Bm); Kafiristan, Rumbur, hillside facing south, 2400 m., Bowes Lyon 675 (pM). Gitetr AGENcy: Baltistan, Kasurmik to Doghani, 2500 m., R. R. Stewart 20799 (GH). KatatT STATE: near Mastung, Kazmi 1324c (pes). QueETTA Drst.: Quetta, Kazmi s. n. (pes); West of Ziarat, 2400 m., Kazmi 1654b (pes); Ziarat, 2400 m., Jafri & Akbar 2071 (sm); between Hindu Bagh and Quetta, Jafri & Akbar 2357 (BM). Kashmir: LADAK: Bod Kharbu, Koelz 6239 (Gu, us). 6. L. semiglabra (Ledeb.) Giirke in Engl. & Prantl, Nat. Pflanzenfam. IV. 3a: 104. 1893; M. Pop. Fl. URSS 19: 430. 1953; Riedl in Rechinger, Fl. Iranica 48: 75. 1967. Echinospermum semiglabrum (Ledeb.) FI, Alt. 1: 204. 1829; Icon. Fl. Rossica 1: t, 28. 1829; Fl. Rossica 3: 158. 1847; DC. Prodr. 10: 138. 1846; Boiss. Fl. Orient. 4: 251. 1875; C. B. Clarke in Hook. f. Fl. Brit. India 4: 163. 1883 Lappula redowskii (Hornem.) Greene var. patula (Lehm.) Nels. & Macbr. subvar. semiglabra (Ledeb.) Brand, Pflanzenr. IV. 252 (Heft 97): 149. 1931. Echinospermum caspicum Fisch. & C. A. Mey. Index Sem. Hort. Petrop. 5: 34. 1838; DC. Prodr. 10: 138. 1846. Tyrer: Habitat in sabulosis ad fi. Irtysch, Ledebour s. n. (LE). Icon.: Ledeb. I. c. 1: ¢. 28. 1829; M. Pop. l. c. 19: t. 21. fig. 2. 1953. Annual, erect herb, 15~50 cm. tall. Stem simple or branched above; stem and branches covered with short, appressed white trichomes. Basal leaves petiolate, oblong-spathulate, obtuse, entire, (including petioles) 40-50 (—60) mm. long, 6-9 mm. broad, upper surface usually glabrous or some- times with few scattered trichomes, lower surface usually loosely, sometimes, especially in the old leaves, densely covered with white, spread- ing trichomes to 1 mm. long, arising from prominent white tuberculate bases; cauline leaves gradually decreasing in size upwards, sessile, more or less lanceolate, acutish. Inflorescence short in flower, much elongated later in fruit, flexuose, bracteate, bracts small. Flowers subsessile, fruits pedicellate; pedicels hairy, erect, to 3 mm. long. Calyx divided to the base, lobes oblong, hairy, erect, 1-1.5 mm. long in flower, enlarged in fruit to 3 mm. long, + 0.5 mm. broad. Corolla bluish-white to light blue, campanulate, 3-3.5 mm. long, 3-4 mm, in diameter; faucal appendages trapeziform. Nutlets narrowly ovoid 4~5 mm. long, margined with uni- seriate glochidiate appendages 2-3 mm. long, slightly dilated at base, dorsal middle area carinate, usually aculeate, outer surface slightly tuber- culate. DIstRIBuTION: URSS, Afghanistan, West Pakistan, Kashmir, northwest 506 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 West Pakistan: Gitcir AcENcy: Baltistan, Thalle La, 2200 m., R. R. Stewart 20584a (GH); rocky barren slopes near south end of Satpura Lake, ca. 4.5 miles south of Skardu, 3000 m., Webster & Nasir (GH), Satpura Nullah, above Skardu, 2700 m., R. R. Stewart 20368 (GH). MAkran Dist.: between Hushab and Panjgur, 20-30 km. from Panjgur, Kazmi 1190b (pes); Lamond 567 (£). 20 miles from Panjgur towards Nag, Kazmi 1243c (pes). ZHop Dist.: Shingarh, 50 km. north of Fort Sandeman, 2000 m., Rechinger 29894 (w); Harsukh 20595 (x). Reported from: Kharan and Quetta Haughes Buller 23159. Kashmir: Miscellaneous: Hab. Tibet Occ. regio temp. 10,000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson 10 (cH); 11,000-12,000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (GH). 7. L. patula (Lehm.) Ascherson ex Giirke in Engl. & Prantl, Nat. Pflan- zenfam. IV. 3a: 107. 1893; M. Pop. Fl. URSS. 19: 436. 1953. Riedl in Rechinger, Fl. Iranica 48: 76. 1967. Echinospermum patulum Lehm. Asperif. 124. 1818; Ledeb. Fl. Alt. 1: 200. 1829; DC. Prodr. 10: 137. 1846; Boiss. Fl. Orient. 4: 250. 1875. Lappula redowskii (Hornem.) Greene var. patula (Lehm.) Nels. & Macbr. Bot. Gaz. 61: 39. 1916; I. M. Johnston, Contr. Gray Herb. 70: 49. 1924. Type: In deserto Caucasico-Caspico, ad Kuman et Terek nec ad Wol- gam inferiorem area Sareptanam colonium et urbem Astrachan, Lehmann S.n,. (HBG). Icon.: Hegi, Ill. Fl. Mitt.-Eur. 5: 3. fig. 3098a. 1927. Annual, branched near the base, to 50 cm. tall; branches long-divaricate. Stem and branches covered with short appressed white trichomes. Basal leaves linear-oblong to linear-spathulate, gradually attenuated towards the base, obtuse, entire, 40-60 mm. long, 6-9 mm. broad, covered on both surfaces, usually densely, (sometimes loosely) with short, white, appressed trichomes, intermixed, more or less, with longer ones, arising from white tuberculate bases; cauline leaves gradually reduced in size upwards. In- florescence short in flower, later much elongated, lax, bracteate, bracts small, leaflike. Flowers subsessile to short pedicellate; pedicels erect, hairy, to 1.5 mm. long in flower, elongated to 2.5 mm. in fruit. Calyx divided to the base, lobes linear to oblong, 1-2 mm. long in flower, much enlarged to 5 mm. long in fruit. Corolla bluish-white to blue, campanulate, 3-3.5 mm. long, 3-4 mm. diameter; faucal appendages trapeziform, prominent. Nutlets ovoid, 3-4 mm. long, margined with uniseriate ap- pendages, appendages 1.5—-3 mm. long, glochidiate at apex, dorsal middle area tuberculate, rarely aculeate, externally slightly tuberculate. DistriBuTION: URSS, Iran, Afghanistan, West Pakistan, Kashmir, northwest India, China. Kashmir: LapAK: Tsaka La, 5000 m., Koelz 2368 (GH); Kharbu to Dras, 3000 m., R. R. Stewart 2115 (cH). Miscellaneous: Hab. Tibet Occ. regio temp., 12,000-14,000 ped., Herb. Ind. Or. Hook. f. & Thoms. Herb. no. 2496849, 2500079 (us); s.n. (GH). 1970} KAZMI, BORAGINACEAE 507 Only two collections of Lappula patula have been made from Kashmir, but the species is to be expected throughout the northern parts of Kashmir and West Pakistan. Lappula patula is very closely related to L. semiglabra from which it may be distinguished by its habit of branching near the base, by its leaves densely hairy on both surfaces, and by its smaller and broader nutlets. In L. semiglabra the stems are branched in the upper part, the leaves are usually glabrous on the upper surface, and the nutlets are narrowly ovoid, 4-5 mm. long, margined with appendages 2-3 mm. ong. 8. L. consanguinea (Fisch. & C. A. Mey.) Giirke in Engl. & Prantl, Nat. Pflanzenfam. IV. 3a: 107. 1893; M. Pop. Fl. URSS 19: 447, 1953. Echinospermum consanguineum Fisch. & C. A. Mey. Index Sem, Hort. Petrop. 5: 35. 1838; DC. Prodr. 10: 137. 1846; Ledeb. Fl. Ross, 3: 157. 1847. E. lappula var. consanguineum (Fisch. & C. A. Mey.) Regel, Bull. Soc. Nat. Moscou 41(1): 89. 1868. Cynoglossospermum consanguineum O. Kuntze, Rev. Gen. Pl. 2: 438, 1891. Lappula echinata var. consanguinea (Fisch. & C. A. Mey.) Brand, Pflanzenr. IV. 252(Heft 97): 140. 1931. Type: Hab. in regionibus altaicus, Bunge s. n. (LE). Icon.: M. Pop. 1. c. #. 22. fig. 9. 1953. Annual or biennial, erect, up to 50 cm. tall, usually branched above, branches flexuose, to 25 cm. long. Stem and branches covered with un- equal, subappressed to patent trichomes to 1.5 mm. long. Basal leaves oblong-lanceolate, entire, obtuse, narrowed towards the base, to 7 cm. long, 11 mm. broad, nerves sunken on the upper surface, prominent below, both surfaces covered with scattered, white, spreading trichomes, to 1 mm. long, arising from prominent tuberculate bases; cauline leaves lanceolate, acutish, gradually reduced in size upwards. Inflorescence Jax, short, elongated in fruit, bracteate. Flowers subsessile to very short pedicellate, pedicels of old fruits pubescent, erect, to 2 mm. long. Calyx divided to the base, lobes linear-oblong, in flower up to 2 mm. long, 0.5 mm. broad, in fruit slightly enlarged, to 3 mm. long and 0.7 mm. broad. Corolla blue with a white eye, infundibuliform, 2.5-3 mm. long; faucal appendages prom- inent. Nutlets ovate, 2-2.5 mm. long, 1-1.5 mm. broad at the base, mar- gined with three series of appendages, appendages glochidiate at apex, slightly dilated at base, rarely confluent to form a wing, appendages of the inner series longest, to 1.5 mm. long; dorsal middle area tuberculate. DistriBution: Turkestan, Altai, West Pakistan, Kashmir. West Pakistan: Gitcrr AGENcy: Lower Rupal Nullah, Nanga Parbat, 2700 m., R. R. Stewart 18862 (cH). at the margin of the nutlets distinguish The t ies of dages : e three series of appendag arbata, which Lappula consanguinea from the closely related species L. b 508 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 has two series of appendages on its nutlets. Sometimes the appendages of the outermost series are very short, but they are always conspicuous. The specimen cited above is identical to the specimens collected from Altai and Turkestan. Brand (/. c. 1931) gives the distribution of L. consanguinea as extending to Eastern Europe, but I did not find any specimen from that area in the European collections. 9, L. barbata (M. Bieb.) Giirke in Engl. & Prantl, Nat. Pflanzenfam. IV. 3a: 107. 1893; Brand, Pflanzenr. IV. 252 (Heft 97): 140. 1931 excl. var. arragonensis; M. Pop. Fl. URSS. 19: 461. 1953; Riedl in Rechinger, Fl. Iranica 48: 72. 1967. Myosotis barbata M. Bieb. Fl. Taur.-Cauc. 1: 121. Echinospermum barbatum Lehm. Asperif. 1: 128. ‘ane DC. Prodr. 10: 137. 1846; Ledeb. Fl. Rossica 3: 156. 1847; Boiss. =e Orient. 4: 250. 1875; C. B. Clarke in Hook. f. Fl. Brit. India 4: 163. Myosotis saxatilis Pall. Nova Act. Petrop. 306. fee: nomen nudum. Echinospermum filiforme Godet in DC. Prodr. 10: 140. 1846. Lappula saxatilis (Pall.) Kusn. Mat. Fl. Kauk. 4(2): 183. 1930, Type: Habitat in Tauriae montibus calcareis circa Karassubasar et ad Caucasum, without citation of collector’s name, s. 7. (LE Icon.: M. Bieb. Cent. Pl. Rar. ¢. 36. 1810. Erect annual or biennial, to 65 cm. tall. Stem simple to branched, branches sometimes divaricate; stem and branches covered with white, soft, appressed to subappressed trichomes. Basal leaves numerous, form- ing a compressed rosette, lanceolate to linear-lanceolate, entire, obtuse to acutish, gradually tapering towards the base, to 7 mm. long and 10 mm. broad, covered on both surfaces with short appressed to subpatent white trichomes, arising from white tuberculate bases; cauline leaves sessile, linear, to 3 mm. broad, in length gradually reduced upwards. Inflorescence axillary or terminal, short in flower, later much elongated. Flowers brac- teate, subsessile to short pedicellate; pedicels hairy, erect to 1.5(—2) mm. long in fruit. Calyx divided to the base, lobes linear-oblong, erect, hairy, 2.5—-3 mm. long in flower, enlarged to 5 mm. long in fruit. Corolla white to light blue with a white or yellow eye, infundibuliform-campanulate, 3.5-7 mm. in diameter, faucal appendages oblong. Nutlets 3-4 mm. long, margined with two series of appendages, appendages glochidiate at apex; middle area tuberculate or rarely aculeate. Lappula barbata is variable in the size of flower and the length of the marginal appendages of the nutlets. Two varieties may be distinguished. 9a. Var. barbata Corolla 5-7 mm. in diameter; marginal appendages of the nutlets more or less equal in length. 1970] KAZMI, BORAGINACEAE 509 DISTRIBUTION OF VAR: Turkey, Caucasus, Turkestan, Irag, Iran, Af- ghanistan. 9b. Var. cariensis (Boiss.) Brand, Pflanzenr. IV. 252 (Heft 97): 141. 1931; Riedl in Rechinger, Fl. Iranica 48: 72. 1967. Echinospermum cariensis Boiss. Diagn. Pl. Orient. 4: 40. 1844; DC. Prodr. 10: 141. 1846. Echinospermum barbatum (M. Bieb.) Lehm. var. cariensis (Boiss.) Boiss. Fl. Orient. 4: 250. 1875. Lappula echinata Gilib. var. cariensis O. Kuntze, Acta Horti Petrop. 10: 214. 1887 Type: Hab. Anatolia, in collibus argillosis ad meridiam Cadmi et ad orientem urbis Danisleh, without citation of collector’s name, s. 7. (G). Corolla 3.5—5 mm. in diameter, marginal appendages of the outer series of nutlets much shorter than those of the inner series. DIstTRIBUTION: Turkey, Syria, Armenia, Iraq, Iran, USSR, Afghanistan, West Pakistan, Kashmir, northwest India. West Pakistan: CHirraL STATE: Barum Gol, above Shokor Shal, 3600 m., Wendelbo sn. (sERG). Quetta Dist.: Urak near Quetta, 1800 m., R. K. Stewart 602b (micH.); near Ziarat, 2500 m., Kazmi 1672b (pes), Rechinger 29392 (w), Fernandes 6269 (cH); Ziarat, Zarghun, Santapau 6419 (GH). WAZIRISTAN AGENcY: Razani, 2500 m., Lester Garland sn. (£). Miscellaneous: Scind, Stocks sn. (GH); Baluchistan, Elliott s.n. (K). Kashmir: Mitsahoi on Ladak Road, 3300 m., R. R. Stewart 9993 (GH); Zaskar, Lagong, 4000 m., Koelz 5408 (GH, US); Kargia, 4000 m., Koelz 5454 (GH, US). 10. L. heteracantha (Ledeb.) Giirke in Engl. & Prantl, Nat. Pflanzen- am. IV. 3a: 107. 1897; M. Pop. Fl. URSS 19: 427. 1953; Riedl. in Rechinger, Fl. Iranica 48: 72. 1967. Echinospermum heteracanthum Ledeb. Suppl. Index Sem. Hort. Dorpat. g. 1823: Fl. Rossica 3: 157. 1847; DC. Prodr. 10: 137. 1846; Boiss, Fl. Orient. 4: 249. 1875. : : Lappula echinata Gilib. var. heteracantha (Ledeb.) O. Kuntze in Acta Horti Petrop. 10: 214. 1887; Brand, Pflanzenr. IV. 252 (Heft 97): 139. 1931, pro parte. : Echinospermum semicinctum Stev. Bull. Soc. Nat. Moscou 24: 605. 1851. Type: Habitat in Astrachan, cultivated from the seed collected from Astrachan, without citation of collector’s name. Icon.: Ledeb. in Eichw. Pl. Casp.-Cauc. t. 21. 1831-1833. Annual or biennial herb, to 50 cm. tall. Stems solitary to many, simple to branched, branches long, divaricate; stem and branches covered with stiff, short or long, usually spreading sometimes subappressed, grayish or white trichomes arising from prominent tuberculate bases; trichomes on 510 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the lower part of the stem falling off with age. Basal leaves usually form- ing rosettes, oblong, gradually narrowed towards the base, entire, obtuse, 2-7 cm. long, 3—-8(—12) mm. broad, covered on both surfaces with stiff, short or long, patent or subpatent grayish trichomes arising from tuber- culate bases; cauline leaves gradually reduced in size upwards; upper leaves sessile, broader than the lower ones, acutish. Inflorescence lax, bracteate, much elongated in fruit. Flowers subsessile to short pedicellate; pedicels much elongated in fruit, to 3 mm. long, pubescent, erect, tapering towards the base. Calyx divided to the base, lobes in flower erect, 1.5—2 mm. long, in fruit spreading, to 3.5 mm. long. Corolla light blue with a white or yellowish eye, salverform, 3-4 mm. long, limbs 3—4 mm. diameter; faucal appendages oblong, conspicuous. Nutlets 2-3 mm. long, margined with two series of appendages, appendages glochidiate at their apices, appendages of the outer series much shorter than those of the inner series; middle dorsal and ventral surfaces tuberculate to minutely aculeate. DIsTRIBUTION: Eastern a Tran, USSR, Central Asia, West Paki- stan, Kashmir, northwest India in Pakistan: Gitcir AGENCY: Nomal, 1700 m., R. R. Stewart 26337 (BM); r, 3000 m., R. R. Stewart 26338 (Bm); Astor valley, 2700-3000 m., pale 22024 (cH); near Chorit, Rupal Nullah, 2700 m., R. R. Stewart 22844 (GH, US); Baltistan, Chatpani Nulla, 3500 m., el 13859 (Kk). Miscella- neous: eet Sinkachh, Harsukh 20594 Kashmir: Zoji Pass, 3300 m., R. R. Stewart ie (GH), 21247 (GH, US); Zoji Pua to Matayan, 3300 m., 'R. R. Stewart 7403 (kK); Dras to Tibet, 3000 m., Gammie (3. 8, 1891) s.n. (xk): Dras to Mistahoi, Ladak Road, R. R. Stewart 22340 (GH); Shingo valley near Gulteri, R. R. Stewart 22231b (cu). Miscella- neous: Hab. Tibet Occ. Regio temp., 12000-14000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (GH). 11. L. myosotis Moench, Meth, 417. 1794; Riedl in Rechinger, FI. Iranica 48: 71. 1967. Myosotis lappula L. Sp. Pl. 131. 1753. Lappula echinata Gilib. Fl. Lithuan. 25. 1781, nomen illegitimum. Echinospermum squarrosum Reichenb. Fl. Germ. Excurs, 345. 1834. Echinospermum ispahanicum Boiss. Diagn. Pl. Orient. 11: 123. 1849. E. lappula (L.) Lehm. Asperif. 1: 121. 1881. Lappula echinata Gilib. var. eu-echinata Brand, Pflanzenr. IV. 252 (Heft 97): 133; 1931. Type: Habitat in Europae argillosis nudis, ruderatis Herb. No. 180.9 (LINN). Icon.: Reichenb. Icon. Fl. Germ. 18: ¢. 128. fig. 2 and 7-19. 1858. Erect annual or biennial herb, Stems solitary or many, usually simple, sometimes branched above; stem and branches covered with short, white trichomes, arising from tuberculate bases. Basal leaves oblong-spathulate, entire, obtuse, gradually or abruptly attenuated into a long petiole, to 7 1970] KAZMI, BORAGINACEAE 511 cm. long and 3-12 mm. broad, covered on both surfaces, loosely or densely, with thin, long, white trichomes, arising from white tuberculate bases; cauline leaves shorter, narrower, acutish. Inflorescence short in flower, later, in fruit, elongated, lax and bracteate. Flowers short pedicellate; pedicels elongated in fruit to 3 mm., erect, hairy. Calyx divided to the base, lobes linear, acutish, hairy, in flower 2—3 mm. long, in fruit narrow- lanceolate, to 4-5 mm. long and 1 mm. broad, spreading to sometimes re- flexed. Corolla blue, salverform, ca. 4 mm. long, limbs 3-4 mm. in di- ameter; faucal appendages oblong. Nutlets 2.5—3 mm. long, margined with two series of appendages, appendages of both series more or less equal in length, glochidiate at apex, middle dorsal area verruculose-granulate, ex- ternally more or less tuberculate. DistrIBuTION: Central and Eastern Europe, Caucasus, Siberia, Aral- Caspian, Dsungare-Tarbagatai, Persia, Afghanistan, West Pakistan, North America. West Pakistan: KurraAM AGENCY: Kurram Valley, Aitchison 198 (GH); Re- ported from: Quetta Dist.: Ziarat, R. R. Stewart 600; Urak, Crookshank 104; Zarghun, Crookshank 368, 384. The closest relative of Lappula myosotis is L. heteracantha, from which it may be easily distinguished by the appendages at the margins of the nutlets being essentially equal in length in both series. 16. Lepechiniella M. Pop. Fl. URSS 19: 713. 1953. Type species: L. transalaica (B. Fedtsch.) M. Pop. (lectotype spe- cies). Annual or perennial herbs. Calyx divided nearly to the base; lobes 5, various, not at all to slightly enlarged in fruit. Corolla white or blue, tubular-infundibuliform, tube short, limbs patent; faucal appendages de- veloped. Stamens included, below the faucal appendages. Gynobase nar- rowly prismatic, tetragonous. Nutlets appendiculate, at the margins ap- pendages free at base or confluent and forming a marginal wing; nutlets ventrally keeled above the minute sub-basal, areola, attached to the gyno- base, free at apex. Style shorter, equalling or little exceeding the apices of nutlets. Species about 14, distributed from Iran to Central Asia, in mountains. KEY TO THE SPECIES ndages free at 3. L. albiflora. » Flowers white, nutlets appendiculate at the margins, appe base, not forming a marginal wing. ......------------+--- Flowers blue, nutlets appendiculate at the margins, appendages partly or completely confluent at base, forming a narrow or broad marginal wing. b. Nutlets broadly winged at margins, wings folded inside, inner margins of the wings scarious, entire to sometimes dentate. ...... 2. L. microcar pa. & $12 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 b. Nutlets narrowly winged at margins, wings not folded inside, inner margins of the wings not scarious, usually dentate, teeth bearing glochidia OE CE trees 28k: nace ota exam bear ews . L. inconspicua. 1. L. inconspicua (Brand) Riedl, Oesterr. Bot. Zeitschr. 110: 517. 1963; in Kgie & Rechinger, Biol. Skr. 13(4): 203. 1963; in Rech- inger, Fl. Iranica 48: 81. 1967 Paracaryum inconspicuum Brand, Repert. Sp. Nov. 13: 549. 1915; Pflanzenr. IV. 252 (Heft 78): 46. 1921. Type: Afghanistan, Griffith 5976 (x-holotype, GH-isotype). Icon.: Ried], 1. c. 48: ¢. 18. fig. 1. 1967. Annual, decumbent to ascendent herb. Stems many from the base, up to 20 cm. long, simple to little branched, slender; stem and branches cov- ered with soft, retrorsely appressed to subpatent trichomes ca. 0.2 mm. long, not arising from tuberculate bases. Basal leaves forming a rosette; basal and lower cauline leaves petiolate, petioles up to 20 mm. long, slender, hairy, with some thick, spreading trichomes at the margins to 2 mm. long, lamina oblong-ovate, to 30 mm. long and 15 mm. broad, gradually or sometimes abruptly narrowed towards the base, obtuse to roundish, loosely covered with white, soft, appressed trichomes to 0.2 mm. long on the upper surface, densely so on the lower surface, on the upper surface these intermixed with scattered long, stout, white trichomes arising from tuberculate bases; upper cauline leaves sessile, gradually reduced in size. Inflorescences short in flower, elongated in fruit, brac- teate. Pedicels short in flower, elongated in fruit, bracteate. Pedicels in flower 1-2 mm. long, in fruit elongated to 10 mm., slender, hairy, recurved. Calyx divided nearly to the base, lobes 1.5 mm. long, hairy, erect, en- larged in fruit, oblong to broadly lanceolate, 2-3 mm. long, 1 mm. broad, pubescent on both sides and margins. Corolla blue, subtubular, 2 mm. long, lobes broadly ovate, patent; faucal appendages subquadrate. Sta- mens included; filaments very short, inserted at the middle of the co- rolla tube. Nutlets not more than 2 mm. long, ovoid, narrowly winged and denticulate at the margins, dorsal middle area tuberculate, tubercles short to long and glochidiate at apex. Style ca. 0.5 mm. long. DIsTRIBUTION: Iran, Afghanistan, West Pakistan. West Pakistan: CHITRAL STATE: Brumboret Gol. N. of Drosh, 1500 m., Bowes Lyon 647 (pm); Stainton 2228 ya Harriss 16387 (Kk); Chitral Mastuj tract, Kaghosi, 1500 m., Stainton 2403 (Bm The specimens cited above differ in some minor details from the type (Griffith 5976) collected in Afghanistan. In the type specimen the stems, petioles, and pedicels are all densely covered with short trichomes, which are not retrorsely appressed but spreading. Sepals in fruit are 3-4 mm. long, oblong, more or less obtuse at apex. In the specimens cited above the short trichomes on the stems, petioles, and pedicels are evidently re- 1970] KAZMI, BORAGINACEAE 513 trorsely appressed and the sepals in fruit do not usually exceed 2.5 mm. (or rarely) 3 mm. They are broadly lanceolate and acutish at their apices. Brand (1. c.) and Riedl (1. c.) have considered the species to be an- nual, but specimens collected by Stainton (2403, 2228) appear to me to be perennials. On examining more material from the area the West Pak- istan plants may prove to be a variation worthy of recognition. 2. L. microcarpa (Boiss.) Riedl, Oesterr. Bot. Zeitschr. 110: Ls Wp 1963; in Kgie & Rechinger, Biol. Skr. 13(4): 203. 1963; in Rech- inger, Fl. Iranica 48: 80. 1967. Paracaryum microcarpum (Boiss.) Diagn. Pl. Orient. 3: 139. 1856; FI. Orient. 4: 255. 1875; C. B. Clarke in Hook. f. Fl. Brit. India 4: 162. 1883; Brand, Pflanzenr. IV. 252 (Heft 78): 46. 1921. Typr: Afghanistan: Hab. in regio Cabulico circa Choky et in sylvaticis Bharowul, Griffith 5975 (x-holotype, GH-isotype). Perennial, procumbent to ascendent herb. Stems many, 10-45 cm. long, simple or branched; stem and branches covered densely with white, pa- tent trichomes to 2 mm. long, arising from prominent tuberculate bases. Basal leaves petiolate, petioles slender, pubescent to 50 mm. long; la- mina oblong-lanceolate to oblong-ovate, entire, obtuse or sometimes acu- tish, to 50 mm. long and 20 mm. broad, covered on both surfaces with white, long, subappressed or, in the old leaves, more or less patent tri- chomes arising from white tuberculate bases; cauline leaves subsessile to sessile, gradually reduced in size upwards. Inflorescence short in flower, much elongated in fruit, lax, bracteate. Pedicels very short in flower, elongated to 13 mm. in fruit, slender, hairy, more or less reflexed. Calyx divided nearly to the base, lobes 1.5—-2.5 mm. long and ca. 0.7 mm. broad in flower, oblong, erect, densely hairy, in fruit oblong-ovate, ob- long and 1.7 mm. broad, pubescent externally, more or less glabrous inside, reflexed. Corolla blue, campanulate, 3-4 mm. long, tube equalling the limb, lobes broadly ovate, spreading, 1 mm. long, 1.7 mm. broad. Anthers included, inserted below the faucal appendages on the corolla tube. Nutlets brown, ovate, 2 mm. long, ca. 1.5 mm. broad, winged at the margins, wings inflexed and again folded outside, inner margins of the wings scarious, entire or sometimes dentate, middle dorsal area tuberculate, tubercles short (to sometimes long), bearing glochidia at apex, external surface smooth to slightly tuberculate. Style 1 mm. long, exceeding the nutlets by 0.5- DistrrpuTion: Afghanistan, West Pakistan, Kashmir. rat, Lowari Pass, 2500 m., Stainton 2558 West Pakistan: CHITRAL STATE: Zia mg aes ss, Siddiqui in R. R. Stewart (pm); Jambatai, Harriss 16385 (K); Lowari Pass, Sid 25448 (pm), Kurram Acency: Kurram Valley, Aitchison 106, 117, 197 (GH). Swat STATE: mountains east of Kalam, 25 miles above Bahrain, 2200 m., 514 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Rodin 5683 (us); Bahrain to Kolaila, 1600-2200 m., Lamond 1761 (E); Rechinger 30693 (w). Reported from Baluchistan hills, Stocks 738, 758; Fort ndeman, Duthie 18931; Harsukh 20601; Khyber Agency, Jamrud hills, ‘Siddi- qui 2460 Kashmir: Kishenganga Valley, Kiran Nullah, 1800-2000 m., R. R. Stewart 17538 (cH); Kiran to Reshna, 1800 m., R. R. & J. D. Stewart 17689 (GH); Islampur, Kazmi 293a (PES); China Kazmi Sn, A sovis Miscellaneous: Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (GH). Reported from: Uri, Meebold 4237; Padar, R. Re Stewart 2916; gato Nullaha, Nath 907. 3. L. albiflora Riedl in Rechinger, Fl. Iranica 48: 83. 1967. Type: Afghanistan: Kataghan, near Salang, 3000 m., Neubaur 4496 (w). Icon.: Riedl, 1. c. ¢. 16. fig. 12; ¢. 18. fig. 5. 1967. Perennial, shrubby herb, to 15 cm. tall. Stems numerous, prostrate, to 25 cm. long, simple or slightly branched, covered loosely or densely with appressed or subpatent trichomes. Basal leaves oblanceolate to spathu- late-lanceolate, entire, acute to obtusish, 30-50 mm. long, 3-6 mm. broad, gradually narrowed towards the petioles; petioles 1/3-1/2 the length of lamina; lower cauline leaves more or less like the basal leaves, upper cauline leaves abruptly reduced in size, sessile, lanceolate to linear-lan- ceolate, 12-20 mm. long, 2—4 mm. broad; all the leaves at first densely, later loosely covered with trichomes. Inflorescence usually terminal, rarely lateral, binate or ternate, congested, short in flower, elongated in fruit to 24 cm., bracteate, lower bracts to 9 mm., upper bracts to 2.5 mm. long, linear-lanceolate. Pedicels 1-1.5 mm. long in flower, elongated in fruit to 4.5 mm. Calyx divided to the base, lobes ca. 3 mm. long in flower, enlarged to 5(—5.5) mm. in fruit, 0. 5 mm. broad, linear, acute, hairy, trichomes subappressed. Corolla white, ‘ifvediballifocni-caninane- late, 44.5 mm. long, tube 1 mm. long, limbs subpatent, 4 mm. in di- ameter, lobes suborbicular, 2 mm. long; faucal appendages 0.7 mm. long, narrow, rounded at apex. Nutlets 2—2.5 mm. long, ovate-pyramidal, brownish, appendiculate at margins, appendages few, free at base, 0.5— 0.8 mm. long, erect, bearing glochidia at apex, dorsal middle area dis- tinctly carinate, otherwise tuberculate. Style 1 mm. long DIsTRIBUTION: Afghanistan, West Pakistan. West Pakistan: Reported from: Kurram Valley: Mont. Sikaram, 3000 m., Aitchison 107 (Riedl, l. c, 48: 83. 1967). I did not see any specimen of Lepechiniella albiflora, but from its original description it appears to be a well distinguished species. In our area it is related to L. inconspicua in the size and form of the nutlets, but differs in having much shorter (up to 4.5 mm.) pedicels in fruit, compared to the much longer (to 10 mm.) pedicels in L. inconspicua, much longer (5 mm.) and narrower (0.5 mm.) calyx lobes compared to 1970] KAZMI, BORAGINACEAE 515 those of the latter species (2-3 mm. long and 1 mm. broad), and large, white corollas. Both Lepechiniella inconspicua and L. albiflora may easily be dis- tinguished from Lepechiniella microcarpa by their wings not folded at the margins of the nutlets. 17. Heterocaryum A. DC. in DC. Prodr. 10: 144. 1846. Echinospermum Lehm. Sect. 1. Heterocaryum (A. DC.) Boiss. FI. Orient. 4: 247. 1875 Lappula § 2. Heterocaryum (A. DC.) Post & Kuntze, Lexicon 216, 1914. Type species: H. laevigatum (Kar. & Kir.) A. DC. (lectotype species). Annual decumbent or erect herbs. Stem, branches, and leaves covered loosely or densely with trichomes usually arising from tuberculate bases, rarely subglabrous. Inflorescence bracteate, elongating in age. Cal divided to the base, lobes enlarged in fruit. Corolla infundibuliform to campanulate with 5 distinct trapeziform faucal appendages. Style very short, stigma capitate. Nutlets 4, homo- or heteromorphic, attached to a narrow columnar gynobase throughout their length, not separable, variously appendiculate or aculeate at margins. Species 7, distributed from Turkey to central Asia. Kev TO THE SPECIES & . Fruits sessile to indistinctly short pedicellate. ..........-. 1. H. subsessile. Fruits distinctly pedicellate, pedicels longer than the fruits, thick, fleshy. b. Plants with few scattered trichomes on the stem and leaf-margins; leaves ovate to oblong-ovate; pedicels usually recurved in fruit; nutlets with two series of appendages at the margins, dorsal middle eye ae 7 . A. laevigatum. . Plants usually densely pubescent on stems, branches, and leaves; leaves linear to linear-lanceolate; pedicels usually erect in fruit; nutlets with a single series of appendages at the margins, dorsal middle area not pubescent. c. Plants robust, one pair of the long decurrent nutlets appendiculate at the margins and the middle dorsal area, appendages of the mid lle area sometimes longer than the marginal ones. .....-.-.-.- 3.H rigidum, c. Plants weak, one pair of the long decurrent nutlets appendiculate at the margins, middle dorsal area plicate, plicae beaded, smooth, rarely appendiculate, appendages (if present) always shorter than the mar- inal ONE ics opine Rumen irk ahe a Ae 4. H. szovitsianum. o 1. H. subsessile Vatke, Zeitschr. Ges. Naturw. 11: 129. 1875; Riedl in Rechinger, Fl. Iranica 48: 85. 1967. Echinospermum oligacanthum Boiss. Fi. Orient. 4: 248. 1875, non Ledeb. 1851. 516 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Lappula echinophora var. sessilis O. Kuntze, Acta Horti Petrop. 10: 215. 1887. Cynoglossospermum oligacanthum (Boiss.) O. Kuntze, Rev. Gen. Pl. 2: 436. 1891 Echinospermum echinophorum (Pall.) Bernm. var. sessile Lipsky, Acta Horti Petrop. 26: 539. 1910. Heterocaryum echinophorum (Pall.) Brand var. oligacanthum (Boiss.) Brand, Pflanzenr. IV. 252 (Heft 97): 95. 1931. Heterocaryum oligacanthum (Boiss.) Bornm. Beih. Bot. Centralbl. 59-B: 306. 1939; M. Pop. in Fl. URSS. 19: 546. 1953. Type: In Persia orientali prope Shahrud et Ssertschah, Bunge s.n. (B). Annual, erect herb to 35 mm. tall. Stems simple or divaricately branched; stem and branches pubescent, trichomes stiff, spreading. Basal and lower cauline leaves short petiolate, linear to linear-lanceolate, ob- tuse to acutish, 10-40 mm. long, 1-3 mm. broad, covered on both surfaces with stiff, appressed to patent, white trichomes arising from tuberculate bases, upper cauline leaves sessile, shorter, and narrower. Inflorescence leafy, short in flower, much elongated in fruit. Flowers sessile, fruits subsessile to very short (to 1.5 mm.) pedicellate. Calyx divided to the base, lobes in flower ca. 1 mm. long, linear, in fruit enlarging to ca. 5 mm. long, 1 mm. broad. Corolla blue, campanulate, ca. 2 mm. long. Fruits narrowly prismatic, 3-(4-5) mm. long, 2-3 mm. in diameter; nutlets heteromorphic, one pair of opposite nutlets longer, decurrent to the short pedicel, ca. 3.5 mm. long, 0.5-0.7 mm. broad, bearing 2-3 appendages at the lateral margins and 3-6 at apex, appendages 1-2 mm. long, glochidiate at apex, middle dorsal area longitudinally plicate, plicae swollen, beaded, glabrous or rarely with 1 or 2 prickles; the other pair of nutlets, exclud- ing wings 2—2.5 mm. long, 0.5—-1 mm. broad, not decurrent to the pedicels, bearing usually 2 appendages on the lateral margins and one at apex, appendages 1—2.5 mm. long, broadly triangular, confluent at base, form- ing marginal wings 0.2-1 mm. broad, puberulous, glochidiate at apex; wings dentate to aculeate at the margins, puberulous on both surfaces; dorsal middle area carinate, glabrous or sometimes slightly tuberculate to aculeolate. Style conspicuous, ca. 0.3 mm. long, stigma capitate. DistRIBUTION: Iran, Afghanistan, Turkestan, Pamir Alaj, West Pakis- West Pakistan: KaLat State: Chauki Guru, between Kalat and Mastung, Jafri & Akbar 1820 (£). Quetta Dist.: Chiltan mountains, between Dulai and Kanak, 1600 m., Kazmi 1547 (PEs); Rechinger 29101 (w); Quetta to Chaman, 30 km. northeast of Quetta, low stony slopes, Lamond 958b (E). 2. H. laevigatum (Kar. & Kir.) DC. Prodr. 10: 145. 1846; Ledeb. FI. Rossica 3: 164. 1847; Brand, Pflanzenr. IV. 252 (Heft 97): 97. 1931; M. Pop. in Fl. URSS, 19: 539. 1953; Riedl in Rechinger, FI. Tranica 48: 88. 1967. Echinospermum laevigatum Kar. & Kir. Bull. Soc. Nat. Moscou 15: 411. 1842; Bunge in Reliq. Lehm. 411. 1851; Boiss. Fl. Orient, 4: 248. 1875. 1970] KAZMI, BORAGINACEAE 517 Type: In montosis apricis Songoriae ad rivulum Ai, nec non in fossis arenosis, prope fontem Sassyk-pastan, Karelin & Kiriloff s.n. (1x-holo- type, GH-isotype). Jcom.: DM. Pop: lc. 19: £..26. fie. 2. 1953, Annual herb to 40 cm. tall. Stem simple or divaricately branched, glabrous or with few scattered, short, subappressed trichomes. Basal leaves few, ovate, subsessile, 15-20 mm. long, 8-12 mm. broad; lower cauline leaves, sessile, oblong-ovate, obtuse, 20-30 mm. long, 6-10 mm. broad, lower surface usually glabrous, upper surface, (especially at the margins) with few, scattered, short, soft, white trichomes arising from minute tuberculate bases; upper cauline leaves similar to the lower, shorter, narrower and acutish. Inflorescence short in flower, later much elongated, bracteate, bracts small. Pedicels 1-3 mm. long in flower, to 16 mm. long in fruit, horizontally patent or later recurved. Calyx divid- ed to the base, ca. 1.5 mm. long in flower, enlarged in fruit, ca. 2.5 mm. long. Corolla blue, minute. Fruits ovoid, to 6 mm. long; nutlets 4, strongly heteromorphic, one pair of opposite nutlets much larger than the other, all nutlets similar in shape and bearing two series of appen- dages at their margins, the outer series composed of 3—4(—5) appendages at the lateral margins and one at the apex, sometimes with one or two very small ones between, appendages more or less dilated at base, 0.5— 2 mm. long, reduced in length from the base of the nutlet to the apex, glochidiate at apex; inner series composed of numerous closely set ap- pendages, not exceeding 0.5 mm, in length, more or less dilated at base, sometimes confluent and forming a narrow lacerated wing, dorsal middle area densely pubescent, carinate. Style very short; stigma capitate. DistripuTion: Iran, Afghanistan, USSR, West Pakistan. West Pakistan: QueTtA Dist.: Sheila Bagh, on rocky slopes, Kazmi 15386 (PEs); between Bostan and Saran Tangi, 1800 m., Kazmi 1503b (pes); Rechinger 29196 (w). The presence of two series of appendages at the margin and the densely pubescent middle dorsal area of the nutlets of Heterocaryum laevigatum, distinguish this species from all the others in our area. 3. H. rigidum DC. Prodr. 10: 145. 1846; Ledeb. Fl. Rossica 3? 163. 1847; M. Pop. Fl. URSS 19: 540. 1953; Riedl in Rechinger, FI. Iranica 48: 87. 1967. Echinospermum heterocaryum Bunge in Mem. Acad. Sci. St. Pétersb. 7: 411. 1854. E. szovitsianum Boiss. Fl. Orient. 4: 248. 1875, pro parte non Fisch. & C. A. Mey. 1835. E. minimum sensu C. B. Clarke in Fl. Brit. India 4: 162. 1883, non Lehm. Heterocaryum echinophorum (Pall.) Brand var. minimum (Lehm.) Brand Pflanzenr. IV. 252 (Heft 97): 95. 1931, pro parte. 518 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Type: Sibiria altaica orientalis, Karelin & Kiriloff 1951 (c-isotype). Icon.: M. Pop. 1. c. ¢. 26. fig. 3. 1953 Annual, erect or decumbent herb, branched above the base, 15-36 cm. tall. Stem and branches brown, to 2 mm. in diameter, covered with an- trorsely appressed, white trichomes to 0.5 mm. long, intermixed with stiff, patent trichomes, arising from tuberculate bases, 1-1.3 mm. long. Basal leaves sessile, linear, to 40 mm. long and 5 mm. broad, obtuse, covered loosely on both surfaces with white, stiff, patent trichomes 0.5—1.3 mm. long, arising from tuberculate bases; middle and upper cauline leaves similar to the basal ones, shorter and narrower. Inflorescence short in flower, much elongated in fruit, bearing fruits 5-15 mm. apart. Pedicels very short in flower, elongated in fruit to 10 mm., 1 mm. thick, gradu- ally attenuated towards the base. Calyx divided to the base, lobes 1.5— 2 mm. long, hairy, elongated in fruit to 3-5 mm. Corolla blue, cam- panulate, 2—2.5 mm. long, sags exceeding the calyx. Fruits ovoid, 5-6 mm. long, 3-4 mm. broad; nutlets 4, heteromorphic, one pair of opposite nutlets oblong, ca. 5 mm. a ng, 1-1.5 mm. broad, decurrent to the pedicel, bearing 3-5 appendages at the lateral margins, 3-5 at their apices, dorsal middle area with two to three parallel rows of 3-7 ap- pendages, appendages to 1.5 mm. long, not at all to slightly dilated at base, puberulous on the lower half, bearing sage at apex; nutlets of the other opposite pair, ovate, 3-4 mm. long, 1.5-2 mm. broad, not decurrent, bearing 3-4 appendages on the ees margins, 1-3 at apex and 1 at base, appendages 0.5-1.5 mm. long, dilated at base, sometimes confluent, forming a narrow wing, glochidiate at apex, dorsal middle area carinate, tuberculate to minutely aculeate, not appendiculate; both types of nutlets usually tuberculate externally. Style very short, stigma capitate. DIsTRIBUTION: Iran, Afghanistan, USSR, West Pakistan. West Pakistan: PESHAWAR Dist.: Peshawar, Deane s.n. (K). Quatra DIST.: Khanai, 1800 m., Duthie s.n. (K); Sariab Road, 5 km. from Quetta, Lamond 926 (£); Quetta, Pishni forest nursery and surroundings, Kazmi 14356 (PES). RAWALPINDI Dist.: Hurroo, Aitchison 1085 (kK). Heterocaryum rigidum is closely related to H. szovitsianum from which it is distinguished by its decurrent pair of opposite nutlets which are ap- pendiculate in the middle dorsal area, with appendages more or less the same as those on the margins. 4. H. szovitsianum (Fisch. & C. A. Mey.) DC. Prodr. 10: 145. 1846; Ledeb. FI. Rossica 3: 163. 1847; M. Pop. Fl. URSS 19: 546. 1953; Riedl in Rechinger, Fl. Iranica 48: 86. 1967 Echinospermum szovitsianum Fisch. & C. A. Mey. in Sem. Hort. Petrop. 2: 36. 1835; Boiss. Fl. Orient. 4: 247. 1875, pro par Heieroceryum pachypodum DC. Prodr, 10: 144. a H. pachypodum DC. var. kotschyanum DC. Prodr. 10: 145. 1846. 1970] KAZMI, BORAGINACEAE 519 H. minimum (Lehm.) DC. var. Szovitsianum Regel, Acta Horti Petrop. 6: 344. 1880. Lappula echinophora var. szovitsiana O. Kuntze, Acta Horti Petrop. 10: 214. L. echinophora var. pachypoda O. Kuntze, /bid. L. szovitsiana (Fisch. & Mey.) Druce, List Brit. Pl. 50. 1908. Heterocaryum echinophorum (Pall.) Brand var. minimum (Lehm.) Brand, Pflanzenr. IV. 252 (Heft 97): 95, 96. 1931, pro parte. H. echinophorum (Pall.) Brand var. pachypodum (DC.) Brand, loc. cit. 96. Type: In Persia boreali ad marginis agrorum, Szovits s.n. (LE). Icon.: M. Pop. 1. c. 19: #. 26. fig. 1. 1953. Annual, erect to subdecumbent branched herb, to 30 cm. tall. Stem and branches covered with stiff, usually patent to sometimes subappressed trichomes to 1.5 mm. long. Basal leaves linear, oblong-spathulate or some- times ovate, obtuse, petiolate, petioles equalling the lamina, (including petioles) 10-30 mm. long, 3-6 mm. broad, covered loosely on both surfaces with stiff, usually patent, white trichomes to 1.5 mm. long, arising from tuberculate bases; cauline leaves sessile, lanceolate to linear-lanceolate, obtuse, to 40 mm. long and 7 mm, broad; upper cauline leaves shorter and narrower. Inflorescence leafy, elongating in age; pedicels 1-3 mm. long in flower, elongated in fruit to 11 mm., hairy. Calyx divided to the base, lobes 3-4 mm. long and 0.5 mm. broad in flower, linear, enlarging in fruit to 11 mm. long and 1.5 mm. broad. Corolla blue, campanulate 2.5—3.5 mm. long. Fruits ovoid, 4-8 mm. long, to 5 mm. broad; nutlets 4, hetero- morphic, one pair of opposite nutlets, oblong, decurrent to the pedicel, 6-7 mm. long, 1-1.5 mm. broad, bearing 2-4 appendages at the lateral, thickened margins and 1-5 at the apices, appendages 0.5—1 mm. Jong, distant, not dilated at base, glochidiate at apex, dorsal middle area carinate, plicate, plicae swollen irregularly at places, glabrous; the other pair of opposite nutlets not decurrent, nutlets oblong, 5—6 mm. long, ca. 1.5 mm. broad, bearing 4-5 appendages at the lateral margins and usually 1 at apex, appendages 1-3 mm. long, much dilated and confluent at base, forming an irregular wing up to 1 mm. broad, glochidiate at apex, marginal wing and lower part of appendages pinkish brown, puberulous; dorsal middle area carinate, tuberculate, tubercles arranged longitudinally, gla- brous or puberulous, rarely aculeolate. Style short, stigma capitate. DistrisuTion: Turkey, Syria, Iraq, Iran, south and southwest USSR, Tien Shan, Afghanistan, West Pakistan, Kashmir. West Pakistan: CHITRAL STATE: Birmogh Lasht, 2600 m., Bowes Lyon 719 (pm); Drosh, 1200 m., Stainton 2271 (BM). QUETTA Dist.: Pishni Ee Nursery and surrounding fields, Kazmi 1425 (PES) ; environ de Quetta, Scam : 1568 (micH); Ziarat to Manna, 2600 m., Jafri & Akbar 2192 (ze), Reporte from: NortH Waziristan AcEeNcY: Datta Khel, Blatter & Fernandes s.n. Kashmir: Miscellaneous: Himal. Bor. Occ., 6000 ped., Herb. Ind. Or. Hook. f. & Thoms., Thomson s.n. (GH). Reported from: Sumbal, Meebold 4197. 520 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Riedl (in Rechinger, Fl. Iranica 48: 89. 1967) has described Hetero- caryum X irregulare, a hybrid between H. rigidum DC. and H. macro- carpum Zak. The new hybrid is based on a specimen (Kabul, Sher Darwasa, 1840m., Gilli 3158) collected in Afghanistan while the only other specimen cited is (Quetta, Sariab, 1700 m., Rechinger 28840) from West Pakistan. I did not see Rechinger 28840 but have examined Kazmi 1435b and Lamond 926, which were collected at the same time and from the same locality with Professor Rechinger. These specimens are H. rigidum and show no affinity with H. macrocarpum. Moreover, as I did not find the other parent in this area, the presence of the hybrid in West Pakistan appears to be quite doubtful. [To be continued | 1970] PRANCE, CHRYSOBALANACEAE 521 THE GENERA OF CHRYSOBALANACEAE IN THE SOUTHEASTERN UNITED STATES 1 GHILLEAN T. PRANCE CHRYSOBALANACEAE R. Brown in Tuckey, Narrat. Exped. Zaire Congo. 433. 1818, nom. cons. Small trees [to large ones], shrubs or subshrubs, with simple, entire, alternate, stipulate, pinnately veined leaves. Inflorescence of terminal and subterminal cymose panicles or terminal and axillary cymules [racemose, spicate, paniculate]. Flowers perfect [rarely polygamous], perigynous, actinomorphic [to markedly zygomorphic]. Sepals 5, regular [often unequal], distinct, imbricate in aestivation, Petals 5 [rarely apetalous}, inserted on margin of disc, equal in size [unequal], imbricate in aestiva- tion [rarely clawed]. Disc always present, forming a lining to floral tube. Stamens numerous, 12-26 [few to numerous, 2-300], inserted around margin of disc [or unilaterally], all fertile [or some reduced to long or short staminodia]; filaments elongated filiform, free or united below up to half their length in small groups [short to elongated, rarely ligulately connate], slightly exserted beyond calyx lobes [included to far exserted |, glabrous or pubescent; anthers bilocular at anthesis, longitudinally dehis- cent; pollen medium sized, with 3 furrows [rarely 4], oblate-spheroidal to subprolate, triangular in polar view, with little wall patterning, scabrous to verrucose. Gynoecium unicarpellate [basically of three, but usually only one developed], inserted at base of floral tube [from base to mouth of floral tube], sessile [rarely with a short gynophore| ; style single, basally attached; stigma 3-lobed [or truncate]; ovary superior, 1-locular with 2 basally attached erect anatropous ovules [rarely 2-locular with one ovule in each locule]. Fruit a fleshy [or dry] drupe. Seed lacking endosperm, embryo large, with plano-convex [or amygdaloid or ruminate] cotyledons. Type GENUS: Chrysobalanus L. An almost exclusively pantropical family of 17 genera and about 430 species; only Chrysobalanus and Licania extend into the southeastern United States. The family is divided into tribes Chrysobalaneae and Hirtelleae; both genera of our area belong to the former. The family has usually been considered to be a subfamily or tribe of i i ject of the * Prepared for a generic flora of the southeastern United States, a projec ; Arnold Arboretum and the Gray Herbarium of Harvard University made possible through the support of the National Science Foundation (Grant GB-6459X, principal : ‘ : i in the first paper (Jour. Arnold published in the series, follows the format established in t } ‘ Arb. 39: 296-346. 1958). The area covered includes esse oe South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, and Louisiana. Bt | are based primarily on the plants of this area, with additional se sanacagy brackets. References that have not been seen and verified are marked by an ES . feook § The plate was drawn by Miss Alma Hochhauser. I wish to than r. ood for his advice and helpful suggestions during the preparation of this manuscript. 522 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 the Rosaceae, but it is markedly distinct from that family, differing in the gynobasic style, the erect ovule, the tendency toward zygomorphic flowers, the rubiaceous stomata, the presence of silica, the pollen morphol- ogy, and in numerous anatomical features, especially the secondary xylem. The Chrysobalanaceae form a well-marked and natural family. Various ad oer rate suggested diverse relationships for the family but it seems to belong to the Rosales between the Rosaceae and Leguminosae. Small and a pes other authors have placed the Chrysobalanaceae in the family Amygdalaceae, which consists of the Chrysobalanaceae and Ros- aceae subfam. Prunoideae. This is, however, an unnatural grouping, and the Prunoideae clearly belong with the rest of the Rosaceae, rather than with the Chrysobalanaceae. The anatomy of these two groups is very different, and the morphological features given above also separate them. The generic limits within the family have recently been redefined on a worldwide basis (Prance, 1969, & in press), and as now defined they are relatively distinct. The result of this work shows that two genera are represented in our area. One species previously considered a member of Chrysobalanus or as the separate genus Geobalanus has been transferred to Licania, a widespread, predominantly neotropical genus. REFERENCES: BAILLON, H. Monographie des Rosacées. Hist. Pl. 1: 345-483. 1869. [Série des Icaquiers, 425-439; Chrysobalaneae, 480-483 BENTHAM, G. Contributions towards a flora of South America. — Enumeration of plants collected by Mr. Schomburgk in British Guiana. Hook. Jour. Bot. 2: 210-223. 1840. [Chrysobalanaceae, 210-222.] & J. D. Hooker. Rosaceae. Gen. Pl. 1: 600-629, 1865. [Treatment by Hooker; Chrysobalaneae, 606-609. | Bonne, G. Sur la constitution du gynécée chez les Chrysobalanées. Compt. Rend. Acad. Sci. Paris 182: 1404-1406. 1926. . Recherches sur le pédicelle et la fleur des Rosacées. 380 pp., 10 pls. bans 1928. CaNDOLLE, A. P. dE. Rosaceae. Prodr. 2: 525-639. 1825. [Tribe Chryso- TNO 525-529. ]| Cronquist, A. Evolution and classification of flowering plants. xii + 396 pp. Boston. 1968. [Rosales, 229-236; Chrysobalanaceae, 235. Duke, J. A. On tropical tree seedlings. I. Seeds, seedlings, systems, and sys- tematics. Ann. Missouri Bot. Gard. 56: 125-161. 1969. [Chrysobalanaceae, 148. EIcHLER, A. W. Rosaceae. Bliithendiagramme 2: 495-514. 1878. [Chrysobal- aneae, 511-514 Focke, W. O. Rosaceae. Nat. Pflanzenfam. III. 3: 1-48. 1888; 49-64. 1891. [ Chrysobalanoideae, 55-60. Fritscn, C. Ueber ee Noeernace der Chrysobalanaceen. Verh. Zool.-Bot. Ges. Wien 38: 93-95. 1 . Beitrage zur ee der Chrysobalanaceen. I. Conspectus generis Licaniae. Ann. Naturh. Hofmus. Wien 4: 33-60. 18 GrRISEBACH, A. H. R. Flora of the British West Indian lidantds: xvi + 789 pp. London. 1860. [Chrysobalaneae, 229-231.] 1970] PRANCE, CHRYSOBALANACEAE 523 GUNDERSEN, A. The families of dicotyledons. xviii + 237 pp. Waltham, Mass. 1950. [Chrysobalanaceae, 109.] Hauman, L. Contribution a l'étude des Chrysobalanoideae africaines. Bull. Jard. Bot. Bruxelles 21: 167-198. 1951. HvutTcuHInson, J. Rosaceae. Gen. Fl. Pl. 1: 174-216. 1964. [Chrysobalaneae, 190-193. ] Kuster, E. Die anatomischen Charaktere der Chrysobalaneen, insbesondere ihre Kieselablagerungen. Bot. Centralbl. 69: 46-54, 97-106, 129-139, 161-169, 193-202, 225-235. pl. 1. 1897. Lrnotey, J. A natural system of botany. ed. 2. xxvi + 526 pp. London. 1836. { Chrysobalanaceae, 158, 159.| Metcuior, H. Chrysobalanaceae. Engler’s Syllab. Pflanzenfam. ed. 12. 2: 219, 220. 1964. Morvittez, F. La trace foliare des Chrysobalanées. Compt. Rend. Acad. Sci. Paris 166: 859-861. 1918. Prance, G. T. The taxonomy and ecology of the Chrysobalanaceae of the Amazon Basin. Jn: H. Lint, ed. Atas do Simpésio sdbre a Biota Amaz6nica 4: 209-228. 1967. . A synopsis of Chrysobalanaceae. (In press.) Oxford. . D. J. Rocers, & F. Wurte. A taximetric study of an angiosperm family: generic delimitation in the Chrysobalanaceae. New Phytol. 68: 1203-1234. 1969. Sma, J. K. Manual of the southeastern flora. xxii + 1554 pp. New York. 1933. (Reprinted by Univ. N. Carolina Press, Chapel Hill.) [Amygdalaceae, 645, 646. tes WetrsTEIN, R. Handbuch der systematischen Botanik. x + 1152 pp. Leipzig & Wien. 1935. [Chrysobalanaceae, 771, 772.] Key TO THE GENERA OF CHRYSOBALANACEAE IN THE SOUTHEASTERN UNITED STATES Plants shrubs or small trees; staminal filaments pubescent, joined in small groups up to half their length; endocarp longitudinally ribbed (costate) ; inflorescences MEUNAIY: ok Ne ein ee SS Ea ee ee 1. Chrysobalanus. Plants suffruticose, low, colonial, spreading by woody underground ae staminal filaments glabrous, connate at base only; endocarp smooth, not be bec : inflorescences terminal and subterminal. .........-----++s-- 0000 2. Licania. 1. Chrysobalanus Linnaeus, Sp. Pl. 1: 514. 1753; Gen. Pl. ed. 5. 299. 1754. Small to large shrubs, or rarely small trees, with coriaceous pinnately veined leaves, often with two glands at base of blade. Inflorescence few flowered, terminal or axillary, either a short raceme of cymules or cymose throughout, or, when consisting of about 6 flowers, a false raceme. Sepals acute, pubescent. Petals equaling sepals, glabrous, white. Stamens 12-2 6, exserted beyond sepals, some often shorter than others, inserted in ; complete circle around disc [slightly unilateral] ; filaments united in igen groups for up to half their length [free almost to base], densely airy. Style inserted at base of ovary, pubescent for most of its length; ovary pilose, inserted at base of floral tube, 1-locular, with two erect ovules. 524 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 A HocHnauser FIGURE 1, CHRYSOBALANACEAE. a-j, Chrysobalanus. C. Jcaco: a, habit, X %; b, flower, X 4; c, flower in vertical section, x 4; d-h, petals, x 4; i, fruit, 1; j, fruit in vertical section, X 1. k-t, Licania. L. Michauxii: k, habit, < %; 1, flower, X 4; m, flower in vertical section, < 4; n-r, petals, X 4; s, fruit, X 1; t, fruit in vertical section, < 1. é 1970] PRANCE, CHRYSOBALANACEAE 525 Fruit a small ellipsoid fleshy drupe, smooth, longitudinally ribbed; endo- carp thin, hard, with 4-8 prominent longitudinal ridges corresponding to lines of fracture that allow the seedling to escape, glabrous within, filled by the large plano-convex cotyledons. Germination hypogeal. Type species: Chrysobalanus Icaco L. (Name derived from Greek chrysos, golden, and dalanos, acorn or fruit.) A primarily tropical genus of four species, two confined to Africa and one to the West Indies. The fourth, Chrysobalanus Icaco, cocoa plum, is a widespread coastal species from subtropical Florida through the Carib- bean and Central America to eastern South America and West Africa. In the southern part of peninsular Florida C. /caco is a common shrub or tree of beaches, sand dunes, coastal hammocks, hammock islands in the Everglades, and cypress-heads. The extreme variation in leaf shape and size and fruit size in Chryso- balanus Icaco has led to the description of numerous taxa throughout its range (e.g., in our area, C. interior Small). There is, however, little cor- relation of the variable characters, and there seems to be no basis for recognition of more than one species. In at least Trinidad, Suriname, and Brazil, the different forms are frequently found growing side by side without any ecological separation (obs. Prance). The leaves vary from extremely small to large and from orbicular to elliptic, and both shape and size are quite variable on the same plant. The very variable fruit is from 0.8 to 4.5 cm. long. The larger fruits are more conspicuously ribbed than the smaller ones, and they tend to have a thicker, more fleshy mesocarp. In addition, the mature fruit may be deep red or purple to black or yellow. (To judge from the name of the genus, the material described by Lin- naeus must have been of the yellow-skinned form.) The fruit is edible, but only the larger, more fleshy fruits borne by some plants make good eating, the differences in fruit type being comparable to those between good and bad varieties of plum (Prunus domestica L.). Where C. Icaco is used commercially there is obviously some selection of plants. For the present, it seems best to maintain only a single polymorphic Species, for it is impossible to subdivide this species on the basis of herbarium material and ecology alone. An experimental study of plants of the pane aah phological types grown in controlled environments should yield muc additional information. Attempts to examine the chromosomes of C. Icaco have thus far proved unsuccessful. : The fruits of Chrysobalanus Icaco are commonly used for aeons Venezuela and Colombia, but only occasionally in our area. Plants of this species are sometimes grown as ornamentals. glia hail is most closely related to Licania, a ee tropical genus. The differences between the two are small, nak ap “i i these genera on a worldwide basis shows that they are best = A * Chrysobalanus differs from Licania in the ridged endocarp of sea : a the hairy filaments that are joined together in small groups, and 1 inflorescence. 526 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 REFERENCES: Under family references see BENTHAM & HOOKER (p. 606), BoNNE (1928, pp. 163, 244-346, 356), Focke (p. 57), GrRISEBACH (p. 229), HuTcHINson, KUSTER (pp. 135-137), PRANCE (1967, p. 216; 1970, in press), and SMALL (p. 645). HaumMANn, L. Contribution a l’étude des Chrysobalanoideae africaines, Bull. Jard. Bot. Bruxelles 21: 167-198. 1951. [C. Icaco & C. orbicularis Schu- Howarp, R. A. Notes on Rosaceae in the Lesser Antilles. Jour. Arnold Arb. 45: 279-283. 1964. [C. cuspidatus Griseb. (Licania oligantha A. C. Sm.), 279-281.] Hurtcuinson, J., & J. M. Datztet. Chrysobalanus, Fl. W. T rop. Afr. ed. 2. 1: 426, 427. 1958. [3 spp.; ed. 2 revised by R. W. J. Kray. SMALL, J. K. Notes on Chrysobalanus Icaco L. Torreya 10: 249. 1910. [Notes yellow, purple, or red fruits, the yellow largest, red smallest. TAvarES, S. Contribuicao para o estudo botanico de “guajeru” (Chrysobalanus Icaco L.). (English summary.) Inst. Pesquisas Pernambuco Agron. Publ. Il. 1. 12 pp. 1957.* 2. Licania Aublet, Hist. Pl. Guiane Frang. 1: 119. pl. 45. 1775. Subshrubs [shrubs to large trees] with coriaceous [membranaceous to chartaceous| pinnately veined leaves. Inflorescences rather lax branched terminal and subterminal cymose panicles [or a sparsely branched panicle of racemes, a panicle, or a spike]. Sepals acute [or rounded], pubescent [to glabrous]. Petals equaling sepals [larger or smaller than sepals, or absent], pubescent [glabrous]. Stamens 14-17 [3-40], exserted slightly beyond sepals [included to far exserted], inserted in a complete circle around disc [unilateral]; filaments united at base only [free to base], glabrous [rarely hairy, very short and included]. Style inserted at base of ovary, glabrous [to densely hairy]: ovary inserted at base of floral tube, usually 1-carpellate but sometimes 2 or 3 carpels developing, glabrous or sparsely hairy [to densely hairy], unilocular, with two erect ovules. Fruit a small ellipsoid drupe [to large and of varying shape], smooth, glabrous [sometimes verrucose, pulverulent or densely pubescent]; endocarp hard, thin [to thick] terete, not ridged, sparsely hairy |to densely hairy within], filled by the large cotyledons. Germination hypogeal. (Including Mo- quilea Aubl., 1775; Hedycrea Schreb., 1789; Geobalanus Small, 1913; and others.) Type species: Licania incana Aublet. (Name a misspelled anagram derived from the local name in French Guiana, caligni.) Primarily a neotropical genus of 151 species, in three subgenera and eight sections, with one species widespread in Malesia and one confined to our area. Licania Michauxii Prance,? gopher-apple or ground-oak, is a common “ *QLicania Michauxii Prance, nom. nov. C. hrysobalanus oblongifolius Michx. Fl. Bor.- Am. 1: 283. 1811, non Licania oblongifolia Standl., 1917; C. retusus Raf. New FI. N. Am. 3: 26. 1838, non L. retusa Pilger, 1914; C. incanus Raf. loc. cit., non L. incana 1970] PRANCE, CHRYSOBALANACEAE 527 suffruticose shrub with spreading underground stems. It is abundant in pinelands and oak scrubland and on sand hills and sand-dunes in Florida and extends westward to Louisiana and northward to South Carolina. The full reasons for treating this species as a Licania instead of a Chrysobalanus will be given elsewhere (Prance, in press). The two genera are very closely related but are distinct when L. Michauxii is transferred from Chryso- balanus to Licania. It falls well within the limits of Licania, and there are no grounds whatever for regarding it as a separate genus. It is a member of subg. Moguitea (Aubl.) Prance (Atas Simp. Biota Amazon. 4: 224. 1967), sect. Moquitea (Aubl.) Prance,® and it is most closely related to L. retifolia Blake, of Mexico. This species has previously been included in both Chrysobalanus L. and Geobalanus Small. Its synonyms include C. oblongifolius Michx., C. retusus Raf., C. incanus Raf., G. oblongifolius (Michx.) Small, G. pallidus Small, and C. pallidus (Small) L. B. Sm. Since the specific epithets of all these combinations are already occupied in Licania (i.e., L. incana Aubl., L. pallida Spruce ex Sagot, L. oblongifolia Standl., and L. retusa Pilger) a new epithet was required. Chrysobalanus pruni- folius Raf. has commonly been referred to this species, but type material has not been found, and the small leaves, few-flowered racemes, and podlike fruit described for C. prunifolius are quite unlike those of L. Michauxii. The differences given by Small for his Geobalanus pallidus (leaves densely white-tomentose beneath, ovary pubescent, drupes 3—4 cm. lo vs. leaves and ovary glabrous, fruit 2—2.5 cm. long in G. oblongifolius) do not hold. There is a gradation in the pubescence of both the ovary and the leaf undersurface, and in many cases the pubescence of the leaf is caducous and the older leaves are glabrous. CArysobalanus incanus Rat., based on the variant noted by Michaux under his C. odblongifolius, repre- sents the same pubescent form, which, although conspicuous in its extreme development, does not seem to be worthy of taxonomic rank. Licania Michauxii has often been said to have a stellate pubescence, but it does not. This erroneous report, attributable to Kiister, has led to the mention of stellate pubescence in connection with the family in several other publications. The mistake is based on a mixed collection of Aubl., 1775; Geobalanus pallidus Small, Fl. Miami 81. 1913, non L. pallida Spruce ex Sagot, 1883, nec L. pallida Britton, 1890. : Mads Aabl, a aa (Aubl.) Prance, comb. nov. Moquilea Aubl. Hist. PI. Guiane Franc. 1: 521. pl. 208. 1775. He This sectional combination, cited without an author, was used under Licania subg. Moquilea (Aubl.) Prance in 1967 (Atas Simp. Biota Amazon, 4: 225), since this was the correct procedure under Art. 22, International Code of Botanical Nomenclature, 966. However, changes adopted at the 11th International Botanical Congress, Seattle, 1969, restrict this tautonymic principle to the subgenus and section that include the type of the genus, but recommend that when a new epithet is needed in other — genera and sections the repetitive principle be retained when possible. In conformity with these changes the combination Licania sect. Moquilea is made here formally and is cited with authors. 528 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 herbarium material. The widely distributed A. H. Curtiss 727 includes, in addition to L. Michauxii, sterile branches of Quercus pumila Walt., a plant that has stellate hairs on the lower leaf surface. Quercus pumila may have the same suffruticose habit as L. Michauxti, and sterile specimens could easily be mistaken for it, except for their stellate pubescence. Licania is most closely related to Chrysobalanus (q.v.). It is also close to the African A frolicania Mildbr. and the Malesian Parastemon A. DC Together the four comprise the tribe Chrysobalaneae, which is distinguished by the regular flowers with a basally inserted gynoecium. REFERENCES: Under family references see BONNE (1928, pp. 164, 347, 348), FrRitscH, HutTcHINSON (p. 191), KUSTER (pp. 137, 194-201), PRANCE (1967, pp. 224- 227; 1970, in press), and SMALL (p. 645, 646, as Geobalanus). Apams, D. W. Gopher-root. Gard. Month. 28: 244. 1886. [Humorous non- scientific letter to editor concerning an underground tree identified in editor’s comment following as iE amar oblongifolius. Adams notes spread of individual plant as 80-100 ft. and underground stems to 2.5 inches in Autes, H. E., C. R. BELL, & A. E. Raprorp. Species new to the flora of North or South ‘Carolina. Rhodora 60: 10-32. 1958. [L. Michauxii, as C. oblongi- folius, in Jasper Co., S. C.; see also Manual of the vascular flora of the Carolinas, 569. | Hooker, J. D. Rosaceae. Ju: C. F. P. von Martius, FI. Brasil. ge 1-76. pls. 1-22. 1867. [Licania, 8-19, pls. 1-4; Moquilea, 19-26, 8.] B. A. KrukKorr CURATOR OF teed BOTANY Tue New York Botanical GARD Bronx, NEw York 10458 1970] MILLER, NEW PARIETARIA FROM MEXICO 529 A NEW SPECIES OF PARIETARIA (URTICACEAE) FROM NORTHEASTERN MEXICO! Norton G. MILyter DURING A GENERAL REVIEW of the collection of Parietaria in the com- bined herbaria of the Arnold Arboretum and Gray Herbarium, an un- described species, variously named P. debilis Forst. f., P. floridana Nutt., P. obtusa Rydb., P. officinalis L., and P. pensylvanica Muhl. ex Willd., was recognized among Mexican material. Examination of additional specimens borrowed from other herbaria has established its presence in three, prob- ably four, states in northeastern México. A genus of about 20 species in two subgenera, Parietaria is largely restricted to temperate and subtropical latitudes (or if nearer the equator, then usually montane), with the bulk of the genus occurring in Europe and North America. Regrettably, no monograph has appeared since pub- lication of Weddell’s treatment (1869) in the De Candolle Prodromus. A major taxonomic problem in this genus, pertaining especially to the widespread species, has been the interpretation of the extensive variability in leaf and bract size, pubescence, and habit. Hedberg (1957), for ex- ample, stressing the intergradations of vegetative characters in the four Parietarias reported from the mountains of east Africa, concluded that three of them could be accommodated within the fourth, the wide-ranging Parietaria debilis. In my own experience, and as has been recently em- phasized by the work of Hinton (1968, 1969) and Townsend (1968), vegetative characters may or may not be satisfactory from the taxonomic standpoint, but achene shape and form of the accrescent perianths develop- ing from both perfect and carpellate flowers (should the latter be present) provide the most stable and, therefore, the most useful diagnostic features. Parietaria decoris N. G. Miller, sp. nov. Herbae perennes (?). Caules ascendentes numerosi vel singulares tri- chomatibus et longis laxisque et brevioribus uncatisque sparse vel dense vestiti. Folia petiolata alterna vel opposita vel in plantis juvenilibus sub- *This is another in a series of peripheral papers arising from research toward a Generic Flora of the Southeastern United States supported by the National Science Foundation (Grant GB-6459X, C. E. Wood, Jr., principal investigator). I thank Dr. Elizabeth A. Shaw for checking and improving the Latin description and the curators of the herbaria at the Texas Research Foundation (11), the University of nian (micH), The New York Botanical Garden (Nv), the University of Texas (TEX), the University of California, Berkeley (uc), the U. S. National Museum (vs), the cul Nacional de Ciencias Biolégicas, Instituto Politécnico Nacional, México, D. ENCB), and the Instituto de Biologia, apie ges Nacional Auténoma de © (mMExu) for sending specimens upon which this study is partly based. The Tnisicsiion is the continuing Gans of Virginia Soa 530 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 opposita; laminae ovatae ad basin cuneatae infra pilosae; nervi laterales prominentes suboppositique pinnatim in paribus duobus dispositi, uno e basi costae, altero e medio exoriente; cystolithi bacilliformes non nisi in pagina supera folii praesentes. Inflorescentiae cymulosae pauciflorae et bracteatae; bracteae discretae linearesque ad apicem obtusae, ciliis et cysto- lithis bacilliformibus instructae, quam perianthium fructum maturum cingens breviores. Flores perfecti; perianthium viride uniseriale ad basin tubulare autem apicem versus lobos 4 libros praebens; stylus 1 apicalis filiformisque stigmate mepielr Achenia fusca subovoidea symmetrica bilat- eraliter, ventricosa, per basin persistentem stylo apiculata. Perianthium accrescens; pars infera (14— 3 longitudinis totae) fusca, 4-angulata et in quoque angulo trichomatibus numerosis lenibusque extrinsecus patentibus praedita; pars supera ex lobis naviculiformibus viridibus, his trichomatibus uncatis cystolithisque bacilliformibus instructis, constata Holotypus: C. H. & M. T. Mueller 645 (cx). Many- or single-stemmed, ascending, perennial (?) herbs. Stems green, sometimes reddish, sparsely to densely covered with long, lax hairs and shorter hooked ones. Leaves petiolate, alternate or opposite to subopposite in young plants; blades ovate, with a cuneate base, size variable, (1.4—) 2.2—5.8(—6.5) cm. long, (0.7—)1.0-3.0(-3.9) cm. wide, weakly pilose; venation pinnate, 2 pairs of prominent, subopposite lateral veins per leaf, one pair originating from near the base of the blade, the other near the middle; cystoliths bacilliform (rarely ovoid or spherical), restricted to the adaxial leaf surface. Inflorescences few-flowered bracteate cymules, two per node, one borne on either side of the petiole base; bracts separate, linear, ciliate, apex blunt, bacilliform cystoliths prominent, bracts shorter than the perianth surrounding the mature fruit. Flowers perfect, strongly protogynous; perianth green, uniseriate, fused basally, but with 4 free, apical lobes, which remain cupped around the stamens until pollen dis- persal; stamens 4, filaments inflexed, anther sacs caducous, filaments re- maining within the enlarging perianth; gynoecium unicarpellate, stipitate; style 1, apical, filiform, constricted near the ovary apex, stigma linear; style (and stigma) falling off before pollen is shed. Achene shiny, brown, + ovoid, 1.0-1.3 mm. long, 0.6—-0.8 mm. wide, bilaterally iyeinctrical, ventricose, stipe off-center in lateral view (see Ficu URE 1, h); style base persisting as a short, hard, + apical projection, Accrescent fruiting peri- anth loosely surrounding the mature achene, both probably dispersed together; continuous basal portion brown, 4-angled, with abundant, soft hairs projecting outward from the angles, apical lobes green, boat-shaped, bearing hooked hairs and bacilliform cystoliths. (Name from Latin, orna- mented, in reference to the lobes of the accrescent perianth.) Specimens examined. xico. COAHUILA: La Mariposa pear: Muzquiz, Marsh 1040 (TEX; see aneen comment), Nuevo LEON: n dense, moist wood, waterway below Alamar, ca. 15 mi. sw. of Galeana, eh ene 1s ae Mueller & Mueller 645 (holotype GH, isotype MICH); growing on side of cliff, oak zone, 1970] MILLER, NEW PARIETARIA FROM MEXICO 531 Quelite, Cerro de la Silla near ea ca. 3800 ft., White 1489 (GH, MICH); Hacienda Vista Hermosa, 35 mi. s. of Mo onterrey, ca. 2400 ft., idem 1567 (mounted ate one plant of don pensylvanica, GH); cliff in limestone canyon, El Diente Canyon, s. of Monterrey, 1200-1300 m., Pennell 16869 (xy, us); wooded ravine, Horse Tail Falls, Santiago Municipality, Chase 7782 ice, MICH); moist cracks in rock, “chute” below cave, Huasteca Canyon, ca. 15 m se. of Monterrey, Ward 5727 (micH); Sierra Madre Mountains, Sehr Mueller & Mueller 58 (GH); Guajuco, Palmer 1266 (cu, Ny) [as P. officinalis L. ?, see S. Watson, Proc. Am. Acad. 18: 155. 1883]; outside entrance to grotto in Canyon Huajuco, e. of Villa Santiago, Johnson & Barkley 15158M (cH, ny, uc); limestone soil of Cafion Huajuco, e. of Villa Santiago in the Rio San Juan valley, idem 15148M (cH), 16152M (ny). TAMAULIPAS: On gravelly sand in open sun, mountains along Rt. 70, ca. 8 mi. s. of Ciudad Victoria, King 4520 (MICH, NY, UC, US); on rich sand near streams or in cafions, near Victoria, 400 m., Runyon 951 (TEx, us); Cerro de los Armadillos, vic. San José, Sierra de San Carlos, Bartlett 10228 (miIcH, us). SAN Luts Portost: Tropical forest, El Salto, Kenoyer & Crum 4031 (a, MICH); Tamasopo Cafion, Pringle 5085 (cH, MEXU); rocky limestone woods, Espinazo del Diablo, Tamasopo Canyon, 800- 900 m., ’ Pennell 17958 (US). The (1) bilaterally symmetrical, + apiculate, ventricose achenes, (2) bacilliform cystoliths borne in cells of the upper leaf epidermis (and best observed in dried plants), and (3) accrescent perianths in which four green, boat-shaped apical lobes contrast with the brown, tubular, basal portion of the calyx serve to distinguish Parietaria decoris. In shape and coloration, the accrescent perianths of this species are similar to those of P. macrophylla Robins. & Greenm., a species of southern México, which, however, in addition to other differences has black rather than brown achenes. During the period of stigma receptivity, the stamens remain minute and the perianth increases only slightly in size. Until explosive pollen dispersal, which occurs sometime after the style has fallen off, the maturing inflexed stamens are confined by the enlarged perianth lobes. The overall lengthening of the perianth following anther dehiscence apparently occurs in the tissues comprising the basal, tubular portion of the perianth. As far as I know, the bacilliform shape of the cystoliths is unique among described North American species, which otherwise have only punctiform markings. Indeed, cystoliths other than the punctiform type are unusual in the 4 or 5 genera and about 25 species that comprise the rest of tribe Parietarieae Gau Leaf-cystoliths i in plants from shaded, forest sites may be up to 8 times longer than wide, but in plants from open habitats where exposure to the sun is maximal (e.g., King 4520), there is a pronounced decrease in the cystolith length/width ratio. Associated with this change is a decrease in leaf size (cf. a, b, & c in Ficure 1). However, in sun-grown plants most leaf-cystoliths remain clearly longer than wide, and the characteristic organization of the lateral veins is retained in the leaves: of such plants. Based on the apparent rarity of bacilliform cystoliths within the genus, the Marsh collection from Coahuila (see above), consisting of only 2 532 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ea (LF 9? d ui). Oe (\sy0 8 eh S Ve oe gy 2 a yy ala eee c Ye ¢ 6 fo oC 0 ela ee [@) 0 Z calf = ° Z = & b FIGURE 1. Parietaria decoris, a, leaf, shade-grown plant (Kenoyer & Crum 4031, A), X 1; b, same (Mueller & Mueller 645, cu, holotype), X 1; c, leaf, plant grown in full sun (King 4520, MIcH), x 1 ; d, cystoliths on adaxial leaf i achene (holotype), x he i achene, rotated 90° counterclockwise from view shown in h (holotype), & 2 juvenile plants without achenes and accrescent perianths, has been pro- visionally named P. decoris. Unfortunately, it is the only specimen from Coahuila assignable to this species to have come to my attention. Parietaria decoris can be placed, somewhat unsatisfactorily, in subg. ParteETArRIA (subg. Euparietaria Komarov in Jarmolenko), members of which have elongated styles and either all flowers of the cymule perfect or the lower ones carpellate and the rest perfect (Jarmolenko, 1936; see also Paclt, 1952). Most species belonging here are perennials. Although Jarmolenko’s map (1941) shows this subgenus to be restricted to Europe and the Middle East, the New World P. macrophylla has an elongated 1970] MILLER, NEW PARIETARIA FROM MEXICO 533 style (with a linear stigma), which is, incidentally, identical to the type occurring in P. decoris. However, the stigmatic apparatus of these two Mexican species differs from the exceptionally thin, glabrous style and the brush-like, apical stigma characteristic of the Old World members of the subgenus. The significance of the two types must await a reevaluation of the infrageneric classification of the genus. Only a very short style, or none at all, supporting a somewhat flattened penicillate stigma, occurs in members of subg. FREIREA (Gaud.) Komaroy in Jarmolenko (later treated as a genus by Jarmolenko, 1941). Lower flowers of an inflorescence are perfect, while upper ones are carpellate. Only annuals, including Parietaria pensylvanica, which ranges across the United States and southern Canada, southward into México, belong to this subgenus. Achenes of P. pensylvanica, although brownish, are non- ventricose. LITERATURE CITED HEDBERG, O. 1957. Afroalpine vascular plants, a taxonomic revision. Symb. t. Upsal. 15. 411 pp. 12 pls. HINTON, B. D. 1968. Parietaria praetermissa (Urticaceae), a new species from the southeastern United States. Sida 3: 191-194 . 1969. Parietaria hespera (Urticaceae), a new species of the southwestern United States. /bid. 3: 293-297. JARMOLENKO, A. V. 1936. Urticaceae. Pp. 384-405. Jn: V. L. Komarov, ed. Flora URSS. Vol. 5. xxvi + 762 pp. Moscow & Leningrad. [Pp. 304-320 in English transl., N. Landau, Flora of the U. S. S. R. Vol. 5. xxvii + 593 pp. Jerusalem. 1970. . 1941, An he at the construction of a system of the genus Parietaria and related forms. (In Russian; English summary.) Acta Inst. Bot. Acad. Sci. URSS. 1. Syst 5: 319-330. Pact, J. 1952. Uber die Identitat von Parietaria ramiflora Monch mit Parie- taria erecta Mertens et Koch (= P. officinalis L.). Phyton Austria 4: 46-50. TOWNSEND, C. C. 1968. Parietaria officinalis and P. judaica. Watsonia 6: 365- 3 WEDDELL, H. A. 1869. Urticaceae. Pp. 32-235%4. Jn: A. De Candolle, ed. Pro- dromus systematis naturalis regni vegetabilis. Vol. 16(1). 1-235%, 236-491. Paris. ARNOLD ARBORETUM Present address: HARVARD UNIVERSITY DEPARTMENT OF BOTANY UNIVERSITY OF NORTH ge CHAPEL Hitt, N. C. 2751 534 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 CHROMOSOME NUMBERS IN THE JUGLANDACEAE A. S. Hans THE JUGLANDACEAE ARE CHARACTERIZED by trees having alternate and pinnate leaves with short-stalked or sessile leaflets. Plants monoecious, flowers unisexual, the staminate flowers in catkins on the twigs of the previous year; perianth typically 4-lobed, sometimes irregularly lobed, 3—many stamens; carpellate flowers solitary or in a small cluster or spike with inferior, bicarpellary, syncarpous, unilocular, or incompletely 2-4- celled ovary having one erect orthotropous ovule, style short with 2 stigmas. Fruit a drupe or a nut. A small family of seven genera and about 60 species (Willis, 1966), distributed mostly in the North Temperate zone and in tropics of both hemispheres. Carya Nutt., the largest genus of the family, has 25 species distributed in eastern North America, Mexico, and eastern Asia. Fifteen species of Juglans L. are met in North America, the Andes, and from southeastern Europe to eastern Asia. The majority of the species of Pterocarya Kunth are confined to China and Japan. Engelhardia Lesch. ex Bl.’ with five species, ranges from the Himalayas to Southeast Asia. Oreomunnea Oerst.,” Platycarya Sieb. & Zucc., and Alfaroa Standl. are small genera distributed i in Mexico, China, and Costa Rica respectively. For its size the family is of much economic value. Juglans regia L. (English or Persian Walnut), J. nigra L. (Black Walnut), and J. cinerea L. (Butternut) produce invaluable and decorative woods which are in much demand for cabinet and furniture making. The species of Carya Nutt. yield tough and elastic wood. To a limited extent, Engelhardia Lesch. ex Bl. is the source of timber in eastern Asia. The nuts of Juglans L. and Carya Nutt. are rich in oil and very nutritious. The significant contribution in the field of cytology of the Juglandaceae had been that of Woodworth (1930), who analyzed the meiotic behavior of several species belonging to three genera. This paper presents chromo- some numbers in the two genera, Juglans L. and the primitive, tropical genus Engelhardia Lesch. ex Bl., in an effort to get a clue to the base number and phylogeny of the family; and as a prerequisite for tree im- provement through a rational breeding program. MATERIAL AND METHODS The material for meiotic studies was collected during 1965, from wild populations near Darjeeling (eastern Himalayas), Digboi (Assam), and * Spellings adapted from Dilmy (1955). ae from Engelhardia Lesch. ex Bl. by Leroy (1955) and Hjelmqvist 1960). 1970] HANS, JUGLANDACEAE 535 Shillong (Khasia and Jaintia hills). The exact source of material is given in TABLE 1. The flower buds were fixed in Carnoys fluid and then trans- ferred to 95 percent alcohol. The anthers were squashed in 1 percent acetocarmine and the slides made permanent in Euparal. Camera lucida drawings were made at a magnification of 1350 %. The voucher speci- mens are deposited in the Herbarium, Panjab University, Botany Depart- ment, Chandigarh, India. OBSERVATIONS The results of cytological studies on three species and two varieties are presented in TABLE 1 Juglans regia L.: A lofty, deciduous tree up to 30 m. in height and, on the average, 3.5 m. in girth. Bark grey, longitudinally fissured. Found in the Himalayas from 900-3000 m., extending eastward to Burma and westward to Baluchistan. Extensively cultivated for its prized wood and edible nut. 3 AO. Figures 1-5. Meiotic chromosomes in Juglans and oa ge all, xX 1350. 1. J. regia: re n = 16; 2. E. spicata var. colebrookeana: ! -I Fag 3. E. spicata: M-II, n = 16: 4. E. spicata var. aceriflora: MI, burghiana: M- 1 % = 16. ie: . E. rox- 536 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Fiower: February to April. Frurr: September to October. At M-II, 16 chromosomes were observed at each pole (FicurE 1) and pollen formation was normal. Engelhardia Lesch. ex Bl.: All the available Indian species and varieties of Engelhardia Lesch. ex Bl. were worked out. Engelhardia “acerifolia Bl.” “E. colebrookiana Lindl.”, as treated by Hooker (1885), are recognized by Manning (1966) as varieties of E. spicata Lesch. ex BI., and his treatment has been followed. Engelhardia spicata Lesch. ex Bl.: A large, deciduous tree up to 30 m. in height. Bark grayish-brown, rough, exfoliating in small flakes. Trees of this species are invariably loaded with climbers, epiphytic ferns, and orchids. Distributed in the sub-Himalayan region from Nepal eastward, and ascending up to 1800 m. LOWER: September to October. Fruir: March to April. Cytological analysis revealed 16 chromosomes at M-II at the two poles (FicurE 3). Meiosis was perfectly normal. Engelhardia spicata Lesch. ex Bl. var. aceriflora (Reinw.) Kds. & Val. (= E. aceriflora (Reinw.) Bl.): A large, deciduous tree in Sikkim, the Himalayas, and Khasia and Jaintia hills. FLower: March to April. Fruir: June to August. Sixteen bivalents were observed at M-I (FicurE 4). Meiosis was normal. Engelhardia spicata Lesch. ex Bl. var. colebrookeana (Lindl. ex Wall.) Kds. & Val. (= E. colebrookeana Lindl. ex Wall.): A small, de- ciduous tree with spreading crown; bark brown, fibrous and light red with white streaks inside. Distributed from Chenab to Nepal, also in Assam, Khasia and Jaintia hills, and Burma. FLoweEr: February to March. Fruit: April to June. Sixteen chromosomes were observed at each pole at A-I (Ficure 2). Meiosis and pollen formation was normal. Engelhardia roxburghiana Wall. (= E. wallichiana Lindl.): A large tree up to 25 m. in height, met in Assam and Khasia and Jaintia hills, growing abundantly in the second storey of tropical wet evergreen forests of Upper Assam, and ascending up to 1000 m. in Khasia and Jaintia hills, where trees grow scattered. Frower: April to May. Fruit: June to July. At M-I, 16 bivalents were revealed (FicurE 5) and meiosis was normal. DISCUSSION All the presently investigated taxa of Juglandaceae uniformly show n = 16, Sixteen is suggested as the base number of the primitive genus TABLE 1. List of the investigated taxa with their localities, chromosome numbers, and results of previous authors CHROMOSOME PREVIOUS NAME LOCALITY NUMBER REPORT 1. Juglans regia Darjeeling: Lopchu 1500 m. 2 = 16 n = 16 (Woodworth, 1930) from Arnold Arboretum; (Delay, 1947); (Mehra & Singh, 1962) from W. Himalayas 2. Engelhardia spicata Darjeeling: Mahanadi 1300 m. n= 16 n = 16 (Mehra & Gill, 1968) from W, Himalayas Lopchu 1500 m. n= 16 3. E. spicata var. aceriflora Darjeeling: Rongtong 300 m. n= 16 4. E. spicata var. colebrookeana Shillong: Garampani 750 m. m= 16 3. E. roxburghiana Digboi: Upper Dehing Reserve Forest 150 m. = 16 Shillong: Nongpoh 350 m. n= 16 [OL6t AVAOVGNVIDAL ‘SNVH Les 538 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Engelhardia. The other genera, Carya, Juglans, and Pterocarya are also based on x = 16 (Darlington & Wylie, 1955). All the known taxa are diploid ee a few tetraploid species of Carya (Woodworth, 1930), thus raising the polyploid figures to about 18 percent. Like the Lauraceae, the Fagaceae, and the conifers, the speciation in the majority of the Juglan- daceae seems to have taken place at the diploid level. Stebbins (1938, 1947, 1950) thinks that x = 10 and the higher base numbers are either primary or secondary numbers of polyploid origin; and he postulated their derivation by polyploidy from woody tropical genera yet unknown cytologically or from the extinct types. The Juglan- daceae with x = 16 could be visualized to have been derived through polyploidy during the oan or early Tertiary period either from the Myricaceae having x = 8 —an interrelationship also suggested by Hjelm- qvist (1948) on morphological grounds, or from the Corylaceae (x = 8 as cytologically evidenced by Woodworth (1930). If the above view holds, Hutchinson’s (1926) system of classification which completely separates the Juglandaceae from the Amentiferae will need some rectification. SUMMARY The cytological investigations of all the available Indian taxa of the Juglandaceae reveal that » = 16 and the plants are diploid. The base number 16 is suggested for the genus Engelhardia. Stebbins’s (1938, 1947, 1950) view of derivation of higher base numbers by polyploidy from unknown woody genera is upheld. Unlike Hutchinson’s (1926) treatment of Juglandaceae, chromosomal data at hand favor the inclusion of the Juglandaceae in the Amentiferae near the Myricaceae and the Corylaceae. ACKNOWLEDGMENTS This study was a part of the wider project dealing with the cyto- morphological investigations of the Himalayan timbers. The project was financed by the U. S. Government with PL 480 funds in India (Grant A7—-FS-12). My sincere thanks are due to the sponsoring authorities. I am thankful to Professor P. N. Mehra, Head, Botany Department, Panjab University, Chandigarh, for his permission to publish this paper. LITERATURE CITED Dariincton, C. D., & A. P. Wytte. 1955. Chromosome Atlas of Flowering Plants. George & Unwin Ltd. 519 Detay, C. 1947, Recherches sur la structure des noyaux quiescent chez les phanérogames. Revue Cytol. Cytophysiol. Vég. 9: 169-223; 10: 103-229. Dirmy, A. 1955. Valid names of the genera Bischoffia BI. and Engelhardia Lesch. Rimba Indonesia 4: 29-37. Hyetmovist, H. 1948. Studies on the floral morphology and phylogeny of the Amentiferae. Bot. Not. Suppl. 2: 1-171. 1970] HANS, JUGLANDACEAE 539 1960. Notes on some names and combinations within the Amentiferae. Ibid. Suppl. 11: 373-380. Hooker, J. D. 1885. The Flora of British India. Vol. 5. L. Reeve & Co. Lon- don HurTcHINson, J. 1926. The Families of Flowering Plants. Vol. 1. Dicotyledons. London. wie - a 1955. Etude sur les Juglandaceae. Mém. Mus, Paris II. B(Bot.). sc E. 1966. New combinations and notes on ih aati (Juglan- daceae) of the Old World. Bull. Torrey Bot. Club 93: 34-52. MEurRd, P, N., & Amar SINGH. 1962. Cytological sbservation on some North Indian trees. Res. Bull. (N.S.) Panjab Univ. 13: i7, & B. S. Gill. 1968. In IOPB chromosome edie report. Taxon 17: ae G. L. 1938. Cytological ae associated — the different growth habits in the dicotyledon : : . ao: 947. Types of polyploids. Their ee and Laie Ad- vances in Genetics 1: 403-429. cage Variation and Evolution in Plants. Columbia Biol. Ser. Columbia Gane, 5. yi ore A Dictionary of the Flowering Plants and Ferns. 7th ed. Revised by H. K. Airy Shaw. Cambridge Univ. Press, 1214 pp. WoopworrtH, R. H. 1030. Meiosis of microsporogenesis in the Juglandaceae. Am. Jour. Bot. 17: 863-869. PANJAB UNIVERSITY Present address: Botany DEPARTMENT TREE IMPROVEMENT RESEARCH CENTRE CHANDIGARH — 14 (INDIA). P. O. Box 121 KITWE, ZAMBIA 540 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 13. PHYTOCHEMICAL SCREENING AND LITERATURE SURVEY G. J. Perstnos, S. K. Curistie, J. M. Brprncer, ANp M. J. LAPIANA AS PART OF A DETAILED STUDY on the restricted flora of the elfin forest on the summit of Pico del Oeste, thirty-five species of plants were screened for the presence of three phytochemical constituents, alkaloids, saponins, and tannins. The literature was reviewed to determine which of these constituents had previously been found in the species cited or in related ones. MATERIAL AND METHODS PREPARATION OF Extracts: The screening procedures for alkaloids, saponins, and tannins were adapted from those described by Wall e¢ al. 1, 2). An extract of each plant sample was prepared by refluxing 50 grams of the air-dried, milled plant material with 300 ml. of 80 percent ethanol for one hour. Each extract was then cooled to room temperature, filtered by suction, and the residue was washed with sufficient 80 percent ethanol to bring the volume of filtrate to 500 ml. Alkaloids. Fifty ml. of each extract was evaporated to dryness using a steam bath after which the residue was stirred with five ml. of 1 percent aqueous hydrochloric acid. One ml. of the filtrate was treated with a few drops of Mayer’s reagent and a separate one ml. portion was treated similarly with silicotungstic acid reagent (12 percent aqueous). Precipita- tion or turbidity with either of these reagents was taken as preliminary evidence for the presence of alkaloids in the extract being evaluated. A confirmatory test, designed to remove non-alkaloidal compounds capable of eliciting ‘‘false-positive” reactions with either of these reagents was con- ducted in the following manner with all extracts giving a preliminary positive test for alkaloids. Two ml. of the acidic aqueous extract, prepared as described above, was treated with 28 percent ammonium hydroxide solution until the solution was distinctly alkaline to litmus paper, and then extracted several times with chloroform. The chloroform extracts were combined and concentrated in vacuo to ca. two ml., then extracted with an equal volume of 1 percent aqueous hydrochloric acid. One ml. of the separated acid extract was treated with a few drops of Mayer’s reagent, and a second one ml. portion was treated with silicotungstic acid reagent as previously described. Tur- bidity or precipitation after the addition of either of these reagents was taken as a confirmed positive test for the presence of alkaloids in the extract. The results of these tests are presented in TABLE 1. 1970] PERSINOS ET AL., ELFIN FOREST, 13 541 Each plant sample was also screened for alkaloids using thin-layer chromatography according to the method of Farnsworth and Euler (3). This procedure was modified only in that the final volume of fraction I (chloroform extract) applied to each thin-layer plate was 30 yl. The results from this test are also presented in TABLE 1. Saponins. Since all saponins, whether steroidal or triterpenoid, will hemolyze red blood cells, utilization of this property is advantageous for detecting this class of compounds in plant material. A red blood cell sus- pension was prepared and standardized against digitonin according to the protocol of Wall et al. (2). One ml. of each plant extract was mixed with 10 ml. of the red blood suspension and the mixtures were allowed to stand for one hour before observing the results. Complete hemolysis of the red blood cells in any instance was taken as evidence for a positive test, the results of which are presented in TABLE 1 Tannins. 100 ml. of the original 80 percent ethanol extract from each plant sample was evaporated to dryness on a steam bath and the residue was stirred with 25 ml. of distilled water and filtered. Two ml. of the filtrate was treated with a few drops of gelatin-salt reagent (1). Pre- cipitation along with a color reaction following the addition of ferric chloride reagent (blue-black, black, green-black) was positive evidence for the presence of tannins, green-black indicating tannins of the catechol type and black indicating tannins of the pyrogallol type. No precipita- tion with the gelatin-salt reagent and a green color with ferric chloride indicated the presence of phenolic compounds but not necessarily tannins. Precipitation with the gelatin-salt reagent and a brown color with ferric chloride indicated no tannins present. SUMMARY Of the thirty-five species of plants tested, alkaloids alone were found in two, alkaloids and tannins in three, saponins alone in one, saponins plus tannins in one, and tannis alone in thirteen. The remaining fifteen species contained none of these compounds although at least three gave tests for phenolic compounds. ae With the exceptions of a report of alkaloids on Lobelia portoricensis and earlier work on Cecropia peltata, no published phytochemical data for any of the species of the test site were found, Our findings of alkaloids in Hillia parasitica and Psychotria berteriana are consistent with earlier reports of the presence of these constituents in other species of the same genera. On the other hand, our sample of //ex sintenisu gave NO: evidence for the presence of caffeine and tannin for which J. paraguartensis 1s well- known, Alkaloids found in other species of Justicia, and the alkaloids, saponin, and tannin reported for other species of Carex, were not found in Justicia martinsoniana or Carex polystachya respectively. JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 542 1q nyjods aye avaovaav'] (s9) = 44 So - ve _ — Ws DUuDIYIDgastas DISNID AVAATILLAY) = 13 = - _ + 4. V/s WUNSOJUIUWADS WUNIPYAISOAY IAP AVANINVU®) (+9) eo Iq + - _ _ - j/98 USMAJUIS DIABUSAS) AVAOVIYANSAD) = 4q/13 + - - _ — W/s DSOmaIDA DIMUDULAULO FT AVAOVOINY = qq 5 oa a = - IW/s psoas DIPLAKD AVAIOVTIYAD ‘ets Iq - — — — -- dm supaas 1149195 (¢9) = = - - ~ - _ dm stsuanbunk stayr0a,q (79-SS) os aq — _ _ _ — V/s pk yavyskjog xasv9 AVAOVAAGAD 3 aq + -- - ~ - ns Dyjofraund DIsDg]D440,] AVANVALSVTAD (+S$-br) — 43 a as 4. ae fe Ws s1suagt40z40g DYaQOT AVIOVIN NVA) = 13 — - _ — - dm USMAqUIS DASatd A AVAOVITAWOUG (¢b-Se) _ ¥q/13 + _ = = _ js Dp1St4 DING2QD I, AVAOVINONOSIG (p£-€T) _ aq + _ _ — a j1/3s usmaqUls Xap] AVAOVITOAINOY (ZI-9) - — ~ = = ie = dm DUDINOSUIZADUL DIDI4SN AVIOVHINVOY “TAY yes it I 2 ,W aasa. AWVN OMILNAIIS "LYT SNINOdVS 3 102d Une woe. gy 4 SMLOIVATY , LavVg SNINNV, » SAIOIVATY [satoads pazelad 10 asay} IO} s}UaNqT}sSUOD snNoLIeA Jo spioda1 0} a1e sadUdIAJaI J1N}L19zT] SuTuseIIg JeIUIayI01AYg JO SINsaYyY “]T AAV], PERSINOS ET AL., ELFIN FOREST, 13 543 1970] ‘uIa13 = £(p) yuasea1 sade, 13 “ypetq = ¥q ‘Yoe[q-uses = {yso} AloyeWIyUOD UO paseq 4q/13 ‘umoiq = 1q, ‘(¢) enpasoid sang pue yyomsuiey, {(¢) poe IT}SBUNZOITIS,, *(1) aanpadsoid ‘yo 4a [Ie II®M, ‘S2AP3] pue suta}s H/4s ‘yuejd ajoym = dm, :Aay (€OI-T0T) (OOT-L6) (96) ($6-26) (16) (06-8) (¢8-08) (6L‘8L) (LL) a iq 49/13 4q/18 4q/18 + Lt++ + + + + ++ ++++ + + + +4 | ++ +41 +41 UNADIIIJUD DIMIDIUAY AVAOVAATIONIZ sisuanbunk vajtg AVaOVOILa punddyiyd sisdouydog GVIOVAVTIAWAH] DyDJIUNG OgID D4akat) avaovaH | DYJUDAINU SOI0]GUKS AVANVIOTAWAS Dyofapiutj4v3 syoydosn py AVAOVLOAVS Sisuamtsoj4og vyyajapuoy DUDIA9}49Q DIAJOYIAS G DaiqispaDg DIT] aVaOVIGNY SNJDALOGO “IBA SNYOf{1gDS sSnYyjUuDIUAD FT aVAOVATO sisuanbut4sog Diuaing usnsy Sayudsd KDI AVAOVLAATL Sisuaqjinbn] visipap AVAOV NISUAJ DyDyJag DId 04198) AVAVAO|] Dpyyog vyry 914 T AVAOVITAN DénauUourk¢ vIu0I1 LY DIDIOaLOf DIUOIT UMNUNDPSKUD WNIUDAIA unsojnuonbs uwniUu0s0IKIDI AVAOVLVWOLSV TAL 544 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ACKNOWLEDGMENT This research was supported, in part, by research grant GB: 3975 to Dr. Richard A. Howard, Harvard University, from the National Science Foundation, Washington, D. C APRONS * Beal © 10. 11. _ nN 13. 14. ARTHUR, 15. 16. LUE 18. 19, 20. Ee, 22, 23. 24. 25, 26. 27, 28. 29. 30. a1; Cay, 32. oo. 34. a5. LITERATURE CITED Watt, M. E., et al. 1954. Jour. Am. Pharm. Assoc., Sci. Ed. 43: 1-7. WALL, M. E., e¢ al. 1952. Anal. Chem. 24: 1337-1341. FARNSWORTH, N. R., & K. L. Euver. 1962. Lloydia 25: 186-195. 0-29. . Mayer, F. F. 1863. ‘Am. Jour. Pharm. 25: 20- : SWANHOLM, C. E., H. Sr. Joun, & P. J. ScHEvER. 1959. Pacific Sci. 13: 295-305. Tsukamotu, T., & Y. Kisurmoro. 1955. Jour. Pharm. Sci. Japan 75: 1565-1567. Menta, P. R. 1960. Indian Jour. Appl. Chem. 64: 603 SpATH, E., & F. KeszTLER-GANDINI. 1960. Monatshefte Chem. 91: 1150, ELST. JAIN, ie & A. K. BHATTACHARYA. 1960. Indian Jour. Appl. Chem. 23: 203- Ota, M. eo &Y. CHEN, 1965. Sea Letters 47: 4167-4170. IKRAM, M., E. S. Hug, S. A. Warsr, & V. Anmap. 1965. Pakistan Jour. Sci. Ind. Res. 8(3): 76-79. . Sippigut, R. H., & I. 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Fac. Ing. Quim. 28: 9-17. Narr, A. G. R., & S. S. SUBRAMANIAN. 1962. Jour. Sci. Ind. Res. (India) 21B: 457, 458. Friese, F. W. 1935. Pharm. Zentralhalle 76: 223-225. VIEHOEVER, A., & C. O. Ewinc. 1921. Jour. Am. Pharm. Assoc. 10: 763- 766. KELLER, O. 1913. Arch. Pharm. 251: 701-713 MAaAcENDIE, F., & J. PELLETIER. 1817. Jour. Pharm. sae 3(2): 145-164. Paul, B. H., & J. J. Cownsy. 1896. Pharm. Jour. Hut, W. H., & C. W. YEE. 1967. Phytochemistry A eis 442. NisHipa, K. 1951. Jour. Japan Forestry Soc. 33: 269-272. NisHipA, K. 1951. ao Bie Meeting Japan. eae 7 312,313. HorHAMMER, L., & S. Rao. 1954. Arch. Pharm. 287: Lynn Index, in press. Ibid. PIREYRE, N. 1953. Compt. Rend. 236: 1595, 1596. PCREQUIN, T. 1954. Compt. Rend. 238: 2125-2127. NATURAL PRopucts RESEARCH LaporaTory [G. J. P. & M. J. L.] 809 VeIrs Mii Roap ROCKVILLE, MARYLAND 20851 P. Ropertet Co. [S. J. C.] NEw York, N. Y. NATIONAL INSTITUTES OF HEALTH [J. M. B.| AND BETHESDA, MARYLA 1970] INDEX 547 INDEX oe in the Southeastern United States, The Gene fee of, 257 Acanthacen, 257-30 — subfa ay ait i ek 281-283 fees Nelsonioideae, 278-280 —subfam. Ruellioideae, 283-309 —-subfam. Thunbergioideae, ae 280 —— age Ruelliinae, 285-292 Acanthus, 258 eeughce Family, 257 desis tiene 208-214 5 20, 2014213 gage er ley 210 —endlicheri, 209 10 a lelocarpa, a Ha, 214 — ligustroides, — ovalifolia, 2 i — ovata, oe — petiolaris, 212 simplicifolia, 209-2 —— subsp. neo- sii 211, 212 — — subsp. petiolaris, 212, subsp. ca age 209-211 Asia 93, 94, 404 —arnottianum, 40 — costatum, » 404, 409 Actinidia, 4 ctinocarya, 139 eerie Nout on the cers aes. of Zanthoxylum (Rutaceae), 4 Prevesti brevicaulis, 66, 67 Adenogramma, 432 Adiantum, 73, 77 — capillus-veneris, 77 — cooperi, Afrolicania, 528 Agavaceae, 490 Agave, 490 Aizoaceae in the Southeastern United States, The Genera of Molluginaceae and, 431 Aizoaceae, 443, —subfam. a se 444, 449 —subfam. Aptenioideae, 444 —subfam. Caryotophoroideae, 444 —subfam. Hymenogynoideae, 444 —subfam. Ruschioideae, 4 — tribe oie: 450 Aizoon, Alfaroa, yen 469, 473, 475 — costaricensis, 465, 466, 469, 472 Alobiellopsis dominicensis, 66 Aloe, 490 Amentiferae, 538 Apetalous Genera of i eicngnatet Floral Morphology and Vascular foie omy of hae Hamamelidaces, The, 3 Aphragmi Ardisia ugullnsi 543 Arnebia pete Laie 286 Arthrost sylidiom tabi 542 542 rugo, 136 Asystasia, 298, 299 Axis of Dracaena fragrans (Agavaceae) The Vascular System in the is- tribution and Development of Second- ary Vascular Tissue, 478 AYENSU, ARD nalysis of t Complex Vascularity in Stems of Dio- scorea composita, 228 Bauerella australiana, 210, 211 — baueri, ueNN breutelian, 63 —lon —lon es 64, 65 roraimensis, 65 548 Bazzania a 65 Beaucarnea, Beloperone, re ttata, 304 Beurreria bes 147 oy BIpIN ex, J. 9 and M. J. Lapiana, G. fe Hades oS. K. Curistiz. The Ecol- ogy of a n Elin Forest in Puerto ne 13; Siveatadiied Screening and L erature Survey, Bignoniaceae, 542 Bignonia sempervirens, 4 Blackburnia pinnata, a: 216 5 ener ot aged and Vascular elie Hamameli- daceae: The eed pti i Ha- mamelidoideae, 310 BocLs, bie INN. The Genera of Mollu- ginaceae and Aizoaceae in the South- panna iid States, 431 Boninia, Botectniceke of West Pakistan and Kash- mir, A Revision of the, 133-184, 367- 402, 499-520 Bothriospermum, 137; 1822186 ellu Br naa A nig Species of Ormosia from, Breeding System of Gelsemium — niaceae), The Systematics and, heme 542 Caccinia para rata, 304 Calophanes, 293, — angusta, 293 Calycogonium squamulosum, 543 Calypogeia cellulosa, 64, 65 —e oe echniaee 65 Calypogeiace hy Calyp thes Oe i, 543 Calyptraria fan te 226, 227 Campanulaceae, 542 Cannabaceae in the pty eee ty United States, The Genera uf a” Cannabis, 186, ae Carex polystachya, 541, 542 Carpet-weed, Carpinus betulus, 472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Carya, 463, 467, 472, 473, 475, 534, 538 — sect. Rhamphocarya, 463 — amara, 469 Raven 465, 467, 468, 472 — ania. 464, 465, 467-469 — illinoensis, 467 Bi omnes 465, 467, 468 467 Central America, Notes on the Genus Galipea (Rutaceae) in, 427 Cephalozia a a 65 Cephalozia Cerato olejeunea ‘Fagellifainis. 64, 67 — mari — patentiadma, 67 — valida, 67 Caristie, $. K., J. M. sane and M. Laprana, G. RSINOS. The Ecology of an Elfin Fo ir in Puerto Rico, 13. Phytochemical Screening and Literature Survey, 54 Chromosome Numbers in the Juglanda- ceae, 534 Chrysobalanaceae, 521 Chrysobalanaceae in the Southeastern United States, ae Genera of, 521 Chrysobalanacea — tribe inv 521, 528 tribe Hirtelleae, 521 Chie Soe 527, 528 ieee Typ ein-endings the Dic notamenl Venation of, 70 Cicaeate, 70-88 ti agrestis, Clevels sional 543 Clock-vine, 274 Clusia oD 542 Coelanthum, Coldenia, 135, a rocum 148 67 parative gra Studies in Dini aceae, V. Leaf Anatomy, 89; VI. Sta ona Young Stem, 403 eine Vascularity in Stems of Dio- scorea composita, Analysis of the, 228 Ss a 1970] Conve, Louis F., and Donatp E. Stone. Seedling Morphology in the Juglanda- ceae, the Cotyledonary Node, 463 eer aeeny 412 Corbichonia, 432 inven 135, ‘139- 145 Me ore 140 naar — ee ae 141 Beier 4 140, 143, 144 — myxa, 139, 142, 143, 145 — obliqua, 140, 142, 143 — — obliqua, 143 , 143 — vestita, ro 144, 145 — wallichii, 143 Cordyline, 490 Cornus gharaf, 140, 141 — Sanguinea, te. 141 Corylaceae, Cotsednery Node, Seedling eile he Juglandaceae, The, es BarparA, Ray MARGARET Peo pak cology of an Elfin est in Puerto Rico, 11. The Leafy ions icae e Pico del Oeste, 56 Craniospermu sangha: 368 ee and ne a, 90, 93-96, 100, 101, 403, 406 — ameri sas 417, 419, 421, 422 Cusparia, 427 Cynoglossospermum consanguineum, 507 Cynoglossum, 139 — diff 182 —Jaxum, 398 — macrophyllum, 397 39 — — var. laxiflorum, 397 INDEX 549 Cyclolejeunea accedens, 62, 67 Sas . wens hier 241-244 , 68 yst ee linea ee oe 543 Davide Davilla, 90, pe ee is 406 06, 417, aa a a a Demidovia, 460 Derris from the Solomon Islands. Studies in the Leguminosae, 11., A New Species of, 251 Derris sect. Dervis. 251, 252 — sect. ora 251, 252 — elegans, — multiflor: = on 252 salomonensis, 251-25 esr ent of Secondary Vascular Tis- sue, Vascular System in the Axis of Dracaena fragrans (Agavaceae) 2. Dis- of Circaeaster, T yDes of Blind Sana in the, 70 DickIs Co omparative ia Hae in Dilleniaceae, VI. Stamens and has Stem, 403 DICKISON IAM omparative Morphological Studies in Dilleniaceae, . Leaf Anatomy, Dicliptera, 300-302 — sect. Dactylostegiae, 301 — sect. Dicliptera, 3 —-sect. Platystegiae, 300 — sect. Sphenostegiae, 301 — halei, viridiflor Didesmande 90 94, 100, 405 Dillenia, . wes v4 any — alata, 417, 419 — aurea, 404 — excelsa, 96, 417, 421 — fagifolia, 404 — indica, 96, 407, 417, 419, 422 a — ovata, 0g, on Th 420 550 Dillenia papuana, 96, 404, 417, 419 — parviflora, 404, 417, 419 — philippinensis, 93, 96, 101, 417, 420 — pulchella, 94, 96, 100, 101 pie sas 93, 96, 100 suffrutic Dileniaese, V. Leai Anatomy, Compara- e Mo oe Studies in, 89; 3, s and Young Stem, Comparative noire ae Studies i in, 403 Dilleniaceae, 89-9 ubfam. Dillenioideae, 404, 409 hie Tetraceroideae, 403 — or Hibbertieae, 409 — tribe Tetracereae, 409 pileanice 412 Dillenites, 411 Dio: pacorea pl So Analysis of the Vas- cularity in Stems o Dioscorea Gam 228- 240 Bisnis 236 — preussii, 23 Bardeen 23 aa unidetata 62, 68 pel hing — ciliosus, 288 biaaicn and Development of Secon- dary Vascular Svan The Vascular nace in the Axis of Dracaena frag- s (Agav sulin = Distlum,. 3135314, 316-324 — buxif ; Paernny cianeeae 406, 417, 419, 420 — guianensis, 96, 10 — lasiogyne, 96 — major, 91, 98, 417, 421 System in the Axis tribution and Development ft Seco dary Vascular Tissue, 478 Dracaena, 478, 482, 490, 491 — fragrans, he 480, 487-490 Drejerella, 30. ttata, esl Drepanolejeunea anoplantha, 68 — crucia nella, s, 68 ae ayy 62, 68 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 onal 292-296 ae er ——an —— cies a 294 Echinospermu — sect. roo n 383 — sect. Heterocaryum, 515 — barbatum, 508, 509 — canum, 390 —cariensis, 509 — caspicum — divaricatum, 502 —echinophorum var. sessile, 516 — filiforme, 508 = saa noi, 398 — laxiflorum, 397 04 = eavite ned: 517,518 —thymifolium, 396 Echiochilon, 136, 183 — kotschyi, 184 — persicum, 183, 184 Ecology of an Elfin Forest in Puerto ew Species of Gonocalyx (Ericaceae) 2213 edoanageye Screening and b icaeeee Survey, Edwardsia hersnseibe, 204 Ehretia, 135, 145-1 oe acuminata var. serrata, 145 — affinis, 147 — Wd. 147 1970] Ehretia floribunda, 147 — laevis, 145, 147, 148 — — floribunda, 147 — — platyphylla, 147 Mires, me — obov aa oom es 147 — punctata, — serrata, 145, Eleocharis eats 542 Elephantorrhiza, 114 — elephantina, 114-125 Elephantorrhiza elephantina, The Mor- and Germination of the seed of, 114 pene ne - Puerto Rico, 11. The Ecol- The pay Hepaticae of Pico del ae 56 ew Species of oo alyx (rice), 221: Notes on Two Spe of Marcgravia, 41; SHO ol ree and Literature oe Exias, THOMA he Genera of Ulma- ceae in the oes United States, ne Tas, THOMA , Notes on the Genu ‘Gone (Ratacese) in Central America, 427 Elm, 18 shia ores i Endoge Sane ser eee 538, 463, 469, 473, 475 — sect. Engelhardia, 469, 473 — sect. Oreomunnea, 463, 469, 473 — sect. asia aa 469, 473 — roxburghiana, Pai ce 466, 469, 470, - a — serrata, 469 pe §36, 537 — — var. aceriflora, 535-53 2: enlebrookenna, 536, 537 =alchian, 53 — elongatum, 374 INDEX 551 par naa cen ai 385, 387 84, 385 subsp. mae var. euvillosum, 386 bare 3 nanum, 386 — — subsp. villosum, 386, 387 Reccemgn hs 390 Sutban 6 Erigenia borinquensis, 543 Esenbeckia Evodia, jee Fagaceae, 472 Fagara ie eas 215, 216 — nadeaudii, 2 — pinnata, 214-216 Fagus oo 472 Fitchia, 4 gn Saree and Vascular Anat- of the Hamamelidaceae: Mortals Genera of Hamamelidoideae, 310 > oO Floras of Norfolk and Lord Howe Islands, I., Notes inaesing to the, 204 Foster, ADRIA S. Types of Blind Vein- uae E in ae Dichotomous Vena- Frullania subtilissima, 67 Frullaniaceae, 6 ULFORD, MARGARET, BARBARA CRANDALL, 0 E aut Satan of Pico del Oeste, 56 seach 444, 449 a, 450 Galipea (Rutaceae) aie America, Notes on the Genus, 552 Galipea, bat tei — guatemalensis, 427 — jasminiflora, aah 428, 430 — nicaraguensis, — panamensis, th 48 Be mien 428 — tri 427 tte es ue Gatesia laete- virens, 301 Gelsemium (Loganiaceae), The Systema- tics and Breeding System of, 1 —sumatranum, 10 Genera of Acanthaceae in the Southeast- ern United States, The, 25 Genera of the Cannabaceae in Vy South- eastern United States, The, 1 Genera of Chrysobalanaceae i in ca South- Genera of Moll oaceae 282 Germination of the Seed of Elephantor- rhiza elephantina, The Morphology a > Gesneriaceae, 542 Gesneria sintenisii, 542 Gieseckia, 436 ILLETT, GEORGE Taxonomic Stat of Protocyrtanidik (Gesneria- eae rey i ae Gisekia, 432, 435, 436 Glinaceae, 43 Glinus, 432, 439, 441, 442 442 Gonocalyx panies A New Species ts) n Elfin Forest in uert co, Conceal concolor 22 2 24 — port: is, 221-225 a sodatek: 222. "924-227 — smilacifolius, 222 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Gramineae, 542 GrEEN, P. S. Notes Relating to the Floras of Norfolk and Lord Howe Islands, Guttiferae, 542 Gynaion oes. 144 Hackberry, 33 Hackelia, 139, 395 eflexa, 3 —— pumila, ee Ss — glochidiata ican Ne aa 397, 398 — meeboldii, 395, 397 — roylei, 398 — stewartii, 395, 399, 400 — thymifolia, 395-397 —uncinata, 395, 398, 399 Haenianthus salicifolius var. obovatus, 543 Halimum, 451 Halimus, 451 Ss he Apetalous masa of H meldideae Floral Morphol- ogy a aia r Anatomy of a 310 anchor Floral Morphology and Vascular Anatomy of the Hamamelida- ceae: The Apetalous Genera of, 310 Hamamelidaceae, 310-365 — tribe Liqui — tribe Rhodolei ideae, 314 — tribe Sy = fea 314 ANS, mosome Numbers in the eli ucee. 534 Haploid and spec Pollen in Hypericum patulum ies heterodonta, 68 —uncinata, 68 Harttey, THomas G., Additional Notes on the Malesian Species of Zanthoxy- Havilandia, Hedycrea, 526 Heliotropium, eh 149-180 — afghan sodas a Shae eri, 151, 168, ea — bacciferum, ‘151, — — subsp. ee et 162, 164 63 164 — — bacciferum, 161, 162, 164 1970] INDEX 553 Heliotropium baccif fartal ,163,164 Heliotropium ramosissimum, 158 —_—— lignosum, 163, — rarifolium, 149, 151, 152 — — tuberculosum, ve 164 — rechingeri, 166, 167 — baluchistanicum, 150, 172, 174 eee 150, 171 — biannulatum, 150, 172-174 — rottler — brahuicum, 151, 166, 167 — eee 158 — brevifolium 3 — sericocarpum, 169 — brocchianum, 178 — strigosum, 149, 152-154 — cabulicum, 150, 169-171 —— subsp. brevifolium, 153, 154 — calcareum, 150, 174, 175 — — subsp. strigosum, 152-154 — chenopodioides, 179 — — brevifolium, 153 — coromandelianum, 178, 180 —subulatum, 151, 157, 158 ——a obovatum, 1 —supinum, 179, 180 —crispum, 151, 158-160 — — var. malabarica, 180 — curassavicum, 149, 178, 179 — syenites, 178 —— obov 79 — taftanicum, 167 ve — tenuiflorum, 176 —— zeylanicuns; 155; 157 — tuberculosum, pe — dasycarpum, 151, 164-167 — turcomanicum, 1 — — dasycarpum, 165, 166 —ulophyllum, 150, es —— gymnostomum, 165, 166 —undulatum var. eT ACRIECE 158 — dicriophorum, 172 — — tuberculosum, 162 — ditrichum, 160 — virens, 179 — eichwaldii, 177 — de 149, 155-157 — — lasiocarpum, 176 —— subsp. paniculatum, 156, 157 — ellipticum, 150, 177 —_—— nar zeylanicum, 156, 1 — — lasiocarpum, 176 Hemigymnia eee — europaeum, 150, 175-177 Hemp, 185, 189-1 —-— europaeum, 176 Hepaticae of ee Oeste, The Leaty. — — lasiocarpum, 176, 177 si Ecology of an Elfin Forest in — fartakense, 163 pee £1556 — fruticosum var. persicum, 183 Slade e, 63 — gillianum, 150, 167, 168 He soo sp mcenee 63 — glaucophyllum, 179 nsilis, 63 — glaucum, 179 a 138, 515 —gymnostomum, 165, 168 — divaricatum, 503 — — aucheriiforme, 168 — echinophorum var. minimum, 517, 519 — khyberianum, 173, 174 — — oligacanthum, 516 — kotschyi, 162 — pachypodum, 519 — kunzii, 178 aviaernats ns, 504 veocDarnaiistlod: 151, 168, 169 — laevigatum, 515, 516 — -sabaag 176 — minimum var. ot 519 — lignosu — oligacanthum, 504, — linifoliur, ae £57 — pachypodum, 518 — littorale, ea — —kotschyanum, 519 — malabaricum, 180 — rigidum, 515, 51 _ ae 149, 154, 155 Soni 51 — — subsp. marifolium, 155 — szovitisianum, 515, 518 — — subsp. wallichii, 155 Heteroscyphus elliottii, 66 — multiflorum, 174 Hibbertia, 89-97, 99, 100, 404, 406 — niloticum, 178 — acicularis, 9 00 — ise sree 158 — eo es 98 ophioglossum, 149, 160, 161 — au ine ipo 150, 178 aiid) 93, 94, 96, 99 — paniculatum, 1 oe — cistifolia, 100 — persicum, 1 158, — coriacea, 101 — popovii sube. eesneare 167, 168 — crenata, 99 554 pas onieerd cuneiformis, 96, 99, 407 — dentata, 407, 417, se 420, ra — exutiacies, 417, 419 Serene 92, 98 — saligna, 92, 99 Riegel eh 100, 404, 407, 414, 417, 19, 422 — sericea, 96, 99 — stirlingii, 101 — subvaginata, 91, 100 — teretifolia, 96, 97 sie kena ee 407, 417, 419 Feocione —unc Hillia praia, 541, 543 a 6, 199, 200 nemannia nore i 542 Aged purslan Howarp, a A. The Ecology of an re Forest in Puerto Rico, 10. Notes on Two Species of ee ine 41 aicahe 186, 188, 196-20 — sect. Hygrophila, 29 Hymenanthera dentata var. oblongifolia, Bis aaiags 218, 2 e-zelandiae, ee lee sorry 220 Hypericum patulum, Haploid and Diploid Pollen in, 247 Hypericum patulum, 247-250 — ‘Sungold’ 248, 249 Hypertelis, 432, 439 Tlex paraguariensis, 541 —sintenisii, 541, 542 Indian chick-weed, 439 Tsotachaceae, 63 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 ee baueri, 210 , 210 ffersonia, 4 Juglandaceae, 469, 472, 534, 538 ae ceae, Chromosome Numbers in , 534 oe ee The Cotyledonary Node, Seedling Morphology in the, _ 463, 467, 469, 472, 473, 475, 534, 535, — ailantifolia, ptt 467-469, 472 — olanchana, 470, — regia, 467, 469, et 535,537 6 — crassifolia, 305 — guttata, 304 — humilis var. scien 303, 305 — martinsoniana, 541, 542 — ovata var. angusta, 303 — — lanceolata, 303 Kashmir, * abot of the Boraginaceae Pakistan and, 133-184, 367- A Revision of the Roruplnncene of West Pakistan and Kashmir, 133-184, 367-402, 499-520 Kingdonia, 76, 77 — uniflora, 80 Laprana, M. J., G. J. Persinos, S. K. Curistie, ‘e oy “BD IDINGER, and. The Ecology of an Elfin Forest in Puerto Rico, 13. Phytochemical Screening and Literature Survey, 5 Lappula, 139, 499, 515 — albiflora, 5 — barbata, 500, 508 mae 500, a 1970] Lappula a 502 — echinata, 509, 510 — macrop. , 39 — microcarpa, 499, 504, 513 — myosotis, 500, 51 — patula, 500, 50 Lasiocaryum, 137, 392, 393 — densiflora, 393, 394 — munroi, 392-394 Lauraceae, 542 Leaf Anatomy, Comparative Morphol- ogical Studies “a Dilleniaceae, V., 89 Leguminosae, 204, 22 Leguminosae, 11. pops in. A New Species of Derris from the Solomon Isla sa Lejeuneaceae, 67 Lepechinella, he S11 Fo ninco idboss 66 — S21, — subg. soe 367 INDEX 350 Lithospermum apa 372 — persicum, 183 — secundiflorum Lobelia ee 541, 542 Lomandra, 490 ONG, ROBERT The Genera of Acan- thaceae in the Southeastern United t tates, 257 Lophocoleaceae, 66 Lophoziaceae, 66 Macarthuria, Malesian ee of Zanthoxylum eal ceae), Addit Pes Notes on the, 4 Malvaceae, 206, 20 Marcgravia, 41-55 Marcgravia, Notes on Two Species of. The oe . an Elfin Forest in Puerto Rico 41. Marcgravia ser tunel 44 Marcgravia subg. Plagiothalamium, 44 — nepenthoides, 46 — rectiflora, 42, 44, 45, 47-55 — sintenisii, 42, 44, 45, 47-55, 542 Marcgraviaceae, 41, 542 92 Melicytus latifolius, 218 vae-zelandiae, 219, 220 —— subsp. centurionis, 219 — ramiflorus sp. oblongifolius, 220 — elongata, 374 556 Mertensia exserta, 368 — moltkioides, 378 378 — strigosa, 372 = eran =e — trollii, nt an Family, 443, 444 Messerschmidia hispida, 157 Mexico, A New Species of Parietaria (Ur- ticaceae) from Northeastern, 529 Miconia foveolata, 543 —pycnoneura, 543 Microcaryum duthieanum, 394 Micropholis garciniaefolia, 543 Microula, 139, 400 — benthamii, 400 — tangutica, 400 — tibetica, 400, 401 Micr “aeeagr carinatum, 65 Millegrana MILLER, vel The Genera of the ORTON G. Cunnahivens ' in the Southeastern United ciate 185 , Norton G. New Species of Pare aria (Urtacee from North- astern Mex 529 nF emia vin 125 Molinadendron, 3 ve 314, 351-354 351 Molluginaceae and oaceae in the moo ene United pais. The Genera of, 431 Molluginaceae — tribe conned: 432 — tribe Gisekieae, 432 — tribe Mollugineae, 432 — tribe O ae, 432 Mollugo, 432, 437-439, 442 Moltkia parviflora, 368 — trollii, 369 Momisia, 33 Moquilea, 526, 527 Moraceae, 543 Mechoiial Studies in Dilleniaceae, Comparative, V. Leaf Anatomy, : VI. — and Young Stem, 4 Morpholo Germination of the Seed a crehunisetitcs elephantina, The, as in the Juglandaceae, the Cotyledonary Node, Seedling, 463 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 Myosotis, 138 — deflexa, 395 — lappula, 510 — longifolia, 390 My Myrtopsis, 428 nai bicuspidatus, 63 Nettle-tr sr bei be aap 68 NEviinc, Lorin » JR, The omagid of an Elfin Forest in Puerto pide. New Species - panne oo, New Species of Derris from the Solomon Islands, A. Studies in the Leguminosae, TE 258 New Species of Gonocalyx (Ericaceae), A. The Ecology of an Elfin Forest in Puerto Rico, 12, 221 New Species of Ornstaia from Brazil, A, 129 New Species of Parietaria (Urticaceae) from Northeastern Mexico, A, 529 New Zealand spinach, 460 Nonea, 136 Norfolk and Lord Howe Islands, I., Notes Relating to the Floras of, 204 Genus Galipea (Rutaceae) ral America, 427 Notes ie to the For pA — and Lord Howe Islands, ste on Two Species of eee avia. The Ecology os beg Elfin Forest in Puerto Rico, 10. Nowellia ee 65 — dominicensis, 65 Ocotea spathulata, 542 Oleaceae, 543 Odontoschisma prostratum, 66 Odontoschismaceae, 66 Omphalodes, 138 — thomsoni, 389 ? Oreocharis, 367 Oreosenia 92, 393 Bos * nroi, pecans yn 534 1970] Ormosia from Brazil, A New Species of, 129 Ormosia friburgensis, 129, 131 —131 . Relationships in the Piriqueta caroliniana~Piriqu - cistoi- des Complex (Turneraceae), OrnpuFF, Rosert. The sacl and aha System of Gelsemium (Loga- eae), rae aa 207, 208 Oxalis clues 207, 208 —— culata, 207, 208 ie cnpenratn 207, 208 — — reptans, 207 — reptans, 207, 208 Pachynema, 405 — dilatatum, 406, my 419 Paracaryum, 138 — olachidiatim 208 — inconspicuum, 512 microcarpum, 513 Scena 528 Parietaria, 529 Parietaria (Urticaceae) from Northeast- ern Mexico, A New Species of, 529 dee subg. Euparietaria, 532 shy hip 532 — ~~ debi ~ decor 9, 5392532 — flori — ee te 531, 532 — obtusa, 52 — offici nalis, 529 Reams 529, 533 Parrot, 313, 314, ae — pers 25,733 a Parrotiopsis, 313, 314, oe — jacquemontiana, 347-351 Pedinogyne, — tibetica, 402 Pelea, 428 Persinos, G cae pene J . M. J. Lara The ees of an Elfin Forest in ghee Rico, 13. Phytochemical Screening and Literature Survey, 5 Pharnaceum, 439 ie deo Screening and Literature Survey, 13. The Ecology of an Elfin Forest in ABs Rico redO Pico del Oeste, The Leafy Hepaticae of. INDEX 557 The Ecology of an Elfin Forest in Puerto nana se Pilea yunquensis, 543 eS ina mein 180 — supin Paes 492-498 Piriqueta caroliniana—Piriqueta cistoides Rig ende (Turneraceae), Relationships Pirie aap 492, 493, 495-497 roliniana, 493, 494, 497 Piriqueta cistoides omplex (Turner ceae), Relati one hs in the Piriqueta caroliniana-, 4 Piriqueta cistoides, 492, 493, 495-497 494 Plagiochila bidens, 66 — bursata, 66 — chinantlana, 66 PI Planera, 21, Platycarya, ne ee , 472, ube ee 534 — strobilacea, 464, 465, 46 Shuiviole 490 Plinthus, 450 Pollen in Hypericum patulum, Haploid and Diploid, 247 Prance, GHILLEAN T. The Gener Chrysobalanaceae in the 5 caueaiers United States, 521 Prionium, 230, 236, 238, 491 — serratum, 489 Prionolejeunea aemula, 64, 68 Prototanda rags maga The Tax- c Status 1 Piracy ae — todaiensis, an Psammotropha, hes oem ae By, 367-383 — anjumiae, 368, 377 — chitralensis, 368, 376 — drummondii, 368, 375 — echioides, 136, 367, 372 — — echioides, 373, 3 — — Jahulensis, 374 558 Pseudomertensia sr 371 — var. trollii, 370 Psychotria ee ea 541, 543 Ptelea pinnata, Pterocarya, ne eis 469, 472, 473, 534, 538 — fraxinifolia, 464 — rhoifolia, 464, 465 repeat 464-466, 470, 474 Puerto Rico, 10. The Ecology of an Elfin Forest in, Notes on Two Species of es ie 41. The Ecology of an Elfin Forest in, Be Leafy Hepaticae of Pico del Oeste, 56 —12. The Ecology of an Elfin Forest in. A new species of Gonocalyx (Erica- ceae), 221 —13. Phytochemical Screening and Lit- erature Survey, 540 Radia Rectolejeunea phyllobola, 68 Relationships in the Piriqueta caroliniana— P. cistoides complex (Turneraceae) , 492 Renealmia antillarum, 543 Revision of the Boraginaceae of West Pakistan and Kashmir, A, 133-184, 367- 402, 499-520 scprmine 481, 483, 490 excelsa, 230, 236, 238, 239, 489 5 Rondeletia portoricensis, 543 Rosaceae, 521, 522 subfam. Prunoideae, 522 JOURNAL OF THE ARNOLD ARBORETUM [vou. 51 Rubiaceae, eo Ruellia, 285-29 — sect. aes 289, 291 caroliniensis var. — hybrida, 288 nedn Rutaceae, 208-218 —tribe Cusparieae, 427 Samara blackburnia, 215 Scapania iecctonnats, 67. 7 405, 406 ae cern 91, 94, 96, 417, 419, 421, 422 Scleria secans, 542 Sclerocaryopsis spinocarpos, 500 Sea purslane, 451, 455 Secondary Vascular Tissue, The Vas e “ge of Piecic aena , Distribution Smilax megalantha, 238 Snymawn, L. and H. P. van DER ScHIJFF. Komphulogs and Germination of the Seed of Elephantorrhiza sehaatiens 11 Socratea, 279, 481 Solenanthus, 139 Sole enmationn: 33 Solomon Islands, A New Species of Derris — the. Studies in the Leguminosae, 1 25 Bibi 205 — sect. ng ag 205 — howinsula, 2 1970] Sophora microphylla, 205 — tetraptera, 204, sp. howinsula, 205 — — howinsula, 205 subsp. macnabiana, 205 Southeastern United States, The of Acanthaceae in the, 257 Seni iacene United States, The Genera Cannabaceae in the, 1 Pe el United States, mo of Chrysobalanaceae in the, ee hari United States, me Genera of Molluginaceae and Aizoaceae in the, 431 ——su Genera Genera Southeastern United States, The Genera of Ulmaceae in the, 18 Stamens and Young Stem. Comparative eg a Studies in Dilleniaceae, VI, Stenandsum, 281-283 sect. Sphaerostenandrium, 283 ran Stenandrium, — dulce var. floridanum, 281, 282 Stone, Donatp E,, and Louis F. Conpe. Seedling Morphology in the Juglanda- ceae, the Cotyledonary Node, 463 STOTLER, Leafy Hepaticae of i del Oeste, 56 Strepsilejeunea involuta Studies in the Le acres 11. A New Species of Derris from the Solomon Islands, 251 Suessenguthiel, 432 Sugarberry, 33 Sattatila > tenuifolia oparrotia Ecidetke 325 a, — sinensis, as, 330-334 — tutcheri ne a Sym plocos me ete ke eae of Gel- Loganiaceae), T 1 Sie lle perfoliata, 66 Tabebuia rigida, 5 Taxilejeunea ae Taxonomic atus of Pres adie (Gesneriaceae), The, 241 INDEX 559 Telaranea sejuncta, 65 Tetracera, 90, 92-94, 99, 403 — akar = hoteieaa: 99 — indica, 96, 99, 100, 407, 417, 419, 422 — leiocarpa, 92, 99 — macrophylla, 407 — ovalifolia, 92, 99 — portobellensis, 92, 99 — scandens, 96, 99, 407 — volubilis, 417, 419 Tetractomia, 42 . Tetragonia, 444 — subg Tetragoniaceae, 4 Theaceae, 543 Tuomas, Joas L. Haploid ni Ph it Pollen in Hypericum patulu TuoTHaTHel, K. eat dies in the yh m- inosae, 11. A ree es? ‘ Derris from the Sant Bland 2 ppc 273-27 sect. Euthunbergia, 274-277 — sect. poecneae 216, 277 ie 274 Thunbergiaceae, 258 Thymelaeaceae, 543 Tipalia limonella, 424 Tomuinson, P. B., and M. H. ZIMMEr- 2 epi email 157 — roylea se 7 — subu at — evan, e. Trachylejeunea a 69 Traxilum, 145 — pratensis, 4 — subgen. Panedasin, 455 Trichodesma, 136 Trigonotis, te 401, 402 560 ee clavata, 401 — tibet 402 penidies, 27 Turneracea raion a tionships in the Piri- queta_ caroliniana—Piriqueta _cistoides complex, 4 Types of Blind Vein-endings in the Dichot- omous Venation of Circaeaster, 70 Ulmaceae, 18-40 in the Southeastern United States, the Genera of, 18 Ulmaceae tribe Celteae, 32-40 — tribe Ulmeae, emer 21-30 t. Blepharocarpus, 23, 24 elea, 23, 24 us, 23 — sect. Trichoptelea, 23, 25 sect. Ulmus, 2 ‘erechccas, 543 VAN DER sls ie H.. P. and a ‘a segie The Morphol ane ‘Ger tion of . seed of oat: erate cuetestng ee sinensis, 141 Anatoury of Vascular Hamameli- daceae: The Apetalous Genera of Ham- amelidoideae. oul Morphology and, 10 Vascular System in the Axis of Dracaena fragrans (Agavaceae) 2. Distribution and siting * of Secondary Vascu- lar Tissue, 478 Vepris simpli 209 Violaceae, 218- Vriesea peg ‘ Water Elm, 31 Water-Willow, 303 JOURNAL OF THE ARNOLD ARBORETUM [voL. 51 West Pakistan and Kashmir, A Revision of the Boraginaceae of, 133-184, 367- 402, 499-52 Xanthorrhoea, 4 Xanthoxylon backbuni, 215 — howeanum, ipsa G. New Species of Ormosia from Brazil, 129 Yeatsia eth -virens, 301 Yucca, 490 Zamia, 77 Zanthoxylum (Rutaceae), Additional Notes on the Malesian Species of, 423 Zanthoxylum, 214-218 —acanthopodium, 425, 426 —avicennae, 423, 426 oa “apa 214-216 — budrunga, 425 Se ean, 426 — forbesii, 424, 42 Ce 425, 426 —limonella, 423, 424 inn onectiele 423, 424, 426 — nadeaudii, 215 —nitidum, 423, 426 — ovalifolium, AS geP — pancheri, 216-21 —pinnatum, 214-2 e — pluviatile, 425, 426 — retroflexum, 424, 426 — rhetsa, 424-427 —scandens, 426 Zz ERMANN, M. H., and P. Tom- LINSON, cular System in the Axis of Dracaena fragrans ein eae) Coca ii est mr visa of ondary Vascular Tissu Zingiberaceat, 543 Zoelleria, 401 Zoopsis antillana, 66 Zoopsidaceae, 66